Bulletin of the British Museum (Natural History). British Museum (Natural History) Converted as part of the ABLE project by Dauvit King London : BM(NH) Continued as: Bulletin of the Natural History Museum. Entomology series Vol.1 (1950) - vol.61 (1992) 2 4 This document has been converted to TEI XML as part of the ABLE project to make it more widely available to biodiversity researchers in a useful format. eng text No corrections have been made in the text. The original source has not been regularized or normalized. Quotation marks have not been processed. They are as in the original DjVu XML document. Hyphens, including end-of-line hyphens, have not been processed. They are as in the original DjVu XML document. The text has been segmented based purely on layout based on page breaks. No language level segmetation, such as sentences, tone-units or graphemic, has been applied. Additional mark up using taXMLit has been applied to the TEI XML based on analysis of the original source through the uBio and OpenCalais web services. (Add comment for fuzzy matching once this has been brought into the final workflow too.) Bulletin of the British Museum (Natural BRITISH Ml(NA1 2v ~ Entomology series Vol 50 1985 British Museum (Natural History)London 1985 Dates of publication of the parts Nol 28 February 1985 No 2 28 March 1985 No 3 . 30 May 1985 ISSN 0524-6431 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset BRITISH(NATUI . pr r GEr ContentsEntomology Volume 50 No 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini(Homoptera)By Peter S. Broomfield 1 No 2 Nymphal taxonomy and systematics of the Psylloidea (Homoptera) By I. M. White & I. D. Hodkinson 153 No 3 The whitefly of New Guinea (Homoptera: Aleyrodidae) ByJ. H.Martin . 303 Bulletin of the British Museum (Natural History) N MWMW9MWW %/ X - Taxonomy of Neotropical Derbidae inthe new tribe Mysidiini (Homoptera) Peter S. Broomfield Entomology series Vol 50 No 1 28 February 1985 The Bulletin of the British Museum (Natural History) , instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History),Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1985 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06008 2 ISSN 0524-6431 Entomology series Vol50Nolppl-152British Museum (Natural History)Cromwell RoadLondon SW7 5BD Issued 28 February 1985 Taxonomy of Neotropical Derbidae in the new tribeMysidiini (Homoptera) Peter S. Broomfield Department of Entomology, British Museum (Natural History), Cromwell Road, LondonSW7 5BD Contents Synopsis Introduction Techniques Acknowledgements Derbinae 3 Key to tribes of Derbinae 4 Mysidiini trib. n Key to genera of Mysidiini Checklist of Mysidiini Mysidia Westwood Pseudomysidia Metcalf 77 Dysimia Muir 87 Dysimiella gen. n 96 Mysidaloides gen. n 98 Neomysidia gen. n 99 Ipsemysidia gen. n 100 Amysidiella gen. n 101 Paramysidia gen. n 103 Symidia Muir 108 References Ill Index 151 Synopsis The subfamily Derbinae is divided into two tribes, the Derbini and Mysidiini, the latter newly described.Six genera and 136 species are described as new, one subspecies is raised to specific status, four specificsynonymies and four combinations are newly established, and one neotype and 17 lectotypes aredesignated. Keys to the tribes, 10 genera and 182 species are provided. Introduction The Derbidae is one of the largest and least-known families of the Fulgoroidea, with probablyless than one-fifth of the species currently recognised. It is world- wide in distribution, and themajority of the genera and species are confined to the tropics. The biology of the Derbidae is little known. The adults are phloem feeders, occurring on awide variety of trees and shrubs, in grass land, and occasionally on cultivated cereals. In theU.S.A., Dozier (1928) recorded them feeding on numerous species of deciduous trees,frequently in moist situations. In the New World tropics they often appear to be randomlyscattered throughout primary and secondary forest, although individual species may occur invery large numbers in plantations. There is little information on their host specificity, and theyare of no known major economic importance. The nymphal stages, which are almost completelyunknown, are frequently associated with decaying vegetation, and are often numerous amongstthe litter of the forest floor or in plantations and orchards. They have been found by the presentauthor in old beetle galleries in rotten timber, which suggests that they feed on fungal exudates. Bull. Br. Mus. nat. Hist. (Ent.) 50 (1): 1-152 Issued 28 February 1985 2 PETER S. BROOMFIELD The family-group name Derbidae was first proposed by Schaum (1850), with Derbe Fabricius(1803) as the type-genus. The family was divided into the Derbinae and Kermesiinae byKirkaldy (1906), but subsequently the latter subfamily was rendered obsolete when Kermesiawas shown to be more correctly placed in the Meenoplidae. West wood (1840) divided Derbeinto seven subgenera: Derbe, Mysidia, Zeugma, Thracia, Phenice, Patara and Cenchrea, all ofwhich have subsequently been raised to generic status. In 1900 Kirkaldy proposed the nameZoraida as a replacement name for Thracia, the latter being preoccupied. Muir (1913), studyingthe Old World fauna, ignored the subfamilies and divided the family into four 'groups' based ontegminal venation. Later (1917) he rearranged these groupings into the subfamilies Derbinae,Otiocerinae, Cenchreinae and Rhotaninae, including in the Derbinae the genera Zoraida,Zeugma, Mysidia and Sikaiana. In 1918 Muir revised his classification as a result of his study ofNew World material. Still using characters of the wing and tegminal venation, he separatedZoraida and related genera into a distinct subfamily, the Zoraidinae, which he divided into thetribes Zoraidini and Sikaianini, and relegated the Cenchreinae, Otiocerinae, Derbinae andRhotaninae to tribes within the Derbinae. This arrangement was confirmed in 1930 and wassubsequently followed by Metcalf (1938). Of Westwood's original seven subgenera of Derbe,Muir (1930) assigned Derbe, Mysidia and Zeugma to the Derbini, though he had recognised(1913) that, with the inclusion of Zeugma, such a grouping was not a natural one. Metcalf (1938)erected Pseudomysidia and included it in the Derbini. This classification was accepted by Metcalf (19456), and revised by Fennah (1952) who, alsousing the venation of the tegmina and wings, omitted the subfamily groupings and divided thefamily directly into the tribes proposed by Muir (1918). Recognising their true affinities hetransferred Zeugma to the Zoraidini, and Symidia and Dysimia from the Cenchreini to theDerbini. However, by ignoring subfamilies this classification restricts the grouping of taxa toonly two levels below that of family, and does not acknowledge the relationship between theZoraidini and Sikaianini. In the present study of the Derbini (sensu Fennah, 1952), the male genitalia have beenexamined in order to evaluate their importance as taxonomic characters in the group; asecondary objective was to investigate the generic groupings, and to propose a classificationwhereby the affinities of these genera might be more clearly expressed. Techniques The characters of the head may occasionally be obscured by deposits of white wax, which may beremoved with a fine paint brush. The measurements in the species descriptions are taken withthe insect in dorsal aspect; the width includes the eyes, and the length is from the base to the apexof the anterior extension. In most cases the vertex and frons merge imperceptibly into eachother, and the length of the vertex is therefore not included. The length of the frons is assumed tobe from its base to a point level with the dorsal margins of the eyes. The proportions of thethorax, and the presence or absence of carinae on the frontal and dorsal surfaces, are importantat specific level. These are best observed under incident light at an angle of 45 to the surfaceunder examination. The length of the abdomen varies according to the condition of thespecimen. For this reason 'whole body' measurements are unreliable and are not provided. The pigmentation of the tegmen and wing is frequently faint and occasionally obscured bywaxy deposits; it is best observed at low magnification in natural light, against a whitebackground. Frequently, specimens stored in alcohol rapidly lose all pigmentation. For thisreason extensive use has been made of characters of the male genitalia, the diagnostic features ofwhich are heavily sclerotised and do not require staining. The method of preparation of the male genitalia is similar to that employed for those in themajority of auchenorrhynchous Homoptera. The abdomen is best removed entire at its junctionwith the thorax. After softening in heated 10% KOH, the abdomen is macerated in glacial aceticacid, cleared in clove oil, cleaned in alcohol, and examined in glycerine. Usually the charactersof the aedeagus and parameres may then be observed without further dissection. The aedeagusis best examined from several aspects, so that permanent preparations are not recommended; TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 3 the lateral aspect is sufficient for examination of the parameres. The female genitalia are notused in the present study. The terms used in the text are those commonly employed for the Fulgoroidea and are basedlargely on those of Kramer (1950). Acknowledgements I thank the following individuals for their assistance in making available to me the material uponwhich much of this study is based, for the opportunity to examine type-material and, in manycases, for most helpful advice. The institutions are referred to in the text by the abbreviationsgiven in parenthesis: Dr N. M011er Andersen, Zoologisk Museum, Copenhagen (ZM); Dr P.Arnaud and Dr D. Rentz, California Academy of Science, San Francisco (CAS); Dr R. G.Fennah, formerly of the Commonwealth Institute of Entomology, London; Dr K. G. A.Hamilton, Agriculture Canada, Ottawa (AC); Dr J. P. Kramer, United States NationalMuseum, Washington D.C. (USNM); Dr I. Lansbury, University Museum, Oxford (UM); DrP. Lindskog, Naturhistoriska Riksmuseet, Stockholm (NR); Dr J. T. Medler, Bernice P. BishopMuseum, Honolulu (BPBM); Dr A. F. Newton, Museum of Comparative Zoology, Harvard(MCZ); Dr L. B. O'Brien, Florida Agricultural & Mechanical University, Tallahassee(FAMU); Dr N. D. Penny, Institute Nacional de Pesquisas da Amazonia, Manaus (INPA); DrR. T. Schuh, American Museum of Natural History, New York (AMNH); Dr C. A. Triplehorn,Ohio State University, Columbus (OSU); Dr G. Wallace and Dr G. Ekis, Carnegie Museum,Pittsburgh (CM). DERBINAE Derbini sensu Fennah, 1952: 109. Species of the New World subfamily Derbinae range from the southern Nearctic region to thesouthern temperature zone of the Neotropical region, with the greatest number recorded fromCentral America and tropical South America. They appear to be restricted to moist, frequentlyforested, habitats and are unknown from either mountainous areas, e.g. the High Andes, orfrom the desert area of Mexico and the arid Pacific coastlands of South America. Throughout the subfamily there appears to be a trend towards smaller size with a correspond-ing reduction in the venation of the tegmina and wings, especially in the medial and cubitalareas. This reduction is paralleled in the Zoraidinae, reaching its fullest extent in the Sikaianini.In the Derbinae this tendency is evident by comparison of the tegminal venation of Derbe, inwhich the medial vein has an average of 12 branches, with that of Symidia, in which the medialvein has only six branches and the species are among the smallest and most specialised in thesubfamily. A second , related , tendency paralleled in the rest of the Derbidae is the lateral compression ofthe vertex and frons between the eyes. The extremes of this trend within the Derbinae are shownby the relatively broad and parallel-sided configuration of the vertex and frons in Derbe, and thestrongly compressed condition of these parts in the more specialised genera Symidia, Mysida-loides and Paramysidia. A third, independent, trend is the shortening of the frons and clypeuswith accompanying obsolescence of the longitudinal carinae of the latter, as in Dysimia,Mysidaloides and Dysimiella in contrast to Derbe. While there is little variation in the proportions of the second antennal segment, except for thegenus Mysidaloides, within the Derbinae, the antennal flagellum tends to migrate from theprimitive apical position, as in Derbe, to a subapical position as in Mysidia and Neomysidia. The male genitalia also show several trends in the Derbinae. The shaft of the aedeagus ishorizontal, basically cylindrical, usually symmetrical and usually with anteriorly directedspine-like and/or flap-like processes subapically on the dorsal surface. The ventral surface israrely armed. The paired parameres are large, usually slightly curved, with their apices directedmedially and their ventral margins closely opposed when at rest. In the frequent absence of anextended subgenital plate, the parameres are presumed to shield the aedeagus ventrally and 4 PETER S. BROOMFIELD posteriorly, while dorsal protection is commonly provided by the posterior extension of the hindmargin of the anal tube. The trends within the male genitalia are complex, though often correlated with reduction ofthe tegmina and constriction of the head. From a comparison with other families of auchenor-rhynchous Homoptera, e.g., Membracidae and Cicadellidae, it is assumed that the heavilyarmed aedeagus bearing simple, paired, spine-like processes on the dorsal surface only, with theventral surface unarmed, and with the grasping function of the parameres little developed, is themore primitive condition. This condition is seen in Derbe, in which there is also a small degree ofasymmetry, and other genera such as Pseudomysidia (Figs 34-80). The tendency for thereduction of the dorsal spine-like processes of the aedeagus, or their replacement completely orpartially by flap-like processes, the occasional development of one or more ventral processes,and an accompanying development of the dorsal process of the paramere, are seen in Mysidiaand reach the extreme degree in Dysimia (Figs 94-129). A secondary modification occurs inParamysidia, Ipsemysidia and Amysidiella in which the dorsal process of the paramere isfrequently obsolete or absent and is often replaced by a small, hook-like, subbasal secondaryprocess accompanied occasionally (Paramysidia) by pronounced asymmetry of the aedeagus. Aseparate tendency is seen in Derbe in which the aedeagus is heavily armed and the often verycomplex parameres show a variety of forms consistent with a grasping function duringcopulation. The relatively primitive characters of Derbe, as seen in the tegminal and wing venation, theproportions of the head, and the distinctly different developmental trends in the male genitalia,set it sufficiently apart from other genera of Derbini (sensu Fennah, 1952) to divide the tribe intotwo groups. This necessitates the elevation of all the tribes recognised by Fennah, exceptSikaianini, to subfamily rank, thereby permitting the division of Derbinae (Derbini sensuFennah, 1952) into Derbini (type-genus Derbe) and Mysidiini (type-genus Mysidia). This actionalso illustrates the close relationship between the Zoraidini and the Sikaianini which remain astribes within the Zoraidinae, as proposed by Muir (1918). The proposed classification of theDerbidae is as follows. Derbidae Derbinae Derbini New World Mysidiini New World Otiocerinae New and Old World Cenchreinae New and Old World Rhotaninae Old WorldZoraidinae Zoraidini Old World Sikaianini New and Old World Key to tribes of Derbinae 1 Head with junction of frons and vertex marked by a distinct transverse carina. Tegmen with 20-23 branches of veins extending to posterior and apical margins; medial vein with basal forknot less than 6-branched; cubital vein with 4-6 branches. All branches of medial and cubitalveins linked by a continuous oblique band of cross-veins. Wing with subcostal and radial veinsfused over c. basal one-third length; total number of branches of medial and cubital veinsattaining posterior and apical margins from 6-1 (Fig . 1 ) DERBINI Schaum 2 Head with frons and vertex merging imperceptibly, lacking a transverse carina. Tegmen with never more than 16 branches of veins extending to posterior and apical margins (Pseudomysi-dia), commonly with 11-13; medial vein with basal fork not more than 3-branched; cubitalvein never with more than 4 branches. Cross-veins of medial and cubital areas not as above.Wing with subcostal vein obsolete; total number of branches of medial and cubital veinsattaining posterior and apical margins not exceeding 5 (Figs 2-11) MYSIDIINI trib. n. (p. 5) The Derbini is monotypic, the genus Derbe consisting of 16 species confined to Central Americaand northern South America. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 5 The Mysidiini is erected to accommodate all the genera of Derbinae except Derbe, i.e.,Mysidia, Pseudomysidia, Dysimia and Symidia, and the newly described Amysidiella, Ipsemysi-dia, Mysidaloides, Neomysidia, Paramysidia and Dysimiella. Species of the tribe extend fromthe southern U.S.A. to Uruguay, with the majority occurring in tropical Central and SouthAmerica. Only one species of Mysidia and eight species of Dysimia have been recorded from theCaribbean Islands, excluding Trinidad. One Dysimia and one Paramysidia species are knownfrom the U.S.A. ; the former appears to have reached south-eastern U.S. A. by 'island hopping'across the Caribbean, while the latter probably first became established in Texas after dispersingthrough Central America. MYSIDIINI trib. n. Type-genus: Mysidia Westwood, 1840: 83. Vertex extending not more than one-half of its length beyond eyes, junction with frons not marked by atransverse carina. Frons longer than wide; lateral margins apically subparallel, diverging subbasally.Pronotum longitudinally constricted at mid-dorsal line, not less than 6 times as wide as long. Tegmen2-0-3-0 times as long as wide; radial and subcostal veins fused subbasally, combined radial, medial andcubital veins with not more than 16 branches. Wing with subcostal vein obsolete; combined radial, medialand cubital veins with not more than six branches. The head in dorsal aspect is distinctly broader than long, its width across the eyes being betweenone and three-fourths greater than its length from the base to the apparent apex of the vertex.The vertex usually extends beyond the eyes for up to one-half its length except in Mysidaloides,in which it terminates approximately level with the anterior margins of the eyes, due to theexceptionally large size of the eyes. The junction of the vertex and frons is smoothly, regularly,and obtusely rounded, except in one species of Dysimia where it is subacute. The transversecarina present in Derbe is absent in the Mysidiini. The basal margin of the vertex is usuallytransverse, but in Dysimia, Symidia, and Dysimiella, and to a lesser extent in Ipsemysidia andParamysidia, it is broadly and deeply incised medially. The frons is laterally constricted and is usually from three to seven times as long as its apicalwidth. In Symidia and Pseudomysidia its length may be up to 20 times its apical width. The ratiobetween the length and width of the frons is important at specific and sometimes also genericlevel. In all genera, with the exception of Symidia and Pseudomysidia, the lateral margins of thefrons are strongly divergent from the level of the ventral margins of the eyes to the junction withthe clypeus; the degree and regularity of this divergence are also important at generic level. Thewidth of the frons at its junction with the clypeus is much greater than its apical width, varyingbetween one-fourth (Pseudomysidia} of to approximately equal to its length (Dysimiella andNeomysidia), with the other genera occurring between these extremes. The genae are broadly rounded in lateral aspect, with the exception of a single Dysimiaspecies (see above), and extend anteriorly beyond the eyes from one-third to one-half thehorizontal diameter of the eye. In Mysidaloides, due to the very large size of the eyes, thisanterior extension is minimal. The eyes are large and reniform, with the ventral surfacesadjacent to the antennae weakly concave, except in Mysidaloides where they are hemispherical. The second antennal segment is club-shaped, approximately one and one-half times as long asits maximum breadth, except in Mysidaloides in which the length is five times the maximumbreadth. In the primitive condition, seen in Symidia, Dysimia, Pseudomysidia, and Dysimiella,the antennal flagellum arises apically from a truncate second segment. In Mysida, Mysidaloides,Amysidiella, Paramysidia, Ipsemysidia, and Neomysidia the flagellum has assumed a subapicalposition, and the apex of the segment is narrowly rounded. The ocelli are commonly distinct and only rarely obscure or obsolete. They are small inDysimia, Symidia, Mysidaloides, Ipsemysidia and Amysidiella, variable in size in Mysidia,Pseudomysidia and Dysimiella, are very large and prominent in Paramysidia and Neomysidia;they are of diagnostic value at species level in Mysidia. In the majority of genera the clypeus is approximately as long as the frons (the term 'clypeus'being used throughout the text to denote the combined para- and ante-clypeus); in Dysimiella, O PETER S. BROOMFIELD however, it is somewhat shorter than the frons, and in Pseudomysidia it is up to one-third longer.Slight variation in the length of the clypeus is of specific diagnostic value in many instances. Inthe primitive condition the clypeus bears distinct and percurrent medial and lateral longitudinalcarinae, as seen in Symidia and Pseudomysidia, and occasionally in Mysidia; these are obsoleteor absent in the remaining genera. The rostrum in the primitive condition extends to the apex of the abdomen, as seen inPseudomysidia and Symidia. In the majority of genera it terminates approximately level with thehind coxae, except in Mysidia where its length is extremely variable and is of value in specificdiagnosis. The pronotum is longitudinally constricted at the mid-dorsal line, with the posterior marginvery broadly and deeply incised. In Mysidia, Paramysidia, Mysidaloides, Dysimiella, Neomysi-dia, Ipsemysidia and Amysidiella, its maximum width is usually 10-20 times its medial length,though in some species of Mysidia it may be so strongly constricted that the width is up to 50times the medial length. In Pseudomysidia, Symidia, and Dysimia this constriction is relativelyslight, and the width of the pronotum may be as little as 6-8 times its length. The ratio of thelength to the breadth of the pronotum varies greatly between species, especially in Mysidia, andis often of considerable diagnostic value. The fronto-lateral surfaces of the pronotum may be distinctly carinate in all genera, thoughthey are not consistently so in Mysidia and Pseudomysidia; in which genera their presence orabsence is of great diagnostic value at species level. In Symidia these carinae are greatly elevatedand foliaceous, and continue along the lateral and ventral margins to form pronounced antennalfoveae; which, although occurring intermittently in other subfamilies of the Derbidiae, areotherwise absent in the Derbinae. The tegulae also vary in the possession of carinae, these beingsometimes present in Mysidia, Pseudomysidia, and Dysimia, although absent in the othergenera. The presence or absence of these carinae may be of value at specific level. Both the discof the mesonotum and the scutellum are approximately as wide as long, their proportionsvarying little between genera; the former occasionally bears three distinct longitudinal carinae,but these are more usually obsolete or absent. The tegmen is from two and one-half to three times as long as its maximum breadth (slightlybroader in Dysimiella) with the radial and sub-costal veins fused over their basal one-third toone-half length. The radial and medial veins are distinct from near their base in Mysidia,Paramysidia, Ipsemysidia, Pseudomysidia, and Mysidaloides, but in Symidia, Dysimiella,Dysimia, Neomysidia, and Amysidiella they may be fused for up to one-quarter of their length.The radial vein is usually two branched, although three branches are present in Symidia, whichalso has the usual seven branches of the medial vein reduced to six. The medial vein ofPseudomysidia is eleven-branched, a condition unique in the tribe and considered to represent aprimitive condition. The number of branches of the cubital vein also varies. The genera Mysidia,Paramysidia, Mysidaloides, Dysimiella, Neomysidia, Ipsemysidia, and Amysidiella have fourbranches; Pseudomysidia and Dysimia have three; and Symidia, perhaps due to the small size ofthe insects, only two. The total number of branches of the radial, medial and cubital veins istherefore usually thirteen, the exceptions being Pseudomysidia with sixteen, Dysimia withtwelve, and Symidia with eleven. In the Derbini the total varies from twenty to twenty-three. The wing is usually approximately one-half the length of the tegmen, but is somewhat longerin Dysimiella and Paramysidia, and occasionally longer in Mysidia. The subcostal vein is fusedthroughout its length with the radial vein, in contrast with the Derbini where it is distinct for thegreater part of its length. The radial vein is also unbranched. The number of branches of themedial vein is variable; being two in the majority of genera, three in Pseudomysidia; in Dysimia,Symidia, and Dysimiella it is unbranched. The cubital vein has either two branches, as inPseudomysidia, Neomysidia and Symidia, or three, as in Amysidiella, Ipsemysidia, Mysidia,Mysidaloides, Paramysidia, Dysimia, and Dysimiella. The total number of branches of themedial and cubital veins therefore varies from three to five, contrasting with the total of six to tenin the Derbini. The head and body are usually pale yellowish brown, rarely dark brown, with brown, black orred markings frequently present, particularly on the head and the frontolateral surfaces of the TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 7 pronotum. The tegmina and wings are usually predominently whitish hyaline but may also beclear, yellowish or smoky brown throughout in some species of Mysidia. They are frequentlymarked with darker spots or transverse bands or with the veins and cross veins dark margined.All markings are consistent within species and are of diagnostic value at this level. The male genitalia are extremely variable in the shape of the aedeagus and parameres, with adistinct correlation in many cases between a reduction in the processes of the former and anincrease in those of the latter. This correlation is well illustrated within Dysimia, where in thespecies numa Fennah the dorsal process of the paramere is but little developed while theaedeagus is the most highly armed in the genus. Both the aedeagus and the parameres are of very great diagnostic importance at specific level,especially in the case of externally similar forms, and occasionally at generic level, i.e.Paramysidia. The primitive condition appears to be that in which the shaft of the aedeagus isarmed subapically with simple, paired, symmetrically arranged, spine-like processes on thedorsal surface only, as in Pseudomysidia, Amysidiella, and a few species of Mysidia. Thedevelopment of additional flap-like processes occurs in Paramysidia, Dysimia, Symidia, Mysi-daloides, Dysimiella, Ipsemysidia, and most species of Mysidia. The possession of flap-likeprocesses only, as seen in Neomysidia and some species of Mysidia, is regarded as being a furtherdevelopment which reaches its most advanced condition in certain species of Dysimia, where thegrasping function of the aedeagal processes can only be minimal and the parameres appear tohave assumed this task during copulation. The ventral surface of the aedeagus is usuallyunarmed, the exceptions being Ipsemysidia and a few species of Mysidia. The development ofasymmetry in the aedeagal processes, present in Symidia and occasionally in Mysidia, reaches itsgreatest extent in Paramysidia, where an unpaired medial dorsal process is also present. The dorsal process of the paramere is well developed in Mysidaloides and Dysimiella, and inthe majority of species of Mysidia and Dysimia. It is less developed in Pseudomysidia andSymidia, and greatly reduced in Amysidiella, Ipsemysidia, Neomysidia, and Paramysidia, and insome species of Mysidia and Dysimia. The reduction, or loss, of the dorsal process is oftencompensated for by the development of a small hook-like secondary process subbasally . Only inMysidia is there the occasional development of an additional process on the ventral surface. The male subgenital plate is usually short with its posterior margin transverse; only inDysimiella is it strongly produced medially. The female genitalia appear not to be of taxonomic value at either the generic or specific level,with the possible exception of the subgenital plate. The variation in this character is however tooslight to be of use diagnostically . Key to genera of Mysidiini 1 Medial vein of tegmen with 11 branches extending to posterior and apical margins (Fig. 11) PSEUDOMYSIDIA Metcalf (p. 77) Medial vein of tegmen with not more than seven branches 2 2(1) Cubital vein of tegmen two-branched; medial vein six-branched (Fig. 5) SYMIDIA Muir (p. 108) Cubital vein of tegmen three- or four-branched ; medial vein seven-branched 3 3(2) Cubital vein of tegmen three-branched (Fig. 9) DYSIMIA Muir (p. 87) Cubital vein of tegmen four-branched 4 4(3) Second antennal segment not longer than 2-5 times maximum width 5 Second antennal segment 5-0 times as long as maximum width (Fig. 16) MYSIDALOIDES gen. n. (p. 98)5(4) Wing with medial vein unbranched (Fig. 3). Male with posterior margin of subgenital plate produced (Fig. 160) DYSIMIELLA gen. n. (p. 96) Wing with medial vein two-branched . Male subgenital plate transverse 6 6(5) Head with length of frons little greater than width at base, c. 2-5 times width at apex (Fig. 31) NEOMYSIDIA gen. n. (p. 99) Proportions of frons not as above 7 7(6) Tegmen with subcostal and radial veins fused over c. basal third of length 8 Tegmen with subcostal and radial veins fused to c. mid-length 9 8 PETER S. BROOMFIELD 8(7) Length of frons not less than twice width at base (Fig. 33). Tegmen with radial and medial veins distinct subbasally (Fig. 7) MYSIDIA Westwood (p. 9) Length of frons less than twice width at base (Fig. 25). Tegmen with radial and medial veins fused over basal sixth of length (Fig. 4) AMYSIDIELLA gen. n. (p. 101) 9(7) Pronotal width less than 10 times length at mid-dorsal line (Fig. 21). Length of frons 3 times width at apex (Fig. 26) IPSEMYSIDIA gen. n.(p. 100) Pronotal width not less than 10 times length at mid-dorsal line (Fig. 20). Length of frons atleast 4 times width at apex (Fig. 27) PARAMYSIDIA gen. n.(p. 103) Checklist of Mysidiini MYSIDHNI trib. n. MYSIDIA Westwoodacidalioides Fowleradamare sp. n.adusta sp. n.agilis sp. n.albicans (Stal)albifasciata sp. n.albipennis Westwood parviceps Fowler syn. n.amaranths sp. n.amazona sp. n.andessp. n.ariasisp. n.asinella sp. n.athena sp. n.augusta sp. n.bt'lln sp. n.bianca sp. n.bibula sp. n.bizzara sp. n.bolivianna sp. n.calliginosa Walker rubra Metcalf syn. n.calypso sp. n.carosella sp. n.cheesemanisp. n.cinerea Fennahclaudata sp. n.clava sp. n.cooper! sp. n.costata (Fabricius)decora sp. n.delicatissima Fowlerdiabola sp. n.diana sp. n.distant! sp. n.distinct a sp. n.dollingisp. n.douglasisp. n.ecuadoria sp. n.enjebetta sp. n.erects sp. n.estfarchina sp. n.etheldreda sp. n.fasciata MetcalfffavUlasp. n.formosa sp. n. fowlerisp. n.fulvodorsalissp. n.fuscofrontalissp. n.fuscomaculatasp. n.geoffreyisp. n.glauca Distantgracilis sp. n.grandis sp. n.harmonia sp. n.havilandisp. n.hengistsp. n.henriettasp. n.hyalina sp. n.immaculata sp. n.infedelis sp. n.inquinata sp. n.insania sp. n.insolita sp. n.intinw sp. n.isteria sp. n.jamesisp. n.josianna sp. n.knightisp. n.kramerisp. n.lacteola sp. n.lactifiora Westwoodlimpida sp. n.liquida sp. n.Uoydisp. n.lucianna sp. n.lucifera sp. n.maculicosta Fowlermaculosa sp. n.magics sp. n.marshallisp. n.minerva sp. n.molests sp. n.musics sp. n.mylesisp. n.nebulosa (Germar)nemorensissp. n.neoasinella sp. n.neonebulosa Muirnigrifrontalissp. n.nigrithoraxsp. n.nitidu sp. n.obscura MetcalfpaUescens Metcalf TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) maculipennissp. n.morris/ sp. n.iiiuir/sp. n.numa Fennahobrienisp. n.pseudomaculata sp. n.putilla Fennahtelfordisp. n. PSEUDOMYSIDIA Metcalfaraguana Fennah slat. n.debora sp. n.delicata sp. n.ecuadoriensis sp. n.fuscovaria Metcalfhind ore sp. n.Juliana sp. n.lepida sp. n.marshallisp. n.obnubilia sp. n.pallida sp. n.palmerisp. n.panamensissp. n.rubidella (Ball) comb. n.similissp. n.trinidadensis sp. n.vest/s sp. n. AMYSIDIELLA gen. n.micare sp. n.pseudomicaresp. n. DYSIMIELLA gen. n.penny! sp. n.williamsisp. n. IPSEMYSIDIA gen. n.beautifica sp. n. MYSIDALOIDESgen. n.trinidadensis sp. n. NEOMYSIDIA gen. n.willisisp. n. PAUAM YSIDIA gen. n.barbara sp. n.boudica sp. n.felixsp. n. mississippiensis (Dozier) comb. n.nigropunctata (Metcalf) comb. n.tessellata sp. n.vulgaris sp. n. pallida (Fabricius)panamensis sp. n.peregrina sp. n.persephone sp. n.perspicua sp. n.polyhymnia sp. n.pseudocostata sp. n.pseudoerecta sp. n.pseudonebulosa Muirpulchella sp. n.punctifera Metcalfpunctum (Fabricius) steinbachi Distant syn. n.quadrifasciata Walkerrichardsisp. n.robust u sp. n.sanguinea sp. n.silvana sp. n.simp In sp. n.squamigera (Fabricius)stall sp. n.stigma (Germar)striata sp. n.subfasciata Westwoodsubfusca Metcalftestacea (Fabricius) citrina Walker syn. n.tikalme sp. n.transversa sp. n.unimaculata sp. n.var/a sp. n.venusta sp. n.v/vfa sp. n.whimper! sp. n.williamsi sp. n. SYMWIA Muirbucaya sp. n./Java Muirpintosamia sp. n.pseudoffava sp. n.withycombei sp . n . DFSIM/A Muirastarte sp. n.distincta sp. n.fennahisp. n.fuscoclypeata Fennahjamaicensis (Distant) comb. n.maculata Muir MYSIDIA WestwoodMysidia Westwood, 1840: 83. Type-species: Derbe pallida Fabricius, by original designation. Width of head in dorsal aspect usually from one-quarter to one-half greater than length, rarely more orless. Vertex extending anterior to eyes for between one-third and one-half its length; lateral margins not 10 PETER S. BROOMFIELD highly elevated, strongly converging from base to level of midline of eyes, thence subparallel to apex; baseshallowly concave. Frons with lateral margins gradually diverging from apex to base; slender, lengthusually 4-7 times width at apex, 2-0-3-5 times width at base, seldom more or less; junction with vertexbroadly rounded, indistinct, lacking a transverse carina; lateral carinae very prominent, often semifoli-naceus. Genae extending anterior to eyes for one-third to one-half horizontal diameter of eye. Eye weaklyreniform, ventral margin adjacent to antennal base weakly concave. Antenna with second segmentclub-shaped, apex narrowly rounded; length usually 1-5-2-5 times maximum width; flagellum arisingsubapically. Ocelli commonly distinct, often small, rarely obscure or obsolete, occasionally very large andprominent. Clypeus broad, not greatly swollen; commonly as long as, or rather longer than, frons; medialcarina frequently obsolete or extending only over c. apical one-half length or less, seldom distinct orpercurrent; lateral carinae commonly not extending over more than basal one-third length, rarely eitherobsolete or distinct throughout. Rostrum often extending to, or beyond, base of subgenital plate; butfrequently terminating at level of hind coxae. Dorsal surface of pronotum very deeply and broadly constricted at midline; width usually 10-20 timesmid-dorsal length, occasionally much greater. Pronto-lateral surfaces often each with a single prominentcarina curving horizontally from adjacent to midline of eye to lateral margin. Tegulae occasionally eachwith a single horizontal carina. Disc of mesonotum often slightly wider than length at midline; medial andtwo lateral longitudinal carinae rarely distinct and percurrent. Tegmen usually 6-12 mm long, rarely distinctly shorter or longer; that of the female usually being up to30 per cent longer than that of the male; length c. 3 times maximum breadth. Medial vein distinct from nearbase; subcostal and radial veins fused over c. one-third length from base. Radial vein with two branchesextending to apical margin; linked to medial vein by a cross-vein at c. two-thirds length, and by anotheradjacent to apical fork. Medial vein forking at c. two-fifths length, and again at midlength; with sevenbranches extending to apical and posterior margins, second and third, and fourth fifth, linked bycross-veins. Cubital vein with four branches extending to posterior margin, first linked to apex of clavusand to second, third to fourth, and fourth to first branch of medial vein by cross-veins (Fig. 7). Wing with length c. one-half to two-thirds that of tegmen. Radial and sub-costal veins fused over ratherless than basal half of length, unbranched; the former linked to medial vein by a cross-vein at c. two-thirdslength. Medial vein with two branches extending to apical margin, linked to cubital vein by a singlecross-vein at c. midlength. Cubital vein with three branches extending to posterior margin. Head and body usually pale yellowish brown, seldom dark, often with distinct markings. Tegmen andwing often hyaline or whitish hyaline, occasionally deep fuscous, frequently with veins and cross-veinsbroadly margined smoky brown, often with distinct transverse bands or with apical and posterior marginssmoky brown. Male genitalia with shaft of aedeagus horizontal, basically cylindrical, usually symmetrical, usuallyslender in lateral aspect, often broadly expanded subapically in vertical aspect. Dorsal surface subapicallyusually with one or two, rarely three or four, pairs of spine or flap-like processes occasionally extendingover lateral surfaces. Lateral and ventral surfaces usually unarmed. Paramere often slender basally,frequently obtusely rounded apically; dorsal surface usually with a well-developed posteriorly producedprocess bearing opposed projections on its posterior surface; often with a distinct secondary process. Analtube often very strongly produced and decurved posteriorly, apex often deeply notched at midline.Subgenital segment with lateral and ventral margins occasionally bearing distinct, posteriorly directedprocesses. Female with posterior margin of subgenital plate commonly transverse or broadly rounded, rarelystrongly produced or shallowly notched medially. Mysidia was erected by Westwood (1840) as a subgenus of Derbe to accommodate the Fabrician speciespallida, squamigera, costata, punctum, testacea and nivea, and his own new species albipennis, lactifloraand subfasciata; further species were added by various authors, mainly Metcalf, Fowler, Distant, Walker,Muir and, more recently, Fennah, bringing the total number to 34. As a result of the present study the distribution includes Brazil (69 species), Trinidad (7), Surinam (5),Guyana (15), French Guiana (2), Venezuela (4), Colombia (10), Ecuador (11), Bolivia (10), Peru (11),Uruguay (1?), Panama (20), Costa Rica (2), Honduras (4), Belize (2), Guatemala (4), Mexico (2) andJamaica (1 species). The localities from which species are recorded more probably reflects, at least in northern SouthAmerica, intensity of collecting rather than diversity of species; with the exception of Trinidad, Jamaica isthe only Caribbean island from which the genus is recorded. Due to the previous confusion and frequentmisidentification, most of the published locality data recorded by Metcalf (1945-6) must be regarded assuspect. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 11 Key to species of Mysidia (based on external characters) It has not been possible to examine the type-material of stigma, while the unique holotypes of cinerea,pallida and pseudonebulosa are badly damaged; these species are therefore omitted from this key. Though external characters are consistent within species, between species they are occasionally slight; inthese instances reference should be made to the structure of the male genitalia. 1 Tegmen entirely dark brown , veins and cross-veins concolorous 2 Tegmen pale or , if predominantly dark brown , with pale markings 6 2 (1) Wing entirely dark brown 3 Wing with a narrow, oblique, pale transverse band. Brazil asinella sp. n. (p. 22) 3 (2) Female legmen more than 11 mm; tegula uniformly pale; fronto-lateral surfaces of pronotum each with a distinct, horizontal, scarlet band. Brazil adusta sp. n. (p. 22) Female tegmen less than 10 mm, or with tegula not uniformly pale 4 4 (3) Tegula uniformly dark brown. Brazil polyhymnia sp. n. (p. 23) Tegula not uniformly dark brown 5 5 (4) Fronto-lateral surfaces of pronotum uniformly pale; tegula with dorsal margins dark brown. Brazil, Guyana, Bolivia, Panama, Peru, Trinidad calliginosa Walker (p. 23) Fronto-lateral surfaces of pronotum reddish dorsally; tegula reddish, dorsal marginsdarker red. Brazil inquinata sp. n. (p. 24) 6 (1) Tegmen and wing pale, veins and cross-veins uniformly pale 7 Tegmen and wing either predominantly dark brownish; or, if pale, with cross-veinsdarker than veins; frequently with dark transverse bands 19 7 (6) Tegmen and wing totally devoid of dark markings; costal cell of tegmen sometimes tinged yellowish brown 8 Tegmen and wing with one or more dark spots 14 8 (7) Tegmen more than 10 mm 11 Tegmen less than 10 mm 9 9 (8) Tegmen less than 8 mm; fronto-lateral surfaces of pronotum prominently carinate. Brazil venusta sp. n. (p. 24) Tegmen more than 8 mm ; fronto-lateral surfaces of pronotum not carinate 10 10 (9) Tegmen with costal cell pale yellowish brown; radial and medial areas hyaline through- out. Guyana richardsi sp. n. (p. 25) Tegmen with costal cell hyaline; radial and medial areas smoky brown apically. Brazil timpida sp. n. (p. 25) 11 (8) Tegmen yellowish hyaline. Brazil robusta sp. n. (p. 26) Tegmen whitish hyaline 12 12 (11) Head and body unicolorous brownish yellow. Tegmen less than 12 mm 13 Head and body with distinct reddish markings. Tegmen more than 13 mm. Peru immaculata sp. n. (p. 27) 13(12) Tegmen with posterior margin weakly tinged smoky brown. Guyana nitida sp. n. (p. 26) Tegmen entirely hyaline. Brazil amazona sp. n. (p. 26) 14 (7) Tegmen with a very prominent dark brown spot extending from costal margin to second branch of cubital vein at one-third length. Peru, Bolivia, Guyana, Brazil punctum (Fabricius) (p. 27)Tegmen not as above 15 15 (14) Tegmen with apical fork of medial vein very narrowly dark brown, not otherwise pigmented. Jamaica hyalina sp. n. (p. 28) Tegmen not as above 16 16 (15) Tegmen less than 8-0 mm 17 Tegmen more than 9-5 mm 18 17 (16) Tegmen with a large, prominent, brown spot between apex of clavus and first branch of cubital vein, otherwise unmarked. Brazil unimaculata sp. n. (p. 28) Tegmen with numerous small dark spots. Brazil ttavilla sp. n. (p. 29) 18 (16) Male tegmen less than 10 mm; tegula distinctly carinate; width of pronotum less than 2 mm . Ecuador athena sp. n. (p. 28) Male tegmen more than 11 mm; tegula with carinae obsolete; width of pronotum morethan 2 mm. Ecuador, Panama, Costa Rica acidaloides Fowler (p. 29) 12 PETER S. BROOMFIELD 19 (6) Tegmen brown with pale transverse bands 20 Tegmen predominantly pale hyaline or, if dark brownish, lacking distinct pale bands 28 20(19) Tegmen with a single, very narrow, pale transverse band. Brazil.... neoasinella sp. n. (p. 30)Tegmen with more than one pale transverse band 21 21 (20) Tegmen with two narrow, pale, transverse bands, both on basal half. Guyana vista sp. n. (p. 30)Tegmen with four pale transverse bands 22 22 (21) Tegmen with all pale transverse bands extending from costal to posterior margins 23 Tegmen without all pale transverse bands extending across entire width 26 23 (22) Tegmen with width of alternating light and dark bands approximately equal 24 Tegmen with pale bands narrower than the dark bands. Ecuador . . . albifasciata sp. n. (p. 31) 24 (23) Tegmen pale brown; head in dorsal aspect little wider than long. Brazil quadrifascia Walker (p. 31)Tegmen dark brown ; head in dorsal aspect with width considerably greater than length 25 25 (24) Male tegmen little more than 7 mm, 3 times maximum width; dorsal surface of abdomen basally dark brown. Brazil, Peru trans versa sp. n. (p. 32) Male tegmen approximately 8 mm, 2-5 times maximum width; dorsal surface ofabdomen pale. Bolivia fulvodorsalis sp. n. (p. 32) 26 (22) Tegmen more than 8 mm ; third pale transverse band faint , broken medially . Brazil musica sp. n. (p. 32) Tegmen less than 8 mm; third pale transverse band extending unbroken from costal toclaval margins 27 27 (26) Tegmen with pale transverse bands narrower than intervening dark areas; disc of mesonotum deep brown. Brazil williamsi sp. n. (p. 33) Tegmen with pale transverse bands as broad as intervening dark bands; disc ofmesonotum pale. Trinidad mylesi sp. n. (p. 33) 28 (19) Tegmen and wing predominantly brownish , veins pale-margined 29 Tegmen and wing predominantly pale or, if largely brownish, with veins broadly edgedbrownish 33 29 (28) Tegmen more than 9 mm 30 Tegmen less than 9 mm 31 30 (29) Tegmen with light and dark markings giving a strongly mottled appearance. Colombia, Ecuador, Brazil varia sp. n. (p. 34) Tegmen not distinctly mottled. Guyana, Brazil tikalme sp. n. (p. 34) 31 (29) Tegmen and wing pale only at margins of veins; fronto-lateral surfaces of pronotum distinctly carinate. Brazil glauca Distant (p. 35) Tegmen and wing with larger cells pale medially; fronto-lateral surfaces of pronotum notdistinctly carinate 32 32 (31) Rostrum hardly extending beyond hind coxae; tegula carinate. Brazil . . gracilis sp. n. (p. 35) Rostrum extending to midlength of abdomen; tegula not carinate. Brazil lucifera sp. n. (p. 35) 33 (28) Tegmen and wing predominantly brownish. Guyana, Brazil havilandi sp. n. (p. 36) Tegmen and wing predominantly pale 34 34 (33) Tegmen with veins and/or cross-veins at least in part broadly margined smoky brown 47 Tegmen with veins and/or cross-veins not broadly margined smoky brown 35 35 (34) Tegmen and/or wing with distinct darker transverse bands 76 Tegmen and wing lacking distinct dark transverse bands 36 36 (35) Scutellum blackish brown. Brazil etheldreda sp. n. (p. 36) Scutellum pale 37 37 (36) Fronto-lateral surfaces of pronotum each with a large , circular , dark brown spot 38 Fronto-lateral surfaces of pronotum either unmarked, or each with a dark bandextending horizontally from adjacent to eye to lateral margin 40 38 (37) Disc of mesonotum with a pair of large, dark brown spots posteriorly. Bolivia, Vene- zuela, Peru liquids sp. n. (p. 37) Disc of mesonotum lacking prominent markings 39 39 (38) Fronto-lateral surfaces of pronotum carinate. Tegmen with a small, dark brown spot at apex of anal vein. Brazil ariasi sp. n. (p. 37) Fronto-lateral surfaces of pronotum not carinate. Tegmen not as above. Trinidad,Guyana, Brazil, Surinam, Peru, Ecuador costata (Fabricius) (p. 40) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 13 40 (37) Pronto-lateral surfaces of pronotum each with a prominent, dark brown, horizontal band 41 Pronto-lateral surfaces of pronotum uniformly pale , or with horizontal bands orange 43 41 (40) Tegmen with costal cell pale, bearing a single dark brown spot 42 Tegmen with subcostal cell yellowish brown, becoming darker distally, with threeirregular brownish spots. Panama fuscofrontalis sp. n. (p. 38) 42 (41) Tegmen less than 10 mm. Pronotum with maximum width less than 20 times length at mid-dorsal line. Guatemala, Belize, Honduras, Brazil albipennis Westwood (p. 38) Tegmen more than 10 mm. Pronotum with maximum width greater than 40 times length at mid-dorsal line. Brazil lactittora Westwood (p. 39) 43(40) Wing broadly brownish at level of radial-medial cross-vein. Ecuador diana sp. n. (p. 43) Wing pale medially 44 44 (43) Tegmen and wing with posterior and apical margins dark smoky brown between veins; wing with a small dark spot adjacent to first branch of cubital vein. Colombia cooper! sp. n. (p. 42)Tegmen and wing not as above 45 45 (44) Tegmen with costal cell hyaline 46 Tegmen with costal cell opaque yellowish brown Panama, Costa Rica: dollingisp. n. (p. 43); Peru, Ecuador, Brazil, Guyana: pseudocos-tatasp. n. (p. 41); Bolivia: biancasp. n. (p. 42) 46 (45) Tegmen more than 9.0 mm. Tegmen and wing with faint yellow transverse bands. French Guiana, Trinidad lacteola sp. n. (p. 39) Tegmen less than 8-5 mm. Tegmen and wing unmarked. Mexico delicatissima Fowler (p. 41) 47 (34) Tegmen with apical fork of medial vein dark brown or black 48 Tegmen with apical fork of medial vein pale 66 48 (47) Pronto-lateral surfaces of pronotum with alternating deep brown and white horizontal bands. Brazil striata sp. n. (p. 43) Pronto-lateral surfaces of pronotum not as above 49 49 (48) Pronto-lateral surfaces of pronotum prominently carinate 50 Pronto-lateral surfaces of pronotum with carinae weak or absent 54 50 (49) Pronto-lateral surfaces of pronotum pale dorsally, deep brown ventrally. Brazil, Peru, Surinam sanguinea sp. n. (p. 44) Pronto-lateral surfaces of pronotum not as above 51 51 (50) Pronto-lateral surfaces of pronotum with carinae narrowly orange . Panama fowleri sp. n. (p. 46)Pronto-lateral surfaces of pronotum unicolorous 52 52 (51) Wing with apex broadly smoky brown. Brazil calypso sp. n. (p. 44) Wing with apex pale 52 53 (52) Ocelli prominent. Tegmen with veins dark brown. Brazil peregrina sp. n. (p. 45) Ocelli obsolete. Tegmen with veins yellowish; cross-veins dark. Brazil, Surinam lucianna sp. n. (p. 45) 54 (49) Tegula uniformly pale 56 Tegula dark, at least in part 55 55 (54) Tegula entirely very dark brown. Brazil squamigera (Fabricius) (p. 40) Tegula dark brown dorsad of base of legmen only. Panama grandissp. n. (p. 46) 56 (54) Pronto-lateral surfaces of pronotum each with a very distinct, narrow, horizontal, orange band. Belize minerva sp. n. (p. 47) Pronto-lateral surfaces of pronotum not as above 57 57 (56) Tegmen and wing with veins and/or cross-veins margined smoky brown, lacking other markings 60 Tegmen and wing with distinct brownish markings in addition to those around veins and cross-veins 58 58 (57) Pronto-lateral surfaces of pronotum each with a horizontal orange band adjacent to eye. 59 Pronto-lateral surfaces of pronotum uniformly pale. Panama punctifera Metcalf (p. 47) 59 (58) Dorsal surface of abdomen with a large, dark brown, spot on either side of midline basally. Guatemala maculicosta Fowler (p. 48) Dorsal surf ace of abdomen uniformly pale. Brazil molestasp. n. (p. 48) 60 (57) Tegmen with posterior margin broadly and continuously smoky brown .Panama obscura Metcalf (p. 49) 14 PETER S. BROOMFIELD Tegmen with posterior margin either hyaline, or narrowly and intermittently smokybrown 61 61 (60) Head with genae pale, unicolorous yellowish brown 64 Head with genae not uniformly pale 62 62 (61) Head with genae narrowly dark reddish brown adjacent to eyes. Brazil Henrietta sp. n. (p. 49)Head with genae dull brownish dorsally 63 63 (62) Tegmen with external margins of medial, radial, and subcostal areas dark smoky brown medially. Belize, Honduras distant! sp. n. (p. 49) Tegmen with posterior and apical margins hyaline. Brazil albicans (Stal) (p. 50) 64 (6 1 ) Tegmen with costal cell hyaline with numerous irregular brown spots 65 Tegmen with costal cell yellowish brown, with a single, brownish, spot at level of point ofseparation of fused subcostal and radial veins. Brazil, Surinam ... nemorensis sp. n. (p. 51) 65 (64) Tegmen densely mottled smoky brown over basal third. Ocelli large. Belize, Honduras insolita sp. n. (p. 51) Tegmen with basal third predominantly hyaline. Ocelli obsolete. Mexico enjebetta sp. n. (p. 51)(66) (47) Pronto-lateral surfaces of pronotum broadly dark brown/black medially. Dorsal surface of abdomen with a large deep red spot. Panama nigrifrontalis sp. n. (p. 52) Pigmentation of pronotum and abdomen not as above 67 67 (66) Fronto-lateral surfaces of pronotum each with a distinct orange band extending horizon- tally from adjacent to eye to lateral margin 68 Fronto-lateral surfaces of pronotum yellowish brown , unmarked 70 68 (67) Tegmen more than 10 mm, mostly smoky brown. Bolivia andes sp. n. (p. 52) Tegmen less than 10 mm, predominantly hyaline 69 69 (68) Clypeus with medial carina percurrent; ocelli prominent; tegula not carinate. Panama. hi hula sp. n. (p. 53)Clypeus with medial carina obsolete; ocelli small; tegula carinate. Ecuador ecuadoria sp. n. (p. 53) 70 (67) Fronto-lateral surf aces of pronotum distinctly carinate 71 Fronto-lateral surfaces of pronotum not carinate 72 71 (70) Fronto-lateral surfaces of pronotum unicolorous pale yellowish brown throughout. Trinidad cheesemani sp. n. (p. 54) Fronto-lateral surfaces of pronotum with carinae narrowly edged reddish brown.Peru augusta sp. n. (p. 54) 72 (70) Tegmen with an irregular brownish transverse band at one-third length, another very irregular band over second fork of medial vein. Ocellismall 74 Tegmen not as above . Ocelli prominent 73 73 (72) Pronotum with width 25 times length at mid-dorsal line. Honduras, Mexico, Panama, CostaRica nebulosa (Germar) (p. 56) Pronotum with width approximately 15 times length at mid-dorsal line 75 74 (72) Tegmen more than 8 mm. Head with vertex pale. Bolivia, Colombia .... erecta sp. n. (p. 55) Tegmen less than 8 mm. Basal angles of vertex dark brown krameri sp. n. (p. 54) 75 (73) Frons less than 6 times width at apex ; clypeus longer than f rons . Ecuador maculosa sp. n. (p. 55)Frons more than 6 times width at apex; clypeus shorter than frons. Bolivia bizzara sp. n. (p. 56) 76 (35) Tegmen and wing yellowish ; tegmen lacking dark transverse bands 77 Tegmen and wing hyaline; tegmen with dark transverse bands 79 77 (76) Wing with two brownish transverse bands. Brazil intima sp. n. (p. 56) Wing with a single dark transverse band 78 78 (77) Wing with dark transverse band extending from costal margin to apex of clavus. Brazil infedelis sp. n. (p. 57) Wing with dark transverse band extending from medial-radial cross-vein to apex ofclavus. Guyana testacea (Fabricius) (p. 57) 79 (76) Thorax and abdomen predominantly dark brown. Brazil diabola sp. n. (p. 58) Thorax and abdomen predominantly pale yellowish brown 80 80 (79) Fronto-lateral surfaces of pronotum distinctly carinate 81 Fronto-lateral surf aces of pronotum not carinate 82 TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 15 81 (80) Wing with two distinct smoky brown transverse bands. Brazil lloydi sp. n. (p. 58) Wing lacking dark transverse bands. Panama fuscomaculatasp. n. (p. 59) 82 (80) Tegula each with a distinct horizontal carina 83 Tegula not carinate 87 83 (82) Pronto-lateral surfaces of pronotum orange at level of eyes. Pronotum with width less than 20 times length at mid-dorsal line 84 Pronto-lateral surfaces of pronotum concolorous dull yellowish. Pronotum with widthgreater than 25 times length at mid-dorsal line 85 84 (83) Tegmen with four faint transverse bands. Brazil adamare sp. n. (p. 60) Tegmen with two faint transverse bands. Peru isteria sp. n. (p. 59) 85 (83) Head with genae adjacent to eyes dark brownish. Panama pallescens Metcalf (p. 60) Head with genae pale throughout 86 86 (85) Tegmen with three irregular smoky brown transverse bands, apex brownish. Ecuador insania sp. n. (p. 60)Tegmen with two very faint, transverse bands, apex hyaline. Panama panamensis sp. n. (p. 61) 87 (82) Tegmen with a single distinct dark transverse band 88 Tegmen with at least two dark transverse bands 90 88 (87) Genae ad j acent to eyes , and posterior margin of pronotum , deep red . Venezuela formosa sp. n. (p. 61)Genae and pronotum not as above 89 89 (88) Disc of mesonotum deep orange. Ecuador whimper! sp. n. (p. 62) Disc of mesonotum yellowish. Brazil stall sp. n. (p. 62) 90 (87) Tegmen with two dark transverse bands 91 Tegmen with three or more dark transverse bands 101 91 (90) Tegmen with distal transverse band extending only over cubital area 92 Tegmen with distal transverse band extending to costal margin 94 92 (91) Pronto-lateral surfaces of pronotum each with a narrow, light orange band extending horizontally from adjacent to eye to lateral margin. Brazil clava sp. n. (p. 62) Pronto-lateral surfaces of pronotum unicolorous yellowish brown 93 93 (92) Pronotum with width less than 20 times length at mid-dorsal line ; fronto-lateral surfaces carinate. Ecuador simpla sp. n. (p. 63) Pronotum with width greater than 30 times length at mid-dorsal line; fronto-lateralsurfaces not carinate. Trinidad, Brazil josiannasp. n. (p. 67) 94 (91) Disc of mesonotum blackish brown. Peru nigrithorax sp. n. (p. 63) Disc of mesonotum pale 95 95 (94) Wing with two smoky brown transverse bands. Panama subfusca Metcalf (p. 64) Wing either with a single transverse band , or such markings absent 96 96 (95) Wing with a distinct transverse band Wing without transverse markings 97 97 (96) Pronotum with width less than 20 times length at mid-dorsal line . Brazil estfarchina sp. n. (p. 64)Pronotum with width more than 20 times length at mid-dorsal line. Brazil subfasciata Westwood (p. 64) 98 (96) Tegmen with claval area dark brown. Panama fasciata Metcalf (p. 65) Tegmen with claval area pale 99 99 (98) Tegmen with distal transverse band at level of second fork of medial vain . Panama douglasi sp. n. (p. 65)Tegmen with distal transverse band at level of radial-medial cross-vein 100 100 (99) Tegmen with distal transverse band much narrower and paler than basal band . Brazil knight i sp. n. (p. 66)Tegmen with distal transverse band as wide as, and little paler, than basal band. Brazil pulchella sp. n. (p. 66) 101 (90) Tegmen with three dark transverse bands 102 Tegmen with four or more dark transverse bands 102(101) Pronotum with width more than 30 times length at mid-dorsal line 103 Pronotum with width less than 30 times length at mid-dorsal line 105 103(102) Wing lacking distinct markings. Female with length of tegmen not less than 11 mm. Peru distinct a sp. n. (p. 66) 16 PETER S. BROOMFIELD Wing with two distinct transverse bands. Female with length of tegmen less than 1 1 mm . 104104(103) Wing with basal transverse band at level of first fork of cubital vein, unbroken. Brazil hengist sp. n. (p. 67)Wing with basal transverse band at level of medial-cubital cross-vein, interrupted medially. Brazil, Trinidad josiannasp. n. (p. 67) 105(102) Disc of mesonotum with a dark brown spot medially. Brazil pseudoerecta sp. n. (p. 68) Disc of mesonotum unicolorous brownish yellow 106 106( 105) Pronotal width not more than 22 times length at mid-dorsal line 107 Pronotal width not less than 25 times length at mid-dorsal line Ill 107(106) Wing with cubital-medial cross-vein broadly margined pale smoky brown, lacking distinct transverse bands. Brazil perspicua sp. n. (p. 68) Wing with one or two distinct transverse bands 108 108(107) Wing with a single transverse band 109 Wing with not less than two transverse bands 110 109(108) Wing with posterior and apical margins dark smoky brown. Brazil, Guyana agilis sp. n. (p. 69) Wing with posterior and apical margins hyaline. Brazil daudatasp. n. (p. 69) 110(108) Wing with two dark transverse bands. Brazil James/ sp. n. (p. 70) Wing with three transverse bands Ill 111(110) Wing with second transverse band extending unbroken from apex of clavus to medial vein.Bolivia carosella sp. n. (p. 70) Wing with second transverse band interrupted between first and second branches of cubital vein. Colombia harmonia sp. n. (p. 70) 112(101) Tegmen with four transverse bands 115 Tegmen with five or six transverse bands 113 113(112) Tegmen with six transverse bands. Peru silvanasp. n. (p. 71) Tegmen with five transverse bands 114 114(113) Tegmen more than 8 mm. Pronotal width more than 25 times length at mid-dorsal line. Brazil decora sp. n. (p. 72) Tegmen less than 8 mm. Pronotal width less than 20 times length at mid-dorsal line. Brazil bellasp. n. (p. 71) 115(1 12) Head with genae adjacent to eyes narrowly very dark brown. Bolivia bolivianna sp. n. (p. 72) Head with genae pale throughout 116 116(115) Ocelli very large and prominent. Brazil persephonesp. n. (p. 73) Ocelli small, not prominent 117 117(116) Wing with three dark transverse bands. Bolivia marshalli sp. n. (p. 73) Wing with two transverse bands, or lacking dark markings 118 118(117) Wing with two pale brownish transverse bands. Guyana neonebulosa Muir (p. 74) Wing lacking distinct transverse bands 119 119(118) Fore tibia and tarsi unicolorous pale yellow 120 Fore tibia with a narrow dark brown band subapically, tarsi brownish. Ecuador amaranths sp. n. (p. 74)120( 1 19) Antenna very long , extending well beyond anterior margins of genae . Surinam magica sp. n. (p. 74)Antenna short, not extending beyond anterior margins of genae. Bolivia, Peru geoffreyi sp. n. (p. 75) Key to species of Mysidia (based on male genitalia) It has not been possible to examine the male genitalia of the following species, which are therefore omittedfrom this key: immaculata, lactiftora, maculicosta, punctifera, quadrifasciata, squamigera, stigma, sub fas-data and subfusca. The genitalia of robusta are damaged, and this species is also omitted. 1 Paramere with a ventral process 2 Paramere lacking a ventral process 11 2 (1) Paramere with primary dorsal process absent (Fig. 391) fowlerisp. n. (p. 46) Paramere with primary dorsal process well developed 3 3 (2) Paramere with ventral process situated subbasally 4 Paramere with ventral process situated at , or distal to , midlength 6 TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 17 4 (3) Shaft of aedeagus with subapical dorsal processes rounded, flap-like (Fig. 174) distant! sp. n. (p. 49)Shaft of aedeagus with subapical dorsal processes long, slender, spine-like 5 5 (4) Shaft of aedeagus with lateral pair of subapical processes short, medial pair apically bifurcate (Fig. 175) claudata sp. n. (p. 69) Shaft of aedeagus with lateral processes long, medial processes simple (Fig. 176) pulchella sp. n. (p. 66) 6 (3) Shaft of aedeagus with subapical processes long and spine-like (Fig. 177) fulvodorsalis sp. n. (p. 32)Shaft of aedeagus with subapical processes flap-like 7 7 (6) Paramere with ventral process small and hook-like (Fig. 396) douglasi sp. n. (p. 65) Paramere with ventral process large and rounded 8 8 (7) Paramere with primary dorsal process vertically directed (Fig . 397) bizzara sp. n. (p . 56) Paramere with primary dorsal process strongly inclined posteriorly 9 9 (8) Paramere with dorsal surface strongly dorsally produced subapically (Fig. 399) 10 Paramere with apex rounded (Fig. 398) jamesi sp. n. (p. 70) 10 (9) Shaft of aedeagus with subapical flap-like processes dorsally directed (Fig. 289) enjebetta sp. n. (p. 51)Shaft of aedeagus with subapical flap-like processes not dorsally directed (Fig. 290) carosella sp. n. (p. 70) 11 (1) Paramere with a small, hook-like, secondary dorsal process subbasally 12 Paramere with secondary dorsal process either large and rounded or absent 14 12 (11) Paramere with primary dorsal process absent (Fig. 401) peregrina sp. n. (p. 45) Paramere with primary dorsal process distinct 13 13 (12) Shaft of aedeagus with lateral subapical processes apically bifurcate (Fig. 183) venusta sp. n. (p. 24)Shaft of aedeagus with lateral subapical processes apically simple (Fig. 184) augusta sp. n. (p. 54) 14 (11) Paramere with dorsal margin strongly produced and folded towards midline along entire length 15 Paramere with dorsal margin not produced throughout 17 15 (14) Shaft of aedeagus with subapical dorsal processes each bearing a small acute spine (Fig. 295) ariasi sp. n. (p. 37) Shaft of aedeagus with subapical dorsal processes greatly produced and apically bifur-cate 16 16 (15) Shaft of aedeagus with apices of subapical dorsal processes almost meeting at midline (Fig. 186) amazona sp. n. (p. 26) Shaft of aedeagus with apices of subapical dorsal processes diverging (Fig. 187) molest a sp. n. (p. 48) 17 (14) Paramere with primary dorsal process large 19 Paramere with primary dorsal process reduced or absent 18 18 (17) Paramere with primary dorsal process obsolete; secondary dorsal process small, bearing three acute spines (Fig. 407) glauca Distant (p. 35) Paramere with primary dorsal process absent; secondary dorsal process large, hook-like,basally directed (Fig. 408) cinerea Fennah (p. 76) 19 (17) Paramere with primary dorsal process produced dorsally and/or posteriorly 29 Paramere with primary dorsal process not produced 20 20 (19) Paramere with primary dorsal process bearing interlocking surfaces 23 Paramere with primary dorsal process lacking interlocking surfaces 21 21 (20) Shaft of aedeagus dorsally with two pairs of acute spine-like subapical processes (Fig. 190) agilissp. n. (p. 69) Shaft of aedeagus dorsally with a single pair of flap-like processes 22 22 (21) Paramere with primary dorsal process bearing a single, posteriorly directed, spine-like projection only (Fig. 410) ecuadoria sp. n. (p. 53) Paramere with primary dorsal process bearing a long, curving, hook-like projection anda small spine (Fig. 411) decora sp. n. (p. 72) 23 (20) Paramere bearing a long, medially directed secondary process distad of primary process (Fig. 412) lacteolasp. n. (p. 39) Paramere lacking a secondary dorsal process 24 18 PETER S. BROOMFIELD 24 (23) Shaft of aedeagus with slender, apically bifurcate, lateral processes (Fig. 194) cheesemani sp. n. (p. 54)Shaft of aedeagus lacking lateral processes 25 25 (24) Shaft of aedeagus with subapical dorsal processes strongly bifurcate 26 Shaft of aedeagus with subapical dorsal processes not bifurcate 27 26 (25) Paramere with anterior component of primary dorsal process strongly curving posterior- ly (Fig. 414) nemorensis sp. n. (p. 51) Paramere with anterior component of primary dorsal process inclined anteriorly (Fig.415) polyhymnia sp. n. (p. 23) 27 (25) Shaft of aedeagus with dorsal components of subapical processes strongly hooked posteriorly (Fig. 307) limpidasp. n. (p. 25) Shaft of aedeagus with dorsal components of subapical processes regularly curvingposteriorly 28 28 (27) Paramere with anterior and posterior components of dorsal process of approximately equal size (Fig. 417) nitida sp. n. (p. 26) Paramere with posterior component of dorsal process larger than the anterior (Fig.418) amarantha sp. n. (p. 74) 29 (19) Paramere with dorsal process slender, greatly produced dorsally, apex hooked, lacking interlocking surf aces 30 Paramere either with dorsal process not greatly produced dorsally, or with one or bothinterlocking surfaces present 31 30 (29) Shaft of aedeagus in dorsal aspect with maximum width greater than two-thirds length (Fig. 201) erectesp. n. (p. 55) Shaft of aedeagus in dorsal aspect with maximum width little greater than one-halflength (Fig. 202) pseudoerecta sp. n. (p. 68) 31 (29) Paramere with dorsal process bearing two interlocking surfaces, or with only the basal surface present 34 Paramere with only the distal interlocking surface present 32 32 (31) Paramere with a large, acute, medially directed process on dorsal surface subapically (Fig. 421) stall sp. n. (p. 62) Paramere not strongly produced subapically 33 33 (32) Shaft of aedeagus with ventral surface subapically bearing numerous small acute spines (Fig.313) knightisp. n. (p. 66) Shaft of aedeagus with ventral surface unarmed (Fig. 314) persephone sp. n. (p. 73) 34 (31) Paramere with basal interlocking surface hook-like , distal surface obsolete (Fig . 424) miner va sp. n. (p. 47) Paramere with both interlocking surfaces well developed 35 35 (34) Paramere with basal interlocking surface greatly produced, curving dorsally and post-eriorly (Fig. 425) harmonia sp. n. (p. 70) Paramere with basal interlocking surface not produced 36 36 (35) Paramere with a large, medially directed, apically acute process covered with very small tooth-like spines on dorsal margin distad of primary dorsal process 37 Paramere lacking a secondary process 38 37 (36) Shaft of aedeagus bearing long, acute, spine-like processes on dorsal surface subapically (Fig. 318) biancasp. n. (p. 42) Shaft of aedeagus with flap-like processes only (Fig. 319) pseudocostata sp. n. (p. 41) 38 (36) Paramere with dorsal process produced posteriorly 42 Paramere with dorsal process not produced posteriorly 39 39 (38) Paramere with dorsal process slender, dorsally directed (Fig. 428) bihula sp. n. (p. 53) Paramere with dorsal process broadly rounded 40 40 (39) Shaft of aedeagus in vertical aspect with length greater than three times maximum width (Fig. 210) panamensis sp. n. (p. 61) Shaft of aedeagus with length less than three times maximum width 41 41 (40) Shaft of aedeagus in lateral aspect with subapical dorsal processes very slender, apices curving dorsally (Fig. 321) isteria sp. n. (p. 59) Shaft of aedeagus in lateral aspect with subapical processes broad, apices inclinedantero-ventrally (Fig. 322) estfarchina sp. n. (p. 64) 42 (38) Paramere with dorsal process bearing a ventrally directed node-like projection at somewhat distad of midlength (Fig. 432) insolitasp. n. (p. 51) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 19 Paramere with dorsal process simple 43 43 (42) Shaft of aedeagus with ventrally directed processes subapically on ventral surf ace 44 Shaft of aedeagus with ventral surface unarmed 49 44 (43) Shaft of aedeagus with subapical process assymmetrical (Fig. 214). Paramere with apex strongly produced dorsally (Fig. 433) athenasp. n. (p. 28) Shaft of aedeagus symmetrical . Paramere with apex not produced dorsally 45 45 (44) Shaft of aedeagus slender , almost parallel-sided in dorsal aspect 46 Shaft of aedeagus considerably expanded subapically 48 46 (45) Shaft of aedeagus with ventral processes only (Fig. 325) sanguinea sp. n. (p. 44) Shaft of aedeagus with both dorsal and ventral processes 47 47 (46) Paramere with apex simple, acute; dorsal process situated at three-fifths length (Fig. 435) acidaloides Fowler (p. 29) Paramere with apex obtusely rounded, bearing an acute, medially curving process;dorsal process subapical (Fig. 436) krameri sp. n. (p. 54) 48 (45) Shaft of aedeagus with two pairs of ventral spine-like processes subapically (Fig . 328) adusta sp. n. (p. 22) Shaft of aedeagus with a single pair of ventrally curving spine-like processes apically(Fig.329) havilandi sp. n. (p. 36) 49 (43) Shaft of aedeagus with subapical processes flap-like and/or spine-like; if flap-like then terminating in long acute spines 61 Shaft of aedeagus with subapical processes flap-like , not terminating in long acute spines 50 50 (49) Shaft of aedeagus with subapical processes each bearing a small , laterally directed , acute spine subbasally (Fig. 219) richardsisp. n. (p. 25) Shaft of aedeagus with subapical processes not as above 51 51 (50) Paramere with a large, medially directed, secondary dorsal process distad of primary process (Fig. 440) costata (Fabricius) (p. 40) Paramere lacking a secondary dorsal process 52 Paramere very broadly rounded, length from base of apodeme to apex very little greater than maximum width (Fig. 441) adamare sp. n. (p. 60) Paramere with length well in excess of 1 -5 times maximum width 53 Shaft of aedeagus in dorsal aspect more than four times maximum width; subapical processes longitudinally aligned and dorsally directed (Fig . 222) simpla sp. n. (p. 63) Shaft of aedeagus in dorsal aspect less than 3-5 times maximum width; subapical processes transversally aligned and anteriorly directed 54 54 (53) Shaft of aedeagus in dorsal aspect less than twice maximum width (Fig . 224) liquids sp. n. (p. 37)Shaft of aedeagus in dorsal aspect not less than 2 5 times maximum width 55 55 (54) Shaft of aedeagus with subapical processes each produced posteriorly and dorsally into a short, acute spine (Fig. 223) marshallisp. n. (p. 73) Shaft of aedeagus with subapical processes not as above 56 56 (55) Shaft of aedeagus in dorsal aspect with subapical processes broadly rounded 59 Shaft of aedeagus with apices of subapical processes transverse , lateral angles acute 57 57 (56) Shaft of aedeagus with apices of subapical processes finely serrated (Fig. 225) fasciata Metcalf (p. 65)Shaft of aedeagus with subapical processes not as above 58 58 (57) Paramere with interlocking surfaces closely opposed , truncate (Fig . 445) hy nl inn sp. n. (p. 28)Paramere with interlocking surfaces acute, spine-like (Fig. 447) diana sp. n. (p. 43) 59 (56) Shaft of aedeagus with subapical processes extending anteriorly for two-fifths length (Fig. 228) albipennis Westwood (p. 38) Shaft of aedeagus with subapical processes extending anteriorly for approximatelyone-quarter length 60 60 (59) Shaft of aedeagus with subapical processes each bearing a large, rounded, medially directed lobe antero-dorsally (Fig. 229) bella sp. n. (p. 71) Shaft of aedeagus with subapical processes simple (Fig. 230) neonebulosa Muir (p. 74) 61 (49) Shaft of aedeagus with all subapical dorsal , processes slender, spine-like 62 Shaft of aedeagus with some or all subapical dorsal processes broad , flap-like 66 62 (61) Shaft of aedeagus with three pairs of subapical processes 63 Shaft of aedeagus with two pairs of subapical processes 64 20 PETER S. BROOMFIELD 63 (62) Shaft of aedeagus with medial pair of subapical processes very long and slender (Fig. 231) nigrithorax sp. n. (p. 63) Shaft of aedeagus with medial pair of subapical processes short (Fig. 232) I licit ITU sp. n. (p. 35) 64 (62) Paramere with dorsal margin narrowly produced towards midline apically (Fig. 453) infedelis sp. n. (p. 51)Paramere with apex regularly rounded 65 65 (64) Paramere very slender from base to approximately midlength (Fig. 454). Shaft of aedeagus with subapical processes broad basally, medial pair strongly diverging (Fig. 234) transversa sp. n. (p. 32) Paramere gradually broadening from base (Fig. 455). Shaft of aedeagus with subapicalprocesses slender from base, apices very weakly diverging (Fig. 235) . . intima sp. n. (p. 56) 66 (61) Shaft of aedeagus with five pairs of subapical processes (Fig. 236) . . caliginosa Walker (p. 23) Shaft of aedeagus with two pairs of subapical processes 67 67 (66) Shaft of aedeagus with medial subapical process slender, longitudinally aligned, bearing numerous, very small, acute tubercules anteriorly and/or dorsally, lateral processes spine-like 68 Shaft of aedeagus with medial subapical process lacking tubercules 70 68 (67) Shaft of aedeagus with lateral subapical processes long , apically serrated (Fig . 346) calypso sp. n. (p. 44)Shaft of aedeagus with subapical processes short , apically acute 69 69 (68) Shaft of aedeagus with lateral spine-like subapical processes very much shorter than medial processes (Fig. 238) wUHamsi sp. n. (p. 33) Shaft of aedeagus with lateral spine-like subapical processes only slightly shorter thanmedial processes (Fig. 239) albifasciata sp. n. (p. 31) 70 (67) Shaft of aedeagus with , in addition to paired lateral processes , a single , dorsally directed , spine-like, medial process subapically (Fig. 389) 71 Shaft of aedeagus with all subapical processes paired 72 71 (70) Shaft of aedeagus with paired spine-like processes long, antero-dorsally directed, situated submedially (Fig. 278) pseudonebulosa Muir (p. 75) Shaft of aedeagus with paired spine-like processes short, laterally directed, situatedlaterally (Fig. 240) josianna sp. n. (p. 67) 72 (70) Shaft of aedeagus with a single pair of slender, spine-like, antero-laterally directed processes subapically on lateral surfaces 73 Shaft of aedeagus not as above 76 73 (72) Shaft of aedeagus with medial pair of subapical processes very large, flap-like, antero- dorsally directed, irregularly rounded anteriorly (Fig. 241) .... nigrifrontalis sp. n. (p. 52)Shaft of aedeagus with medial subapical processes not as above 74 74 (73) Shaft of aedeagus with lateral subapical processes short, hooked, anteriorly directed (Fig. 242) c/avasp. n. (p. 62) Shaft of aedeagus with lateral subapical processes long, slender 75 75 (74) Shaft of aedeagus with lateral subapical processes deeply serrated apically; medial processes each with a long, acute, anteriorly directed, spine-like projection dorsally (Fig. 354) inquinata sp. n. (p. 24) Shaft of aedeagus with lateral subapical processes apically acute; medial processes not asabove (Fig. 355) gracilis sp. n. (p. 35) 76 (72) Paramere with dorsal margin strongly produced apically, inclined ventro-medially (Fig. 466) fuscofrontalis sp. n. (p. 38) Paramere with dorsal margin not produced apically 77 77 (76) Paramere with primary dorsal process situated at three-quarters length (Fig. 467) formosa sp. n. (p. 61)Paramere with primary dorsal process situated at, or basad of, mid-length 78 78 (77) Shaft of aedeagus asymmetrical , apex strongly rotated clockwise (Fig . 358) cooper/ sp. n. (p. 42)Shaft of aedeagus symmetrical , not apically rotated 79 79 (78) Shaft of aedeagus with a pair of long, ventrally directed, lateral processes subapically 80 Shaft of aedeagus with subapical processes not ventrally inclined 81 80 (79) Paramere with apex acute , interlocking surfaces small and acute (Fig . 469) punctum (Fabricius) (p. 27) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 21 Paramere with apex broadly rounded, interlocking surfaces large and opposed (Fig.470) magica sp. n. (p. 74) 81 (79) Shaft of aedeagus with a pair of large, horizontal, flap-like subapical processes strongly overlapping medially 82 Shaft of aedeagus with subapical processes not strongly overlapping 85 82 (81) Shaft of aedeagus with subapical processes each bearing a slender, dorsally directed, spine-like projection medially ; anterior angles acute , greatly produced anteriorly 83 Shaft of aedeagus with flap-like processes lacking medial spine-like projections, notproduced anteriorly 84 83 (82) Shaft of aedeagus with anterior extensions of flap-like processes broad, overlapping basally (Fig. 250) bolivianna sp. n. (p. 72) Shaft of aedeagus with anterior extensions of flap-like processes slender, not over-lapping basally (Fig. 251) silvana sp. n. (p. 71) 84 (82) Shaft of aedeagus with subapical flap-like processes each bearing a slender, curving, spine-like projection basally (Fig. 252) hengist sp. n. (p. 67) Shaft of aedeagus with subapical processes basally unarmed (Fig. 253). musics sp. n. (p. 32) 85 (81) Shaft of aedeagus with a very broad, transverse, posteriorly directed, flap-like process subbasally, extending to midlength on dorsal surface (Fig. 365) . . . etheldreda sp. n. (p. 36)Shaft of aedeagus not as above 86 86 (85) Shaft of aedeagus with a medial subapical process in addition to paired lateral processes (Fig. 255) dollingisp. n. (p. 43) Shaft of aedeagus with paired subapical processes only 87 87 (86) Shaft of aedeagus with a single pair of subapical processes laterally, not extending over dorsal surface 88 Shaft of aedeagus with subapical processes on , or extending over , dorsal surface 89 88 (87) Shaft of aedeagus with subapical processes triangular (Fig. 256) diabola sp. n. (p. 58) Shaft of aedeagus with subapical processes long and slender (Fig. 257) neoasinella sp. n. (p. 30) 89 (87) Shaft of aedeagus with a pair of very large, longitudinally aligned, flap-like processes subapically , each bearing two small , anteriorly directed , tooth-like pro j ections 90 Shaft of aedeagus not as above 93 90 (89) Shaft of aedeagus in lateral aspect with posterior tooth-like projection larger than the anterior (Fig. 369) andessp. n. (p. 52) Shaft of aedeagus with posterior tooth-like projection not the larger 91 91 (90) Shaft of aedeagus with posterior tooth-like projection much smaller than the anterior (Fig.370) tikalmesp. n. (p. 34) Shaft of aedeagus with tooth-like projections of equal size 92 92 (91) Shaft of aedeagus in dorsal aspect strongly tapering from base to apex (Fig. 260) obscura Metcalf (p. 49)Shaft of aedeagus with lateral margins subparallel (Fig. 261) varia sp. n. (p. 34) 93 (89) Paramere with apex broadly , if occasionally , somewhat irregularly , rounded 95 Paramere with apex somewhat produced dorsally , inclined towards mid-line 94 94 (93) Shaft of aedeagus with a single pair of massive, flap-like, strongly asymmetrically arranged subapical processes (Fig. 262) lucianna sp. n. (p. 45) Shaft of aedeagus with two pairs of symmetrically arranged, spine-like subapicalprocesses (Fig. 263) fuscomaculata sp. n. (p. 59) 95 (93) Shaft of aedeagus with one pair of longitudinally aligned, dorsally directed, anteriorly acutely angled, flap-like processes adjacent to midline (Fig. 264) ttoydi sp. n. (p. 58) Shaft of aedeagus not as above 96 96 (95) Shaft of aedeagus in dorsal aspect with length not greater than twice maximum width 97 Shaft of aedeagus in dorsal aspect with length greater than two and one-half timesmaximum width 99 97 (96) Paramere with basal interlocking surface very large , distal surface obsolete (Fig . 486) insania sp. n. (p. 60)Paramere with interlocking surfaces of approximately equal size 98 98 (97) Paramere with dorsal process very small (Fig. 487). Shaft of aedeagus with apices of subapical processes slender, dorsally directed (Fig. 377) nebulosa (Germar) (p. 56) Paramere with dorsal process large (Fig. 488). Shaft of aedeagus with apices of subapicalprocesses broad, anteriorly directed (Fig. 378) perspicua sp. n. (p. 68) 22 PETER S. BROOMFIELD 99 (96) Paramere with dorsal surface subapically produced into a rounded, medially directed secondary process bearing numerous robust spines 100 Paramere lacking a distinct subapical secondary process 101 100(99) Shaft of aedeagus with subapical processes simple, triangular (Fig. 268) fiavilla sp. n. (p. 29)Shaft of aedeagus with subapical processes rounded, each bearing a long, slender,antero-ventrally directed, spine-like projection at midlength on anterior margin (Fig. 269) unimaculate sp. n. (p. 28) 101(100) Shaft of aedeagus with subapical processes long, slender, spine-like, dorsally or anterior-ly directed 102 Shaft of aedeagus with subapical processes broad or , if slender , ventrally directed 105 102(101) Shaft of aedeagus with subapical processes curving anteriorly 103 Shaft of aedeagus with subapical processes straight, dorsally directed (Fig. 381) maculosa sp. n. (p. 55) 103(102) Paramere with dorsal process situated at midlength (Fig. 492) distincta sp. n. (p. 66) Paramere with dorsal process situated somewhat distad of midlength 104 104(103) Shaft of aedeagus with subapical processes strongly diverging (Fig. 272). Paramere with interlocking surfaces truncate, distant from each other (Fig. 493) whimper! sp. n. (p. 62)Shaft of aedeagus with subapical processes weakly diverging (Fig. 273). Paramere with interlocking surfaces acute, closely opposed (Fig. 494) testacea (Fabricius) (p. 57) 105(101) Paramere with dorsal process situated basad of midlength, very long and slenderly produced posteriorly (Fig. 495) geoffreyisp. n. (p. 75) Paramere with dorsal process situated at approximately midlength 106 106(105) Shaft of aedeagus with subapical processes longitudinally aligned, dorsally directed,massive, each produced posteriorly into a short, acute, ventrally directed, spine-like projection (Fig. 386) delicatissima Fowler (p. 41) Shaft of aedeagus with subapical processes transversally aligned , anteriorly directed 107 107(106) Shaft of aedeagus with subapical processes each bearing a long, slender, anteriorly directed, spine-like projection basally (Fig. 276) mylesi sp. n. (p. 33) Shaft of aedeagus lacking slender projections 108 108(107) Shaft of aedeagus with subapical processes deeply notched apically (Fig. 277) Henrietta sp. n. (p. 49)Shaft of aedeagus with subapical processes apically acute (Fig. 279) . . . albicans (Stal) (p. 50) Mysidia asinella sp. n. Female: head 0-97 mm long, 1-12 mm wide; pronotum 2-52 mm wide; legmen 12-00-12-50 mm long; wing7-05 mm long. Male unknown. Length of frons 5 times width at apex, c. 2-5 times width at base; ocelli small, distinct; clypeus c. as long asfrons; rostrum terminating immediately posterior to hind coxae. Pronotal width 18 times mid-dorsallength; fronto-lateral surfaces weakly carinate; tegula not carinate. Head bright scarlet, antenna yellow. Pronotum with mid-dorsal and fronto-lateral surfaces scarlet;scutellum scarlet; disc of mesonotum brown; abdomen brownish dorsally, tinged scarlet apically. Tegmenand wing dark brownish, veins brown, posterior margins very narrowly scarlet. Tegmen unmarked, basenarrowly scarlet; costal vein and branches, and apical part of subcostal vein and its branches reddish;radial-subcostal cross-vein, medial-radial apical cross-vein, and bases of fifth and sixth branches of medialvein white. Wing with a narrow, oblique, transverse, whitish band extending from costal to posteriormargins immediately distad of radial-medial cross-vein; otherwise unmarked. MATERIAL EXAMINED Holotype $, Brazil: Belem, Para, vi.1924 (Williams) (BMNH).Paratypes, Brazil: 4 $ , Breves, Lower Amazon (INPA; BMNH). In the absence of males, asinella is readily distinguished by the bright scarlet pigmentation of thehead and thorax, and by the single pale band on the otherwise dark brownish wing. Mysidia adusta sp. n. (Figs 217, 328, 437) Male: head 0-67 mm long, 1-03 mm wide; pronotum 2-37 mm wide; tegmen 9-80 mm long; wing 5-90 mmlong. Female: tegmen 11-70 mm long. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 23 Length of frons c. 4-5 times width at apex, c. 3 times width at base; ocelli large, prominent; clypeus aslong as frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 14 times length atmid-dorsal line; fronto-lateral surfaces and tegula not carinate. Head, excluding antenna, rostrum, and extreme baso-ventral margins usually scarlet; ocelli concol-orous. Fronto-lateral surfaces of pronotum each with a broad, scarlet band extending horizontally fromadjacent to dorsal margin of eye to lateral margin; tegula and disc of mesonotum pale brownish; scutellumirregularly tinged dull pink. Tegmen and wing dark smoky brown; veins and cross-veins dark brown;lacking prominent markings. Tegmen with veins over basal third narrowly edged yellowish hyaline; apicalforks of medial vein, second radial-medial cross- vein, and radial-subcostal cross-vein white; costal andposterior marginal veins very narrowly crimson. Wing unmarked; posterior marginal vein narrowlycrimson. Shaft of aedeagus broad ; apex with a pair of opposed , flap-like processes extending from ventral surface ;dorso-lateral surfaces subapically each with a long, slender, spine-like process; ventral surface subapicallywith two pairs of small, transversally aligned spines. Paramere with apex very broadly rounded; dorsalprocess situated at three-quarters length, large, strongly produced posteriorly. Subgenital plate producedmedially into a rounded, posteriorly directed, lobe bearing a fringe of long, erect, spine-like hairs. MATERIAL EXAMINED Holotype cT, Brazil: Amazonas, 120 km E. of Tapuruquara, 19.L1978 (Penny) (INPA).Paratype. Brazil: 1 $, Amazonas, Manaus (BMNH). M. adusta is readily distinguished by the pigmentation of the head and thorax, and the darkbrown tegmen and wing, both lacking transverse markings. Mysidia polyhymnia sp. n. (Figs 196, 306, 415) Male: head 0-61 mm long, 0-86 mm wide; pronotum 1-97 mm wide; tegmen 9-00 mm long; wing 5-10 mmlong. Female unknown. Length of frons c. 5 times width at apex, 2-33 times width at base; ocelli small, not prominent; clypeus c.0-33 longer than frons; rostrum extending to base of subgenital plate. Pronotal width c. 13 times mid-dorsallength, fronto-lateral surfaces and tegula distinctly carinate. Fronto-lateral surfaces of pronotum dorsal to upper margins of eyes brownish; tegula dark brown; dorsalsurface of abdomen tinged red basally . Tegmen and wing smoky brownish, veins dark brown. Tegmen withcross-veins pale, apical fork of medial vein surrounded by a very small white spot, costal margin basallydark brown. Wing unmarked. Shaft of aedeagus slender in lateral aspect, greatly expanded laterally; dorsal surface at approximatelytwo-thirds length with a pair of large processes, each terminating posteriorly in a long, slender, curvingspine, and anteriorly in a shorter, straighter spine. Paramere very robust; apex very obtusely rounded,almost truncate; dorsal process situated at three-fifths length, reduced, proximal component slender,inclined antero-dorsally and terminating in a medially directed hook, distal component short and rounded. MATERIAL EXAMINEDHolotype cf , Brazil: Amazon, Fonteboa (BMNH). The pigmentation of the tegmen and wing is closely similar to that of caliginosa and inquinata,but polyhymnia is distinguished by its larger size, relatively obscure ocelli, carinate tegula, andby the structure of the male genitalia. Mysidia caliginosa Walker(Figs 236, 348, 456, 463) Mysidia caliginosa Walker, 1858: 98. Holotype $, BRAZIL (BMNH) [examined].Mysidia rubra Metcalf, 1945: 128. Holotype cT, GUYANA (AMNH) [examined]. Syn. n. Male: head 0-60 mm long, 0-80 mm wide; pronotum 1-60 mm wide; tegmen 7-00-7-65 mm long; wing 4 -50mm long. Female: tegmen 7-20-9-80 mm long. Length of frons c. 5 times width at apex, 2-5 times width at base; ocelli very large and prominent; clypeusslightly longer than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width from10-12 times mid-dorsal length; fronto-lateral surfaces and tegula with carinae weak or obsolete. 24 PETER S. BROOMFIELD Frons and genae dorsally scarlet; ocelli yellow, broadly edged scarlet. Dorsal surfaces of thorax andabdomen from pale brown to deep reddish brown, usually scarlet, abdomen rarely blackish; tegula withdorsal margin broadly dark brown. Tegmen and wing dark brownish, unmarked; veins and cross-veinsdark brown; posterior marginal veins very narrowly crimson. Tegmen with costal, subcostal and radialveins frequently tinged crimson. Shaft of aedeagus cylindrical; dorsal surface subapically with a pair of large, flap-like processes, eachterminating in a slender, curving spine; lateral surfaces each with four slender spine-like processes.Paramere slender; apex narrowly rounded; dorsal process well developed, situated somewhat distad ofmid-length, apex weakly produced posteriorly; dorsal surface subasally with a group of short robust spines. MATERIAL EXAMINED Holotype $ (caliginosa), Brazil: Santarem (Bates) (BMNH). Holotype cf (rubrd), Guyana: Kartabo,Bartica District, 1920 (AMNH). Guyana: 1 CT, 2 $ (BMNH). Brazil: 1 $, Santarem (Bates) (BMNH); 2 $ (BMNH). Bolivia: 1 cf(BMNH). Colombia: 1 $ (BMNH). Ecuador: 2 cf , 2 $ (BMNH). Panama: 1 cf (FAMU). Peru: 1 cf(FAMU). Surinam: 1 (FAMU). Trinidad: 1 $ (BMNH). The holotype of rubra Metcalf has the tegmina and wings damaged or missing; the genitalia arepreserved in balsam and are not accessible for detailed study. This species is readily distinguished by the dark brown, unmarked, tegmen and wing, thepigmentation of the head and body, and by the structure of the male genitalia. Mysidia inquinata sp. n. (Figs 243, 354, 464) Male: head 0-62 mm long, 0-67 mm wide; pronotum 1-47 mm wide; tegmen 7-22-7-25 mm long; wing4-34mm long. Female: tegmen 8-70 mm long. Length of frons c. 6 times width at apex, c. 3 times width at base; ocelli very large and prominent; clypeusslightly longer than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 14times mid-dorsal length; fronto-lateral surfaces and tegula with carinae weak or obsolete. Frons and genae anterior to eyes usually tinged deep crimson; lateral surfaces of clypeus weakly tingedcrimson. Fronto-lateral surfaces of pronotum dorsad to level of eyes often suffused reddish; tegula reddish,dorsal margins dark reddish brown; disc of mesonotum reddish brown; dorsal surface of abdomenoccasionally dark brown. Tegmen and wing uniformly dark brownish, unmarked; veins and cross- veinsdark brown; posterior margins often very narrowly crimson. Tegmen with apical branches of subcostal andradial veins bright crimson or white. Shaft of aedeagus cylindrical; dorsal surface subapically with a pair of large, flap-like processes, eachbearing a long spine subbasally on dorsal surface, and numerous very small and blunt spines on internal andventral surfaces; lateral surfaces each with a long, apically serrated, process subapically. Paramere slender;apex broadly rounded; dorsal process situated at three-fifths length, apex strongly produced posteriorly;dorsal surface at one-quarter length with a rounded, internally directed secondary process bearingnumerous short, robust spines. MATERIAL EXAMINED Holotype cf , Brazil: Amazon, Fonteboa (BMNH).Paratypes. Brazil: 14 cf, 11 $, Amazonas (BMNH; INPA). The tegminal and wing pigmentation of inquinata is very similar to that of caliginosa andpolyhymnia, but it differs from the former in the proportions and pigmentation of the head andpronotum, and from the latter by its much smaller size, and from both in the structure of themale genitalia. Mysidia venusta sp. n. (Figs 183, 293, 402) Male: head 0-55 mm long, 0-69 mm wide; pronotum 1-47 mm wide; tegmen 6-20 mm long; wing 3-40 mmlong. Female unknown. Length of frons 4 times width at apex, 2-66 times width at base; ocelli small, indistinct; clypeus as long asfrons; rostrum extending to posterior surface of hind coxae. Pronotal width slightly less than 12 timeslength at mid-dorsal line; fronto-lateral carinae prominent; tegula not carinate. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 25 Head and body unmarked . Tegmen and wing whitish hyaline ; veins pale yellow , otherwise totally devoidof pigmentation. Shaft of aedeagus basally slender, broadly expanded subapically; dorsal surface subapically with a pair oflarge, apically shallowly bifurcate processes; a pair of rounded, flap-like processes; at midline, a single,apically shallowly concave, flap-like process. Paramere with ventral margin somewhat dorsally producedapically; dorsal process very reduced, situated at approximately midlength, not at all produced dorsally orposteriorly; dorsal surface at one-third length with a simple hook-like process, basally with a rounded lobebearing numerous robust spines; ventral surface basally produced into a small rounded lobe bearingnumerous robust spines. MATERIAL EXAMINEDHolotype cf , Brazil: Amazon, Rio Autaz, x (Roman) (NR). The male genitalia bear a close resemblance to those of cheesemani, but venusta is readilydistinguished by the complete absence of tegminal and wing pigmentation. Mysidia richardsisp. n. (Figs 219, 331, 439) Male: head 0-73 mm long, 0-90 mm wide; pronotum 2-10 mm wide; tegmen 8.92 mm long; wing 5-20 mmlong. Female unknown. Length of frons 6-5 times width at apex, c. 2-5 times width at base; ocelli obsolete; clypeus as long asfrons; rostrum terminating immediately posterior to hind coxae. Pronotal width 20 times mid-dorsallength; fronto-lateral surfaces and tegulae with carinae obsolete or absent. Head and body unmarked. Tegmen and wing whitish hyaline, veins and cross-veins pale brownishyellow. Tegmen with costal cell pale smoky yellow; posterior margin very pale yellowish brown. Wing withposterior margin very narrowly and faintly yellowish. Shaft of aedeagus with dorsal surface subapically bearing a pair of large flap-like lobes extendinganteriorly to midlength, each with a small lateral spine at c. midlength; ventral surface at midlength with anantero-ventrally directed, apically rounded process at each lateral angle. Paramere slender, apex narrowlyrounded; dorsal process situated slightly basad of midlength, posteriorly produced, with a large, rounded,medially directed process on internal surface at three-quarters length. MATERIAL EXAMINEDHolotype cf , Guyana: Blairmont, ix.1923 (Williams) (BMNH). This species is distinguished by the lack of distinct markings on the head, body, tegmen andwing, and by the structure of the male genitalia. Mysidia Hmpida sp. n. (Figs 197, 307, 416) Male: head 0-63 mm long, 0-88 mm wide; pronotum 1-87 mm wide; tegmen 8-33-8-80 mm long; wing 4-75mm long. Female unknown. Length of frons 5 times width at apex, twice width at base; ocelli small, obscure; clypeus one-third longerthan frons; rostrum extending beyond hind coxae. Pronotal width 30 times mid-dorsal length; fronto-lateral surfaces not carinate; tegula distinctly carinate. Head and body unmarked. Tegmen and wing whitish hyaline basally, weakly tinged yellowish frommidlength, veins yellow. Tegmen with posterior and apical margins, cross-veins and branches of veinsbroadly edged pale fuscous, the last coalescing at midlength to form a very indistinct, pale, transverseband. Wing unmarked. Shaft of aedeagus very broad in dorsal aspect; apex transverse, flap-like, strongly produced dorsally;lateral surfaces subapically each with a very large, rounded, dorsally directed, flap-like process; dorsalsurface subapically with a pair of very large, flap-like processes medially, apex of each with posterior angleproduced into a long spine. Paramere robust, apex irregularly rounded; dorsal process situated somewhatdistad of midlength, small, apex not produced posteriorly. MATERIAL EXAMINED Holotype cf , Brazil: Mato Grosso, 1250'S 5147'W, cerradao, 2.iii.l968 (Richards) (BMNH).Paratype. Brazil: 1 cf , Para, Belem (BMNH). 26 PETER S. BROOMFIELD This species appears closely related to amarantha and nitida, but differs in the detailed structureof the male genitalia and in the pigmentation of the tegmen and wing. Mysidia robusta sp. n. (Fig 501) Male: head 0-84 mm long, 1-20 mm wide; pronotum 3-02 mm wide; tegmen 13-60 mm long; wing 8-50 mmlong. Female unknown. Length of frons 5 times width at apex, 2-33 times width at base; ocelli small, distinct; clypeus slightlylonger than frons; rostrum terminating slightly posterior to hind coxae. Pronotal width c. 10 timesmid-dorsal length; fronto-lateral surfaces and tegula not distinctly carinate. Head and thorax tinged reddish on dorsal surfaces, ocelli crimson. Tegmen and wing hyaline; veinsyellowish brown; posterior margins crimson, narrowly edged smoky brown. Tegmen with medial, cubitaland anal veins basally dark brown. Wing unmarked. Shaft of aedeagus broad, robust; dorsal surface subapically with a pair of large, flap-like processesadjacent to midline and extending over apex onto ventral surface. Paramere robust; apex broadly rounded;dorsal process situated at approximately two-thirds length, large, posteriorly produced; dorsal surfacesubbasally produced towards midline. Subgenital plate produced medially into a pair of short, broad lobescovered in very small obtuse spines. MATERIAL EXAMINEDHolotype cf , Brazil: Amazon, Fonteboa (BMNH). Amongst the largest species of the genus, robusta is readily distinguished by the yellowishhyaline pigmentation of the otherwise unmarked tegmen and wing. Mysidia nitida sp. n. (Figs 198, 308, 417) Male: head 0-62 mm long, 0-94 mm wide; pronotum 2-00 mm wide; tegmen 10.20 mm long; wing 5-85 mmlong. Female unknown. Length of frons 6-25 times width at apex, 2-5 times width at base; ocelli small, obscure; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 24 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale yellowish. Tegmen withposterior margin weakly tinged smoky brown; with a very indistinct, smoky brown, transverse band at levelof first fork of cubital vein, and a very faint, intermittent, brownish, transverse band at level ofmedial-cubital cross-vein. Wing with posterior margin very weakly tinged smoky brown, otherwiseunmarked. Shaft of aedeagus slender in lateral aspect, broadly expanded laterally; dorsal surface subapically with apair of large, flap-like processes, each bearing a large spine on postero-dorsal surface. Paramere veryrobust; apex obtusely rounded; dorsal process directed towards midline, not produced posteriorly. MATERIAL EXAMINED Holotype cf, Guyana: Amazon-Courantyne Divide, head of Oronoque River, 1937 (Beddington)(BMNH). The male genitalia show a similarity with those of amarantha, but the external characters aredistinct. Mysidia amazona sp. n. (Figs 186, 296, 405) Male: head 0-76 mm long, 1-10 mm wide; pronotum 2-40 mm wide; tegmen 10-04-11-05 mm long; wing6-00 mm long. Female unknown. Length of frons 6 times width at apex, c. 2-5 times width at base; ocelli small, distinct; clypeus one-sixthlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 19 times mid-dorsallength; fronto-lateral surfaces not distinctly carinate; tegula with carinae distinct. Head and body dark brownish, unmarked. Tegmen and wing hyaline, veins yellow. Tegmen with costaland subcostal areas pale brownish. Wing unmarked. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 27 Shaft of aedeagus very broad in dorsal aspect; apex recurving; lateral surfaces each with a very large,flap-like process extending over dorsal surface and overlapping at midline, each with anterior marginproduced dorsally into a slender secondary process with apical angles produced into acute spines.Paramere very robust, broadly rounded, apex obtuse; dorsal margin strongly curved medially andventrally, considerably expanded from mid-length to apex; dorsal process very small, situated at two-fifthslength, apex not produced. MATERIAL EXAMINED Holotype d", Brazil: Amazon, Rio Autaz, 31.x. 1914 (Roman) (NR).Paratype. 1 Cf , same data as holotype (BMNH). This species is readily distinguished by its large size, lack of pigmentation, and by the structure ofthe male genitalia. Mysidia punctum (Fabricius)(Figs 248, 359, 469) Derbe punctum Fabricius, 1803: 82. LECTOTYPE $, CENTRAL AMERICA (ZM), here designated [ex-amined]. Mysidia punctum (Fabricius) Westwood, 1840: 83. Mysidia steinbachi Distant, 1907: 396. LECTOTYPE cf , BOLIVIA (BMNH), here designated [examined].Syn. n. Male: head 0-55 mm long, 0-76 mm wide; pronotum 1-60 mm wide; tegmen 7-50-8-00 mm long; wing 4-52mm long. Female: tegmen 8-50-10-20 mm long. Length of frons slightly less than 8 times width at apex, 3 times width at base; ocelli small, indistinct;clypeus c. as long as frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 15-20times mid-dorsal length; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale brown, cross-veins weaklyedged yellowish brown. Tegmen with a prominent dark brown/black spot at one-third length, extendingfrom costal margin almost to second branch of cubital vein, a very indistinct, irregular, pale smoky brown,transverse band at midlength extending from costal margin over one-half width. Wing with cross-veinsweakly brownish; costal margin with a pale brown spot at one-third length. Shaft of aedeagus very greatly expanded dorso-ventrally and laterally over apical half; dorsal surfacesubapically with a pair of flap-like processes, each bearing a spine-like projection; ventral surfacesubapically with a pair of broad apically truncate processes. Paramere massive, tapering from midlength toacutely rounded apex; dorsal process situated at one-quarter length; dorsal surface at midlength with alarge, flap-like, roughly rectangular secondary process. MATERIAL EXAMINED Lectotype $ (punctum), Central America (Schmidt) (ZM). Lectotype cf (steinbachi), Bolivia: 1904(Steinbach) (BMNH) (badly damaged). Trinidad : 1 cf , St George (BMNH) . Peru : 2 cf , Callanga (BMNH) . Bolivia : 1 $ , San Antonio (BMNH) .Guyana: 3 $, Bartica (BMNH). Brazil: 1 cf (NR). Central America: 1 cf (paralectotype of punctum) (ZM)(head missing). This species is readily distinguished by the prominent dark brown spot on the costal area of thetegmen. Mysidia immaculata sp. n. Female: head 0-80 mm long, 1-25 mm wide; pronotum 2-90 mm wide; tegmen 14-80 mm long; wing 10-00mm long. Male unknown. Length of frons slightly greater than 5 times width at apex, 2-5 times width at base; ocelli obsolete;clypeus slightly longer than frons; rostrum terminating at level of hind coxae. Pronotal width 17 timesmid-dorsal length; fronto-lateral surfaces and tegula prominently carinate. Dorsal surfaces of head and thorax brownish; genae adjacent to eyes and fronto-lateral surfaces ofpronotum at level of eyes reddish; abdomen dorsally with a small red spot on either side of midline onsegments five and six, ventral surface and lateral margins reddish basally; median and posterior femorasubapically tinged reddish. Tegmen and wing whitish hyaline, veins pale yellow. Tegmen with apicalmargin very narrowly reddish brown. Wing unmarked. 28 PETER S. BROOMFIELD MATERIAL EXAMINEDHolotype $, Peru: Callanga (BMNH). One of the largest species of the genus, immaculata is distinguished by the lack of pigmentationof the tegmen and wing, and by the reddish pigmentation of the head and body. Mysidia hyalina sp. n. (Figs 226, 337, 445) Male: head 0-55 mm long, 0-82 wide; pronotum 1-66 mm wide; tegmen 8-68-8-48 mm long; wing 5-40 mmlong. Female unknown. Length of frons 6 times width at apex, twice width at base; ocelli very small, distinct; clypeus slightlylonger than frons; rostrum terminating at level of hind coxae. Pronotal width 13 times mid-dorsal length,fronto-lateral surfaces without carinae; tegula basally carinate. Vertex deep red; genae level with midline of eyes dark brown; fronto-lateral surfaces of pronotum eachwith a broad, horizontal, orange band extending from adjacent to eye to lateral margin; tegula with ventralmargin dull brown. Tegmen and wing hyaline, veins yellowish. Tegmen with cross-veins and forks of veinspale brown; apical fork of medial vein narrowly dark brown; claval margin with a small brown spot at levelof point of fusion of anal veins, a small brown apot adjacent to claval suture at level of first fork of medialvein. Wing with a small pale brown spot adjacent to claval suture at midlength; radial-medial cross-veinbrown. Shaft of aedeagus very slender; dorsal surface subapically with a pair of large flap-like processes.Paramere very robust; apex obtusely rounded; dorsal process situated at one-third length, small, apexstrongly produced posteriorly; dorsal surface subbasally produced, bearing numerous, long, robust spines. 'MATERIAL EXAMINED Holotype d", Jamaica: Moneague, ii.1904 (Walsingham) (BMNH).Paratype. 1 C?, same data as holotype (BMNH). This species is readily distinguished by the pigmentation of the head and pronotum, the relativelack of pigmentation of the tegmen and wing, and by the structure of the male genitalia. Mysidia unimaculata sp. n. (Figs 269, 380, 490) Male: head 0-50 mm long, 0-66 mm wide; pronotum 1-28 mm wide; tegmen 6-12 mm long; wing 3-65 mmlong. Female unknown. Length of frons 6-5 times width at apex, twice width at base; ocelli distinct; clypeus c. as long as frons;rostrum terminating immediately posterior to hind coxae. Pronotal width 15 times mid-dorsal length,fronto-lateral carinae distinct; tegulae weakly carinate. Head and body unmarked; disc of mesonotum with lateral angles brown. Tegmen and wing whitishhyaline. Tegmen with a prominent brown spot between apex of clavus and first branch of cubital vein,otherwise unmarked. Wing with a large brown spot between clavus and cubital vein at midlength,otherwise unmarked. Shaft of aedeagus slender, slightly expanded apically; dorsal surface subapically with a pair of large,broadly rounded, flap-like processes, each bearing a long tapering spine near base on anterior surface.Paramere very slender basally, broadening towards apex; dorsal process situated slightly distad ofmid-length, very large, not greatly produced; dorsal surface at one-quarter length with a short roundedsecondary process bearing numerous robust spines. MATERIAL EXAMINEDHolotype C?, Brazil: Para, Jabaty, v.1924 (Williams) (BMNH). This species is readily distinguished by the unique pigmentation of the tegmen and wing, and bythe structure of the male genitalia. Mysidia athena sp. n. (Figs 214, 324, 433) Male: head 0-69 mm long, 0-96 mm wide; pronotum 1-90 mm wide; tegmen 9-77-10-00 mm long; wing 6-40mm long. Female unknown. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDHNI (HOMOPTERA) 29 Length of frons c. 5 times width at apex, 2-5 times width at base; ocelli obsolete; clypeus c. as long asfrons; rostrum terminating at level of hind coxae. Pronotal width 15 times mid-dorsal length, fronto-lateralcarinae distinct; tegulae with carinae obsolete or absent. Genae each with a small dark brown spot adjacent to dorsal margin of eye, occasionally extending ontofrons, with a similar marking level with mid-line of eye; antenna deep red; lateral surfaces of paraclypeusdeep red; fronto-lateral surfaces of pronotum each with a deep red horizontal band extending fromadjacent to eye to lateral margin; apices of anterior and medial coxae broadly deep red. Tegmen and wingwhitish hyaline, veins and cross-veins yellowish. Tegmen with a small dark brown spot on costal celladjacent to first fork of radial vein; clavus with a larger dark brown spot at apex, and a a smaller spot at levelof point of fusion of anal veins. Wing with a dark brown spot on first branch of cubital vein at midlength;posterior margin with a single, semi-circular, dark brown spot between each branch of anal, claval andmedial veins. Shaft of aedeagus slender, cylindrical, asymmetrical; dorsal surface apically with a large, twisted,flap-like process terminating anteriorly in a blunt point; left dorso-lateral surface subapically with a largetriangular process; right lateral surface subapically with a large flap- like process terminating posteriorly inan acute spine-like lobe. Paramere very large; apex acute, strongly produced dorsally; dorsal process welldeveloped, situated at midlength, apex postero-dorsally directed. MATERIAL EXAMINED Holotype cf , Ecuador: Cachabe, i.1897 (Rosenberg) (BMNH).Paratype. 1 cf , same data as holotype (BMNH). The tegminal and wing pigmentation of this species closely resembles that of acidaloides, but theprominent markings of the head and thorax, and the structure of the male genitalia, render itreadily distinguishable. Mysidia Havilla sp. n. (Figs 268, 379, 489) Male: head 0-42 mm long, 0-65 mm wide; pronotum 1-40 mm wide; tegmen 6-12-6-38 mm long; wing3-65 mm long. Female unknown. Length of frons 7-5 times width at apex, c. 1-5 times width at base; ocelli small, obscure; rostrumextending to anterior surface of hind coxae. Pronotal width 13 times mid-dorsal length, fronto-lateralcarinae very prominent; tegula weakly carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale. Tegmen with a small darkbrown spot adjacent to point of separation of fused medial and radial-subcostal veins, another at point ofseparation of fused radial and subcostal veins, a third at first fork of cubital vein, another on posteriormargin at midlength of clavus, a fifth adjacent to medial vein at level of first fork of cubital vein, anotheradjacent to apex of clavus, and a seventh over apical fork of radial vein; cell between second and thirdbranches of medial vein with a pale brown spot medially. Wing with a large dark brown spot adjacent toclaval fold at midlength; cells of claval area and first cubital cell each with a dark brown spot on posteriormargin. Shaft of aedeagus broadly laterally expanded subapically; dorsal surface subapically with a pair ofapically acute flap-like processes. Paramere broadest at midlength, apex narrowly rounded; dorsal processslightly distad of midlength, produced posteriorly; dorsal surface subbasally with a broad secondaryprocess bearing numerous, large, robust spines. MATERIAL EXAMINED Holotype cT, Brazil: Nictheroy, iv.1924 (Williams) (BMNH).Paratypes. Brazil: 1 cf , Nictheroy; 1 cf , Rezende (BMNH). This species is distinguished by the intricate pattern of small dark spots on the tegmen and wing,and by the structure of the male genitalia. Mysidia acidalioides Fowler (Figs 7, 15,33,216,326,435)Mysidia acidalioides Fowler, 1900: 72. LECTOTYPE $ , PANAMA (BMNH), here designated [examined]. Male: head 0-80 mm long, 1-10 mm wide; pronotum 2-30 mm wide; tegmen 11-00 mm long; wing 7-30 mmlong. Female: tegmen 11-10-12-60 mm long. 30 PETER S. BROOMFIELD Length of frons c. 4-5 times width at apex, c. twice width at base; ocelli obsolete; clypeus c. as long asfrons; rostrum terminating at level of anterior surface of hind coxae. Pronotal width c. 16 times mid-dorsallength, fronto-lateral carinae very prominent; tegulae distinctly carinate. Vertex dark brown between carinae; frons with a broad, often broken, dark brown transverse band atlevel of midline of eyes; antenna often reddish; paraclypeus and lateral surfaces of anteclypeus red;fronto-lateral surfaces of pronotum each with a broad, deep red, horizontal band extending from level ofmidline of adjacent eye to just above ventral margin; fore and mid coxae bright red over apical half; hindfemur reddish apically. Tegmen and wing whitish hyaline, veins pale yellow. Tegmen with a small brownspot on costal cell at level of point of separation of fused subcostal and radial veins; clavus with a large,irregular, brown spot at apex, and a smaller spot on exterior margin adjacent to point of fusion of analveins; posterior and apical margins narrowly and weakly brownish, somewhat darker between apicalbranches of medial and radial veins. Wing with a small, somewhat indistinct, brown marking betweenclaval suture and midlength of first branch of cubital vein; posterior margin with a dark brown spot betweeneach branch of anal, cubital and medial veins. Shaft of aedeagus slender, cylindrical ; dorsal surface subapically with a pair of triangular processes, eachproduced posteriorly into a slender, rounded, flap extending beyond apex of shaft; ventral surface apicallyproduced into two pairs of large, acute, triangular processes. Paramere slender, broadest at three-fifthslength, apex acutely rounded; dorsal process robust, situated at three-fifths length. MATERIAL EXAMINED Lectotype C?, Panama: V. de Chiriqui, 2500-4000 ft (Champion) (BMNH). Panama: 2 $ (paralecto-types), same data as lectotype (BMNH); 5 cf , 6 $ (USNM; FAMU; CAS). Belize: 1 $ (FAMU). The species here designated as lectotype bears Fowler's handwritten determination label and the BMNH'type' label. This species is readily distinguished by the very slight pigmentation of the tegmen and wingcombined with the distinctive markings of the head and thorax; from athena it is separated by itslarger size and the structure of the male genitalia. Mysidia neoasinella sp. n. (Figs 257, 368, 477) Male: head 0-70 mm long, 1-02 mm wide; pronotum 2-10 mm wide; tegmen 9-70 mm long; wing 6-00 mmlong. Female unknown. Length of frons 6 times width at apex, c. three times width at base; ocelli very prominent; clypeus slightlylonger than frons; rostrum terminating somewhat posterior to hind coxae. Pronotal width 12-5 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Ocelli crimson; dorsal and lateral surfaces of pronotum irregularly pale crimson; tegulae tinged orange,margins narrowly brownish; disc of mesonotum pale yellowish brown, darker between lateral carinae.Tegmen and wing dark smoky brown. Tegmen with veins narrowly edged hyaline from base to level of apexof claval area; with a very narrow and indistinct, hyaline, transverse band extending from costal margin atlevel of second fork of cubital vein to posterior margin at apex of clavus. Wing with costal cell narrowlyhyaline basally; with a very narrow and indistinct, hyaline, transverse band at approximately two-thirdslength extending from costal to posterior margins. Shaft of aedeagus slender, cylindrical, slightly expanded laterally over apical third; lateral surfaces eachwith a large flap-like process produced antero-dorsally into a long, slightly curving spine. Paramere short,rounded; apex acute; dorsal process situated slightly distad of mid-length, strongly produced posteriorly;dorsal surface subbasally roundly produced, bearing numerous, short, robust, spines. Lateral margins ofsubgenital segment each produced into a single, long, broad, apically rounded, posteriorly directed lobe atmidline. MATERIAL EXAMINEDHolotype cf, Brazil: Amazonas, P. das Laranjeiras, viii.-ix.1981 (Arias) (INPA). This species is distinguished by the pigmentation of the thorax and the structure of the malegenitalia. Mysidia vista sp. n. Female: head 0-63 mm long, 0-71 mm wide; pronotum 1-25 mm wide; tegmen 6-88-7-68 mm long; wing3-85 mm long. Male unknown. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 31 Length of frons 6-5 times width at apex, 3 times width at base; ocelli very large, prominent; clypeus aslong as frons; rostrum extending almost to base of subgenital plate. Pronotal width 15 times mid-dorsallength; fronto-lateral carinae weak; tegula not carinate. Genae anterior and dorsal to eyes orange. Pronotum with dorso-lateral margins and a large, circular spotadjacent to eye on each fronto-lateral surface orange; scutellum with baso-lateral angles tinged orange;abdomen posteriorly orange on mid-dorsal line. Tegmen and wing smoky brown, veins pale brown.Tegmen with a narrow, transverse, whitish band extending from costal margin to apex of clavus atone-third length, another, fainter band at one-fifth length; brown pigmentation gradually becoming fainterfrom base to apex. Wing with brown pigmentation weakening from base; posterior and apical margins clearwhitish hyaline. MATERIAL EXAMINED Holotype $, Guyana: Blairmont, x.1923 (Williams) (BMNH).Paratypes. Guyana: 1 $, Blairmont; 1 $, New Amsterdam (BMNH). In the absence of males this species is most readily distinguished by the pigmentation of thethorax, tegmen and wing. Mysidia albifasciata sp. n. (Figs 239, 350, 459) Male: head 0-59 mm long; 0-71 mm wide; pronotum 1-28 mm wide; tegmen 7-00 mm long; wing 3-70 mmlong. Female: tegmen 7-40 mm long. Length of frons 6 times width at apex, 3-5 times width at base; ocelli large, prominent; clypeus as long asfrons; rostrum terminating immediately basad of subgenital plate. Pronotal width 15 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Ocelli occasionally deep red; pronotum with fronto-lateral surfaces tinged red adjacent to eyes; disc ofmesonotum occasionally blackish brown. Tegmen smoky brown; veins pale brownish yellow, irregularlymottled white over basal quarter; a transverse, hyaline band extending from costal to posterior margins atone-third length, another at midlength, and a third at three-quarters; apex, beyond apical fork of medialvein, irregularly hyaline. Wing whitish hyaline with a pale, smoky brown, transverse band at mid-length;apical third pale brownish. Shaft of aedeagus slender, cylindrical; dorsal surface subapically with a pair of broad, apically acuteprocesses adjacent to midline bearing small obtuse spines; laterally, a pair of slender, spine-like processes.Paramere slender, apex narrowly rounded; dorsal process situated at two-thirds length, strongly produceddorsally and posteriorly; dorsal surface subbasally with a broad, rounded secondary process. MATERIAL EXAMINED Holotype cf , Ecuador: Mera, l-2.ii.1923 (Williams) (BMNH).Paratype. Ecuador: 1 $, Tena (BMNH). This species is readily distinguished by the pigmentation of the tegmen and wing, and by thestructure of the male genitalia. Mysidia quadrifascia WalkerMysidia quadrifascia Walker, 1858: 97. Holotype 9, BRAZIL (BMNH) [examined]. Female : head 66 mm long ,0-67 mm wide ; pronotum 1 55 mm wide ; tegmen 7 65 mm long , wing 4 25 mmlong. Male unknown. Length of frons c. 5 times width at apex, c. 3 times width at base; ocelli very prominent; clypeus slightlylonger than frons; rostrum terminating level with midlength of abdomen. Pronotal width c. 11 timesmid-dorsal length; fronto-lateral surfaces and tegula without distinct carinae. Ocelli very dark crimson; disc of mesonotum brown. Tegmen and wing hyaline, veins and cross-veinspale brownish yellow. Tegmen pale smoky brown basally, with a smoky brown transverse band at level offirst fork of cubital vein , a broader band at level of first and second forks of medial vein , and another at levelof radial-medial cross-vein; apical margin very pale smoky brown. Wing with a very pale, irregular, smokybrown, transverse band at midlength; apical third pale smoky brown. MATERIAL EXAMINEDHolotype $, Brazil: Santarem (Bates) (BMNH). This species is distinguished by the prominent dark pigmentation of the tegmen and wing. 32 PETER S. BROOMFIELD Mysidia transversa sp. n. (Figs 234, 345, 454) Male: head 0-62 mm long, 0-78 mm wide; pronotum 1-44 mm wide; tegmen 7-20 mm long; wing 3-85 mmlong. Female unknown. Length of frons 5 times width at apex, 3-33 times width at base; ocelli prominent; clypeus slightly longerthan frons; rostrum terminating immediately basad of apex of subgenital plate. Pronotal width c. 14 timesmid-dorsal length, fronto-lateral carinae absent; tegula with very weak carinae. Ocelli scarlet; fronto-lateral surfaces of pronotum occasionally each with a scarlet spot adjacent to eye;disc of mesonotum and dorsal surface of abdomen dark brown. Tegmen and wing whitish hyaline, veinspale brownish. Tegmen with a broad, smoky brown, transverse band at one-eighth length, another at levelof first fork of cubital vein, another, broader, band immediately distad of medial-cubital cross-vein, afourth at approximately two-thirds length; apical area, distad of last fork of radial vein, broadly smokybrown. Wing smoky brown basally; with a broad smoky brown band extending obliquely from costalmargin to posterior margin at midlength; apical quarter smoky brown. Shaft of aedeagus cylindrical; dorsal surface subapically with two pairs of robust, anteriorly directed,spine-like processes. Paramere very slender basally, becoming abruptly expanded at midlength, apexobtusely rounded; dorsal process situated immediately distad of midlength, long, slender, apex stronglyproduced. MATERIAL EXAMINED Holotype cf , Peru: Iquitos, Rio Chinchicuy 1-5 km, 27.xi.1972 (Waldo) (FAMU).Paratypes. Peru: 1 cf (BMNH). Brazil: 3 cf , 1 $, Amazonas (INPA; BMNH). Superficially this species resembles quadrifascia, but differs in the proportions of the head andpronotum and in the pigmentation of the tegmen. Mysidia fulvodorsalissp. n. (Figs 177, 286, 395) Male: head 0-63 mm long, 0-74 mm wide; pronotum 1-51 mm wide; tegmen 7-90-8-20 mm long; wing 4-25mm long. Female: tegmen 8-00-8-40 mm long. Length of frons c. 6 times width at apex, 3-33 times width at base; ocelli prominent; clypeus slightlylonger than frons; rostrum extending to base of subgenital segment. Pronotal width 12 times mid-dorsallength, fronto-lateral carinae absent, tegula weakly carinate. Ocelli crimson ; disc of mesonotum brown . Tegmen and wing whitish hyaline . Tegmen with a broad , darksmoky brown, transverse band near base, another at level of first fork of cubital vein, another at midlength,a fourth at level of radial-medial cross-vein; apical area, distad of last fork of radial vein, dark smokybrown. Wing with a broad, transverse, smoky brown band extending from medial vein to posterior marginat midlength; apical third dark smoky brown. Shaft of aedeagus somewhat laterally expanded subapically; dorsal surface subapically with a pair oflong, spine-like processes laterally; a pair of long, slender, processes at midline. Paramere basally slender,becoming broadly expanded over distal half length, apex obtusely rounded; dorsal process situated atthree-fifths length, apex slender; dorsal surface at one-fifth length with a small, conical, secondary processbearing numerous small spines; ventral surface at midlength with a rounded process bearing numerouslong spines. MATERIAL EXAMINED Holotype cf , Bolivia: Cbb., Villa Tunari, 31.iii.1978 (O'Brien) (FAMU).Paratypes. 1 cf , 3 $ , same data as holotype (FAMU; BMNH). This species is distinguished by the pronotal and tegminal pigmentation, and by the structure ofthe male genitalia. Mysidia musics sp. n. (Figs 253, 364, 473) Male: head 0-67 mm long, 0-95 mm wide; pronotum 1-80 mm wide; tegmen 8-40 mm long; wing 5-20 mmlong. Female: tegmen 8-80 mm long.Length of frons 5 times width at apex, 2-5 times width at base; ocelli large, prominent; clypeus c. as long TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 33 as frons; rostrum extending well beyond hind coxae. Pronotal width 20 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Genae and fronto-lateral surfaces of pronotum adjacent to eyes weakly pale orange; disc of mesonotumbrownish. Tegmen and wing smoky brown. Tegmen with veins brownish; with a narrow, whitish,transverse band at one-fifth length, another at two-fifths; a third at three-fifths; an irregular, broken, bandat approximately mid-length; posterior margin whitish from apex of first branch of cubital vein to fifthbranch of medial vein; apical fork of radial vein irregularly pale. Wing pale, whitish hyaline, veins pale;with an irregular brownish, transverse band extending from costal margin to base of cubital vein; anirregular brownish band extending obliquely from medial-cubital cross-vein to posterior margin; a veryfaint band extending from radial-medial cross-vein almost to posterior margin. Shaft of aedeagus broadly laterally expanded subapically ; dorsal surface subapically with a pair of large,adpressed, overlapping, flap-like processes, each terminating anteriorly in a tapering spine -near midline.Paramere massive, apex broadly rounded; dorsal process greatly reduced, situated at one-third length;dorsal surface at two-thirds length with a large, conical, secondary process. MATERIAL EXAMINED Holotype cT, Brazil: Para, Jabaty, v.1924 (Williams) (BMNH).Paratype. 1 $ , same data as holotype (BMNH). This species is distinguished by the pigmentation of the tegmen and wing. Mysidia williamsisp. n. (Figs 238, 349, 458) Male: head 0-52 mm long, 0-76 mm wide; pronotum 1-34 mm wide; tegmen 6-46 mm long; wing 3-60 mmlong. Female: tegmen 6-40-7-00 mm long. Length of frons 5-5 times width at apex, 3 times width at base; ocelli very large and prominent; clypeusslightly longer than frons; rostrum terminating slightly basad of subgenital plate. Pronotal width c. 14 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Dorsal surface of mesonotum and abdomen brownish. Tegmen and wing dark, smoky brown, veinsbrownish yellow. Tegmen with a narrow, whitish, transverse band extending from costal margin to clavalsuture at one-sixth length, another extending from costal margin to apex of clavus at one-third, anotherextending from costa to hind margin somewhat distad of mid-length; an irregular whitish area aroundapical fork of medial vein extending broadly to costal margin; apical area between first and fifth branches ofmedial vein hyaline. Wing with an irregular pale band running transversely from costal margin at level ofradial-medial cross-vein, becoming broader and less distinct towards posterior margin at level of first twobranches of cubital vein. Shaft of aedeagus slender, cylindrical, slightly asymmetrical; dorsal surface subapically with a pair oflong, flap-like processes bearing clusters of very small, tooth-like spines apically; dorso-lateral surfacessubapically each with a small, spine-like process. Paramere slender; dorsal process large, situated attwo-thirds length, strongly produced dorsally and posteriorly; dorsal surface subbasally with a cluster ofshort, robust spines; ventral surface subbasally with several long, robust spines. MATERIAL EXAMINED Holotype $, Brazil: Para, Jabaty, v.1924 (Williams) (BMNH).Paratypes. Brazil: 1 cT, 4 $, Amazonas (INPA; BMNH). This species is readily distinguished by the pigmentation of the tegmen and wing, and by thestructure of the male genitalia. Mysidia mylesisp. n. (Figs 276, 387, 497) Male: head 0-52 mm long, 0-69 mm wide; pronotum 1-20 mm wide; tegmen 5-60 mm long; wing 3-20 mmlong. Female: tegmen 6-40 mm long. Length of frons 5-5 times width at apex, 3 times width at base; ocelli large, prominent; clypeus c. as longas frons; rostrum extending to base of subgenital segment. Pronotal width c. 12 times mid-dorsal length,fronto-lateral carinae absent; tegula with weak carinae. Ocelli bright scarlet; fronto-lateral surfaces of pronotum adjacent to eyes pale reddish; abdomen withdorsal surface tinged reddish. Tegmen and wing whitish hyaline, veins pale brown. Tegmen with basal and 34 PETER S. BROOMFIELD claval areas smoky brown, a smoky brown transverse band at level of first fork of cubital vein, another atmidlength; apical third smoky brown, with anterior margin apically, and posterior margin between first andfifth branches of medial vein broadly pale. Wing pale smoky brown over basal half and apical third. Shaft of aedeagus slender, cylindrical, slightly asymmetrical; dorsal surface subapically with two pairs ofacutely pointed processes. Frame re slender; dorsal process situated slightly distad of midlength, apexstrongly produced posteriorly. MATERIAL EXAMINED Holotype cf , Trinidad: Caura, on Parthenium sp., 2.viii.l976 (Noyes) (BMNH).Paratype. 1 $, same data as holotype (BMNH). This species is readily distinguished by the pigmentation of the legmen and wing, and by therelatively simple structure of the aedeagus. Mysidiu varia sp. n. (Figs 261, 372, 482) Male: head 0-65 mm long, 1-05 mm wide; pronotum 2-39 mm wide; tegmen 9-40-10-54 mm long; wing6-00 mm long. Female: tegmen 10-20-11-40 mm long. Length of frons 5 times width at apex, 3-33 times width at base; ocelli obsolete; clypeus slightly longerthan frons; rostrum extending beyond apex of subgenital plate. Pronotal width 11 times mid-dorsal length,fronto-lateral surfaces weakly carinate; tegula not carinate. Head and pronotum often tinged pale orange. Tegmen smoky brown; veins dark brown, narrowlyboardered whitish hyaline; central areas of cubital cells and larger medial cells irregularly whitish hyaline.Wing predominantly whitish hyaline, veins dark brown, cells irregularly smoky brown medially. Shaft of aedeagus cylindrical, somewhat expanded apically; dorsal surface subapically with a pair offlap-like processes, each produced into two acute, spine-like processes dorsally. Paramere very long andslender; apex narrowly rounded; dorsal process situated somewhat distad of two-thirds length, littleproduced posteriorly; dorsal surface at one-fifth length with an obtusely rounded, secondary process. MATERIAL EXAMINED Holotype cf , Colombia: Caqueta, Yuruyacu, 70 km SW. Florencia, 22.i.l979 (Cooper) (BMNH). Paratypes. Colombia: 1 cf , same data as holotype (BMNH). Guyana: 1 cf , Essequibo River (BMNH).Ecuador: 8 cf , 6 $, Tena (BMNH). Brazil: 1 cf , Amazonas (INPA). Though closely related to tikalme, varia may be distinguished readily by the pigmentation of thetegmen and wing, and by the detailed structure of the male genitalia. Mysidiu tikalme sp. n. (Figs 259, 370, 480) Male: head 0-63 mm long, 0-99 mm wide; pronotum 2-30 mm wide; tegmen 11.00 mm long; wing 6-30 mmlong. Female: tegmen 11-00-11-25 mm long. Length of frons c. 6 times width at apex, 4-5 times width at base; ocelli obsolete; clypeus as long as frons;rostrum extending beyond base of genital segment. Pronotal width c. 12 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Genae occasionally tinged crimson; fronto-lateral surfaces of pronotum and disc and ventral surfaces ofmesonotum frequently tinged deep crimson. Tegmen and wing hyaline, veins dark brown, central areas ofcells broadly dark brown, with a narrow hyaline margin adjacent to veins. Shaft of aedeagus simple; dorsal surface subapically with a pair of flap-like processes, each bearing twoacute spines dorsally. Paramere slender; apex narrowly rounded; dorsal process situated at two-thirdslength, small, little produced posteriorly; dorsal surface at one-fifth length with a slender secondaryprocess bearing a row of long, robust spines. MATERIAL EXAMINED Holotype cf, Guyana: confluence of Oronoque and New rivers, 650 ft, ix-xii.1937 (Rosenberg)(BMNH). Paratypes. Brazil: 1 $, Para; 1 $, Amazonas (BMNH). This species is distinguished by the pigmentation of the tegmen and wing, and by the structure ofthe male genitalia. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 35 Mysidia glauca Distant (Figs 188, 298, 407)Mysidia glauca Distant, 1907: 397. LECTOTYPE cf , BRAZIL (BMNH), here designated [examined]. Male: head 0-55 mm long, 0-80 mm wide; pronotum 1-60 mm wide; legmen 7-10 mm long; wing 4-20 mmlong. Female unknown. Length of frons c. 5 times width at apex, c. 3 times width at base; ocelli obsolete; clypeus slightly longerthan frons; rostrum terminating slightly posterior to hind coxae. Pronotal width 13 times mid-dorsal length,fronto-lateral carinae distinct; tegula not carinate. Central area of frons, and genae dorsal to eyes and anterior to antennae, pale crimson; pronotumdorsally, around carinae on fronto-lateral surfaces, and anterior to fore coxae, pale crimson; mesonotumwith lateral surfaces and coxae pale pinkish; posterior abdominal segments brown. Tegmen and wing palesmoky brown, unmarked, veins dark brown. Tegmen with costal vein narrowly crimson. Shaft of aedeagus laterally expanded subapically; dorsal surface subapically with a pair of parallelflap-like processes; a pair of dorso-lateral flaps, each terminating anteriorly in an obtuse point. Paramereslender; apex broadly rounded; dorsal process situated at two-thirds length, rounded, not posteriorlyproduced, bearing a single, blunt spine; dorsal surface at one-third length with a low, rounded, secondaryprocess bearing three robust spines. MATERIAL EXAMINEDLectotype cf , Brazil: Parana de Buyassu, Lower Amazon, 18. i. 1896 (Austin) (BMNH). This species is distinguished by the pigmentation of the head and body, the absence of markingson the wing and tegmen, and by the unique structure of the paramere. Mysidiagradlissp. n. (Figs 244, 355, 465) Male: head 0-53 mm long, 0-76 mm wide; pronotum 1-53 mm wide; tegmen 7-65-7-90 mm long; wing4-80 mm long. Female: tegmen 8-50 mm long. Length of frons 5-5 times width at apex, c. twice width at base; ocelli small, distinct; clypeus slightlylonger than frons; rostrum extending slightly beyond hind coxae. Pronotal width c. 15 times mid-dorsallength, fronto-lateral carinae absent; tegula distinctly carinate. Head and body tinged pale crimson; genae adjacent to eyes broadly crimson; pronotum dorsally, andadjacent to eyes on fronto-lateral surfaces, crimson. Tegmen and wing whitish hyaline; veins dark brown,edged pale smoky hyaline; cells distant from veins smoky brownish, pale hyaline medially. Wing with avery irregular, indistinct, pale brownish, transverse band at midlength; posterior and apical marginsbetween veins broadly smoky brown. Shaft of aedeagus expanded subapically; lateral surfaces each with a slender spine-like processsubapically; dorsal surface subapically with a pair of large flap-like processes. Paramere slender, parallel-sided; apex rounded; dorsal process situated slightly distad of midlength, strongly produced posteriorly;dorsal surface basally with a large, flap-like secondary process bearing numerous, large, robust spines. MATERIAL EXAMINED Holotype cf , Brazil: Rio de Janeiro, i.1924 (Williams) (BMNH).Paratypes. Brazil: 2 cf , 2 9, same data as holotype; Rezende (BMNH). This species is distinguished by the pigmentation of the head, pronotum, tegmen and wing, andby the structure of the male genitalia. Mysidia lucifera sp. n. (Figs 232, 343, 452) Male: head 0-56 mm long, 0-73 mm wide; pronotum 1-43 mm wide; tegmen 7-65 mm long; wing 4-25 mmlong. Female: tegmen 8-50 mm long. Length of frons 7 times width at apex, c. 2-5 times width at base; ocelli indistinct; clypeus slightly longerthan frons; rostrum extending to midlength of abdomen. Pronotal width c. 14 times mid-dorsal length;fronto-lateral surfaces and tegula lacking distinct carinae. Head and body tinged pale crimson; female with abdomen tinged crimson. Tegmen and wing whitish 36 PETER S. BROOMFIELD hyaline, weakly and irregularly mottled pale smoky brown, veins and cross-veins brown. Tegmen with avery faint, pale brown, transverse band at midlength. Shaft of aedeagus somewhat expanded over apical third; dorsal surface subapically with a pair of broad,apically bifid processes laterally; lateral surfaces each with a long, spine-like process subapically. Paramereslender; apex actutely rounded; dorsal process situated at two-thirds length, large, strongly producedposteriorly; dorsal surface at one-quarter length with a prominent, rounded, secondary process bearingnumerous, long, robust spines. MATERIAL EXAMINED Holotype cf , Brazil: Rezende, Estado de Rio, ii.1924 (Williams) (BMNH).Paratype. 1 $, same data as holotype (BMNH). This species is only readily distinguished by reference to the structure of the male genitalia. Mysidia havilandisp. n. (Figs 218, 329, 438) Male: head 0-59 mm long, 0-80 mm wide; pronotum 1-55 mm wide; tegmen 6-80-7-44 mm long; wing4-40 mm long. Female: tegmen 8-24 mm long. Length of f rons 5 5 times width at apex ,2-25 times width at base ; ocelli large , prominent ; clypeus slightlylonger than frons; rostrum extending well beyond hind coxae. Pronotal width 15 times mid-dorsal length;fronto-lateral carinae prominent; tegula not carinate. Head and body predominantly brown; base of vertex often crimson; base of antenna narrowly red;pronotum reddish dorsally, occasionally on fronto-lateral surfaces also; disc of mesonotum brown;scutellum deep red or reddish brown, adjacent surfaces of metanotum dark brown; abdomen deep red toblackish brown. Tegmen and wing pale brownish hyaline, narrowly crimson basally, apical and posteriormargins very narrowly crimson; veins reddish brown, very broadly edged smoky brown. Tegmen withcostal vein and apices of subcostal and radial veins crimson, an indistinct, brown, transverse bandextending from costal margin to first fork of cubital vein. Wing with posterior margins of cells narrowlysmoky brown; first anal vein crimson. Shaft of aedeagus slender, cylindrical, dorso-laterally expanded into a pair of large, curving, spine-likeprocesses; apex with a pair of small curved processes. Paramere slender; dorsal process situated somewhatdistad of midlength, robust; dorsal surface somewhat expanded at one-third length, bearing long robustspines. MATERIAL EXAMINED Holotype cf , Guyana: Tumatumari, 19.vii.1923 (Williams) (BMNH). Paratypes. Guyana: 4 cf, 1 $, same data as holotype (BMNH). Brazil: 3 cf, 5 $, Amazonas (INPA;NR). This species is distinguished by the dark pigmentation of the body, tegmen and wing, and by thestructure of the male genitalia. Mysidia etheldreda sp. n. (Figs 254, 365, 475) Male: head 0-65 mm long, 1-00 mm wide; pronotum 2-14 mm wide; tegmen 9-60 mm long; wing 5-44 mmlong. Female unknown. Length of frons c. 1 times width at apex, 2-5 times width at base; ocelli small, distinct; clypeus c.one-third longer than frons; rostrum extending to base of subgenital plate. Pronotal width c. 24 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Disc of mesonotum dark brown, scutellum blackish brown. Tegmen and wing predominantly whitishhyaline, irregularly marked smoky brown; veins yellow; apical margins between branches of medial veinsvery broadly smoky brown. Tegmen pale smoky brown over basal quarter; medial and cubital areas palebrown from approximately one-third length to midlength; costal and radial areas brownish subapically.Wing narrowly brownish basally; a narrow and irregular, obliquely curving, smoky brown, transverse bandimmediately basad of first fork of cubital vein; second fork of cubital vein and cubital-medial cross-veinedged smoky brown; first branch of cubital vein subapically broadly edged smoky brown. Shaft of aedeagus robust; apex strongly produced, curving antero-dorsad; lateral surfaces subapicallyeach with a large flap-like process extending over dorsal surface, each process with a pair of anteriorly TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 37 directed spines at internal angle; dorso-lateral surfaces each with a triangular flap-like process atmidlength; dorsal surface subbasally with a broad transverse process. Paramere broad; apex obtuselyrounded; dorsal process situated at midlength, small, weakly produced posteriorly. Subgenital lateralsternites each with posterior margin produced into a very long, slender, parallel-sided, horizontallydirected lobe. MATERIAL EXAMINEDHolotype cf , Brazil: Amazonas, P. das Laranjeiras, vii.-viii.1981 (Arias) (INPA). This species, though closely related to estfarchina and molesta, is readily distinguished by thepigmentation of the tegmen and wing. Mysidia liquida sp. n. (Figs 224, 335, 443) Male: head 0-65 mm long, 0-82 mm wide; pronotum 1-60 mm wide; tegmen 9-35-10-20 mm long; wing5-44 mm long. Female: tegmen 11-05-13-60 mm long. Length of frons c. 6 times width at apex, c. 3 times width at base; ocelli distinct; clypeus one-sixth longerthan frons; rostrum extending to base of subgenital plate. Pronotal width 40 times mid-dorsal length;fronto-lateral carinae indistinct; tegulae distinctly carinate. Genae anterior to eyes tinged reddish orange; fronto-lateral surfaces of pronotum each with a large,dark brown irregularly circular spot at level of eye and adjacent to lateral margin; tegula dark brownmedially; disc of mesonotum with a pair of large, prominent, roughly circular, dark brown spotsposteriorly. Tegmen and wing pale, whitish hyaline, lacking prominent markings; veins pale; cross-veinspale brownish, narrowly margined smoky brown. Tegmen with costal area tinged brownish yellow,narrowly dark brown basally. Wing unmarked. Shaft of aedeagus greatly expanded subapically; dorso-lateral surfaces each with a very large, flap-likeprocess extending over dorsal surface, lacking spine-like secondary processes. Paramere slender; apexacute; dorsal process strongly produced posteriorly. MATERIAL EXAMINED Holotype cf , Bolivia: Bueyas [?] (BMNH). Paratypes. Bolivia: 2 cf, 1 $, same data as holotype; Buena Vista (BMNH; FAMU). Venezuela: 1 cf(NR). Peru: 1 ?, Callanga (BMNH). This species is distinguished by the dark spots on the fronto-lateral surfaces of the pronotum, thealmost complete absence of pigmentation on the tegmen and wing, and by the massive aedeagus. Mysidia ariasisp. n. (Figs 185, 295, 404) Male: head 0-63 mm long, 1-08 mm wide; pronotum 2-06 mm wide; tegmen 9-40 mm long; wing 5-60 mmlong. Female unknown. Length of frons c. 6 times width at apex, 2-5 times width at base; ocelli obsolete; clypeus c. as long asfrons; rostrum extending to base of sugenital plate. Pronotal width c. 24 times mid-dorsal length;fronto-lateral carinae distinct; tegula not carinate. Fronto-lateral surfaces of pronotum at level of eyes each with a prominent, dark brown, roughly circularspot, not reaching either internal or lateral margins; tegula tinged brownish on ventral margins; fore tibiaand tarsus brownish. Tegmen and wing hyaline, without distinct markings. Tegmen with costal area tingedyellowish brown; apical and posterior margins weakly smoky brown; cross-veins brownish; apex of analvein with a small, triangular, brownish spot. Wing with posterior and apical margins indistinctly smokybrown; radial-medial cross-vein narrowly edged smoky brown. Shaft of aedeagus broad; dorso-lateral surfaces subapically each with a large flap- like process extendingover ventral surfaces, dorsally produced into a rounded lobe bearing a small conical projection. Paramereslender; apex acutely rounded; dorsal process situated slightly basad of mid-length, large, apex stronglyproduced posteriorly; dorsal surface distad of process somewhat produced. MATERIAL EXAMINEDHolotype cf , Brazil: Amazonas, P. das Laranjeiras, viii.-ix.1981 (Arias) (INPA). 38 PETER S. BROOMFIELD This species is distinguished by the prominent dark spots on the fronto-lateral surfaces of thepronotum and the lack of distinct markings on the tegmen and wing; from liquida it is separatedby the structure of the male genitalia. Mysidia fuscofrontalissp. n. (Figs 245, 356, 466) Male: head 0-67 mm long, 0-88 mm wide; pronotum 2-37 mm wide; tegmen 8-90 mm long; wing 5-10 mmlong. Female unknown. Length of frons c. 4 times width at apex, 2-25 times width at base; ocelli large, not prominent; clypeusslightly longer than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 28times mid-dorsal length, fronto-lateral carinae absent; tegula with carinae very prominent. Head with base of vertex, and genae from base to level of ventral margins of eyes, bright scarlet;fronto-lateral surfaces of pronotum each with a narrow, bright scarlet, horizontal band extending fromadjacent to base of head to lateral margin. Tegmen and wing whitish hyaline; veins, excepting subcostaland radial veins of tegmen, pale; cross-veins dark brown; posterior and apical margins broadly smokybrown between veins. Tegmen with subcostal and radial veins brown over greater part of length basally;clavus with a small, dark brown spot between anal veins, another slightly distad of point of fusion of analveins, another subapically; cubital area with four, irregular brownish spots; costal cell dark brown betweencross-veins, costal margin narrowly scarlet; medial vein with bases of first and second branches, and apicalfork, dark brown, fifth and sixth branches each with two, evenly spaced, dark brown spots. Wing with firstfork of cubital vein dark brown; with an irregular, dark brown spot over midlength of first cubital branch;second and third cubital branches, and both medial branches, dark brown from base to immediately priorto posterior margin. Shaft of aedeagus slender; lateral surfaces subapically each with a large flap-like process, producedantero-dorsally into a long curving spine, and with a small spine ventrally. Paramere robust; apex broadlyrounded, with a long, curving, flap-like process dorsally; dorsal process small, situated at midlength,posteriorly produced. MATERIAL EXAMINEDHolotype cf , Panama: Las Cumbres, 6.vi.l976 (Woldo) (FAMU). This species is readily distinguished by the pigmentation of the head, pronotum, tegmen andwing. Mysidia albipennis Westwood(Figs 228, 339, 448) Mysidia albipennis Westwood, 1840: 83. LECTOTYPE cf , BRAZIL (UM), here designated [examined].Mysidia parviceps Fowler, 1900: 73. LECTOTYPE $, GUATEMALA (BMNH), here designated [ex-amined]. Syn. n. Male: head 0-63 mm long, 0-80 mm wide; pronotum 1-82 mm wide; tegmen 8-40-9-00 mm long; wing5-60 mm long. Female: tegmen 8-84-9-20 mm long. Length of frons c. 6 times width at apex, c. twice width at base; ocelli distinct; clypeus c. as long as frons;rostrum terminating at level of hind coxae. Pronotal width c. 14 times mid-dorsal length; fronto-lateralsurfaces and tegula not distinctly carinate. Vertex often scarlet from base to level of dorsal margins of eyes; genae each with a narrow, horizontal,dark brown/black band extending from adjacent to dorsal margin of eye to anterior margin; fronto-lateralsurfaces of pronotum each with a broad reddish band extending horizontally from adjacent to eye to lateralmargin, this band often incorporating a large black spot in its ventral margin; abdomen with a small,circular, black spot on either side of midline on dorsal surface of fifth segment. Tegmen and wing whitishhyaline, veins very pale brownish. Tegmen with cross-veins and first and second forks of medial vein darkbrown; cells between branches of cubital and medial veins each with a smoky brown semicircular spot onposterior and apical margins; apical fork of medial vein covered by a prominent, circular, dark brown spot;clavus with an irregular dark brown spot between anal veins subbasally, another between fused anal veinsand claval suture at two-thirds length, another between fused anal veins and posterior margin, and a fourthat apex of claval suture. Wing with apical and posterior margins narrowly smoky brown between veins;radial-medial cross-vein dark brown, an irregular, dark brown spot between medial and cubital veins attwo-fifths length, and another between first branch of cubital vein and first anal vein. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 39 Shaft of aedeagus slender, laterally expanded over apical half; dorsal surface with a pair of large flap-likeprocesses arising subapically and extending anteriorly to midlength. Paramere with apex very obtuselyrounded; dorsal process situated at two-fifths length, small, with apex slender and strongly producedposteriorly; dorsal surface subbasally with a group of short robust spines. MATERIAL EXAMINED Lectotype C? (albipennis), Brazil: Vera Cruz (UM). Lectotype 9 (parviceps), Guatemala: Zapota(Champion) (BMNH). Honduras: 6 cf , 8 $ , Lancertillo (FAMU; BMNH). Belize: 1 $ (FAMU). Westwood did not indicate the number of specimens in the type-series of albipennis; the singlemale available for study is here designated as lectotype. The three female specimens of parvicepsdescribed by Fowler are not conspecific. The specimen here designated as lectotype has thetegmina damaged; it bears Fowler's handwritten 'type' label. This species is readily distinguished by the distinctive pigmentation of the head and pronotum,the mottled appearance of the tegmen and wing, and by the structure of the male genitalia. Mysidia lactittora WestwoodMysidia lactiflora Westwood, 1840: 83. LECTOTYPE $ , BRAZIL (UM), here designated [examined]. Female: head 0-75 mm long, 0-98 mm wide; pronotum 2-63 wide; tegmen 12-07 mm long; wing 7-00 mmlong. Male unknown. Length of frons 10 times width at apex, 2-5 times width at base; ocelli small, distinct; clypeus one quarterlonger than frons; rostrum only just reaching hind coxae. Pronotal width c. 50 times mid-dorsal length,fronto-lateral carinae absent; tegula prominently carinate. Base of vertex reddish; genae from level of dorsal margins of eyes to level of midline of eyes dull crimson;fronto-lateral surfaces of pronotum each with a prominent, broad, brownish band extending horizontallyfrom adjacent to eye to lateral margin, each band deep red along dorsal margin; tegulae ventral to carinaedeep brownish. Tegmen and wing almost hyaline, veins pale yellow, cross-veins brownish. Tegmen withfirst, second and apical forks of medial vein pale brownish; costal cell yellowish brown, bearing a small,prominent, roughly circular, dark brown spot at level of first fork of cubital vein, another similar spotadjacent to point of separation of fused subcostal and radial veins, a third, smaller spot at c. one-thirdlength; with a small, roughly circular, prominent, dark brown spot over cross-vein linking second and thirdbranches of medial vein; a small dark brown spot on anal margin somewhat distad of point of fusion of analveins. Wing with cross- veins slightly darker brown than those of tegmen; a small, irregular, indistinct,brownish spot over first branch of cubital vein somewhat basad of midlength. MATERIAL EXAMINEDLectotype $ , Brazil: no further data (UM). The lectotype has the abdomen partially eaten away, and the left tegmen and wing glued inplace; Westwood's 'type' label gives the name as 'lactiflorea'. This species, in the absence of reference to the male genitalia, is most readily distinguished byits large size, the markings on the fronto-lateral surfaces of the pronotum, and the paucity ofmarkings on the tegmina and wings, in particular the absence of a dark spot over the apical forkof the medial vein of the tegmen. Mysidia lacteola sp. n. (Figs 193, 303, 412) Male: head 0-88 mm long; 1-01 mm wide; pronotum 1-90 mm wide; tegmen 9-90-11-20 mm long; wing5-80 mm long. Female: tegmen 11-20 mm long. Length of frons 4-5 times width at apex, c. twice width at base; ocelli obscure; clypeus c. one-fifth longerthan frons; rostrum extending to base of subgenital plate. Pronotal width c. 47 times mid-dorsal length,fronto-lateral carinae absent; tegula strongly carinate basally. Antenna, frons at level of eyes, and fronto-lateral surfaces of pronotum occasionally tinged crimson.Tegmen and wing whitish hyaline, veins and cross-veins very pale brown. Tegmen with a pale brownishtransverse band at level of first fork of cubital vein, another at level of medial-cubital cross-vein, and a thirdslightly distad of midlength; posterior and apical margins narrowly and faintly edged pale brown. Wingwith a very faint, brownish, transverse band at midlength, and a second at level of radial-medial cross-vein. 40 PETER S. BROOMFIELD Shaft of aedeagus slender, slightly expanded towards apex; dorso-lateral surfaces each expandedsubapically into a low flap-like process; dorsal surface subapically with a slender spine on either side ofmidline. Paramere robust, dorsal process small, situated at two-fifths length, not posteriorly produced;dorsal surface at three-fifths length with a long, slender, apically rounded secondary process. MATERIAL EXAMINED Holotype cf , French Guiana: Mana River, v.1917 (CM). Paratypes. French Guiana: 1 $, same data as holotype (BMNH). Trinidad: 1 d" (USNM). Brazil: 5 C?,15 9, Taracua (NR; BMNH). The structure of the male genitalia, in particular the paramere, is very distinctive. Mysidia squamigera (Fabricius) Derbe squamigera Fabricius, 1803: 81. LECTOTYPE $, CENTRAL AMERICA (ZM), here designated [examined] .Mysidia squamigera (Fabricius) Westwood, 1840: 83. Female: head 0-75 mm long, 1-08 mm wide; pronotum 2-37 mm wide; legmen 10-50 mm long; wing6-40 mm long. Male unknown. Length of frons 7 times width at apex, c. 3 times width at base; ocelli small, distinct; clypeus c. one-fifthlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 50 times mid-dorsallength, fronto-lateral carinae absent; tegula prominently carinate. Antenna and genae irregularly orange; fronto-lateral surfaces of pronotum very dark brown dorsally andlaterally; tegula dark brown with dorsal margin narrowly pale; disc of mesonotum smoky brown overposterior three-quarters length; abdomen with dorsal surface narrowly smoky brown adjacent to midline.Tegmen and wing predominantly whitish hyaline, veins pale yellow or brown, cross-veins and posteriormargins broadly smoky brown. Tegmen with costal and radial areas very dark brown, interrupted bynarrow, irregular, yellow, transverse bands basad and distad of level of point of separation of fusedsubcostal and radial veins, and slightly basad of two-thirds length; base, including clavus, dark brown; abroad, transverse, dark brown band at one-sixth length; a narrower and more broken band over first fork ofcubital vein; a more distinct, somewhat oblique band extending from medial vein to apex of clavus;indistinct irregular markings over radial-medial cross- vein and apical fork of radial vein. Wing with anirregular, smoky brown, transverse band at c. two-fifths length. MATERIAL EXAMINED Lectotype $, Central America (Schmidt) (ZM). Brazil: 1 $ (INPA). The second syntype is also female, but is not conspecific. The lectotype is damaged and its left tegmen ismissing. The pigmentation of this species is very distinctive, and readily separates it from costata withwhich it is frequently confused. Mysidia costata (Fabricius)(Figs 220, 330, 440) Derbe costata Fabricius, 1803: 81. LECTOTYPE 9, CENTRAL AMERICA (ZM), here designated [ex-amined] .Mysidia costata (Fabricius) Westwood, 1840: 83. Male: head 0-74 mm long, 1-05 mm wide; pronotum 2-31 mm wide; tegmen 8-10-11-25 mm long; wing6-30 mm long. Female: tegmen 10-60-13-00 mm long. Length of frons 4-5 times width at apex, c. twice width at base; ocelli small, distinct; clypeus one-thirdlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 22 times mid-dorsallength; fronto-lateral surfaces and tegulae not carinate. Fronto-lateral surfaces of pronotum each with a large, roughly circular, dark brown spot, c. as large aseye, situated adjacent to lateral margin, well distant from eye; tegula concolorous with, or slightly paler,than eye; disc of mesonotum usually brownish. Tegmen and wing whitish hyaline, veins pale yellow,cross-veins pale brownish, posterior margins between veins smoky brown. Tegmen with costal cell palebrownish, darker at base; otherwise unmarked. Wing unmarked. Shaft of aedeagus broad, tapering from base to apex in dorsal aspect; dorsal surface subapically with a TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 41 pair of large flap-like processes, each strongly produced anteriorly into a long slender lobe bearing a smalltooth-like spine laterally at apex; ventral surface subapically produced into a long, transverse, spine-likeprocess. Paramere slender; apex acutely rounded; dorsal process well developed, situated at approxi-mately one-third length, posteriorly produced; dorsal surface at two-thirds length strongly and roundlyproduced into a medially directed lobe. MATERIAL EXAMINED Lectotype $ , Central America: no further data (Schmidt) (ZM).21 cf , 27 9 from various localities in Guyana, Trinidad, Brazil, Surinam, Peru and Ecuador. Many of the above specimens were incorrectly determined as squamigera which is readilydistinguished from costata by the strongly pigmented tegmina. Mysidiapseudocostatasp. n. (Figs 208, 319, 427) Male: head 0-55 mm long, 0-88 mm wide; pronotum 2-75 mm wide; tegmen 9-35-10-20 mm long; wing5-53 mm long. Female: tegmen 10-20-11-90 mm long. Length of frons c. 1 times width at apex, c. 2-5 times width at base; ocelli small, obscure; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 23 times mid-dorsallength; fronto-lateral carinae absent. Pronto-lateral surfaces of pronotum each with a broad orange-brown band extending horizontally fromadjacent to eye to lateral margin; tegula pale brownish. Tegmen and wing whitish hyaline; veins very pale;cross-veins slightly darker, brownish. Tegmen with costal cell yellowish brown, otherwise unmarked. Wingunmarked. Shaft of aedeagus basally slender, gradually broadening towards apex; lateral surfaces subapically eachwith a large, flap-like, process bearing a small, tooth-like projection on external surface. Paramere slender;apex narrowly rounded; dorsal process large, situated somewhat basad of mid-length, strongly producedposteriorly; dorsal surface at three-fifths length produced into a large, apically acute secondary processdensely covered with small tooth-like spines. MATERIAL EXAMINED Holotype cf , Guyana: Bugaba, 800-1500 ft (Champion) (BMNH). Paratypes. 4 cf, 2 $, same data as holotype; San Isidro; Blairmont (BMNH). Brazil: 1 9, Campinas(BMNH). The holotype, the other two specimens from the type-locality, and the single specimen from SanIsidro, all collected by Champion, are part of the Biologia Centrali Americana material, andwere previously determined as 'costata Fowler'. Externally this species closely resembles costata but is distinguished by the fronto-lateralsurfaces of the pronotum which have a horizontal reddish band; in costata the fronto-lateralsurfaces bear a circular dark brown spot. The structure of the male genitalia is also distinct. Mysidia delicatissima Fowler (Figs 275, 386, 496)Mysidia delicatissima Fowler, 1900: 74. LECTOTYPE cf , MEXICO (BMNH), here designated [examined]. Male: head 0-48 mm long, 0-68 mm wide; pronotum 1-30 mm wide; tegmen 6-80 mm long; wing 4-40 mmlong. Female unknown. Length of frons 5 times width at apex, c. twice width at base; ocelli obsolete; clypeus as long as frons;rostrum terminating at level of hind coxae. Pronotal width 32 times mid-dorsal length; fronto-lateralsurfaces and tegula not carinate. Genae each with a brownish spot at level of eye; abdomen with an oval dark brown spot on dorsal surfaceat either side of midline on fifth and sixth segments. Tegmen and wing whitish hyaline, veins pale,cross-veins brownish, posterior margins pale smoky brown. Tegmen with medial forks brown. Shaft of aedeagus laterally expanded over apical half; dorsal surface subapically with a pair of very largeflap-like processes, each terminating anteriorly in an acute spine. Paramere robust; apex broadly rounded;dorsal process very large, prominent, situated at midlength. 42 PETER S. BROOMFIELD MATERIAL EXAMINEDLectotype O", Mexico: Teapa, Tabasco, iii (Smith) (BMNH). The size and the relative lack of pigmentation of the tegmina and wing, and the structure of themale genitalia, readily distinguish this species. Mysidia bianca sp. n. (Figs 207, 318, 426) Male: head 0-60 mm long, 0-90 mm wide; pronotum 1-93 mm wide; legmen 10-03 mm long; wing 5-95 mmlong. Female unknown. Length of frons 5 times width at apex, 2-5 times width at base; ocelli small, indistinct; clypeus slightlylonger than frons; rostrum terminating slightly posterior to hind coxae. Pronotal width 16 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Genae dorsad of eyes, and vertex, brownish; fronto-lateral surfaces of pronotum brownish yellow.Tegmen and wing whitish hyaline, cross-veins narrowly edged smoky brown. Tegmen with costal marginnarrowly brownish; costal cell with a small dark spot adjacent to fork of fused subcostal and radial veins,and another similar spot somewhat basad; a small indistinct spot at apex of clavus, and another at apex ofanal vein. Wing lacking distinct markings, posterior margin weakly edged smoky brown. Shaft of aedeagus broad, expanded over apical third; dorsal surface subapically with a pair of long,curving, spine-like processes, and a pair of flap-like processes, each terminating in an acute point and a pairof ventrally directed projections laterally. Paramere slender; dorsal process large, situated at midlength;dorsal surface at three-quarters length with a large secondary process bearing numerous short robustspines. MATERIAL EXAMINEDHolotype cT, Bolivia: Prov. del Sara (Steinbach) (CM). This species can be distinguished most readily by the structure of the male genitalia. Mysidia cooperisp. n. (Figs 247, 358, 468) Male: head 0-59 mm long, 0-88 mm wide; pronotum 2-06 mm wide; tegmen 8-60 mm long; wing 5-61 mmlong. Female unknown. Length of frons 5 times width at apex ,1-66 times width at base ; ocelli small , obscure ; clypeus c. as long asfrons; rostrum extending fractionally beyond hind coxae. Pronotal width 16 times mid-dorsal length,fronto-lateral carinae very prominent; tegula with carinae weak. Frons and genae at level of eyes irregularly brownish; fronto-lateral surfaces of pronotum at level of eyesbroadly pale orange. Tegmen and wing whitish hyaline, veins pale, cross-veins very dark brown. Tegmenwith costal margin narrowly black basally; costal cell with a prominent blackish spot at level of point ofseparation of subcostal and radial veins; clavus with a blackish brown spot at apex of anal vein; apical andposterior margins with semicircular smoky brown spots between veins. Wing with radial-medial cross-veinvery dark brown; a dark brown spot between cubital vein and claval suture at midlength; a dark spot onposterior margin at apex of clavus; semi-circular dark spots on posterior margin between branches ofcubital, medial and radial veins. Shaft of aedeagus rotated 90 clockwise in anterior aspect; right lateral surface subapically produced intoa pair of flap-like processes bearing very numerous, small, conical spines, ventral process terminatinganteriorly in an acute spine. Paramere slender, apex acute; dorsal process situated at midlength, robust,apex little produced. MATERIAL EXAMINEDHolotype d", Colombia: Putumayo, Mocoa, 550 m, 16.viii.1978 (Cooper] (BMNH). This species is readily distinguished by the pigmentation and by the unique structure of theaedeagus. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 43 Mysidia diana sp. n. (Figs 227, 338, 447) Male: head 0-63 mm long, 0-80 mm wide; pronotum 1-82 mm wide; tegmen 9-35 mm long; wing 5-53 mmlong. Female: tegmen 11-22 mm long. Length of frons 6-5 times width at apex, 2-66 times width at base; ocelli small, distinct; clypeusone-quarter longer than frons; rostrum extending to base of subgenital plate. Pronotal width 18 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with costal margin smoky brown;an irregular brownish band extending over first fork of cubital vein almost to vanal fold; indistinct palebrown mottling immediately distad of midlength; a small, prominent, dark brown spot over bases offourth, fifth and sixth branches of medial vein; a smaller dark spot on margin over apex of fifth branch ofmedial vein; cross-veins narrowly dark brown. Wing with anal area irregularly mottled smoky brownextending over first and second forks of cubital vein; a very large smoky brown area covering radial-medialcross-vein and branches of medial vein extending to apical margin between branches of medial and cubitalveins. Shaft of aedeagus robust; dorsal surface subapically with a pair of very large, apically truncate, flap-likeprocesses. Paramere broad, apex rounded; dorsal process situated immediately basad of midlength,strongly produced posteriorly; internal lateral surface with a low curving ridge bearing robust spines atone-quarter length. MATERIAL EXAMINED Holotype cf , Ecuador: Feltons, 12 km Napo, nr Tema, 8.iv.l923 (Williams) (BMNH).Paratype. Ecuador: 1 $, Tema (BMNH). This species is readily distinguished by the pigmentation of the wing and by the structure of themale genitalia. Mysidia dollingisp. n. (Figs 255, 366, 476) Male: head 0-63 mm long, 0-95 mm wide; pronotum 1-97 mm wide; tegmen 9-35 mm long; wing 5-10 mmlong. Female: tegmen 10-00-11-40 mm long. Length of frons c. 1 times width at apex, 2-5 times width at base; ocelli indistinct; clypeus one-fifth longerthan frons; rostrum extending to base of subgenital plate. Pronotal width 19 times mid-dorsal length,fronto-lateral carinae absent; tegula prominently carinate. Head with a broad orange band extending horizontally from anterior margin of genae to eye, andcontinuing over fronto-lateral surface of pronotum to lateral margin. Tegmen and wing whitish hyaline,posterior and apical margins pale smoky brown, cross-veins dark brown; veins intermittently narrowlyedged smoky brown. Tegmen with costal cell pale brownish, with a small dark brown spot at level of firstfork of cubital vein and another immediately basad of second fork; claval margin with a small brown spotadjacent to point of fusion of anal veins and another at apex of anal vein. Shaft of aedeagus basally slender, becoming laterally expanded towards apex; dorsal surface subapicallywith a pair of slender spine-like processes and a pair of lateral flap-like processes, each terminating in asmall spine. Paramere very robust; apex broadly rounded; dorsal process situated somewhat distad ofmidlength, apex produced posteriorly; dorsal surface at one-quarter length with a rounded flap-likesecondary process bearing numerous, very long, robust spines. MATERIAL EXAMINED Holotype cf , Panama: Canal Zone, Barro Colorado, 8.viii.l967 (O'Brien) (FAMU). Paratypes. Panama: 1 Cf, 4 $, Canal Zone (FAMU; BMNH). Costa Rica: 4 $, Guan, 5 miles SE.Liberia (FAMU; BMNH). Mysidia striata sp. n. Female : head 50 mm long ,0-71 mm wide ; pronotum 1 40 mm wide ; tegmen 6 80 mm long ; wing 3 80 mmlong. Male unknown. Length of frons 6 times width at apex, c. twice width at base; ocelli very large, prominent; clypeus slightlylonger than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 14 timesmid-dorsal length; fronto-lateral carinae distinct; tegula without carinae. 44 PETER S. BROOMFIELD Genae dorsad of eyes tinged reddish; pronotum white, fronto-lateral surfaces each with a broad, darkreddish brown band extending horizontally dorsad of carina from adjacent to eye to lateral margin; anothersimilar band terminating immediately prior to reaching lateral margin at level of ventral margin of eye;meso- and metanotum yellowish brown dorsally, tinged reddish ventrally; coxae reddish; abdomendorsally deep brown, paler at mid-dorsal line, ventral surface deep reddish. Tegmen and wing yellowishhyaline, veins yellow. Tegmen with cross-veins edged pale brownish; a narrow, brown, transverse bandextending from costal margin to apex of clavus; a large dark brownish spot at three-quarters lengthextending from costal margin over medial-radial cross-vein. Wing with a broad, brown, transverse bandextending obliquely from costal to posterior margin at level of cross- veins; apical two-fifths brown, medialand apical cells broadly hyaline medially. MATERIAL EXAMINED Holotype $, Brazil: Para, Jabaty, v.1924 (Williams) (BMNH).Paratype. 1 $, same data as holotype (BMNH). In the absence of males, this species is very readily distinguished by its pigmentation, especiallythe unique double horizontal dark bands on the fronto-lateral surfaces of the pronotum. Mysidia sanguinea sp. n. (Figs 215, 325, 434) Male: head 0-71 mm long, 0-90 mm wide; pronotum 1-68 mm wide; tegmen 7-20-8-30 mm long; wing4-33 mm long. Female: tegmen 7-40-8-40 mm long. Length of frons 5-5 times width at apex, 2-5 times width at base; ocelli large, prominent; clypeus as longas frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 17 times mid-dorsallength, fronto-lateral carinae highly elevated; tegula not distinctly carinate. Frons, genae ventral to lower margins of eyes, and vertex deep reddish brown; ocelli orange;fronto-lateral surfaces of pronotum ventral to midline of eyes, fore tibia, apices of mid and hind tibia, anddisc of mesonotum dark reddish brown; pronotum dorso-laterally, under surfaces of mesonotum,metanotum, and abdomen, and legs pale brownish; dorsal surface of abdomen dark reddish brown/black,pregenital segment narrowly crimson posteriorly. Tegmen and wing whitish hyaline, veins dark brown/black. Tegmen with a large, irregular, dark-brown spot near base, another at apex of clavus; a transversedark band extending from costal margin to first fork of cubital vein, linking with another dark spot at firstfork of medial vein, and extending to cover second cubital vein and medial-cubital cross-vein; apical forksof medial and radial veins narrowly edged orange or yellow; posterior margin between veins narrowly darkbrown. Wing with an irregular, obliquely transverse dark brownish band covering forks and cross-veins;branches of medial and cubital veins narrowly edged dark brown; posterior margin narrowly and distinctlydark. Male genitalia with shaft of aedeagus slender; ventral surface subapically with a pair of slender spines;dorsal surface unarmed. Paramere basally slender, becoming expanded from midlength; apex obtuselyrounded; dorsal process large, robust, situated at two-thirds length; dorsal surface at one-third length witha large, obtusely rounded, secondary process bearing numerous short robust spines. MATERIAL EXAMINED Holotype cf , Brazil: Vila Amazonas, Amapa, 21.iii.1963 (Ross) (CAS). Paratypes. Brazil: 6 cf, 13 $, Amazonas (CAS; INPA; BMNH). Peru: 1 cf, Tingo Maria (CAS).Surinam: 1 cf , Brokopondo (FAMU). This species is readily distinguished by the striking pigmentation of the head, body, tegmina andwings, and by the unique structure of the male genitalia, in particular the aedeagus. Mysidia calypso sp. n. (Figs 237, 146, 457) Male: head 0-53 mm long, 0-78 mm wide; pronotum 1-72 mm wide; tegmen 8-84 mm long; wing 5-10 mmlong. Female unknown. Length of frons c. 1 times width at apex, c. 2-5 width at base; ocelli very large, prominent; clypeus c. aslong as frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 20 timesmid-dorsal length; fronto-lateral carinae distinct; tegula not carinate. Disc of mesonotum and dorsal surface of abdomen dark brown; ocelli yellow, narrowly edged scarlet. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 45 Tegmen and wing whitish hyaline, veins dark brown. Tegmen with cross- veins narrowly edged dark brown;posterior margin between veins broadly brownish; basal cells irregularly smoky brown medially; apicalcells and cells on posterior margin irregularly edged smoky brown; clavus with a dark spot subapically; adark brown spot between apex of clavus and first branch of cubital vein. Wing with apex of clavus, adjacentarea at level of first branch of cubital vein, and apical one-fifth length broadly dark smoky brown. Shaft of aedeagus slender; dorsal surface subapically with a pair of long apically serrated processeslaterally, and a pair of broad flap-like processes bearing numerous small conical spines medially, each witha small spine basally; ventral surface with a pair of short, apically acute, flap-like processes subapically.Paramere very slender; apex obliquely truncate; dorsal process arising slightly basad of midlength, greatlyproduced posteriorly; dorsal surface at one-fifth length with a broadly rounded secondary process bearingnumerous short robust spines. MATERIAL EXAMINEDHolotype cT, Brazil: Rezende, Estado de Rio, ii.1924 (Williams) (BMNH). This species is distinguished by the tegminal, and especially the wing, pigmentation, and by thestructure of the male genitalia. My si din lucianna sp. n. (Figs 262, 373, 481) Male: head 0-54 mm long, 0-90 mm wide; pronotum 1-89 mm wide; tegmen 8-80-9-10 mm long; wing5-35 mm long. Female: tegmen 9-35-9-84 mm long. Length of frons c. 4 times width at apex, c. twice width at base; ocelli obsolete; clypeus c. one-thirdlonger than frons; rostrum terminating at level of mid coxae. Pronotal width 13 times mid-dorsal length;fronto-lateral carinae very prominent; tegula distinctly carinate. Frons with a narrow, transverse, dark brown band at level of dorsal margins of eyes; vertex mottled darkbrown; fronto-lateral surfaces of pronotum each with a broad pale orange band extending horizontallyfrom adjacent to eye to lateral margin. Tegmen and wing whitish hyaline, cross-veins narrowly dark brown.Tegmen with a small dark brown spot on costal margin slightly basad of one-third length; an irregular palebrown area around apical forks of radial and medial veins; a small brown spot on claval margin at apex ofanal vein; a paler brown spot at level of junction of anal veins; apical margin between veins with smallsmoky brown spots. Wing with a transverse brown marking over cubital vein at midlength; posteriormargin with dark brown spots between anal veins, between branches of cubital vein, and between cubitaland medial branches. Shaft of aedeagus slightly asymmetrical, somewhat expanded over apical half; dorsal surface subapicallywith a pair of large flap-like processes bearing numerous , extremely small , tooth-like projections ; left-handprocess terminating anteriorly in a curved spine. Paramere long and slender; apex acute; dorsal processrobust, not greatly produced posteriorly, situated at midlength; dorsal surface at approximately one-fifthlength with a small, blunt, secondary process bearing numerous long slender spines. MATERIAL EXAMINED Holotype cf , Brazil: Belem, Para, vi.1924 (Williams) (BMNH).Paratypes. Brazil: 15 C?, 22 $ (BMNH; FAMU; INPA). Surinam: 1 cf , S. Kraka (FAMU). This species is most readily determined by the pigmentation of the head and pronotum, and bythe structure of the male genitalia. Mysidia peregrins sp. n. (Figs 182, 292, 401) Male: head 0-52 mm long, 0-82 mm wide; pronotum 2-12 mm wide; tegmen 7-65 mm long; wing 4-42 mmlong. Female unknown. Length of frons 4 times width at apex, 2-5 times width at base; ocelli large, prominent; clypeus c. as longas frons; rostrum terminating at level of hind coxae. Pronotal width c. 9 times mid-dorsal length,fronto-lateral carinae very prominent; tegula not carinate. Fronto-lateral surfaces of pronotum each with a brownish band extending horizontally from adjacent toeye to lateral margin; disc of mesonotum whitish anteriorly, gradually darkening posteriorly. Tegmen andwing whitish hyaline. Tegmen with cross-veins and branches of veins dark brown; cells, especially thoseadjacent to costal margin, densely mottled smoky brown; a small, distinct black spot over apical fork of 46 PETER S. BROOMFIELD medial vein; a small, but very distinct, black spot in each of the five apical cells. Wing with veins darkbrown; cells faintly and sparsely mottled smoky brown. Shaft of aedeagus cylindrical; apex slightly expanded, curving dorsally; ventral surface subapically withnumerous, extremely small, blunt spines; dorsal surface subapically with a pair of long spines laterally, anda pair of hooked spine-like processes adjacent to midline. Paramere basally slender, expanded apically;dorsal process reduced to a simple, posteriorly directed hook at one-third length; internal surface with alow, transverse, flap-like process bearing robust spines at one-quarter length. MATERIAL EXAMINEDHolotype cf , Brazil: Mato Grosso, Barra do Tapirape, 30.xii.1952 (Malkin) (CAS). This species is readily distinguished by the prominent black spot on the tegmen; and by theunique structure of the male genitalia in which the complex armature of the aedeagus is coupledwith the great reduction in the development of the dorsal process of the paramere. Mysidia fowlerisp. n. (Figs 173, 282, 391) Male: head 0-44 mm long, 0-61 mm wide; pronotum 1-64 mm wide; tegmen 5-50-5-90 mm long; wing3-05 mm long. Female: tegmen 6-20-6-40 mm long. Length of frons 4 times width at apex, twice width at base; ocelli large, not prominent; clypeusone-quarter longer than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width11 times mid-dorsal length, fronto-lateral carinae prominent; tegula distinctly carinate. Pronto-lateral surfaces of pronotum each with an orange band extending horizontally from adjacent toeye to lateral margin. Tegmen and wing clear hyaline, costal margins and basal areas predominantly darkbrown, veins and cross-veins broadly margined dark smoky brown. Shaft of aedeagus slender in dorsal aspect; ventral surface at midlength and subapically thickly coveredwith tiny blunt spines; dorsal surfaces at two-thirds length with a pair of rounded flap-like processes and apair of slender spines. Paramere with dorsal process situated at one-quarter length, greatly producedposteriorly; a large flap-like process on internal surface basally; dorsal surface over apical half stronglycurved towards midline, bearing numerous short robust spines; ventral surface subbasally with numerouslong slender spines, at midlength with a long, slender, apically rounded process. MATERIAL EXAMINED Holotype cf , Panama: C.Z., Fort Kobbe, 24.vi.1976 (Riley) (FAMU).Paratypes. 1 d", 3 $, same data as holotype (FAMU; BMNH). The external characters confirm the placement of this species in Mysidia, but the male genitaliaare not characteristic of the genus. Mysidiagrandissp. n. Female: head 0-80 mm long, 1-05 mm wide; pronotum 2-56 mm wide; tegmen 13-25-13-40 mm long; wing7-80 mm long. Male unknown. Length of frons c. 6 times width at apex, 4 times width at base; ocelli large, not prominent; clypeusone-sixth longer than frons; rostrum extending to base of pregenital segment. Pronotal width 24 timesmid-dorsal length; fronto-lateral carinae absent; tegula with distinct carinae. Genae occasionally tinged brownish between eye and anterior margin; fronto-lateral surfaces ofpronotum occasionally tinged with orange laterally; tegula dorsad to carinae dark brown; disc ofmesonotum with a broad, dark brown, transverse band medially; metanotum with a large, circular, darkbrown spot on either side at base of scutellum. Tegmen and wing whitish hyaline, veins pale brown,cross-veins edged smoky hyaline, posterior and apical margins with smoky brown spots between veins.Tegmen with costal cell white, with a small, dark brown spot at level of subcostal-radial fork, and anothersomewhat basad; basal half of cells between radial and medial veins, and entire cell between medial andcubital veins boldly and irregularly mottled dark brown and yellow, adjacent cells mottled smoky hyaline;radial vein with a prominent, roughly circular, blackish brown spot over apical fork; fifth and sixth radialbranches linked by a paler, irregular, brownish spot; first cubital branch with a large, irregular, brown spotextending to claval margin. Wing with a large, irregular, broad, smoky brown band extending from cubitalvein to anal margin at approximately one-fifth length. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 47 MATERIAL EXAMINED Holotype $, Panama: Chiriqui, Fortuna, 82 15'W 8 44'N, 17.V.1978 (O'Brien & Marshall) (FAMU).Paratype. 1 $ , same data as holotype (BMNH). This species is readily distinguished by its large size and by the pigmentation of the thorax andtegmina. Mysidiu minerva sp. n. (Figs 205, 315, 424) Male: head 0-55 mm long, 0-63 mm wide; pronotum 1-25 mm wide; tegmen 5-70-6-55 mm long; wing3-80 mm long. Female unknown. Length of frons 5 times width at apex, twice width at base; ocelli large, prominent; clypeus slightly shorterthan frons; rostrum terminating level with hind coxae. Pronotal width 20 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Genae and frons at level of eyes pale reddish brown; fronto-lateral surfaces of pronotum each with aprominent bright orange band extending horizontally from adjacent to eye to lateral margin; dorsal surfaceof abdomen basally mottled bright orange. Tegmen and wing hyaline. Tegmen with veins pale brown;cross-veins and forks of veins dark brown, very narrowly edged smoky brown; with two roughly circulardark smoky brown spots on costal cell subbasally; apex of clavus with a large, irregular, smoky brown spot.Wing with veins over apical two-thirds length dark brown; apex of claval area narrowly dark smoky brown. Shaft of aedeagus slender, apex laterally expanded; dorsal surface subapically with a pair of large,rounded, flap-like processes, each bearing a small spine-like projection subbasally. Paramere with apexbroadly and irregularly rounded; dorsal process large, situated at two-thirds length, strongly producedpostero-dorsally. MATERIAL EXAMINED Holotype cf , Belize: Toledo District, Salamanca, 25 miles NW. Punta Gorda, 28.viii.1978 (Broomfield)(BMNH). Paratypes. Belize: 1 cf , Belize; 1 cf , Orange Walk District (FAMU). This species is most readily distinguished by the very sparse pigmentation of the tegmen andwing, coupled with the bright orange bands on the pronotum, and by the structure of the malegenitalia. Mysidia punctifera MetcalfMysidia punctifera Metcalf, 1938: 313. Holotype $, PANAMA (MCZ) [examined]. Female: head 0-71 mm long, 0-92 mm wide; pronotum 2-00 mm wide; tegmen 9-35 mm long; wing 5-53 mmlong. Male unknown. Length of frons 7 times width at apex, 3 times width at base; ocelli large, not prominent; clypeus slightlylonger than frons; rostrum terminating at level of hind coxae. Pronotal width 19 times mid-dorsal length;fronto-lateral carinae absent; tegula weakly carinate basally. Genae each with a narrow, pale brown band extending from adjacent to midline of eye to anteriormargin; ocelli pale; fronto-lateral surfaces of pronotum broadly pale orange from adjacent to eyes tolateral margins; disc of mesonotum irregularly brown posteriorly between lateral carinae; dorsal surface ofabdomen devoid of dark markings. Tegmen and wing predominantly whitish hyaline; veins yellowish;cross-veins dark brown, narrowly edged dark smoky brown, Tegmen with costal cell irregularly mottleddark brown over basal half; radial, medial and cubital areas densely mottled dark brown and yellow overbasal two-fifths; claval area narrowly dark brown basally, with a large, prominent, irregular, dark brownspot between apex and first branch of cubital vein; medial area at slightly distad of two-fifths length, andagain at level of second fork, broadly and irregularly dark brown, the latter marking extending anteriorly tosubcostal vein and, indistinctly, posteriorly to adjacent to claval apex; apical fork of medial vein with alarge, prominent, dark brown spot; posterior and apical margins with small pale brownish spots intermit-tently between veins. Wing with a pale, indistinct, narrow, transverse brownish band at level of first fork ofcubital vein; a large, indistinct, pale brown spot over fork of medial vein; a distinct, dark brown, roughlycircular spot between first fork of cubital vein and claval suture at approximately midlength; posteriormargin between anal veins and branches of cubital veins medially dark brown, the former very prominentlyso. 48 PETER S. BROOMFIELD MATERIAL EXAMINEDHolotype $, Panama: Canal Zone, Barro Colorado, 15.vii.1924 (Banks) (MCZ). The antennae are unusually long, extending for one-half their length beyond anterior margins ofgenae. The species is also distinguished by the dark mottling and the prominent dark spot at theapex of the clavus of the tegmen. Mysidia maculicosta Fowler Mysidia maculicosta Fowler, 1900: 73. LECTOTYPE $>, GUATEMALA (BMNH), here designated [ex-amined]. Female: head 0-75 mm long, 0-88 mm wide; pronotum 1-80 mm wide; tegmen 9-60 mm long; wing 6-00 mmlong. Male unknown. Length of frons 7-5 times width at apex, 2-66 yimes width at base; ocelli large, prominent; clypeus slightlyshorter than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 30 timesmid-dorsal length, fronto-lateral carinae absent; tegula distinctly carinate. Genae reddish at level of dorsal margins of eyes; fronto-lateral surfaces of pronotum each with a broad,horizontal, reddish brown band extending from adjacent to eye to lateral margin; abdomen dorsally with alarge dark brown spot at midline over first three segments. Tegmen and wing whitish hyaline, veins pale,cross-veins narrowly edged dark brown. Tegmen with costal margin irregularly brownish from base to levelof first fork of cubital vein, this marking extending posteriorly to cover base of first branch of cubital vein;apical fork of medial vein covered by an irregular dark brown spot; clavus with a brown spot at apex; fifthand sixth branches of medial vein each with a brown spot near base; posterior margin weakly brownishbetween veins. Wing with irregular brownish markings on cross-veins and posterior margin. MATERIAL EXAMINEDLectotype $, Guatemala: Pantaleon, 1700ft (Champion) (BMNH). There is also a male from Costa Rica in the type-series; it is very badly damaged, with the head,pronotum and right wing and tegmen missing, and it is very doubtful if it represents this species.The specimen here designated as lectotype bears Fowler's handwritten 'type' label. The speciesis distinguished by the pigmentation of the pronotum and abdomen. Mysidia molests sp. n. (Figs 187, 297, 406) Male: head 0-69 mm long, 1-01 mm wide; pronotum 2-14 mm wide; tegmen 9-35 mm long; wing 5-52 mmlong. Female: tegmen 9-35-10-20 mm long. Length of frons 6-5 times width at apex, 2-33 times width at base; ocelli small, obscure; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 17 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Disc of mesonotum with a large dark brown spot posteriorly. Tegmen and wing whitish hyaline, veinsyellow, posterior and apical margins broadly smoky brown. Tegmen heavily mottled dark brownish, thesemarkings coalescing to form irregular transverse bands at one-third and two-thirds length. Wing withbranches of cubital vein broadly and irregularly edged smoky brown; apical third smoky brown over radialand medial veins. Shaft of aedeagus broadly laterally expanded; apex strongly produced dorsally; dorsal surface subapical-ly with a pair of large flap-like processes extending over lateral surfaces, each bearing a single, deeplybifurcate projection. Paramere very robust, greatly expanded dorso-ventrally, longitudinally foldedtowards midline; dorsal process situated at two-fifths length, ventrally directed, small, not posteriorlyproduced. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, vii-viii.1981 (Arias) (INPA).Paratypes. 4 , same data as holotype (INPA; BMNH). Closely related to estfarchina, molesta is distinguished by the dark spot on the mesonotum andby the pigmentation of the tegmen and wing. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 49 Mysidia obscura Metcalf (Figs 260, 371, 483)Mysidia obscura Metcalf, 1938: 317. Holotype $, PANAMA (USNM) [examined]. Male: head 0-84 mm long, 1-05 mm wide; pronotum 2-25 mm wide; tegmen 9-70 mm long; wing 5-44 mmlong. Female: tegmen 11-05-12-00 mm long. Length of frons c. 5-5 times width at apex, 3-5 times width at base; ocelli obsolete; clypeus slightly longerthan frons; rostrum extending to subgenital plate. Pronotal width c. 14 times mid-dorsal length; fronto-lateral surfaces and tegula distinctly carinate. Head and body unmarked. Tegmen and wing whitish hyaline, cross-veins and posterior and apicalmargins broadly edged smoky brown. Tegmen with central areas of cells irregularly smoky brown, thesemarkings coalescing with those bordering cross-veins to form very irregular transverse bands. Wing with anirregular, smoky brown, transverse band at c. midlength, and another over radial-medial cross- vein. Shaft of aedeagus very broad in lateral aspect; dorsal surface subapically with a pair of large,longitudinally aligned, dorsally directed, flap-like processes, each bearing a pair of short, robust, spine-likeprojections dorsally. Paramere slender; apex acutely rounded; dorsal process situated slightly distad ofmidlength, little produced posteriorly; dorsal surface subbasally with a blunt secondary process bearingrobust spines. MATERIAL EXAMINED Holotype $, Panama: Porto Bello, 27.ii.1911 (Busck) (USNM).Panama: 1 cT, 1 $, Barro Colorado (CAS; BMNH). This species is distinguished by its large size, the mottled appearance of the tegmina and wings,and by the structure of the male genitalia. Mysidia henrietta sp. n. (Figs 277, 388, 498) Male: head 0-59 mm long, 0-73 mm wide; pronotum 1-57 mm wide; tegmen 7-65-8-10 mm long; wing4-68 mm long. Female: tegmen 8-90 mm long. Length of frons 8 times width at apex, twice width at base; ocelli small, indistinct; length of clypeus equalto that of frons; rostrum terminating at level of hind coxae. Pronotal width 19 times mid-dorsal length;fronto-lateral carinae obsolete; tegula distinctly carinate. Genae each with a narrow reddish or dark brown band extending from adjacent to eye to anteriormargin; fronto-lateral surfaces of pronotum each with a pale, often indistinct, horizontal band extendingfrom midline of eye to lateral margin. Tegmen and wing whitish hyaline, cross- veins dark brown. Tegmenwith a small, prominent, dark brown spot over apical fork of medial vein; a small, often indistinct, palebrown spot on clavus adjacent to junction of anal veins. Wing unmarked. Male genitalia with shaft of aedeagus slender, broadest at midlength; dorsal surface subapically with apair of apically bifurcate flap-like processes. Paramere massive; apex rounded; dorsal process situated atmidlength, slender, strongly produced posteriorly. MATERIAL EXAMINED Holotype cf , Brazil: Nietheroy, iv.1924 (Williams) (BMNH).Paratypes. Brazil: 4 cf , 7 $, same data as holotype; Itaparica; Bahia (BMNH; NR). This species is readily distinguished by the pigmentation of the tegmen and wing, and by thestructure of the male genitalia. Mysidia distant! sp. n. (Figs 174, 283, 392) Male: head 0-59 mm long, 0-75 mm wide; pronotum 1-60 mm wide; tegmen 7-80-8-50 mm long; wing4-70 mm long. Female: tegmen 8-90-10-00 mm long. Length of frons c. 7-5 times width at apex, slightly less than 3 times width at base; ocelli distinct, notprominent; clypeus as long as frons; rostrum extending to base of subgenital plate. Pronotal width slightlygreater than 19 times mid-dorsal length; fronto-lateral surfaces not carinate; tegula carinate basally. Genae anterior to eyes often brownish; fronto-lateral surfaces of pronotum and tegula often orange at 50 PETER S. BROOMFIELD level of eyes; dorsal surface of abdomen seldom with two pairs of small dark brown spots adjacent tomidline basally. Tegmen and wing almost hyaline, very weakly tinged whitish, veins yellowish; cross-veinsdark brown, edged smoky brown; posterior and apical margins broadly smoky brown between veins.Tegmen weakly and irregularly mottled smoky brown between branches of medial and cubital veins; clavusoften with a small dark brown spot level with point of fusion of anal veins; apical fork of medial veinnarrowly and distinctly dark brown or black. Wing with radial-medial cross-vein and adjacent branchesvery dark brown. Male genitalia with shaft of aedeagus broad in lateral aspect; dorsal surface at midlength with a pair oflarge flap-like processes extending almost to apex; lateral surfaces each with a flap-like process extendingfrom subapically almost to base. Paramere robust; apex broadly rounded; dorsal process situated slightlydistad of midlength, strongly curved postero-ventrally. MATERIAL EXAMINED Holotype cf , Honduras: Sta. B. 13 km SE. El Mochito, 22.vii.1977 (O'Brien & Marshall) (FAMU). Paratypes. Honduras: 2 cf, 3 $, same data as holotype (FAMU; BMNH). Belize: 1 cf, Belize Distr.(FAMU). This species closely resembles insolita, but differs in the almost hyaline tegmina and wings, thedark pigmentation of the veins of the wing around the radial-medial cross-vein, and in thestructure of the male genitalia. Mysidia albicans Stal(Figs 279, 390, 500) Derbe albicans Stal, 1855: 191. LECTOTYPE cf , BRAZIL (NR), here designated [examined].Mysidia albicans (Stal) Stal 1856: 163. Male: head 0-61 mm long, 0-71 mm wide; pronotum 2-40 mm wide; tegmen 8-40 mm long; wing 5-30 mmlong. Female unknown. Length of frons slightly less than 6 times width at apex, 2-25 times width at base; ocelli small, notprominent; length of clypeus one-fifth greater than that of frons; rostrum extending somewhat behindposterior coxae. Pronotal width 27 times mid-dorsal length; fronto-lateral surfaces not carinate. Genae at level of eyes dull brownish; fronto-lateral surfaces of pronotum broadly and very indistinctlypale brown at level of eyes. Tegmen and wing whitish hyaline, veins pale yellow. Tegmen with cross-veinsand branches of medial vein narrowly dark brown; clavus with a faint brownish spot subbasally, andanother, more distant, spot adjacent to point of fusion of anal veins; costal cell with a small, dark brownspot at one-sixth length, and another at point of separation of subcostal and radial veins, the latter spotbeing somewhat fainter and extending transversely over medial vein; medial vein with a small, veryprominent, roughly circular, dark brown/black spot over apical fork; a very faint and irregular, palebrownish, transverse band extending from costal margin to third branch of cubital vein slightly distad ofmidlength. Wing with radial-medial cross-vein brownish; very indistinctly mottled pale brown at level offirst and second forks of cubital vein; a very irregular, broken, pale brownish, transverse band at levelof fork of radial vein; an irregular, brownish spot between first branch of cubital vein and claval suture attwo-thirds length of latter. Male genitalia with shaft of aedeagus slender; lateral surfaces subapically each with a large, flap-likeprocess extending over dorsal surface and produced anteriorly into an acute spine. Paramere robust; apexbroadly rounded; dorsal process situated somewhat distad of midlength, strongly produced posteriorly;dorsal surface subbasally bearing numerous short, robust spines. MATERIAL EXAMINEDLectotype cf , BrazU (Westermari) (NR). The single specimen available for study is damaged and the tegulae are missing; it appears to beteneral but is believed to be free of distortion. In his description Stal did not cite the number ofspecimens, neither did he designate a holotype; the specimen listed above is here designated aslectotype. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 51 Mysidianemorensissp. n. (Figs 195, 305, 414) Male: head 0-59 mm long, 0-86 mm wide; pronotum 1-95 mm wide; legmen 7-91 mm long; wing 4-50 mmlong. Female: tegmen 9-70 mm long. Length of frons rather less than 5 times width at apex, slightly greater than twice width at base; ocellismall, distinct; clypeus one-quarter longer than frons; rostrum extending to base of subgenital plate.Pronotal width 22 times mid-dorsal length; fronto-lateral carinae obsolete; tegula with distinct carinae. Pronto-lateral surfaces of pronotum usually each with a broad yellowish brown band extendinghorizontally from adjacent to eye to lateral margin. Tegmen and wing whitish hyaline, irregularly mottledsmoky brown around forks of veins and cross-veins and on posterior and apical margins; veins pale brown.Tegmen with an irregular, smoky brown, transverse band at level of first fork of cubital vein. Wingunmarked. Male genitalia with shaft of aedeagus broadly laterally expanded distad of midlength; dorsal surfacesubapically with a pair of large flap-like processes, broadening apically, each produced anteriorly andposteriorly into a long curving spine-like projection. Paramere robust; apex obtusely rounded; dorsalprocess situated at mid-length, not posteriorly produced. MATERIAL EXAMINED Holotype cf , Surinam: Brokopondo, 30.U969 (O'Brien) (FAMU). Paratypes. Surinam: 1 cf , 2 $ , same data as holotype (FAMU; BMNH). Brazil: 11 cf , 12 9 , Amazonas(INPA; BMNH). Not readily distinguished by external characters, this species can be recognized most easily bythe complex armature of the aedeagus and the undeveloped paramere. Mysidia insolita sp. n. (Figs 213, 323, 432) Male: head 0-61 mm long, 0-75 mm wide; pronotum 1-95 mm wide; tegmen 6-80-8-10 mm long; wing4-50 mm long. Female: tegmen 9-35 mm long. Length of frons 6-5 times width at apex, slightly less than 2-5 times width at base; ocelli large, notprominent; clypeus slightly longer than frons; rostrum extending to base of subgenital plate. Pronotalwidth slightly less than 18 times mid-dorsal length; fronto-lateral surfaces and tegula not carinate. Genae tinged orange-brown; fronto-lateral surfaces of pronotum each with a broad, horizontal, orangeband extending from adjacent to eye to lateral margin. Tegmen and wing whitish hyaline, veins yellow;cross-veins brown, broadly edged smoky brown; posterior and apical margins broadly smoky brownbetween veins. Tegmen with costal and radial cells mottled brown; with an irregular, smoky brown,transverse band at one-quarter length; apical fork of medial vein narrowly very dark brown. Wingirregularly mottled smoky brown. Male genitalia with shaft of aedeagus broad; dorsal surface subapically with a pair of large flap-likeprocesses adjacent to midline and extending anteriorly to base. Paramere slender; apex acute; dorsalprocess situated at one-third length, produced postero-dorsally, with a ventrally directed lobe at mid-length. MATERIAL EXAMINED Holotype cf , Honduras: Cortes, 27 km S. Potrerillos, 8.viii.l977 (O'Brien & Marshall) (FAMU).Paratypes. Honduras: 1 cf , same data as holotype (BMNH). Belize: 1 <j> , Altun Ha (FAMU). Bearing a close superficial resemblance to distanti, this species is distinguished by the densermottling of the tegmen and wing, the paler veins of the latter, and by the structure of the malegenitalia. Mysidia enjebetta sp. n. (Figs 281, 289, 399) Male: head 0-63 mm long, 0-80 mm wide; pronotum 1-70 mm wide; tegmen 7-90 mm long; wing 4-85 mmlong. Female unknown.Length of frons 6-5 times width at apex, slightly less than 3 times width at base; ocelli obsolete; clypeus as 52 PETER S. BROOMFIELD long as frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 27 timesmid-dorsal length; fronto-lateral surfaces not carinate; tegula weakly carinate. Lateral carinae of frons tinged brownish at level of eyes; fronto-lateral surfaces of pronotum each with abroad orange band extending horizontally from adjacent to eye to lateral margin; lateral surfaces ofmesonotum tinged crimson ventral to bases of tegmina. Tegmen and wing whitish hyaline; veins paleyellowish brown; cross-veins and forks of veins dark brown; posterior and apical margins smoky brownbetween veins. Tegmen with cross-veins broadly margined smoky brown; apical fork of medial vein veryprominently dark brown; non-apical cells narrowly and irregularly pale smoky brown medially. Wing withirregularly spaced, roughly circular, smoky brown spots between cubital vein and claval suture; apical cellsirregularly tinged smoky brown medially. Male genitalia with shaft of aedeagus broad; dorsal surface subapically with a pair of very large, flap-likeprocesses; lateral surfaces each with a very large, flap-like process over apical half length. Paramere withapex acutely rounded; dorsal process situated slightly basad of mid-length, strongly produced; dorsalsurface at three-quarters length strongly and roundly produced into an internally directed secondaryprocess. MATERIAL EXAMINEDHolotype cT, Mexico: Tepic, Nayarit, 13.iii.1957 (Dreisbach) (USNM). This species is distinguished by the combination of pronotal and tegminal pigmentation, and bythe structure of the male genitalia. Mysidia nigrifrontalis sp. n. (Figs 241, 352, 461) Male: head 0-52 mm long, 0-65 mm wide; pronotum 1-30 mm wide; tegmen 6-55 mm long; wing 3-91 mmlong. Female: tegmen 8-33-8-92 mm long. Length of frons 5 times width at apex, c. 3 times width at base; ocelli very prominent; clypeus slightlyshorter than frons; rostrum extending to base of pregenital segment. Pronotal width slightly greater than12 times mid-dorsal length; fronto-lateral surfaces weakly carinate; tegula not carinate. Ocelli scarlet; fronto-lateral surfaces of pronotum each with a very large dark brown/black marking,considerably broader than eye, tapering gradually from adjacent to eye to lateral margin; abdomen with avery large, circular, scarlet spot on dorsal surface subapically; metanotum suffused smoky brown on eitherside of scutellum. Tegmen and wing whitish hyaline, veins pale brown, cross-veins edged dark smokybrown, otherwise unmarked. Male genitalia with shaft of aedeagus slender, parallel-sided in dorsal aspect, somewhat expandedsubapically in lateral aspect; ventro-lateral surfaces each with a rounded lobe bearing small obtuse spines;dorsal surfaces with a pair of very large, rounded, flap-like processes subapically; lateral surfaces each witha single, long, bifurcate process subapically. Paramere robust; apex broadly rounded; dorsal process large,situated subapically, strongly produced posteriorly; dorsal surface over basal one-third length withnumerous, short, robust spines; ventral surface over basal half length with numerous, long, slender spines. MATERIAL EXAMINED Holotype $ , Panama: Chiriqui, Fortuna, 82 15'W 8 44'N, S.v.1978 (O'Brien & Marshall) (FAMU).Paratypes. 2 $ , same data as holotype (FAMU; BMNH). This species is readily distinguished by the very striking pigmentation of the thorax andabdomen, and by the unique structure of the male genitalia. Mysidia andessp. n. (Figs 258, 369, 479) Male: head 0-73 mm long, 1-05 mm wide; pronotum 2-14 mm wide; tegmen 10-20-10-45 mm long; wing6-00 mm long. Female unknown. Length of frons 4-25 times width at apex, 3-33 times width at base; ocelli obsolete; clypeus as long asfrons; rostrum extending to base of subgenital plate. Pronotal width c. 11 times mid-dorsal length;fronto-lateral surfaces not carinate; tegula weakly carinate. Dorsal surfaces of pronotum tinged scarlet; disc of mesonotum dark brown posteriorly. Tegmen andwing smoky brown, veins brown, cells with hyaline spots medially; otherwise unmarked. male genitalia with shaft of aedeagus robust, apically truncate; dorsal surface subapically with two pairs TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 53 of short, spine-like processes. Paramere slender; apex acutely rounded; dorsal process situated attwo-thirds length, small, strongly produced posteriorly; dorsal surface basally with a rounded secondaryprocess bearing short robust spines. MATERIAL EXAMINED Holotype d", Bolivia: La Paz, Rio Beni, San Buenaventura, 270 km, 22.iv.1979 (Cooper) (BMNH).Paratypes. Bolivia: 2 cf , Prov. del Sara (Steinbach) (CM). This species is distinguished by its large size, the pigmentation of the tegmina and wing, and bythe very reduced armature of the aedeagus. Mysidia bibula sp. n. (Figs 209, 317, 428) Male: head 0-63 mm long, 0-78 mm wide; pronotum 1-70 mm wide; tegmen 7-90-8-70 mm long; wing5-10 mm long. Female: tegmen 8-80-9-40 mm long. Length of frons c. 7 times width at apex, 2-5 times width at base; ocelli very large, prominent; length ofclypeus slightly greater than that of frons; rostrum extending to base of subgenital segment. Pronotal width18 times mid-dorsal length; fronto-lateral surfaces and tegula devoid of carinae. Genae dorsad of ocelli and fronto-lateral surfaces of pronotum at level of eyes tinged orange; disc ofmesonotum brownish yellow. Tegmen and wing whitish hyaline, lacking distinct dark markings; veins pale;tinged pale smoky brown around cross-veins and, irregularly, around veins, within cells, and adjacent toposterior and apical margins. Male genitalia with shaft of aedeagus slender in lateral aspect; apex acute; dorsal surface subapicallywith a pair of large, flap-like processes, each terminating in a long, spine-like projection; lateral surfaces atthree-fifths length each with a narrow, flap-like process. Paramere apically rounded; dorsal processsituated at one-third length, slender, produced dorsally; dorsal surface at one-quarter length with a clusterof short, tooth-like spines. MATERIAL EXAMINED Holotype cf , Panama: Las Cumbres, 14.V.1978 (O'Brien) (FAMU). Paratypes. Panama: 8 cf, 15 $, same data as holotype; Tocumen; Chorrera; Villa Real (FAMU;USNM; BMNH). Though lacking distinctive external characters, this species may be distinguished by the structureof the aedeagus: Mysidia ecuadoria sp. n. (Figs 191, 301, 410) Male: head 0-52 mm long, 0-63 mm wide; pronotum 1-32 mm wide; tegmen 6-37-7-31 mm long; wing4-10 mm long. Female: tegmen 7.90 mm long. Length of frons 6-5 times width at apex, 2-66 times width at base; ocelli not prominent; clypeus slightlyshorter than frons; rostrum terminating at midlength of abdomen. Pronotal width c. 13 times mid-dorsallength; fronto-lateral surfaces not carinate; tegula with distinct carinae. Fronto-lateral surfaces of pronotum each with a broad, horizontal, brown band extending from adjacentto eye to lateral margin. Tegmen and wing whitish hyaline. Tegmen with branches of medial and cubitalveins dark brown, narrowly edged smoky brown; basal third of length mottled smoky brown. Wing withveins and cross-veins in apical half pale brown. Male genitalia with shaft of aedeagus laterally expanded over apical two-fifths length; dorsal surfacesubapically with a pair of rounded flap-like processes adjacent to midline; ventral surface with a transverseprocess at two-thirds length. Paramere slender, apex acute; dorsal process slightly distad of midlength, notproduced; dorsal surface near base with a rounded secondary process bearing numerous, long, robustspines. MATERIAL EXAMINED Holotype cf , Ecuador: Mera, 1-2. ii. 1923 (Williams) (BMNH).Paratypes. 9 cf , 13 $, same data as holotype (BMNH). This species is most readily distinguished by the male genitalia. 54 PETER S. BROOMFIELD Mysidia cheesemanisp. n. (Figs 194, 304, 413) Male: head 0-59 mm long, 0-73 mm wide; pronotum 1-45 mm wide; tegmen 6-00-6-80 mm long; wing3-40 mm long. Female: tegmen 6-40-7-60 mm long. Length of frons 4 times width at apex, 2-5 times width at base; ocelli distinct; clypeus slightly shorter thanfrons; rostrum extending to midlength of subgenital plate. Pronotal width c. 13 times mid-dorsal length;fronto-lateral carinae distinct; tegula not carinate. Head and body unmarked; abdomen occasionally tinged reddish dorsally. Tegmen and wing whitishhyaline, veins and cross-veins edged smoky brown, otherwise unmarked. Male genitalia with shaft of aedeagus laterally expanded over apical half length; dorsal surface with apair of transverse flap-like processes subapically; with a pair of long bifid processes at two-thirds length; aslender bifid process medially at two-fifths length. Paramere broad, shallowly concave apically; dorsalprocess small, situated at midlength, not posteriorly produced; ventral surface basally with a rounded lobebearing long slender spines. MATERIAL EXAMINED Holotype cf , Trinidad: Palo Seco, 21.ix.1919 (Williams) (BMNH). Paratypes. Trinidad: 69 cf, 48 $, same data as holotype; Caura, on Parthenium sp. (BMNH).Venezuela: 3 cf , Bolivar (BMNH). Guyana: 8 cf , 5 $, Blairmont (BMNH). External characters are unreliable in this species, and reference must be made to the malegenitalia. Mysidia augusta sp. n. (Figs 184, 294, 403) Male: head 0-46 mm long, 0-63 mm wide; pronotum 1-40 mm wide; tegmen 6-20-6-30 mm long; wing3-57 mm long. Female: tegmen 7-00 mm long. Length of frons 4 times width at apex, 3 times width at base; ocelli small, often obscure; length of clypeusc. three-quarters that of frons; rostrum extending to base of subgenital plate; fronto-lateral surfaces withcarinae prominent; tegula not carinate. Fronto-lateral surfaces of pronotum with carinae narrowly edged brownish. Tegmen and wing whitishhyaline, veins and cross-veins broadly edged smoky brown. Wing with irregular, smoky brown, transversebands at one-half and three-quarters length. Male genitalia with shaft of aedeagus slender in dorsal aspect, apex somewhat expanded; dorsal surfaceat midlength with a slender bifurcate process; lateral surfaces at three-quarters length each with aspine-like process extending anteriorly to base of dorsal process. Paramere robust; apex obtusely rounded;dorsal process situated slightly distad of midlength, little produced; dorsal surface at one-third length withan acute, hook-like, secondary process. MATERIAL EXAMINED Holotype cf , Peru: Tingo Maria, Los Cuevos road, 2000 ft, 10.viii.1971 (Broomfield) (BMNH).Paratypes. Peru: 3 cf , 1 $, Tingo Maria (BMNH; CAS). Though not readily distinguished by external characters, this species can be recognized by thestructure of the male genitalia. Mysidia kramerisp. n. (Figs 280, 327, 436) Male: head 0-42 mm long, 0-71 mm wide; pronotum 1-36 mm wide; tegmen 6-40 mm long; wing 3-40 mmlong. Female unknown. Length of frons 6-5 times width at apex, 3 times width at base; ocelli small, not prominent; clypeus c. aslong as frons; rostrum extending to apex of abdomen. Pronotal width 28 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Vertex with basal angles dark brown; ocelli pale; disc of mesonotum irregularly brownish. Tegmen andwing hyaline; veins pale yellow; cross-veins pale brown; both veins and cross-veins broadly edged smokybrown. Tegmen with a broad, irregular, brownish, transverse band extending from claval margin over firstfork of cubital vein to costal margin at c. one-third length; with irregular smoky brown markings around TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 55 forks of medial vein; posterior and apical margins very broadly and indistinctly pale smoky brown. Wingwith an indistinct smoky brown transverse band over radial-medial cross-vein. Male genitalia with shaft of aedeagus slender; dorsal surface subapically with a pair of large flap-likeprocesses adjacent to midline; ventral surface apically produced into a pair of rounded processes,numerous, very small, tooth-like spines laterally at three-quarters length. Paramere slender; apex obtuselyrounded, produced into an acute hook-like process; dorsal process situated subapically, well developed;dorsal surface subbasally produced, bearing numerous long robust spines. MATERIAL EXAMINEDHolotype d", Panama: Cerro Campana, 19.ix.1951 (Blanton) (USNM). Though the male genitalia, in particular the aedeagus, resemble those oilloydi, the tegminal andwing pigmentation readily distinguish this species. Mysidiu erects sp. n. (Figs 201, 310, 419) Male: head 0-63 mm long, 0-82 mm wide; pronotum 1-90 mm wide; tegmen 8-25-8-90 mm long; wing5-10 mm long. Female unknown. Length of frons 6 times width at apex, 2-5 times width at base; ocelli small; clypeus slightly longer thanfrons; rostrum extending to base of subgenital segment. Pronotal width 22 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, cross-veins broadly edged smoky brown,veins irregularly and intermittently edged smoky brown. Tegmen with an irregular, curving, brownish,transverse band at one-third length, and another over second fork of medial vein; apical and posteriormargins broadly edged pale smoky brown. Wing with apical and posterior margins irregularly edged smokybrown. Male genitalia with shaft of aedeagus very greatly laterally expanded; dorsal surface subapically with apair of large, rounded, flap-like processes extending over lateral surfaces; a pair of long acute processesmedially. Paramere very robust; apex obliquely truncate; dorsal process situated slightly basad ofmidlength, produced dorsally into a long, slender, spine-like projection. MATERIAL EXAMINED Holotype cT, Bolivia: La Paz, Rio Beni, 270 m, San Buenaventura, 22.iv.1979 (Cooper) (BMNH).Paratype. Columbia: 1 cT, Caqueta, Yuruyacu (BMNH). This species, though lacking distinctive external characters, is readily distinguished by the dorsalextension of the paramere. Mysidia maculosa sp. n. (Figs 270, 381, 491) Male: head 0-73 mm long, 0-71 mm wide; pronotum 1-60 mm wide; tegmen 8-33 mm long; wing 5-10 mmlong. Female: tegmen 8-50 mm long. Length of frons 6-5 times width at apex, c. 4 times width at base; ocelli large, prominent; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 13 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with branches of medial andcubital veins broadly edged smoky brown; basal third of length mottled smoky brownish. Wing with veinson apical half pale brownish. Male genitalia with shaft of aedeagus somewhat swollen subapically; dorsal surface subapically with apair of slender processes. Paramere very slender, constricted at one-third length, apex obtusely rounded;dorsal process situated slightly distad of one-third length, posteriorly produced; dorsal surface subbasallywith a cluster of small robust spines; ventral surface over basal half length with numerous similar spines. MATERIAL EXAMINED Holotype d", Ecuador: Mera, 1-2. ii. 1923 (Williams) (BMNH).Paratypes. Ecuador: 3 $, Mera (BMNH). This species is distinguished by its lack of tegminal and wing pigmentation, and by the verysimple armature of the aedeagus. 56 PETER S. BROOMFIELD Mysidia nebulosa (Germar) (Figs 266, 377, 487) Derbe nebulosa Germar, 1830: 56. Syntypes, BRAZIL [examined by L. B. O'Brien].[Derbe pallida Fabricius; Spinola, 1839: 379. Misidentification.]Mysidia nebulosa (Germar) Schaum, 1850: 70. Male: head 0-57 mm long, 0-74 mm wide; pronotum 1-58 mm wide; tegmen 7-20-7-85 mm long; wing4-10 mm long. Female: tegmen 8-00-8-80 mm long. Length of frons c. 5 times width at apex, 2-25 times width at base; ocelli large, distinct; clypeusone-quarter longer than frons; rostrum extending to base of subgenital plate. Pronotal width 25 timesmid-dorsal length; fronto-lateral surfaces and tegula not distinctly carinate. Pronto-lateral surfaces of pronotum occasionally pale orange at level of eye; abdomen with dorsalsurface often brown or orange. Tegmen and wing whitish hyaline; veins and cross- veins yellowish,irregularly and broadly edged smoky brown; posterior and apical margins broadly smoky brown betweenveins. Tegmen with basal third deeply mottled smoky brown. Wing unmarked. Male genitalia with shaft of aedeagus laterally expanded over apical half; dorsal surface subapically witha pair of flap-like processes, each terminating dorsally in a laterally curving spine. Paramere robust; apexvery obtusely rounded; dorsal process situated somewhat basad of midlength, very small, little producedposteriorly. MATERIAL EXAMINED Honduras: 22 C?, 13 $ , Comayagua (FAMU; BMNH). Mexico: 1 $ , Cintalapa (FAMU). Panama: 1 cf ,Canal Zone (FAMU). Costa Rica: 1 cT, 2 $ , Guan (FAMU; BMNH). Ecuador: 2 cf, 2 $ , Tema (FAMU;BMNH). Though this species was described from specimens from Brazil, the redescription above and thedrawings of the male genitalia are based on the specimens from Honduras listed above; thesespecimens were compared with the type-material by Dr Lois B. O'Brien. The species isdistinguished by the tesselated tegmen. Mysidia bizzara sp. n. (Figs 179, 288, 397) Male: head 0-46 mm long, 0-65 mm wide; pronotum 1-30 mm wide; tegmen 6-80 mm long; wing 3-90 mmlong. Female unknown. Length of frons 6-5 times width at apex; 2-66 times width at base; ocelli distinct; clypeus slightly shorterthan frons; rostrum extending to base of subgenital plate. Pronotal width c. 15 times length mid-dorsally;fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, faintly smoky brown around cross-veins.Tegmen weakly mottled Smoky brown over basal third, these markings coalescing to form a broad,irregular, transverse band over first fork of cubital vein. Wing unmarked. Male genitalia with aedeagus greatly expanded laterally, widest at one-third length, dorso-ventrallycompressed; dorsal surface subapically with a pair of flap-like processes extending over apical half.Paramere with apex very obtusely rounded; dorsal processes situated slightly distad of midlength; dorsalsurface strongly produced apically; ventral surface slightly distad of midlength with a large internallydirected lobe. MATERIAL EXAMINEDHolotype d", Bolivia: Prov. del Sara (SteinbacK) (CM). The lack of distinctive external characters makes the determination of this species largelydependent upon examination of the male genitalia. Mysidia intima sp. n. (Figs 235, 347, 455) Male: head 0-63 mm long, 0-84 mm wide; pronotum 1-65 mm wide; tegmen 7-70 mm long; wing 4-70 mmlong. Female unknown.Length of frons 4-5 times width at apex, 3 times width at base; ocelli obsolete; clypeus one-quarter longer TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 57 than frons; rostrum extending to base of subgenital plate. Pronotal width 20 times mid-dorsal length;fronto-lateral surfaces and tegula not distinctly carinate. Head and body unmarked. Tegmen and wing hyaline, veins yellow, posterior and apical marginsnarrowly and weakly pale yellowish brown, tegmen otherwise unmarked. Wing with a very indistinct paleyellowish brown transverse band over radial-medial cross-vein, and an even fainter band at two-fifthslength. Male genitalia with shaft of aedeagus slender; dorsal surface subapically with a pair of long, slightlycurving, spine-like processes adjacent to midline; lateral surfaces each with a slender spine-like processsubapically. Paramere with apex broadly rounded; dorsal process situated at midlength, slender, stronglyproduced postero-dorsally. MATERIAL EXAMINEDHolotype cf , Brazil: Rio Negro, Umarituba, 22.iv.[?] (Roman) (NR). This species is distinguished by the lack of pigmentation (even the wing markings are extremelyfaint) and by the simple spines on the aedeagus. Mysidia infedelis sp. n. (Figs 233, 344, 453) Male: head 0-67 mm long, 0-92 mm wide; pronotum 1-90 mm wide; tegmen 8-50 mm long; wing 4-70 mmlong. Female: tegmen 9-01 mm long. Length of frons c. 5 times width at apex, 3-5 times width at base; ocelli obsolete; clypeus slightly longerthan froms; rostrum extending to base of subgenital plate. Pronotal width 15 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Vertex, dorsal area and margins of frons dark brownish; dorsal and dorso-lateral surfaces of pronotumand tegula darker brownish. Tegmen bright yellowish hyaline, veins yellow, costal margin orange;anterior, apical and posterior margins regularly and narrowly dark smoky brown; claval margin unmarked.Wing with basal and claval areas hyaline, becoming tinged with yellow towards apex; veins yellow;posterior margins broadly and very distinctly edged dark smoky brown from first branch of cubital vein toapex; with a prominent, dark brownish, transverse band at level of apex of clavus, and another similarthough rather oblique band at level of radial-medial cross-vein. Male genitalia with shaft of aedeagus very slender in dorsal aspect; dorsal surface subapically with a pairof slender spines laterally, and a pair of long, somewhat overlapping, spine-like processes medially.Paramere slender; apex broadly rounded, with an acute, medially directed, hook-like projection dorsally;internal surface produced medially into a pair of acute, dorsally directed lobes; dorsal process situated atthree-fifths length, slender, strongly produced postero-dorsally; dorsal surface subbasally somewhatproduced. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. des Laranjeiras, viii.-ix.1981 (Arias) (INPA).Paratypes. 1 cf , 2 $ , same data as holotype (INPA; BMNH). This species is most readily distinguished by the yellowish pigmentation of the tegmen and by thedark transverse bands on the wing. Mysidia testacea (Fabricius) (Figs 273, 384, 494) Derbe testacea Fabricius, 1803: 82. LECTOTYPE cf, CENTRAL AMERICA (ZM), here designated [ex-amined]. Mysidia testacea (Fabricius) Westwood, 1840: 83.Mysidia citrina Walker, 1858: 98. Holotype cf , BRAZIL (BMNH) [examined]. Syn. n. Male: head 0-65 mm long, 0-73 mm wide; pronotum 1-42 mm wide; tegmen 6-43-7-65 mm long; wing4-42 mm long. Female: tegmen 8-00 mm long. Length of frons c. 5 times width at apex, 3 times width at base; ocelli distinct; clypeus slightly longer thanfrons; rostrum extending to base of subgenital plate. Pronotal width 8-5 times mid-dorsal length;fronto-lateral surfaces and tegula with carinae obsolete. Dorsal surfaces of head and body suffused scarlet. Tegmen and wing yellowish hyaline, posterior 58 PETER S. BROOMFIELD margins broadly dark fuscus. Tegmen with costal vein narrowly scarlet. Wing with a prominent fuscousband extending obliquely from radial-medial cross-vein to apex of clavus. Shaft of aedeagus slender in dorsal aspect; a pair of large flap-like processes enclosing ventral and lateralsurfaces subapically; dorsal surface subapically with a pair of curving spine-like processes adjacent tomidline. Paramere robust; apex obtusely rounded; dorsal process situated at three-fifths length, stronglyproduced posteriorly. MATERIAL EXAMINED Central America: 1 cf (lectotype of testacea) (Schmidt) (ZM). Brazil: 1 cf (holotype of citrina),Santarem (Bates) (BMNH). Guyana: 1 cf , Kartabo; 1 9, no data (BMNH). The tegmina of the testacea lectotype are missing, but its very distinctive wing pigmentation andthe structure of the genitalia indicate that it is conspecific with citrina, thus confirming thesynonymy of the latter. Mysidia diabola sp. n. (Figs 256, 367, 478) Male: head 0-55 mm long, 0-67 mm wide; pronotum 1-30 mm wide; legmen 6-00 mm long; wing 3-40 mmlong. Female unknown. Length of frons c. 7 times width at apex, twice width at base; ocelli small, distinct; clypeus slightly longerthan frons; rostrum terminating just short of base of subgenital plate. Pronotal width 12-5 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head with vertex and adjacent surfaces of genae pale orange; frons apically yellowish brown, crimsonbetween lateral carinae ventrad to midline of eyes, reddish brown basally; clypeus dark reddish brown,median and lateral carinae pale crimson. Pronotum dark brown, broadly white along posterior and ventralmargins at level of midline of eyes; tegular dark brown, ventral margins paler; mesonotum dark brownanteriorly and laterally, becoming paler towards posterior margin; ventro-lateral surfaces pale crimson;lateral margins of scutellum and dorsal surface of abdomen dark brown. Tegmen and wing yellowishhyaline, veins yellow, cross-veins broadly edged dark smoky brown. Tegmen with posterior and apicalmargins narrowly edged smoky brown; with a prominent, narrow, brown band extending from costal toposterior margin at one-third length; area around apices of branches of cubital vein broadly and irregularlydark brown; with a large, irregular, dark brown marking extending from costal margin to first branch ofmedial vein at between two-thirds and three-quarters length; apex smoky brown between first branch ofradial vein and seventh branch of medial vein. Wing with a narrow, irregular, transverse, smoky brownband extending from first fork of cubital vein to posterior margin; a very broad, dark brown band extendingfrom costal to posterior margins between one-half and three-quarters length; apex irregularly dark brownaround apices of radial and medial veins. Shaft of aedeagus with dorsal surface unarmed; lateral surfaces at c. two-thirds length each with atriangular flap-like process. Paramere slender; apex narrowly rounded; dorsal process situated slightlybasad of three-quarters length, large, strongly produced dorsally and posteriorly; dorsal surface subapical-ly produced, bearing several short robust spines. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, 5.viii.l981 (Arias) (INPA).Paratype. Brazil: 1 cf , Amazonas (BMNH). This species is readily distinguished by the pigmentation of the head, pronotum, tegmen andwing; the proportions of the head are also distinctive. Mysidia Iloydisp. n. (Figs 264, 375, 485) Male: head 0-63 mm long, 0-76 mm wide; pronotum 1-36 mm wide; tegmen 6-80 mm long; wing 3-80 mmlong. Female unknown. Length of frons 6-5 times width at apex, 3-25 times width at base; ocelli small, not prominent; clypeus aslong as frons; rostrum terminating at level of hind coxae. Pronotal width 13 times mid-dorsal length;fronto-lateral carinae distinct; tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale brown. Tegmen with a narrow, TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 59 pale brownish, transverse band at one-third length; another, less distinct, at c. midlength. Wing with anindistinct, pale brown, transverse band over first fork of cubital vein, and another over radial-medialcross-vein. Shaft of aedeagus slender in lateral aspect, somewhat laterally expanded; dorsal surface subapically witha pair of flap-like processes with apices acute and spine-like. Paramere broad; apex obtusely rounded;dorsal process situated at two-thirds length, produced postero-dorsally; dorsal surface somewhat producedsubbasally into a rounded lobe bearing robust spines. MATERIAL EXAMINEDHolotype cf , Brazil: Para, Jabaty, v.1924 (Williams) (BMNH). Though this species closely resembles josianna, the proportions of the head and the structure ofthe male genitalia render it distinct. Mysidia isteria sp. n. (Figs 211, 321, 430) Male: head 0-61 mm long, 0-90 mm wide; pronotum 1-95 mm wide; tegmen 8-50 mm long; wing 4-70 mmlong. Female unknown. Length of f rons c. 5 times width at apex , c. 2 5 times width at base ; ocelli small , obscure ; clypeus one-fifthlonger than frons; rostrum extending to base of subgenital plate. Pronotal width c. 19 times mid-dorsallength; fronto-lateral carinae absent; tegula prominently carinate. Pronto-lateral surfaces of pronotum broadly pale orange at level of eyes. Tegmen and wing whitishhyaline; veins and cross-veins yellowish, cross-veins narrowly edged pale smoky brown. Tegmen with anindistinct, narrow, transverse, smoky brown band at one-third length; another, broader, but even fainterband immediately distad of midlength. Wing unmarked. Shaft of aedeagus very short and broad; apex truncate; a pair of large flap-like processes extending overdorsal and lateral surfaces from apex almost to midlength, each bearing a single, long, spine-like processdorsally. Paramere very large in relation to aedeagus, slender, apex acute; dorsal process situated atslightly less than one-third length, large, not posteriorly produced; dorsal surface distad of midlengthstrongly produced and directed towards midline. MATERIAL EXAMINEDHolotype cf , Peru: Iquitos 5 km, Marine road, 24.xi.1972 (Woldd) (FAMU). In the structure of the male genitalia this species most closely resembles panamensis, but in theproportions of the head and body, and in the detailed structure of the aedeagus, it is quitedistinct. Mysidia fuscomaculata sp. n. (Figs 263, 374, 484) Male: head 0-54 mm long, 0-60 mm wide; pronotum 1-47 mm wide; tegmen 6-80 mm long; wing 3-60 mmlong. Female unknown. Length of frons 9 times width at apex, c. twice width at base; ocelli obscure; clypeus as long as frons;rostrum extending to base of abdomen. Pronotal width 20 times mid-dorsal length; fronto-lateral carinaedistinct. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale. Tegmen with a broad, smokybrown, transverse band extending from costal margin to apex of clavus; posterior margin very pale smokybrown. Wing very pale smoky brown on apical and posterior margins. Shaft of aedeagus slender; dorsal surface subapically with a pair of flap-like processes laterally, eachterminating in a long curving spine; lateral surfaces each with a small spine subapically. Paramere robust;dorsal process large, situated at two-thirds length, somewhat produced; dorsal surface at one-quarterlength with a large rounded secondary process bearing numerous, large, robust spines. MATERIAL EXAMINEDHolotype cf , Panama: Cerro Campana, 2700 ft, 23.V.1978 (O'Brien & Marshall) (FAMU). This species is distinguished by the tegminal pigmentation, apically very narrow frons, and bythe structure of the male genitalia. 60 PETER S. BROOMFIELD Mysidia adamare sp. n. (Figs 221, 332, 441) Male: head 0-63 mm long, 0-84 mm wide; pronotum 1-90 mm wide; legmen 8-84 mm long; wing 5-10 mmlong. Female unknown. Length of frons 5-5 times width at apex, 2-33 times width at base; ocelli distinct; clypeus one-third longerthan frons; rostrum extending to base of subgenital plate. Pronotal width 13 times mid-dorsal length,fronto-lateral surfaces not carinate; tegula distinctly carinate. Pronto-lateral surfaces of pronotum each with a broad, horizontal, orange band extending from adjacentto eye to lateral margin; fore and mid femora narrowly scarlet apically. Tegmen and wing whitish hyaline,veins yellow. Tegmen with a faint, brownish, transverse band at one-seventh length; a broader, moreprominent band slightly distad of one-quarter length; weaker, indistinct, bands over first and second forksof medial vein. Wing with a very pale, brownish, transverse band at midlength and another overradial-medial cross-vein. Shaft of aedeagus slender; lateral surfaces each with a large flap-like process extending over dorsalsurface and slightly overlapping at midline; dorsal surface subapically with a pair of large flap-likeprocesses medially. Paramere extremely robust, almost circular in lateral aspect; dorsal processes small,situated slightly basad of midlength; dorsal surface subbasally with a large, rounded, secondary processbearing numerous robust spines. MATERIAL EXAMINEDHolotype C?, Brazil: Mato Grosso, 1250'S 5147'W, 16.iv.1968 (Richards) (BMNH). This species is most readily distinguished by reference to the male genitalia, in particular to thevery broadly rounded paramere. Mysidia pallescens MetcalfMysidia pallescens Metcalf, 1938: 315. Holotype 9> PANAMA (MCZ) [examined]. Female: head 0-73 mm long, 0-88 mm wide; pronotum 2-05 mm wide; tegmen 8-70-9-35 mm long; wing5-30 mm long. Male unknown. Length of frons 5 times width at apex, 2-5 times width at base; ocelli small, distinct; clypeus c. as long asfrons; rostrum extending almost to apex of subgenital plate. Pronotal width 34 times mid-dorsal length,fronto-lateral carinae absent; tegula distinctly carinate basally. Genae adjacent to eyes brownish; posterior angle of mesonotal disc brownish. Tegmen and wing whitishhyaline, veins brownish yellow, veins and cross-veins irregularly edged smoky brown. Tegmen over c.basal third and over subcostal and medial cells mottled brownish; a distinct, smoky brown, transverse bandover first fork of cubital vein; another, more irregular and less distinct, pale brownish, transverse band oversecond fork of medial vein; area between these bands, and between second band and apex, indistinctly andirregularly mottled pale brownish. Wing with a pale brown transverse band over first fork of cubital vein. MATERIAL EXAMINED Panama: 2 $ (holotype and paratype), Canal Zone, Barro Colorado, 17.vii.1924 (Banks) (MCZ); 3 9,Canal Zone (FAMU; BMNH). This species may be distinguished by the tegminal pigmentation, and by the relatively large headand broad pronotum. Mysidia insania sp. n. (Figs 265, 376, 486) Male: head 0-67 mm long, 0-75 mm wide; pronotum 1-78 mm wide; tegmen 8-84 mm long; wing 4-93 mmlong. Female unknown. Length of frons 6 times width at apex, c. 2-5 times width at base; ocelli not prominent; clypeus one-fifthlonger than frons; rostrum terminating slightly posterior to hind coxae. Pronotal width 28 times mid-dorsallength, fronto-lateral surfaces not carinate; tegula distinctly carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with basal one-fifth lengthirregularly smoky brown; with a brownish transverse band at one-third length; another, more irregularband over medial-cubital cross-vein; and a third slightly distad of midlength, from thence to apex very pale TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 61 brownish; posterior margin tinged smoky brown. Wing with a very irregular, smoky brown, transverseband at midlength; apical third smoky brown. Shaft of aedeagus slender, becoming greatly laterally expanded subapically; dorsal surface subapicallywith a pair of large, adpressed, flap-like processes, each terminating in a large curving spine. Paramererobust; apex broadly rounded; dorsal process situated slightly basad of midlength, weakly producedposteriorly. MATERIAL EXAMINEDHolotype cf , Ecuador: Morona, Santiago, Cordillera de Cutucu, 1000 m, 21.X.1978 (Cooper) (BMNH). Though rather similar to some other species in external characters, the male genitalia showinsania to be quite distinct. Mysidiapanamensissp. n. (Figs 210, 320, 429) Male: head 0-53 mm long, 0-76 mm wide; pronotum 1-64 mm wide; tegmen 8-00 mm long; wing damaged.Female unknown. Length of frons 4-5 times width at apex, 4 times width at base; ocelli very small, distinct; clypeus slightlylonger than frons, rostrum extending to base of subgenital plate. Pronotal width 26 times mid-dorsallength, fronto-lateral carinae absent; tegula distinctly carinate. Head and body unmarked. Tegmen and wing hyaline. Tegmen with a very faint, smoky brown,transverse band over first fork of cubital vein; another immediately distad of midlength; cross-veins verynarrowly edged smoky brown. Wing with cross-veins very indistinctly edged smoky brown. Shaft of aedeagus greatly expanded dorso-ventrally over apical half length; dorsal surface subapicallywith a pair of very large, hooked, processes. Paramere slender; apex acutely rounded; dorsal processsituated at one-third length, not posteriorly produced. MATERIAL EXAMINED Holotype cf , Panama: 6 miles E. Porto Bello, 6.H.1930 (Zschokke) (CAS).Paratype. Panama: 1 cf , Darien, Santa Fe (FAMU). This species is readily distinguished by the combination of hyaline tegmina and wings, and by themassive structure of the aedeagus. Mysidia formosa sp. n. (Figs 246, 357, 467) Male: head 0-53 mm long, 0-67 mm wide; pronotum 1-55 mm wide; tegmen 7-10 mm long; wing 4-42 mmlong. Female unknown. Length of frons c. 1 times width at apex, 2-25 times width at base; ocelli very large, prominent; clypeusone-quarter longer than frons; rostrum extending to base of subgenital plate. Pronotal width c. 12 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Genae anterior to eyes crimson; pronotum with posterior margins crimson. Tegmen and wing almosthyaline, veins dark brown. Tegmen weakly smoky brown over basal one-fifth length; with a broad, palesmoky brown, transverse band over first fork of cubital vein. Wing with a large, irregular, pale smokybrown spot on anal area at one-fifth length; first fork of cubital vein and radial-medial cross-vein broadlyand weakly edged smoky brown. Shaft of aedeagus very slender; apex strongly recurved over dorsal surface, partially obscuring a singlepair of large flap-like processes, each of which bears a hook-like spine on the dorsal surface subapically.Paramere slender, strongly constricted at midlength; apex obtusely and irregularly rounded, bearing ahook-like process on the internal surface; dorsal process situated at four-fifths length, strongly produceddorsally and posteriorly; dorsal surface between one-quarter and one-third length with a prominentsecondary process bearing numerous, short, robust spines. MATERIAL EXAMINED Holotype cf , Venezuela: Carabobo, San Esteban, 8 km SE. Puerto Cabello, 100 m, 7.V.1978 (O'Brien &Marshall) (FAMU). This species is readily distinguished by the unique pigmentation of the head, pronotum, tegmenand wing, and by the structure of the male genitalia. 62 PETER S. BROOMFIELD Mysidia whimperisp. n. (Figs 272, 383, 493) Male: head 0-53 mm long, 0-57 mm wide; pronotum 1-22 mm wide; tegmen 6-40 mm long; wing 3-40 mmlong. Female unknown. Length of frons c. 6 times width at apex, c. 4 times width at base; ocelli small, distinct; clypeus as long asfrons; rostrum extending to base of subgenital plate. Pronotal width c. 12 times mid-dorsal length,fronto-lateral surfaces and tegula not distinctly carinate. Pronto-lateral surfaces of pronotum adjacent to eyes, and disc of mesonotum, pale reddish brown.Tegmen and wing whitish hyaline. Tegmen with a faint, smoky brown, transverse band over first fork ofcubital vein. Wing with an obscure, smoky brown, transverse band at midlength. Shaft of aedeagus subapically expanded into a pair of flap-like processes extending over lateral anddorsal surfaces, each terminating antero-dorsally in a long spine. Paramere with apex obtusely rounded;dorsal process situated at three-fifths length, posteriorly produced; dorsal surface at one-fifth length with asmall rounded secondary process bearing numerous short spines. MATERIAL EXAMINEDHolotype cf , Ecuador: 8 km NE. Puyo, 28.iv.1978 (O'Brien & Marshall) (FAMU). The markings of the tegmina and wings are extremely faint, the species is therefore most readilydetermined by reference to the male genitalia. Mysidia stali sp. n. (Figs 200, 312, 421) Male: head 0-61 mm long, 0-75 mm wide; pronotum 1-63 mm wide; tegmen 8-00 mm long; wing 4-10 mmlong. Female unknown. Length of frons 5-5 times width at apex, c. 3 times width at base; ocelli distinct; clypeus slightly longerthan frons; rostrum extending slightly beyond hind coxae. Pronotal width 13 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins yellow. Tegmen with costal marginbroadly dark brown at c. one-quarter length; with a broad brownish band extending transversally over firstfork of cubital vein to posterior margin. Wing with a very faint, indistinct, brownish, transverse bandslightly distad of one-third length; and a more prominent band over radial-medial cross-vein. Shaft of aedeagus slender; dorsal surface subapically with a pair of long spine-like processes; lateralsurfaces subapically each with a spine-like process; ventral surface with a slender, narrowly bifurcate,process subbasally. Paramere with apex acute; dorsal process situated at c. midlength, produced dorsally;dorsal surface with a robust hook-like process subapically, subbasally with a low, rounded, secondaryprocess bearing numerous short spines; ventral surface subbasally with very numerous, small, tooth-likespines. MATERIAL EXAMINED Holotype cf, Brazil: Amazon, Rio Autaz (Roman) (NR).Paratypes. 1 cT, 1 $, same data as holotype (NR; BMNH). This species is distinguished by the tegminal pigmentation, the proportions of the head andpronotum, and by the structure of the male genitalia, especially the paramere. Mysidia clava sp. n. (Figs 242, 353, 462) Male: head 0-48 mm long, 0-65 mm wide; pronotum 1-10 mm wide; tegmen 5-80 mm long; wing 3-40 mmlong. Female unknown. Length of frons 7 times width at apex, 2-33 times width at base; ocelli small, obscure; clypeus c. as long asfrons; rostrum extending to base of subgenital plate. Pronotal width c. 18 times mid-dorsal length;fronto-lateral surfaces and tegula with carinae weak or obsolete. Fronto-lateral surfaces of pronotum each with a narrow, bright orange band extending horizontally fromadjacent to midline of eye to lateral margin. Tegmen and wing whitish hyaline. Tegmen with a narrow, veryfaint, transverse, smoky brown band over first fork of claval vein; an irregular, pale, smoky brown areaadjacent to apex of clavus. Wing with posterior and apical margins pale brownish grey. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 63 Shaft of aedeagus very slender, gradually tapering over c. basal half length , somewhat expanded towardsapex; ventral surface bearing numerous transverse rows of small blunt spines; lateral margins each with along hooked process at three-quarters length; dorsal surface with a pair of slender processes at two-thirdslength, a large, flap-like, bifurcate process at midline. Paramere slender; dorsal process situated slightlydistad of midlength, posteriorly produced; dorsal surface subbasally with a rounded secondary processbearing numerous robust spines. MATERIAL EXAMINEDHolotype cT, Brazil: Belem, Para, v.1924 (Williams) (BMNH). Its small size, relative lack of tegminal pigmentation, and the structure of the male genitaliareadily distinguish this species. Mysidia simpla sp. n. (Figs 222, 333, 442) Male: head 0-53 mm long, 0-73 mm wide; pronotum 1-43 mm wide; tegmen 6-80-7-40 mm long; wing4-16 mm long. Female: tegmen 7-60-8-50 mm long. Length of frons 6-25 times width at apex, 3-33 times width at base; ocelli small, obscure; clypeus as longas frons; rostrum extending to base of subgenital plate. Pronotal width c. 14 times mid-dorsal length,fronto-lateral carinae distinct; tegulae not carinate. Lateral carinae of frons oft' ;n brownish; apex of disc of mesonotum occasionally reddish. Tegmen andwing pale brownish hyaline, veing pale brown. Tegmen pale brown basally; a brownish transverse band atone-third length; a fainter band at midlength. Wing with a pale brownish transverse band over first fork ofcubital vein, another over radial-medial cross-vein. Shaft of aedeagus slender; dorsal surface subapically with a pair of parallel flap-like processes. Paramereslender; dorsal process situated at three-fifths length, small, posteriorly produced; dorsal surface sub-basally with a small conical secondary process bearing robust spines. MATERIAL EXAMINED Holotype cT, Ecuador: Tena, 4.iv.l923 (Williams) (BMNH).Paratypes, Ecuador: 1 cf , 2 $ , 18 km S. Tena (FAMU; BMNH). Though this species closely resembles lloydi, the proportions of the head, and the very reducedarmature of the aedeagus render it distinct. Mysidia nigrithorax sp. n. (Figs 231, 342, 451) Male: head 0-57 mm long, 0-82 mm wide; pronotum 1-50 mm wide; tegmen 7-65 mm long; wing 3-80 mmlong. Female: tegmen 8-00 mm long. Length of frons c. 5 times width at apex, 3 times width at base; ocelli prominent; clypeus slightly longerthan frons; rostrum extending almost to base of subgenital plate. Pronotal width 10 times mid-dorsallength; fronto-lateral surfaces and tegulae not carinate. Ocelli scarlet; disc of mesonotum dark brown; dorsal surface of abdomen brown. Tegmen and wingwhitish hyaline, posterior and apical margins broadly smoky brown. Tegmen with a narrow, transverse,brownish band over first fork of cubital vein, another much broader band at midlength. Wing with anarrow, transverse, brownish band at midlength, apex broadly smoky brown. Shaft of aedeagus somewhat expanded subapically; dorsal surface subapically with a pair of very longspine-like processes, a pair of much shorter spines, and laterally a pair of narrow flap-like processes.Paramere very slender; apex narrowly rounded; dorsal process situated at three-fifths length, slender,posteriorly produced; dorsal surface subbasally with a rounded secondary process bearing numerous shortrobust spines; ventral surface at midlength with a low flap-like process. MATERIAL EXAMINED Holotype cf , Peru: Tingo Maria, 13.vii.1968 (O'Brien) (FAMU).Paratypes. 1 cf , 3 $ , same data as holotype (FAMU; BMNH). This species is distinguished by the pigmentation and by the structure of the male genitalia. 64 PETER S. BROOMFIELD Mysidia subfusca MetcalfMysidia subfusca Metcalf, 1938: 315. Holotype $, PANAMA (MCZ) [examined]. Female: head 0-69 mm long, 0-88 mm wide; pronotum 1 -72 mm wide; tegmen 8-85 mm long; wing 5-00 mmlong. Male unknown. Length of frons 8 times width at apex, 3-5 times width at base; ocelli large, distinct; clypeus c. as long asfrons; rostrum extending almost to base of subgenital plate. Pronotal width c. 13 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Head and pronotum weakly tinged reddish. Tegmen and wing whitish hyaline, veins and cross-veinsyellowish brown. Tegmen irregularly smoky brown over basal one-fifth length; with a narrow, smokybrown, transverse band immediately distad of first fork of cubital vein; broadly, irregularly, and faintlybrownish from immediately distad of second fork of cubital vein to c. three-fifths length; apical one-thirdlength irregularly pale smoky brownish. Wing with a very pale, smoky brown, transverse band over firstfork of cubital vein, another over radial-medial cross-vein. MATERIAL EXAMINEDHolotype $, Panama: C.Z., Barro Colorado, 26.vi.1924 (Banks) (MCZ). This species is very close to pallescens from which, in the absence of male genitalia forcomparison, it is most readily distinguished by the proportions of the head and pronotum, andby the two transverse bands on the tegmen. Mysidia estfarchina sp. n. (Figs 212, 322, 431) Male: head 0-63 mm long, 1-01 mm wide; pronotum 2-00 mm wide; tegmen 9-35-10-20 mm long; wing5-10 mm long. Female unknown. Length of frons c. 6 times width at apex, c. 2-5 times width at base; ocelli small, distinct; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 12 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing basally hyaline, veins pale, posterior and apical marginsbroadly and irregularly smoky brown between apices of veins, cross- veins narrowly edged smoky brown.Tegmen with an irregular, pale brownish, transverse band slightly basad of one-third length; another, morebroken and paler band at midlength. Wing lacking distinct tranverse markings; branches of veinsirregularly edged smoky brown. Shaft of aedeagus massively expanded subapically; dorsal surface subapically with a pair of very large,flap-like, apically acute and strongly diverging processes adjacent to midline; lateral surfaces subapicallyeach with a large flap-like process extending over ventral surface. Paramere robust; apex acute; dorsalprocess situated slightly distad of one-third length, not posteriorly produced; dorsal surface stronglyproduced and inclined towards midline; ventro-lateral surface subbasally with very numerous, tiny,tooth-like spines. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, 30.vii.1981 (Arias) (INPA).Paratypes. 5 cf, same data as holotype (INPA; BMNH; NR). This species is distinguished by the pigmentation of the tegmina and wing, and by the structure ofthe male genitalia. Mysidia subfasciata Westwood Mysidia subfasciata Westwood, 1840: 83. LECTOTYPE (? sex), BRAZIL (BMNH), here designated[examined] . Head 0-62 mm long, 0-80 mm wide; pronotum 1-78 mm wide; tegmen 8-75 mm long; wing 4-85 mm long.The abdomen is missing, and the sex of the unique type-specimen is therefore unknown. Length of frons 10 times width at apex, 3-33 times width at base; ocelli small, distinct; clypeus slightlyshorter than frons; rostrum damaged. Pronotal width 28 times mid-dorsal length; fronto-lateral surfacesand tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins and cross-veins yellowish. Tegmenwith an indistinct very pale brown spot over cubital vein at midlength between base and first fork; a faint, TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 65 irregular, pale brown, transverse band over first fork of cubital vein; a very faint and indistinct, pale browntransverse band over first fork of medial vein. Wing lacking distinct markings. MATERIAL EXAMINEDLectotype, Brazil: Para [?] (Burchell [?]) (BMNH). This is the only specimen available for study, and the abdomen is missing and the rostrumdamaged; it bears Westwood's handwritten determination label. The species is distinguished bythe very narrow frons and the tegminal markings. Mysidia fasciata Metcalf (Figs 225, 336, 446)Mysidia fasciata Metcalf, 1938: 314. Holotype $, PANAMA (MCZ). Male: head 0-67 mm long, 0-80 mm wide; pronotum 1-68 mm wide; legmen 7-20-7-65 mm long; wing4-10 mm long. Female: tegmen 7-20-8-50 mm long. Length of frons 7 times width at apex, c. 3 times width at base; ocelli distinct; clypeus one-sixth longerthan frons; rostrum extending to base of subgenital plate. Pronotal width 10 times mid-dorsal length;fronto-lateral surfaces and carinae not carinate. Ocelli narrowly edged crimson; disc of mesonotum often deep yellowish brown, occasionally with anarrow, longitudinal, dark brown band on either side of midline; fronto-lateral surfaces of pronotum rarelydistinctly crimson; dorsal surface of abdomen rarely tinged reddish. Tegmen and wing whitish hyaline.Tegmen with veins and cross-veins brown; costal area pale smoky brown; claval area dark brown; anarrow, dark brown, transverse band over first fork of cubital vein; a broader, paler, less distinct, smokybrown, transverse band between first and second forks of medial vein; slightly less than apical one-halflength entirely smoky greyish brown. Wing with posterior and apical margins broadly smoky brown; a darkbrownish transverse band at c. three-quarters length. Shaft of aedeagus slender; dorsal surface subapically with a pair of large, finely serrated, flap-likeprocesses extending to just short of midlength; ventro-lateral surfaces each with a ventrally directedflap-like process at three-quarters length. Paramere basally slender, broadening abruptly to very obtuselyrounded apex; dorsal process situated slightly distad of midlength, produced postero- ventrally. MATERIAL EXAMINED Allotype cf , Panama: C.Z., Barro Colorado, 21. vi. 1924 (Banks) (MCZ).Panama: 14 cT, 7 $ , various localities in Canal Zone (USNM; CAS; FAMU; BMNH). The genitalia of the allotype are damaged , and a preparation was not made of this specimen . Thetegminal and wing markings of this species are distinctive. Mysidia douglasi sp. n. (Figs 178, 287, 396). Male: head 0-50 mm long, 0-67 mm wide; pronotum 1-32 mm wide; tegmen 6-97 mm long; wing 3-77 mmlong. Female unknown. Length of frons c. 8 times width at apex, c. 3 times width at base; ocelli large, distinct; clypeus slightlylonger than frons; rostrum extending to apex of abdomen. Pronotal width 21 times mid-dorsal length;fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with a pale brownish transverseband over first fork of cubital vein; another similar band at midlength. Wing with a pale brown transverseband over first fork of cubital vein and another over radial-medial cross-vein. Shaft of aedeagus greatly expanded laterally; dorsal surface subapically with a pair of long, adpressed,flap-like processes, each terminating in an acute spine; a pair of small triangular processes at midlineimmediately distad of midlength. Paramere with apex obtusely rounded; dorsal process large, situatedslightly distad of midlength, posteriorly produced; ventral surface at two-thirds length with a smallhook-like process. MATERIAL EXAMINEDHolotype cT, Panama: Gatun Lake, x.1931 (Zschokke) (CAS). The external characters alone are not considered sufficient for the positive determination of thisspecies; reference should therefore be made to the male genitalia. 66 PETER S. BROOMFIELD Mysidia knight isp. n. (Figs 203, 313, 422) Male: head 0-53 mm long, 0-68 mm wide; pronotum 1-15 mm wide; tegmen 6-05 mm long; wing 3-40 mmlong. Female unknown. Length of frons 4-5 times width at apex, c. 3 times width at base; ocelli very large, prominent; clypeusslightly longer than frons; rostrum extending to apex of abdomen. Pronotal width 10 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale. Tegmen with a prominent,broad, dark brown, transverse band extending from costal margin to claval suture over first fork of cubitalvein; a paler, less distinct, more irregular, smoky brown band over radial-medial cross-vein. Wing with afaint, irregular, pale smoky brown, transverse band at midlength and at three-quarters. Shaft of aedeagus very slender; dorsal surface subapically with two pairs of long spine-like processes;ventral surface with two longitudinal rows of small obtuse spines. Paramere slender, medially constricted,apex obtusely rounded; dorsal process situated at two-thirds length, strongly produced posteriorly; dorsalsurface subbasally with a large rounded secondary process bearing numerous small robust spines. MATERIAL EXAMINEDHolotype cf , Brazil: Mato Grosso, 1249'S 5145'W, 18.xii.1968 (Knight) (BMNH). Very similar to pulchella, this species is distinguished by the pale apex to the wing and by thestructure of the paramere. Mysidia pulchella sp. n. (Figs 176, 285, 394) Male: head 0-53 mm long, 0-70 mm wide; pronotum 1-26 mm wide; tegmen 6-40 mm long; wing 3-57 mmlong. Female unknown. Length of frons slightly greater than 4-5 times width at apex, 3 times width at base; ocelli large,prominent; clypeus as long as frons; rostrum extending little beyond hind coxae. Pronotal width 12 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Head and body unmarked; ocelli broadly and irregularly edged scarlet. Tegmen and wing whitishhyaline, veins pale. Tegmen with a prominent, broad, dark brown, transverse band extending from costalto posterior margins over first fork of cubital vein; another over radial-medial cross- vein. Wing with a pale,irregular, smoky brown, transverse band extending from first fork of cubital vein to posterior margin;apical one-quarter length broadly pale smoky brown. Shaft of aedeagus slender; dorsal surface subapically with two pairs of large spine-like processes; ventralsurface at two-thirds length with a cluster of small obtuse spines. Paramere very slender, stronglyconstricted subbasally; dorsal process small, situated somewhat basad of midlength, little producedposteriorly; dorsal surface subbasally with a slender secondary process bearing numerous small robustspines; ventral surface at one-third length with a slender process bearing numerous robust spines. MATERIAL EXAMINEDHolotype cT, Brazil: Mato Grosso, 1250'S 5147'W, 17.X.1968 (Richards) (BMNH). Though very similar to the preceding species, pulchella is distinguished by the markings of thewing and by the structure of the paramere. Mysidia distincta sp. n. (Figs 271, 382, 492) Male: head 0-73 mm long, 0-88 mm wide; pronotum 2-00 mm wide; tegmen 9-40-10-20 mm long; wing5-95 mm long. Female: tegmen 11-56 mm long. Length of frons 5-5 times width at apex, c. 3 times width at base; ocelli small, distinct; clypeus c. one-fifthlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 47 times mid-dorsallength, fronto-lateral carinae absent; tegula weakly carinate. Genae brown level with anterior margins of eyes; ocelli narrowly edged scarlet; disc of mesonotumcommonly dark brown. Tegmen and wing whitish hyaline. Tegmen pale smoky brown basally; a distinct,brownish, transverse band at c. one-third length, another at midlength; area between these bands with a TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 67 much fainter, indistinct, transverse band medially; posterior margin very pale brownish. Wing weakly andindistinctly pale brown over cross-veins; posterior margin very faintly pale brownish. Shaft of aedeagus broadly laterally expanded over apical one-half length; dorsal surface with a pair offlap-like processes over apical two-fifths length, a pair of very long hook-like processes subapically.Paramere robust, apically truncate; dorsal process situated somewhat basad of midlength, stronglyproduced posteriorly. MATERIAL EXAMINEDHolotype cf , Peru: Callanga (BMNH).Paratypes. 1 cf , 2 $?, same data as holotype (BMNH). The dark brown mesonotal disc, the tegminal pigmentation, and the structure of the malegenitalia distinguish this species. Mysidia hengistsp. n. (Figs 252, 363, 474) Male: head 0-57 mm long, 0-76 mm wide; pronotum 1-53 mm wide; tegmen 7-32-7-90 mm long; wing4-00 mm long. Female: tegmen 8-15 mm long. Length of frons, slightly less than 7 times width at apex, slightly less than twice width at base; ocelli small,obscure; clypeus one-quarter longer than frons; rostrum extending to base of subgenital plate. Pronotalwidth 37 times mid-dorsal length; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing hyaline, veins pale yellow. Tegmen with a smoky browntransverse band extending from costal margin to apex of anal vein between first and second forks of cubitalvein; another much fainter and irregular band extending from medial-cubital cross-vein to posteriormargin; another pale band extending from costal to posterior margins at level of second fork of medialvein; posterior margin narrowly and very indistinctly tinged smoky brown between branches of cubitalvein. Wing with a pale smoky brown, transverse band extending from costal to posterior margin at level offirst fork of cubital vein; another slightly darker band extending obliquely from around radial-medialcross-vein to posterior margin; posterior and apical margins faintly smoky brown between veins. Shaft of aedeagus broad; lateral surfaces subapically each with a large flap-like process extending overdorsal surface and strongly overlapping at mid-dorsal line, each process bearing a long, slightly curvingspine on its antero-dorsal surface, and a somewhat shorter spine on its posterior margin. Paramere veryrobust; apex very obtusely rounded, somewhat produced dorsally; dorsal process situated at midlength,little produced. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, 30.vii.1981 (Arias) (INPA).Paratypes. 2 cf , 1 <j>, same data as holotype (INPA; BMNH). This species is distinguished by the pigmentation of the tegmina and wings, and by the structureof the aedeagus. Mysidia josiannasp. n. (Figs 240, 351, 460) Male: head 0-59 mm long, 0-80 mm wide; pronotum 1-75 mm wide; tegmen 7-10-8-15 mm long; wing4-10 mm long. Female unknown. Length of frons 5-5 times width at apex, 2-5 times width at base; ocelli small, distinct; clypeus as long asfrons; rostrum extending to subgenital plate. Pronotal width 35 times mid-dorsal length; fronto-lateralsurfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale brown. Tegmen pale smokybrown over basal half length; a broad, dark brown, transverse band immediately basad of second fork ofcubital vein; another somewhat fainter band over medial-cubital cross-vein; occasionally another very faintand indistinct band immediately distad of midlength; posterior margin between branches of cubital veinand first branch of medial vein edged smoky brown. Wing with cross-veins and forks of veins irregularlyedged pale smoky brown; posterior margin between branches of cubital vein and medial vein broadlysmoky brown. Shaft of aedeagus laterally expanded over apical three-fifths length; lateral surfaces subapically eachwith a large flap-like process bearing a long acute spine; dorsal surface with a single, medial, spine-like 68 PETER S. BROOMFIELD process slightly distad of midlength. Paramere slender; dorsal process situated slightly basad of midlength,well developed. MATERIAL EXAMINED Holotype cf , Trinidad: Arima Valley, Arima 10 miles, 15.vii.1976 (Noyes) (BMNH). Paratypes. Trinidad: 3 cf, Mount Tucuche and Aripo Valley (BMNH). Brazil: 1 cf , Para, Jabaty(BMNH). This species, lacking definitive external characters, is readily distinguished by the structure ofthe male genitalia. Mysidia pseudoerecta sp. n. (Figs 202, 311, 420) Male: head 0-65 mm long, 0-90 mm wide; pronotum 1-90 mm wide; legmen 8-50 mm long; wing 5-00 mmlong. Female unknown. Length of frons 6 times width at apex, 2-5 times width at base; ocelli small, obscure; clypeus slightlylonger than frons; rostrum extending somewhat beyond posterior coxae. Pronotal width 25 timesmid-dorsal length; fron to-lateral surfaces and tegula not carinate. Disc of mesonotum with a roughly circular dark brown spot medially. Tegmen and wing clear hyaline,veins pale yellow. Tegmen with posterior and apical margins broadly pale smoky brown; a pale smokybrown spot in angle of anal veins at point of fusion; another similar spot on cubital vein at midlengthbetween base and first fork; a distinct brown transverse band immediately basad of one-third length; apaler, irregular, less distinct band at level of medial-cubital cross-vein; a rather more distinct transverseband at level of second fork of medial vein. Wing with posterior and apical margins broadly pale smokybrown; a weak, very irregular, broken, smoky brown transverse band somewhat basad of midlength; aneven less distinct smoky marking around medial-cubital cross-vein, extending to posterior margin; atransverse band extending from radial-medial cross-vein to posterior margin. Shaft of aedeagus considerably expanded laterally over apical one-third length; lateral surfacessubapically each with a large flap-like process extending over dorsal surface and terminating anteriorly in aslender spine. Paramere robust, apex broadly rounded; dorsal process situated at c. midlength, slender,greatly produced vertically, apex acute and narrowly recurved. MATERIAL EXAMINEDHolotype cf , Brazil: Amazonas, P. das Laranjeiras, viii.-ix.1981 (Arias) (INPA). The paramere and aedeagus of this species closely resemble those of erecta, but the tegminal andwing pigmentation are quite distinct. Mysidia perspicua sp . n . (Figs 267, 378, 488) Male: head 0-61 mm long, 0-78 mm wide; pronotum 1-47 mm wide; tegmen 7-65 mm long; wing 3-83 mmlong. Female unknown. Length of frons 8 times width at apex, 2-5 times width at base; ocelli small, obscure; clypeus one-quarterlonger than frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 14 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Pronto-lateral surfaces of pronotum each with a broad pale orange band extending horizontally fromadjacent to eye to lateral margin. Tegmen and wing whitish hyaline, veins pale brown. Tegmen with a veryfaint, smoky brown, transverse band at one-third length; another less distinct band slightly distad ofmidlength. Wing unmarked. Shaft of aedeagus greatly expanded subapically; dorsal surface subapically with a pair of very largeflap-like processes, each terminating anteriorly in a large spine-like projection. Paramere very broad, apexobtusely rounded; dorsal process situated at midlength, strongly produced posteriorly. MATERIAL EXAMINEDHolotype cf , Brazil: Para, Jabaty, v.1924 (Williams) (BMNH). Though externally very similar to simpla, this species is readily distinguished by the structure ofthe male genitalia. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 69 Mysidia agilissp. n. (Figs 190, 300, 409) Male: head 0-52 mm long, 0-65 mm wide; pronotum 1-15 mm wide; tegmen 5-40-5-95 mm long; wing3-40 mm long. Female: tegmen 5-50 mm long. Length of frons 7 times width at apex, slightly greater than twice width at base; ocelli small, notprominent; clypeus one-quarter longer than frons; rostrum extending to base of subgenital segment.Pronotal width 11 times mid-dorsal length; fronto-lateral surfaces and tegula not distinctly carinate. Head and body unmarked. Tegmen and wing hyaline, only very weakly tinged whitish. Tegmen with afaint, transverse, smoky brown band at level of first fork of cubital vein; another less distinct, moreirregular band immediately distad of mid-length; a very indistinct and irregular band over apical branchesof medial vein; cross-veins weakly edged smoky brown. Wing with a prominent, transverse, smoky-brownband at level of radial-medial cross-vein; posterior and apical margins broadly smoky brown. Shaft of aedeagus ventrally and laterally expanded apically, with a pair of slender processes dorsally;dorsal surface subapically with a large flap-like process at midline, a pair of slender, spine-like processeslaterally. Paramere broad, robust; apex obtusely rounded; dorsal process situated slightly distad ofmidlength, apex vertically directed, not produced posteriorly. MATERIAL EXAMINED Holotype c?, Guyana: Tumatumari, 19.vii.1923 (Williams) (BMNH).Paratypes. Brazil: 1 c?, 2 $, Amazonas, P. das Laranjeiras (INPA; BMNH). The structure of the male genitalia is very distinctive in this species; the heavily armed aedeaguscoupled with the relatively undeveloped paramere, and the pigmentation of the tegmen andwing render it easily distinguishable. Mysidia claudata sp. n. (Figs 175, 284, 393) Male: head 0-50 mm long, 0-74 mm wide; pronotum 1-26 mm wide; tegmen 5-60-6-03 mm long; wing3-45 mm long. Female: tegmen 6-12-6-80 mm long. Length of frons c. 6 times width at apex, twice width at base; ocelli small, obscure; clypeus c. as long asfrons; rostrum extending to level of hind coxae. Pronotal width 15 times mid-dorsal length; fronto-lateralsurfaces and tegula not carinate. Fronto-lateral surfaces of pronotum rarely with a bright orange band extending horizontally fromadjacent to midline of eye to lateral margin; females with last abdominal segment very narrowly orange atventro-lateral angles. Tegmen and wing very faintly smoky hyaline, veins pale. Tegmen with a smallbrownish spot between subco'stal vein and costal margin at one-sixth length; a narrow, irregular, transverseband extending from costal to claval margin at one-third length; a brownish band extending from secondbranch of cubital vein to claval margin slightly basad of midlength; a very faint and ill-defined transverseband over apical forks of radial and medial veins. Wing with a very faint, broken, oblique, smoky browntransverse band at approximately midlength; apical third of length with veins and branches of veins broadlyedged smoky brown. Shaft of aedeagus slender; lateral surfaces subapically each produced into a flap-like process extendingbasad from apex to midlength, terminating anteriorly in long, curving, lateral process at midlength; dorsalsurface with five pairs of short, triangular, lateral spines subapically, and a large rounded process mediallyat three-fifths length. Paramere slender, apex truncate; dorsal process situated at two-thirds length,strongly produced posteriorly; dorsal surface subapically with a large secondary process bearing numerousshort robust spines; internal surface narrowly produced and extended dorsally at one-third length;ventro-lateral surface at one-third length with a large, triangular, hook-like process. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, 6.viii.l981 (Arias) (INPA).Paratypes. 8 C?, 30 $, same locality as holotype (INPA; BMNH). This species is distinguished by the pigmentation of the tegmen and wing, and by the complexstructure of the male genitalia. 70 PETER S. BROOMFIELD Mysidia jamesisp. n. (Figs 181, 291, 398) Male: head 0-53 mm long, 0-61 mm wide; pronotum 1-24 mm wide; tegmen 6-12 mm long; wing 3-50 mmlong. Female unknown. Length of frons 6 times width at apex, c. 3 times width at base; ocelli small, distinct; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width c. 14 times mid-dorsallength, fronto-lateral carinae absent; tegula weakly carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with a pale brownish transverseband over first fork of cubital vein; a paler, irregular, marking over medial-cubital cross-vein; a weaktransverse band immediately distad of midlength; basal one-fifth length irregularly mottled pale smokybrown. Wing with a very faint, brownish, transverse band at midlength, another over radial-medialcross- vein. Shaft of aedeagus slender in lateral aspect, very broad in dorsal aspect; dorsal surface expanded laterallyinto a pair of flap-like processes; with a pair of low flap- like processes adjacent to midline, each terminatingin an acute spine posteriorly. Paramere robust; dorsal process situated slightly distad of midlength, greatlyproduced posteriorly; dorsal surface subbasally with numerous small robust spines; ventral surface atmidlength with a rounded projection bearing a tuft of long robust spines. MATERIAL EXAMINEDHolotype C?, Brazil: Para, Jubaty, v.1924 (Williams) (BMNH). Mysidia carosella sp. n. (Figs 180, 290, 400) Male: head 0-57 mm long, 0-67 mm wide; pronotum 1-47 mm wide; tegmen 6-80 mm long; wing 3-80 mmlong. Female unknown. Length of frons 8 times width at apex, 2-5 times width at base; ocelli small, not prominent; clypeusslightly longer than frons; rostrum extending to base of subgenital plate. Pronotal width 25 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Genae and fronto-lateral surfaces of pronotum adjacent to eyes tinged orange. Tegmen and wing whitishhyaline; veins, cross-veins and forks of veins pale yellowish brown. Tegmen with radial, medial and clavalareas irregularly smoky brown over basal one-quarter length; an irregular brownish band extending fromcostal to claval margins at level of first fork of cubital vein; a very irregular, somewhat oblique, brownishband extending from medial vein to apex of clavus at level of first fork of medial vein; a broad brownishband extending from costal to posterior margins at level of second fork of medial vein; apical one-thirdlength irregularly mottled brownish around veins and forks of veins. Wing with an indistinct, palebrownish, transverse band extending from costal margin to claval suture at one-quarter length; a darkertransverse band extending from medial vein to apex of clavus immediately posterior to first fork of cubitalvein; a broad, rather irregular, transverse, brownish band at level of radial-medial cross- vein; apicalmargin narrowly pale smoky brown. Shaft of aedeagus slender; apex broadly produced dorsally and anteriorly; dorso-lateral margins eachsubapically produced into a large, flap-like process extending to just short of midlength; dorsal surface atone-third length with a pair of rounded flap-like processes. Paramere complex, constricted medially; apexbroadly rounded, strongly produced dorsally; dorsal process situated at c. two-thirds length, stronglyproduced dorsally and posteriorly; internal ventral surface with a rounded node at two-thirds length,bearing numerous, small, tooth-like spines subbasally. MATERIAL EXAMINEDHolotype cT, Bolivia: 3 miles N. Buena Vista, 26.iii.1978 (O'Brien) (FAMU). No single external character distinguishes this species, but the male genitalia are distinctive. Mysidia harmonia sp. n. (Figs 206, 316, 425) Male: head 0-50 mm long, 0-75 mm wide; pronotum 1-51 mm wide; tegmen 7-35 mm long; wing 4-25 mmlong. Female unknown. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 71 Length of frons c. 6 times width at apex, 2-5 times width at base; ocelli small, distinct; clypeus one-thirdlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 26 times mid-dorsallength, fronto-lateral carinae obsolete; tegula weakly carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with veins and cross-veinsbroadly edged very pale brownish; basal area irregularly mottled pale brown; an irregular, brownish,transverse band at one-third length; another, fainter, band over medial-cubital cross-vein; a third, verynarrow, band over second fork of medial vein. Wing with an irregular, smoky hyaline, transverse band overfirst fork of cubital vein; another more broken band over medial-cubital cross-vein; a third overradial-medial cross- vein. Shaft of aedeagus laterally expanded subapically; dorsal surface subapically with a pair of very large,acute, flap-like processes. Paramere broadly rounded apically; dorsal process small, situated at c.midlength, dorsally produced, laterally bifurcate apically; with a secondary process subbasally bearing asingle rounded projection. MATERIAL EXAMINEDHolotype c?, Colombia: Putumayo, La Hormiga, 6.ix.l978 (Cooper) (BMNH). In the absence of distinctive external characters, this species is most readily distinguished by themale genitalia. Mysidia silvana sp. n. (Figs 251, 362, 472) Male: head 0-55 mm long, 0-82 mm wide; pronotum 1-80 mm wide; legmen 9-35 mm long; wing 5-10 mmlong. Female unknown. Length of frons c. 6 times width at apex, c. 3 times width at base; ocelli distinct; clypeus one-fifth longerthan frons; rostrum extending to base of subgenital plate. Pronotal width 28 times mid-dorsal length,fronto-lateral carinae absent; tegula weakly carinate. Head unmarked. Disc of mesonotum dark brown. Tegmen and wing whitish hyaline. Tegmen withcross-veins broadly edged pale smoky brown; an irregular, pale brown, transverse band at one-tenthlength; another at one-fifth length; a third over first fork of cubital vein; another at two-fifths length; a fifthat midlength; another over second fork of medial vein; a seventh much fainter band at three-quarterslength. Wing with a broken, irregular, indistinct, pale brownish, transverse band at midlength; anotherover first fork of medial vein. Shaft of aedeagus slender, greatly expanded subapically; dorsal surface subapically with a pair of verylarge flap-like processes, each terminating in a long spine and bearing at midlength an erect spine.Paramere very robust, apex obtusely rounded; dorsal process situated at two-fifths length, posteriorlyproduced; dorsal surface at two-thirds length produced into a large, medially directed, flap-like secondaryprocess. MATERIAL EXAMINEDHolotype $, Peru: Tingo Maria, 13.vii.1968 (O'Brien) (FAMU). The dark brown mesonotal disc and the structure of the male genitalia distinguish this species. Mysidia bella sp. n. (Figs 229, 340, 449) Male: head 0-63 mm long; 0-82 mm wide; pronotum 1-36 mm wide; legmen 6-80 mm long; wing 4-00 mmlong. Female unknown. Length of frons c. 6 times width at apex, slightly less than 2-5 times widlh al base; ocelli small andobscure; clypeus slighlly longer lhan frons; roslrum exlending lo base of subgenilal plale. Pronolal widlh c.15 limes mid-dorsal lenglh; fronlo-laleral surfaces and legula nol carinale. Head and body unmarked. Tegmen and wing whilish hyaline, veins pale. Tegmen wilh an indislincl,irregular, brownish, Iransverse band subbasally; anolher more broken band exlending from medial vein toclaval margin at approximately one-sixth length; another more continuous band extending from costalmargin to apex of anal vein at one-quarler lenglh; anolher irregular band exlending from second fork ofcubilal vein lo apex of clavus al Ihree-eighlhs lenglh; a very indislincl, pale, Iransverse band exlendingfrom coslal margin lo apex of firsl branch of cubilal vein slighlly dislad of midlenglh. Wing with anirregular, smoky brown band extending transversely from medial vein to apex of clavus at one-third length, 72 PETER S. BROOMFIELD and produced narrowly basad over apices of anal veins; another broader band extending from radial-medial cross-vein to posterior margin at three-quarters length, extending narrowly along posterior marginover apices of first and second branches of cubital vein. Shaft of aedeagus slender in lateral aspect; lateral surfaces subapically each with a large, flap-like processextending over dorsal surface, bearing rounded projection dorsally; ventral surface at two-thirds lengthwith a small, triangular process at midline. Paramere robust; apex obtusely rounded; dorsal process large,situated at midlength, strongly produced dorsally and posteriorly. MATERIAL EXAMINEDHolotype cf , Brazil: Amazonas, P. das Laranjeiras, 30.vii.1981 (Arias) (INPA). This species is distinguished by the five transverse bands on the tegmen, the two transversebands on the wing, and by the structure of the male genitalia. Mysidia decora sp. n. (Figs 192, 302, 411) Male: head 0-59 mm long, 0-71 mm wide; pronotum 1-64 mm wide; tegmen 8-25 mm long; wing 4-70 mmlong. Female unknown. Length of frons slightly greater than 6 times width at apex, slightly less than 3 times width at base; ocellidistinct; clypeus slightly shorter than frons; rostrum extending to apex of abdomen. Pronotal width25 times mid-dorsal length; fronto-lateral surfaces and tegula not carinate. Pronto-lateral surfaces of pronotum weakly tinged orange. Tegmen and wing whitish hyaline, veins paleyellow, cross-veins and forks of veins broadly edged pale smoky brown. Tegmen with a narrow, smokybrown, transverse band at one-tenth length; another at one-quarter length; another at one-third length;with less distinct and more irregular bands at two-fifths and midlength. Wing with an irregular, indistinct,smoky brown, transverse band at midlength, another over radial-medial cross- vein. Shaft of aedeagus broadly expanded from midlength; lateral surfaces subapically each with a largeflap-like process; dorsal surface subapically with a pair of long slender processes medially. Paramererobust, apex very obtusely rounded; dorsal process situated at midlength, small, little producedposteriorly. MATERIAL EXAMINEDHolotype cf , Brazil: Mato Grosso, 12 50'S 51 47'W, 6.iv.l968 (Richards) (BMNH). This species, though closely related to nitida, limpida and amarantha, is readily distinguished bythe structure of the male genitalia, especially the paramere, and by the tegminal pigmentation. Mysidia boliviana sp. n. (Figs 250, 361, 471) Male: head 0-67 mm long, 0-88 mm wide; pronotum 1-90 mm wide; tegmen 9-25 mm long; wing 5-20 mmlong. Female unknown. Length of frons c. 5 times width at apex, 2-5 times width at base; ocelli distinct; clypeus as long as frons;rostrum extending to posterior margins of hind coxae. Pronotal width 23 times mid-dorsal length,fronto-lateral surfaces and tegula not carinate. Genae adjacent to eyes and dorsal to ocelli broadly dark brownish. Tegmen and wing whitish hyaline;veins pale, yellowish. Tegmen with a narrow, faint, pale brown band at approximately one-eighth length; avery broad, dark brown, transverse band over first and second forks of cibutal vein; a very faint, broken,pale brownish transverse band extending from immediately basad of medial-cubital cross-vein to posteriormargin; a broad, darker brownish, transverse band extending from costal to posterior margins immediatelydistad of second fork of medial vein. Wing with an irregular, pale brownish, transverse band immediatelydistad of first cubital fork; another, similar band extending from radial-medial cross-vein to posteriormargin. Shaft of aedeagus slender, somewhat expanded over apical one-quarter length; lateral surfacessubapically each with a large flap-like process extending over dorsal surface and strongly over-lapping atmid-dorsal line, each produced anteriorly into a long spine, bearing mid-dorsally a small spine. Paramererobust; apex broadly rounded; dorsal process situated slightly distad of midlength, strongly producedposteriorly; dorsal surface at c. three-fifths length strongly, conically and dorsally produced. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 73 MATERIAL EXAMINEDHolotype cT, Bolivia: Buenavista, 400 m (Steinbach) (CM). The male genitalia of this species closely resemble those ofsilvana, but it is readily distinguishedby the prominent dark bands on the tegmina. Mysidiapersephonesp. n. (Figs 204, 314, 423) Male: head 0-57 mm long, 0-75 mm wide; pronotum 1-28 mm wide; tegmen 6-38 mm long; wing 3-57 mmlong. Female unknown. Length of frons c. 5 times width at apex, c. 3 times width at base; ocelli very large and prominent; clypeusone-quarter longer than frons; rostrum terminating slightly posterior to hind-coxae. Pronotal width c. 12times mid-dorsal length; fronto-lateral surfaces and tegula without distinct carinae. Head and body unmarked; ocelli narrowly edged reddish. Tegmen and wing whitish hyaline, veins pale.Tegmen with a broad, pale brownish, transverse band at one-seventh length; another, similar bandextending from costal to claval margins over first fork of cubital vein; a very faint brownish band betweenfirst and second forks of medial vein; another similar band extending from radial-medial cross-vein almostto apical fork of radial vein; with an irregular and indistinct, pale brownish spot over apical branches ofradial and medial veins. Wing with a faint, pale brownish spot over first branch of cubital vein at midlength;apical one-third length very faintly brownish. Shaft of aedeagus slender; dorsal surface subapically with a pair of long, spine-like processes; dorso-lateral surfaces each with a curving spine-like process. Paramere slender; apex acutely rounded; dorsalprocess situated immediately distad of midlength, weakly produced dorsally and posteriorly; dorsal surfacesubbasally with a rounded projection bearing numerous short robust spines. MATERIAL EXAMINED Holotype d" r Brazil: Rio Uaupes, Taracua, IS.iii (Roman) (NR).Paratype. 1 d", same data as holotype (BMNH). The structure of the aedeagus closely resembles that of intima but that of the paramere is quitedistinct; the species is also distinguished by the prominent ocelli and by the tegminal pigmenta-tion. Mysidiamarshallisp. n. (Figs 223, 334, 444) Male: head 0-69 mm long, 0-48 mm wide; pronotum 1-72 mm wide; tegmen 8-10-8-50 mm long; wing4-68 mm long. Female unknown. Length of frons 5 times width at apex, c. twice width at base; ocelli small, distinct; clypeus one-thirdlonger than frons; rostrum extending to base of subgenital plate. Pronotal width c. 20 times mid-dorsallength; fronto-lateral surfaces and tegula without distinct carinae. Head and body unmarked. Tegmen and wing whitish hyaline, veins pale brown. Tegmen withalternating transverse bands of pale and rather darker smoky brown over basal one-third length, enclosinga small, circular, pale spot between cubital vein and clavus; with an indistinct, smoky brown, transverseband over first fork of medial vein; another darker band over second fork of medial vein; apical two-fifthslength very pale smoky brown. Wing with an irregular, smoky brown, transverse band over first fork ofcubital vein; another broken band at midlength; a third over radial-medial cross-vein; apex smoky brown. Shaft of aedeagus greatly expanded from two-fifths length to apex; lateral surfaces each with a very largeflap-like process extending over dorsal surface. Paramere robust, apex broadly rounded; dorsal processsituated at midlength, small, produced posteriorly. MATERIAL EXAMINED Holotype cT, Bolivia: Cbb., Pto., San Francisco, 19 miles NW. Villa Tunari, l.iv.1978 (O'Brien &Marshall) (FAMU). Paratype. 1 d", same data as holotype (BMNH). Though the pigmentation of the wing is distinctive, reference should also be made to the malegenitalia in determination of this species. 74 PETER S. BROOMFIELD Mysidia neonebulosa Muir (Figs 230, 341, 450)Mfysidia] neonebulosa Muir, 1918: 424. Holotype cf , GUYANA (OSU) [examined]. Male: head 0-56 mm long, 0-80 mm wide; pronotum 1-50 mm wide; tegmen 6-43-8-50 mm long; wing4-00 mm long. Female: tegmen 7-70 mm long. Length of frons 7 times width at apex, c. 2-5 times width at base; ocelli small, not prominent; clypeus aslong as frons; rostrum extending to base of subgenital plate. Pronotal width 18 times mid-dorsal length;fronto-lateral surfaces not carinate; tegula with weak carinae. Pronto-lateral surfaces of pronotum each with a broad orange band extending horizontally fromadjacent to midline of eye to lateral margin. Tegmen and wing whitish hyaline. Tegmen with a faint,irregular, pale brownish, transverse band at one-eighth length, another at level of first fork of cubital vein,another at level of medial-cubital cross-vein, a fourth, very faint band immediately distad of midlength.Wing with an indistinct, very pale brownish, transverse band at midlength, another at three-quarterslength. Shaft of aedeagus somewhat expanded distad of midlength; dorsal surface subapically with a pair ofbroad, flap-like processes; ventro-lateral surfaces each with a short, rounded flap-like process subapically.Paramere broad, apex obtusely rounded; dorsal process situated immediately distad of midlength, apexstrongly produced posteriorly. MATERIAL EXAMINED Holotype cf , Guyana: Bartica, 14.vii.1901 (Parish) (OSU).Guyana: 1 cf , 2 $, Kartabo (BMNH). Brazil: 4 cf , Para, Jabaty (BMNH). The holotype has one tegmen and both wings missing. A small delicate species; the markings ofthe tegmen and wing are often very faint; the structure of the male genitalia is, however, quitedistinctive. Mysidia amarantha sp. n. (Figs 199, 309, 418) Male: head 0-59 mm long, 0-80 mm wide; pronotum 2-00 mm wide; tegmen 8-50-9-70 mm long; wing5-00 mm long. Female: tegmen 9-30 mm long. Length of frons c. 5-5 times width at apex, c. 2-5 times width at base; ocelli small, distinct; clypeusone-quarter longer than frons; rostrum extending to midlength of abdomen. Pronotal width 20 timesmid-dorsal length; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline. Tegmen with a pale brownish transverseband at one-third length; another, more irregular and less distinct band over medial-cubital cross-vein; athird, more broken band over radial-medial cross- vein; cross- veins very weakly edged smoky brown. Wingwith an indistinct transverse band at two-fifths length; another over radial-medial cross-vein, smokybrown. Shaft of aedeagus greatly laterally expanded over apical two-fifths length; dorsal surface subapically witha pair of large flap-like processes, each with anterior margin adjacent to midline produced into a curvingspine. Paramere broad, apex obtusely rounded; dorsal process situated at midlength, not posteriorlyproduced. MATERIAL EXAMINED Holotype cf , Ecuador: Napo, Muyana, 5 km SW. of Tena, 27.xi.1978 (Cooper) (BMNH). Paratypes. French Guiana: 1 cf, Mana River (CM). Brazil: 2 cf, 1 $, Amazonas, P. das Laranjeiras(INPA; BMNH). This species is only reliably distinguished by reference to the male genitalia. Mysidia magica sp. n. (Figs 249, 360, 470) Male: head 0-63 mm long, 0-84 mm wide; pronotum 1-40 mm wide; tegmen 7-60-8-00 mm long; wing4-30 mm long. Female: tegmen 8-65-9-60 mm long. Length of frons 7 times width at apex, c. 3 times width at base; ocelli small, distinct; clypeus one-quarter TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 75 longer than frons; rostrum extending to base of subgenital plate. Pronotal width 22 times mid-dorsallength; fronto-lateral surfaces and tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins yellowish. Tegmen with a narrow,pale brownish, transverse band over point of fusion of anal veins; another more distinct band extendingfrom costal margin to apex of clavus; a third, fainter and more broken band extending from medial-cubitalcross-vein to posterior margin; a fourth very faint band extending from costal to posterior margins at c.midlength. Wing occasionally with a very pale, indistinct, irregular, smoky brown, transverse band overfirst fork of medial vein, over radial-medial cross-vein, and a third over second fork of cubital vein. Shaft of aedeagus apically expanded; dorsal surface subapically with a pair of conical processes, eachterminating in a large spine; lateral surfaces subapically each with a large spine and a large roundedprocess. Paramere short and broad; dorsal process situated at midlength. MATERIAL EXAMINED Holotype cf , Surinam: Brokopondo, 29.L1969 (O'Brien) (FAMU).Paratypes. Surinam: 1 cf , 3 $ , data as holotype and 17 km S. of Kraka (FAMU; BMNH). This species is distinguished by the four transverse bands of the legmen, the three bands of thewing, and by the structure of the male genitalia. Mysidia geoffreyi sp. n. (Figs 274, 385, 495) Male: head 0-63 mm long, 0-78 mm wide; pronotum 1-62 mm wide; tegmen 8-80-8-90 mm long; wing5-10 mm long. Female: tegmen 9-77 mm long. Length of frons 6 times width at apex, 2-33 times width at base; ocelli small, distinct; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width c. 15 times mid-dorsallength, fronto-lateral carinae absent; tegula weakly carinate. Fronto-lateral surfaces of pronotum occasionally each with a broad, pale orange, horizontal bandextending from adjacent to eye to lateral margin. Tegmen and wing whitish hyaline. Tegmen with anirregular, often very pale, brownish, transverse band slightly basad of one-third length; a fainter, broken,band immediately distad of midlength; irregularly mottled pale brown basally and over first fork of medialvein; cross- veins very narrowly and irregularly edged pale brownish. Wing with irregular and indistinct,pale brown, transverse bands at one-third length and over radial-medial cross-vein. Shaft of aedeagus greatly laterally expanded; dorsal surface over apical two-fifths length with a pair ofbroad flap-like lobes, a pair of overlapping apically acute processes at midline. Paramere broad, apicallyrounded; dorsal process situated at one-third length, greatly produced posteriorly. MATERIAL EXAMINED Holotype cf , Bolivia: Pando, Provenir, 9.vii.l979 (Cooper) (BMNH).Paratype. Peru: 2 cf , 2 $, Callanga (BMNH). Lacking distinctive external characters, this species is most readily distinguished by the structureof the male genitalia. Mysidia pseudonebulosa Muir (Figs 278, 389, 499)Mysidia pseudonebulosa Muir, 1918: 423. Holotype cf , GUYANA (OU) [examined]. Male: head 0-63 mm long, 0-80 mm wide; pronotum 1-76 mm wide; tegmen 8-50 mm long. Femaleunknown. Length of frons 7 times width at apex, 3 times width at base; ocelli large, prominent; rostrum extendingbeyond base of subgenital plate. Pronotal width 21 times mid-dorsal length; fronto-lateral surfaces andtegula not carinate. Genae at level of eyes broadly tinged orange ; fronto-lateral surfaces of pronotum broadly and irregularlyorange from level of dorsal margins of eyes to lateral margins. Tegmen whitish hyaline, with a brownishtransverse band at immediately basad of second fork of cubital vein, another immediately distad of secondfork of medial vein. Shaft of aedeagus, laterally expanded over apical two-fifths length; lateral surfaces subapically eachbearing a large, flap-like process extending over dorsal surface, each bearing a long, curving, spine-like 76 PETER S. BROOMFIELD projection; dorsal surface at three-quarters length with a single process medially. Paramere robust; apexbroadly rounded; dorsal process situated at midlength, strongly produced posteriorly; dorsal marginsubapically somewhat produced and medially^nclined. MATERIAL EXAMINEDHolotype cT, Guyana: Bartica, 9.V.1901 (Osborri) (OU). The clypeus is obscured and, due to the fragile and badly damaged condition of the uniquespecimen available for study, it is considered inadvisable to remount it. The right tegmen ismounted on a card below the specimen with the markings obscured by glue; it would appear,however, that more dark transverse bands may be present than the two noted above. The lefttegmen and both wings are missing. Due to the lack of distinctive external characters, this species may only be positivelydistinguished by reference to the male genitalia. Mysidia cinerea Fennah (Figs 189, 299, 408)Mysidia cinerea Fennah, 1945: 439. Holotype d", TRINIDAD (USNM) [examined]. Male: head 0-46 mm long, 0-63 mm wide; pronotum 1-95 mm wide; tegmen 6-00 mm long; wing 3-50 mmlong. Length of frons c. 5 times width at apex, 2-5 times width at base; ocelli distinct; clypeus c. as long as frons;rostrum extending slightly beyond hind coxae. Pronotal width 11 times mid-dorsal length, fronto-lateralcarinae absent; tegula prominently carinate. Head and body unmarked. Tegmen and wing clear hyaline; veins and cross-veins dull yellowish, veryfaintly margined pale smoky brown. Shaft of aedeagus with lateral surfaces subapically each bearing a large flap-like process extending overdorsal surface; ventral surface with a pair of short triangular processes somewhat basad of two-thirdslength. Paramere with apex regularly rounded; dorsal process large, situated slightly basad of midlength,reduced to a large, curving, posteriorly directed hook. MATERIAL EXAMINEDHolotype cT, Trinidad: Northern Range, 12.vi.1942 (Fennah) (USNM). The paratype noted by Fennah as being in the BMNH is presumed lost. The holotype, probablydue to having been previously stored in alcohol, has lost most of its pigmentation, which Fennahdescribed thus: . . . eyes red; tegmina hyaline, all veins faintly and broadly overlain with brown, a clear ellipsoidalspot near apical fork of M, with the veins tinged brown at fork, veins otherwise concolorous; wingspale hyaline, veins irregularly pale brown, apical cells clouded near margin, veins concolorous.Insect in life powdered pearly gray. Due to the extremely shrivelled condition of the type, the above measurements are largelyestimated, and the species can only be determined by reference to the male genitalia. Nomina dubia Mysidia pallida (Fabricius) Derbe pallida Fabricius, 1803: 81. LECTOTYPE , CENTRAL AMERICA (UZM), here designated [ex-amined].Mysidia pallida (Fabricius) Westwood, 1840: 83. Female: head 0-78 mm long, 0-96 mm wide; pronotum 2-31 mm wide; tegmen 9-70 mm long; wing 5-40 mmlong. Male unknown. Length of frons 5 times width at apex, c. twice width at base; ocelli small, distinct; clypeus one-fifthlonger than frons; rostrum unknown. Pronotal width 25 times mid-dorsal length, lacking fronto-lateralcarinae; tegula prominently carinate. Fronto-lateral surfaces of pronotum each with a narrow orange band extending horizontally fromadjacent to eye to lateral margin; tegula ventral to carina dark brown [?]; disc of mesonotum and dorsal * TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDHNI (HOMOPTERA) 77 surface of abdomen brownish. Tegmen and wing clear hyaline; veins and cross-veins pale brown, narrowlyedged brown. Tegmen with costal area yellowish brown; basal one-fifth length irregularly pale brownish; apale brownish transverse band at slightly distad of first fork of cubital vein; another adjacent to first fork ofmedial vein; a third adjacent to second fork of medial vein; a fourth, very faint band extending from costalmargin to radial-medial cross-vein; apical fork of radial vein dark brown. Wing with a pale, indistinct,brownish, transverse band immediately distad of first fork of cubital vein; an even fainter band overradial-medial cross-vein. MATERIAL EXAMINED Lectotype $, Central America: (Schmidt) (UZM).Paralectotype. Central America: 1 $ (UZM). The lectotype appears to be teneral and the pigmentation described above is by no meanscertain. The paralectotype lacks both tegmina and it may not be conspecific with the lectotype.M. pallida is known only from these two females, neither of which is in good condition; 'pallida'is therefore regarded as a nomen dubium and is omitted from both keys. Mysidia stigma Germar Derbe stigma Germar, 1830: 56. Syntypes, URUGUAY [not examined].Mysidia stigma (Germar) Schaum, 1850: 70. It has not been possible to examine the type-material of this species, which therefore cannot beredescribed and is omitted from the keys. Germar 's description is as follows: alba, elytris puncto ante apicem venisque transversis nigris. Habitat in Monte Video. Mus. deWinthem. Caput, thorax et abdomen alba. Elytra alba, opaca: puncta ante angulum anticumserieque venarum transversali nigris. Alae albae striga una alterave transversa nigra. It would appear from the above description that the species is probably correctly placed inMysidia; distinct transverse bands on the tegmina and wings are not a feature of the genusDerbe, nor within the other subfamilies of the Neotropical Derbidae. This is the only speciesrecorded from as far south as Uruguay. Species previously transferred from Mysidia Heronax elatior (Fowler) Mysidia elatior Fowler, 1900: 73.Heronax elatior (Fowler) Muir, 1918: 230. Examination of Fowler's type-material confirms Muir's transfer of this species from theDerbinae. Neocenchrea spreta (Fowler) Mysidia spreta Fowler, 1900: 74.Basileocephalus spretus (Fowler) Muir, 1918: 230.Neocenchrea spreta (Fowler) Metcalf, 1938: 331. Examination of the type-material confirms the transfer of the species from the Derbinae. PSEUDOMYSIDIA MetcalfPseudomysidia Metcalf, 1938: 317. Type-species: Pseudomysidia fuscovaria Metcalf, by monotypy. Width of head in dorsal aspect slightly less than one-third greater than length. Vertex with lateral marginsstrongly converging from base to level of anterior margins of eyes, then very gradually converging tojunction with frons; extending beyond anterior margins of eyes for up to one-half length; basal margintransverse; lateral carinae distinct, but not foliate; junction with frons broadly and regularly rounded.Frons with lateral margins subparallel from apex to level of midline of eyes, then gradually and regularlydiverging to base; very slender, length 12-17 times width at apex, c. 3-0-4-5 times width at base; lateralcarinae very prominent subbasally. Genae extending anterior to eyes for from one-third to one-half 78 PETER S. BROOMFIELD horizontal diameter of eye. Antenna with second segment club-shaped, c. twice as long as maximumbreadth; apex transverse; flagellum arising apically. Ocelli small, distinct, occasionally prominent. Clypeusslender, length up to one-third greater than that of frons, 3-5-4-5 times width at base; medial carina distinctover c. apical three-quarters length; lateral carinae distinct and percurrent. Rostrum extending to frombase of subgenital plate to slightly beyond apex of abdomen. Pronotal width 6-5-11-0 times mid-dorsal length; very deeply, broadly and regularly incised basally.Pronto-lateral surfaces usually each with a distinct, rarely obsolete, horizontal carina extending fromadjacent to midline of eye to lateral margin. Tegula rarely weakly carinate. Disc of mesonotum c. as long aswide; medial and lateral carinae usually distinct, extending over apical half to four-fifths length, rarelyobsolete. Tegmen length usually 5-50-6-80 mm; those of females being slightly longer than those of males. Medialvein becoming distinct from fused radial and subcostal veins at c. one-eighth length; radial and subcostalveins separating slightly basad of midlength. Radial vein with two branches extending to apical margin.Medial vein with 11 branches extending to apical and posterior margins, linked to radial vein by cross-veinsat three-quarters length and subapically; cross-veins between first and third, fourth and fifth, six andseventh, and eighth and ninth branches. Cubital vein with three branches extending to posterior margin;first linked to claval suture and to second, second to third, and third to first branch of medial vein bycross- veins. Wing c. half as long as tegmen. Subcostal and radial veins fused over basal one-third length; radial veinunbranched, linked to medial vein by a single oblique cross-vein somewhat distad of midlength. Medialvein distinct from base, with three branches extending to apical and posterior margins. Cubital vein withtwo branches extending to posterior margin, second linked to first medial by a cross-vein. Head and thorax predominantly pale yellowish brown, often with dorsal surfaces, genae, and lateralsurfaces of clypeus tinged reddish; ocelli often bright red; frons with lateral carinae rarely dark brown;fronto-lateral surfaces of pronotum often tinged reddish. Dorsal surface of abdomen, at least in part, brightred. Tegmen and wing whitish or hyaline; veins usually pale, occasionally with cross-veins and forks ofveins brownish; veins and cross-veins usually edged smoky brown, these markings frequently very faint,often coalescing to form very irregular transverse bands; posterior and apical margins often broadly smokybrown, prominent markings absent. Male genitalia with shaft of aedeagus symmetrical, horizontal, slender in lateral aspect, basallycylindrical; dorsal surface subapically with 4-6 pairs of mainly horizontal, anteriorly directed, occasionallystrongly forked, serrated or apically bifurcate spine-like processes; ventral surface unarmed. Paramereslender, never very robust; basal apodeme one-quarter to slightly less than half total length; apex usuallyacutely rounded, narrowly inclined towards midline; dorsal process situated at or basad of midlength,simple, usually not produced posteriorly, very rarely with interlocking processes, usually short, broad andapically truncate, lacking a secondary dorsal process; ventral surface usually with numerous long robustspines subbasally. Anal tube little produced, c. as long as broad; apex commonly rounded, deeply notchedmedially. Female with posterior margin of subgenital plate frequently strongly produced; apex broadly rounded,transverse, or shallowly concave. The tegminal venation, the structure of the male genitalia and the apical position of the antennalflagellum tend to indicate that Pseudomysidia is the least specialised of the mysidiine genera, and that withDysimia, Dysimiella and Symidia it diverged from the more common trend of development within thetribe, as exemplified by Mysidia, at a comparatively early stage in the development of the group. The 11branches of the medial vein of the tegmen distinguish the genus from all others in the Mysidiini. Although the aedeagal characters show continuity within the genus, two species-groups are proposed,based on the structure of the paramere. The/kscovan'o-group. Paramere with dorsal process slender, dorsally produced, bearing rudimentaryinterlocking processes which are most highly developed in Juliana. This group also includes palmeri,rubidella, debora and hindore. Thepanamensis-group. Paramere with dorsal process very much reduced, broad, apically truncate, withno suggestion of interlocking processes. This group includes all species not included above. Distributed from Mexico to Costa Rica, Trinidad, Panama, Brazil, Bolivia, Venezuela and Ecuador. Key to species of Pseudomysidia (based on external characters) The external differences between species are often very slight; where possible, reference should be made tothe structure of the male genitalia. 1 Tegmen with claval area dark. Venezuela araguana Fennah (p. 81) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 79 Tegmen with claval area pale 2 2 (1) Tegmen and wing with cross-veins and forks of veins strongly and broadly margined dark smoky brown 12 Tegmen with cross- veins and forks of veins weakly margined pale smoky brown; appear-ance predominantly hyaline 3 3 (2) Abdomen with dorsal surface predominantly bright red 4 Abdomen with dorsal surface brown or yellow , rarely narrowly red basally at midline 6 4 (3) Pronotum with fronto-lateral surfaces distinctly carinate. Panama palmer! sp. n. (p. 82) Pronotum with fronto-lateral carinae weak or absent 5 5(4) Male tegmen less than 6 mm. Mexico rubidella Ball (in part) (p. 80) Male tegmen greater than 6 mm. Panama Juliana sp. n. (p. 81) 6(3) Male tegmen not greater than 5 mm. Panama fuscovaria Metcalf (p. 80) Male tegmen greater than 5 mm 7 7 (6) Fronto-lateral surfaces of pronotum distinctly carinate 10 Fronto-lateral surfaces of pronotum with carinae obsolete 8 8 (7) Tegmen with veins and cross-veins pale yellow. Trinidad trinidadensis sp. n. (p. 82) Tegmen with veins and cross-veins brownish 9 9 (8) Junction of frons and vertex usually dark brown. Tegmen with pigmentation around cross-veins and forks of veins coalescing to form two broken, very irregular, transverse bands. Costa Rica s/mi/issp. n. (p. 83) Junction of frons and vertex unmarked. Tegmen with four very irregular transverse bands. Mexico rubidella Ball (in part) (p. 80) 10 (7) Tegmen with cross-veins and forks of veins dark brown. Costa Rica .... marshalli sp. n. (p. 83) Tegmen with cross-veins and forks of veins pale 11 11(10) Head pale yellowish throughout. Panama hindoresp. n. (p. 83) Head with genae dorsal and ventral to eyes bright scarlet. Brazil vestis sp. n. (p. 84) 12 (2) Fronto-lateral surfaces of pronotum tinged reddish 13 Fronto-lateral surfaces of pronotum yellowish throughout 15 13(12) Fronto-lateral surfaces of pronotum weakly carinate 14 Fronto-lateral surfaces of pronotum strongly carinate. Ecuador pallida sp. n. (p. 84) 14(13) Male tegmen less than 6 mm. Panama panamensis sp. n. (p. 84) Male tegmen greater than 6 mm . Ecuador ecuadoriensis sp. n. (p. 85) 15(12) Pronotal width 11 times length. Costa Rica deborasp. n. (p. 85) Pronotal width not greater than 8 times length 16 16(15) Pronotal width 8 times length. Tegmen with posterior and apical margins broadly smoky brown. Bolivia lepida sp. n. (p. 86) Pronotal width distinctly less than 8 times length. Tegmen with posterior and apicalmargins pale. Ecuador delicata sp. n. (p. 86), obnubilia sp. n. (p. 86) Key to species of Pseudomysidia (based on male genitalia) It has not been possible to examine a male of lepida which is therefore omitted from this key. 1 Paramere with dorsal process greatly reduced, situated basad of midlength, short, broad, and apically truncate 7 Paramere with dorsal process produced dorsally, situated at midlength, long, slender,apically acute 2 2 (1) Aedeagus with four pairs of subapical spines (Fig. 34) palmeri sp. n. (p. 82) Aedeagus with five or six pairs of subapical spines 3 3 (2) Aedeagus with five pairs of subapical spines (Fig. 35) rubidella Ball (p. 80) Aedeagus with six pairs of subapical spines 4 4 (3) Paramere with interlocking surfaces 5 Paramere without interlocking surfaces 6 5 (4) Aedeagus with longest pair of subapical spines curving laterally and apically serrated (Fig. 36) Juliana sp. n. (p. 81) Aedeagus with longest pair of subapical spines anteriorly directed and apically acute (Fig.37) deborasp. n. (p. 85) 6 (4) Aedeagus with third pair of subapical spines serrated, lateral spines short (Fig. 38) similis sp. n. (p. 83) 80 PETER S. BROOMFIELD Aedeagus with third pair of subapical spines not serrated, lateral spines long (Fig. 39) hi ml ore sp. n. (p. 83) 7 (1) Aedeagus with three pairs of subapical spines 8 Aedeagus with four or more pairs of subapical spines 9 8 (7) Aedeagus with medial pair of subapical spines strongly branched and dorsally serrated (Fig. 40) panamensis sp. n. (p. 84) Aedeagus with medial pair of subapical spines shallowly forked at apex (Fig. 41) fuscovaria Metcalf (p. 80) 9 (7) Aedeagus with four pairs of subapical spines 10 Aedeagus with at least five pairs of subapical spines 11 10(9) Aedeagus with third pair of subapical spines apically forked (Fig. 42) pallidasp.n. (p. 84) Aedeagus with all spines simple (Fig. 43) araguana Fennah (p. 81) 11 (9) Aedeagus with five pairs of subapical spines 13 Aedeagus with six pairs of subapical spines 12 12(11) Aedeagus with fifth pair of subapical spines strongly branched (Fig. 44) marshalli sp. n. (p. 83) Aedeagus with fifth pair of subapical spines not as above (Fig. 45) vestis sp. n. (p. 84) 13(1 1) Aedeagus with subapical spines strongly curving laterally (Fig. 46) trinidadensis sp. n. (p. 82) Aedeagus with subapical spines anteriorly directed 14 14(13) Aedeagus with subapical spines strongly curving dorsally (Fig. 63) ecuadoriensis sp. n. (p. 85) Aedeagus with spines not as above 15 15(14) Aedeagus with medial pair of subapical spines very slender, fourth pair longest (Fig. 48) delicata sp. n. (p. 86) Aedeagus with medial pair of subapical spines robust and longest (Fig. 49) ohnuhilia sp. n. (p. 86) Pseudomysidia fuscovaria Metcalf (Figs 11, 41, 57, 73)Pseudomysidia fuscovaria Metcalf, 1938: 317. Holotype cf , PANAMA (MCZ) [examined]. Male: head 0-44 mm long, 0-48 mm wide; pronotum 0-90 mm wide; tegmen 4-75 mm long; wing 2-58 mmlong. Female: tegmen 5-70 mm long. Length of frons 16 times width at apex, 3-5 times width at base; ocelli distinct; clypeus slightly longer thanfrons; rostrum extending to base of subgenital plate. Pronotal width 7 times mid-dorsal length; fronto-lateral surfaces and tegula with carinae obsolete or absent. Genae and fronto-lateral surfaces of pronotum occasionally tinged orange; ocelli crimson; scutellum anddorsal surfaces of abdomen occasionally tinged pale crimson. Tegmen and wing almost hyaline, faintlytinged whitish, veins yellowish brown; cross- veins and forks of veins darker brown, narrowly edged palesmoky brown; without transverse markings. Shaft of aedeagus slender; dorsal surface subapically with three pairs of spine-like processes, medial pairapically bifid. Paramere slender, apex narrowly rounded; dorsal process situated at midlength, notposteriorly produced. MATERIAL EXAMINED Holotype cf , Panama: Canal Zone, Barro Colorado, 15.vii.1924 (Banks) (MCZ).Paratypes. Panama: 7 cf , 12 $ same data as holotype (MCZ); 1 tf, 1 $ (FAMU; BMNH). This species is readily distinguished by its small size, lack of pigmentation on the tegmen andwing, and by the structure of the male genitalia. Pseudomysidia rubidella (Ball) comb. n. (Figs 35, 51, 67)Mysidia rubidella Ball, 1928: 199. Holotype cf , MEXICO (USNM) [examined]. Male: head 0-46 mm long, 0-42 mm wide; pronotum 0-92 mm wide; tegmen 5-60-5-70 mm long; wing3-00 mm long. Female: tegmen 6-30-6-45 mm long. Length of frons 11 times width at apex, 4 times width at base; ocelli small, distinct; clypeus slightly longerthan frons; rostrum terminating immediately posterior to hind coxae. Pronotal width 9 times mid-dorsallength; fronto-lateral surfaces and tegula weakly carinate. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 81 Head and body often bright scarlet with frons, clypeus, tegula and legs yellowish brown; otherwiseyellowish brown throughout. Tegmen and wing whitish hyaline, veins and cross-veins yellowish brown.Tegmen with cross-veins and forks of veins darker brown, broadly and irregularly edged smoky brown,these markings coalescing to form a very broken transverse band at level of second fork of cubital vein, andanother at each of first, second and third branches of medial vein. Wing with a faint, oblique, smoky browntransverse band at level of radial-medial cross- vein; apical and posterior margins weakly tinged greyishbrown. Shaft of aedeagus slender; dorsal surface subapically with five pairs of long spine-like processes.Paramere broadly rounded apically; dorsal process situated at midlength, dorsally directed. MATERIAL EXAMINED Holotype cf , Mexico: Verz Cruz, Presido, vi. (Barrett) (USNM).Mexico: 3 cf , 5 $ (AC; USNM; BMNH). Honduras: 5 cf (FAMU; BMNH). The transfer of this species from Mysidia is based on the 11 branches of the medial vein of thetegmen and on the structure of the male genitali. It is distinguished by its relatively large size,frequently bright red pigmentation, and by the structure of the male genitalia. Pseudomysidia Juliana sp. n. (Figs 36, 52, 68) Male: head 0-52 mm long, 0-54 mm wide; pronotum 1-22 mm wide; tegmen 6-12-6-46 mm long; wing3-40 mm long. Female: tegmen 6-31-6-48 mm long. Length of frons 13 times width at apex, 4-5 times width at base; ocelli very prominent; clypeus slightlylonger than frons; rostrum extending beyond base of subgenital plate. Pronotal width 8 times mid-dorsallength, fronto-lateral carinae weak or obsolete; tegula not carinate. Genae ventral to eyes frequently reddish, ocelli crimson, lateral margins of frons at level of eyes rarelydark brown; fronto-lateral surfaces of pronotum occasionally reddish; dorsal surface of abdomen,excluding genital segment, often deep red. Tegmen and wing whitish hyaline, veins yellow; cross-veins andforks of veins brownish, narrowly and irregularly edged pale smoky brown; posterior margins weakly andindistinctly tinged smoky grey. Shaft of aedeagus basally slender; dorsal surface subapically with six pairs of spine-like processes, thosefourth from midline with apices narrowly serrated. Paramere slender, apex acute; dorsal process situatedat midlength, long and slender; ventral surface subbasally with numerous long slender spines. MATERIAL EXAMINED Holotype cT, Panama: Chiriqui, Fortuna, 8215'W 844'N, 19.V.1978 (O'Brien & Marshall) (FAMU).Paratypes. Panama: 6 cf , 3 $, data as holotype (FAMU; BMNH). The relatively complex structure of the paramere closely resembles that of some species ofMysidia; the venation of the tegmen, however, and the structure of the aedeagus leave no doubtas to the correct placement of the species in Pseudomysidia. It is distinguished by its relativelylarge size, reddish pigmentation, and by the structure of the male genitalia. Pseudomysidia araguana Fennah stat. n.(Figs 43, 59, 75) Pseudomysidia fuscovaria Metcalf ssp. araguana Fennah, 1952: 123. Holotype $, VENEZUELA (BMNH)[examined]. Male: head 0-48 mm long, 0-53 mm wide; pronotum 1-05 mm wide; tegmen 5-80 mm long; wing 2-80 mmlong. Female: tegmen 6-40 mm long. Length of frons c. 13 times width at apex, 4 times width at base; ocelli small, distinct; clypeus one-fifthlonger than frons; rostrum extending to midlength of abdomen. Pronotal width 8 times mid-dorsal length,fronto-lateral carinae distinct; tegula not carinate. Genae weakly tinged reddish around crimson ocelli; fronto-lateral surfaces of pronotum adjacent to eyesreddish, disc of mesonotum brownish, abdomen broadly crimson along midline. Tegmen and wing whitishhyaline, veins yellowish. Tegmen with cross-veins and forks of veins narrowly edged smoky brown; clavalarea broadly blackish brown; with an irregular, dark brown, transverse band extending from second fork ofclaval vein to posterior margin, and another extending from first fork of cubital vein obliquely across 82 PETER S. BROOMFIELD cubital cross-veins; costal area with six faint, evenly spaced, pale brownish bands extending from medialvein to anterior margin. Wing unmarked. Shaft of aedeagus slender; dorsal surface subapically with four pairs of spine-like processes, thoseadjacent to midline longest. Paramere slender, apex acute; dorsal process situated somewhat distad ofmidlength, broad, apex truncate; ventral surface subbasally with long, robust spines. MATERIAL EXAMINED Holotype $, Venezuela: Aragua, Rancho Grande, 1949 (Racenis) (BMNH).Venezuela: 1 d" , nr Maracay (AMNH). Fennah's description is in error in ascribing to this species only 10 branches to the medial vein ofthe tegmen the left tegmen of the type, though damaged, shows eleven.This species is unique in having the claval area of the tegmen darkly pigmented. Pseudomysidiapalmerisp. n. (Figs 34, 50, 66) Male: head 0-54 mm long, 0-54 mm wide; pronotum 1-30 mm wide; tegmen 6-17 mm long; wing 3-48 mmlong. Female unknown. Length of frons c. 16 times width at apex, c. 3 times width at base; ocelli prominent; clypeus slightlylonger than frons; rostrum terminating at level of genital segment. Pronotal width c. 8 times mid-dorsallength, fronto-lateral carinae distinct; tegula each with two weak carinae. Genae ventral to eyes orange, ocelli narrowly edged crimson; fronto-lateral surfaces of pronotumventral to dorsal margins of eyes orange; abdomen dull reddish. Tegmen and wing very weakly tingedwhitish, veins yellow; cross-veins and forks of veins brownish, the latter narrowly edged smoky brown.Wing with posterior margin narrowly edged pale smoky grey. Shaft of aedeagus slender, weakly expanded subapically; dorsal surface subapically with four pairs ofspine-like processes; medial pair long, adjacent pair with apices weakly serrated, next pair stronglycurving. Paramere with apex obtusely rounded; dorsal process large, situated at midlength, apex decurvedand bearing numerous small spines; dorsal surface subbasally with a row of long, slender spines. MATERIAL EXAMINED Holotype d", Panama: Chir. Las Lagunas, 2-5 miles W. El Volcan, 4400 ft (O'Brien & Marshall)(FAMU). This species is distinguished by the reddish pigmentation of the abdomen, the carination of thefronto-lateral surfaces of the pronotum, and by the structure of the male genitalia. Pseudomysidia trinidadensissp. n. (Figs 46, 62, 78) Male: head 0-46 mm long, 0-57 mm wide; pronotum 1-07 mm wide; tegmen 5-70 mm long; wing 2-77 mmlong. Female unknown. Length of frons c. 12 times width at apex, 3-5 times width at base; ocelli small; clypeus c. one-third longerthan frons; rostrum extending to apex of abdomen. Pronotal width 6-5 times mid-dorsal length; fronto-lateral surfaces and tegula not carinate. Genae around ocelli and dorsad of eyes pale crimson; fronto-lateral surfaces of pronotum and disc ofmesonotum pale orange. Tegmen and wing whitish hyaline, veins and cross-veins pale yellow; cross-veinsand forks of veins broadly and irregularly edged very pale brownish. Tegmen with a broad, irregular, faint,brownish transverse band extending from cubital vein to posterior margin at one-quarter length. Shaft of aedeagus short, broad; dorsal surface subapically with four pairs of spine-like processes, medialpair each with a curving spine at midlength. Paramere slender, apex acute; dorsal process broad, truncate,situated at one-third length. MATERIAL EXAMINEDHolotype cf , Trinidad: Mt Harris (Withy combe) (BMNH). This species is distinguished by the paleness of the markings of the tegmen and wing, and by thestructure of the aedeagus. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 83 Pseudomysidia similis sp. n. (Figs 38, 54, 70) Male: head 0-46 mm long, 0-48 mm wide; pronotum 0-95 mm wide; legmen 5-40-5-60 mm long; wing2-60 mm long. Female: tegmen 6-00-6-70 mm long. Length of frons 14 times width at apex , 4 times width at base ; ocelli small , distinct ; clypeus slightly longerthan frons; rostrum extending to base of subgenital segment. Pronotal width 8 times mid-dorsal length;fronto-lateral surfaces and tegula with carinae obsolete or absent. Junction of vertex and frons often tinged brownish, ocelli yellowish brown or red. Tegmen and wingwhitish hyaline, veins yellowish, cross-veins broadly and irregularly edged smoky brown. Tegmen withpigmentation around cross-veins coalescing to form an irregular transverse band at one-quarter length, andat approximately two-fifths length. Wing with veins narrowly edged smoky brown apically. Shaft of aedeagus gradually broadening towards apex; dorsal surface subapically with six pairs ofspine-like processes, third pair from lateral margins finely serrated. Paramere slender basally; apexbroadly rounded; dorsal process situated at midlength, strongly produced posteriorly. MATERIAL EXAMINED Holotype cf , Panama: Barro Colorado, C.Z., 5.vii.l971 (Woldd) (FAMU).Paratype. 1 $ , same data as holotype (BMNH). This species is distinguished by the pigmentation of the head and tegmen, and by the structure ofthe male genitalia. Pseudomysidia marshallisp. n. (Figs 44, 60, 76) Male: head 0-46 mm long, 0-55 mm wide; pronotum 1-07 mm wide; tegmen 5-86 -6-03 mm long; wing3-00 mm long. Female: tegmen 6-70-7-00 mm long. Length of frons 13 times width at apex, 3-5 times width at base; ocelli small, occasionally obscure;clypeus one-fifth longer than frons; rostrum terminating slightly basad of subgenital segment. Pronotalwidth c. 1 times mid-dorsal length, fronto-lateral carinae distinct; tegula not carinate. Genae rarely tinged crimson ventral to eyes; ocelli commonly bright red. Tegmen and wing whitishhyaline, posterior margins broadly hyaline, veins pale; cross-veins and forks of veins darker, broadly edgedsmoky brown. Tegmen with markings around forks of veins coalescing to form very irregular transversebands at level of second fork of cubital vein, and at level of each of first, second and third forks of medialvein, the latter two not extending to posterior margin. Wing with a very irregular, pale smoky transverseband at level of radial-medial cross- vein. Shaft of aedeagus slender; dorsal surface subapically with four pairs of spine-like processes, medial pairtrifurcate. Paramere slender, apex acute; dorsal process situated at one-third length, truncate, weaklyproduced anteriorly; ventral surface over basal half length with long robust spines. MATERIAL EXAMINED Holotype cf , Costa Rica: Turrialba, 21. vi. 1974 (O'Brien & Marshall) (FAMU).Paratypes. Costa Rica: 1 cT, 12 $ (FAMU; BMNH). This species is most readily distinguished by the structure of the aedeagus. Pseudomysidia hindore sp. n. (Figs 39, 55, 71) Male: head 0-38 mm long, 0-42 mm wide; pronotum 1-05 mm wide; tegmen 5-10-5-70 mm long; wing2-80 mm long. Female unknown. Length of frons 12 times width at apex, c. 3 times width at base; ocelli obscure; clypeus c. as long as frons;rostrum extending to apex of abdomen. Pronotal width 8-5 times mid-dorsal length, fronto-lateral carinaedistinct; tegula not carinate. Head and body unmarked. Tegmen and wing whitish hyaline, veins and cross- veins pale yellowishbrown; cross-veins irregularly edged pale smoky brown. Tegmen with a weak, irregular, smoky brown,transverse band at one-third length. Shaft of aedeagus broadly expanded over apical one-half length; dorsal surface subapically with six pairs 84 PETER S. BROOMFIELD of spine-like processes, medial pair apically serrated, lateral pair inclined ventrally. Paramere slender,apex acutely rounded; dorsal process situated at midlength, large, posteriorly produced. MATERIAL EXAMINED Holotype cf , Panama: Turdi River, San Bias, i.1979 (Operation Drake Expedition) (BMNH).Paratype. Panama: 1 cf , C.Z., 7 km SW. Gatun Lock (FAMU). Though lacking distinctive pigmentation, this species may be readily distinguished by thestructure of the aedeagus. Pseudomysidiavestissp.n. (Figs 45, 61, 77) Male: head 0-42 mm long, 0-57 mm wide; pronotum 1-10 mm wide; tegmen 6-00 mm long; wing 2-50 mmlong. Female unknown. Length of frons c. 12 times width at apex, 3-5 times width at base; ocelli small, distinct; clypeusone-quarter longer than frons; rostrum extending to base of subgenital plate. Pronotal width c. 9 timesmid-dorsal length, fronto-lateral carinae distinct; tegula with carinae obscure. Head with genae dorsal and ventral to eyes, and lateral margins of clypeus bright scarlet; ocelli scarlet.Tegmen and wing clear hyaline, veins yellowish. Tegmen with an indistinct, irregular, oblique, smokybrown, transverse band at level of each of first, second and third forks of medial vein. Wing with a broad,faint, smoky brown, transverse band at one-third and two-thirds length; apex weakly smoky hyaline. Shaft of aedeagus robust; with six pairs of spine-like processes, of which the next to lateral pair arelongest. Paramere slender; apex narrowly rounded; dorsal process situated at approximately one-thirdlength, apically truncate. MATERIAL EXAMINED Holotype cf , Brazil: Amazon, Rio Purus, i.1915 (Roman) (NR).Paratype. Brazil: 1 cf , Amazon, S. Gabriel (BMNH). This species is distinguished by the pigmentation of the head and tegmen, and by the six pairs ofprocesses of the aedeagus. Pseudomysidia pallida sp. n. (Figs 14, 30, 42, 58, 74) Male: head 0-52 mm long, 0-50 mm wide; pronotum 1-06 mm wide; tegmen 5-90-6-12 mm long; wing2-90 mm long. Female: tegmen 6-20-6-80 mm long. Length of frons 15 times width at apex, 3 times width at base; ocelli small, distinct; clypeus slightly longerthan frons; rostrum terminating c. level with apex of abdomen. Pronotal width 7-5 times mid-dorsal length,fronto-lateral carinae distinct; tegula not carinate. Genae tinged reddish dorsal to eyes and around ocelli and bases of antennae, lateral surfaces of clypeusreddish brown; fronto-lateral surfaces of pronotum reddish, each with a dark reddish brown spot at level ofand adjacent to eye. Tegmen and wing whitish hyaline, veins pale brown. Tegmen with veins andcross-veins broadly and irregularly edged brownish. Wing with radial-medial cross-vein and adjacent forkof medial vein broadly edged smoky hyaline. Shaft of aedeagus slender, broadening subapically; dorsal surface subapically with four pairs of robustspine-like processes of which the longest pair are apically bifid. Paramere slender, apex acutely rounded;dorsal process situated at one-quarter length, large, truncate, weakly inclined anteriorly. MATERIAL EXAMINED Holotype cf , Ecuador: Mera, l-2.ii.1923 (Williams) (BMNH).Paratypes. 13 cf , 11 $, same data as holotype (BMNH; FAMU). This species is distinguished by the pigmentation of the head and pronotum, and by the structureof the aedeagus. Pseudomysidia panamensissp. n. (Figs 40, 56, 72) Male: head 0-40 mm long, 0-48 mm wide; pronotum 1-05 mm wide; tegmen 5-00-5-52 mm long; wing2-40 mm long. Female: tegmen 6-10 mm long. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 85 Length of frons 13 times width at apex, 3-5 times width at base; ocelli small, distinct; clypeus as long asfrons; rostrum extending to subgenital plate. Pronotal width 8-5 times mid-dorsal length, fronto-lateralcarinae weak; tegula not carinate. Genae crimson dorsad of eyes and from level of anterior margins of eyes to ventral margins, ocelli red.Fronto-lateral surfaces of pronotum crimson at and ventral to level of eyes; lateral surfaces of mesonotumand metanotum weakly crimson; disc of mesonotum pale brownish; dorsal surface of abdomen darkbrown, tinged red. Tegmen and wing whitish hyaline, veins and cross-veins pale brown; cross-veins andbranches of veins very broadly edged dark smoky brown. Tegmen with brownish markings coalescing toform a very irregular transverse band at level of first fork of cubital vein, with less distinct bands at level ofeach of first, second and third forks of medial vein. Wing with an irregular, transverse, smoky brown bandat level of radial-medial cross-vein, a weaker band near costal margin at level of first fork of cubital vein;apex faintly tinged smoky brown. Shaft of aedeagus slender throughout; dorsal surface subapically with three pairs of spine-like processes;medial pair very long and robust, bifurcate from midlength. Paramere basally slender, apex acutelyrounded; dorsal process situated slightly basad of midlength, truncate. MATERIAL EXAMINED Holotype cf , Panama: Cerro Campana, 29.vi.1974 (O'Brien & Marshall) (FAMU).Paratypes. 5 cf , 1 $, same data as holotype (FAMU; BMNH). This species is most readily distinguished by the structure of the aedeagus. Pseudomysidia ecuadorensis sp. n. (Figs 47, 63, 79) Male: head 0-46 mm long, 0-57 mm wide; pronotum 1-25 mm wide; tegmen 6-40-6-80 mm long; wing3-33 mm long. Female unknown. Length of frons 15 times width at apex; 3-33 times width at base; ocelli very small, distinct; clypeus c. aslong as frons; rostrum extending to midlength of genital segment. Pronotal width 7-5 times mid-dorsallength, fronto-lateral carinae weak; tegula not carinate. Genae pale crimson dorsad to eyes and around ocelli, ocelli crimson; fronto-lateral surfaces of pronotumreddish from level of eyes to ventral margins. Tegmen and wing whitish hyaline, veins and cross-veinsyellowish brown; cross-veins and forks of veins irregularly and broadly edged smoky brown. Tegmen withmarkings coalescing to produce a mottled appearance, forming an oblique, irregular, transverse band atlevel of second fork of cubital vein and at each of first, second and third forks of medial vein; posterior andapical margins broadly pale smoky brown. Wing with a faint, oblique, smoky brown transverse band atlevel of radial-medial cross-vein and at one-third length; apex broadly pale smoky brown. Shaft of aedeagus slender, gradually expanded to apex; dorsal surface subapically with five pairs ofspine-like processes. Paramere slender; apex narrowly rounded; dorsal process situated at one-quarterlength, truncate, apex antero-laterally directed. MATERIAL EXAMINED Holotype cf Ecuador: Tena, 14.ii.1923 (Williams) (BMNH).Paratypes. Ecuador: 2 cf , Tena (BMNH). This species is distinguished by the dark mottled appearance of the tegmen, and by the structureof the aedeagus. Pseudomysidia debora sp. n. (Figs 37, 53, 69) Male: head 0-40 mm long, 0-55 mm wide; pronotum 1-13 mm wide; tegmen 6-30 mm long; wing 3-10 mmlong. Female: tegmen 7-00-7-30 mm long. Length of frons 12 times width at apex, 4 times width at base; ocelli small, distinct; clypeus one-quarterlonger than frons; rostrum extending to base of subgenital plate. Pronotal width 11 times mid-dorsallength, fronto-lateral carinae very weak; tegula not carinate. Ocelli narrowly edged crimson. Tegmen and wing whitish hyaline, veins pale yellow; cross-veins andforks of veins brownish, broadly edged smoky brown. Tegmen with a brownish spot on radial cell at level offirst fork of medial vein, apex of clavus irregularly pale smoky brown. Wing with posterior and apicalmargins broadly and irregularly very pale smoky brown. 86 PETER S. BROOMFIELD Shaft of aedeagus slender; dorsal surface subapically with six pairs of spine-like processes, medial pairlongest with external surfaces basally serrated. Paramere basally slender, apex acutely rounded; dorsalprocess situated at approximately midlength, produced dorsally; ventral surface basally with numerousvery long, robust, spines. MATERIAL EXAMINED Holotype cf , Costa Rica: Turrialba, 28.V.1957 (Cartwright) (USNM).Paratypes. 4 $, same data as holotype (USNM; BMNH). This species is most readily distinguished by the structure of the male genitalia. Pseudomysidia lepida sp. n. Female: head 0-53 mm long, 0-59 mm wide; pronotum 1-32 mm wide; tegmen 6-63-6-80 mm long; wing3-40 mm long. Male unknown. Length of frons 16 times width at apex, c. 4 times width at base; ocelli small, prominent; clypeus slightlylonger than frons; rostrum extending to base of subgenital plate. Pronotal width 8 times mid-dorsal length,fronto-lateral carinae distinct; tegula not carinate. Genae dorsad and ventral to eyes orange; ocelli crimson. Tegmen and wing whitish hyaline, veinsirregularly alternating yellow and dark brown; cross- veins dark brown, these and forks of veins broadlyedged dark brownish grey. Tegmen with markings coalescing to form irregular, broad, transverse bands atone-seventh length, at level of first fork of cubital vein, at level of first, second and third forks of medialvein, and at somewhat basad of apical fork of medial vein. Wing with posterior and apical margins broadlydark greyish brown; a broad, irregular, transverse band at three-eighths length and another, oblique bandat level of medial-radial cross-vein. MATERIAL EXAMINED Holotype $, Bolivia: Beni, Rio Beni, Rurrenabaque, 270 m, 21.vii.1979 (Cooper) (BMNH).Paratypes. 1 $, same data as holotype (BMNH). This species is readily distinguished by the very striking markings of the tegmen and wing. Pseudomysidia obnubilia sp. n. (Figs 49, 65, 81) Male: head 0-52 mm long, 0-60 mm wide; pronotum 1-24 mm wide; tegmen 5-70-6-40 mm long; wing2-84 mm long. Female: tegmen 6-12-6-88 mm long. Length of frons 14 times width at apex, 3-33 times width at base; ocelli small, prominent; clypeus slightlylonger than frons; rostrum extending to apex of abdomen. Pronotal width 6-5 times mid-dorsal length,fronto-lateral carinae distinct; tegula not carinate. Genae dorsad and ventrad of eyes orange, ocelli crimson; abdomen occasionally red on mid-dorsal line.Tegmen and wing whitish hyaline, veins yellow, cross-veins dark brown. Tegmen with veins andcross-veins broadly and irregularly edged smoky brown, these markings coalescing to form very irregulartransverse bands at one-sixth length, at level of first fork of cubital vein, at each of first and second branchesof medial vein and at two-fifths length. Wing with posterior and apical margins broadly smoky brown; abroad, irregular, smoky brown, transverse band at two-thirds length. Shaft of aedeagus slender, with five pairs of long, spine-like processes. Paramere slender, apex acutelyrounded; dorsal process situated at one-third length, truncate. MATERIAL EXAMINED Holotype cf , Ecuador: 18 km S. Tena, 28.iv.1978 (O'Brien & Marshall) (FAMU).Paratypes. 3 cf , 4 <j>, same data as holotype (FAMU; BMNH). Externally very similar to delicata, this species is readily distinguished by the male genitalia. Pseudomysidia delicata sp. n. (Figs 48, 64, 80) Male: head 0-52 mm long, 0-55 mm wide; pronotum 1-10 mm wide; tegmen 5-60-6-40 mm long; wing2-80 mm long. Female: tegmen 5-85-6-40 mm long. Length of frons 12 times width at apex, 2-75 times width at base; ocelli small, distinct; clypeus TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 87 one-quarter longer than frons; rostrum extending to base of genital segment. Pronotal width c. 1 timesmid-dorsal length, fronto-lateral carinae distinct; tegula not carinate. Genae crimson dorsal to eyes and from level of anterior margins of eyes to ventral margins; clypeus withlateral margins often pale crimson or brown; ocelli red. Tegmen and wing whitish hyaline, veins pale;cross-veins and bases of branches of veins dark brown, irregularly broadly edged smoky brown; posteriorand apical margins broadly pale brownish. Tegmen with dark markings coalescing to form irregulartransverse bands at each of first , second and third branches of medial vein. Wing with an irregular, oblique,transverse, smoky brown band at level of medial-cubital cross-vein and at one-third length; apex broadlysmoky brown. Shaft of aedeagus slender, slightly expanded subapically; dorsal surface subapically with five pairs ofrobust, spine-like processes. Paramere basally slender, apex obtusely rounded; dorsal process situated atone-third length, truncate. MATERIAL EXAMINED Holotype cf , Ecuador: Tena, 4.iii.l923 (Williams) (BMNH).Paratypes. 10 cf , 21 $, same locality as holotype (BMNH). Though externally similar to obnubilia, this species is readily distinguished by the structure ofthe aedeagus. DYSIMIA MuirDysimia Muir, 1924: 462. Type-species: Dysimia maculata Muir, by monotypy. Head in dorsal aspect one-quarter to two-thirds wider than long. Lateral margins of vertex stronglyconverging from base to level of anterior margins of eyes, then subparallel to apex, carinae very prominent,extending beyond anterior margins of eyes for one-quarter to one-third length; basal margin very deeplyincised; junction with frons broadly rounded. Frons 6-9 times as long as wide at apex, 1-5-2-0 times widthat base; lateral margins strongly and regularly diverging from apex to base. Genae extending anterior toeyes for one-third to two-thirds horizontal diameter of eye. Antenna with second segment club-shaped,twice as long as broad; apex transverse; flagellum arising medially. Ocelli small, usually distinct;occasionally obscure or obsolete. Clypeus short, broad; length from three-quarters of to equal that offrons, from c. equal to up to one-half greater than width at base; medial carina obsolete or absent; lateralcarinae usually obsolete or absent, occasionally weak and extending over basal c. one-third length.Rostrum usually terminating at level of posterior surfaces of hind coxae, occasionally slightly shorter. Pronotal width 8-12 times mid-dorsal length; fronto-lateral surfaces each with a single, distinct,occasionally prominent carina extending horizontally from adjacent to eye to lateral margin. Tegula rarelycarinate. Disc of mesonotum c. as long as wide; medial carina percurrent, often weak or obsolete, rarelyprominent; lateral carinae commonly obsolete or absent, rarely distinct. Tegmen commonly 3-20-4-40 mm long, rarely more than 6-00 mm long. Medial vein separating fromfused subcostal and radial veins at c. one-quarter length; subcostal and radial veins separating at somewhatbasad of midlength. Radial vein with two branches extending to apical margin. Medial vein with sevenbranches extending to apical and posterior margins; with cross- veins between first and second and secondand third branches. Cubital vein with three branches extending to posterior margin; anterior branch linkedby a cross-vein to first medial vein at c. midlength (Fig. 9.) Length of wing greater than one-half that of legmen. Subcostal and radial veins fused over rather morethan basal one-half length. Radial vein unbranched. Medial vein distinct from base, unbranched, linked toradial vein by a cross-vein at c. two-thirds length. Cubital vein with three branches extending to posteriormargin, linked to medial vein by a cross-vein at slightly distad of midlength. Head and body pale, yellowish or brownish. Genae frequently each with a brown band extendinghorizontally from level of dorsal margin of eye to anterior margin, a similar band at level of ventral marginof eye, rarely unmarked; area around ocelli occasionally tinged brownish. Frons and clypeus rarely withdark markings. Fronto-lateral surfaces of pronotum usually each with a broad brown band extendinghorizontally from adjacent to eye to lateral margin. Tegmen and wing whitish hyaline; veins pale yellow;cross-veins and forks of veins brownish, often broadly edged smoky brown; apical and posterior marginalveins often crimson, or flecked with red. Tegmen with a large, often prominent, black or brownish, roughlycircular spot at one-fifth to one-third length over first branch of cubital vein; often with smaller but equallydistinct spots between cubital vein and clavus, adjacent to point of separation of subcostal and radial veinsand around apical fork of medial vein. Wing often with a large, prominent, circular, dark brown spotbetween cubital vein and clavus at c. one-third length; where marking is absent, often with a pale browntransverse band at one-third length. 88 PETER S. BROOMFIELD Male genitalia with shaft of aedeagus horizontal, slender, cylindrical, usually symmetrical; dorsalsurface subapically with a pair of large, anteriorly directed, flap-like processes, often bearing from one tothree pairs of large, curving, anteriorly or antero-laterally directed spines; ventral surface unarmed.Paramere commonly slender, rarely very robust; apex frequently acutely rounded and directed towardsmidline; basal apodeme not more than one-third total length; dorsal process situated at or distad tomidlength, usually well developed with apex strongly produced posteriorly; interlocking surfaces usuallywell developed, situated basally or at midlength, rarely reduced or absent; dorsal surface basad of mainprocess usually with a large, internally directed, secondary process; ventral surface unarmed. Anal tubewith posterior margin rounded, more or less strongly produced, often deeply notched at midline. Female with posterior margin of subgenital plate more or less strongly produced medially, broadly andregularly convex, rarely truncate apically. The genus is distinguished by the seven branches of the medial vein of the tegmen, the threebranches of the cubital vein of the wing and the apical position of the antennal flagellum. Thislast character and the proportions of the frons and clypeus show a closer affinity to Pseudomy-sidia than to Mysidia, though it is apparently considerably more specialised than the former. The male genitalia show three possible diverging lines of development within the genus, whichallow the postulation of the following species groups. The maculata-gToup. Species where the shaft of the aedeagus is lacking large spines, with acorresponding increase in the size and armature of the paramere. The group includes distincta,fennahi, pseudomaculata and telfordi, and is probably the most highly specialized of the three. The astarte-group. Species where the shaft of the aedeagus bears strong, heavily chitinized,spine-like processes, and the paramere shows a lesser degree of development. This groupincludes morrisi, muiri, obrieni, maculipennis and jamaicensis . The numa-group. This monotypic group shows a separate line of development from theabove. The aedeagus is heavily armed with three pairs of long, acute, spine-like processes, andthe dorsal process of the paramere is greatly reduced and can have little grasping function. Due to the absence of males, fuscoclypea ta andputilla are omitted from the above groups. The genus is distributed from U.S.A. (Florida) through Central America to Ecuador andVenezuela. Key to species of Dysimia (based on external characters) D. putilla is omitted from this key due to the fragmentary condition of the unique holotype. In someinstances the differences between species are slight and, where possible, reference should be made to thestructure of the male genitalia. 1 Tegmen exceeding 6 mm. Jamaica jamaicensis (Distant) (p. 91) Tegmen less than 5 mm 2 2(1) Wing with a prominent, dark brown, spot adjacent to cubital vein 6 Wing lacking a distinct spot adjacent to cubital vein 3 3(2) Genae with dark brown markings adjacent to eye 4 Genae unmarked, pale brownish throughout. Florida pseudomaculata sp. n. (p. 91) 4(3) Tegmen with cross-veins and apical veins dark brown. Costa Rica obrieni sp. n. (p. 92) Tegmen with cross-veins and apical veins pale 5 5(4) Male tegmen in excess of 4-00 mm; female with tegmen in excess of 4-50 mm. Tegmen with posterior margin narrowly scarlet. Ecuador morrisi sp. n. (p. 93) Neither sex with tegmen exceeding 4-00 mm. Tegmen with posterior margin pale. Jamaica muiri sp. n. (p. 92)6(2) Clypeus narrowly pale basally, thence dark brown to junction with paraclypeus. Venezuela fuscoclypeata Fennah (p. 90) Clypeus pale throughout 7 7(6) Genae each with one or two dark brown bands extending horizontally from adjacent to eye to anterior margin. Pronotum dorsad of eyes unmarked 8 Genae unmarked. Pronotum dorsad of base of head with two very pale fuscous, parallel, transverse bands. Cayman Islands numa Fennah (p. 90) 8(7) Abdomen with a large, dark brown, spot on either side of midline on dorsal surface. Puerto Rico distincta sp. n. (p. 93), fennahisp. n. (p. 94), telfordisp. n. (p. 94) Abdomen with dorsal surface unmarked . . 9 TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 89 9(8) Disc of mesonotum ventrolaterally marked brownish. Venezuela astartesp. n. (p. 95) Disc of mesonotum unicolorous yellowish throughout 10 10(9) Pronto-lateral surfaces of pronotum ventral to level of dorsal margins of eye dark brown. Ecuador macuJipennis sp. n. (p. 95) Pronto-lateral surfaces of pronotum pale brown throughout. Puerto Rico maculate Muir (p. 89) Key to species of Dysimia (based on male genitalia) It has not been possible to examine males oifuscoclypeata andputilla, which are omitted from this key. 1 Paramere with dorsal process bearing a large hook-like spine on dorsal surface (Fig. 1 18) distincte sp. n. (p. 93)Paramere with dorsal process unarmed on dorsal surface 2 2 (1) Shaft of aedeagus with three pairs of spine-like processes on dorsal surface subapically (Fig. 95). Paramere with dorsal process reduced (Fig. 119) numa Fennah (p. 90) Shaft of aedeagus with not more than two pairs of spine-like processes. Paramere withdorsal process well developed 3 3 (2) Shaft of aedeagus devoid of large spine-like processes 4 Shaft of aedeagus bearing one or two pairs of large, anteriorly directed, spine-like processes subapically 8 4(3) Shaft of aedeagus with dorsal surface with a pair of anteriorly directed , flap-like , processessubapically, each bearing a very small, antero-laterally directed spine (Fig. 96) maculate Muir (p. 89)Shaft of aedeagus with flap-like processes unarmed 5 5 (4) Paramere slender, apex acute, dorsal process situated at mid-length (Fig. 121) 6 Paramere robust , dorsal process arising well distad of mid-length (Fig. 123) 7 6 (5) Paramere with apex of dorsal process horizontally directed; inter-locking surfaces poorly developed (Fig. 121) pseudomaculate sp. n. (p. 91) Paramere with apex of dorsal process inclined ventrally; interlocking surfaces welldeveloped (Fig. 122) muiri sp. n. (p. 92) 7 (5) Paramere very robust; apex strongly curved towards midline; dorsal process very long and slender; interlocking surfaces situated basally (Fig. 123) fennahi sp. n. (p. 94) Paramere relatively small; apex rounded, not curved towards midline; dorsal processrobust; interlocking surfaces situated at mid-length (Fig. 124) telfordi sp. n. (p. 94) 8 (3) Shaft of aedeagus with two pairs of spines subapically on dorsal surface (Fig. 101) macuJipennis sp. n. (p. 95)Shaft of aedeagus with one pair or three spines subapically on dorsal surface 9 9 (8) Shaft of aedeagus bearing, in addition to a pair of long, slender, acute spines subapically, a single, short, apically obtuse spine at midline (Fig. 102) obrieni sp. n. (p. 92) Shaft of aedeagus with a single pair of long, curving, spines only 10 10 (9) Shaft of aedeagus long and slender ; apical spines simple , long and curving 11 Shaft of aedeagus short and broad; apical spines overlaying large, flap-like, anteriorlydirected processes, each of which bears a short, antero-laterally directed, apically bifurcated, subsidiary process baso-laterally (Fig. 103) jamaicensis Distant (p. 91) 11(10) Paramere with a large, rounded, secondary process on dorsal surface subbasally (Fig. 128) morrisi sp. n. (p. 93)Paramere without a secondary process (Fig. 129) asterte sp. n. (p. 95) Dysimia maculata Muir (Figs 9, 13,29,96, 108, 120)Dysimia maculata Muir, 1924: 463. Holotype C?, PUERTO Rico (BPBM) [examined]. Male: head 0-30 mm long, 0-40 mm wide; pronotum 0-80 mm wide; tegmen 3-60-3-75 mm long; wing2-30 mm long. Female: tegmen 3-80-4-40 mm long. Length of frons c. 8 times width at apex, 1-66 times width at base; ocelli small, distinct; clypeus c.three-quarters length of frons. Pronotal width c. 12 times mid-dorsal length. Genae each with a broad, brown band at level of eye, darker at dorsal and ventral margins, extendinghorizontally to anterior margin; area around ocelli frequently brownish; frons with base occasionally darkbrown. Pronto-lateral surfaces of pronotum each with a brownish band extending horizontally from 90 PETER S. BROOMFIELD adjacent to eye to external margin. Tegmen and wing with bases of branches of veins and cross- veinsbrownish, cross-veins edged pale smoky brown; posterior and apical margins frequently very narrowlyedged scarlet. Tegmen with a large, circular, dark brown spot over first branch of cubital vein at one-thirdlength; a smaller spot between cubital vein and clavus at one-sixth length; another adjacent to andimmediately basad of subcostal and fused radial-medial fork; a third small spot, more irregular and lessdistinct, around fused subcostal-radial-medial vein at one-third length. Wing with a prominent, circular,dark brown spot between cubital vein and clavus at one-quarter length. Shaft of aedeagus slender , with two pairs of flap- like processes , the external pair each with a small , blunt ,spine on ventral surface. Paramere with dorsal process situated slightly distad of midlength, slender,strongly produced postero-dorsally; dorsal surface at one-quarter length with a slender, regularly tapering,secondary process bearing long, robust, spines. MATERIAL EXAMINED Holotype cf , Puerto Rico: Rio Piedras, viii.1923 (Wolcott) (BPBM).Paratypes. Puerto Rico: 26 cf , 13 $, same data as holotype (BMNH); 1 cf (USNM). This species has also been recorded from Haiti (Dozier, 1922). The specimens recorded by Ball(1928) from Florida are the newly described species pseudomaculata (p. 00). The two species arereadily distinguished by the absence of the dark spot on the wing of pseudomaculata, and by themale genitalia. Dysimht fuscoclypeata Fennah Dysimia fuscodypeata Fennah, 1952: 124. Holotype $, VENEZUELA: Aragua, Rancho Grande, 1100 m,l.x.1950 (Yepez) (lost). NEOTYPE $, VENEZUELA, here designated (BMNH) [examined]. Female: head 0-30 mm long, 0-42 mm wide; pronotum 0-86 mm wide; legmen 4-40 mm long; wing 2-60 mmlong. Male unknown. Length of frons 7 times width at apex, 1-5 times width at base; ocelli distinct; clypeus c. as long as frons,1-5 times width at base. Pronotal width c. 8 times mid-dorsal length; tegula weakly carinate. Frons with apical half somewhat darker than basal half, with a dark brown transverse band at level ofdorsal margin of eye and another at midline; genae irregularly reddish brown, with a dull brown bandrunning from level of midline of eye to anterior margin; clypeus with basal quarter whitish, thence brown tojunction with pale yellowish paraclypeus. Pronto-lateral surfaces of pronotum each with a broad brownband extending horizontally from adjacent to eye, terminating short of external margin; metanotum withfronto-lateral surfaces at level of clypeus pale brown. Tegmen and wing with cross-veins narrowly edgedsmoky brown. Tegmen with a very large, prominent, dark brown spot on first branch of cubital vein atone-third length; a much smaller spot adjacent and anterior to medial vein at equi-distance from base; twosmall spots at one-seventh length, one between fused subcostal, radial and medial veins and costal margin,and the other immediately posterior to cubital vein. Wing with a single, irregular, dark brown spot attwo-fifths length, adjacent to and posterior to cubital vein. MATERIAL EXAMINEDNeotype $ , Venezuela: Aragua, Camino, Choroni, 950 m, montane forest, 24.iii.1949 (Box) (BMNH). This is the only specimen available for study. A pin with a label bearing the data of the holotypeis in the BMNH, but the specimen is missing. With Dr Fennah's agreement I therefore designatethe single paratype as a neotype. As the name suggests, fuscodypeata is readily distinguished by the dark pigmentation of theclypeus. Dysimia numa Fennah (Figs 95, 107, 119)Dysimia numa Fennah, 1971: 324. Holotype cf , CAYMAN ISLANDS (BMNH) [examined]. Male: head 0-28 mm long, 0-40 mm wide; pronotum 0-72 mm wide; tegmen 3-60-4-20 mm long; wing2-00 mm long. Female: tegmen 4-00-4-60 mm long. Length of frons 6 times width at apex, 1-5 times width at base; ocelli small, distinct; clypeus as long asfrons. Pronotal width 10 times mid-dorsal length. Head unmarked; pronotum with a pale fuscous transverse band parallel to posterior margin on dorsal TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 91 surface; fronto-lateral surfaces each with a very faint fuscous band extending horizontally from adjacent todorsal margin of eye to lateral margin. Tegmen and wing with cross-veins and forks of veins faintly edgedbrownish. Tegmen with a large, prominent, roughly circular, brown spot over first branch of cubital vein atthree-fifths length; a smaller, paler, irregular marking over apical fork of medial vein; two smaller spots,one between cubital vein and clavus, the other between point of separation of medial vein from fusedsubcostal-radial veins and costal margin; an irregular spot over medial and fused subcostal-radial veins atthree-eighths length. Wing with a small dark brown spot immediately basad of one-third length, adjacentto cubital vein. Shaft of aedeagus laterally expanded over apical one-third length, with three pairs of large spine-likeprocesses. Paramere slender; apex acute; dorsal process situated at two-thirds length, greatly reduced,with a posteriorly directed spine on posterior surface at midlength; dorsal surface at one-third length withan apically rounded secondary process. MATERIAL EXAMINED Holotype cf , Grand Cayman: N. Coast, N. Side, Hut Rd, 15.vii.1938 (Lewis & Thompson) (BMNH).Paratypes. Cayman Brae: 38 cf , 33 $ , Statae [?] Bay (BMNH). In his description Fennah erroneously refers to a single pair of spines on the aedeagus; hisillustration correctly shows three. This species is distinguished by the lack of dark pigmentation on the genae and pronotum, andby the structure of the male genitalia. Dysimiajamaicensis Distant comb. n.(Figs 103, 115, 127) Mysidia jamaicensis Distant, 1907: 396. LECTOTYPE cf , JAMAICA (BMNH), here designated [ex-amined] . Male: head 0-38 mm long, 0-59 mm wide; pronotum 1-20 mm wide; tegmen 6-40 mm long; wing 3-50 mmlong. Female unknown. Length of frons 6 times width at apex, twice width at base; ocelli small, distinct; clypeus two-thirds longerthan frons. Pronotal width 12 times mid-dorsal length; tegula carinate. Genae and fronto-lateral surfaces of pronotum at level of eyes brown. Tegmen and wing with veins andcross-veins narrowly and irregularly edged pale brownish, posterior margins between veins broadly smokybrown, posterior marginal veins crimson. Tegmen with a large irregular brown spot over fork of secondbranch of medial vein; a smaller spot over apical fork of medial vein; another larger spot between firstbranch of cubital vein and vanal fold at approximately three-quarters length of clavus. Wing withcross-veins brown; an indistinct, broken, pale brownish, transverse band at one-third length. Shaft of aedeagus bearing a pair of large, flap-like processes, each with an adjacent spine-like processdorsally and a small, apically bifurcate spine laterally. Paramere very slender; apex acute; dorsal processsituated at midlength, apex posteriorly directed and reduced. MATERIAL EXAMINEDLectotype cf , Jamaica: Moneague, 10.ii.1905 (Nicholl) (BMNH). This species is included in Dysimia because of the tegminal venation and the proportions of thehead. Its relatively large size and long wings, with the reduction in the dorsal process of theparamere and complex armature of the aedeagus, readily distinguish it. Dysimia pseudomaculata sp. n. (Figs 97, 109, 121)[Dysimia maculata Muir sensu Ball, 1928: 199. Misidentification.] Male: head 0-30 mm long, 0-40 mm wide; pronotum 0-84 mm wide; tegmen 3-58-3-78 mm long; wing2-40 mm long. Female: tegmen 4-20-4-40 mm long. Length of frons 6 times width at apex, 1-5 times width at base; ocelli small or obsolete; clypeus slightlyshorter than frons. Pronotal width 10 times mid-dorsal length. Head ventral to dorsal margins of eyes brown, genae lacking distinct darker markings. Fronto-lateralsurfaces of pronotum ventral to horizontal carinae pale dull brownish. Tegmen and wing with veins andcross-veins predominantly pale brown; marginal veins very narrowly crimson, broadly and irregularly 92 PETER S. BROOMFIELD edged pale smoky brown. Tegmen with a very large, prominent, circular, dark brown spot over first branchof cubital vein immediately basad of level of apex of clavus; a small, dark brown spot between cubital veinand claval suture subbasally; a somewhat smaller spot on costal cell immediately basad of point ofseparation of fused radial and medial veins; a small, indistinct spot over medial vein at one-third length; anirregular, fainter spot covering apical forks of medial and radial veins; bases of branches of medial veinoccasionally dark brown. Wing lacking a dark brown spot between cubital vein and clavus; irregularlytinged pale smoky brown at approximately one-third length, level of cubital fork, and over apicalone-quarter length. Shaft of aedeagus strongly dorsally and laterally expanded subapically, devoid of spines. Paramereslender; apex acute; dorsal process situated at midlength, apex strongly produced posteriorly; dorsalsurface slightly basad of one-third length with a secondary process bearing a few long, robust, spines. MATERIAL EXAMINED Holotype cT, U.S.A.: Florida, Sanford, 19.xi.1927 (Ball) (USNM).Paratypes. 4 cf , 3 $, same data as holotype (USNM; BMNH). The genitalic differences are consistent, though not great, but the lack of a prominent dark spoton the wing of pseudomaculata renders the species readily distinguishable. Dysimia muiri sp. n. (Figs 98, 110, 122) Male: head 0-24 mm long, 0-41 mm wide; pronotum 0-70 mm wide; tegmen 3-50-3-60 mm long; wing2-15 mm long. Female: tegmen 3-60-4-00 mm long. Length of frons 8 times width at apex, twice width at base; ocelli obscure; clypeus slightly shorter thanfrons. Pronotal width 10 times mid-dorsal length. Genae each with a horizontal dark brown band extending from adjacent to dorsal margin of eye toanterior margin, a similar band at level of ventral margin of eye; frons occasionally dark brown betweenlateral carinae; fronto-lateral surfaces of pronotum each with a pale brown horizontal band extending fromadjacent to eye to lateral margin. Tegmen and wing with veins and cross-veins prominently edgedgrey-brown, apical and posterior margins fuscous between veins. Tegmen with a very large, prominent,roughly circular, dark brown spot near posterior margin between first branch of cubital vein and clavus; asmaller, brownish spot on medial vein at equal distance from base. Wing lacking distinct spots. Shaft of aedeagus slender; apex laterally expanded, devoid of spines or flap-like processes. Paramereslender, apex acutely rounded; dorsal process large, situated at midlength, greatly produced posteriorly;dorsal surface subbasally with a membranous secondary process. MATERIAL EXAMINED Holotype d", Jamaica: Portland, Somerset Falls, 8.xii.l975 (O'Brien & Marshall) (FAMU).Paratypes. 28 O", 19 $ , same data as holotype (FAMU; BMNH). This species is readily distinguished by the pigmentation of the tegmen, lack of dark spots on thewing, and by the structure of the male genitalia. Dysimia obrienisp. n. (Figs 102, 114, 126) Male: head 0-26 mm long, 0-36 mm wide; pronotum 0-70 mm wide; tegmen 3-60 mm long; wing 2-00 mmlong. Female: tegmen 3-90-4-00 mm long. Length of frons 7 times width at apex, c. twice width at base; ocelli obsolete; clypeus slightly shorter thanfrons. Pronotal width 10 times mid-dorsal length. Genae each with a dark brown band extending from level of dorsal margin of eye to anterior margin, asimilar band level with ventral margin of eye. Tegmen and wing with veins, cross-veins and posteriormargins narrowly edged smoky brown; apical cells fuscous; posterior marginal veins very narrowly scarlet.Tegmen with a prominent, roughly circular, dark brown spot on first branch of cubital vein at level of apexof clavus; a smaller spot near base of cubital vein; another immediately basad of point of separation offused subcostal, radial and medial veins; a fourth at c. midlength. Wing lacking conspicuous spots. Shaft of aedeagus slightly asymmetrical; with a pair of large, flap-like processes, each bearing a single,large, curving spine; a slender acute spine medially. Paramere very slender; dorsal process situated atmidlength, apex produced postero-dorsally; dorsal surface subbasally with a conical secondary process. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 93 MATERIAL EXAMINED Holotype cf , Costa Rica: Turialba, 2l.vi.l914 (O'Brien & Marshall) (FAMU).Paratypes. 3 $ , same data as holotype (FAMU; BMNH). This species is readily distinguished by the prominent brown spot on the first branch of thecubital vein of the tegmen coupled with the absence of a corresponding spot on the wing, and bythe structure of the male genitalia. Dysimia morrisisp. n. (Figs 104, 116, 128) Male: head 0-29 mm long, 0-47 mm wide; pronotum 0-88 mm wide; tegmen 4-20-4-30 mm long; wing2-40 mm long. Female: tegmen 4-50-4-60 mm long. Length of frons 8 times width at apex, 1-5 times width at base; ocelli indistinct; clypeus slightly shorterthan frons. Pronotal width slightly less than 10 times mid-dorsal width. Genae each with a broad, dark brown band extending horizontally from level of dorsal margin of eye toanterior margin, another at level of ventral margin of eye. Pronto-lateral surfaces of pronotum each with abroad brownish band extending from adjacent to eye to exterior margin. Tegmen and wing with cross veinsnarrowly edged pale brown, posterior margins very narrowly scarlet. Tegmen with a large, roughlycircular, dark brown spot on first branch of cubital vein at level with apex of clavus; a much smaller spot onmedial vein at same level; another at two-thirds length of fused subcostal, radial, and medial veins; anotherof intermediate size between cubital vein and vanal fold at midlength between base and first fork; areaaround apical fork of medial vein irregularly pale brown. Wing lacking distinct dark spots. Shaft of aedeagus slightly asymmetrical; a pair of large, flap-like processes, each bearing a single, large,curving spine mid-dorsally. Paramere slender, apex narrowly rounded; dorsal process large, situated atmidlength, apex slender, strongly produced posteriorly; dorsal surface at one-quarter length with a largerounded secondary process bearing long robust spines. MATERIAL EXAMINED Holotype cT, Ecuador: Naranjapata, 1850 ft, xii.1922 (Williams) (BMNH).Paratypes. 4 cf , 4 $ , same data as holotype (BMNH). The pigmentation of this species closely resembles that of obrieni, but it is distinguished by itslarger size and by the structure of the male genitalia. Dysimia distinctasp. n. (Figs 94, 106, 118) Male: head 0-30 mm long, 0-44 mm wide; pronotum 0-76 mm wide; tegmen 3-60-3-80 mm long; wing2-20 mm long. Female unknown. Length of frons c. 8 times width at apex, twice width at base; ocelli distinct; clypeus c. three-quarterslength of frons. Pronotal width c. 9 times mid-dorsal length. Genae each with a dark brown band extending horizontally from level of dorsal margin of eye to anteriormargin; another similar band at level of ventral margin of eye. Pronto-lateral surfaces of pronotum eachwith a broad brown band extending from adjacent to eye horizontally to exterior margin; tegula withventral margin narrowly brown; abdomen with a large, circular, dark brown spot on either side of midlinesubapically on dorsal surface. Tegmen and wing with cross-veins broadly edged very pale brown. Tegmenwith anterior and posterior margins very narrowly and intermittently tinged reddish; a large, circular, darkbrown spot on cubital vein at one-third length; a somewhat smaller spot on medial vein equidistant frombase; an irregular spot over apical fork of medial vein; two smaller spots, one adjacent to fused subcostal,radial and medial veins at one-fifth length, the other posterior to cubital vein equidistant from base. Wingwith posterior margin narrowly tinged reddish, a small brown spot immediately posterior to cubital vein atone-third length. Shaft of aedeagus lacking spine-like processes, laterally somewhat expanded over apical one-half length;dorsal surface subapically with a transverse, membranous, flap-like process, partially overlying a pair ofslender, projections situated one on either side of midline. Paramere complex, robust; dorsal process verylarge, situated immediately distad of mid-length, apex strongly produced posteriorly, bearing a large,robust, internally curving, hook-like process dorsally; dorsal surface basad of midlength with a shallowlybifurcate secondary process. 94 PETER S. BROOMFIELD MATERIAL EXAMINED Holotype cf , Puerto Rico: 8 miles E. of Mayaguez, 9.ii.l969 (O'Brien) (FAMU).Paratype. Puerto Rico: 1 cf , Rio Piedras, 4.xii.l968 (Telford & Medina) (BMNH). Although very similar in external characters to fennahi and telfordi, this species is readilydistinguished by the unique development of the paramere, which here reaches a degree ofcomplexity not seen elsewhere during the present study. Dysimia fennahi sp. n. Male: head 0-29 mm long, 0-42 mm wide; pronotum 0-80 mm wide; tegmen 3-60 mm long; wing 2-00 mmlong. Female unknown. Length of frons c. 8 times width at apex, twice width at base; ocelli obscure; clypeus three-quarterslength of frons. Pronotal width slightly less than 8 times mid-dorsal length. Genae each with a dark brown band running horizontally from level of dorsal margin of eye to anteriormargin , another similar parallel band at level of ventral margin of eye . Pronto-lateral surfaces of pronotumeach with a broad dark brown band running horizontally from adjacent to eye to exterior margin; tegulawith ventral margin broadly dark brown; abdomen with a large, rather irregular, dark brown spot on eitherside of midline subbasally on dorsal surface. Tegmen and wing with veins and cross-veins narrowly andintermittently edged brownish, anterior and posterior margins very narrowly and irregularly flecked palereddish. Tegmen with a large, roughly circular, dark brown spot on first branch of cubital vein atapproximately one-third length; a somewhat smaller spot on medial vein equidistant from base; anirregularly shaped, brownish marking over apical fork of medial vein; two small, dark brown spots, oneadjacent to fused subcostal, radial and medial veins, the other on cubital vein subbasally. Wing with asmall, circular, dark brown spot adjacent to cubital vein at one-third length. Shaft of aedeagus with apical third length weakly expanded dorso-late rally; dorsal surface subapicallywith a large, transverse, flap-like process and a pair of spine-like processes adjacent to midline. Parameremassive; apex acutely rounded; dorsal process situated at two-thirds length, apex slender and stronglyproduced postero-dorsally; dorsal surface with a well-developed secondary process slightly basad ofmidlength. MATERIAL EXAMINEDHolotype cf , Puerto Rico: 8 miles E. of Mayaguez, 9.H.1969 (O'Brien) (FAMU). Externally, this species closely resembles telfordi, from which it is most readily distinguished bythe structure of the male genitalia, where the reduction in the armature of the aedeagus isaccompanied by massive development of the paramere. Dysimia telfordi sp. n. (Figs 100, 112, 124) Male: head 0-26 mm long, 0-42 mm wide; pronotum 0-75 mm wide; tegmen 3-40-3-60 mm long; wing2-16 mm long. Female unknown. Length of frons 7 times width at apex, twice width at base; ocelli indistinct; clypeus three-quarters lengthof frons. Pronotal width c. 8 times mid-dorsal length. Genae each with a dark brown band extending horizontally from adjacent to dorsal margin of eye toanterior margin, a similar band at level of ventral margin of eye, these bands continued over frons;fronto-lateral surfaces of pronotum each with a broad pale brown band extending horizontally fromadjacent to eye to lateral margin; tegula with ventral margin dark brown; abdomen with a large, roughlycircular, dark brown spot on either side of midline subapically on dorsal surface. Tegmen and wing withcross-veins narrowly edged pale smoky brown. Tegmen with anterior and posterior marginal veinscrimson; a large, circular, dark brown spot on first branch of cubital vein at one-third length; a rathersimilar spot on medial vein equidistant from base; another over apical fork of medial vein; two smallerspots, one immediately anterior to point of separation of fused subcostal, radial and medial veins, the otherposterior to cubital vein subbasally. Wing with posterior marginal vein crimson; a small, distinct, circular,brown spot immediately posterior to cubital vein at one-third length. Shaft of aedeagus weakly expanded over apical two-fifths length; dorsal surface subapically with atransverse, flap-like, membranous projection, partially obscuring a pair of short, blunt, spine-likeprocesses medially. Paramere robust; dorsal process situated at three-quarters length, strongly producedposteriorly; dorsal surface at midlength with an apically bifurcate secondary process. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 95 MATERIAL EXAMINED Holotype cf , Puerto Rico: Rio Piedras, 4.xii.l968 (Telford & Medina) (FAMU).Paratypes. 5 cf , 1 $ , same data as holotype (FAMU; BMNH). Closely resembling fennahi in external characters, this species may be distinguished by itssmaller size; though the structure of the aedeagus differs only slightly, that of the paramere isquite distinct. Dysimia astarte sp. n. (Figs 105, 117, 129) Male: head 0-29 mm long, 0-40 mm wide; pronotum 0-74 mm wide; tegmen 3-80-3-95 mm long; wing2-30 mm long. Female: tegmen 4-10-4-40 mm long. Length of frons c. 6 times width at base; c. 2-5 times width at base; ocelli large, not prominent; clypeustwo-thirds length of frons. Pronotal width 9 times mid-dorsal length. Genae each bearing an irregular dark brown band adjacent to dorsal margin of eye, this band oftenextending onto frons, a similar, broader band level with midline of eye; ocelli pale; apices of secondantennal segments brownish. Pronto-lateral surfaces of pronotum adjacent to eyes narrowly and irregular-ly dark brown; mesonotum with a large, irregular, pale brown spot on each side ventro-laterally. Tegmenand wing with cross-veins brownish, narrowly edged pale brown, marginal veins yellowish. Tegmen veryfaintly and irregularly mottled greyish brown distad of medial-cubital cross-vein; with a small, circular,dark brown spot on subcostal cell, between cubital vein and claval suture at one-sixth length, another overmedial vein at slightly basad of one-third length, a fourth, rather larger and more irregular spot over firstbranch of cubital vein at one-third length. Wing with a small, irregular, brown spot between cubital veinand claval suture at slightly basad of two-fifths length. Shaft of aedeagus not apically expanded ; with a pair of large , flap-like processes extending ventrally overlateral surfaces; a pair of acute, spine-like processes adjacent to midline. Paramere slender; apex acute;dorsal process large, situated at approximately midlength. MATERIAL EXAMINED Holotype cf , Venezuela: San Esteban, Carabobo, 2.ii.l920 (Williamson & Ditzler} (USNM).Paratypes. 3 cf , 2 $, same data as holotype (USNM; BMNH). This species is readily distinguished by the pigmentation of the fronto-lateral surfaces of thepronotum, and by the structure of the male genitalia. Dysimia maculipennissp. n. (Figs 101, 113, 125) Male: head 0-22 mm long, 0.38 mm wide; pronotum 0-68 mm wide; tegmen 3-20 mm long; wing 1.88 mmlong. Female: tegmen 3-80-4-10 mm long. Length of frons 8 times width at apex , twice width at base ; ocelli small , distinct ; clypeus two-thirds lengthof frons. Pronotal width 10 times mid-dorsal length. Genae at level of eyes each with a broad, horizontal brown band, darkest at upper and lower margins,extending from adjacent to eye to anterior margin and continuing across frons; head dorsad of uppermargins of eyes very pale, whitish. Fronto-lateral surfaces of pronotum whitish dorsally, each with a broad,dark brown band extending horizontally from adjacent to eye to external margin. Tegmen and wing withbases of branches and cross-veins pale brown, weakly bordered smoky brown; very narrowly edgedcrimson on posterior and apical margins. Tegmen with a large, circular, dark brown spot on cubital vein atone-third length; two smaller spots at one-sixth length, one on cubital vein, the other exterior and adjacentto fused subcostal, radial and medial veins; an intermediately sized spot over cross-vein linking second andthird branches of medial vein; a small, irregular marking around separation of subcostal and fused radialand medial veins; posterior margin irregularly pale brown; apical margin with semi-circular brown spotsbetween branches of radial and medial veins. Wing with a pale brown spot immediately distad of first forkof cubital vein. Shaft of aedeagus slender in dorsal aspect; a pair of large, flap-like processes, each bearing a pair of long,robust spines dorso-laterally. Paramere slender; apex narrowly rounded; dorsal process situated atmidlength, apex strongly produced posteriorly, dorsal surface at one-quarter length with an irregularlyrounded secondary process bearing scattered, robust spines. 96 PETER S. BROOMFIELD MATERIAL EXAMINED Holotype cf , Ecuador, Tena, 4.iii.l923 (Williams) (BMNH).Paratypes. 1 C?, 2 $ , same data as holotype (BMNH; BPBM). The four specimens in the series bear 'paratype' labels similar to those used by Muir, and theseries is labelled as 'maculipennis Muir'; however, a search of the literature reveals no record ofthis name ever having been published by Muir. The name maculipennis is used here to avoid anypossible confusion. This species is distinguished by the pigmentation of the head and tegmen, and by the structureof the male genitalia. Nomen dubium Dysimia putilla FennahDysimia putilla Fennah, 1952: 124. Holotype <j>, ST LUCIA (BMNH) [examined]. Female: tegmen 4-40 mm long; wing (damaged) 2+ mm long. Head lost. Pronotum with fronto-lateral surfaces distinctly carinate. Tegmen and wing with veins pale brown. Tegmen with a small, circular, brownish spot on medial vein atone- third length; another, slightly larger spot on first fork of cubital vein equidistant from base; anotheradjacent to apical fork of medial vein; smaller and less distinct pale brownish markings at level ofseparation of medial from fused radial and subcostal veins, and at midlength of cubital vein. Wing with asmall, roughly circular, pale brown spot adjacent to and basad of first fork of cubital vein. MATERIAL EXAMINEDHolotype $, St. Lucia: Quilesse, mountain forest, 1000 ft, 22.ii.1941 (Fennah) (BMNH). This species was described from a unique female of which only a few fragments stored in alcoholnow remain, thus making an adequate re-description impossible. Fennah stressed the similarityof the species to maculata, but also the differences in its markings; in his description he stated: Basad portion of frons, excluding the secretory pits, a single narrow stripe on side of head beforeeyes, a minute spot on genae above base of antennae, dark fuscous. A broad area on lateral lobes ofpro no turn, posterior margin of tegulae, and a slight suffusion on mesonotum light sepia-brown tofuscous; abdomen with third segment sublaterally, fourth prominently laterally and with four shortantero-posterior stripes between them, fifth with four paler spots, and sixth with two pale submedianspots and two spots laterally fuscous. and [Tegminal] veins interruptedly and diffuselyoverlain with fuscous. The subgenital sternite is figured, and described as: . . . produced caudad in a subequilaterallytriangular lobe with the apex shortly truncate. DYSIMIELLA gen. n.Type-species: Dysimiella penny isp. n. Width of head one-quarter to two-thirds greater than length in dorsal aspect. Vertex c. 1 -25 times as long aswide; lateral margins strongly and regularly converging from base to level of anterior margins of eyes,thence parallel to junction with frons; extending for c. one-third its length beyond anterior margins of eyes;base very deeply incised medially; lateral carinae very prominent. Length of frons 9 times width at apex, c.twice width at base; junction with vertex broadly and regularly rounded; lateral margins from apex initiallygradually and regularly diverging, then abruptly and strongly diverging to base; carinae very prominent.Genae extending anterior to eyes for c. one-third to one-half horizontal diameter of eye. Second antennalsegment club shaped, c. 1-5 times as long as wide; apex truncate; flagellum arising apically. Ocelliprominent or obsolete. Clypeus short and broad, shorter than frons, as long as basal width; medial carinaobsolete; lateral carinae distinct basally. Rostrum not extending beyond hind coxae. Pronotal width 11-24 times mid-dorsal length, strongly constricted medially; fronto-lateral surfaces eachwith a single carina curving horizontally from adjacent to midline of eye to lateral margin. Tegula withcarina weak or absent. Disc of mesonotum slightly wider than long; medial carina weak; lateral carinaeobsolete or absent. Tegmen short and broad; length 4-00-5-20 mm, little greater than twice width. Cubital vein with fourbranches extending to posterior margin; first and second, and third and fourth linked by cross- veins. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 97 Medial vein fused with radial and subcostal veins over basal one-fifth length; forking at two-fifths andthree-fifths length and apically; with seven branches extending to posterior and apical margins; first forklinked to cubital vein by a short cross- vein subbasally; linked to radial vein by a cross- vein slightly distad ofsecond fork. Radial vein fused to subcostal vein over basal two-fifths length; two branches extending toapical margin; linked to subcostal vein by a cross-vein subapically. Wing short and broad; length two-thirds that of legmen, twice maximum width; apex broadly truncate.Cubital vein with three branches extending to posterior margin. Medial vein unbranched, linked to cubitalby a cross-vein slightly distad of the second fork of the latter. Radial vein unbranched, linked to medial by asingle cross-vein slightly basad of three-quarters length. Male genitalia with shaft of aedeagus horizontal, cylindrical, symmetrical, heavily armed on dorsalsurface subapically. Paramere with dorsal process well developed. Subgenital plate with posterior marginnarrowly and prominently produced at midline. Dysimiella is represented in Brazil and Guyana. The venation of the tegmen and the distinctiveproportions of the head and pronotum indicate a close relationship to Dysimia; the structure ofthe male genitalia, in particular the subgenital plate, is unique. Key to species of Dysimiella 1 Tegmen and wing with distinct dark transverse bands. Aedeagus with three pairs of spines (Fig. 154) williamsi sp. n. (p. 97) - Tegmen and wing without distinct transverse markings. Aedeagus with large flap-like processes and a single pair of long slender spines (Fig. 155) penny! sp. n. (p. 97) Dysimiella williamsi sp. n. (Figs 3, 17,32, 154, 156, 158) Male: head 0-38 mm long, 0-48 mm wide; pronotum 0-90 mm wide; tegmen 4-00-4-20 mm long; wing2-65 mm long. Female: tegmen 4-60-4-80 mm long. Length of frons 9 times width at apex, c. twice width at base; ocelli obsolete; clypeus slightly shorter thanfrons. Pronotal width c. 11 times mid-dorsal length, fronto-lateral carinae distinct; tegula weakly carinate. Genae each with a brown band extending horizontally from adjacent to centre of eye to anterior margin.Fronto-lateral surfaces of pronotum each with a brown band extending horizontally from adjacent tomidline of eye to lateral margin, and continued over lower surface of tegula. Tegmen and wing whitishhyaline, cross- veins edged dark brown. Tegmen with a broad, irregular, pale brownish, transverse band atone-fifth length; apical fork of medial vein covered by a large, very prominent, dark brown spot; costal andposterior margins very narrowly, regularly intermittently, flecked with crimson. Wing with a distinct, verypale brown, transverse band at one-quarter length, posterior margin very narrowly flecked with crimson. Shaft of aedeagus slender, with three pairs of strong, curving spines subapically. Paramere robust; dorsalprocess situated slightly distad of midlength, apex strongly produced postero-dorsally; dorsal surfacewithout a secondary process. Subgenital plate with posterior margin strongly and narrowly producedmedially into a very long, slender, apically acute spine. MATERIAL EXAMINED Holotype cf , Brazil: Para, Jabaty, v.1924 (Williams) (BMNH).Paratypes. Brazil: 11 cf , 17 $, same data as holotype; nr Manaus (BMNH; INPA). This species is readily distinguished by the prominent markings on the tegmen, and by thestructure of the male genitalia. Dysimiella penny i sp. n. (Figs 155, 157, 159, 160) Male: head 0-34 mm long, 0-55 mm wide; pronotum 1-01 mm wide; tegmen 4-62 mm long; wing 3-15 mmlong. Female: tegmen 5-10 mm long. Length of frons 9 times width at apex, 1-66 times width at base; ocelli large, prominent; clypeusthree-fifths as long as frons. Pronotal width 24 times mid-dorsal length, fronto-lateral carinae weak; tegulanot carinate. Head and body pale yellowish brown; lateral carinae of frons, tibiae and tarsi darker brownish. Tegmenand wing hyaline, veins pale. Tegmen with veins and cross-veins narrowly edged pale smoky brown, with a 98 PETER S. BROOMFIELD darker brownish spot over point of separation of fused radial and subcostal veins, another over apical forksof radial and medial veins, apical cells between branches of radial and medial veins each with a slender,longitudinal, brownish stripe medially. Wing unmarked. Female with subgenital plate dark brownish,ventro-lateral angles of adjacent segment brownish black. Shaft of aedeagus slender, broadening slightly at two-thirds length; a large, rounded, flap-like processmedially, bearing on each side posteriorly a very long curving spine; lateral surfaces each produced into anacute, ventrally directed flap. Paramere slender; apex narrowly rounded; dorsal process situated attwo-thirds length, dorsally produced but hardly inclined posteriorly; dorsal surface strongly and obtuselyproduced at midlength, acutely produced subapically. Subgenital plate narrowly produced medially into along, slender, apically somewhat expanded and deeply and narrowly notched, spine-liked process. MATERIAL EXAMINED Holotype cf , Brazil: Amazonas, P. das Laranjeiras, 28.vii.1981 (Arias) (INPA).Paratypes. 2 $, same data as holotype (INPA; BMNH). This species is readily distinguished by the lack of a distinct transverse band on the tegmen, andby the structure of the male genitalia, including the medial process of the subgenital plate. MYSIDALOIDESgen. n. Type-species: Mysidaloides trinidadensis sp. n. Width of head in dorsal aspect greater than 1-5 times length. Vertex hardly extending anterior to eyes;lateral margins weakly carinate, very abruptly converging from base to level of midline of eyes, thengradually and regularly converging to apex; basal margin transverse; junction with frons broadly andregularly rounded. Frons exceeding narrow apically; length greater than 10 times width at apex, less than 3times width at base; lateral margins strongly carinate, parallel from apex to level of ventral margins of eyes,then strongly diverging to base. Genae extending anterior to eyes for one-eighth horizontal diameter ofeye. Eyes extremely large, prominent, almost hemispherical. Antenna cylindrical; second segment morethan 5 times as long as wide; flagellum arising at c. two-thirds length; apex acutely rounded. Ocelli verysmall, not prominent. Rostrum terminating somewhat posterior to hind coxae. Pronotal width more than 20 times mid-doral length; very deeply and regularly constricted medially;fronto-lateral surfaces each with a distinct carina curving horizontally from adjacent to midline of eye tolateral margin. Tegula not carinate. Disc of mesonotum broader than long; not distinctly carinate. Tegmen 3 times as long as wide. Medial vein separating from fused radial and subcostal veins atone-eighth length; radial and subcostal veins separating slightly distad of one-third length. Radial vein withtwo branches extending to apical margin, linked to medial vein by a cross- vein at two-thirds length and atlevel of apical fork. Medial vein forking slightly basad of, and again slightly distad of, midlength; with sevenbranches extending to apical and posterior margins, second and third, and fourth and fifth linked bycross-veins. Cubital vein with four branches extending to posterior margin, first linked to apex of clavusand to second, second to third, third to fourth, and fourth to first branch of medial vein by cross-veins. Length of wing slightly greater than one-half of that of tegmen. Radial and subcostal veins fused over c.basal one-third length. Radial vein unbranched, linked to medial vein by a cross-vein at two-thirds length.Medial vein distinct throughout, with two branches extending to apical margin, linked to third branch ofcubital vein by a cross-vein slightly distad of midlength. Cubital vein with three branches extending toposterior margin. Head and body uniformly pale, lacking distinct markings. Tegmen and wing hyaline, also lacking distinctmarkings. Male genitalia with shaft of aedeagus horizontal, cylindrical, symmetrical, slender; dorsal surfacesubapically bearing well-developed flap-like and spine-like processes. Paramere with dorsal process welldeveloped, without a distinct secondary process; ventral surface unarmed. Anal tube little producedposteriorly. Female with posterior margin of subgenital plate not produced posteriorly. The venation of the tegmen and wing of this genus is similar to that of Mysidia, but it differs inhead characters, in particular the large size of the eyes, the very slender apical portion of thefrons and the extreme length of the antennae. The genus is monotypic and is known from Brazil,Guyana and Trinidad. 99Mysidaloides trinidadensissp. n. (Figs 10, 16,24, 130, 131,132) Male: head 0-38 mm long, 0-65 mm wide; pronotum 1-32 mm wide; tegmen 6-30 mm long; wing 3-40 mmlong. Female: tegmen 7-40 mm long. Length of frons 12 times width at apex, 2-33 times width at base; ocelli indistinct; clypeus as long as frons.Pronotal width 22 times mid-dorsal length, fronto-lateral carinae distinct; tegula without carinae. Head and body unmarked. Tegmen and wing hyaline, veins pale yellow. Tegmen unmarked except for avery indistinct, pale smoky, transverse band extending from second branch of cubital vein to apex ofclavus. Wing with posterior margin between branches of cubital vein and apex of radial vein pale smokybrown, otherwise unmarked. Shaft of aedeagus somewhat expanded subapically; dorsal surface subapically with a pair of large,flap-like processes extending to midlength, each terminating in an antero-dorsally directed projection atmidline; a pair of diverging spines at midlength. Paramere basally slender, apex broadly rounded; dorsalprocess situated slightly distad of midlength, not greatly produced posteriorly; dorsal surface at approx-imately one-third length with a low, rounded projection bearing a cluster of robust, internally directedspines. MATERIAL EXAMINED Holotype cT, Guyana: Tumatumari, 19.vii.1923 (Williams) (BMNH). Paratypes. Guyana: 1 cf , same data as holotype (BMNH). Trinidad: 1 cf , Aripo Valley (BMNH).Brazil: 9 cf , Amazonas, Manaus (INPA; BMNH). NEOMYSIDIA gen. n. Type-species: Neomysidia willisisp. n. Width of head considerably greater than length in dorsal aspect. Vertex slightly longer than wide at base;extending less than one-quarter its length beyond anterior margins of eyes; lateral carinae low, verygradually converging from base to apex; basal margin very weakly concave; junction with frons broadlyand regularly rounded. Frons parallel-sided from apex to immediately above base, thence lateral marginsstrongly divergent; c. 2-5 times as long as width at apex, little longer than width at base. Genae extendingbeyond anterior margins of eyes for one-third horizontal diameter of eye. Second antennal segment ovate,c. twice as long as broad; apex rounded; flagellum arising subapically. Ocelli large and distinct. Clypeusbroad, rounded, longer than frons; length one and two-thirds width at base; lacking distinct medial andlateral carinae. Rostrum terminating at level of mid coxae. Pronotal width slightly greater than 7 times length mid-dorsally; not strongly constricted medially;fronto-lateral surfaces each with a distinct carina curving horizontally from adjacent to eye to lateralmargin. Tegula not carinate. Disc of mesonotum c. 1-5 times as wide as long; broadly triangular; medialcarina distinct only at midlength; lateral carinae absent or obsolete. Tegmen length c. 6 mm, 2-5 times maximum breadth. Medial vein distinct from fused radial andsubcostal veins throughout; forking slightly basad and slightly distad of midlength, with seven branchesextending to posterior and apical margins. Radial and subcostal veins separating at midlength; radial withtwo branches extending to apical margin; linked to subcostal vein by a cross-vein subapically, and to medialvein at c. two-fifths length. Cubital vein with four branches extending to posterior margin, the first andsecond, and third and fourth linked by cross-veins (Fig. 6). Wing almost twice as long as wide, more than half length of tegmen; apex rounded. Subcostal and radialveins fused from base to slightly basad of midlength, unbranched; radial vein linked to medial by across-vein slightly distad of midlength. Medial vein with two branches extending to post-apical margin;linked to cubital vein by a cross-vein at c. midlength. Cubital vein two branched. Male genitalia with shaft of aedeagus horizontal, symmetrical; dorsal surface heavily armed subapically.Paramere with dorsal process reduced; dorsal surface with a secondary process subbasally. This genus is readily distinguished by the proportions of the head; especially the very short andbroad frons, the relatively unconstricted dorsal surface of the pronotum, and the short,triangular, mesonotal disc.Known only from Brazil. 100 PETER S. BROOMFIELD Neomysidia willisisp. n. (Figs 6, 18,31,161,165, 169) Male: head 0-42 mm long, 0-71 mm wide; pronotum 1-58 mm wide; legmen 6-20 mm long; wing 3-50 mmlong. Female unknown. Length of frons 2-5 times width at apex, one-quarter greater than width at base; ocelli large, distinct;clypeus one-third longer than frons. Pronotal width 7-5 times mid-dorsal length. Vertex, genae ventral to eyes, antennae, base of frons, lateral surfaces of clypeus, a small spot onfronto-lateral surfaces of pronotum above each eye, and dorsal margins of tegula dark brown. Tegmen andwing hyaline, veins pale yellowish, cross- veins very narrowly dark brown, posterior and apical marginswith dark brown spots over apices of veins. Tegmen with radial, medial and cubital areas basally irregularlymottled dark smoky brown ; an irregular dark brown spot immediately distad of apex of clavus ; a very dark ,almost black spot over apical fork of medial vein. Wing with a short, oblique, irregular, transverse smokybrown band over medial-cubital cross- vein; a broad, indistinct, clique, pale smoky brown, transverse bandat somewhat distad of three-quarters length. Shaft of aedeagus robust; a pair of massive flap-like processes situated one on either side of midline, eachterminating anteriorly in a small acute point. Paramere robust; apex acutely rounded; dorsal processsituated at midlength, greatly reduced; present only as a slight projection; dorsal surface subbasally with acurving posteriorly directed, hook-like secondary process; ventral and lateral surfaces subbasally with verynumerous, tiny, tooth-like projections. MATERIAL EXAMINEDHolotype cT, Brazil: Amazonas, P. das Laranjeiras, 28.vii.1981 (Arias) (INPA). IPSEMYSIDIA gen. n. Type-species: Ipsemysidia beautifica sp. n. Width of head in dorsal aspect greater than 1-5 times length. Vertex little longer than width at base,extending less than one-third its length beyond anterior margins of eyes; lateral margins not highlyelevated, very gradually and regularly converging from base; basal margin shallowly concave; junctionwith frons weakly angulate. Frons c. 3 times as long as width at apex, c. twice width at base; lateral marginsweakly diverging from apex to level of midline of eyes, then very strongly diverging to base. Genaeextending anterior to eyes for c. half horizontal diameter of eye. Second antennal segment c. 1-5 times aslong as broad, rounded; flagellum arising subapically. Ocelli distinct, not prominent. Clypeus short,broadly rounded, little longer than frons; medial and lateral carinae obsolete or absent. Rostrum notextending beyond hind coxae. Pronotal width less than 10 times mid-dorsal length; dorsal surface little constricted medially; fronto-lateral surfaces each with a somewhat sinuate carina extending horizontally from adjacent to midline of eyeto lateral margin. Tegula not carinate. Disc of mesonotum considerably wider than long; medial carinaeweak, lateral carinae absent. Tegmen more than 6 mm long, c. 3 times maximum width. Fused radial and subcostal veins separating atc. midlength. Medial vein distinct from base, forking at c. midlength and two-thirds length, with sevenbranches extending to posterior and apical margins, first branch linked to cubital vein, second to third bycross-veins. Radial vein with two branches extending to apical margin, linked to medial by a cross- veinslightly distad of two-thirds length. Cubital vein with four branches extending to posterior margin, first andsecond, and third and fourth linked by cross-veins. Length of wing more than half that of tegmen, c. twice maximum width; apex acutely rounded. Subcostaland radial veins fused from base to c. midlength, unbranched; radial linked to medial vein by a cross-veinslightly distad of two-thirds length. Medial vein with two branches extending to post-apical margin, linkedto cubital by a cross-vein at level of second fork of the latter. Cubital vein with three branches extending toposterior margin. Male genitalia with shaft of aedeagus symmetrical, horizontal; dorsal surface heavily armed subapically,including a single process at midline. Paramere with dorsal process reduced; a hook-like secondary processon dorsal surface subbasally. Though resembling Neomysidia in the development of the paramere, Ipsemysidia is readilydistinguished by external and aedeagal characters. It is recorded from Brazil and Panama. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 101 Ipsemysidia beautifies sp. n.(Figs 8, 21, 26, 162, 166, 170) Male: head 0-42 mm long, 0-65 mm wide; pronotum 1-50 mm wide; tegmen 6-00 mm long; wing 3-60 mmlong. Female: tegmen 7-20 mm long. Length of frons c. 3 times width at apex, c. twice width at base; ocelli distinct; clypeus c. as long as frons;rostrum terminating at level of mid-coxae. Pronotal width 7 times mid-dorsal length, fronto-lateral carinaevery prominent; tegula not carinate. Head unmarked. Fronto-lateral surfaces of pronotum basally reddish; abdomen with a small dark brownspot on either side of mid-dorsal line . Tegmen and wing whitish hyaline , veins yellow , cross-veins and forksof veins dark brown; posterior margin irregularly dark brown. Tegmen with branches of cubital vein andtwo basal branches of medial vein each with at least one small black tubercule at c. midlength; medial veinwith apical forks blackish brown; branches of apical vein each with a small, circular, dark brown spotsubapically; posterior branch of radial vein similarly marked; apical cells of radial and medial veins eachwith an irregular brownish spot medially at two-thirds length; area between radial vein and costal marginirregularly mottled brownish, these markings extending posteriorly over first forks of medial and cubitalveins; clavus with a brown spot adjacent to point of fusion of anal veins, another smaller spot subapically.Wing with radial and medial branches each bearing a small black tubercule; medial vein basad ofmedial-cubital cross-vein with two to four similar tubercules; cubital vein with two tubercules between firstand second forks, third branch occasionally with a single tubercule subbasally; a large, irregular, brownishspot between first branch of cubital vein and claval suture somewhat distad of base of former; an irregular,oblique, smoky brown band over radial and medial areas at five-sixths length. Shaft of aedeagus bearing a pair of large, apically rounded, flap-like processes laterally, and a single,slender, curving, spine-like process medially; ventral surface with a pair of long, closely opposed, flap-likeprocesses subapically. Paramere slender basally, broadly expanded towards regularly rounded apex;dorsal process situated at approximately three-quarters length, very weakly produced postero-dorsally;dorsal surface at one-quarter length with a large, posteriorly directed, hook-like secondary process; ventralsurface subbasally with a group of long, robust spines. MATERIAL EXAMINED Holotype C?, Brazil: Rondonia, Porto Velio, 22.ii.1979 (Campbell} (INPA).Paratypes. Brazil: 1 $, same data as holotype (BMNH). Panama: 1 d", Tocumen (USNM). AMYSIDIELLA gen. n. Type-species: Amysidiella micare sp. n. Width of head in dorsal aspect half to three-quarters greater than length. Vertex extending for less thanone-quarter its length beyond anterior margins of eyes; lateral margins elevated, gradually convergingfrom base; basal margin transverse, very weakly concave; junction with frons broadly and regularlyrounded. Length of frons c. 5 times width at apex, less than twice width at base; lateral margins graduallydiverging from apex to level of ocelli, then strongly diverging to base. Genae extending anterior to eyes forless than one-third horizontal diameter of eye. Second antennal segment club-shaped, not longer thantwice maximum width; flagellum arising subapically. Ocelli distinct, not prominent. Clypeus, short, broad,rounded; length c. equal to that of frons, 1-5 times width at base; lacking medial and lateral carinae.Rostrum not extending beyond hind-coxae. Pronotal width 11-14 times mid-dorsal length; fronto-lateral surfaces each with a weak carina extendinghorizontally from adjacent to eye to lateral margin. Tegula not carinate. Disc of mesonotum c. as wide aslong; medial carina weak or obsolete; lateral carinae absent. Length of tegmen 5-90-7-20 mm, slightly less than 3 times maximum width. Medial vein separating fromfused radial and subcostal veins at c. one-sixth length; with seven branches extending to post-apical margin.Radial and subcostal veins fused over c. basal one-third length; radial with two branches extending toapical margin, linked to medial by cross-veins at slightly distad of second fork of the latter, and to subcostalsubapically. Cubital vein with four branches extending to posterior margin; first and second, and third andfourth linked by cross-veins. Length of wing slightly greater than half that of tegmen; apex acutely rounded. Cubital vein with threebranches extending to posterior margin, linked by a cross-vein to medial at level of second fork. Medialvein two-branched; linked to unbranched radial vein by a cross- vein at c. two-thirds length. Head and body yellowish. Frons and genae at level of eyes reddish. Fronto-lateral surfaces of pronotumand ventro-lateral surfaces of mesonotum each with a distinct horizontal reddish band. Tegmen and wing 102 PETER S. BROOMFIELD faintly whitish hyaline, veins pale, lacking prominent pigmentation. Tegmen with a brownish spot adjacentto costal margin subbasally; another darker irregular marking extending anteriorly from claval margin at c.one-third length. Wing either unmarked, or with a very faint, irregular transverse band. Male genitalia with shaft of aedeagus horizontal, symmetrical, subapically expanded, robust in lateralaspect; dorsal and lateral surfaces with prominent spine-like processes; ventral surfaces unarmed.Paramere with dorsal process reduced to a small, posteriorly directed, hook-like projection subapically.Posterior margin of subgenital plate transverse. Anal tube strongly produced posteriorly, postero-lateralangles expanded. Female with posterior margin of subgenital plate regularly produced, apically truncate. Amysidiella is distinguished by a combination of the proportions of the head, pronotum, legmenand wing, and by the structure of the male genitalia, consisting of a very heavily armed aedeaguswith a paramere in which the dorsal process is almost obsolete. Recorded from Brazil andGuyana. Key to species of Amysidiella 1 Pronotum with maximum width in excess of 13 times length at mid-dorsal line. Aedeagus with lateral spines short, arising basad of mid-length (Fig. 164) micare sp. n. (p. 102) Pronotum with maximum width less than 12 times length at mid-dorsal line. Aedeagus withlateral spines very long, arising distad of mid-length (Fig. 163) pseudonucaresp. n. (p. 102) Amysidiella micare sp. n. (Figs 4, 22, 25, 164, 167, 171) Male: head 0-40 mm long, 0-69 mm wide; pronotum 1-18 mm wide; tegmen 5-95-6-12 mm long; wing3-40 mm long. Female: tegmen 7-14 mm long. Length of frons 5-5 times width at apex, 1-5 times width at base; ocelli small, distinct; clypeus as long asfrons. Pronotal width 14 times mid-dorsal length. Frons with a scarlet spot at level of eyes, often extending onto adjacent surfaces of genae. Pronto-lateralsurfaces of pronotum each with a narrow scarlet band extending horizontally from level of ventral marginof eye to lateral margin; male with apices of posterior lobes of anal tube dark brown/black. Tegmen with apale brownish spot between costal margin and point of separation of radial and subcostal veins; anotherlarger, irregular marking at approximately one-third length extending from costal margin over first fork ofcubital vein; an irregular band extending from apex of clavus to second branch of cubital vein. Wing with avery weak transverse band at level of medial-cubital cross-vein. Shaft of aedeagus broadly expanded over apical one-half length; lateral surfaces subapically each with alarge, flap-like process terminating apically in a curving spine; dorsal surface at three-quarters length with apair of small, curving, spines at midline, and laterally at rather over midlength, with a pair of long, slender,curving, spine-like processes. Paramere slender; dorsal process situated at three-fifths length, greatlyreduced, consisting of a single, short, posteriorly directed, hooked spine; dorsal surface subbasally with aflap-like process bearing numerous long, robust spines. MATERIAL EXAMINED Holotype C?, Brazil: Belem, Para, v;1924 (Williams) (BMNH).Paratypes. Brazil: 2 cT, 1 9, same data as holotype (BMNH). Guyana: 4 cf , Blairmont (BMNH). This species is readily distinguished by its pigmentation, especially the dark posterior lobes ofthe anal tube in the male, and by the reduced armature of the paramere. Amysidiella pseudomicaresp. n. (Figs 163, 168, 172) Male: head 0-44 mm long, head 0-67 mm wide; pronotum 1-10 mm wide; tegmen 6-00 mm long; wing3-35 mm long. Female: tegmen 6-60 mm long. Length of frons c. 5 times width at apex, one and two-thirds width at base; ocelli large, not prominent;clypeus as long as frons. Pronotal width 11 times mid-dorsal length. Frons pale crimson at level of eyes; fronto-lateral surfaces of pronotum each with a narrow, pale crimsonband extending horizontally from adjacent to midline of eye to lateral margin; mesonotum with a similarband dorsad of bases of mid coxae. Tegmen with a small, distinct, brownish spot on costal margin at level of TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 103 point of separation of medial and fused radial-subcostal veins; a distinct, irregular, transverse, brownishband extending from costal margin over radial and medial veins at one-third length and terminating at baseof third branch of cubital vein; an irregular, pale brownish spot extending from first-second cubitalcross-vein to claval margin. Wing usually unmarked; occasionally with an indistinct, pale brownish,transverse band at midlength. Shaft of aedeagus robust, strongly laterally expanded over apical one-third length; dorsal surfacesubapically with a pair of short, spine-like processes, a pair of long, slender, curving, processes; lateralsurfaces subapically each with a large, dorsally directed, flap-like process, and a very long, slender,spine-like, horizontal process. Paramere very slender; apex acutely rounded; dorsal process situatedslightly basad of three-quarters length, reduced to a small, posteriorly directed hook with a low, roundedprojection distally; dorsal surface subapically with a large, rounded, secondary process bearing numerouslong, robust, spines. MATERIAL EXAMINED Holotype d", Brazil: Rio Uapes, 5.iii (Roman) (NR).Paratype. 1 $, same data as holotype (BMNH). This species, though closely related to micare, may be distinguished by the greater lateralexpansion and the length, position and inclination of the spines of the aedeagus. PARAM YSIDIA gen. n. Type-species: Mysidia mississippiensis Dozier, by present designation. Head in dorsal aspect as wide as or up to one-third wider than long. Vertex extending beyond anteriormargins of eyes for from one-third to one-half its length; lateral margins gradually converging from base tolevel of anterior margins of eyes, thence subparallel to apex; lateral carinae very prominent, almostfoliaceous apically; basal margin shallowly concave; junction with frons broadly and regularly rounded.Frons c. 4-6 times as long as wide at apex, c. twice width at base; lateral margins subparallel from apex tolevel of ocelli, then gradually diverging to base, or very gradually and regularly diverging from apex andabruptly curving outwards subbasally. Genae extending beyond eyes for two-fifths to one-half horizontaldiameter of eye. Antenna short; second segment club-shaped, c. twice as long as broad; apex weaklystepped; flagellum arising subapically. Ocelli commonly large, distinct; rarely prominent. Clypeus short,broadly rounded; length c. equal to that of frons, twice width at base; medial carina generally obsolete orabsent; lateral carinae obsolete, or only distinct subbasally. Rostrum usually terminating immediatelyposterior to hind coxae, rarely extending to midlength of abdomen. Pronotal width from 10-13 times mid-dorsal length; fronto-lateral surfaces each with a prominent carinacurving horizontally from adjacent to eye to lateral margin. Tegula not carinate, or with carina obsolete.Disc of mesonotum slightly wider than long; medial carina often weak or obsolete, rarely distinct andpercurrent; lateral carinae usually absent, rarely distinct. Tegmen commonly from 5 -00-6 -50 mm long, rarely exceeding 7-00 mm long. Medial vein distinct fromnear base, with seven branches extending to posterior and apical margins, second branch linked to third,and fourth to fifth by cross-veins. Fused radial and subcostal veins separating at c. midlength; radial withtwo branches extending to apical margin, linked to medial by a single cross-vein at two-thirds length.Cubital vein with four branches extending to posterior margin; first and second, and third and fourth linkedby cross-veins. Length of wing from three-fifths to three-quarters that of tegmen. Subcostal and radial veins fused overbasal one-third length, unbranched. Radial vein linked to second fork of medial vein by a single cross-veinat two-thirds length. Medial vein with two branches extending to apical margin, linked to cubital vein by across-vein at two-thirds length. Cubital vein with three branches. Head and body predominantly yellowish brown, often very pale; genae adjacent to ocelli often tingedreddish; pronotum with fronto-lateral surfaces often deep yellow over carinae; dorsal surface of abdomenrarely red or dark brown. Tegmen and wing whitish hyaline; veins yellowish or dark brown; cross- veins andforks of veins usually dark brown; veins and cross-veins often edged smoky brown, giving a mottledappearance. Tegmen rarely with an irregular, smoky brown, transverse band. Wing commonly with twoweak, very irregular, smoky brown, transverse bands; posterior and apical margins often broadly palebrown. Male genitalia with shaft of aedeagus slender, horizontal, cylindrical, somewhat laterally expandedsubapically; dorsal surface subapically with a pair of very asymmetrical, dorsally directed, longitudinallyaligned processes, of which that on the right is flap-like, that on the left commonly slender and spine-like;dorsal surface at c. midlength with a single, usually slender, dorsally directed, apically bifid secondary 104 PETER S. BROOMFIELD apodeme medially; lateral margins often each with an antero-laterally directed apically bifid process at c.midlength; ventral surface unarmed. Paramere commonly with apex broadly rounded; dorsal processsituated subapically, greatly reduced; dorsal surface subbasally with a small hook-like secondary process;rarely with apex acute or dorsal process robust. Anal tube not greatly extended, apex commonly notchedmedially.Female with posterior margin of subgenital plate frequently produced medially. Paramysidia differs from Mysidia as follows: the armature of the aedeagus is strongly asymmet-rical and includes a prominent process at midlength on the mid-dorsal line; the development ofthe dorsal process of the paramere is usually strongly reduced, its function possibly being, atleast partially, taken over by the hook-like secondary process. Distributed from the U.S.A. (Mississippi, Florida, Texas, Louisiana) to Costa Rica, Hon-duras, El Salvador, Panama, Brazil and Peru. From this distribution it would appear that speciesof the genus probably reached the U.S. A. via Central America, though it has not been recordedfrom Mexico. This shows a marked contrast with Dysimia, which almost certainly found its wayinto North America by island-hopping across the Caribbean. Key to species of Paramysidia (based on external characters) Due to the extreme external similarity of many species, reference should be made, where possible, to the structure of the male genitalia. 1 Tegmen with cross- veins and forks of veins not distinctly margined smoky brown. South- easternU.S.A mississippiensis (Dozier)(p. 105) Tegmen with cross-veins and forks of veins broadly margined smoky brown 2 2(1) Tegmen with veins yellowish or pale brown 3 Tegmen with veins dark brown 4 3(2) Clypeus with length less than that of f rons ; genae often tinged orange ; f ronto-lateral surfaces of pronotum with carinae orange. Peru vulgaris sp. n. (p. 106) Clypeus with length equal to that of frons; head and body unmarked. Brazil .... felix sp. n. (p. 106)4(2) Clypeus with length not greater than that of frons; frons with length less than 5 times width at apex 5 Clypeus with length one-third greater than that of frons; frons with length almost 6 times width at apex. Honduras, Costa Rica, El Salvador barbara sp. n. (p. 106) 5(4) Width of pronotum 11 times length at mid-dorsal line 6 Width of pronotum greater than 13 times length at mid-dorsal line. Panama boudica sp. n. (p. 107) 6(5) Rostrum extending to mid-length of abdomen. Costa Rica tessellatasp. n.(p. 107) Rostrum terminating immediately posterior to hind coxae. Costa Rica, Panama nigropunctata (Metcalf) (p. 105) Key to species of Paramysidia (based on male genitalia) 1 Paramere with dorsal process strongly reduced 2 Paramere with dorsal process very robust (Fig. 147) vulgaris sp. n. (p. 106) 2(1) Paramere with apex broadly rounded 3 Paramere with apex acutely rounded (Fig. 148) felix sp. n. (p. 106) 3(2) Aedeagus with lateral processes 4 Aedeagus lacking lateral processes (Fig. 135) nigropunctata (Metcalf) (p. 105) 4(3) Aedeagus with right dorsal process slender , distinctly longer than broad 5 Aedeagus with right dorsal process massive, as long as broad (Fig. 143) mississippiensis (Dozier) (p. 105) 5(4) Aedeagus with right dorsal process distinctly longer than left dorsal process 6 Aedeagus with right dorsal process very short, shorter than left dorsal process (Fig. 144) boudica sp. n. (p. 107)6(5) Paramere with secondary dorsal process apically acute (Fig. 152); aedeagus with right dorsal process apically acute (Fig. 145) tessellata sp. n. (p. 107) Paramere with secondary dorsal process apically serrate (Fig. 153); aedeagus with rightdorsal process apically rounded (Fig. 146) barbara sp. n. (p. 106) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 105 Paramysidia mississippiensis (Dozier) comb. n. (Figs 20, 27, 136, 143, 150)Mysidia mississippiensis Dozier, 1922: 82. Holotype $, U.S.A. (USNM) [examined]. Male: head 0-52 mm long; 0-69 mm wide; pronotum 1-56 mm wide; legmen 6-00-7-50 mm long; wing4-42 mm long. Female: tegmen 6-80-7-30 mm long. Length of frons 4 times width at apex, twice width at base; ocelli large, not prominent; clypeus as long asfrons. Pronotal width 11 times mid-dorsal length; tegula weakly carinate. Genae adjacent to ocelli often with an orange or dull reddish circular spot; fronto-lateral surfaces ofpronotum with carinae broadly deep yellow. Tegmen and wings with veins yellowish brown; cross- veinsand branches of veins dark brown. Tegmen with cross-veins each surrounded by a roughly circular smokybrown spot; posterior margin narrowly smoky brown; basal three-eighths of length, and costal and radialcells, mottled smoky brown; area between first branch of cubital vein and apex of clavus broadly smokybrown. Wing with area distad of radial-medial cross-vein irregularly mottled smoky brown. Shaft of aedeagus basally slender, broadly laterally expanded over apical half length; dorsal surface atthree-quarters length with a pair of large processes, that on the right of midline massive, hooked, anteriorlydirected; lateral processes present, apically bifid; dorsal apodeme situated at three-fifths length, slender,weakly curving. Paramere slender at base, becoming greatly expanded towards obtusely rounded apex;dorsal process situated at rather over two-fifths length, slender, apex posteriorly produced. MATERIAL EXAMINED Holotype $, U.S.A.: Mississippi, Leland, 15.ix.1921 (Drake) (USNM). U.S.A.: 4 cf, 4 $, Mississippi (FAMU; BMNH); 6 cf, 4 $, Louisiana (FAMU; BMNH); 2 cf, Texas(FAMU; USNM); 2 cf , Florida (USNM). This species is most readily distinguished by the pigmentation of the tegmen and wing, and bythe structure of the male genitalia. Paramysidia nigropunctata (Metcalf) comb. n. (Figs 135, 142, 149)Mysidia nigropunctata Metcalf, 1938: 316. Holotype $, PANAMA (MCZ) [examined]. Male: head 0-52 mm long, 0-61 mm wide; pronotum 1-40 mm wide; tegmen 5-60-6-30 mm long; wing3-55 mm long. Female: tegmen 6-60 mm long. Length of frons 4 times width at apex, 2-5 times width at base; ocelli large, prominent; clypeus c. as longas frons. Pronotal width 11 times mid-dorsal length. Genae around ocelli occasionally pale orange; abdomen with dorsal surface usually dark brown.Tegmen and wing with veins and cross-veins dark brown, broadly and irregularly edged brownish. Wingwith an obscure, irregular, smoky brown, transverse band over radial-medial cross-vein; another muchnarrower band over first fork of cubital vein; posterior and apical margins irregularly edged smoky brown. Shaft of aedeagus somewhat laterally expanded at two-thirds length; dorsal surface with a broadhook-like process on the right side, and a slender spine-like process on the left; ventro-lateral surfacesunarmed; mid-dorsal process slender, curving, situated at midlength. Paramere robust; apex broadlyrounded; dorsal process simple, situated at three-quarters length. MATERIAL EXAMINED Holotype $, Panama: C.Z., Barro Colorado, 13.vii.1924 (Banks) (MCZ). Panama: 7 cf , 6 $ (including 2 cf , 1 $ paratypes) (BMNH; FAMU; CAS; MCZ). Costa Rica: 7 cf , 5 $(BMNH; FAMU; USNM). Nicaragua: 1 $ (USNM). In his description Metcalf refers to the holotype as female; however, a male specimen bears theMCZ 'type' label, while the female is labelled as 'allotype'. In external characters this species resembles boudica, but it is readily distinguished by thestructure of the male genitalia. 106 PETER S. BROOMFIELD Paramysidia vulgarissp. n. (Figs 2, 133,140, 147) Male: head 0-46 mm long, 0-59 mm wide; pronotum 1-20 mm wide; tegmen 5-00-6-00 mm long; wing3-00 mm long. Female: tegmen 6-00-7-25 mm long. Length of frons c. 5 times width at apex, c. twice width at base; ocelli large, distinct; clypeus slightlyshorter than frons. Pronotal width c. 12 times mid-dorsal length. Genae often orange around ocelli; second antennal segment rarely black apically; fronto-lateral carinaenarrowly orange. Tegmen with radial and medial veins pale, other veins and cross-veins brown; cross-veinsand forks of veins broadly edged smoky brown to give an irregular mottled appearance; without distincttransverse bands; apical branches of medial vein dark brown at midlength. Wing with veins alternately paleand dark; with a broad, smoky brown, transverse band at midlength, another slightly distad of three-quarters length. Shaft of aedeagus slender basally, broader from midlength; dorsal surface subapically with a large,apically acute, flap-like process on the right side and a long, slender, acute, spine-like process on the left;lateral surfaces each with a weakly bifurcate process; mid-dorsal process situated at midlength, slender,curving antero-dorsally. Paramere robust; apex obtusely rounded; dorsal process large, situated somewhatdistad of midlength, not posteriorly produced. MATERIAL EXAMINED Holotype C?, Brazil: Para, Jabaty, v.1924 (Williams) (BMNH). Paratypes. Brazil: 8 cT, 8 9 , same data as holotype; Mato Grosso; Bahia Iguassu (BMNH; NR). Peru: 63Cf , 64 $ , Ivitas, 60 km W. of Pucallpa (BMNH; FAMU). The structure of the paramere of this species is unique and readily separates it from all others inthe genus. Paramysidia felixsp. n. (Figs 134, 141,148) Male: head 0-46 mm long, 0-61 mm wide; pronotum 1-30 mm wide; tegmen 5-30 mm long; wing 3-06 mmlong. Female unknown. Length of frons c. 6 times width at apex, c. twice width at base; ocelli large, not prominent; clypeus aslong as frons. Pronotal width c. 10 times mid-dorsal length. Head and body unmarked. Tegmen and wing with veins and cross-veins pale yellow, broadly andirregularly edged pale brown. Tegmen with markings coalescing to form a broad, indistinct, transverseband over first fork of cubital vein; narrower, very broken, bands at midlength and three-quarters length.Wing with a narrow, pale brownish, transverse band slightly basad of midlength; another broader bandover radial-medial cross-vein; posterior and apical margins very faintly and irregularly tinged smokybrown. Shaft of aedeagus slender, not laterally expanded; dorsal surface with a pair of large flap-like processes,each terminating anteriorly in a long slender spine; lateral surfaces at three-fifths length each with a broadapically bifid process; mid-dorsal process situated at three-fifths length, very broad in lateral aspect,curving, with a large acute spine on posterior surface at midlength. Paramere slender; apex narrowlyrounded; dorsal process robust, situated somewhat distad of midlength, posteriorly produced. MATERIAL EXAMINED Holotype cf , Brazil: Jabaty, Para, v.1924 (Williams) (BMNH).Paratype. 1 cf , same data as holotype (BMNH). This species is distinguished by its very pale pigmentation, and by the structure of the malegenitalia. It is regarded as having the least specialized genitalia in the genus; the shaft of theaedeagus is almost symmetrical, but the presence of the mid-dorsal process confirms itsplacement; the paramere bears a subbasal, hook-like, secondary process. Paramysidia barbara sp. n. (Figs 139, 146, 153) Male: head 0-42 mm long, 0-42 mm wide; pronotum 1-03 mm wide; tegmen 5-60-6-40 mm long; wing3-15 mm long. Female: tegmen 6-40-6-90 mm long. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 107 Length of frons c. 6 times width at apex, c. twice width at base; ocelli large, not prominent; clypeusone-third longer than frons. Pronotal width 12 times mid-dorsal length. Head and body unmarked. Tegmen and wing hyaline; veins and cross-veins dark brown, narrowly andirregularly edged smoky brown. Tegmen with subcostal, radial, medial and cubital areas irregularlymottled smoky brown; posterior margin narrowly smoky brown. Wing with an indistinct, irregular, smokybrown, transverse band immediately distad of radial-medial cross-vein. Shaft of aedeagus with a large, slightly hooked, flap-like process to the right of mid-dorsal line, and along, slender, slightly curving, spine-like process to the left; mid-dorsal process situated at two-thirdslength, curving, strongly bifurcate apically; lateral surfaces each with a short apically shallowly bifurcateprocess. Paramere robust; apex very obtusely rounded; dorsal process situated subapically, obsolete. MATERIAL EXAMINED Holotype d" , Honduras: Santa Barbara, Chumbagua, 26.vii.1966 (Malta) (FAMU). Paratypes. Honduras: 7 d", 7 $ , same data as holotype (FAMU; BMNH). Costa Rica: 1 $ , Villa Neilly(FAMU). El Salvador: 2 C?, 4 $, San Miguel; San Pedro Perulapan (FAMU; BMNH). This species is distinguished by the very narrow head, and by the structure of the male genitalia. Paramysidia boudica sp. n. (Figs 137, 144, 151) Male: head 0-48 mm long, 0-67 mm wide; pronotum 1-40 mm wide; tegmen 6-40 mm long; wing 3-60 mmlong. Female unknown. Length of frons 4-5 times width at apex, 2-5 times width at base; ocelli large, distinct; clypeus slightlyshorter than frons. Pronotal width c. 13 times mid-dorsal length. Genae around ocelli and anterior to bases of antennae tinged pale orange. Pronto-lateral surfaces ofpronotum with carinae narrowly edged pale orange. Tegmen and wing with veins and cross-veins darkbrown, broadly and irregularly edged smoky brown. Tegmen with base narrowly smoky brown. Wing withan irregular, broad, smoky brown band extending transversely from costal margin to apex of clavus at levelof first fork of cubital vein; another, very broad band at level of radial-medial cross-vein; posterior andapical margins broadly smoky brown. Shaft of aedeagus slender; dorsal process to right of midline broad, tapering, hook-like; ventro-lateralsurfaces each with an apically bifid, spine-like process at midlength; dorsal apodeme situated slightlydorsad of midlength, sinuate, very slender. Paramere very robust; apex very obtusely rounded; dorsalprocess situated subapically, strongly reduced; secondary dorsal process situated at approximatelyone-third length, small, hook-like. MATERIAL EXAMINEDHolotype C?, Panama: 1924 (Cheeseman) (BMNH). This species is readily distinguished by the markings of the tegmen and wing, and by thestructure of the male genitalia. Paramysidia tessellata sp. n. (Figs 138, 145, 152) Male: head 0-46 mm long, 0-57 mm wide; pronotum 1-36 mm wide; tegmen 6-00-6-80 mm long; wing3-65 mm long. Female unknown. Length of frons 4 times width at apex, c. twice width at base; ocelli large, distinct; clypeus as long asfrons. Pronotal width 11 times mid-dorsal length. Genae often tinged deep yellow around ocelli. Pronto-lateral surfaces of pronotum with carinae broadlydeep yellow; abdomen with dorsal surface usually tinged red or brown. Tegmen and wing with veins andcross-veins dark brown, broadly and very irregularly edged smoky brown. Wing with a very faint smokyhyaline transverse band at one-third length, a much more distinct smoky brown band at level of first fork ofcubital vein, a third band at level of radial-medial cross-vein; posterior and apical margins irregularly edgedsmoky brown. Shaft of aedeagus broad, strongly laterally expanded from midlength to apex; dorsal surface atthree-quarters length with right-hand process very robust and hook-like; lateral surfaces each with anapically bifurcate, spine-like process at midlength; dorsal apodeme situated slightly basad of midlength,slender, slightly curving. Paramere robust; apex broadly expanded; dorsal process simple, situated atthree-quarters length. 108 PETER S. BROOMFIELD MATERIAL EXAMINED Holotype cf , Costa Rica: Guan, 3 km NW. Liberia, 500 ft, 12.vii.1974 (O'Brien & Marshall) (FAMU).Paratypes. 8 cf , data as holotype (FAMU; BMNH). This species is distinguished by the three transverse bands on the wing, the pigmentation of theabdomen, and by the structure of the male genitalia. SYMIDIA MuirSymidia Muir, 1918: 234. Type-species: Symidiaflava Muir, by monotypy. Head in dorsal aspect distinctly wider than long. Vertex triangular, extending for one-quarter to one-thirdits length beyond anterior margins of eyes, lateral carinae very prominent; posterior margin very broadlyand deeply incised; junction with frons commonly broadly rounded, rarely acutely angled. Frons extremelynarrow, length 18-20 times width at apex, 2-3 times width at base; c. parallel-sided from apex toimmediately above base, then very abruptly laterally expanded. Genae extending anterior to eyes forone-third to two-fifths horizontal diameter of eye. Second antennal segment club-shaped, c. as long aswide; apex truncate, flagellum arising apically. Ocelli very small, usually distinct. Clypeus c. as long asfrons; medial carina usually distinct over greater part of length from base; lateral carinae usuallypercurrent. Rostrum extending at least to base of subgenital plate. Pronotal width 8-16 times mid-dorsal length; fronto-lateral surfaces each with a very highly elevated,foliaceous carina curving horizontally from adjacent to midline of eye to lateral margin, continuingventrally and internally along lateral and ventral margins to genae at level of base of antenna (Fennah,1952, refers to this apparent encirclement of the antennal base as an 'antennal fovea'). Tegula not carinate.Disc of mesonotum slightly broader than long; medial and lateral carinae usually distinct; rarely obscure,extending from anterior margin to midlength or almost to hind margin. Tegmen 4 -90-6 -00 mm long. Subcostal and radial veins fused from base over c. one-third length. Radialvein with three branches extending to apical margin. Medial vein separating from fused radial andsubcostal veins at one-sixth length; with six branches extending to apical and posterior margins. Cubitalvein with two branches extending to posterior margin, linked to medial vein by a cross-vein at c. one-thirdlength (Fig. 5). Wing not more than c. half length of tegmen, often considerably shorter. Radial, medial and cubitalveins distinct throughout. Radial and medial veins unbranched, linked by a cross-vein at slightly distad oftwo-thirds length. Cubital vein two-branched, linked to medial vein by a cross-vein at two-thirds length. Head and body predominantly pale brownish yellow; abdomen occasionally darker. Genae often withred or orange markings dorsal or ventral to eyes. Tegmen and wing usually pale whitish hyaline, veins pale;often with pale brownish markings which may coalesce to form irregular and indistinct transverse bands. Male genitalia with shaft of aedeagus horizontal, cylindrical, variably asymmetrical; dorsal surfacesubapically with three or four pairs of large, anteriorly directed processes; ventral surface usually unarmed.Paramere with dorsal process situated subapically, usually rounded, never greatly produced, interlockingsurfaces situated basally, often reduced; ventral surface subbasally with numerous, very small, obtusespines, or ridged. Anal tube moderately produced posteriorly, somewhat laterally expanded, apicallynotched or bifurcate. Subgenital plate with posterior margin transverse, or with a small triangularprojection medially. Female with posterior margin of subgenital plate medially produced, occasionally greatly so. Symidia is regarded as being a highly developed off-shoot of the Mysidiini because of thereduction of the tegminal and wing venation, the great expansion of the fronto-lateral carinae ofthe pronotum, the asymmetry of the aedeagus, and the simple form of the dorsal process of theparamere, while still retaining the apical position of the antennal flagellum. Of the species available for study, flava is regarded as the most primitive because of its veryextensive range and relatively unspecialized aedeagal development. The genus is recorded from Trinidad, Guyana, Brazil, Ecuador and Peru. Key to species of Symidia (based on external characters) 1 Junction of vertex and frons acutely angled in lateral aspect flava Muir (p. 109) Junction of vertex and frons broadly and regularly rounded in lateral aspect 2(1) Tegmen with oblique transverse bands dark and prominent pintosamia sp. n.(p. 109) Tegmen with oblique transverse bands indistinct 3 TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 109 3(2) Genae ventral to eyes scarlet. Abdomen often dark brown pseudoffava sp. n. (p. 110) Genae ventral to eyes occasionally brownish yellow, never distinctly reddish; occasionally pale orange at level of midline of eyes. Abdomen pale 4 4(3) Wing with distinct, brownish, transverse bands at two-fifths and two-thirds length. Genaeextending anterior to eyes for two-thirds horizontal diameter of eye. Female with posteriormargin of subgenital plate produced medially beyond apex of abdomen, apex obtusely rounded. Ecuador bucaya sp. n. (p. Ill) Wing with a very faint, brown, transverse band at one-third length, another much moredistinct band at three-quarters length. Genae extending anterior to eyes for only two-fifthshorizontal diameter of eye. Female with posterior margin of subgenital plate bearing asmall triangular spine medially, not greatly produced posteriorly. Brazil withycombei sp. n. (p. 110) Key to species of Symidia (based on male genitalia) It has not been possible to examine a male ofpintosamia which is therefore omitted from this key. 1 Shaf t of aedeagus with ventral surf ace unarmed (Fig. 86) pseudoffavasp. n.(p. 110) Shaft of aedeagus with ventral surface bearing spine-like processes subapically 2 2(1) Paramere with apex obtusely rounded, dorsal process somewhat inclined posteriorly (Fig. 91) bucaya sp. n.(p. Ill) Paramere with apex acutely rounded, dorsal process not posteriorly inclined 3 3(2) Shaft of aedeagus with ventral surface bearing a single, anteriorly directed process, situated to right of midline subapically (Fig. 88) ttava Muir (p. 109) Shaft of aedeagus with ventral surface bearing three posteriorly directed spine-like processes(Fig. 89) withycombei sp. n. (p. 110) Symidia tiava Muir (Figs 5, 19, 28, 84, 88, 92)Symidia flava Muir, 1918: 234. LECTOTYPE d", GUYANA (BMNH), here designated [examined]. Male: head 0-38 mm long, 0-53 mm wide; pronotum 1-05 mm wide; tegmen 5-00-5-30 mm long; wing2-23 mm long. Female: tegmen 5-10-5-50 mm long. Junction of vertex and frons acutely angled; length of frons 19 times width at apex, c. 3 times width atbase; ocelli very small, obscure; clypeus c. as long as frons; rostrum extending to apex of abdomen.Pronotal width 10 times mid-dorsal length. Genae each with an orange or red band extending horizontally from adjacent to dorsal margin of eye tojunction of vertex and frons. Tegmen and wing whitish hyaline, veins pale yellow. Tegmen with pale,smoky brown markings coalescing to form irregular and intermittent transverse bands at one-third, mid-and three-quarters length. Wing with very faint, smoky brownish bands at two-fifths and two-thirds length;apex pale smoky brown. Shaft of aedeagus broad; dorsal surface subapically with three pairs of processes, the second pairflap-like and strongly hooked apically; ventral surface subapically with a single, slender, flap-like process.Paramere slender, apex narrowly rounded; dorsal process situated slightly basad of three-quarters length,low and rounded; ventral surface subbasally with numerous small tooth-like spines. MATERIAL EXAMINED Holotype cf , Guyana: Demerara R., 20.iii.1913 (Muir) (BMNH). Guyana: 21 cT, 19 9 (including 4 d", 5 9 paratypes) (BMNH). Brazil: 11 C?, 14 $ (BMNH). Trinidad: 9Cf , 16 9 (BMNH). Ecuador: 7 cf , 4 9 (BMNH). This species is readily distinguished by the acutely angled junction of the vertex and frons, andthe adjacent markings on the genae. Symidia pintosamia sp. n. Female: head 0-44 mm long, 0-55 mm wide; pronotum 1-20 mm wide; tegmen 5-60-6-00 mm long; wing2-40 mm long. Male unknown. Junction of vertex and frons broadly rounded; length of frons 20 times width at apex, 3 times width at 110 PETER S. BROOMFIELD base; ocelli very small, distinct; clypeus as long as frons; rostrum extending well beyond apex of abdomen.Pronotal width 11-5 times mid-dorsal length. Genae ventral to eyes orange; ocelli reddish. Tegmen and wing almost hyaline, veins yellow. Tegmenwith a broad, oblique, brown, transverse band at level of separation of fused subcostal and radial veins,another at level of radial-medial cross-vein; apex broadly smoky brown. Wing with a broad, smoky brown,transverse band at four-fifths length. MATERIAL EXAMINED Holotype <j>, Peru: eastern foothills of Andes, 1 km S. Tingo Maria, 2000 ft, 16.viii.1971 (Broomfield)(BMNH). Paratype. 1 $ , same data as holotype (BMNH). This species is distinguished by the prominent dark markings on the tegmen, and by the extremelength of the rostrum. Symidiapseudoflavasp. n. (Figs 82, 86, 90) Male: head 0-40 mm long, 0-50 mm wide; pronotum 1-11 mm wide; tegmen 5-10-5-30 mm long; wing2-55 mm long. Female: tegmen 5-35-5-95 mm long. Junction of vertex and frons broadly rounded; length of frons c. 16 times width at apex, 2-33 times widthat base; ocelli small, distinct; clypeus as long as frons; rostrum extending to apex of abdomen. Pronotalwidth 13 times mid-dorsal length. Genae ventral to eyes scarlet; abdomen often dark brown. Tegmen and wing whitish hyaline, veins verypale. Tegmen with a broad, pale brown, transverse band immediately distad of medial-cubital cross-vein;an ill-defined, oblique, pale brown, transverse band slightly distad of midlength; apical quarter lengthirregularly mottled pale brownish. Wing with a distinct, pale brown, transverse band at four-fifths length. Shaft of aedeagus slender; dorsal surface subapically with four pairs of large processes, anterior pair verylong and broad, apices hooked; ventral surface unarmed. Paramere with apex very obtusely rounded;dorsal process situated at two-thirds length, large, rounded, with a dorsally aligned, heavily spined ridge oninternal surface; ventral surface subbasally with numerous small, tooth-like spines. MATERIAL EXAMINED Holotype cf , Ecuador: Tena, 29.iii.1923 (Williams) (BMNH).Paratypes. Ecuador: 3 cT, 5 $ , Tena (BMNH). This species is distinguished by the scarlet markings on the genae, the pigmentation of thetegmen and wing, and by the structure of the male genitalia. Symidia withy combeisp. n. (Figs 85, 89, 93) Male: head 0-38 mm long, 0-53 mm wide; pronotum 1-18 mm wide; tegmen 5-10 mm long; wing 2-50 mmlong. Female: tegmen 5-30-5-70 mm long. Junction of vertex and frons obtusely rounded; length of frons 18 times width at apex; 3 times width atbase; ocelli very small, distinct; clypeus slightly longer than frons; rostrum extending to base of subgenitalplate. Pronotal width 14 times mid-dorsal length. Genae and fronto-lateral surfaces of pronotum pale, ocelli red, genae at level of eyes rarely pale orange.Tegmen and wing whitish hyaline, veins pale yellow. Tegmen broadly and irregularly mottled brownisharound cross-veins and forks of veins, these markings coalescing at one-third and two-thirds length to formoblique, intermittent transverse bands. Wing with a distinct, broad, brown, transverse band at three-quarters length, a much fainter band at one-third length not extending to posterior margin. Shaft of aedeagus slender; dorsal surface subapically with four pairs of processes, fourth pair large,flap-like, apically hooked; single, spine-like process partially obscured by the paired processes; ventralsurface subapically with a pair of spine-like processes; at some distance basad of apex, a single spine-likeprocess on left side. Paramere slender, apex acute; dorsal process small, situated at three-quarters length,apex truncate and tuberculose. MATERIAL EXAMINED Holotype cf , Brazil: Rezende, Estado de Rio, ii.1924 (Williams) (BMNH).Paratypes. 1 cf , 6 $ , same data as holotype (BMNH). TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 111 This species is distinguished by the lack of reddish pigmentation ventral to the eye, the roundedjunction of the vertex and frons, and by the structure of the male genitalia. Symidia bucaya sp. n. (Figs 83, 87, 91) Male: head 0-33 mm long, 0-50 mm wide; pronotum 1-05 mm wide; tegmen 4-90-5-10 mm long; wing2-12 mm long. Female: tegmen 5-40-5-78 mm long. Junction of vertex and frons obtusely rounded; length of frons 19 times width at apex, 3 times width atbase; ocelli very small, distinct; clypeus c. as long as frons; rostrum terminating level with apex ofabdomen. Pronotal width 16 times mid-dorsal length. Genae ventral to eyes, and clypeus brownish yellow; disc of mesonotum and abdomen brownish, thelatter occasionally tinged reddish. Tegmen and wing almost hyaline, veins and cross-veins pale yellow.Tegmen with irregular brownish mottlings coalescing to form intermittent, oblique, transverse bands atlevel of second fork of medial vein and at two-thirds length. Wing with a broad, pale brown, transverseband at two-fifths length, another at two-thirds length. Shaft of aedeagus subapically bearing four pairs of processes on dorsal surface, fourth pair large,flap-like, apically bifurcate; ventral surface, on the right side only, subapically with a short spine-likeprocess. Paramere with apex obtusely rounded; dorsal process situated at three-quarters length, large,rounded; ventral surface subbasally with numerous small, tooth-like spines. MATERIAL EXAMINED Holotype cf , Ecuador: Bucay, 1000 ft, 7.X.1922 (Williams} (BMNH).Paratypes. 6 cf, 9 9 , same data as holotype (BMNH). This species is distinguished by the absence of reddish pigmentation on the genae, the markingsof the tegmen and wing, and by the structure of the male genitalia. References Ball, E. D. 1928. Some new genera and species of N. A. Derbidae with notes on others (Fulgoridae). Canadian Entomologist 60: 196-201.Dozier, H. L. 1922. A synopsis of the genus Stenocranus , and a new species of Mysidia. Ohio Journal of Science 22: 69-82.1931. New and interesting West Indian Hompotera. American Museum Novitates, New York. no. 510: 1-24. Fabricius, J. C. 1803. Systema Rhyngotorum x + 314pp. Brunsvigae.Fennah, R. G. 1945. The Fulgoroidea, or Lanternflies, of Trinidad and adjacent parts of South America. Proceedings of the United States National Museum 95: 411-520.1952. On the generic classification of Derbidae (Fulgoroidea), with descriptions of new Neotropical species. Transactions of the Royal Entomological Society of London 103: 109-170. 1971. Fulgoroidea from the Cayman Islands and adjacent areas. Journal of Natural History 5: 299-342. Fowler, W. W. 1900. Biologia Centrali-Americana. Rhynchota 1: 1-147.Germar, E. F. 1830. Thon's Entomologisches Archives 2: 56. Jena.Kirkaldy, G. W. 1900. Bibliographical and nomenclatorial notes on the Rhynchota (I). Entomologist 33: 242. 1906. Leaf hoppers and their natural enemies (IX. Leaf Hoppers, Hemiptera). Bulletin of the Hawaiian Sugar Planters' Association Experimental Station 1: 287-479. Kramer, S. 1950. The morphology and phytogeny ofAuchenorrhynchous Homoptera (Insecta). vii + 111 pp. Urbana.Metcalf, Z. P. 1938. The Fulgorina of Barro Colorado and other parts of Panama. Bulletin of the Museum of Comparative Zoology, Harvard 82: 277-423. 1945a. Fulgoroidea (Homoptera) of Kartabo, Bartica District, British Guiana. Zoologica 30: 125-143. 1945ft. General catalogue of the Homoptera. Fulgoroidea 4 (4): 1-212. Muir, F. E. S. 1913. On some new species of leaf-hoppers (II). Bulletin of the Hawaiian Sugar Planters' Association Experimental Station 3: 28-91.1917. The Derbidae of the Philippine Islands. Philippine Journal of Science 12: 49-104. 112 PETER S. BROOMFIELD 1918a. Notes on the Derbidae in the British museum Collection, II, Derbinae. Entomologists Monthly Magazine 54: 228-243. 1918ft. Homopterous notes (II). Proceedings of the Hawaiian Entomological Society 3: 414-429. 1924. New and little known fulgorids from the West Indies (Homoptera). Proceedings of the Hawaiian Entomological Society 5: 461-472.Schaum, H. R. 1850. Allgemeine Encyklopddie Ersch und GruberSl: 64-69.Spinola, M. M. 1839. Sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des Rhyngotes. Annales de la Societe Entomologique de France 8: 133-454.Westwood, J. 0. 1840. Observations on the genus Derbe of Fabricius. Proceedings of the Linnean Society of London 1: 82-85.1842a. Descriptions of some new species of exotic homopterous insects, Transactions of the Linnean Society of London 19: 1-18. 18426. Descriptions of several new homopterous insects belonging to various subgenera of Derbe of Fabricius. Transactions of the Linnean Society of London 19: 19-22. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 113 Figs 1-9 The genera of the Derbinae, tegmen and wing: 1 , Derbe westwoodi; 2, Paramysidia vulgaris; 3,Dysimiella williamsi; 4, Amysidiella micare; 5, Symidia flava; 6, Neomysidia willisi; 7, Mysidiaacidaloides; 8, Ipsemysidia beautifica; 9, Dysimia maculata. Derbe, Paramysidia, Mysidia, and Ipsemysi-dia are shown to one-half scale. Sc = subcostal area; R = radial area; M = medial area; Cu = cubitalarea; A = anal area. 114 PETER S. BROOMFIELD 22 Figs 10-22 The genera of the Derbinae. 10,11, legmen and wing of (10) Mysidaloides trinidadensis; (11)Pseudomysidia fuscovaria. 12-22, head, pronotum and mesonotum in dorsal aspect of (12) Derbewestwoodi; (13) Dysimia maculata; (14) Pseudomysidia pallida; (15) Mysidia acidaloides; (16) Mysida-loides trinidadensis; (17) Dysimiella williamsi; (18) Neomysidia willisi; (19) Symidia flava; (20) Para-mysidia mississippiensis; (21) Ipsemysidia beautifica; (22) Amysidiella micare. Derbe and Mysidia areshown to one-half scale. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 115 28 32 34 36 Figs 23-38 The genera of the Derbinae. 23-33, head in frontal aspect of (23) Derbe westwoodi; (24)Mysidaloides trinidadensis; (25) Amysidiella micare; (26) Ipsemysidia beautifica; (27) Paramysidiamississippiensis; (28) Symidia flava (including fronto-lateral surfaces of pronotum); (29) Dysimiamaculata; (30) Pseudomysidia pallida; (31) Neomysidia willisi; (32) Dysimiella williamsi; (33) Mysidiaacidaloides. Derbe and Mysidia are shown to one-half scale. Pseudomysidia species. 34-38, dorsal viewof aedeagus of (34) palmeri; (35) rubidella; (36) Juliana; (37) debora; (38) similis. 116 PETER S. BROOMFIELD Figs 39-53 Pseudomysidia species. 39-49, dorsal view of aedeagus of (39) hindore; (40) panamensis; (41)fuscovaria; (42) pallida; (43) araguana; (44) marshalli; (45) vestis; (46) trinidadensis; (47) ecuadoriensis;(48) delicata; (49) obnubilia. 50-53, left lateral view of aedeagus of (50) palmeri; (51) rubidella; (52)Juliana; (53) debora. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 117 Figs 54-65 Left lateral view of aedeagus of Pseudomysidia species. 54, similis; 55, hindore; 56,panamensis; 57, fuscovaria; 58, pallida; 59, araguana; 60, marshalli; 61, vestis; 62, trinidadensis; 63,ecuadoriensis; 64, delicata; 65, obnubilia. 118 PETER S. BROOMFIELD 77 Figs 66-78 Left lateral view of paramere of Pseudomysidia species . 66 , palmeri ; 67 , rubidella ; 68 , Juliana ;69, debora; 70, similis; 71, hindore; 72, panamensis; 73, fuscovaria; 74, pallida; 75, araguana; 76,marshalli; 77, ve^^is; 78, trinidadensis . TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 119 91 Figs 79-91 Pseudomysidia species. 79-81, left lateral view of paramere of (79) ecuadoriensis; (80)delicata; (81) obnubilia. 82-91. Symidia species. 82-85, dorsal view of aedeagus of (82) pseudoflava; (83)bucaya; (84) flava; (85) withycombei. 86-89, left lateral view of aedeagus of (86) pseudoflava; (87)bucaya; (88) flava; (89) withycombei. 90, 91, left lateral view of paramere of (90) pseudoflava; (91)bucaya. 120 PETER S. BROOMFIELD Figs 92-107 92,93, Symidia species , left lateral view of paramere of (92)flava ; (93) withy combei. 94-107 ,Dysimia species. 94-105, dorsal view of aedeagus of (94) distincta; (95) numa; (96) maculata; (97)pseudomaculata; (98) muiri; (99) fennahi; (100) telfordi; (101) maculipennis; (102) obrieni; (103)jamaicensis; (104) morrisi; (105) astarte. 106. 107, left lateral view of aedeagus of (106) distincta; (107) numa. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 121 Figs 108-119 Dysimia species. 108-117, left lateral view of aedeagus of (108) maculata; (lW))pseudoma-culata- (110) muiri; (111) fennahi; (112) telfordi; (113) maculipennis; (114) obrieni; (115) jamaicensis;(116) morrisi; (117) astarte. 118, 119, left lateral view of paramere of (118) distincta; (119) numa. 122 PETER S. BROOMFIELD Figs 120-132 120-129, Dysimia species, left lateral view of paramere of (120) maculata; (121) pseudoma-culata; (122) muiri; (123) fennahi; (124) telfordi; (125) maculipennis; (126) obrieni; (127) jamaicensis;(128) morrisi; (129) astarte. 130-132, Mysidaloides trinidadensis. (130) dorsal view of aedeagus; (131)left lateral view of aedeagus; (132) left lateral view of paramere. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 123 Figs 133-147 Paramysidia species. 133-139, dorsal view of aedeagus of (133) vulgaris; (134) felix; (135)nigropunctata; (136) mississippiensis; (137) boudica; (138) tessellata; (139) barbara. 140-146, left lateralview of aedeagus: (140) vulgaris; (141) felix; (142) nigropunctata; (143) mississippiensis; (144) boudica;(145) tessellata; (146) barbara. 147, left lateral view of paramere of vulgaris. 124 PETER S. BROOMFIELD 158 Figs 148-160 148-153, Paramysidia species, left lateral view of paramere of (148) /e/u; (149) nigropunc-tata; (150) mississippiensis; (151) boudica; (152) tessellata; (153) barbara. 154-160, Dysimiella species.154, 155, dorsal view of aedeagus of (154) williamsi; (155)pennyi. 156, 157, left lateral view of aedeagusof (156) williamsi; (I51)pennyi. 158, 159, left lateral view of paramere of (158) williamsi; (I59)pennyi.160, ventral view of male subgenital plate ofpennyi. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 125 Figs 161-172 161-164, dorsal view of aedeagus of (161) Neomysidia willisi; (162) Ipsemysidia beautifica;(163) Amysidiella pseudomicare; (164) A. micare. 165-168, left lateral view of aedeagus of (165) N.willisi; (166) /. beautifica; (167) A. micare; (168) A. pseudomicare. 169-172, left lateral view of paramereof (169) N. willisi; (170) /. beautifica; (171) A. micare; (172) A. pseudomicare. 126 PETER S. BROOMFIELD 186 Figs 173-189 Mysidia species, dorsal view of aedeagus. 173, fowleri; 174, distanti; 175, claudata; 176,pulchella; lll,fuscodorsalis; 178, douglasi; 179, bizzara; 180, carosella; l&I,jamesi; lS2,peregrina; 183,venusta; 184, augusta; 185, ariasi; 186, amazona; 187, molesta; 188, glauca; 189, cinerea. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 127 205 Figs 190-205 Mysidia species, dorsal view of aedeagus. 190, agilis; 191, ecuadoria; 192, decora; 193,lacteola; 194, cheesemani; 195, nemorensis; 196, poly hymnia; 197, limpida; 198, nitida; 199, amarantha;200, stall; 201, erecta; 202, pseudoerecta; 203, knighti; 2Q4,persephone; 205, minerva. 128 PETER S. BROOMFIELD Figs 206-223 Mysidia species, dorsal view of aedeagus. 206, harmonia; 207, bianco; 208, pseudocostata;209, bibula; 210, panamensis; 211, isteria; 212, estfarchina; 213, insolita; 214, athena; 215, sanguinea;216, acidaloides; 217, adusta; 218, havilandi; 219, richardsi; 220, costata; 22l,adamare; 222, simpla; 223, TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 129 227 235 238 239 Figs 224-242 Mysidia species, dorsal view of aedeagus. 224, liquida; 225, fasciata; 226, hyalina; 227,diana; 228, albipennis; 229, bella; 230, neonebulosa; 231, nigrithorax; 232, lucifera; 233, infedelis; 234,transversa; 235, intima; 236, caliginosa; 237, calypso; 238, williamsi\ 239, albifasciata; 24QJosianna; 241,nigrifrontalis; 242, c/ava. 130 PETER S. BROOMFIELD 259 Figs 243-259 Mysidia species , dorsal view of aedeagus . 243 , inquinata ; 244 , gracilis ; 245 , fuscofrontalis ;246,formosa; 247, cooperi; 248, punctum; 249, magica; 250, boliviano; 251,silvana; 252, hengist; 253,musica; 254, etheldreda; 255, dollingi; 256, diabola; 257, neoasinella; 258, andes; 259, tikalme. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDHNI (HOMOPTERA) 131 265 269 270 276 Figs 26ft-277 Mysidia species, dorsal view of aedeagus. 260, obscura; 261, varia; 262, lucianna; 263,fuscomaculata; 264, lloydi; 265, insania; 266, nebulosa; 267, perspicua; 26S,flavilla; 269, unimaculata;270, maculosa; 21\,distincta\ 272, whimperi; 273, testacea; 274,geoffreyi; 215,delicatissima; 276, my/es/. 132 PETER S. BROOMFIELD Figs 277-289 Mysidia species. 277-281, dorsal view of aedeagusof (277) henrietta; (278) pseudonebulosa;(279) albicans; (280) krameri; (281) enjebetta. 282-289, left lateral view of aedeagus of (282) fowleri;(283) distantly (284) daudata; (285) pulcella; (286) fuscodorsalis; (287) douglasi;(288) bizzara; (289) TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 133 Figs 290-303 Mysidia species, left lateral view of aedeagus. 290, carosella; 291, jamesi; 292, peregrina;293, venusta; 294, augusta; 295, ariasi; 296, amazona; 297, molesta; 298, glauca; 299, cinerea; 300, agilis;3Ql,ecuadoria; 302, decora; 303, lacteola. 134 PETER S. BROOMFIELD Figs 304-316 Mysidia species, left lateral view of aedeagus. 304, cheesemani; 305, nemorensis; 306,polyhymnia; 307, llmplda; 308, nitida; 309, amarantha; 310, erecta; 311 , pseudoerecta; 312, stall; 313,knighti; 3l4,persephone; 315, minerva; 316, harmonia. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 135 Figs 317-331 Mysidia species, left lateral view of aedeagus. 317, bibula; 318, bianco; 319, pseudocostata;320, panamensis; 321, isteria; 322, estfarchina; 323, insolita; 324, athena; 325, sanguinea; 326, acida-loides; 327, krameri; 328, adttfta; 329, havilandi; 330, costata; 331, richardsi. 136 PETER S. BROOMFIELD Figs 332-345 Mysidia species, left lateral view of aedeagus. 332, adamare; 333, simpla; 334, marshalli;335, liquida; 336,fasciata; 337, hyalina; 338, diana; 339, albipennis; 340, bella; 341, neonebulosa; 342,nigrithorax; 343, lucifera; 344, infedelis; 345, transversa. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 137 346 Figs 346-357 Mysidia species, left lateral view of aedeagus. 346, calypso; 347, intima\ 348, caliginosa;349, williamsi; 350, albifasciata; 351, josianna; 352, nigrifrontalis; 353, clava; 354, inquinata; 355,gracilis; 356,fuscofrontalis; 351,formosa. 138 PETER S. BROOMFIELD Figs 358-370 Mysidia species, left lateral view of aedeagus. 358, cooperi; 359,punctum; 360, magica; 361,boliviano; 362,-silvana; 363, hengist; 364, musica; 365, etheldreda; 366, dollingi; 367, diabola; 368,neoasinella; 369, andes; 370, tikalme. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 139 Figs 371-383 Mysidia species, left lateral view of aedeagus. 371, obscura; 372, varia; 373, lucianna; 374,fuscomaculata; 375, lloydi; 376, insania; 377, nebulosa; 378, perspicua; 379 , flavilla; 380, unimaculata;381, maculosa; 382, distincta; 383, whimperi. 140 PETER S. BROOMFIELD Figs 384-395 Mysidia species. 384-390, left lateral view of aedeagus of (384) testacea; (385) geoffreyi;(386) delicatissima; (387) mylesi; (388) Henrietta; (389) pseudonebulosa; (390) albicans. 391-395, leftlateral view of paramere of (39l)fowleri; (392) distanti; (393) daudata; (394) pulchella; (395)fulvodorsa- TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDHNI (HOMOPTERA) 141 Figs 396-406 Mysidia species , left lateral view of par amere . 396 , douglasi ;397,bizzara;398, jamesi ; 399 ,enjebetta; 400, carosella; 401, peregrina; 402, venusta; 403, augusta; 404, ariasi; 405, amazona; 406,molesta. 142 PETER S. BROOMFIELD Figs 407-417 Mysidia species, left lateral view of paramere. 407, glauca; 408, cinerea; 409, agilis; 410,ecuadoria; 411, decora; 412, lacteola; 413, cheesemani; 414, nemorensis; 4l5,polyhymnia; 416, limpida;417, nitida. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDHNI (HOMOPTERA) 143 Figs 418-427 Mysidia species, left lateral view of paramere. 418, amarantha; 419, erecta; 420,pseudoerec-ta; 421, s tali; 422, knighti; 423,persephone; 424, minerva; 425, harmonia; 426, bianco; 427 , pseudocosta-ta. 144 PETER S. BROOMFIELD Figs 428-439 Mysidia species, left lateral view of paramere. 428, bibula; 429, panamensis; 430, isteria;431, estfar china; 432, insolita; 433, athena; 434, sanguinea; 435, acidaloides; 436, krameri; 437, adusta;438, havilandi; 439, richardsi. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 145 Figs 440-451 Mysidia species, left lateral view of paramere. 440, costata; 441 , adamare; 442, simpla; 443,liquida; 444, marshalli; 445, hyalina; 446, fasciata; 447, diana; 448, albipennis; 449, bella; 450,neonebulosa; 451, nigrithorax. 146 PETER S. BROOMFIELD Figs 452-463 Mysidia species. 452-462, left lateral view of paramere of (452), lucifera; (453), infedelis;(454), transversa; (455), intima; (456), caliginosa; (457), calypso; (458), williamsi; (459), albifasciata;(46Q),josianna; (461), nigrifrontalis; (462), clava. 463, lateral view of posterior projection of pregenitaltergite of male of caliginosa. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 147 Figs 464-473 Mysidia species, left lateral view of paramere. 464, inquinata; 465, gracilis; 466,fuscofron-talis; 467 , formosa; 468, cooperi; 469 , punctum; 470, magica; 471, boliviano; 472, silvana; 473, musica. 148 PETER S. BROOMFIELD Figs 474-483 Mysidia species, left lateral view of paramere. 474, hengist\ 475, etheldreda; 476, dollingi;477, neoasinella; 478, diabola; 479, andes; 480, tikalme; 481, lucianna, 482, varia; 483, obscura. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) 149 Figs 484-^494 Mysidia species , left lateral view of paramere . 484 , fuscomaculata ; 485 , lloydi ; 486 , insania ;487, nebulosa; 488, perspicua; 489, flavilla; 490, unimaculata; 491, maculosa; 492, distincta; 493,whimperi; 494, testacea. 150 PETER S. BROOMFIELD Figs 495-501 Mysidia species, left lateral view of paramere. 495, geoffreyi; 496, delicatissima;'497 , mylesi;498, Henrietta; 499,pseudonebulosa; 500, albicans; 501, robusta. TAXONOMY OF NEOTROPICAL DERBIDAE IN THE NEW TRIBE MYSIDIINI (HOMOPTERA) Index Synonyms are in italics; page references to descriptions are in bold. 151 acidaloides 29adamare 60adusta 22agilis 69albicans 50albifasciata 31albipennis 38amarantha 26, 72, 74amazona 26Amysidiella 4, 101andes 52araguana 81ariasi 37asinella 22astarte 95athena 28, 30augusta 54 barbara 106beautifica 101bella 71bianca 42bibula 53bizzara 56boliviana 72boudica 105, 107bucaya 111 caliginosa 23, 24calypso 44carosella 70Cenchrea 2Cenchreinae 2Cenchreini 2cheesemani 25, 54cinerea 76citrina 57claudata 69clava 62cooped 42costata 40, 41 debora 85decora 72delicata 86delicatissima 41Derbe 2Derbidae 1Derbinae 2, 3, 4Derbini 4diabola 58diana 43distant! 49, 51distincta (Dysimia) 93distincta (Mysidia) 66dollingi 43douglasi 65Dysimia 3, 5, 87, 104Dysimiella 3,5,% ecuadoria 53ecuadoriensis 85 elatior 77enjebetta 51erecta 55, 68estfarchina 37, 48, 64etheldreda 36 fasciata 65felix 106fennahi 94flava 108, 109flavilla 29formosa 61fowled 46fulvodorsalis 32fuscoclypeata 90fuscofrontalis 38fuscomaculata 59fuscovaria 77, 80, 81 geoffreyi 75glauca 35gracilis 35grandis 46 harmonia 70havilandi 36hengist 67Henrietta 49Heronax 77hindore 83hyalina 28 immaculata 27infedelis 57inquinata 23, 24insania 60insolita 50, 51intima 56, 73Ipsemysidia 4, 5, 100isteria 59 jamaicensis 91jamesi 70josianna 59, 67Juliana 81 Kermesia 2Kermesiinae 2knighti 66krameri 54 lacteola 39lactiflora 39lepida 86limpida 25, 72liquida, 37lloydi, 55, 58lucianna 45lucifera 35 maculata 87, 89 maculicosta 48maculipennis 95maculosa 55magica 74 marshalli (Mysidia) 73marshalli (Pseudomysidia) 83micare 101, 102, 102minerva 47 mississippiensis 103, 105molesta 37, 48morrisi 93muiri 92musica 32mylesi 33 Mysidaloides 3, 5, 98Mysidia 2, 4, 9, 104Mysidiini 1, 4, 5 nebulosa 56nemorensis 51neoasinella 30Neocenchrea 77Neomysidia 3, 99, 100neonebulosa 74nigrifrontalis 52nigrithorax 63nigropunctata 105nitida 26, 72numa 90 obnubilia 86, 87obrieni 92, 93obscura 49Otiocerinae 2 pallescens 60, 64 pallida (Mysidia) 9, 76 pallida (Pseudomysidia) 84 palmeri 82 panamensis (Mysidia) 59, 61 panamcnsis (Pseudomysidia) 84 Paramysidia 3, 5, 103 parviceps 38 Patara 2 pennyi 96, 97 peregrina 45 persephone 73 perspicua 68 pintosamia 109 Phenice 2 polyhymnia 23, 24 pseudocostata 41 pseudoerecta 68 pseudoflava 110 pseudomaculata 90, 91 pseudomicare 102 Pseudomysidia 2, 4, 77 pseudonebulosa 75 punctifera 47 pulchella 66 punctum 27 152putilla % quadri fascia 31, 32 Rhotaninae 2richardsi 25robusta 26rubidella 80rubra, 23, 24 sanguinea 44Sikaiana 2Sikaianini 2, 4silvana 71, 73simil is 83simpla 63, 68spreta 77squamigera 40, 41 PETER S. BROOMFIELD stall 62steinbachi 27stigma 77striata 43subfasciata 64subfusca 64Symidia 2, 5, 108 telfordi 94 tessellata 107 testacea 57 Thracia 2 tikalme 34 transversa 32 trinidadensis (Mysidaloides) 98, 99 trinidadensis (Pseudomysidia) 82 uni maculata 28 varia 34venusta 24vestis 84vista 30vulgaris 106 whimperi 62williamsi (Dysimiella) 97williamsi (Mysidia) 33willisi 99, 100withycombei 110 Zeugma 2Zoraida 2Zoraidinae 2, 4Zoraidini 2, 4 British Museum (Natural History) Milkweed butterflies: their cladistics and biology P. R. Ackery & R. I. Vane- Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects oftheir biology have stimulated far more attention than can be justified by species numbersalone. In recent years, an expansive literature has grown, considering aspects of theircourtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry andgenetics. The popularity of danaines among biologists can certainly be attributed to thiscombination, within one small group, of so many of the factors that make butterflies such aninteresting group to study. The obvious need to place this wealth of biological data within anacceptable systematic framework provided the impetus for this volume. Started eight years ago within the conventions of evolution by natural selection andHennig's phylogenetic systematics, the book is now largely about natural history (what theanimals have and do, where they live and how they develop) and natural groups - as revealedby a form of analysis approaching that practised by the new school of 'transformed cladistics'.The authors have prepared a handbook that will appeal to a wide range of biologists, frommuseum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography.ISBN 565 00893 5. 1984. Price 50. Titles to be published in Volume 50 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). By Peter S. Broomfield Nymphal taxonomy and systematics of the Psylloidea (Homoptera). By I. M. White & I. D. Hodkinson The whitefly of New Guinea (Homoptera: Aleyrodidae). ByJ. H.Martin Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester . - Nymphal taxonomy and systematics ofthe Psylloidea (Homoptera) I. M. White & I. D. Hodkinson Entomology series Vol 50 No 2 28 March 1985 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History),Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) Tustees of the British Museum (Natural History), 1985 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06009 Entomology series ISSN 0524-6431 Vol 50 No 2 pp 153-301 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 28 March 1985 Nymphal taxonomy and systematics of the Psylloidea(Homoptera) I. M. White Commonwealth Institute of Entomology, c/o British Museum (Natural History), CromwellRoad, London SW7 5BD I. D. Hodkinson Department of Biology, Liverpool Polytechnic, Byrom Street, Liverpool L3 3AF Contents Synopsis 153 Introduction 154 Methods of illustration 157 Material examined 157 Morphology of final instar nymphs 168 Selection of nymphal characters 179 Phenetic analysis of nymphs 212 Incorporation of adult characters 224 Cladistic analysis 228 Host-plant considerations 255 Zoogeographic evidence 258 Classification and phylogeny 261 Nymphal phenetic classification 261 General classification 263 Possible phylogeny 265 Keys 275 Key construction 275 Artificial key to families 275 Keys to subfamilies and genera of Aphalaridae 278 Key to genera and species groups of Spondyliaspididae 284 Key to subfamilies and genera of Psyllidae 285 Key to genera of Calophyidae 288 Key to genera of Phacopteronidae 288 Key to genera of Homotomidae 288 Key to subfamilies of Carsidaridae 289 Key to genera of Triozidae 289 Acknowledgements 290 References 291 Synopsis The objectives of this study are to form a phenetic classification of Psylloidea using nymphal characters; tocompare phenetic relationships suggested by nymphs with relationships suggested by adults; to combinenymphal data with existing adult data and produce a new classification; to devise keys based upon nymphalcharacters; to produce a predictive model (a cladogram) which describes the probable evolutionary historyof the Psylloidea. Nymphs of 303 species belonging to 94 genera were examined: these represented every existing psyllidfamily and included material from all zoogeographic regions. Phenetic analyses were carried out using cluster analysis and ordination. A summary phenetic classifica-tion of nymphs is presented which defines four major groups: species with sectasetae (mainly Triozidae);species with lanceolate setae (mainly Aphalaridae); species with capitate setae (most Psyllidae); specieswithout sectasetae, lanceolate setae or capitate setae (some species of each family). Bull. Br. Mus. not. Hist. (Ent.) 50 (2): 153-301 Issued 28 March 1985 154 I. M. WHITE AND I. D. HODKINSON Construction of an evolutionary ground plan for the Psylloidea was aided by the phenetic analyses. Acladogram of 106 psyllid genera and subgenera, based on this ground plan and using adult nymphalcharacters, is presented. Host-plant and zoogeographic evidence in association with the cladogram suggestthat the modern psyllids evolved from an ancestor associated with the plant order Rutales in Gondwana-land. General trends in the cladogram indicate that the following six families derived from an extinct ancestralgroup: Triozidae; Carsidaridae; Homotomidae (= Carsidaridae auctt., partim); Phacopteronidae (=Carsidaridae auctt., partim); Calophyidae (= Carsidaridae auctt., partim); Aphalaridae. The Spondylias-pididae and Psyllidae appear to have evolved from a common aphalarid ancestor. The main contributions made to psyllid systematics are: phenetic groups of nymphs are defined moreprecisely than they were previously; phenetic and cladistic studies indicate characters of especial value informing psyllid taxa of rank above the generic category, e.g. tarsal arolium structure; a theory of the originand evolution of psyllids is proposed incorporating information on nymphal and adult characters plushost-plant and zoogeographic data; a revised classification based on a cladogram incorporating taxa fromall zoogeographic regions is presented; keys to genera are produced based on nymphal characters; at thetheoretical level the method of cladogram construction is an advancement on that of Hennig (1966); ninenew family group taxa are proposed. Introduction Psyllids or jumping plant-lice (Homoptera, Sternorrhyncha, Psylloidea) are small (1-5 mmlong) phloem-sucking insects which breed almost exclusively upon perennial dicotyledonousplants (Eastop, 1972; Hodkinson, 1974). A review of psyllid biology is given by Hodkinson(1974). Nomenclatural history The first psyllid described was Chermes alni Linnaeus, 1758 (= Psylla aim), the type-species ofthe group. Four years later Geoff roy described Psylla, which is the type-genus, ChermesLinnaeus, 1758 having been suppressed and P. alni designated as the type-species by theInternational Commission on Zoological Nomenclature (Eastop, 1963 and Opinion 731, Bull,zool. Nom. 22: 86-87, 1965). The first major contribution to psyllid systematics was by Forster(1848) who described the genera Aphalara, Euphyllura, Rhinocola, Spanioneura and Trioza. Low (1879) produced the first formalized classification of the psyllids which he regarded asone family, the Psyllidae, comprised of four defined subfamilies: Liviinae (containing the genusLivid), Aphalarinae (containing the genera Aphalara, Euphyllura, Psyllopsis and Rhinocola),Psyllinae (Alloeoneura, Amblyrhina, Arytaina, Calophya, Diaphorina, Floria, Homotoma,Psylla and Spanioneura) and Triozinae (Bactericera and Trioza). Subsequently, Scott (1882)erected the subfamily Livillinae (for which he only lists Creiis) and the family Prionocnemidae(Carsidara and Tyora) but neither of these are valid as they are not based on recognised genera.Low's subfamilies were raised to family status by Edwards (1896) but this was not generallyaccepted for another 60 years. Schwarz (1898) erected a further subfamily, the Spondyliaspinae, for the genus Spondyliaspis ,and several genera, such as Carsidara, Tyora and Ciriacremum, were placed in the subfamilyCiriacreminae by Enderlein (1910). However, Crawford (1911) separated Ciriacremum fromCarsidara and erected a new subfamily, the Carsidarinae, to include such genera as Carsidaraand Tyora. Later, Aulmann (1913) listed six subfamilies: Psyllinae (e.g. Calophya, Diaphorina,Mycopsylla, Pauropsylla and Psylla), Triozinae (e.g. Bactericera and Trioza) , Aphalarinae (e.g.Aphalara, Cardiaspis (= Cardiaspina) , Euphalerus, Euphyllura and Phytolyma), Liviinae,Ciriacreminae (e.g. Carsidara, Ciriacremum and Phacopteron) and Spondyliaspinae. The classification was further revised by Crawford (1914) who recognised six subfamilies:Liviinae (Aphalara, Aphalaroida, Livia and Rhinocola) , Pauropsyllinae (Calophya, Heteropsyl-la, Paurocephala and Pauropsylla), Carsidarinae (Carsidara, Epicarsa, Freysuila sensu Schwarz(= Mastigimas) and Rhinopsylla) , Ceriacreminae (Ceriacremum (= Ciriacremum)), Triozinae(e.g. Trioza) and Psyllinae (e.g. Euphalerus, Euphyllura, Pachypsylla and Psylla). A similar NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 155 classification was presented by Pflugfelder (1941) except that Liviinae and Aphalarinae (includ-ing Diaphorina and Psyllopsis) were again separated. The Spondyliaspinae was first properly defined by Heslop-Harrison who reviewed thesubfamily groupings and separated the subfamilies Aphalarinae, Ciriacreminae (includingBactericera, Carsidara, Ciriacremum and Pauropsylla), Liviinae, Psyllinae, Spondyliaspinae,and Triozinae in a key (Heslop-Harrison 1949, 1951, 1954, 1958, 1959). TRIOZINAE-BACTERICERINAE -HOMOTOMINAE CARSIDARINAE -TENAPHALARINAE -PHACOPTERONINAE -LEPTYNOPTERINAE -PAUROPSYLLINAE CALOPHYINAE -LIVIINAE -PAUROCEPHALINAE -APHALARINAE -ANOMONEURINAE -SPONDYLIASPIDINAE-PACHYPSYLLINAE -CIRIACREMINAE-ARYTAININAE TRIOZIDAE CARSIDARIDAE JLIVIIDAE APHALARIDAE SPONDYLIA5PIDIDAE PSYLLINAE PSYLLIDAE Fig. 1 Cladogram showing the relationships of subfamilies according to Becker-Migdisova (1973). Superfamily status was given to the psyllids by Handlirsch (1903) and the subfamily units wereagain promoted to families by Vondracek (1957) who recognised the Aphalaridae (e.g.Aphalara, Paurocephala and Pauropsylla), Carsidaridae, Liviidae, Psyllidae (e.g. Calophya,Ciriacremum, Diaphorina, Psylla and Psyllopsis), Spondyliaspidae and Triozidae. Vondracek(1963) later replaced the Carsidaridae by the Ciriacremidae (which included Bactericera,Ciriacremum, Syndesmophlebia ( Klieniella) and Triozamia) and expanded the content ofSpondyliaspididae to include Anomalopsylla, Phytolyma and Tainarys. 156 I. M. WHITE AND I. D. HODKINSON The classification of Klimaszewski (1964) differed from that of Vondracek in certain respects:Bactericerinae (e.g. Triozamid) was moved to Triozidae, Anomalopsyllinae (e.g. Phytolymaand Tainarys) to the Aphalaridae, and Ciriacremum to the Psyllidae. Genera such as Carsidara,Homotoma and Tenaphalara were placed in the Carsidaridae. A similar classification forPalaearctic genera is given by Loginova (1964ft). The most recent comprehensive study of psyllid systematics was undertaken by Becker-Migdisova (1973), who produced a classification similar to that of Klimaszewski. SubsequentlyLoginova (1972, 1973, 19740, 1974ft, 1975, 19760, 1976ft, 1977) has revised several subfamiliesand tribes and since completion of this work, has produced a paper on nymphal morphology(Loginova, 1982). Recently Eastop (1978) presented a classification (attributed to D. Hollis) in which only twofamilies, the Psyllidae (including Aphalara, Calophya, Diaphorina, Livia, Psylla and Spondy-liaspis) and the Triozidae (including Carsidara and Trioza) were proposed. Contemporary classifications of the Psylloidea (Vondracek, 1957; Klimaszewski, 1964;Becker-Migdisova, 1973) are based on suggested phyletic trees and a summary of the tree ofBecker-Migdisova is given in Fig. 1. No attempt has been made to produce cladograms for thePsylloidea as a whole, but they have been constructed for a few genera and tribes, namelyGlycaspis (Moore, 1970), Psylla (Burckhardt, 1979), Strophingia (Hodkinson, 1981) andCiriacremini (Hollis, 1976). Comprehensive faunal surveys have been made for several temperate and subtropicalregions. They include Alaska (Hodkinson, 1978), Australia (Tuthill & Taylor, 1955), centralEurope (Haupt, 1935), Czechoslovakia (Vondracek, 1957), European U.S.S.R. (Loginova,1964a), Great Britain (Hodkinson & White, 1979ft; White & Hodkinson, 1982), Mallorca(Hodkinson & Hollis, 1981), New Zealand (Tuthill, 1952), North America (Crawford, 1914;Tuthill, 1943), Poland (Klimaszewski, 1969, 1975), Rumania (Dobreanu & Manolache, 1962),Spain (Gomez, 19560, 1956ft, 1960) and Switzerland (Schaefer, 1949). The only relativelycomplete faunal surveys of major tropical areas are for India (Mathur, 1975) and Taiwan (Yang,1984). Other substantial faunistic papers on psyllids cover Central Africa (Vondracek, 1963), SouthAfrica (Pettey, 1924, 1925, 1933; Capener, 1968, 1970, 1973), Central America (Crawford,1914; Caldwell, 19440, 1944ft; Tuthill, 1944, 1945, 1950), South America (Crawford, 1914, 1925;Lima, 1942; Tuthill, 1959, 19640), Borneo (Crawford, 1920), Hawaii (Zimmerman, 1948),Japan (Miyatake, 1963, 1964), Micronesia (Tuthill, 1964ft), Philippines (Uichanco, 1921:Miyatake, 1971, 1972) and Puerto Rico (Caldwell & Martorell, 1952). Comprehensive bibliog-raphies of taxonomic literature are available for the Palaearctic (Klimaszewski, 1973) andNeotropical (Hodkinson & White, 1981) regions and the Austro-Oriental, Pacific andHawaiian regions (Hodkinson, 1983). Aims of present study A classification such as that of Becker-Migdisova (1973) is adequate for studies on temperatepsyllid faunas. However, an increasing knowledge of tropical psyllids has brought with it arealisation that a classification which has been largely based on a knowledge of north temperatepsyllids is perhaps inappropriate when applied to tropical forms. This problem cannot be resolved simply by defining more major groups; new information of asuitable nature for incorporation in a systematic study is required. A study of nymphalmorphology is one possible source of such information. The aims of this work were: to investigate the phenetic taxonomic relationships of psyllids assuggested by nymphal data; to compare phenetic relationships based upon nymphal or adultdata; to pool nymphal data with existing adult data and produce a new predictive classification,i.e. a classification which has maximal likelihood of predicting unknown character states; toproduce a predictive model (a cladogram) against which other forms of data can be compared,and to use this model to derive a theory as to the age, possible origin and ancestral host of thepsyllids; to write provisional keys for the nymphs of psyllids. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 157 Over 2000 species of Psylloidea have been described and for nymphal data to be of any valuethe nymphal stages of a few hundred species, representing as many genera as possible, had to beexamined and described. This was facilitated by the use of numerical description and com-puterised data-handling techniques. Use of numerical taxonomic methods The use of phenetic methods indicates, within the bounds of the characters used, which taxa aremost similar to other taxa, without any characters being empirically weighted. They can initiatenew ideas, they are resistant to preconceived ideas, and they can help in deciding whether anattribute is ancestral or derived. The first application of numerical techniques to psyllid taxonomy was a study, based uponinadequate data, of Polish Trioza adults (Klimaszewski, 1967). Recently, Hodkinson (1981)used principal component analysis in a study of Strophingia adults. Previous studies of nymphal psy II ids Many descriptions of nymphal psyllids have been published and these are listed by White (1980).Prior to 1920 these were generally colour descriptions, such as those of Scott (18860, 18866,1886c), although a few authors, such as Low (1876, 1884, 1886), presented outline drawings. Thefirst descriptions of any taxonomic value were those of Ferris (1923, 1924, 1925, 1926, 1928a,19286) who also presented a phenetic classification of the nymphs (Ferris, 1925) which was laterexpanded by Rahman (1932). There are only three keys to nymphal psyllids: Swedish species ofPsylla (Ossiannilsson, 1970), subgenera of Psy Ha (Loginova, 1978) and the British Psylloidea(White & Hodkinson, 1982). Good nymphal descriptions of almost half the species known fromthe Indian subcontinent have been provided by Mathur (1975). Methods of illustration Nymphal morphology In whole nymph drawings (Figs 2-4) and in most illustrations of anal pore fields (Figs 97-160) thedorsal view is shown to the left of the body mid-line and the ventral view to the right. Minimum spanning networks (MSN) MSN's use abbreviated generic names with numbers denoting species (Table 1). In Figs 178 and182 a summary of each MSN is given in which species are only labelled with the initial letter ofthe family to which they belong in the classification of Becker-Migdisova (1973) (A - Aphalar-idae, C-Carsidaridae, L-Liviidae, P-Psyllidae, S-Spondyliaspididae, T-Triozidae). Phenograms Phenograms are labelled with the full names of species. Species adjacent in the phenogram butbelonging to different Becker-Migdisova families are spaced further apart than species of thesame family so as to give a visual impression of which clusters are highly congruent with thefamilies of Becker-Migdisova (congruent clusters appear more tightly packed than incongruentclusters). The initial letter of the family of Becker-Migdisova (1973) to which each speciesbelongs is indicated in the phenogram. Material examinedTaxonomic and zoogeographic coverage The final instar nymphs of 301 species were examined (Table 1). Two species descriptions(Tetragonocephala sp. from Ferris, 1926 and Togepsylla matswnurana from Miyatake, 1970)were incorporated from the literature as they were considered valuable additions to the study. 158 I. M. WHITE AND I. D. HODKINSON Figs 2-4 Psylloidea nymphs, morphological features of the three types of nymph defined by Rahman(1932). 2, psylline type nymph; 3, pauropsylline type nymph; 4, triozine type nymph, (a - antenna; apl -anal plate ; c - coxa ; cl - clypeus ; cp - cephaloprothorax ; cpl - caudal plate ; cr - circum-anal pore ring ; e -eye; f- femur; fw - forewing-pad; h - humeral lobe; hw - hindwing-pad; 1 - labium; ms - mesothoracicsclerites; mt - metathoracic sclerites; ss - sectasetae arranged 4 + 4 on abdomen margin; ts - tibiotarsus;t - tarsal segment II.) Modified from White & Hodkinson (1982). This represents a total species coverage of approximately 15 per cent and a generic coverage ofabout 43 per cent. For purposes of comparison the genera examined are fitted as closely as possible to thephylogenetic tree proposed by Becker-Migdisova (Fig. 1 and Table 1). The generic and specific representation across each recognised family and each zoogeographicregion are given in Tables 2 and 3. The absolute total number of genera and species is not givenbecause taxonomic position and status of many groups is uncertain; the Liviidae, however,contains only one genus. Recognition of final instar nymphs In all cases where the life-cycle of a psyllid has been carefully studied five nymphal instars wererecorded (e.g. Mathur, 1975). The most certain method of recognising a fifth instar nymphal NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 159 Table 1 Species examined listed according to the classification of Becker-Migdisova (1973) . Type-speciesof genera and subgenera, when examined, are indicated by an asterisk. Genera not examined byBecker-Migdisova are indicated by t. Abbreviated generic and subgeneric names of up to eight smallcapital letters are given on the right of each generic and subgeneric name. These abbreviations are used inthe minimum spanning network diagrams, in combination with the species numbers. Species numbers aregiven in parentheses on the right-hand side of the table. Depositories of material are indicated after each species name. Locality data for each specimen are givenby White (1980). Abbreviations of depositories BMAG Bolton Museum and Art Gallery, Bolton, Lanes. BMNH British Museum (Natural History), London. FRI Forest Research Institute, Dehra Dun, India. IDH I. D. Hodkinson (private collection), Liverpool. LC Loyola College, Madras, India. TRL Tasmanian Regional Laboratory, Commonwealth Scientific and Industrial Research Orga- nisation, Hobart, Tasmania, Australia. UC University of California, Davies, California, U.S.A. USD AC United States Department of Food and Agriculture (California), Sacramento, California,U.S.A. USNM United States National Museum of Natural History, Entomological Collection at the Systema-tic Entomology Laboratory, USDA, Beltsville, Maryland, U.S.A. APHALARIDAEPaurocephalinaeRhinocolini Rhinocola Forster, 1848 RHINOCOL*aceris (Linnaeus, 1758) [BMNH] ^LeurolophusTuthill, 1942 LEUROLOP *vittatus Tuthill, 1942 [USNM]Euphyllurini Euphyllura Forster, 1848 EUPHYLLU taethiopica Silvestri, 1934 [BMNH] (1) olivina (Costa, 1939) [BMNH] (2) *phillyreae Forster, 1848 [BMNH] (3) ^NeophylluraLoginova, 1973 NEOPHYLLArbutophila Loginova, 1973 *arbuti (Schwarz, 1904) [USNM] (1)Neophyllura Loginova, 1973 *arctostaphyli (Schwarz, 1904) [USNM] (2) bicolor (Martin, 1931) [USNM] (3)Paurocephalini Paurocephala Crawford, 1914 PAUROCEP gossypii Russell, 1943 [BMNH] (1) urenae Russell , 1946 [USNM] (2) StrophingiaEnderltm, 1914 STROPHIN dnereae Hodkinson, 1971 [BMNH] (1) *ericae (Curtis, 1835) [BMNH] (2) AgonoscenaEnderlem, 1914 AGONOSCE A. sp. (A) [USNM] (1) A. sp. (B) [BMNH] (2) A. sp. (C) [BMNH] (3) Aphalaroida Crawford, 1914 [USD AC] APHROIDA inermis Crawford, 1914 [USD AC] (1) ?near pithecolobia Crawford, 1914 [USNM] (2) tCaraarofcttcenaHaupt, 1935 CAMAROTO *speciosa (Flor, 1861) [BMNH] (1) lunicolor Loginova & Parfent'ev, 1958 [BMNH] (2) ^Moraniella Loginova, 1972 MORANIEL*calodendri (Moran, 1968) [BMNH] 160 I. M. WHITE AND I. D. HODKINSON ^ParaphalaroidaLoginova, 1972 PARAPHAL *fremontiae (Klyver, 1930) [USD AC]AphalarinaeAphalarini Aphalara Forster, 1848 APHALARA curta Caldwell, 1937 [USNM] (1) exilis (Weber & Mohr, 1804) [BMNH] (2) monticola Hodkinson, 1973 [IDH] (3) "tnubifera Patch, 1912 [USNM] (4) persicaria Caldwell, 1937 [USNM] (5) polygoni Forster, 1848 [BMNH] (6) rumicis Mally, 1894 [USNM] (7) simila Caldwell, 1937 [USNM] (8) CraspedoleptaEnderlein, 1921 CRASPEDO langustipennis (Crawford, 1911) [USNM] (1) *artemisiae (Forster, 1848) [BMNH] (2) Ivancouverensis (Klyver, 1931) [USNM] (3) Iconstricta (Caldwell, 1936) [USNM] (4) furcata (Caldwell ,1936) [USNM] (5) minuta (Caldwell ,1938) [USNM] (6) minutissima (Crawford, 1914) [USNM] (7) nebulosa (Zetterstedt, 1828) [IDH] (8) nervosa (Forster, 1848) [BMAG] (9) sonchi (Forster, 1848) [IDH] (10) suaedae (Crawford, 1914) [USNM] (11) subpunctata (Forster, 1848) [BMNH] (12) tveaziei (Patch, 1911) [USNM] (13) ^GyropsyllaBrethes, 1921 GYROPSYL ilicis (Ashmead, 1881) [USNM] (1) spegazziniana (Lizer, 1917) [USNM] (2)Colposceniini ColposceniaEndertein, 1929 COLPOSCEC. sp. [USNM] CrosrimzLoginova,1964 CRASTINA llinavuorii Loginova, 1974 [BMNH]AnomalopsyllinaeApsyllini Apsylla Crawford, 1912 APSYLLA *cistellata (Buckton, 1896) [LC]Anomalopsyllini Tfl/mzrysBrethes, 1920 TAINARYS *sc/um Brethes, 1920 [USNM]Phytolymini Phytolyma Scott, 1882 PHYTOLYM fusca Alibert, 1947 [BMNH] (1) *lata (Walker, 1852) [BMNH] (2) minuta (Hollis, 1973) [BMNH] (3) CARSIDARIDAE Calophyinae Calophya Low, 1878 CALOPHYA 1 calif ornica Schwarz, 1904 [USD AC] (1) dubia Crawford, 1914 [USNM] (2) flavida Schwarz, 1904 [USNM] (3) nigripennis Riley, 1883 [USNM] (4) *rhois (Low, 1877) [IDH] (5) schini Tuthill , 1959 [USNM] (6) triozomima Schwarz, 1904 [USNM] (7) rotundipennis White & Hodkinson, 1980 [BMNH] (8) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 161 PauropsyllinaeMicroceropsylliniMicroceropsylla Boselli , 1930 MICROCER M. sp. [BMNH]PelmatobrachiaEnderlein, 1921 PELMATOB P. sp. [LC]Pauropsyllini PauropsyllaRubsaamen, 1899 PAUROPSY beesoni Laing, 1930 [BMNH] (1) depressa Crawford, 1912 [LC] (2) trichaeta Pettey, 1924 [BMNH] (3) Leptynopterinae Leptynoptera Crawford, 1919 LEPTYNOP *sulfurea Crawford, 1919 [BMNH] Phacopterinae Pseudophacopterini PseudophacopteronEnderlein, 1921 PSEUDOPH floccosa (Crawford, 1915) [USNM] (1) sp. (A) [BMNH] (2) sp. (B) [BMNH] (3) Phacopterini PhacopteronBuckton, 1894 PHACOPTE *lentiginosum Buckton, 1894 [USDAC]TenaphalarinaeTenaphalariniProtyora Kieffer, 1906 PROTYORA *sterculiae (Froggatt, 1901) [BMNH] TenaphalaraKuwayama, 1907 TENAPHAL tacutipennis Kuwayama, 1907 [BMNH] (1) malayensis Crawford, 1919 [BMNH] (2) sp. [BMNH] (3) Togepsyllini Togepsylla Kuwayama, 1931 TOGEPSYL matsumurana Kuwayama, 1949 [described from Miyatake, 1970]DiclidophlebiiniDididophlebia Crawford, 1920 DICLIDOP teastopi Vondracek, 1963 [BMNH]Mastigimatini MastigimasEnderlein, 1921 MASTIGIM cedrelae (Schwarz, 1899) [BMNH] (1) sp.(A)[IDH] (2) sp. (B) [BMNH] (3) CarsidarinaeMesohomotominiEpicarsa Crawford ,1911 EPICARSA E. sp. [USNM] Mesohomotoma Kuwayama, 1907 MESOHOMO hibisci (Froggatt, 1901) [BMNH] (1) tessmanni (Aulmann, 1912) [BMNH] (2) ParacarsidaraHeslop-Harrison, 1960 PARACARS dugesii (Low, 1886) [USNM] (1) gigantea (Crawford, 1911) [BMNH] (2) sp. [USNM] (3) Unplaced Wharatiana Mathur, 1974 BHARATIA *octospinosa Mathur, 1974 [BMNH] 162 I. M. WHITE AND I. D. HODKINSON HomotominaeSynoziini SynozaEnderlein, 1918 SYNOZA floccosa Ferris, 1928 [UC] (1) sp. [USNM] (2)Dynopsyllini Macrohomotoma Kuwayama, 1907 MACROHOM *gladiatum Kuwayama, 1907 [BMNH] (1) striata Crawford, 1925 [BMNH] (2) MycopsyllaFroggatt, 1901 MYCOPSYL rfici (Tryon, 1895) [BMNH] (1) gardenensis Bhanotar, Ghosh & Ghosh, 1972 [BMNH] (2) ^Pseudoeriopsylla Newstead ,1911 PSEUDOER AryosaeNewstead, 1911 [BMNH]Homotomini Homotoma Guerin-Meneville, 1834 HOMOTOMA *ficus (Linnaeus, 1767) [BMNH] (1) indica (Mathur, 1975) [FRI] (2) LIVIIDAE Livia Latreille , 1 804 LIVI A coloradensis Crawford, 1914 [USNM] (1) crefeldensis (Mink, 1855) [BMNH] (2) *juncorum (Latreille , 1798) [IDH] (3) maculipennis (Fitch, 1857) [USNM] (4) vernalis Fitch, 1851 [USNM] (5) PSYLLIDAE CiriacreminaeCiriacremini CmacrewwmEnderlein, 1910 CIRIACRE capeneri Hollis, 1976 [BMNH] (1) capense Enderlein, 1923 [BMNH] (2) harteni Hollis, 1976 [BMNH] (3) julbernardioides Hollis, 1976 [BMNH] (4)Anomoneurini AnomoneuraSchwaTZ, 1896 ANOMONEU *mori Schwarz, 1896 [USNM]ArytaininaeDiaphorini DiaphorinaLov/,1879 DIAPHORI albomaculata Capener, 1970 [BMNH] (1) cardiac Crawford , 1924 [BMNH] (2) chobauti Puton, 1898 [BMNH] (3) citri Kuwayama, 1907 [BMNH] (4) clutiae Capener, 1970 [IDH] (5) florea Capener, 1970 [IDH] (6) punctulata (Pettey, 1924) [BMNH] (7) *putonii Low, 1878 [USNM] (8) solani Capener, 1970 [IDH] (9) ^Pennavena Capener, 1968 PENNAVEN *fabulosa Capener, 1968 [IDH]Arytainini ArytainaForster, 1848 ARYTAINA*genistae (Latreille, 1804) [BMAG] ^Acizzia Heslop-Harrison ,1961 ACIZZIA acaciae (Maskell, 1894) [BMNH] (1) acaciaebaileyanae (Froggatt, 1901) [BMNH] (2) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 163 hakeae (Tuthill, 1952) [BMNH]russellae Webb & Moran, 1974 [BMNH]uncatoides (Ferris & Klyver, 1932) [BMNH] ^Amorphicola Heslop-Harrison, 1961*amorphae (Mally, 1894) [USNM] ^ArytainillaLoginova, 1972 cytisi(Puton, 1873) [BMNH]HakaniLoginova, 1972 [BMNH]spartiicola (Sulc, 1907) [BMNH]spartiophila (Forster, 1848) [BMNH] ^Ceanothia Heslop-Harrison, 1961 aculeata (Crawford, 1914) [USNM]*ceanothi (Crawford, 1914) [USNM] ^EuceropsyllaBoselli, 1929 cayeyensis (Caldwell, 1942) [USNM]minuticona (Crawford, 1914) [USNM]*russoiBoselli, 1949 [USNM]sp. [USNM] \Euglyptoneura Heslop-Harrison, 1961fuscipennis (Crawford, 1914) [USNM]robusta (Crawford, 1914) [USNM]sp. [USNM] Gloria Low, 1879 variegata Low, 1881 [BMNH] Mnsnesia Tuthill, 1964 glabruscuta (Caldwell, 1942) [USNM] ^Purshivora Heslop-Harrison, 1961chelifera (Crawford, 1914) [USNM]pubescens (Crawford, 1914) [USNM]Euphalerini Colophorina Capener, 1973 *cassiae Capener, 1973 [BMNH] EuphalerusSchwarz, 1904 gallicolus Ferris, 1928 [UC]jugovenosus Tuthill, 1937 [USNM]nidifex Schwarz, 1904 [USNM]rugipennis Crawford, 1914 [USNM]to/Ufl/usTuthffl, 1937 [USNM]vermiculosus Crawford, 1914 [USNM]sp. (A) [BMNH]sp. (B) [BMNH]sp.(C)[LC]sp. (D) [BMNH]Psyllopseini Psyllopsis Low, 1879 fraxini (Linnaeus, 1761) [BMAG]Jraxinicola (Forster, 1848) [BMNH]Psyllinae Psylla Geoffrey, 1862AsphagidellaEnderlein, 1921 *buxi (Linnaeus, 1758) [BMNH]BaeopelmaEnderlem, 1926 foersteri Flor, 1861 [BMNH]Cac0/w_y//flOssiannilsson, 1970*ma//(Schmidberger, 1836) [IDH]peregrina Forster, 1848 [BMNH]sorbi (Linnaeus, 1758) [IDH]stricklandi (Caldwell, 1939) [USNM]w/ro/ Forster, 1848 [BMNH] (3)(4)(5) AMORPHIC ARYNILLA(1) (2)(3)(4) CEANOTHI(1) (2) EUCEROPS(1) (2)(3)(4) EUGLYPTO(1) (2)(3) FLORIA INSNESIA PURSHIVO(1) (2) COLOPHOR EUPHALER(1) (2)(3)(4)(5)(6)(7)(8)(9)(10) PSYLLOPS(1) (2) P-ASPHAGP-BAEOPE P-CACOPS(1) (2)(3)(4)(5) 164 I. M. WHITE AND I. D. HODKINSON Hepatopsylla Ossiannilsson, 1970albagena (Caldwell, 1938) [USNM]ambigua6rster, 1848 [IDH]brunneipennis Edwards, 1896 [BMNH]moscovita Andrianova, 1848 [BMNH]myrtilli Wagner, 1847 [IDH]*nigrita (Zetterstedt, 1828) [IDH]Iparallela Crawford, 1914 [USNM]palmeniLow, 1883 [IDH]pulchra (Zetterstedt, 1840) [BMNH]pyri (Linnaeus, 1758) [USNM]pyricola Forster, 1848 [BMNH]salicetiFoTster, 1848 [BMNH]visci Curtis, 1835 [BMNH] ? Hepatopsylla Ossiannilsson, 1970alba Crawford, 1914 [USNM]americana Crawford, 1914 [USNM]annulata Fitch, 1851 [USNM]brevistigmata Patch, 1912 [USNM]Icoryli Patch, 1912 [USNM]/iflwa/flTuthill, 1944 [IDH]hirsutaTuthill 1938 [USNM]magnicauda Crawford, 1914 [USNM]minor v.flava Crawford, 1914 [USNM]minuta Crawford, 1914 [USNM]negundinis Mally, 1895 [USNM]rhododendriPuton, 1871 [USNM]sinuata Crawford, 1914 [USNM]subspiculata Hodkinson, 1976 [IDH] Osmopsylla Loginova, 1978 *ribesiae Crawford, 1911 [USNM] Psylla Geoffrey, 1862 *alni Linnaeus, 1758 [BMNH]betulaenanae Ossiannilsson, 1970 [IDH]carpinicola Crawford, 1914 [USNM]floccosa Patch, 1909 [USNM]galeaformis Patch, 1911 [USNM]striata Patch, 1911 [USNM]trimaculata Crawford, 1911 [USNM] Thamnopsylla Loginova, 1978melanoneura Forster, 1848 [BMNH]*pyrisuga Forster, 1848 [BMNH]rhamnicola Scott, 1876 [BMAG] 1 'Thamnopsylla Loginova, 1978magna Crawford, 1914 [USNM]mediaTutm, 1943 [USNM]/>rwm'(Scopoli, 1763) [IDH]sp. near simlae Crawford, 1912 [BMNH] subgenus not certainphoradendraeTuthill 1939 [USNM]pulchella Low, 1878 [BMNH]"\Spanioneura Forster, 1848 *fonscolombii Forster, 1848 [BMNH]Unplaced t4/"epm*Tuthill,1959 A. sp. [USNM]t/7/ps_y//aKuwayama, 1907E. sp. (A) [BMNH]E. sp. (B) [BMNH] P-HEPATO (14) (15)(16)(17)(18)(19)(20)(21)(22)(23)(24)(25)(26)(27) P-OSMOPS O-PSYLLA(1) (2)(3)(4)(5)(6)(7) P-THAMNO(1) (2)(3) (4)(5)(6)(7) PSYLLA(1) (2) SPANIONE AREPUNA EPIPSYLL(1) (2) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 165 fFreysuilaAleman, 1887 FREYSUIL F. sp. [USNM] l>Heteropsylla Crawford, 1914 HETEROPS incisa (Sulc, 1914) [BMNH] (1) *texana Crawford, 1914 [USD AC] (2) "flsogonoceraiaTuthill, 1964 ISOGONOC divergipennis White & Hodkinson, 1980 [BMNH] ^Mitrapsy lla Crawford, 1914 MITRAPSY Ideserata Caldwell, 1944 [USNM] "fNeopsylliaCaldwell, 1947 NEOPSYLL erythrinae (Lizer, 1918) [USNM] (1) sp. [USNM] (2) ^Pexopsylla Jensen ,1957 PEXOPSYL *cercocarpi Jensen, 1957 [USNM] ^PlatycoryphaTuthill, 1945 PLATYCOR *princepsTuihi\\, 1945 [USNM] "\Retroacizzia Heslop-Harrison, 1961 RETACIZZ *flrttertrtfltaHeslop-Harrison, 1961 [BMNH] ^Trigonon Crawford, 1920 TRIGONON *longicornis (Crawford, 1919) [USNM] SPONDYLIASPIDIDAE Spondyliaspidinae SpondyliaspisSignoret, 1879 SPONDYLI5. sp. [BMNH] ^Cardiaspina Crawford ,1911 CARDIASP albitextura Taylor, 1962 [BMNH] (1) demitexta Taylor, 1962 [IDH] (2) squamula Taylor, 1962 [TRL] (3) sp. [TRL] (4) tOw Scott, 1882 CREIISC. sp. [TRL] ^Ctenarytaina Ferris & Klyver, 1932 CTENARYT* eucalypti (Maskell, 1890) [BMNH] ^EucalyptolymaFroggatt, 1901 EUCALYPTE. sp. [BMNH] fGlycaspis Taylor 1960 GLYCASPI baileyi Moore, 1961 [BMNH] (1) imponens Moore, 1961 [BMNH] (2) rivalis Moore, 1961 [BMNH] (3) aggregata Moore , 1961 [BMNH] (4) conflecta Moore, 1961 [BMNH] (5) conserta Moore, 1961 [BMNH] (6) cyanoreios Moore, 1961 [BMNH] (7) orientalis Moore, 1961 [BMNH] (8) salebrosa Moore, 1961 [BMNH] (9) \Phellopsylla Taylor 1960 PHELLOPS P. sp. [TRL]Pachypsyllinae Pachypsylla Riley, 1883 PACHYPSY Iceltidisgemma Riley, 1883 [USD AC] (1) celtidismamma (Riley, 1876) [USD AC] (2) celtidisvesiculum Riley, 1883 [USD AC] (3) japonica Miyatake , 1968 [USNM] (4) *venusta (Osten-Sacken, 1861) [USD AC] (5) Tetragonocephala Crawford, 1914 TETRAGONT. sp. [described from Ferris, 1926] 166 I. M. WHITE AND I. D. HODKINSON TRIOZIDAE BactericerinaeTriozamiini Triozamia Vondracek, 1963 TRIOZAMI *lamborni (Newstead, 1913) [BMNH]Unplaced Neolithus Scott, 1882 NEOLITHU N. sp. [BMNH]TriozinaeEutriozini Trichochermes Kirkaldy, 1904 TRICHOCH * walked (Forster, 1848) [BMNH]Paracomecini "\Leuronota Crawford, 1914 LEURONOT michoacana Ferris, 1828 [UC]Triozini Egeirotrioza Boselli , 1 93 1 EGEIROTR *ceardiv. euphratica Boselli, 1931 [BMNH] (1) verucifica Loginova, 1965 [BMNH] (2) sp.(A)[BMNH] (3) sp. (B) [BMNH] (4) Paratrioza Crawford ,1911 PARATRIO arbolensis Crawford, 1910 [USNM] (1) cockerelli (Sulc, 1909) [USNM] (2) lavaterae (Van Duzee, 1925) [USNM] (3) maculipennis (Crawford, 1910) [USNM] (4)Trioza Forster, 1848 BactericeraPuton, 1876 T-BACTER crithmi Low, 1880 [BMAG] (1) Icurvatinervis Forster, 1848 [BMNH] (2) Inigricornis Forster, 1848 [USNM] (3) salicivora Reuter, 1876 [USNM] (4)IBactericera Puton, 1876 atkasookensis Hodkinson, 1978 [IDH] (5) aylmeriae Patch, 1912 [USNM] (6) frontalis Crawford, 1910 [USNM] (7) obtusa Patch, 1911 [IDH] (8) tripunctata (Fitch, 1851) [USNM] (9) Heterotrioza Dobreanu & Manolache , 1962 T-HETERO alacris Flor, 1861 [BMNH] (1) albiventris Forster, 1848 [BMAG] (2) remota Forster, 1848 [IDH] (3) chenopodii Reuter, 1876 [BMNH] (4)1 Heterotrioza Dobreanu & Manolache, 1962 lobata Crawford, 1914 [USNM] (5) magnoliae (Ashmead, 1881) [USNM] (6) minuta Crawford, 1910 [USNM] (7) obsoleta (Buckton, 1900) [BMNH] (8) litseaelBordage, 1914 [BMNH] (9) Megatrioza Crawford, 1915 T-MEGATR diospyri (Ashmead, 1881) [USNM] (1) hirsuta (Crawford, 1912) [BMNH] (2) incidata Tuthill, 1945 [USNM] (3) palmicola Crawford, 1918 [USNM] (4) vitiensis (Kirkaldy, 1907) [BMNH] (5) Trioza Forster, 1848 T-TRIOZA cinnamomi (Boselli, 1930) [BMNH] (1) Imarginepunctata Flor, 1861 [USNM] (2) *urticae (Linnaeus, 1758) [BMNH] (3) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 167 ?7nozflF6rster, 1848 albifrons Crawford, 1910 [USNM] (4) bakeri Crawford, 1910 [USNM] (5) beameri Tuthill, 1939 [USNM] (6) erytreae (Del Guercio, 1918) [BMNH] (7) falcata (Ferris & Klyver, 1932) [BMNH] (8) panacis Maskell, 1890 [BMNH] (9) phoradendrae Tuthill, 1939 [USNM] (10) quadripunctata Crawford, 1910 [USNM] (11) vitreoradiata (Maskell, 1879) [BMNH] (12) subgenus not certain TRIOZA lanceps Tuthill , 1944 [USNM] (1) sp. [USNM] (2) ^Aacanthocnema Tuthill & Taylor, 1955 AACANTHO*7casuarinae (Froggatt, 1901) [IDH] t Ceropsylla Riley , 1 884 CEROPS YL Imartorelli Caldwell, 1942 [USNM] (1) Isideroxyli Riley, 1883 [UC] (2) sp. [USNM] (3) ^Crawforda Caldwell, 1940 CRAWFORDtriopsyllina Caldwell, 1940 [USNM] t//eva/ievaKirkaldy, 1902 HEVAHEVAswezeyi Crawford, 1928 [USNM] Mtuwayama Crawford, 1911 KUWAYAMApisonia Caldwell, 1940 [USNM] ^Swezeyana Caldwell, 1940 SWEZEYANelongagena Caldwell, 1940 [USNM] t Triozoida Crawford ,1911 TRIOZOIDsilvestris Tuthill, 1959 [USNM] Table 2 Numbers of genera and species examined in each psyllid family. skin was by the presence of a pharate adult within. Such specimens had at least one tarsalsegment separate from the tibiotarsus of the hindleg (and usually the fore and midlegs as well). Itwas therefore assumed that the maximum differentiation of tarsi from the tibia occurred in thefinal instar. As a further check the size of the final instar nymph was compared with the adult ofthe same species. With experience the size of the wing-pads relative to the body proved to be afurther confirmatory character. 168 I. M. WHITE AND I. D. HODKINSON Table 3 Numbers of species and genera, and total genera recorded from each zoogeo-graphic region. 'Figure includes species collected outside the region. Morphology of final instar nymphs Pflugfelder (1941) gives a general account of nymphal morphology. The following account isintended as an explanation of terminology, a discussion of possible homology and an indicationof the taxonomic distribution of attributes within the superfamily Psylloidea. Major body partsare shown in Figures 2-4. General form Nymphal psyllids are generally dorso-ventrally flattened, a condition reaching its maximumexpression in pit-gall inhabiting species, e.g. many Trioza spp. (Fig. 4). Ferris (1925) definedtwo morphological types of nymph, i.e. 'psylline' and 'triozine'. This division was made on thebasis of wing-pad shape. The 'psylline' type (Fig. 2) does not have the fore wing-pad producedanteriorly into a prominent humeral lobe and the apical extremity projects prominently from thecontour of the body. In the 'triozine' type there is an anteriorly produced humeral lobe (Fig. 4)to the forewing-pad and the margins of the pads are confluent with the body margin. Rahman(1932) defined a third type of nymph in which the humeral lobe is present but not producedanteriorly and the forewing-pad margin is parallel to the general body contour. He termed thisthe 'pauropsylline' type (Fig. 3) and gave examples of each of the three nymphal forms, e.g.Aphalara calthae and Pauropsylla depressa ('pauropsylline'), Paracarsidara gigantea and Psyllaalni ('psylline') plus Diaphorina citri and Trioza urticae ('triozine'). These descriptions ofmorphological types were found to be inadequate and emphasis was placed upon the charactersdescribed below. Head The main dorsal sclerite or pair of sclerites generally extend posteriorly so that the hind margin isposterior to the procoxae. This suggests that these sclerites are derived from a fusion of thevertex and part of the pronotum at least. This fusion is least expressed in the Spondyliaspididae,whereas complete fusion is observed in many Carsidaridae and Triozidae, especially thoseforming pit-galls. The length, basal position and number of apparent segments of the antennae vary within thegroup. Free-living forms such as Psylla alni have long antennae with many apparent segmentsand the antennal base is on the margin of the head. Most Triozidae and Calophya spp. whichinhabit pit-galls have short antennae, often with a total of only one apparent segment, and the NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 169 Figs 5, 6 Body measurements. 5, psylline type nymph; 6, triozine type nymph. (AL- antenna length; BB- body breadth; BL - body length; WL - forewing-pad length.) Modified from White & Hodkinson(1982). base is usually ventrally placed (Fig. 4). Adults normally have 10 antennal segments withrhinaria or antennal sensoria at the apices of segments IV, VI, VIII and IX. Most nymphs alsohave four rhinaria which, when 10 antennal segments are present, occur on segments IV, VI,VIII and IX. In those nymphs with fewer than 10 apparent segments fine sutures can sometimesbe seen suggesting the positions where further division of the antennae will occur. However, toavoid implication of homology between nymphs and adults, the term 'division' will be used andthe divisions will be numbered with arabic numerals. Rhinaria positions are assumed to behomologous with those of the adult and the first, second, third and fourth rhinaria are referred toas rhinaria IV, VI, VIII and IX respectively. Some nymphal Carsidaridae (Microceropsylla sp. and Calophya rotundipennis) appear tohave fewer than four rhinaria. Many adult Aphalaridae have six rhinaria placed singly at theapices of segments IV, V, VI, VII, VIII and IX. However, their nymphs have four, five or sixrhinaria, e.g. the adults of Craspedolepta artemisiae and C. angustipennis have four and fiverespectively. The mouthparts are of the normal sternorrhynchous type, i.e. the apex of the clypeus extendsto the mesothorax. The labium is two-segmented. Thorax The prothoracic tergum is assumed to be the area posterior to the eyes and anterior to thethoracic area which is continuous with the dorsal surface of the mesothoracic or fore wing-pads.Most of the sclerites are generally fused with the head to form the 'cephaloprothorax'. TheAphalaridae and Psyllidae normally have at least 2 + 2 sclerites between the mesothorax andcephaloprothorax (Fig. 9) whereas most Triozidae have 1-1-1 (Fig. 10) or no sclerites of theprothorax. 170 I. M. WHITE AND I. D. HODKINSON Figs 7-10 Circum-anal pore rings, antenna and prothorax forms. 7, circum-anal pore ring terminologyand measurement; 8, antenna not extending beyond head margin, character N35; 9,2 + 2 prothoracicsclerites (shaded), character N7; 10, 1 + 1 prothoracic sclerite (shaded), character N7. (a -antenna; an-anus; cl - clypeus; cp - cephaloprothorax; fw - forewing-pad; ir - inner circum-anal pore ring; or - outercircum-anal pore ring.) Fig. 7 from White & Hodkinson (1982). The mesothoracic tergum is taken to be the area between the lines of attachment of thefore wing-pads with the body. The sclerites are completely fused in many Triozidae, especiallythose forming pit-galls. Some species oiEuphalerus and Pachypsylla have large numbers of verysmall sclerites. In general the sclerites may be divided into laterals and medials. Nymphs of mostPsyllidae and Spondyliaspididae have small laterals and small medial sclerites (Fig. 11). Thenymphs of Aphalara and Diaphorina have elongate small lateral sclerites and at least 2 + 2enlarged medials (Fig. 12) and Paurocephala have no laterals but at least 2 + 2 large medials(Fig. 13). Egeirotrioza and Homotoma have 1 + 1 medial sclerites only (Fig. 14). The metathor-acic tergum is assumed to be the area between the lines of attachment with the hindwing-padswith the body. The sclerites of the metathoracic tergum take the same form as those of themesothoracic tergum. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 171 11 13 Figs 11-14 Thoracic morphology, character N8. 11, small lateral and small medial sclerites (shaded); 12,elongate lateral (adjacent to wing-pad) and 2 + 2 large medial sclerites on both meso- and metathorax;13, no lateral and 2 + 2 large medial sclerites on both meso- and metathorax; 14, no lateral and 1 + 1large medial sclerite on both meso- and metathorax. (fw - forewing-pad; hw - hindwing-pad.) Many species have paired depressions on each thoracic tergite. These are very well developedon the meso- and metathoracic tergites of Aacanthocnema casuarinae. The dorsal thoracic andabdominal areas of some Calophya spp. are covered in long processes (Fig. 144) while otherspecies of the genus have apparent 'perforations' in their dorsal sclerites. The forewing-pad is extended anteriorly as a humeral lobe in many Aphalaridae, Carsidar-idae, Diaphorina and Triozidae. In Aphalara and Diaphorina the humeral angle is normallyplaced away from the head margin (Fig. 17) while in Triozidae the humeral lobe may be adjacentto the eye (Fig. 19). In most species the apical (posterior) angle of the forewing-pad is adjacentor exterior to the margin of the hindwing-pad (Fig. 20). However, in some Calophya spp. andTriozidae the apical angle is interior to the hindwing-pad margin, and the margins of the 172 I. M. WHITE AND I. D. HODKINSON 15 16 19 Figs 15-19 Humeral lobe development, character Nl. 15, forewing-pad without a humeral lobe; 16,humeral lobe present, but not extending anterior to procoxa; 17, humeral lobe anterior to procoxa; 18,humeral lobe anterior to posterior margin of eye; 19, humeral lobe anterior to eye. (C - level of anteriormargin of procoxa; cp - cephaloprothorax; fw - forewing-pad; H - level of humerus; pc - procoxa.) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 173 21 22 23 Figs 20-26 Wing-pad and tarsal arolium forms. 20, apex of forewing-pad exterior to margin of hindwing-pad, character N2; 21, apex of forewing-pad interior to margin of hindwing-pad, but apex of hindwing-pad external to abdomen margin, characters N2 and N3; 22, apex of forewing-pad adjacent to margin ofhindwing-pad and wing-pad margins confluent, character N2; 23, apex of forewing-pad interior tomargin of hindwing-pad, apex of hindwing-pad interior to margin of abdomen and all margins confluent,characters N2 and N3; 24, triangular tarsal arolium, character N41; 25, triangular and petiolate arolium,character N4; 26, almost circular arolium, character N5. (a -arolium; ab- abdomen; fw- forewing-pad;hw - hindwing-pad; tc - tarsal claw.) 174 I. M. WHITE AND I. D. HODKINSON forewing-pad and hindwing-pad are confluent (Fig. 22). Furthermore, the hindwing-pad marginmay also be confluent with the abdominal margin (Fig. 23). These features are especially welldeveloped in species which form pit-galls. Adults of Leptynoptera sulfurea and Trioza diospyrihave very reduced hindwings and thus the nymphs have reduced hindwing-pads (Fig. 48).Wing-pad tracheation is sometimes observed just before adult moult and Heslop-Harrison(1951) illustrates this for Psylla sp. and Trioza sp. Psyllid nymphal legs lack such elaborate features as meracanthi, genual spurs, metatibialspines and metabasitarsal spines which are seen in the adult. Of the nymphs studies only two, i.e.Paraphalaroida fremontiae and Togepsylla matsumurana, appear to have articulate tarsi, as inthe adult (Fig. 55). Pachypsylla celtididisgemma and Pelmatobrachia sp. also have two-segmented tarsi but they appear non articulate (Fig. 52). Most species have a single apparenttarsal segment on each leg. This appears to be homologous with segment II of the adult becausethe apparent 'tibia' is often constricted sub-apically at a point assumed to represent the divisionline between the tibia and segment I (Fig. 53). The division line is often marked by simple setae.Because this division line is not always visible the apparent tibia and tarsus will hereafter becalled the tibiotarsus and the one apparent tarsal segment will be called a division. Most specieshave a pad between the tarsal claws. Ferris (1923, 1925, 1926), Ferris & Hyatt (1923) andRahman (1932) term this a pulvillus. Ferris (1928a, 19286) uses the word empodium and Lai(1937) was inconsistent. However, the correct term is arolium (Imms, 1957; Chapman, 1971).Arolia are very reduced, or absent, in some Carsidaridae, while in many Triozidae they are atleast semicircular (Fig. 26) and in most Psyllidae the arolia are petiolate (Fig. 25). Abdomen The abdomen is assumed to be any body part posterior to the join of the metathoracic tergumand hindwing-pads. Any short broad free sclerites near the base of the abdomen are discountedwhen describing the sclerites of the abdomen as they may equally well be of thoracic origin. Inmost species free sclerites and transverse rows of setae indicate the segments. In these speciesthe sclerites are arranged at least 1 + 1 per segment but in the apical area the individual scleritesare often fused to form a caudal plate (Figs 2-4). Some Carsidaridae have completelymembraneous abdomens while in many Triozidae the caudal plate covers the whole abdomen(Fig. 4). With this confusing set of possibilities no attempt has been made to homologise theabdominal sclerites. The ventral abdominal surface of most species has 2 + 2 sclerites on eachsegment anterior to the anal plate. The anal plate is formed by a fusion of the individual scleritesin the apical area. The most lateral free sclerites surround the spiracles. Because the true base ofthe abdomen is poorly defined and spiracles can 'migrate' or 'float' (Heslop-Harrison, 19520) orbe reduced in number (Matsuda, 1976) the possible homology of the spiracles was not studied. The abdomen of Paurocephala spp. has large apical tubercles (Fig. 114) and many Spondylias-pidinae have lateral bulges, coincident with each apparent segment (Fig. 28). Some species haveapical 'teeth' (Fig. 122). In most Pachypsylla spp. the central 'tooth' or 'teeth' are larger than themost lateral 'teeth' (Fig. 125) while in Euphalerus gallicolus the most lateral 'teeth' are enlarged(Fig. 120). Some species of each family have the anal opening posterior (i.e. apical) while others have aventral anal opening. In many species the sex of the final instar nymphs can be determined by theshape of the suture which extends anteriorly from the anus (Ball & Jensen, 1966; Ossiannilsson,1970; Hodkinson, 1973). Most species have an anal pore-field which usually surrounds the anus as a ring (Figs 7,130-142). This ring, which is often double, is called the circum-anal pore ring by Ferris (1928a).The following types of anal pore-fields have been observed. (i) Circum-anal ring only (Fig. 111). (ii) Circum-anal ring plus two additional rings placed laterally to it (Fig. 110). (iii) Two rings each lateral to the anus and no circum-anal ring (Fig. 150). (iv) Four rings and no circum-anal ring (Fig. 120). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 175 27 n 30 31 Figs 27-33 Abdomen shapes and setal types. 27, abdomen margin with lateral bulges, character N44; 28,abdomen margin serrate (typical of many Spondylaspididae), character N44; 29, capitate seta, showinghollow structure (as observed with a scanning electron microscope); 30, rod seta; 31, lanceolate seta; 32,sectaseta, pointed and without a ring; 33, sectaseta, pointed and with a ring. 176 I. M. WHITE AND I. D. HODKINSON (v) Outer circum-anal ring broken at two or more places (Fig. 97). (vi) Circum-anal ring very small and remainder of pore-field arranged as bands (Fig. 155). (vii) Circum-anal ring absent - otherwise as (vi) .(viii) Circum-anal ring plus round or ovoid groups of pores which are probably derived from an outer ring (Fig. 99).(ix) Small groups of pores (Fig. 154). (x) Individual or grouped pores arranged in rings (Fig. 123) or broken rings (Fig. 119). (xi) Small groups of pores dispersed in anal region, no circum-anal ring (Fig. 129). Ferris (1928a) discussed the homology of the anal pore-field. He concluded that the pore ringsof Euphalerus gallicolus (iv) were not homologous with the circum-anal ring. However, specieswere observed whose circum-anal rings were constricted, suggesting a tendency to break intoseparate rings, e.g. Livia spp. feeding on Carex (Fig. Ill) and Macrohomotoma gladiatum (Fig.148). This breakage of the circum-anal ring is complete in Livia spp. feeding onJuncus (Fig. 110)and in M. striata (Fig. 149). This suggests that the pore rings of E. gallicolus (Fig. 120) may be ofsimilar origin, i.e. homologous with the circum-anal ring. Groups of pores (type x) mayrepresent a reduction of pore rings (of type ii, iii or iv) as in Eucalyptolyma sp. (Fig. 119).Further breakdown may result in isolated pore groups (type xi) as in many Spondyliasipidinae(Fig. 129). The homology of pore bands (type vi & vii), e.g. Mesohomotoma hibisci (Fig. 155),remains uncertain although they may derive from the outer circum-anal ring. Setal types and chaetotaxy Simple setae, including the ring-based setae of Lai (1937), are simple articulated hairs showing ageneral distribution over many body parts in all major groups. It may be possible to homologisethe positions of some of these setae within a few groups. Because this study is concerned withformation of, and the relationships between higher groups, these setae are not furtherconsidered. Capitate setae (Ossiannilsson, 1970) or spatulate setae (Klyver, 1931; Lai, 1937) are defined asany seta which is apically dilated. Under the SEM the apex appears to be 'cup-like' (Fig. 29) andbroken setae reveal a hollow structure. Capitate setae are found on the head, dorsal surface ofthe thorax, wing-pads, tibia and abdomen of many Psyllidae. These setae are also found singly orin pairs at the tarsal apices of many psyllids and they are best developed in most Spondyliaspidi-nae(Fig. 38). Clavate setae (Ferris, 1923) are very short setae with a narrow base broadening gradually tomaximum breadth just prior to the blunt apex (Fig. 172). Clavate setae are observed in Arepunasp., many Diaphorina spp. and some Triozidae. In some individual species of Diaphorina theyoccur together with lanceolate setae while in some Triozidae they occur together with sectasetaeor scales. Clavate setae are very small and difficult to observe. In reality they may be modifiedlanceolate setae, sectasetae or scales. Sectasetae (Ferris, 1923; Boselli, 1929; Lai, 1937). These include dagger-shaped and spear-shaped setae (Lai, 1937), and dagger-like setae (Klyver, 1931). Sectasetae are defined as setaehaving an angle (Fig. 32) or ring (Figs 33, 34), around their circumference, in the basal thirdwhich is visible under phase contrast. Sectasetae arranged in the 1 + 1, 2 + 2, 3 + 3 and 4 + 4pattern, spaced from the next by more than their own length (Fig. 37), along the abdomenmargin occur in two forms. In Ciriacremum spp., Euceropsylla spp., Insnesia glabruscuta andIsogonoceraia divergipennis they are tubular. However, these sectasetae are pointed in otherPsyllidae. Species with this precise arrangement of abdomen margin sectasetae normally lacksectasetae on other body areas. The only exceptions are Neopsyllia spp. and Platycoryphaprinceps each of which have a single sectaseta on the hindwing-pad margin. Pointed sectasetae (Figs 32-33) arranged in large numbers on many body areas are a feature ofsome Aphalaridae, some Carsidaridae and some Triozidae. However, on most Triozidae the NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 177 Figs 34-37 Setal types and chaetotactic arrangements. 34, sectaseta, truncate and with a ring; 35, truncatesectasetae on a body margin, adjacent to each other, character N25; 36, scales on a body margin (typicalof many Hawaiian Triozidae); 37, the positions in which up to 4 + 4 setae are placed on the abdomenmargin of many Psyllidae, and the three types of setae which occupy these positions, namely lanceolate(e.g. Mitrapsylla deseratd), pointed sectaseta (e.g. many Psylla spp.) and the tubular shaped truncatesectaseta of Ciriacremum spp. (ab - abdomen; 1 - lanceolate seta; ss - sectaseta; ts - tubular sectaseta). 178 I. M. WHITE AND I. D. HODKINSON ss 41 Figs 38-41 Chaetotactic arrangements and antennae. 38, a pair of capitate setae at the apex of a tarsus(these setae are, especially well developed in many Spondyliaspididae), characters N20-N22; 39, acapitate seta placed behind the eye, character N13. Antennae. 40, Calophya californica; 41, Gyropsyllaspegazziniana. (c- capitate seta; cp-cephaloprothorax; fw-forewing-pad; r-rhinaria; ss-sectaseta; tc-tarsalclaw.) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 179 sectasetae are truncate (Fig. 34). Three distribution patterns of sectasetae could be recognisedon the antennae. One row on the opposite side to the rhinaria occurs in many Calophya spp.(Fig. 40) while one row adjacent to the rhinaria occurs in Diclidophlebia eastopi. Species withmore than one row (Fig. 45) are Moraniella calodendri, Paraphalaroida fremontiae,Paurocephala spp. and Togepsylla matsumurana. Lanceolate setae (Ferris, 1923; Boselli, 1929; Rahman, 1932; Lai, 1937) are defined as stout setaewith a convex profile and a constricted base (Fig. 31). The maximum breadth of the seta isnormally in the basal two-thirds. Lanceolate setae are a feature of most Aphalaridae, Diaphor-ini and Psyllopsis spp. They are also observed in a few Carsidaridae (e.g. Epicarsa sp.,Mycopsylla fid and Pseudophacopteron floccosa) and some Spondyliaspididae (Ctenarytainaeucalypti, Eucalyptolyma sp. and Phellopsylla sp.). Homotoma ficus has lanceolate setae basedupon tall tubercles (Fig. 171). Heteropsylla spp. have lanceolate setae arranged 3 + 3 in positions resembling the similarlyplaced sectasetae in many species of Psyllidae (Fig. 37). These lanceolate setae are treated ashomologous with similarly placed sectasetae and not with other lanceolate setae. Scales are defined as broad, apparently flat setae with a narrow base (Fig. 36) which are placedmarginally on some New World and Hawaiian Triozidae. Rod setae are long, parallel or subparallel-sided, with a constricted base (Fig. 30), and they arefound covering the bodies of Aphalaroida pithecolobia, Euglyptoneura robusta, Pexopsyllacercocarpi and Psylla ulmi. All attempts to recognise homology of setal positions across the whole of the Psylloidea failedexcept in one case, i.e. a single capitate seta placed laterally or sublaterally behind each eye(Fig. 39). In general the specialised setae occur in large numbers in any one body area. It isassumed that at least some of the setae in one body area of one species are homologous withsome of the same type of seta in the same body area in any other species. The only exception tothis rule is the sectasetae and lanceolate setae on the margin of the abdomen. Two arrangementsof these setae were recognised: a set of one to four pairs spaced well apart (Fig. 37) and secondlymuch larger numbers with no obvious individual positions. Selection of nymphal characters A set of 88 ordered multistate and two-state characters were initially defined. From thesecharacters those most likely to be of value in forming major groups were selected for a detailedseries of analyses. For character selection the ordered multistate characters were receded into 120 additivetwo-state characters (Sneath & Sokal, 1973). Characters which correlated with large numbers ofother characters were identified by the SUMRAT information statistic (Legendre & Rogers,1972). Only those characters having a SUMRAT value in excess of the mean value wereaccepted (Baum, 1977). Characters selected by this method were almost identical with thoseselected by an examination of character eigenvector values in a principal component analysis,carried out on the original character covariance matrix (see Davies & Boratynski, 1979 formethod). After these initial analyses, 45 selected two-state characters remained and these werecombined to form 34 ordered multistate and two-state characters (Table 4). Rejected charactersare summarised in Table 5. Once the number of characters has been reduced to 34, many species are identical as definedby the selected character set. Forty-eight species groups were formed, and one representativespecies was chosen from each (Table 6). The 134 species which remain distinct are also listed inTable 6, making a total of 182 selected species. All subsequent phenetic analyses are conductedusing the 34 selected characters and the 182 selected species. 180 I. M. WHITE AND I. D. HODKINSON Figs 42-47 Antennae. 42, Microceropsylla sp.; 43, Moraniella calodendri; 44, Mycopsylla gardenensis;45, Paurocephala gossypii; 46, Pauropsylla depressa; 47, Phacopteron lentiginosum. (r- rhinaria.) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 181 48 49 52 Figs 48-53 Wing-pads, hind tibia and tarsi. 48, Leptynoptera sulfurea wing pads, showing reducedhindwing-pad corresponding to the dipterous adult. Wing-pads and chaetotaxy, 49, Pauropsyllatrichaeta; 50, Trioza chenopodii. Hind tibia and tarsi, 51, Camarotoscena unicolor showing tibiotarsus(tibia + tarsal segment I) and the differentiated tarsal segment II; 52, Pelmatobrachia sp. showing thetibia and both tarsal segments differentiated; 53, Phytolymafusca showing tibiotarsus and tarsal segmentII. (fw - forewing-pad; hw - hindwing-pad.) 182 I. M. WHITE AND I. D. HODKINSON 55 Figs 54-66 Hind tibia and tarsi, and tarsal arolia (uniguitractor shown in black). Hind tibia and tarsi, 54,1 Pseudophacopteron floccosa (IChineura sp.) showing tibiotarsus, tarsal segment II and associatedsetae; 55, Togepsylla sp. showing both tarsal segments separate to tibia. Tarsal arolia of Aphalarinae: 56,Aphalara persicaria', 57, Colposcenia sp.; 58, Gyropsylla ilicis. Tarsal arolium of Aphalaroidinae. 59,Aphalaroida ? pithecolobia. Tarsal arolium of Diaphorininae: 60, Diaphorina solani; 61, Psyllopsisfraxinicola. Tarsal arolium of Euphylurini: 62, Euphyllura ? aethiopica; 63, Neophyllura (Arbutophila)arbuti; 64, N. (N.) arctostaphyli. Tarsal arolium of Paraphalaroidini: 65, Dididophlebia eastopi; 66,Paraphalaroidafremontiae. Scale line represents 0-05 mm. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 183 67 Figs 67-82 Tarsal arolia (unguitractor shown in black). Liviinae: 67, Livia maculipennis; 68, L. vernalis.Paurocephalinae: 69, Camarotoscena unicolor; 70, Paurocephala urenae. Strophingiinae: 71, Strophin-gia ericae. Rhinocolinae: 72, Agonoscena sp. (A);. 73, Leurolophus vittatus; 74, Moraniella calodendri;75, Rhinocola aceris; 76, Tainarys schini. Euphalerinae: 77, Euphalerus nidifex; 78, Retroacizziaantennata. Psyllidae: 79, Acizzia acaciaebaileyanae; 80, A. uncatoides; 81, Anomoneura mori; 82,Ciriacremumjulbernardioides. Scale line represents 0-05 mm. 184 I. M. WHITE AND I. D. HODKINSON 90 95 96 Figs 83-96 Tarsal arolia (uniguitractor shown in black). Psyllidae: 83, Epipsylla sp. (A); 84, Epipsylla sp.(B); 85, Euceropsylla cayeyensis; 86, Freysuila sp.; 87, Insnesia disjuncta; 88, Isogonoceraia sp.; 89,Mitrapsylla deserata; 90, Neopsyllia erythinae; 91, Platycorypha princeps; 92, Psylla parallela; 93,Trigonon longicornis. Phacopteronidae: 94, Pseudophacopteron sp. (A). Triozidae: 95, Egeirotrioza sp.(A); 96, Trioza hirsuta. Scale line represents 0-05 mm. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 185 100 101 102 103 Figs 97-103 Anal pore-fields of Apalaridae. 97, Agonoscena sp. (A); 98, Aphalara polygoni; 99,Camarotoscena speciosa; 100, C. unicolor (broken line indicates position of abdomen margin); 101,Craspedolepta nebulosa; 102, C. suaedae; 103, C. subpunctata. 186 I. M. WHITE AND I. D. HODKINSON 104 105 106 107 Figs 104-107 Anal pore-fields of Aphalaridae. 104, Craspedolepta ? vancouverensis; 105, Ctenarytainaeucalypti (inset shows detail of pore-field); 106, Diclidophlebia ? eastopi; 107, Euphyllura ? aethiopica. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 187 108 109 110 Figs 108-111 Anal pore-fields of Aphalaridae. 108, Gyropsylla ilicis; 109, Leurolophus vittatus; 110,Liviamaculipennis; 111, L. v emails. 188 I. M. WHITE AND I. D. HODKINSON 112 114 113 115 Figs 112-115 Anal pore-fields of Aphalaridae. 112, Neophyllura bicolor; 113, Paraphalaroida fremon-tiae; 114, Paurocephala gossypii; 115, Psyllopsisfraxinicola. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 189 116 117 Figs 116-118 Anal pore-fields. Aphalaridae: 116, Strophingia cinereae. Spondyliaspididae: 117, Col-ophorina cassiae; 118, Creiis sp. (insets show pore details and 'tooth' at apex of abdomen). 190 I. M. WHITE AND I. D. HODKINSON 120 Figs 119, 120 Anal pore-fields. Aphalaridae: 119, 'Eucalyptolyma' sp. (inset shows detail of pore field).Spondyliaspididae: 120, Euphalerus gallicolus . NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 191 121 122 Figs 121-123 Anal pore-fields of Spondyliaspididae. 121, Euphalerus jugovenosus; 122, E. nidifex; 123,Euphalerus sp. (A). 192 I. M. WHITE AND I. D. HODKINSON 125 126 Figs 124-126 Anal pore-fields and abdomen apex shapes of Spondyliaspididae. 124, Glycaspis baileyi(inset shows detail of pore-field); 125, Pachypsylla celtidismamma, which lacks a pore-field; 126, P.japonica (inset shows detail of pore-field). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 193 127 128 o Figs 127-129 Anal pore-fields and abdomen apex shapes of Spondyliaspididae. 127, Phellopsylla sp.(insect shows detail of a pore group); 128, Retroacizzia antennata, which lacks a pore-field; 129,Spondyliaspis sp. (inset shows detail of two of the pore groups). 194 I. M. WHITE AND I. D. HODKINSON Figs 130-133 Anal pore-fields and abdomen chaetotaxy of Psyllidae. 130, Anomoneura mori; 131,' Euphalerus' sp. (C); 132, Euglyptoneura fuscipennis; 133, Mitrapsylla deserata (inset shows detail offunnel setae). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 195 136 137 138 Figs 134-138 Anal pore-fields of Psyllidae. 134, Psylla buxi; 135, P. foersteri; 136, P. galeaformis; 137, P.phoradendrae; 138, P. pulchella. 196 I. M. WHITE AND I. D. HODKINSON 0000^ 139 140 142 Figs 139-142 Anal pore-fields of Psyllidae. 139, Psylla saliceti; 140, P. simlae; 141, P. sorbi (inset showspore structure of part of outer circum-anal ring); 142. Spanioneurafonscolombii. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 197 145 Figs 143-145 Anal pore-fields and abdominal structures of Calophyidae and Phacopteronidae.Calophyidae: 143, Apsylla cistellata (inset shows pore structure of parts of the pore-field which is outsidethe circum-anal rings); 144, Calophya californica, showing the long processes which cover the dorsalsurfaces of the abdomen and thorax. Phacopteronidae: 145, Bharatiana octospinosa. 198 I. M. WHITE AND I. D. HODKINSON 146 ,0o< 147 148 149 Figs 146-149 Anal pore-fields. Phacopteronidae: 146, Phacopteron lentiginosum (anus crosshatched);147, Pseudophacopteron sp. (B). Homotomidae: 148, Macrohomotoma gladiatum; 149, M. striatum. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 199 151 152 Figs 150-152 Anal pore-fields of Homotomidae. 150, Mycopsylla ? fid; 151, M. gardenensis; 152,Pseudoeriopsylla nyasae. 200 I. M. WHITE AND I. D. HODKINSON 155 Figs 153-155 Anal pore-fields and abdomen chaetotaxy. Homotomidae: 153, Synoza floccosa (insetshows abdomen margin sectaseta). Carsidaridae: 154, Mastigimas cedrelae; 155, Mesohomotomahibisci. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 201 156 157 * * **' /< ' Figs 156-158 Anal pore-fields and abdomen margin shapes of Triozidae. 156, Leuronota michoacana;157, Neolithus sp., showing shape of abdomen apex; 158, Triozamia lamborni (inset shows detail of apore area). 202 I. M. WHITE AND I. D. HODKINSON 0-05 0-05 164 Figs 159-164 Anal pore-fields and chaetotaxy. Triozidae: 159, Trioza alacris, with details of circum-analpore ring and abdomen margin setae; 160, Triozoida silvestris, circum-anal pore rings. Aphalaridae:161, Aphalaroida pithecolobia, abdomen margin rod setae; 162, Phytolyma minuta, circum-anal porerings and abdomen margin lanceolate setae; 163, Crastina linavuorii, abdomen margin lanceolate setae;164, Aphalara persicaria, abdomen margin lanceolate setae. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) JO-02 165 203 166 0-02 0-02 167 Figs 165-167 Chaetotaxy of Aphalaridae. 165, Diaphorina putonii, abdomen margin lanceolate setae;166, Moraniella calodendri, forewing-pad margin sectasetae; 167, Tainarys schini, abdomen marginlanceolate setae. 204 I. M. WHITE AND I. D. HODKINSON 169 0-01 .0-01 172 005 173 Figs 168-173 Chaetotaxy. Aphalaridae: 168, Rhinocola aceris, abdomen margin with truncate lanceolatesetae. Spondyliaspididae: 169, Arepuna sp., abdomen margin clavate setae. Psyllidae: 170, Pexopsyllacercocarpi, abdomen margin rod and sectasetae. Homotomidae: 171, Homotoma ficus , lanceolate setamounted on a tubercle. Triozidae: 172, Crawforda triopsyllina, forewing-pad dorsal surface clavate seta;173, Hevahevaswezeyi, abdomen margin scales. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 205 176 0-02 J \ 0-02 177 Figs 174-177 Abdomen margin scales of Triozidae. 174, Ceropsylla sideroxyli; 175, Kuwayama pisonia;176, Swezeyana elongagena; 111, Triozapalmicola, showing both scales and sectasetae. 206 I. M. WHITE AND I. D. HODKINSON Table 4 Selected nymphal characters. Selected characters are numbered N1-N34. All the character states in the original list of 88 characters aretabulated. However, some character states became combined by the selection procedure and, hence, insome characters two consecutive states are marked with the same value. Characters describing shape and position Nl . Humeral lobe (in the case of variability the code chosen was the highest observed) . No humeral lobe (Fig. 15). =0 Humeral lobe present, anterior margin of forewing-pad not extending anterior to procoxa(Fig. 16). = 1 Humeral lobe present, anterior margin of forewing-pad anterior to procoxa and posterior toeye (Fig. 17). =2 Humeral lobe present, anterior margin of forewing-pad anterior to posterior of eye andposterior to anterior of eye (Fig. 18). =3 Humeral lobe present, anterior margin of forewing-pad anterior to eye (Fig. 19). =4 N2 . Forewing-pad ; position of apex . Apex exterior to margin of hindwing-pad (Fig. 20). =0 Apex adjacent or interior to margin of hindwing-pad, margin of forewing-pad not confluentwith margin of hindwing-pad (Fig. 21). = Apex adjacent or interior to margin of hindwing-pad, margin of forewing-pad confluent withmargin of hindwing-pad (Fig. 22). =1 N3 . Hindwing-pad ; position of apex . Apex exterior to margin of abdomen (Fig. 21). = Apex adjacent or interior to margin of abdomen, margin of hindwing-pad not confluent withmargin of abdomen (Fig. 22). = Apex adjacent or interior to margin of abdomen, margin of hindwing-pad confluent withmargin of abdomen (Fig. 23). =1 N4-N5 . Form of tarsal arolium (clarifying characters of N41 ) . N4. Triangular and petiolate (Fig. 25).No = Yes = 1 N5. Disc-like or more than semicircular (Fig. 26).No = Yes = 1 N6. Anal opening ventral.No = Yes = 1 Characters of sclerite fusion N7. Prothorax dorsal sclerites. Extent of fusion with head. Prothorax dorsal sclerites completely separate from head. = Prothorax dorsal sclerites partly fused with head. At least 2 + 2 sclerites of prothoraxseparate to cephaloprothorax (Fig. 9). =0 Prothorax dorsal sclerites fused with head. 1 + 1 sclerite of prothorax separate to cephalo-prothorax (Fig. 10). =1Prothorax dorsal sclerites fused with head. = 2 N8. Mesothorax and metathorax dorsal surface sclerite arrangement (species which are mem-braneous or completely sclerotized, i.e., the separate sclerites are undifferentiated, arecoded zero). Medial sclerites (more than 1 + 1) small. Lateral sclerites small (Fig. 11). =0 Medial sclerites (more than 1 + 1) large. Lateral sclerites small (Fig. 12). = 1 Medial sclerites (more than 1 + 1) large. Lateral sclerites absent (Fig. 13). = 1 Medial sclerite (1 + 1) large. Lateral sclerites absent (Fig. 14). = 1 N9. Abdomen dorsal surface with many free sclerites. Caudal plate, if present, covering less thanwhole abdomen (small transverse sclerites at base of abdomen are discounted) (Fig. 2).No = Yes = 1 N10. Abdomen dorsal surface heavily sclerotized (caudal plate). At most with a few smalltransverse sclerites at the base of the abdomen (Fig. 4).No = Yes = 1 NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 207 Chaetotaxy characters Twelve following characters (N11-N22). Capitate setae present.No = Yes = 1 Nil. Head margin capitate setae. N12. Antenna with capitate setae close to rhinaria IV and VI. N13. Capitate seta placed laterally or sublaterally behind eye (Fig. 39). N14. Thorax dorsal capitate setae. N15. Forewing-pad dorsal capitate setae. N16. Forewing-pad margin capitate setae. N17. Hindwing-pad margin capitate setae. N18. Abdomen dorsal capitate setae. N19. Abdomen margin capitate setae. N20-N22 . Tarsal apex with 2 capitate setae (Fig . 38) . N20. Fore tarsi. N21. Midtarsi. N22. Hindtarsi. Three following characters (N23-N25). Lanceolate setae present.No = Yes = 1 N23. Forewing-pad margin lanceolate setae.N24. Hindwing-pad margin lanceolate setae. N25. Abdomen margin lanceolate setae present, numbering more than 4 + 4, or if fewer than5 + 5 they are separated by less than their own length. Eight following characters (N26-N33). Sectasetae. Up to three 'present' states were recognised in the initial coding. Absent. = Present and pointed (Figs 32, 33). = 1 Present and truncate (Fig. 34). =2 Present, truncate and each seta adjacent to the next or separated by less than one-quarter its maximum breadth from the next seta (Fig. 35). =3 N26. Head margin sectasetae. Absent = 0; Pointed = 1; Truncate = 2; Adjacent = 2 N27. Head dorsal sectasetae.Absent = 0; Pointed = 1. N28 . Mesothoracic and metathoracic dorsal sectasetae .Absent = 0; Pointed = 1; Truncate = 2. N29. Forewing-pad dorsal sectasetae. Absent = 0; Pointed = 1; Truncate = 2. N30. Forewing-pad margin sectasetae. Absent = 0; Pointed = 1; Truncate = 2; Adjacent = 2. N3 1 . Hindwing-pad margin sectasetae . Absent = 0; Pointed = 1; Truncate = 2; Adjacent = 2. N32. Abdomen dorsal surface sectasetae. Absent = 0; Pointed = 1; Truncate = 2. N33. Abdomen margin sectasetae numbering more than 4 + 4, of if less than 5 + 5 they are eachseparated from the next by less than their own length.Absent = 0; Pointed = 1; Truncate = 2; Adjacent = 2. N34. Abdomen margin sectasetae or lanceolate setae present and numbering 1 + 1, 2 + 2, 3 + 3,or 4 + 4 and each separated by more than their own length (Fig. 37).No = Yes = 1 208 I. M. WHITE AND I. D. HODKINSON Table 5 Rejected nymphal characters. Rejected characters are numbered N35-N88. Values are not given against the states, which are separated by a '/'. Characters describing shape and positionN35. Antenna base position. On head margin, or if ventral antenna apex extends beyond head margin (Fig. 4)/Ventral, antenna apex not extending beyond margin of head (Fig. 8). N36. Antenna with one rhinarium (Fig. 42). No/Yes. N37. Antenna with five rhinaria. No/Yes. N38. Antenna with six rhinaria. No/Yes. N39. Thorax with pairs of large depressed areas on each segment. No/Yes. N40. Hindwing-pad very small (Fig. 48). No/Yes. N41. Tarsal arolia. Very reduced or apparently absent/triangular (Figs 24, 25) or disc-like (Fig. 26). N42. Thoracic and abdominal dorsal sclerites with large 'perforations'. No/Yes. N43. Thoracic and abdominal dorsal surfaces with cuticular processes (Fig. 144). No/Yes. N44. Abdominal segments laterally bulging or serrate. No. Margin evenly shaped/Lateral bulges (Fig.27)/Serrate (Fig. 28). N45. Apical margin of abdomen with 'tooth-like' processes. No/Yes (plus the two following clarifyingcharacters). N46. No medial 'tooth' (Figs 120, 122). No/Yes. N47. Medial 'tooth' present (Fig. 125). No/Yes. N48. Anal pore-field present (in any form). No/Yes (plus 6 following clarifying characters). N49. Circum-anal ring with two additional rings placed laterally to it (Fig. 121). No/Yes. N50 Two rings each lateral to the anus, no circum-anal ring (Fig. 150). No/Yes. N51. Four rings, no circum-anal ring (Fig. 120). No/Yes. N52. Outer circum-anal ring broken at two or more places. No/Yes, but remaining in form of acircum-anal ring (Fig. 97)/Bands dispersed. Not forming a circum-anal ring. A small circum-analring may remain which is assumed to derive from an inner ring (Fig. 155)/Bands dispersed andbroken. Not forming a circum-anal ring. A small circum-anal ring may remain which is assumed toderive from an inner ring (Fig. 154). N53. As N52 but describing absence of small circum-anal ring. Present/ Absent. N54. Outer circum-anal ring broken into single pores or small groups of pores. No/Yes, pores or groupsof pores in the form of rings (Fig. 117)/Yes, pores or groups of pores dispersed (Fig. 129). Characters of sclerite fusion N55. Mesothorax and metathorax dorsal surface. Numerous very small sclerites. No/Yes. Three following characters (N56-N58): Tibio-tarsal fusion of each leg. Two tarsal segments free, articulate (Fig. 55)/Two tarsal segmentsfree, not articulate (Fig. 52)/One tarsal segment free, (segment II) not articulate (Fig. 51)/No tarsalsegment free. N56. Foreleg (as above). N57. Midleg (as above). N58. Hindleg (as above). N59. Abdomen dorsal surface membraneous. No/Yes. Chaetotaxy characters N60. Head dorsal capitate setae present. No/Yes. N61 . Antenna segment I inner apical angle capitate seta present. No/Yes.N62. Abdomen ventral capitate setae present. No/Yes.N63. Rod setae on body surface present. No/Yes. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 209 Nine following characters (N64-N72). Clavate setae present. No/Yes. N64. Head margin clavate setae. N65. Head dorsal clavate setae. N66. Antenna segment I inner apical angle clavate seta. N67. Thorax dorsal clavate setae. N68. Forewing-pad dorsal clavate setae. N69. Forewing-pad margin clavate setae. N70. Hindwing-pad margin clavate setae. N71 . Abdomen dorsal clavate setae . N72. Abdomen margin clavate setae. Six following characters (N73-N78). Lanceolate setae present. No/Yes. N73. Head margin lanceolate setae. N74. Head dorsal lanceolate setae. N75 . Antenna segment II with lanceolate setae . N76. Thorax dorsal lanceolate setae. N77. Forewing-pad dorsal lanceolate setae. N78. Abdomen dorsal lanceolate setae. N79. Lanceolate setae present and placed on tall tubercles (Fig. 171). No/Yes. N80. Antenna with one row of sectasetae located on the opposite margin of the antenna to the rhinaria(Fig. 40). No/Yes. N81. Antenna with more than one row of sectasetae (Fig. 45). No/Yes. N82. Antenna with one row of sectasetae located on the same margin as the rhinaria. No/Yes. N83. Tibia with stout setae (Fig. 51). No/Yes. N84. Abdomen margin sectasetae present and based on large clustered tubercles (Fig. 114). No/Yes. N85. Scales present on body margin (Fig. 36). No/Yes. Three clarifying characters of N34.N86 . Sectasetae (of N34 type) tubular . No/Yes .N87. Lanceolate setae (of N34 type) present. No/Yes.N88. Sectasetae (of N34 type) pointed . No /Yes . Table 6 Groups of species identical with the selected nymphal characters. The species chosen to representthe group is named at the top of each list. 1. Aacanthocnema casuarinae 6. Calophya triozomima Ceropsylla matorelli Calophya dubia 2. Acizzia acaciae 1. Ceanothia ceanothi Acizzia acaciaeb alley anae Euphalerus tantillus Psylla phoradendrae Psylla simlae 3. Agonoscenasp. (C). 8. Colophorina cassiae Agonoscena sp. (B). Epipsylla sp. (B). Strophingia cinereae 9. Colopsceniasp. 4. Apsylla cistellata Craspedolepta minuta Mastigimas cedrelae C. minutisimma M. sp. (B) Diaphorina cardiae Mesohomotoma tessmanni D. chobauti Tenaphalara malay ensis 10. Creiissp. 5. Calophya nigripennis Livia coloradensis Calophya flavida L. maculipennis 210 I. M. WHITE AND I. D. HODKINSON 11. Ctenarytaina eucalypti Craspedolepta subpunctata 12. Diaphorina citri Craspedolepta constrictaDiaphorina florea 13 . Diaphorina putonii Diaphorina clutiae D. punctulata 14. Euceropsylla russoi Ciriacremum capenseEuceropsylla minuticona E. sp. 1 5 . Euphalerus jugo venosus E. rugipennisE. vermiculosusPsylla betulaenanaeP. carpinicolaP. floccosaP. galeaformisP. striataP. trimaculata 16. Glycaspis baileyi Cardiaspina albitexturaC. sp. C. squamulaGlycaspis spp. 17. Gyropsylla ilicis Gyropsylla spegazziniana 18. Insnesia glabruscuta Mitrapsylla deserata 19. Mycopsyllafici Macrohomotoma gladiatumMycopsylla gardenensis 20. Neophyllura arctostaphyli Neophyllura arbuti 21 . Neopsyllia erythrinae Euglyptoneura sp.Neopsyllia sp.Platycorypha princeps 22. Pachypsylla venusta Pachypsylla spp.Tetragonocephala sp. 23. Paracarsidara gigantea Bharatiana octospinosaMastigimas sp. (A)Mesohomotoma hibisciParacarsidara spp.Tenaphalara acutipennis 24. Paratrioza cocker elli Paratrioza arbolensisP. maculipennisTrioza curvatinervisT. minutaT. salicivora 25. Paurocephala gossypii Paurocephala urenae 26. Pennavena fabulosa Craspedolepta furcataC. nervosa 27. Psylla alba Psylla sinuata 28. Psylla alni Psylla buxi Spanioneura fonscolombii 29. Psylla brevistigmata Psylla minutaP. parallela 30. Psylla brunneipennis Psylla coryliP. hirsutaP. moscovitaPurshivora chelifera 31. Psylla mail Euglyptoneura robustaPexopsylla cercocarpiPsylla ribesiae 32. Psylla media Arytainilla hakani 33. Psylla minor Psylla magnicauda 34. Psylla nigrita Psylla saliceti 35. Psylla pulchra Psylla hamata 36. Psylla pyricola Psylla myrtilliP. visci 37. Psylla pyrisuga Arytainilla cytisiPsylla melanoneura 38. Psyllopsis fraxinicola Aphalaroida pithecolobiaPsyllopsis fraxini 39. Spondyliaspissp. Cardiaspina densitexta 40. Strophingia ericae Aphalara curta A. nubifera A. polygoni A. simila Craspedolepta augustipennis C. artemisiae C. sonchi C. suaedae C. vancouverensis C. veaziei Tainarys schini 41. Swezeyana elongagena Trioza palmicola 42. Trichocherm.es walkeri Trioza phoradendrae 43. Trioza albifrons Paratrioza lavateraeTrioza beameriT. quadripunctata 44. Trioza albiventris Trioza atkasookensisT. crithmi NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 211 45. Trioza chenopodii Trioza litseae 46. Trioza diospyri Trioza bakeriT. frontalis 47. Trioza incidata Trioza tripunctata 48. Trioza marginepunctata Trioza vitiensisT. sp. The following species have unique descriptions:Acizzia hakeaeA. russellaeA. uncatoidesAgonoscena sp. (A)Amorphicola amorphaeAnomoneura moriAphalara exilisA. monticolaA. persicariaA. rumicisAphalaroida inermisArepuna sp.Arytaina genistaeArytainilla spartiicolaA. spartiophilaCalophya calif ornicaC. rhoisC. schiniC. rotundipennisC. sp. Camarotoscena spp.Ceanothia aculeataCeropsylla sideroxyliC. sp. Ciriacremum capeneriC. harteni C. julbernardioidesCraspedolepta nebulosaCrastina linavuoriiCraw for da triopsyllinaDiaphorina albomaculata D. solani Diclidophlebia eastopiEgeirotrioza spp.Epicarsa sp.Epipsylla sp. (A)Eucalyptolyma sp.Euceropsylla cayeyensisEuglyptoneura fuscipennisEuphalerus gallicolus E. nidifexE. sp. (A).E. sp. (B).E. sp. (C).E. sp. (D).Euphyllura spp.Floria variegataFreysuila sp.Heteropsylla spp.Hevaheva swezeyiHomotoma spp. Isogonoceraia divergipennis Kuwayama pisonia Leptynoptera sulfurea Leurolophus vittatus Leuronota michoacana Livia crefeldensis L. juncorum L. vernalis Macrohomotoma striata Microceropsylla sp. Moraniella calodendri Neolithus sp. Neophyllura bicolor Paraphalaroida fremontiae Pauropsylla spp. Pelmatobrachia sp. Phacopteron lentiginosum Phellopsylla sp. Phytolyma spp. Protyora sterculiae Pseudoeriopsylla nyasae Pseudophacopteron spp. Psylla albagena P. ambigua P. americana P. annulata P. foersteri P. magna P. negundinis P. palmeni P. peregrina P. pruni P. pulchella P. pyri P. rhamnicola P. rhododendri P. stricklandi P. sorbi P. subspiculata P. ulmi Purshivora pubescens Retroacizzia antennata Rhinocola aceris Synoza spp. Tenaphalara sp. Togepsylla matsumurana Trigonon longicornis Trioza alacris T. aylmeriae T. anceps T. cinnamomi 212 I. M. WHITE AND I. D. HODKINSON T. erytreae T. remota T. falcata T. urticae T. hirsuta T. vitreoradiata T. lobata Triozamia lamborni T. magnoliae Triozoida silvestris T. nigricornis indet. (A). T. obsoleta indet. (B). T. obtusa indet. (C). T. panacis Phenetic analysis of nymphs The selected characters are used for these analyses of 182 selected species and resemblance wasmeasured using the mean character difference (Cain & Harrison, 1958). Minimum spanning network (MSN) The MSN (Fig. 178) was constructed using an algorithm given by Farris (1970). In a MSN a set oft taxa are joined by the shortest possible set of linkages (numbering t - 1) and it indicates whichspecies are phenetically most similar to each other. The main features of the MSN relative to theBecker-Migdisova (1973) families are as follows. 1. The following genera of Psyllidae form one group (Fig. 178a): Acizzia, Amorphicola,Anomoneura, Arepuna, Arytaina, Arytainilla, Ceanothia, Ciriacremum, Colophorina, Eucer-opsylla, Euglyptoneura, Floria, Freysuila, Heteropsylla, Insnesia, Isogonoceraia, Mitrapsylla,Neopsyllia, Pexopsylla, Platycorypha, Psylla, Purshivora, Retroacizzia, Spanioneura and Trigo-non plus Epipsylla sp. (B) and many Euphalerus spp. Genera of Psyllidae not included in thisgroup are: Diaphorina, Pennavena and Psyllopsis plus Epipsylla sp. (A). , Euphalerus gallicolus,E. nidifex and E. sp. (A). 2. The following genera of Triozidae form one group (Fig. 178c): Aacanthocnema, Ceropsylla,Crawforda, Hevaheva, Kuwayama, Paratrioza, Swezeyana, Trichochemes, Trioza (minus 2spp.) and Triozoida. Genera of Triozidae not included in this group are: Egeirotrioza,Leuronota, Neolithus and Triozamia plus Trioza alacris and T. hirsuta. 3. All members of the Liviidae form one group; however, two are included with Creiis sp.(Fig. 178b). 4. The following genera of Aphalaridae form one group (Fig. 178b): Agonoscena, Aphalara,Colposcenia, Craspedolepta, Crastina, Euphyllura (minus E. phillyreae), Leurolophus,Neophyllura, Rhinocola, Strophingia and Tainarys plus Aphalaroida pithecolobia and Camar-otoscena speciosa. Genera of Aphalaridae not included in this group are: Apsylla, Gyropsylla,Moraniella, Paraphalaroida, Paurocephala and Phytolyma plus Aphalaroida inermis, Camar-otoscena unicolor and Euphyllura phillyreae . 5. The following genera of Spondyliaspididae form one group (Fig. 178c): Cardiaspina,Eucalyptolyma, Glycaspis and Spondyliaspis . 6. A second group of Spondyliaspididae consists of the genera (Fig. 178b) Creiis, Pachypsylla,Phellopsylla and Tetragonocephala. 1. The Carsidaridae failed to form as one major group. In the MSN many genera and species are not placed with the majority of their family. The maindiscordant features are as follows. 1. Aphalaroida inermis and Paraphalaroida (Aphalaridae), and Diclidophlebia (Carsidaridae)are placed with the Psyllidae (Fig. 178a) because they share the feature of a petiolate tarsalarolium. 2. Calophya schini, Calophya rotundipennis and Microceropsylla (Carsidaridae) are placedwith the Triozidae (Fig. 178c). These species have a well-developed humeral lobe of the NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 213 fore-wing pad like most Triozidae. Calophya schini is surrounded by pointed sectasetae likeCrawforda to which it is linked, whereas Calophya rotundipennis and Microceropsylla lacksectasetae like Kuwayama pisonia to which they link. 3. Creiis (Spondyliaspididae) is identical to some species of Livia (Liviidae), as defined by theselected characters. This is because these species have almost entirely zero character states.Euphalerus sp. (A) (Psyllidae) also links to Livia. 4. Diaphorina, Pennavena and Psyllcpsis (Psyllidae), Ctenarytaina (Spondyliaspididae) plusEpicarsa, Homotoma ficus, Phacopteron and Pseudophacopteron floccosa (Carsidaridae) areplaced with most Aphalaridae (Fig. 178b). Group 4 is a collection of species with lanceolatesetae. 5. Euphalerus nidifex (Psyllidae) is linked with Glycaspis (Spondyliaspididae). 6. Euphalerus gallicolus (Psyllidae) is linked with Creiis (Spondyliaspididae). 7. Most taxa with pointed sectasetae form one group (Fig. 178b): most Calophya spp.,Camarotoscena unicolor, most Egeirotrioza spp., Homotoma indica, Leuronota, Moraniella,Paurocephala, Synoza floccosa and Trioza alacris. The remaining genera and species lack specialised chaetotaxy. They are placed near to thecentre of the minimum spanning network and the distinct groups branch from them. Average linkage phenogram (AL) Further insight into the resemblances between nymphs is provided by a hierarchical representa-tion of data as provided by an average linkage phenogram. An average linkage phenogram was constructed using the 'weighted pair group method witharithmetic averages' (Fig. 179) which was computed by the 'JOIN' algorithm of Hartigan (1975). Most of the groups formed in the minimum spanning network were also recognised by averagelinkage. Three major clusters were formed (Figs 179b-d). Cluster 1 (Fig. 179b). This includes members of the family Psyllidae which have capitate setae:Acizzia hakeae, A. uncatoides, Amorphicola, Arytaina, Arytainilla, Ceanothia, Ciriacremum,Euceropsylla, Euphalerus tantillus, E. sp. (B)., E. sp. (C)., Fiona, Freysuila, Heteropsylla,Insnesia, Isogonoceraia, Mitrapsylla, Psylla (minus subgen. Asphagidella, subgen. Psylla, P.annulata, P. mali, P. phoradendrae & P. ribesiae), Purshivora and Trigonon. The remainingPsyllidae are in cluster 3 (except Retroacizzia). Cluster 2 (Fig. 179c). This contains taxa with sectasetae. It also includes Retroacizzia:Camarotoscena unicolor, Moraniella, Paraphalaroida and Paurocephala (Aphalaridae),Calophya, Diclidophlebia, Homotoma, Leptynoptera, Microceropsylla, Pauropsylla depressa,P. trichaeta, Synoza floccosa and Togepsylla (Carsidaridae), Retroacizzia (Psyllidae), Aacan-thocnema, Ceropsylla, Crawforda, Egeirotrioza, Hevaheva, Kuwayama, Leuronota, Neolithus,Paratrioza, Swezeyana, Trichochermes, Trioza (minus T. hirsuta), Triozoidaandindet. sp. (A).(Triozidae). Cluster 3 (Fig. 179d,e). This includes taxa with lanceolate setae plus most taxa which lackcapitate setae and sectasetae: Agonoscena, Aphalara, Aphalaroida, Apsylla, Camarotoscenaspeciosa, Colposcenia, Craspedolepta, Crastina, Euphyllura, Gyropsylla, Leurolophus,Neophyllura, Phytolyma, Rhinocola, Strophingia and Tainarys (Aphalaridae), Bharatiana,Epicarsa, Macrohomotoma, Mastigimas, Mesohomotoma, Mycopsylla, Paracarsidara, Paurop-sylla beesoni, Pelmatobrachia, Phacopteron, Protyora, Pseudoeriopsylla, Pseudophacopteron,Synoza sp. and Tenaphalara (Carsidaridae), Livia (Liviidae), Acizzia acaciae, A. aca-ciaebaileyanae, A. russellae, Anomoneura, Arepuna, Colophorina, Diaphorina, Epipsylla,Euglyptoneura, Euphalerus gallicolus, E. jugovenosus, E. nidifex, E. rugipennis, E. vermiculo-sus, E. sp. (A)., E. sp. (D).,Neopsyllia, Pennavena, Pexopsylla, Platycorpha, some Psylla spp.(subgen. Asphagidella, subgen. Psylla, P. annulata, P. mali, P. phoradendrae and P. ribesiae)and Psyllopsis (Psyllidae), Cardiaspina, Creiis, Ctenarytaina, Eucalyptolyma, Glycaspis,Pachypsylla, Phellopsylla, Spondyliaspis and Tetragonocephala (Spondyliaspididae). Cluster 3 (Fig. 179d) is clearly least congruent with the classification of Becker-Migdisova(1973). Capitate setae (N 11 - N 19) and sectasetae (N 26- N 33) are described by nine and eight 214 I. M. WHITE AND I. D. HODKINSON selected characters respectively. However, lanceolate setae (N 23-N 25) are only described bythree selected characters and therefore contribute less weight to the classification than eithercapitate setae or sectasetae. Hence most taxa which lack any of these three major setal types arephenetically closer to taxa with lanceolate setae than to taxa with capitate setae or sectasetae andcluster 3 is produced. Detailed analysis of the characters which are responsible for groups is more easily performedusing principal component analysis. Principal component analysis (PC) Principal component analysis, like other forms of ordination, aims to transform a data matrix,whose variance is in many dimensions, into a matrix with most of the variance explained by a P-BAEOPE P-HEPATO (8) P-HEPATO (DP-HEPATO (14) P-HEPATO (17) EUCEROPS d) P-HEPATO (27) ARYTAINA P-HEPATO (9) P-HEPATO (3) EUCEROPS (3) CIRIACRE (4) CIRIACRE d)FREYSUIL-^^ JACIZZIA (5) P-HEPATO d5) AMORPHIC CEANOTHI (2) HETEROPS d) P-HEPATO (6)' \ PURSHIVO (2) P-HEPATO (22) P-HEPATO (24) CEANOTHI (1) P-THAMNO (5) P-CACOPS (4) HETEROPS (2) P-CACOPS (5) EUPHALER (8)^ EUPHALER (9) ACIZZIA (3) P-HEPATO (25) TRIGONON DICLIDOP PARAPHAL. FLORIAP-HEPATO dO) ISOGONOC INSNESIA P-CACOPS (3) P-THAMNO (4)P-HEPATO dl) P-THAMNO (2)/ IILLA (3) ^P-THAMNO (3) ARYNILLAP-CACOPS (2) P-THAMNO (6) P-HEPATO (2) ARYNILLA (4) CIRIACRE (3) ACIZZIA (4) P-HEPATO (16) -ACIZZIA (D APHROIDA (1) AREPUNA \ P-PSYLLA (1) -EUPHALER (2) EUGLYPTO (D P-CACOPS (D NEOPSYLL (D P-P-P P-P p P-P-Pp p p-p V A-A-P- P-P-P-P P-P / P P P Fig 178a Part of a minimum spanning network of 182 selected species with 34 nymphal characters.Internode lengths are not drawn in proportion to taxonomic distance and for convenience the network isdivided into 3 sections. Continued in Figs 178b-c. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 215 very few dimensions. In this case 34 dimensions (because there are 34 characters) would beneeded to illustrate the spatial relationships of 182 species. However, when principal componentanalysis was carried out almost 60% (58-2%) of the variance was accounted for by just threedimensions, i.e. a three dimensional figure of spatial relationships of the species could illustratemore than half of the variance in the data. Other forms of ordination, such as principalcoordinate analysis, tend to yield very similar results (Boratynski & Davies, 1971) and are notpractical when the number of taxa is large and in excess of the number of characters (Rohlf ,1972). For this analysis data were scaled by ranging and eigenvectors were extracted from correlationand covariance matrices for the first 10 principal components, which accounted for most of thevariance (82-8% when extracted from a correlation matrix and 83-7% from a covariance GYROPSYL (IV^PHYTOLYM (2 )^ p H YTOLYM (D a 'PACHYPSY (5) EUPHYLLU (3) PAUROPSY (DTRIOZAMIUPHALER (D PHELLOPSPSEUDOER T-HETERO (1) LEURONOT SYNOZA (D CALOPHYA (1) CAMAROTO (2) EUPHALER (7) LIVIA (2) CALOPHYA (?) CALOPHYA < 4 >PSEUDOPH (2) LIVIA (5) LIVIA (3) MYCOPSYL (D HOMOTOMA (2) MORANIEL CALOPHYA (9) EGEIROTR (2) PSEUDOPH (D CAMA CRASPEDO (8) CTEN APHALARA (?) STRO ALA ROTO (D PHACOPTE EGEIROTR (3)EGEIROTR (D DIAPHORI (9) DIAPHORI (8) PENNAVEN DIAPHORI (4) 'HIN (2)^ AGONOSCE (3) RHINOCOL APHALARA (2) COLPOSCE HOMOTOMA (1) INDET (B) APHALARA (5) c -S-A C r L s L-L-5 C-A-C C-A-C'P-P ^A-AC DIAPHORI (D PSYLLOPS (2) EUPHYLLU (D NEOPHYLL (3) EUPHYLLU (2) NEOPHYLL (2) LEUROLOP AGONOSCE (D A-A-A Fig. 178b Part of a minimum spanning network of 182 species; continued in Figs 178a and 178c. 216 I. M. WHITE AND I. D. HODKINSON matrix). Only the analysis based upon the covariance matrix is considered further (thisexplained 34-3, 16-8 and 7-1% variance on each of the first three axes). To reduce the number of species to a number which could be easily represented visually anaverage linkage phenogram was generated across PCs I to X, using average distance (White,1980). This produced 68 clusters below phenon level 0-2, which is a convenient number of pointsfor visual representation in three dimensions. The position of one species from each of theseclusters was plotted relative to PCs I to III (Fig. 180). Each species is labelled with the initialletter of the family to which it is assigned by Becker-Migdisova (1973). The species content of each of the three main 'arms' of the trifurcate scatter roughlycorresponds with that of the minimum spanning network (Fig. 178) and the average linkagephenogram (Fig. 179). Again the three main groupings are dominated by the Aphalaridae,Psyllidae and Triozidae. Furthermore, the major incongruences with the classification ofBecker-Migdisova (1973) are the same. For example those Psyllidae and Carsidaridae, whichare placed with the Aphalaridae, have lanceolate setae, e.g. Diaphorina, Psyllopsis and T-MEGATR (2)-^ EP1PSYLL (D PELMATOB PARACARS (2)APSYLLAPHYTOLYM (3) - TENAPHAL (3) - PROTYORA - NEOLITHU EUCALYPT^^EUPHALER (3)^SPONDYLI'^*'^ GLYCASPI (1) PAUROPSY (2) HEVAHEVA PSEUDOPH O) MACR'OHOM (2) SYNOZA (2) SWEZEYAN MICROCER CALOPHYA ( SJ^KUWA^AMA CEROPSYL (2) TR10ZOID. LEPTYNO EGEIROTR (4) CRAW CALOPHYA (6) T ~t AACANTHO CEROPSYL (3) INDET (A) A (\ _ 5 I T I A-C-C-T T-TRIOZA (2) C I PAUROPSY (3) T-HETERO (2) Q__Q J C-C T T T-BACTER (3>^T-MEGATR (3) "j^J- J_ Q 1CT T T T-TRIOZA (9) PARATRIO ( 2 ) T-MEGATR d) T-TRIOZA d) -|- T-TRIOZA (7) C TTRICHOCH. j -p q- i-T-T-T T-BACTER (6) T T T-HETERO (5) T-BACTER (8) T-HETERO (4) T-HETERO (6) J .J ^ J I III I I T T c T-HETERO (3) T-TRIOZA (4) TOGEPSYL J j T-TRIOZA d2) T-HETERO (8) Fig. 178c Part of a minimum spanning network of 182 species; continued in Figs 178a-b. 0-4 0-3 0-2 DISTANCE 01 0-0 Fig. 179a Average linkage phenogram of 182 selected species with 34 nymphal characters; key diagramshowing linkages to Figs 179b-d. CIRIACREMUM HARTENIHETEROPSYLLA TEXANAFREYSUILA SP,HETEROPSYLLA INCISACIRIACREMUM CAPENERIPSYLLA (T. ) MEDIACEANOTHIA CEANOTHICEANOTHIA ACULEATAEUCEROPSYLLA CAYEYENSISC1RIACREMUM JULBERNARDIOIDESEUCEROPSYLLA RUSSOIAMORPHICOLA AMORPHAEPSYLLA ' H -> BREVISTIGMATAPSYLLA HI. ) PALMENIPSYLLA < H - ) ALBAGENAPSYLLA ( H -> SUBSPICULATAPSYLLA (H- > ALBAPSYLLA (H- ) NIGRITAPSYLLA < B - > FOERSTERIARYTAINA GENISTAEPURSHIVORA PUBESCENSPSYLLA ; H - > NEGUNDINISPSYLLA (C. > STRICKLANDIPSYLLA 'H- > MINORPSYLLA ' H - 1 PULCHRAPSYLLA (H -> BRUNNEIPENNISPSYLLA < H - > AMERICANAACIZZIA UNCATOIDESISOGONOCERAIA SP.INSNESIA GLABRUSCUTAPSYLLA PULCHELLAARYTAINILLA SPARTIOPHILAPSYLLA < T - ) MAGNAFLORIA VARIEGATAPSYLLA < c - ^ SORB!PSYLLA (T. ) PRUNIARYTAINILLA SPARTIICOLAPSYLLA (C. ) ULMIPSYLLA ' ) PYRIPSYLLA (T. ) PYRISUGAPSYLLA <T- ) RHAMNICOLAEUPHALERUS SP. < c >EUPHALERUS SP. < B 'TRIGONON LONG I CORN ISPSYLLA ( c - > PEREGRINAPSYLLA < H . ) AMBIGUAACIZZIA HAKEAEPSYLLA (H- ) PYRICOLAPSYLLA lH.) RHODODENDRI 02 00 Fig. 179b Part of a phenogram of 182 species; continued from Fig. 179a. 218 I. M. WHITE AND I. D. HODKINSON 02 0-1 00 DISTANCE Fig. 179c Part of a phenogram of 182 species; continued from Fig. 179a. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 219 -NEOPHYLLURA ARCTOSTAPHYL I-LEUROLOPHUb VITTATUS-AGONOSCENA SP . (A) SYNOZA SP, PSEUDOPHACOPTERON SP. (B)MACROHOMOTOMA STRIATA EUCALYPTOLYMA SP.SPONDYLIASPIS SP.GLYCASPIS BAILEY! EUPHALERUS NIDIFEXPROTYORA STERCULIAEPHYTOLYMA MINUTATENAPHALARA SP . APSYLLA C! STELLA TA-EUPHYLLURA PHILLYREAE -EPICARSA SP,-PAUROPSYLLA BEESONI-PSEUDOERIOPSYLLA NYASAE -INDET (C)-PELMAT03RACHIA SP . -PHYTOLYMA FUSCA-PHYTOLYMA LATA-GYROPSYLLA ILICIS -TRIOZA (M. ) HIRSUTA-PARACARSIDARA GIGANTEA-EPIPSYLLA SP. (A> ] P 3 C ] A J C "I . J A DC AJ ] T] C 02 0-1DISTANCE 00 Fig. 179d Part of a phenogram of 182 species; continued in Fig. 179e and from Fig. 179a. Epicarsa. The Carsidaridae and Aphalaridae placed with Triozidae have sectasetae, e.g.Pauropsylla trichaeta and Paurocephala spp. However, the major advantage of principalcomponent analysis is that it permits the analysis of characters that are responsible for theformation of major groupings. Characters with absolute eigenvector values on PCs I to III of at least the mean eigenvectorvalue (0-17) are listed in Table 7. These are the characters which largely control the placing ofspecies on PCs I to III. On PC I (Table 7) the positive eigenvectors account for sectasetae (other than N 34) and shape(N 7) while the large negative values correspond to the petiolate tarsal arolium and capitatesetae. The present states of those characters are mainly associated with Triozidae (positivevalues) and Psyllidae (negative values). On PC II the positive values are those applying toTriozidae and Psyllidae while the negative ones relate to the Aphalaridae (plus Diaphorini andPsyllopsis). PC III has a very high positive value for anus position, while other characters withpositive values apply to Aphalaridae (plus Diaphorini and Psyllopsis). The remaining negativevalue is a shape character which relates to the Triozidae. 220 I. M. WHITE AND I. D. HODKINSON 01 DISTANCE 00 Fig. 179e Part of a phenogram of 182 species; continued from Fig. 179d. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 221 ^^ 8&$ 0) O C ex c o c S O ^ J3 '5 C-a CX O <UPH Pi t/3 > l-i t/5 Q g c "S> rt , '*5 887 >-.!> O ra 53 u C vc ex co (/5 f . . C -U ^ r" s s> C -O CM O <U 00 C ^ ^ JS ti-i <-| cc o ex g .2 <N JS *j 4) C ^ Ofl"* 3 g_ H 1 -i i; I) -C i_ O CL oo -i- m <_, ^N O JZ t^ eb S H 222 I. M. WHITE AND I. D. HODKINSON Table 7 Absolute eigenvector values in excess of the mean on principalcomponent I, II and III. Character PCI PC II PC III When the eigenvector values (EVs) of the 34 selected characters are plotted in threedimensions they form eight major groups (Fig. 181). 1. Characters N7, N26, N30, N31 and N33 describe the fusion of the head and prothorax andmarginal positions of sectasetae. These characters have positive states for most Triozidae. 2. Characters Nl, N2, N3, N5, N10, N27, N28, N29, and N32 describe the humeral lobeposition, position of wing-pad apices, disc-like tarsal arolium, sclerotization of dorsal surface ofabdomen and dorsal positions of sectasetae. The taxonomic distribution of positive states ofthese characters is similar to those in group 1 . 3. Character N6 refers to the position of the anus. The extreme lack of compatibility of thischaracter with most other characters causes it to be placed alone. All previously recognisedmajor taxa have some species with each state of this character. 4. Characters Nil, N12, N13, N14, N15, N16, N17, N18, N19 and N34 are positions of capitatesetae and positioned abdominal margin lanceolate setae/sectasetae. These are exclusivelyfeatures of the Psyllidae. As with the sectasetae the dorsal surface setal positions fall nearer thezero eigenvector values than the marginal setal positions. 5. Character N4 described the presence or absence of a petiole on the tarsal arolium. Thisfeature is present in most Psyllidae as well as a few other groups. 6. Characters N20, N21 and N22 refer to the pair of tarsal capitate setae which are bestdeveloped in the Spondyliaspidinae. 7. Characters N8, N23, N24 and N25 refer to the large thoracic sclerites and marginal lanceolatesetae typical of the Aphalaridae. Character groups 1 , 2 and 3, groups 4 and 5, plus groups 6 and 7 each form one of three 'arms'of a trifurcate scatter (Fig. 181). Group 8 however, consisting only of characters N9 (free dorsalsclerites on abdomen), does not fit this trifurcate scatter because it is a feature of bothAphalaridae and Psyllidae. Overall, the scatter of characters shown in Fig. 181 mirrors the trifurcate scatter of species(Fig. 180). This is because only the data matrix is required to transform the charactereigenvector values to principal component values. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 223 . a _ _ J - -V 224 I. M. WHITE AND I. D. HODKINSON Incorporation of adult charactersSelection of characters Examination of the adults of the species studied was beyond the scope of this work. However, itwas desirable to study adult characters as the interpretation of possible cladistic relationshipsrequires the incorporation of characters controlled by different selection pressures. Charactersselected from two rather than one life cycle stages are more likely to meet this criterion.Furthermore, such a study should provide insight into the underlying causes of the partialcongruence between the nymphal phenetic relationships and the classification of Becker-Migdisova (1973), which was based almost exclusively upon adult data. Twenty-seven adult characters were coded largely from the literature (Tables 8, 9). Majorsources of information included those of Crawford (1914), Eastop (1958), Heslop-Harrison(1959), Hodkinson & White (19796), Klimaszewski (1964), Loginova (19640, 1972), Mathur(1975), Tuthill (1943, 1959, 19640), Tuthill & Taylor (1955) and Zimmerman (1948). Whendescriptions are coded in this way several problems occur (Young & Watson, 1970). Thegreatest problem was the lack of consistency in the conventional approach to description whichoften made it impossible to distinguish between a genuine absence and mere failure to record thestate of the character concerned. The least recorded characters are marked by an asterisk in thecharacter listings (Tables 8, 9). In several cases the states of characters were deduced fromfamily or tribal descriptions. The 27 characters were coded identically for all species within each genus except forEuphalerus and Pauropsylla. These genera are variable intra-generically and the species weregrouped as follows.Euphalerus spp. received four different group descriptions: a. E. nidifex, E. tantillus, E. sp. (A), E. sp. (B) and E. sp. (D). b. E. gallicolus c. E. jugovenosus, E. rugipennis and E. vermiculosus d. E. sp. (C) Pauropsylla spp. received two different group descriptions: a. P. beesoni b. P. depressa and P. trichaeta The 27 adult characters were combined with the original 88 nymphal characters (Tables 4, 5)and the ordered multistate characters were receded into additive two-state code to give 33 adultand 120 nymphal characters. The SUMRAT information statistic was applied and 18 adult and40 nymphal two-state characters were selected. These were then combined to form 14 adult(Table 8) and 30 nymphal (Table 10) multistate and two-state characters. Several groups of species were identical with respect to the selected adult plus nymphalcharacters. Forty-seven species groups were formed and one representative species was chosenfrom each (Table 11). A further 161 species remained distinct (Table 11) making a total of 208'taxa' available for further analysis. Cluster analysis of combined adult and nymphal data A MSN was generated across the data (Fig. 182). It was not possible to place an average linkage(AL) phenogram across the combined data because the required computing time was in excessof that available. Instead, as the first 10 principal components (PCs), extracted from a betweencharacter covariance matrix, explain most of the variance (79%), an AL phenogram calculatedacross them should approximate an AL phenogram using the untreated combined data. Such anAL phenogram using average distance is given in Fig. 183.Five major clusters are formed by the AL. 1 (Fig. 183b). The members of the family Psyllidae whose nymphs have capitate setae. 2 (Fig. 183c). The members of the families Triozidae and Carsidaridae which are typified byadult 'triozine' wing venation (Character A6) and nymphal sectasetae. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) TRIOZAMI 225 HEPATO (15) P-HEF P-THAMNO (7) PURSHIVO (!) CEANOTHI (1) ACIZZIA (5).^^HETEROPS (2)-** CEANOTHI (2) PURSHIVO (2) HETEROPS (1) CIRIACRE (D CIRIACRE (2) EUCEROPS (3) CIRIACRE (*) EUCEROPS (D ARYNILLA (2). ARYTAINA ARYNILLA (1) ARYNILLA (4) ARYNILLA ( 3 ) FLOR1A Fig. 182a Part of a minimum spanning network of 208 selected species with 14 adult and 30 nymphalcharacters. Internode lengths are not drawn in proportion to taxonomic distance; for convenience thenetwork is divided into three sections. Continued in Figs 182b-c. 226 I. M. WHITE AND I. D. HODKINSON PARACARS (2)i PROTYORA MESOHOMO (D TENEPHAL (D TENAPHAL (3) MASTIGIM (1) APSYLLA BHARAT1A CALOPHYA (5) CALOPHYA (7) CALOPHYA (9) PARAPHAL CALOPHYA (2 DICLIDOP MORANIEb CALOPHYA (3)^CALOPHYA (6)CALOPHYA (1) HOMOTOMA (2)_ HOMOTOMA (1) cp-cc-c-cc A-C-C c-c A-t C ~ C ? rc C A-C-C P-P-P-C-L c fcp^ ^-A-A-A-A-C ~T T~ AA A-^A-A |c K A-C j-AP-A-A DIAPHORI (8) AREPUNA^ PSEUDOPH (2) LIVIA (5)- DIAPHORI (2) DIAPHORI (9) a STROPHIN (2) CRAST I NA COLPOSCE PHIN (2) AGON! APHALARA ( ? )y .APHALARA (6). APHALARA (2) APHALARA (3) AGONOSCE (3) LEUROLOP NEOPHYLL (2) NEOPHYLL ( 1 > EPICARSA AGONOSCE (2) EUPHYLLU (3) CRASPEDO (6) CAMAROTO ( 1 ) CAMAROTO (2 )^ PAUROCEP (D CRASPEDO (1) PSEUDOPH Cl) APHALARA (5) CRASPEDO (12) CRASPEDO (8) PENNAVEN . CRASPEDQ (5) CRASPEDO (4) Fig. 182b Part of a minimum spanning network of 208 species; continued in Figs 182a and 182c. 3 (Fig. 183d). The members of this cluster are typified by numerous adult metatibial spines,usually with a lack of pronounced adult genal cones and the presence of nymphal lanceolatesetae. It includes many Aphalaridae, all Liviidae, some Carsidaridae (many Homotominae andPhacopterinae) and a few Psyllidae (Diaphorini and Psyllopsis). 4 (Fig. 183e). This cluster contains species whose nymphs have pointed sectasetae together withthose lacking capitate setae, lanceolate setae and truncate sectasetae. The incorporated taxa areAphalaridae (e.g. Paraphalaroida) and Carsidaridae (e.g. Calophyd) whose nymphs have NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) SYNOZA (D SYNOZA (2) NEOL1THU " PAUROPSY (1) - PAUROPSY (2) EGEIROTR (D T-MEGATR (4) SMEZEYAN EGEIROTR (3) TRIOZOID CEROPSYL (2) TRIOZA (D EGEIROTR (2) T-TRIOZA (2) AACANTHO- T-BACTER (9) T-BACTER (3) -T-TRIOZA (7) T-TR 1 OZA (8) T-MEGATR (2) PAUROPSY (3) CEROPSYL. (D CEROPSYL (3) T-TRIOZA (5) T-BACTER (2) T-TR OZA (10) TRICHOCH- CTER (6) T-TR 1 OZA (3) T-HETERO (D T-BA T-HETERO (6) - T-HETERO () T-BACTER (8) PARATRIO (3)i T-TR PARATRIO (2) PARATRIO (D C-C-T-*b LEURONOT C~~C T T-CTT T T-HETERO (3) OZA () T-HETERO (5) T T TT T-TR OZA (12) T-HETERO (8) TT-- 227 Fig. 182c Part of a minimum spanning network of 208 species; continued in Figs 182a-b. pointed sectasetae, Psyllidae lacking capitate setae (e.g. Acizzia acaciae) and species with nospecialised setae belonging to the families Carsidaridae (e.g. Tenaphalard) and Spondyliaspidi-dae (e.g. Glycaspis). The genera Ctenarytaina, Eucalyptolyma and Phellopsylla (all Spondylias-pididae are exceptional in that their nymphs do have lanceolate setae). 5 (Fig. 183a) . This cluster is the genus Euglyptoneura which is a long distance from other speciesand hence are not included in any larger hierarchical cluster. However, in the MSN this genus isjoined to Psylla. In general, these clusters are very similar to those formed by analysis of nymphal data only. The incorporation of adult data now enables observations to be made of the causes ofcongruence and incongruence between adult and nymphal resemblances. A between character correlation matrix was generated across the selected adult plus nymphaldata. The high correlations (r greater than 0-5) were as follows. 1 . Triozidae attributes of adults and nymphs correlate (A3, A5, A6 and A12 with Nl , N7, N10,N26, N30, N31 and N33), e.g. the adult wing venation and nymphal sectasetae. 228 I. M. WHITE AND I. D. HODKINSON 2. Aphalaridae attributes of adults and nymphs correlate (All with N8, N23, N24 and N25), i.e.large numbers of adult metatibial spines with, for example, nymphal lanceolate setae. 3. Psyllinae attributes of adults and nymphs correlate (A4 with N4, N16, N17 and N19), i.e. adiagonal suture between the epimeron and episternum of the adult with, on the nymph, apetiolate tarsal arolium and capitate setae. These correlated sets of characters, whose positive states roughly define currently recognisedhigher taxa, are the characters which weight the analyses towards complete congruence with theexisting classification. One observed form of incongruence is that groups of species that are regarded as genera onthe basis of adult morphology often fail to cluster when nymphal data are analysed. In theminimum spanning network (MSN) (Fig. 182) 11 genera failed to cluster. These were Acizzia,Aphalaroida, Ceropsylla, Ciriacremum, Craspedolepta, Egeirotrioza, Euphalerus, Paratrioza,Pauropsylla, Pseudophacopteron and Trioza. A total of 21 other genera, of which more than onespecies was examined, are clustered. It is tentatively concluded that nymphal dissimilarity withina genus does not usually outweigh adult similarity. Cladistic analysisGround plan construction Prior to carrying out a cladistic analysis a ground plan was formed, i.e. a description of thehypothetical ancestor to present day Psylloidea. Various 'directional arguments' have beenproposed for deducing which character states are primitive and therefore belong to the groundplan, and the methods are reviewed by de Jong (1980) and Arnold (1981). The favouredtechnique is known as OUT-GROUP COMPARISON, i.e. a character state that is not restricted to asingle monophyletic group is likely to be ancestral. To apply the out-group criterion a previouslysuggested phylogeny is needed. Watrous & Wheeler (1981) noted that there could be circularityinvolved in forming monophyletic groups from directional arguments based upon monophyleticgroups. Instead a previous classification can be used and in this study directional arguments werebased on the results of the phenetic analyses presented earlier. For example, pointed sectasetaeare present in many clusters in any given phenogram and so they appear to be an ancestral c bde -EUGLYPTONEURA ROBUSTA-EUGLYPTONEURA SP .-EUGLYPTONFURA FUSCIPENNIS 0-6 0-5 0-4 0-3 02 01 00 Fig. 183a Average linkage phenogram of 208 selected species. Distances were calculated from the first 10principal components derived from 14 adult and 30 nymphal characters; key diagram showing linkages toFigs 183d-e. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 229 feature. Conversely, truncate sectasetae, capitate setae and lanceolate setae each occur in asingle large cluster and are therefore assumed to be derived features.A summary of the ground plan is as follows. ADULT Head rounded. Genal cones absent. Anteoccipital lobes present. Antenna with narrow flagellarsegments and rhinaria on segments III, IV, V, VI, VII, VIII, IX. Suture between epimeron and episternum vertical. Forewing: coriaceous, rhomboidal in shape, costalbreak present, pterostigma present, nodal line present and veins Cu+M with a common stalk after thebranching of R from the R+M+Cu stalk. Hind leg: meracanthus well developed, genual spine present,apex of tibia with a crown of many (c. 12) spines and apex of tarsal segment I with two spines. Proctiger of male bipartite. EUPHALERUS SP. (OEUPHALERUS SP. (B)FREYSUILA SP.EUPHALERUS TANTILLUSAMORPHICOLA AMORPHAEFLORIA VARIEGATA.ARYTAINILLA SPARTIOPHILA.ARYTAINILLA SPARTIICCLA.ARYTAINILLA CYTISIARYTAINILLA HAKANI-ARYTAINA GENISTAEACIZZiA UNCATOIDESEUCERCPSYLLA P.USSOIEUCEROPSYLLA CAYEYENSISHETERCPSYLLA INCISACIRIACREMUM CAPENSE IRIACREMUM JULEERNARDI 01 DESCIRIACREMUM CAPENERITRIGONON LONGICORNISPSYLLA < T - i PYRISUGA ) RHAMNICOLA) PRUNI) PEREGRINAi RHODODENDRI PYRICOLA PSYLLA ( H - ' PALMENIPSYLLA ( ) ALBAGENAPURSHIVORA PUBESCENSPURSHIVORA CHELIFERAPSYLLA (T. ) SIMLAEPSYLLA (T. ) MEDIAPSYLLA ' H. ) AMERICANAPSYLLA ;T - > MAGNAPSYLLA (B.) FCERSTERI CC. ) STRICKLANDI(H. ) NIGRITA ^ MAGMICAUDA ) BRUNNEIPENNIS) NEGUNDINIS . ) PYRI PSYLLAPSYLLA PSYLLAPSYLLAPSYLLAPSYLLA PSYLLA (H. ) SUBSPICULATA (H. ) ALBA CEANCTHIA CEANOTHIEANOTHIA ACULEATAISOGONOCERAIA SP.MITRAPSYLLA DESERATAINSNESIA GLABRUSCUTAHETEROPSYLLA TEXANACIRIACREML'M HARTENIACIZZIA HAKEAE 0-3 0-1 0-0 0-2DISTANCEFig. 183b Part of a phenogram of 208 species; continued from Fig. 183a. 230 I. M. WHITE AND I. D. HODKINSON TOGEPSYLLA MATSUMURANAHOMOTOMA FICUSHOMOTOMA INDICAPAUROPSYLLA BEESONISYNOZA FLOCCOSASYNOZA SP. EGEIROTRIOZA VERUCIFICAEGEIROTRIOZA SP, (A)EGEIROTRIOZA CEARDITRIOZA ( ) HIRSUTATRIOZA ANCEPSLEURONOTA MICHOACANA PAUROPSYLLA DEPRESSA NEOLITHUS SP.TRIOZOIDA SILVESTRISEGEIROTRIOZA SP, (B)TRIOZA ( M - ) PALM I COLA LEPTYNOPTERA SULFUREA SWEZEYANA ELONGAGENACRAWFORDA TRIOPSYLLINAKUWAYAMA PI SON I AHEVAHEVA SWEZEYICEROPSYLLA SIDEROXYLITRIOZA < > MAGNOLIAETRIOZA (H. ) ALACRISTRIOZA <H. ) REMOTATRIOZA (B. ) OBTUSATRIOZA (H. ) LOBATATRIOZA ( ) OBSOLETATRIOZA (T. ) FALCATATRIOZA (T. ) ERYTREAETRIOZA (T.) PHORADENDRAETRIOZA (B.) CURVATINERVISTRIOZA ( T - > BAKER ITRIOZA < T - ' URT1CAETRIOZA (T. ) VITREORADIATATRIOZA <B- ) AYLMERIAETR!OZA 'H. ) CHENCPODI ITRIOZA (T. ) ALBIFRONSPARATRIOZA LAVATERAETRICHOCHERMES WALKER IPARATRIOZA COCKERELLIPARATRIOZA ARBOLENSISTRIOZA (B.) TRIPUNCTATATRIOZA (B. ) NIGRICORNISTRIOZA (T.) HARGINEPUNCTATATRIOZA CH. ) ALBIVENTRISCEROPSYLLA SP . T D C ]' ] C PAUROPSYLLA TRICHAETA 05 0-3 02 DISTANCE 01 0-0 Fig. 183c Part of a phenogram of 208 species; continued from Fig. 183a. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 231 CRASTINA LINAVUORI ICOLPOSCENIA SP,APHALARA PERSICARIACRASPEDOLEPTA NEBULOSACRASPEDOLEPTA SUBPUNCTATACRASPEDOLEPTA FURCATACRASPEDOLEPTA CONSTRICTAAPHALARA MONTICOLACRASPEDOLEPTA MINUTACRASPEDOLEPTA ANGUSTI PENN I SAPHALARA RUMICISAPHALARA POLYGONIAPHALARA EXILIS -MYCOPSYLLA FICI-MACROHOMOTOMA STRIATA-MACROHOMOTOMA GLADIATUM-PSEUDOERIOPS/LLA NYASAE -PHYTOLY.'IA LATA-PHYTOLYMA MINUTA-PHYTOLYMA FUSCA-GYROPSYLLA ILICIS -PSEUDOPHACOPTERON SP, (B)-PSEUDOPHACOPTERON SP . (A) -PENNAVENA FABULCSA -LIVIA VERNALIS-LIVIA JUNCORUM-LIVIA CREFELDENSIS-LIVIA COLORADENSIS -NEOPHYLLURA ARBUTI-NEOPHYLLURA ARCTOSTAPHYLI-LEUROLOPHUS VITTATUS -EPICARSA SP.-PHACOPTERON LENTIG1NOSUM -DIAPHORINA CARDIAE-DIAPHORINA ALBOMACULATA-PSYLLCPSIS FRAXINICOLA -APHALAROIDA PITHECOLOBIA-APHALAROIDA INERMIS -DIAPHORINA PUTONI I-DIAPHORINA CITRI-DIAPHORINA SOLAN I -PSEUDOPHACOPTERON FLOCCOSA -CAMARCTOSCENA SPECIOSA-EUPHYLLURA PHILLYREAE-EUPHYLLURA CL1VINA-EUPHYLLURA AETHIOPICA-RHINOCOLA ACERIS-STROPHINGIA ERICAE-AGONOSCENA SP, (C)-AGONOSCENA SP. < B) 0-A 03 0-2DISTANCE 01 00 Fig. 183d Part of a phenogram of 208 species; continued from Fig. 183a. 232 I. M. WHITE AND I. D. HODKINSON1 1 05 0-4 03 02 DISTANCE 0-1 -MICROCEROPSYLLA SP.-CALOPHYA SP. (A)-CALOPHYA SCHINI -TENAPHALARA SP.-TENAPHALARA ACUTIPENNIS-PARACARSIDARA GIGANTEA-MASTIGIMAS CEDRELAE-BHARATIANA OCTOSPINOSA-PELMATOBRACH1A SP ,-PROTYORA STERCULIAE-MESOHOMOTOMA HIEISCI -APSYLLA CISTELLATA-CAMAROTOSCENA UN I COLOR-PAUROCEPHALA GOSSYPII-MORANIELLA CALODENDRI -CALOPHYA SP. (B)-CALOPHYA RHOIS-CALOPHYA TRIOZOfllMA :ALOPHYA CALIFORNICA-CALOPHYA DUB I A-CALOPHYA FLAVIDA -PEXOPSYLLA CERCOCARPI -PSYLLA <C. ) MALI-PSYLLA PHORADENDRAE-PSYLLA (H- > ANNULATA-PS V LLA PULCHELLA-PSYLLA < H - ) AMBIGUA-PSYLLA (P-) BETULAENANAE-PSYLLA (P. ) ALNI-RETROACIZZIA ANTENNATA-NEOPSYLLIA ERYTHRINAE-EUPHALERUS GALLICOLUS -FARAPHALAROIDA FREMONTIAE-D1CLIDOPHLEBIA EASTOPI -EUPHALERUS NIDIFEX-COLOFHORINA CASSIAE -SPONDYLIASPIS SP.-GLYCASPIS BAILEYI-TETRAGONOCEPHALA SP.-PACHYPSYLLA VENUSTA-PHELLOPSYLLA SP.-EUCALYPTOLYMA SP.-CARDIASPINA DENSITEXTA-CARD1ASPINA ALB1TEXTURA -EPIPSYLLA SP. (A)-EPIPSYLLA SP. (B)-ANOMONEURA MORI-EUPHALERUS SP. (A)-EUPHALERUS SP, (D) -CTENARY7AINA EUCALYPTI -AREPUNA SP,-ACIZZIA ACACIAE-ACIZZIA RUSSELLAE 00 3 A 3 C 3 S IP Fig. 183e Part of a phenogram of 208 species; continued from Fig. 183a. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 233 Table 8 Selected adult characters. Selected characters are numbered A1-A14. All the character states in the original list of 27 characters aretabulated. However, some character states become combined by the selection procedure and, hence, insome characters two consecutive states are marked with the same value. An asterisk indicates a characterwhich was poorly recorded in the literature. Head charactersAl. Formofgenae. Genae not swollen = Genae swollen but not conical = 1 Genae conical, frons not enveloped by genae = 2 Genae conical, frons enveloped by genae = 3 *A2. Antenna with rhinaria on following segments. III, IV, V, VI, VII, VIII, IX =0 IV, V, VI, VII, VIII, IX =0IV, V, VI, VIII, IX = 1IV, VI, VIII, IX =2III, VI, VIII, IX =2 Thorax characters *A3. Pronotum vertically or subvertically inclined, and laterally constricted. Often completelyor partly concealed by head.No = Yes = 1 *A4. Suture between epimeron and episternum. Horizontal = Vertical (dorsally terminating at mid point of pronotal lateral margin) = Diagonal (dorsally terminating at posterior of pronotal lateral margin) = 1 Forewing characters A5. Forewing with apex acute or acutely rounded. Costal margin curved. M 1+2 terminating ator anterior to apex. No = Yes = 1 A6. Cu+M+R or M+R common stalk present.No = Yes = 1 *A7. Costal break absent. No = Yes = 1 A8. Pterostigma absent. No = Yes = 1 A9. C2 not terminating adjacent to Cu lbNo = Yes = 1 Hind-leg characters *A10. Metatibia with basal (genual) spine present.No = Yes = 1 *A11. Metatibial apical spines or platellae numbering more than five.No = Yes = 1 *A12. Metatarsal spines. Absent = One present = 1 Two present = 2 Male genitalia charactersA13. Male proctiger unipartite.No = Yes = 1 A14. Male proctiger with long caudal lobes present in at least some species of the genus.No = Yes = 1 234 I. M. WHITE AND I. D. HODKINSON Table 9 Rejected adult characters. Rejected characters are numbered A15-A27. Values are not given against the states, which are separated by a V. An asterisk indicates a character which was poorly recorded in the literature. Head characters A15. Vertex with cleft and antennae based upon apices of blunt vertex lobes. No/Yes. A16. Vertex produced into lobes and enveloping genae. No/Yes. A17. Preoccipital lobes present. No/Yes. A18. Preoccular tubercules present: No/Yes. A19. Antenna segment II greatly enlarged. No/Yes. A20. Apical antennal spines longer than antennal segment III. No/Yes. A21. Clypeus long and cylindrical, projecting to anterior margin of head. No/Yes. Forewing characters A22. Forewing with apex acute or acutely rounded. Costal margin curved. M 1+2 terminating posterior to apex. No/Yes. A23 . Nodal line absent . No/Yes . A24. R-Mi+2 cross vein or anastomosis present. No/Yes.A25 . R M (bifurcation of M\ + 2 and M 3 + 4 ) cross vein present . No /Yes . Hind-leg characters *A26. Meracanthus reduced or absent. No/Yes. Male genitalia characters A27. Male subgenital plate with hypovalves. No/Yes. Table 10 Nymphal characters selected after incorporation of adult characters. Characters previously selected and now reselected: Nl , N3, N4, N5, N6, N7, N8, N9, N10, Nil, N13, N14,N15, N16, N17, N18, N19, N23, N24, N25, N26, N27, N28, N30, N31, N32, N33 and N34. Two furthercharacters were selected, as follows. - Outer circum-anal pore ring broken at two or more places (modified N52). No = Yes = 1 - Forewing-pad dorsal surface sectasetae (modified N29). Absent = Pointed = 1 Truncate = 1 Table 11 Groups of species identical with the selected adult plus nymphal characters. The species chosento represent the group is named at the top of each list. 1. Acizzia acaciae 8. Craspedolepta furcata Acizzia acaciaebaileyanae Craspedolepta nervosa 2. Agonoscenasp. (B). 9. Craspedolepta minuta Agonoscena sp. (A). Craspedolepta minutissima 3. Agonoscenasp. (C). 10. Diaphorina cardiae Strophingia cinereae Diaphorina chobauti 4. Aphalara polygoni 11. Diaphorina citri Aphalara curta Diaphorina florea A. nubifera 12. Diaphorina putoniiA. simila Diaphorina clutiae 5. Calophyaflavida D. punctulata Calophya nigripennis 13. Euceropsylla russoi 6. Cardiaspina albitextura Euceropsylla minuticona Cardiaspina spinifera E. sp. C. squamula 14. Euphalerus nidifex Creiis sp . Euphalerus jugo venosus 7. Craspedolepta augustipennis E. rugipennis Craspedolepta artemisiae E. vermiculosus C. sonchi 15. Glycaspis bailey iC. suaedae Glycaspis spp. C. vancouverensis 16. Gyropsylla ilicisC. veaziei Gyropsylla spp. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 235 17. Livia color adensis . Livia maculipennis 18. Mastigimas cedrelae Mastigimas spp. 19. Mesohomotoma hibisci Mesohomotoma spp. 20. Mycopsyllafici Mycopsylla spp. 21. Neophyllura arctostaphyli Neophyllura bicolor 22. Neopsyllia erythrinae Neopsyllia spp.Platycorypha princeps 23. Pachypsylla venusta Pachypsylla spp. 24. Paracarsidara gigantea Paracarsidara spp. 25. Paratrioza cockerelli Paratrioza maculipennis 26. Paurocephala gossypii Paurocephala urenae 27. Psyllaalba Psylla sinuata 28. Psylla alni Psylla buxi Spanioneura fonscolombii 29. Psylla betulaenanae Psylla carpinicolaP. floccosaP. galeaformisP. striataP. trimaculata 30. Psylla brunneipennis Psylla brevistigmata P. coryli P. hamata P. hirsuta P. minuta P. moscovita P. parallela P. pulchra The following species have unique descriptions:Aacanthocnema casuarinaAcizzia hakeaeA. russellaeA. uncatoidesAmorphicola amorphaeAnomoneura moriAphalara exilisA. monticolaA. persicariaA. rumicisAphalaroida spp.Apsylla cistellataArepuna sp.Arytaina genistaeArytainilla spp. 31. Psylla magnicauda Psylla minor 32. Psylla mali Psylla ribesiae 33. Psylla nigrita Psylla saliceti 34. Psylla pyri Psylla ulmi 35. Psylla pyricola Psylla myrtilliP. visci 36. Psylla pyrisuga Psylla melanoneuraP. sorbi 37. Psyllopsis fraxinicola Psyllopsis spp. 38. Strop hingia ericae Tainarys schini 39. Tenaphalara acutipennis Tenaphalara malayensis 40. Trioza albifrons Trioza beamedT. quadripunctata 41. Trioza albiventris Trioza atkasookensisT. crithmi 42. Trioza bakeri Trioza cinnamomiT. diospyriT. frontalis 43. Trioza chenopodii Trioza litseae 44. Trioza curvatinervis Trioza minutaT. salicivora 45. Trioza erytreae Trioza panacis 46. Trioza marginepunctata Trioza vitiensisT. sp. 47. Trioza tripunctata Trioza incidata Bharatiana octospinosaCalophya calif ornicaC. dubiaC. rhoisC. schiniC. triozomimaC. rotundipennisC. sp. Camarotoscena spp.Cardiaspina densitextaCeanothia spp.Ceropsylla spp.Ciriacremum spp.Colophorina cassiaeColposcenia sp. 236 I. M. WHITE AND I. D. HODKINSON Craspedolepta constrictaC. nebulosa C. subpunctataCrastina linavuoriiCrawforda triopsyllinaCtenarytaina eucalyptiDiaphorina albomaculata D. solani Diclidophlebia eastopiEgeirotrioza spp.Epicarsa sp.Epipsylla spp.Eucalyptolyma sp.Euceropsylla cayeyensisEuglyptoneura spp.Euphalerus gallicolus E. tantillusE. sp. (A).E. sp. (B).E. sp. (C).E. sp. (D).Euphyllura spp.Floria variegataFreysuila sp.Heteropsylla spp.Hevaheva swezeyiHomotoma spp.Insnesia glabruscutaIsogonoceraia divergipennisKuwayama pisoniaLeptynoptera sulfureaLeurolophus vittatusLeuronota michoacanaLivia crefeldensis L. juncorumL. vernalis Macrohomotoma spp.Microceropsylla sp.Mitrapsylla deserataMoraniella calodendriNeolithus sp.Neophyllura arbutiParaphalaroida fremontiaeParatrioza arbolensisP. lavateraePauropsylla spp.Pelmatobrachia sp.Pennavena fabulosaPexopsylla cercocarpiPhacopteron lentiginosumPhellopsylla sp. Phytolyma spp. Protyora sterculiae Pseudoeriopsylla nyasae Pseudophacopteron spp. Psylla albagena P. ambigua P. americana P. annulata P. foersteri P. magna P. media P. negundinis P. palmeni P. peregrina P. phoradendrae P. pruni P. pulchella P. rhamnicola P. rhododendri P. simlae P. stricklandi P. subspiculata Purshivora spp. Retroacizzia antennata Rhinocola aceris Spondyliaspis sp. Swezeyana elongagena Synoza spp. Tenaphalara sp. Tetragonocephala sp. Togepsylla matsumurana Trichochermes walkeri Trigonon longicornis Trioza alacris T. anceps T. aylmeriae T. falcata T. hirsuta T. lobata T. magnoliae T. nigricornis T. obsoleta T. obtusa T. palmicola T. phoradendrae T. remota T. urticae T. vitreoradiata Triozamia lamborni Triozoida silvestris NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 237 NYMPH Prothorax completely separate to head. Mesothorax and metathorax with well-defined medial andlateral sclerites. Abdomen with well-defined sclerites and a small caudal plate. Anus ventral andsurrounded by a pore field of uncertain form. Whole body (dorsal, margin, including wing-pads, and antennae) covered in pointed sectasetae andsimple setae. It is interesting to note that pointed sectasetae appear to be an ancestral attribute. However,Becker-Migdisova (1973) believed the ancestral nymph lacked sectasetae and was similar toTenaphalara. This would imply that sectasetae, which are a highly complex structure, must haveevolved several times. Becker-Migdisova fails to explain this unparsimonious assumption. Wagner tree An attempt was made to analyse cladistic relationships by constructing a Wagner tree (Farris,1970). Characters were selected by phyletic weighting (Cain & Harrison, 1960), so that only onecharacter out of a set of characters that might be functionally or ecologically correlated was usedin the analysis. This weighting left only 17 characters, which were inadequate for meaningfulanalysis (White, 1980). However, this analysis did indicate a need to re-examine the structure ofthe tarsal arolium. Reanalysis of tarsal arolium structure One large branch of the Wagner tree was initially defined by the presence of a petiolate tarsalarolium in the nymph and the branch included all the Psyllidae except Diaphorini and Psyllopsis.Furthermore, the presence or absence of a petiolate tarsal arolium in the nymph, received veryhigh eigenvector and SUMRAT values in previous analyses, and the presence state appears tobe largely confined to the Psyllidae. However, it was also recorded as present in a fewAphalaridae (Aphalara persicaria and Paurocephald) and Triozidae (Trioza hirsuta). Uponre-examination the finer structure of the arolium became apparent. Most species have asclerotized 'rod' or 'rods' running longitudinally from the base of the arolium (shown black inFigs 59-94). The homology of this structure is uncertain, but it will be referred to as anunguitractor. Some species lack any visible arolium (most Spondyliaspididae), while others lackany visible unguitractor (Aphalarinae (Aphalaridae) and Egeirotrioza (Triozidae)) (Fig. 95). Ashort unguitractor is probably the ancestral state (Fig. 67) . It is greatly elongated in almost all thePsyllidae, Diclidophlebia (Carsidaridae), Aphalaroida, Camarotoscena, Euphyllurini, Para-phalaroida and Paurocephala (Aphalaridae) (Figs 59, 62-66, 69, 70, 79-93). Some species haveretained a membrane adjacent to the long unguitractor: Diaphorini (Fig. 60) and Euphyllurini(Figs 62-64) while in others it is lost (Fig. 59) . Careful analysis of arolium form was found to be ofgreat value in cladistic analysis of species which otherwise differ little from the ground plan. Cladistic method The Wagner tree analysis indicated that a method of cladogram construction was needed whichused all of the available characters. To overcome the problem of GAIN characters being of greatervalue than LOSS characters a two-phase method of cladogram construction was devised. A GAIN character is here defined as one whose derived state is the presence of an attribute. ALOSS character is defined as one whose derived state is the absence of an attribute. Losscharacters should be accorded low weight because a structure could be lost in many distantlyrelated phyletic lines (Mayr, 1969). When a high proportion of characters are of the loss typecertain modifications to the cladistic principles of Camin & Sokal (1965) and Hennig (1966) mustbe applied. A total of 159 adult and nymphal characters was divided into two sets (Table 12): 1, gaincharacters and, 2, loss characters. Using gain characters only, the most parsimonious cladogramwas formed. The loss characters were then added to the cladogram such that each group of taxa 238 I. M. WHITE AND I. D. HODKINSON which was divided by loss characters only had maximum parsimony. However, the losscharacters were not most parsimonious over the entire tree. When a section of the tree could only be structured by loss characters and many equallyparsimonious solutions were possible, the characters were weighted and added to the tree inorder of decreasing weight. The weights were the sum of the mutual information (Legendre &Rogers, 1972) values for each character with all other characters in the section of tree understudy. This method is similar in principle to using the compatibility method of Le Quesne (1969) . In the illustration of the cladogram (Figs 184-196) the following convention was adopted. 1. Gain characters are marked by squares; black for derived and white for ancestral. 2. Loss characters are marked by circles; black for derived and white for ancestral. This enablesclades which are only defined by loss characters to be instantly recognisable. 3. Each ancestor is numbered, e.g. the ancestral psyllid is number one. All descendants are saidto belong to clade one. There are 94 ancestors giving rise to 106 taxa of generic, subgeneric andin some cases species level groupings. Formation of initial branches in the cladogram (Fig. 185) A few of the characters used by Schlee (1969) and Szelegiewicz (1971) are included at the base ofthe cladogram, to define both the Psylloidea and their supposed sister clade the Aleyrodoidea(Fig. 184). Certain major clades (2, 6, 13, 15, 17, 32, 37 and 39) are defined by complex gain characters.Apsylla, Bharatiana, Livia, Mastigimas and Strophingia are separated from the ground planalmost entirely by loss characters and are not easily placed. Apsylla adults have very long apical antennal spines, which is a gain character ancestral toclade 39 (Fig. 190), e.g. Calophya, although Pseudophacopteron (clade 37, Fig. 189) also has thisattribute. Apsylla is therefore placed as a sister group to clade 39 (clade 10 is formed).Bharatiana adults have a fairly large clypeus and large lateral ocelli, though not as pronounced asin clade 37, e.g. Phacopteron. With hesitation, Bharatiana is placed as a sister group to clade 37(clade 8 is formed). Mastigimas nymphs have broken bands of anal pores, a feature which couldbe derived from bands of anal pores, an attribute which is ancestral to clade 32, e.g.Paracarsidara (Fig. 187). Although anal pore bands also occur in Epicarsa, Mastigimas is placedas a sister group to clade 32 (clade 4 is formed). Livia and Strophingia are only separated fromthe ground plan by loss characters and cannot be placed with any clade so far formed. The ten-way furcation from the ground plan (to clades 2, 4, 6, 8, 10, 13, 15 and 17 plus Liviaand Strophingia) was resolved using weighting and the resulting branches are shown in Fig. 185. Becker-Migdisova (1973) proposed that the ancestral psyllid gave rise to three separate lines(a CarsidaralTrioza line, an Aphalara line and a Psylla line). However, Klimaszewski (1964)proposed a bifurcation into a CarsidaralTrioza line and an Aphalara/ Psylla line. Furthermore,Vondracek (1957) suggests a Spondyliaspisl CarsidaralTrioza line and a Calophya/ Psylla/ Apha-lara line. The present cladogram agrees with a line typified by Trioza and Carsidara (cf.Paracarsidara} and includes some of the groups which these authors have referred to Carsidar-idae, such as Calophya, Phacopteron and Homotoma. However, the present cladogramprovides no justification for the Aphalaridae, except as a collection of groups phenetically closeto the ground plan from which other clades can be derived. The Psyllidae arise from one suchgroup (clade 17). Clade descriptions Clade 2 (Fig. 186) corresponds with the Triozidae plus the Leptynopterinae and Pauropsyllini ofBecker-Migdisova (1973). However, this is not the Pauropsyllini of Loginova (1972), many ofwhich are placed in clade 17, e.g. Paurocephala (Figs. 193-196). The genera Leptynoptera andPauropsylla have a typical Trioza type adult pronotum, wing venation and sometimes nymphalform. They are placed in clade 18 with other genera possessing these features. The pronotalstructure of Trioza is also found, to a lesser extent of development, in Homotoma (clade 6, NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 239 184 185 Figs 184, 185 Clades and 1. 184, clade 0, including the Psylloidea (clade 1) and Aleyrodoidea; 185, clade1, the Psylloidea. Details of clades 2, 4, 6, 8, 10, 13, 15 and 17 are illustrated in Figs 186-196. 240 I. M. WHITE AND I. D. HODKINSON Fig. 188) and Microceropsylla (clade 10, Fig. 190). Clade 2 can be divided into three majorsections. (i) Clade 2 (minus clade 18) (Fig. 186a), e.g. Trichochermes includes those genera which lackthe Trioza type adult pronotum and the fusion of the dorsal surface of the nymphal abdomen. (ii) Clade 18 (minus clade 23) (Fig. 186b) contains genera in which the nymphs are fairlyelongate, the hindwing-pad margin is not confluent with the abdomen margin and the marginalsectasetae are normally well spaced apart. (iii) Clade 23 (Fig. 186c) contains genera which have 'disc'-shaped nymphs, often withmarginal scales, dorsal clavate setae or closely spaced marginal sectasetae. Clade 4 (Fig. 187) is equivalent to the Carsidarinae and Tenaphalarinae of Becker-Migdisova(1973) and Klimaszewski (1964). Clade 32 is defined by the presence of a cross-vein in the adultfore wing. This is subdivided into Tenaphalara (with an extra cross vein) and clade 31 (with adeeply cleft adult head). Figs 186a, b Clades 2 and 18. 186a, clade 2, continued in Figs 186b, c; 186b, clade 18, continued in Fig.186c and from Fig. 186a. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 241 Fig. 186c Clade 23; continued from Fig. 186b. Clade 6 (Fig. 188) is the Homotominae of Becker-Migdisova (1973). Clade 33 represents agroup in need of some revision, for some Homotoma spp. not examined in this study may beplaced with Synoza in the present cladogram, e.g. H. gressitti Miyatake has no M+Cu vein in thewing. Miyatake (1974) revised some Homotoma spp. but overlooked Synoza and was unawareof undescribed African species of the group (in coll. British Museum (Natural History)). Clade 8 (Fig. 189) is the Phacopterinae of Becker-Migdisova (1973) plus lEpicarsa andBharatiana. The latter is only tentatively placed. The nymph examined of the former waslabelled as being found with an adult close to Epicarsa, from Brazil. The adult characters in thepresent cladogram are largely those given by Crawford (1911); Ferris (19286) described anEpicarsa from Mexico, but there is some doubt about its true identity. Lima & Guitton (1962)described another Brazilian member of clade 37 (Phacosemoides sicki). The Pacific genusChineura Tuthill should also be placed here. In the cladogram Pseudophacopteron nymphs aredescribed as having lanceolate setae. This only applies to material labelled as ?P. floccosa fromGuam: these setae are lost in other Pseudophacopteron spp. examined. P. floccosa is a Sri Lankaspecies which is unlikely to occur on Guam. Since wing form in this group is distinct it is assumedthat the material from Guam is a member of clade 37. Guam is the type locality of Chineurapaucivena Tuthill which may have been confused with Pseudophacopteron. Clade 10 (Fig. 190) includes Calophya plus some genera referred to Pauropsyllinae: Micro-ceropsyllini and Anomalopsyllinae: Apsyllini by Becker-Migdisova (1973). Loginova (1972)places Microceropsylla and Pelmatobrachia in the Pauropsyllini. The genus Calophya itself isreferred to the Carsidaridae by Becker-Migdisova (1973), the Pauropsyllinae by Crawford(1914), and the Psyllidae by many authors (Klimaszewski, 1964; Dobreanu & Manolache, 1962;Hodkinson & White, 19796). Several species which probably belong within clade 34 are still 242 I. M. WHITE AND I. D. HODKINSON 187 188 Figs 187, 188 Clades 4 and 6. 187, clade 4; 188, clade 6. placed in Pauropsylla, e.g. P. longispiculata Mathur, P. maculata Mathur and P. verrucosaMathur. There is little apparent difference in general form between some Calophya s.str.,Neocalophya Miyatake (a subgenus of Calophya), Paracalophya Tuthill and HolotriozaBrethes. The entire group is in urgent need of revision as indicated by Miyatake (1971). Livia and Strophingia (Fig. 185) must be regarded as morphologically primitive genera,although it is not intended to imply that they are very ancient. They have retained the groundplan facies and have no gain characters in common with other clades and their true cladisticrelations are uncertain. Cladistic relations within the genus Strophingia are postulated byHodkinson(1981). Clade 13 (Fig. 191) contains species with 'Aphalarid' facies, i.e. species which are pheneticallyclose to the ground plan but possess a caudal lobe on the adult male proctiger. The caudal lobealso occurs in clade 83 and it is possible that it has evolved more than once among the species inclade 13. Although clade 13 may not be a monophyletic group, clade 41 is partly defined by theunique feature of a 'tooth', lobe or hook-like structure on the ventral edge of the caudal lobe.Hence, only the position of Phytolyma is dubious. Phytolyma has in the past been assigned tomany different groups. Heslop-Harrison (1952b) placed it near Rhinocola (clade 15), but laterhe (Heslop-Harrison, 1958) added it to the Pauropsyllini. Both Vondracek (1963) and Becker-Migdisova (1973) referred it to the Anomalopsyllinae, though the former regarded thatsubfamily as belonging to the Spondyliaspididae, while the latter placed it in the Aphalaridae. Inreality Phytolyma is probably a relic genus retaining many features of the ground plan, abipartite male proctiger and preoccipital lobes in the adult. Hollis (1973) stated that the tropical NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 243 12612714314770 93 i 89 189 X 190 Figs 189, 190 Clades 8 and 10. 189, clade 8; 190, clade 10. components of this group are poorly known; they probably derived from basic psyllid stockearlier than temperate forms, and are difficult to place in the existing classification. Clade 43 is defined by the diagonal suture between the adult epimeron and episternum. Clade42 is the tribe Stigmaphalarini of Vondracek (1957) and Colposceniini of Becker-Migdisova(1973). Clade 40 is the Aphalarini of most authors. Clade 15 (Fig. 192) contains species which have retained many ground plan features:Moraniella nymphs are surrounded by pointed sectasetae and adult males of Tainarys have abipartite proctiger. However, the form of the tarsal arolium which defines clade 15 is unique.Nevertheless, the branching within the clade is based on loss characters only, and weights wereapplied. Clade 17 (Figs 193-196) contains over half of the species studied and it is defined by theelongation of the unguitractor in the tarsal arolium of the nymph. The major division is intoclades 48 and 53 (Fig. 193). Clade 53 is defined by nymphal capitate setae, but excludes those species where the nymph hasretained numerous sectasetae or where the sectasetae are reduced to lanceolate setae. Clade 48is formed by adult loss characters, which are in common to all of these morphologically moreprimitive species. It is further divided into clades 47 and 50. Clade 47 (Fig. 194), defined by the presence of 1 + 1 pore rings (or a derivable feature) on thenymphal abdomen, is divided into clades 60 and 62. Clade 62 contains the genera Diclidophlebiaand Paraphalaroida. Paraphalaroida contains one species (P. fremontiae) which, prior to therevision of Loginova (1972), was regarded as a Paurocephala sp. Diclidophlebia was referred tothe Carsidaridae: Tenaphalarinae, Diclidophlebiini by Becker-Migdisova (1973). The genus 244 I. M. WHITE AND I. D. HODKINSON 192 Figs 191, 192 Clades 13 and 15. 191, clade 13; 192, clade 15. Togepsylla (omitted from this cladogram due to lack of information on certain character states,e.g. the form of the tarsal arolium) may belong to clade 62 because its adult head structure issimilar to Diclidophlebia (Becker-Migdisova, 1973). Certain species still referred to the genusPaurocephala, such as P. tuxtlaensis Conconi from Mexico and P. menoni Mathur from India,probably belong to clade 62. Clade 60 is divided into clades 59 and 61 by the development of adult vertex lobes as opposedto genal cones and elongated wing cells. Clade 59 is the Euphyllurini of Loginova (1973) and partof the Euphyllurini of Becker-Migdisova (1973). Clade 61 is referred to the Spondyliaspididae by most authors (Becker-Migdisova, 1973;Hodkinson & White, 19796; the Spondyliaspidinae by Heslop-Harrison, 1954). Unlike thenymphs of other Spondyliaspididae (clade 55, Fig. 195) examined, Ctenarytaina and Eucalypto-lyma have marginal lanceolate setae and, therefore, fit the cladogram outside of clade 56. Bothare characterised by the reduction or absence of tarsal arolia and if they do belong to clade 17 thismust represent a secondary loss. The pore field on the nymphal abdomen of Eucalyptolyma sp.and Ctenarytaina eucalypti may be derived from 1 + 1 pore rings, the defining character of clade NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 245 Fig. 193 Clade 17, giving rise to clades 47, 55 and 56 which are illustrated by Figs 194, 195 and 196respectively. 47, and hence clade 61 is placed in a position of maximum parsimony. Nymphs of Ctenarytainathysanum Ferris & Klyver, which we have subsequently seen, lack the abdominal pore field. Theposition of Eucalyptolyma requires further investigation since recently acquired material of ?fuscipennis Froggatt nymphs are of a structure concordant with clade 61 while those of E.maideni Froggatt are structurally close to clade 68. Two Indian species (Euphyllura caudataMathur and E. concolor Mathur) may also belong to clade 61 on the basis of the pore field of thenymphal abdomen and the structure of the adult female proctiger. 246 I. M. WHITE AND I. D. HODKINSON Fig. 194 Clade 47. Clade 50 (Fig. 193) is a maximum parsimony collection, based on loss characters, of generabelonging to clade 17 but not clades 47 or 53. It is divided into clades 49 and 51. Clade 51 isdefined by the presence of genal cones, a feature which occurs many times in the cladogram, andclade 49 is only formed by loss characters. Clade 49 (Fig. 193) contains the genera Camarotoscena and Paurocephala, the former ofwhich was regarded as a subgenus of Paurocephala by Vondracek (1957). Most authors haveplaced these genera in the tribe Paurocephalini of the Aphalaridae: Paurocephalinae (Becker-Migdisova, 1973; Klimaszewski, 1964). Because the general facies of the adult head is similar toPauropsylla (clade 25, Fig. 186b) many authors placed Paurocephala in the Pauropsyllinae(Crawford, 1914; Loginova, 1972; Mathur, 1975). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 247 Clade 51 (Fig. 193) is the Psyllidae: Arytaininae, Diaphorinini of Vondracek (1957),Dobreanu & Manolache (1962) and Klimaszewski (1975). Becker-Migdisova (1973) placedPsyllopsis in the Psyllidae: Arytaininae, Psyllopseini. Pflugfelder (1941) placed Diaphorina andPsyllopsis in the Aphalaridae, and Low (1879) placed them in the Psyllidae and Aphalaridaerespectively. The nymphs of Psyllopsis, Diaphorina and Pennavena have an 'Aphalara' faciesand are surrounded by marginal lanceolate setae. The long unguitractor of the nymphal aroliumsuggests that these genera belong in clade 17. However, the presence of lanceolate setaeexcludes these genera from clade 56 where most former authors have placed them. Further-more, the adults retain a crown of about 10 spines at the apex of the metatibia, a feature alwaysreduced in clade 57 (Fig. 196). Clade 53 (Figs 193-196) begins with a major transition between members of clade 17 with'Aphalara' and 'Psylla' type facies. The nymph of Aphalaroida is in many respects similar toEuphalerus or Acizzia while the adult is phenetically similar to Strophingia. The nymph ofAphalaroida pithecolobia is covered by rod setae, similar to Euglyptoneura robusta andPexopsylla cercocarpi (both Clade 56, Fig. 193). The position of E. robusta (clade 82, Fig. 196e)suggests that rod setae are modified capitate setae and therefore, in the cladogram, rod setae arenot differentiated from capitate setae. The adult oiArepuna has a wing of a Euphalerus type and the nymph is surrounded by clavatesetae. These could be very reduced sectasetae, which would place Arepuna outside of clade 56,or reduced capitate setae, which would place it anywhere in clade 53. A larger number of lossesmust be proposed if Arepuna is to be placed within clade 56 rather than as a sister group to it. Clade 55 (Fig. 195) is initially defined by the presence, in the nymph, of 1 + 1 pore ringsadditional to the circum-anal ring. It is assumed that these rings become split to form the 2 + 2rings which initially define clade 66. Even without this assumption the contents of clades 64 and66 would still arise from close to the start of clade 56 (Fig. 193). In clade 63 the preoccipital lobesare lost and the 1 + 1 pore rings become areas of separated pores. Although nymphal capitatesetae are regarded as lost in clade 55, they may be retained in some species, such as Psyllabengalensis Mathur, which were not examined but appear to belong to clade 64. Clade 66 is initially defined both by the presence of 2 + 2 nymphal pore rings and a serrateapex to the nymphal abdomen. Clade 65 is a collection of species in which the serrate apex to theabdomen is retained but the pore rings are often lost or reduced to small groups of pores. Thisreduction could be derived from 1 + 1 or 2 + 2 pore rings and species which belong to clade 55,but are excluded from clades 64 and 67, are grouped for convenience into clade 65. Clade 65consists of the following taxa: clade 72, Euphalerus nidifex, Pachypsylla spp. (other than P.japonica), Phellopsylla and Retroacizzia all of which are only separated from ancestor 65 by losscharacters. The details of clade 65 were constructed by weighting. Phellopsylla belongs to theSpondyliaspididae of all authors, clade 73 to the Spondyliaspididae: Pachypsyllinae of Becker-Migdisova (1973) and clade 74 to the Psyllidae. Ancestor 67 (Fig. 195a) marks a transition. Clade 67 is initially defined by having enlargedouter teeth on the serrate apex of the nymphal abdomen, as in Euphalerus gallicolus. Clade 68 isa collection of species with a pointed cauda in the nymph: Creiis has both a pointed cauda and1 + 1 tooth-like structures near the apex of the abdomen. These are treated as being homologouswith the enlarged outer teeth in E. gallicolus. Further evidence for the inclusion of clade 68 in 66is provided by the fact that lerp-forming species are confined to clades 65 and 68. Nymphs of species in clade 68 have weakly sclerotized abdomens and the caudal plate isrudimentary. This implies either that a large caudal plate has been derived separately in severalbranches, or that a reversal to separate segments has occurred. This appears to contradictDollo's Law. However, the genotype must contain coding for all abdominal segments since theyoccur in the adult, i.e. Dollo's Law is not broken at the level of the genotype. Clade 57 (Fig. 196) is defined by loss characters only and contains genera which belong withinclade 56 but not 55. With the exception of clade 51 (Fig. 193), and some Euphalerus spp.(including the type-species of the genus, E. nidifex} and Retroacizzia which have been assignedto clade 55 (Fig. 195), it is the Psyllidae of most authors. Certain clades (78, 82 and 85) are defined by gain characters leaving the genera Acizzia, 248 I. M. WHITE AND I. D. HODKINSON Figs 195a, b Clades 55 and 65. 195a, clade 55, continued in Fig. 195b; 195b, clade 65, continued from Fig.195a. Amorphicola, Anomoneura, Arytaina, Arytainilla, Epipsylla, Floria, Mitrapsylla, Trigonon anda few species referred to the genus Euphalerus unplaced. Character 43 (caudal lobe on adultmale proctiger) was incompatible with character 1 (very broad head) and was initially ignoredbecause it occurs in other apparently unrelated groups such as Aphalara and may also haveevolved many times within clade 57. Character 13 (position of antennal insertions) was alsoomitted initially since a tendency for the antennal bases to move back not only occurs in all of NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 249 clade 82 (Fig. 196e), Arytaina, Arytainilla and Floria but also in a few species of other clades,such as Ciriacremum nigripes Hollis. Character 106 (broad anal ring) is also ignored since thisoccurs in Psylla s.str. (clade 82) as well as in Anomoneura and Epipsylla, and its derivation isuncertain. The details of clade 57 were then constructed by weighting to form five major clades(75, 76, 78, 80 and 82); characters 43, 13 and 106 were then replaced. Clade 75 (Fig. 196b): the first branch, from the general line in clade 57, forms clade 75. Thelargest genus in this clade is Acizzia, some species of which, such as the type-species (A.acaciae), differ from the description used in the cladogram in that they have lost one spine fromthe apex of adult basal metatarsus. Furthermore, there is a very high diversity of nymphal formin the genus. Neopsyllia and Platycorypha, two genera which apparently differ only in therelative length of the caudal lobes of the adult male proctiger, are the only taxa in clade 57 toretain nymphal sectasetae on the hindwing-pad margins. Freysuila is placed as a sister group toNeopsyllia and Platycorypha on the basis of the very broad adult head and, therefore, asecondary loss of the caudal lobes is assumed to occur in Freysuila. Mitrapsylla deserata nymphshave lanceolate setae on the dorsal surface indicating the retention of dorsal sectasetae, or thederivative lanceolate setae, well into clade 57. The abdomen margin sectasetae (character 114)are reduced to lanceolate setae in M. deserata, a feature known elsewhere only in Heteropsylla(clade 78). In the remainder of clade 57 (clade 77) the only remaining sectasetae are (up to 4 -I- 4in number) on the abdomen margin (character 114). 75 Figs 196a, b Clades 57 and 75. 196a, clade 57, continued in Figs 196b-e; 196b, clade 75, continued fromFig. 196a. 250 I. M. WHITE AND I. D. HODKINSON Clade 76 (Fig. 196c). In the character weighting procedure character 76 (loss of genual spine)received the greatest weight and defined clade 76, which includes Amorphicola, Anomoneura,Epipsylla and some species referred to the genus Euphalerus. Euphalerus spp. are placed in bothmajor branches of clade 56 (Fig. 193) and these separate groups may only be recognisable in thenymphal stages. Some species at present referred to the genus Psylla, such as P. hyalina Mathurand P. oblonga Mathur, probably belong to clade 76. Adults of Anomoneura and Epipsylla areradically different in facies due to such characters as the presence of forewing cross veins in theformer and very long genal cones in the latter. The nymphs, however, differ only in the presenceof abdominal sectasetae in A. mori. Clade 78 (Fig. 196d) is defined initially by the shape of the adult male paramere (character 46)and by the fact that antenna segment III is not the longest. Both these characters aresubsequently lost by many species. There is also a tendency for the base of the pterostigma to bebroader than the length of the vein R between the R/Rs fork and the fl/pterostigma basepositions. This feature reaches its maximum development in certain Kleiniella spp. (Hollis,1976). The African genus Kleiniella Aulmann is one of several genera which probably belong toclade 90 but whose nymphs are unknown; others being Delina Blanchard (South America),Palmapenna Hollis (Africa) and Panisopelma Enderlein (South America). Clade 89 is definedby tubular abdominal sectasetae (character 115) on the nymphs. The adults of Euceropsylla andHeteropsylla differ only in the development of the genae, despite radically different nymphs. Anabsence of genal cones in Heteropsylla has previously caused it to be referred to the Pauropsylli-nae by many authors, who also include Paurocephala in that group. There is also a tendency forthe genal cones to be reduced in Ciriacremum, though only in some species. Contrary to theopinion of Becker-Migdisova (1973) Ciriacremum spp. have neither rudimentary genal conesnor a bipartite male proctiger. Further details of the cladistic relations of clade 90 are given byHollis (1976). Many Neotropical species at present referred to the genus Psylla, such as P.forcipata Tuthill, P. fuscinodulus Enderlein and P. ingae Tuthill, probably belong close to Figs 196c, d Clades 76 and 78. 196c, clade 76, continued from Fig. 196a; 196d, clade 78, continued fromFig. 196a. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 251 Fig. 196e Clade 82; continued from Fig. 196a. Euceropsylla in clade 78. Many of the described species of Euceropsylla are very similar and afull revision is required. The genus Arytaina may contain some Pacific species belonging close toInsnesia or Isogonoceraia. Clade 81 (Fig. 196a) is formed by replacing character 13 (antennal base position) which wasomitted in the initial weighted analysis of clade 57 and is divided into clades 80 and 82. Clade 80 (Fig. 196a) is the Arytainini of Becker-Migdisova (1973). Loginova (19760, 1977)divided the Arytaininae into the tribes Arytainini (containing Arytaina and Florid) and theCyamophilini (containing Arytainilla, plus Acizzia of clade 75 and Amorphicola of clade 76).The character used to separate the tribes was the absence and presence of the costal break in thefore wing, respectively. However, this character is variable in the type-species of Floria(Hodkinson & White, 1979a) and is therefore a poor character on which to base tribal groups.Heslop-Harrison (1961) included within the Arytainini, Amorphicola (clade 76), Ceanothia,Euglyptoneura and Purshivora (all clade 82) together with Acizzia (clade 75) and clade 80. Nymphs of Amblyrhina torifrons Low which we recently collected are, within the bounds ofthe characters used in the cladogram, identical to Arytaina. Other genera which probably fitclade 80 are Alloeoneura Low and Livilla Curtis. Clade 82 (Fig. 196e) is the genus Psylla s.l. (minus species which have already been referred toother clades) plus Spanioneura and three North American genera, Ceanothia, Euglyptoneuraand Purshivora, placed in the Arytainini by Heslop-Harrison (1961). Additional information,on the number of gonads, is available for a few of the subgenera of Psylla shown in clade 82 and 252 I. M. WHITE AND I. D. HODKINSON Table 12 Characters used in the cladogram. There are four categories of characters used in the cladogram: 50 adult gain, 32 adult loss, 35 nymphal gainand 41 nymphal loss. In some cases shape changes could not easily be categorized as gain or loss. In general,such characters were described as gains, especially if of a complex nature. However, when a shape changewas lacking in compatibility with complex type gain characters, i.e. liable to be multiply derived, it waslisted with the loss type characters. Only the derived state of each character is given in the followingtabulation. Adult 'gain' characters 1. Head, with eyes, more than six times as broad as vertex is long. 2. Vertex deeply cleft and antennae based upon apices of blunt vertex lobes. 3. Genae swollen. 4. Genae formed into cones which are in the same plane as the vertex. 5. Genae formed into cones which are deflected ventrally from plane of vertex. 6. Vertex produced into lobes and enveloping genae. 7. Vertex mid-line paralleled by closely proximal ridges.9. Preocular tubercles present. 10. Lateral ocelli at extreme posterior margin of head and very prominent. 11. Clypeus large. 12. Clypeus produced anteriorly. 13. Antennal insertions high on vertex, not on front vertex/genal area. 14. Antennal flageller segments (III-X) as broad as basal two segments. 15. Antennal segment II greatly enlarged. 16. Antennal segment VIII longer than segment III. 17. Antennal apical spines very long (at least as long as segments IX and X together or segment III andoften almost as long as whole antenna). 18. Pronotum vertically or subvertically inclined, and laterally constricted. Often completely or partlyconcealed by head. 19. Suture between episternum and epimeron diagonal (dorsally terminating at posterior of pronotallateral margin). 20. Suture between episternum and epimeron horizontal. 21. Wing very broad subapically (Pauropsylla shape). 22. Wing of Spanioneura shape. 23. Wing very elongate with veins straight and almost parallel. 24. Forewing with apex acute or acutely rounded. Costal margin curved. M 1+2 terminating at or anteriorto apex. 25. Forewing with apex acute or acutely rounded. Costal margin curved. M 1+2 terminating posterior toapex. 26. Wing thick or coriaceous, not membraneous. Shape rhomboidal. 27. Wing veins broad. 28. Costa broad. 29. Lenticula costal field. 30. Veins Rs and M sinuate (Neophyllura subgenus Arbutophila}. 31. Cells cui and m 2 very elongate. Wing fairly broad (Auchmerina, Caradocia, Freysuila, Geijerolymaand Macrocorsa wing forms). 32. Subcosta and costa coalesced. 33. Veins R, M and Cu\ with a common stem. 34. Veins R, M and Cu\ separated from common stem at one point or veins R and M with a common stalkafter the branching of C^. 35. Cu 2 terminating at a point well separated from Ci b , and often closer to the wing base (this separationoccurs in many clades but is only well expressed in clade 22, e.g. Trioza). 36. Cross veins: pterostigma-Rs. 37. Cross vein : R-Rs. 38. Cross vein: R-M (bifurcation of M 1+2 and M 3 + 4 ) cross vein. 39. Cross vein: R-Mi +2 (or anastomosis of R and M 1+2 ). 40. Well-developed radular spinules. 41. Mid tibia with a dark heavily sclerotized band around apex. 42. Sperm pump of form common to Psylloidea and Aleyrodoidea. 43. Male proctiger expanded posteriorly to form caudal lobes. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 253 44. Tooth'-like armature placed ventrally in basal half of caudal lobe. 45. Paramere with inner spiniform process (Purshivord). 46. Paramere bifid when viewed posteriorly (e.g. Heteropsylld) . 47. Paramere bifid when viewed laterally (Egeirotriozd). 48. Long thin paramere (Arytainilla). 49. Stout paramere with a heavily sclerotized and thickened blunt apex (Arytaind). 50. Female dorsal valve short, rounded down and densely covered by long thin setae (Pachypsyllajaponica and Tetragonocephala sp.). 51. Female dorsal valve (proctiger) with lanceolate setae arranged along margin (Ctenarytaina, Euphyl-lura caudata and E. concolor) . Adult 'loss' characters 52. Genal cones greatly reduced or lost (derived from character 5). 53. Anteoccipital lobes absent. 54. Antenna without very narrow (Mastigimas type) flagellum (segments III to X). 55 . Antenna less than twice as long as head breadth (a character of secondary loss which is only applied toclade82,e.g. Psylla). 56. Rhinarium absent from segment III. 57. Rhinarium absent from segment IV. 58. Rhinarium absent from segment V. 59. Rhinarium absent from segment VII. 60. Wing apex rounded, remaining coriaceous (unless combined with character 61) (derived fromcharacter 26). 61. Wing membraneous (derived from characters 26 or 60). 62. Euphalerine wing (derived from character 26). Generally maculate. 63. Wing apex rounded (derived from character 62). 64. Wing apex rounded (derived from character 24). 65. Wing apex rounded (derived from character 25). 66. Branches of M and Cu reduced. 67. Cell cu 2 (claval field) and anal vein absent. 68. Cell cui absent. 69. Nodal (fold) line absent. 70. Costal break absent. 71. Pterostigma reduced (only applied to clade 82, e.g. Psylla). 72. Pterostigma absent or very reduced. 73. Hind wing very reduced or absent. 74. Meracanthus very small. 75. Meracanthus absent. 76. Metatibia basal (genual) spine absent. 77. Fewer than six spines at apex of metatibia. 78. Metatarsus segment I with one apical spine. 79. Metatarsus segment I with no apical spines. 80. Male proctiger not bipartite (segment X and XI of abdomen fused to produced a unipartiteproctiger). 81 . Tooth'-like armature in basal half of caudal lobe absent. (Applies only to clade 41 . Secondary loss ofcharacter 44.) 82. Caudal lobe absent (secondary loss of character 43). 83. Paramere of bifid form (character 46) secondarily lost. Nymphal 'gain' characters 84. General body form broader than long (Homotoma, Macrohomotoma, Mycopsylla and Pseudoeriop-sylla). 85. General body form rounded (facies of e.g. Ceropsylla). 86. General body form very elongate (facies of Aacanthocnema). 87. Antenna short and narrowed evenly to apex (Fig. 44). 88. Head and prothorax completely fused dorsally. 89. Humeral lobe of forewing-pad anteriorly produced to an extreme which is anterior to the procoxa(Figs 17-19). 90. Unguitractor long (Figs 77-94). 91. Arolium with a long petiole (Figs 65, 66). 254 I. M. WHITE AND I. D. HODKINSON 92. Arolium broader than long, without petiole and with a pair of darkened areas (Figs 72-76). 93. Arolium base/petiole apex with a semicircular membraneous area (Fig. 65). 94. Abdomen with all dorsal sclerites fused with caudal plate. 95. Abdominal apex serrate (Fig. 122). 96. Abdominal apex with large 'teeth' at lateral extremities of serrate area (follows from character 95)(Figs 118, 120). 97. Abdominal apex with large medial 'teeth' (Fig. 125) (from character 95). 98. Abdominal apex with a pair of apical 'teeth'. 99. Abdominal segments produced laterally as rounded or 'tooth' like projections (Fig. 118). 100. Cauda pointed (Fig. 118). 101. Lingula present. 102. Circum-anal ring constricted either side of anus (Fig. Ill) or broken into three rings (Livid) (Fig.110). 103. Circum-anal ring constricted either side of anus (Fig. 148) or broken into three rings (Fig. 149). 104. Anal pore-field arranged as bands (Fig. 155). 105. Specialised circum-anal ring (subgenus Baeopelma oiPsylld) present (Fig. 135). 106. Broad circum-anal ring present (Figs 134, 136, 142). 107. Specialised shape of broad circum-anal ring (Fig. 136). 108. Anal pore-field (other than circum-anal ring) arranged as 1 + 1 rings or of derivable form (discountcharacters 102 and 103). Rings placed ventrally or dorso-ventrally, i.e. each ring is partly dorsal andpartly ventral) (Figs 106, 107, 112, 113, 117, 119, 121). 109. Anal pore field (other than circum-anal ring) arranged as 2 + 2 rings or of derivable form. The ringsare arranged ventrally 1 + 1 and dorsally 1 + 1 (Fig. 120). 110. Sectasetae (marginal and dorsal) trunate (Fig. 34). 111. Abdominal sectasetae arranged on large tubercles (Fig. 114). 112. Body margin surrounded by long scales (probably derived from sectasetae) (Fig. 173). 113. Body margin surrounded by broad scales (probably derived from sectasetae) (Figs 174-176). 114. Most abdominal margin sectasetae lost, leaving a distinct arrangement (treated as a gain characterbecause of complexity) of up to 4 + 4 sectasetae. Secondary loss may reduce this number to 3 + 3,2 + 2orl + l(Fig.37). 115. Sectasetae of character 1 14 type truncated to form tubes which are normally based on slight tubercles(Fig.37[ts]). 116. Lanceolate setae (marginal) greatly elongated. 117. Very thin lanceolate setae (assumed to derive from thickened simple setae) present. 118. Capitate setae present on body plus wing-pad margins and dorsal surfaces (Fig. 29). Nymphal 'loss' characters 119. General body form not broader than long (secondary loss of character 84). 120. Thorax dorsal surface with distinct sclerites (at least medials); lateral sclerites small or absent (Figs12-14). 121. Tarsal claws absent. 122. Unguitractor not visible with optical microscope. Arolium present (Figs 56-58). 123. Basal area of arolium reduced to a thin membrane (Figs 60, 64). 124. Basal membrane of arolium absent (derived from character 123) (Fig. 59) (or reduced; Fig. 62). 125. Unguitractor and arolium not visible with optical microscope. 126. Dorsal surface of abdomen lacking distinct sclerites (membrane only anterior to caudal plate area). 127. Anus at posterior of abdomen, not on ventral surface. 128. Abdominal apical teeth absent (secondary loss of 95 and 96). 129. Anal pore-field bands broken into round areas of pores (derived from 104) (Fig. 154). 130. Broad outer area of circum-anal ring broken into round or ovoid areas of pores (Figs 97, 99, 100). 131. Anal pore-field of type described by character 108 broken into round areas of pores (Fig. 113). 132. Anal pore-field of type described by character 108 reduced to narrow bands (Fig. 107). 133. Anal pore-field of type described by character 108 with pores reduced as in Fig. 117. 134. Small groups of pores in abdominal areas such that a reduction of rings as described by character 108may have occurred (used in clades containing character 108 but not 109) (Fig. 119). 135. Small groups of pores in abdominal areas such that a reduction of rings as described by character 108may have occurred (used in clades containing characters 108 and 109) (Fig. 129). 136. Anal pore-field in broken rings, probably derived from character 108, as in Fig. 112. 137. Circum-anal pore ring of type described by character 103 with pores widely separated. 138. Outer circum-anal pore ring reduced to a single row of pores (only applied to clade 82) (Fig. 139). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 255 139. Circum-anal pore ring absent. 140. Anal pore-field absent. 141. Anal pore-field of 1 + 1 rings very reduced (derived from character 134), or absent. 142. Circum-anal pore ring reduced to a few large pores (Fig. 146). 143. Body margin without sectasetae or derivable structures. 144. Body dorsal surface without sectasetae or derivable structures. 145. Body margin (other than abdomen) without sectasetae except for a small number around thehindwing-pad. 146. Abdomen margin without sectasetae (derived from 114). 147. Antennae without sectasetae or derivable structures. 148. Hindwing-pad without marginal sectasetae (derived from character 145). 149. Pointed sectasetae (derived from character 110). 150. Body and wing-pad surfaces (antennae, dorsal body surface and body margin) with lanceolate setae(assumed to derive from reduced sectasetae). 151. Abdominal 4 + 4 (or fewer) sectasetae positions with lanceolate setae (assumed to derive fromcharacter 114) (Fig. 37). 152. Dorsal surface of body and wing-pads with lanceolate setae (assumed to derive from reducedsectasetae). Character only applied when other sectasetae characters are in a derived state. 153. Clavate setae present on body (probably very small sectasetae or lanceolate setae and thereforeregarded as a loss character). 154. Body margin lanceolate setae absent (derived from character 150). 155. Dorsal surface of body without lanceolate setae (derived from character 150). 156. Antennae without lanceolate setae (derived from character 150). 157. Clavate setae absent (derived from character 153). 158. Body margin without capitate setae (except in some species which retain one seta behind each eye)(derived from character 118). 159. Body dorsal surface without capitate setae (derived from character 118). this has been incorporated in a cladogram of these subgenera by Burckhardt (1979). The detailsof clade 82 are largely governed by loss characters and it is very unlikely that any cladogram ofthe subgenera of Psylla, based upon present knowledge, will approximate its true cladistichistory. Host-plant considerations Psyllids are monophagous or narrowly polyphagous and breed almost exclusively upon angio-sperms. Eastop (1972) considered the plant family level relations of 847 species of Psylloidea, ofwhich only 8 were associated with the Monocotyledoneae and the remainder (99%) with theDicotyledoneae. In this study the probable hosts of 298 of the 303 species examined wereknown. Of these, only five species of Livia, on Juncus and Car ex in the Holarctic region, andTrioza palmicola, on an endemic Hawaiian palm, were associated with monocotyledons, andvery few species are associated with annual or biennial herbs. Closely related psyllid species usually occur on closely related host-plants, i.e. psyllid cladesare usually restricted to definite angiosperm taxa (Table 13) (Hodkinson, 1974). Individualspecies of Psylloidea usually occur on host-plants of only one genus and almost exclusively of onefamily. Examples of psyllids breeding on host-plants in separate families are rare. Empirical observation suggests that certain psyllid taxa have a narrow taxonomic distributionof host-plants while others have a broad distribution. It is instructive to examine the taxonomicdistribution of host-plants, for certain psyllid clades, across the 28 plant orders of relevance tothis study. Such an analysis was performed for clades in which at least one terminal taxondescends directly from the ancestor of the clade. The null hypothesis is as follows: 256 I. M. WHITE AND I. D. HODKINSON __ c_ b~d where a = no. psyllid species in clade x associated with plant order y;b = no. psyllids in all clades associated with plant order y;c = no. psyllids in clade x;d = no. psyllids in all clades. The deviation from the regular distribution was measured by the Kolmogorov-Smirnovtwo-sample test (with two tails of significance). This was converted to a x value, by anapproximation, with two degrees of freedom (Siegel, 1956). It is expected that x 2 is underesti-mated for any clade with less than 40 species, that is all except 2 and 53, which makes the testconservative (Siegel, 1956), i.e. the significance level may be underestimated. This test (Table14) indicates that most clades have a taxonomic distribution of host-plants which is significantlynon-regular. The variance (s 2 ) and the mean (x) number of psyllid species in each of the 28host-plant orders were calculated for each clade shown to depart significantly from a regulardistribution. These values were expressed as a ratio (Table 14) which is a measure of dispersion,such that the greater the value the more clumped the host-plant distribution. Clades with verysmall variance-mean ratios, and with large sample sizes (more than 10 psyllid species) are clade51, e.g. Diaphorina, and clade 2, e.g. Trioza. The clade with the most clumped, that is mostrestricted host distribution, is clade 53, e.g. Psylla. This does not imply that genera such as Trioza lack distinct groups feeding upon relatedgroups of plants; for example one subgroup of Trioza is exclusively associated with the plantgenus Salix (Salicaceae). Taxa such as clade 53, e.g. Psylla, of which 49% feed on Resales, andmost of those on Fabaceae, differ from taxa such as clade 2 (e.g. Trioza) in that distinct hostassociations exist at a suprageneric rather than subgeneric level. If the cladogram roughlyrepresents the true cladistic history of the Psylloidea, then in clade 53 (e.g. Psylla) morphologi-cal divergence exceeds host-plant choice divergence. However, in clade 2 (e.g. Trioza)host-plant choice has undergone more evolutionary changes than morphological form. The cladogram was assumed to be a true record of the cladistic history of the Psylloidea and anattempt was made to find the most parsimonious fit of the host relationships to the cladogram. Clade 2 (Fig. 186) (e.g. Trioza) is a large highly polyphagous taxon. Widely separate branchtips feed on plant taxa such as Annonales, Moraceae and Salicaceae. At this stage no hypothesiscan be made about the ancestral host of clade 2. Clade 4 (Fig. 187) is associated with Malvales (e.g. Paracarsidara) and Rutales: Meliaceae(Mastigimas). Either of these plant groups could represent the ancestral host of clade 4. Clade 6 (Fig. 188) is associated exclusively with Ficus (Moraceae), the most likely ancestralhost of clade 6. Clade 8 (Fig. 189) is associated with two families of Rutales, i.e. Burseraceae and Meliaceae.The host of Epicarsa, however, is unknown. Clade 10 (Fig. 190) is associated with the Rutales (mainly Anacardiaceae, plus Burseraceaeand Rutaceae), e.g. Calophya. It is now reasonable to suggest that Rutales-feeding is an ancestral feature retained by disjunctgroups of the above clades and, by the parsimony criterion, is the most likely ancestral host ofclade 9. The association with Ficus evolved with clade 6, with Malvales in clade 32 and theancestral host of clade 2 remains unknown. In the remaining branches of the cladogram Liviaand Strophingia are associated with Commelinales (Cyperaceae and Juncaceae) and Ericaceaerespectively; none feeds on Rutales. Clade 13 (Fig. 191) contains several groups with distinct host relations: Phytolyma onMoraceae, Gyropsylla on Ilex (Aquifoliaceae) and Nectandra (Lauraceae), clade 42 on Tamarixand Myricaria (Tamaricaceae) and clade 40 on herbs. Clade 40 (Aphalara and Craspedolepta)has several distinct groups of species restricted to certain families or genera of plants. Aphalaralive on Brassicaceae, Polygonaceae and Ranunculaceae, while Craspedolepta are associated NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 257 Table 13 Psyllid taxa which, in the present study, are restricted to specified taxa of angiosperms. Anasterisk marks entries which are known to occur on other angiosperm taxa, when psyllid species notcovered in the present survey are considered. i. Psyllid taxa restricted to plant orders. TAXON HOST ORDER Clades 8, 15Clade 32Livia ii. Psyllid taxa restricted to plant families.TAXON HOST FAMILY Rutales Malvales Commelinales HOST ORDER with Asteraceae, Chenopodiaceae and Onagraceae. The ancestral host of clade 13 is uncertainand none feeds on Rutales. Species in clade 15 (Fig. 192) feed exclusively on Rutales and the ancestral host is assumed tobe a species of Rutales. Clade 17 (Fig. 192) is divided into two major taxa; clades 48 and 53. Clade 8 (Fig. 193) has a high diversity of host relationships; clade 58 on Ericaceae, Euphylluraon Oleaceae, clade 61 on Myrtaceae, Onagraceae and Rutaceae, clade 62 on Sterculiaceae andMelastomataceae, Paurocephala on Malvaceae and Moraceae, Camarotoscena on Salicaceae,Psyllopsis on Oleaceae, Diaphorina on several families (e.g. Rutaceae and Solanaceae), andPennavena on Loganiaceae. The ancestral host is most likely to be a plant taxon associated withmore than one branch tip, i.e. Malvales (Malvaceae and Sterculiaceae), Oleaceae or Rutales(Rutaceae). Clade 53 (Fig. 193) is associated with Resales except for: Arepuna (Solanaceae), Euphalerusjugovenosus group (p. 224) (Rhamnaceae), Phellopsylla (Myrtaceae), clade 73 (Ulmaceae), E.gallicolus (Rhamnaceae), clade 68 (Myrtaceae), Trigonon (host unknown), Freysuila sp.(Solanaceae), one Acizzia sp. (A. hakeae, Proteaceae), Anomoneura (Moraceae), someInsnesia spp. (Euphorbiaceae), and many species in clade 82 (e.g. Betulaceae, Rhamnaceae andSalicaceae). Of those species on Resales, most (63%) are associated with Fabaceae. 258 I. M. WHITE AND I. D. HODKINSON Table 14 Values of x 2 approximation to Kolmogorov-Smirnov test, variance/mean ratio, and mostfavoured host-plant order, of selected clades. Significance levels: *** P< 0-001, ** P<0-01 &*P + 0-05. Only Euphalerus tantillus (Rosaceae) and most members of clade 82 (Buxaceae, Rosaceaeand Saxifragaceae) are not. It is assumed that Fabaceae-feeding is ancestral to clade 53. Nospecies occur on Rutales. From Fabaceae host changes to Myrtaceae, Rhamnaceae, Rosaceaeand Solanaceae must have occurred more than once. The evidence suggests that this hasoccurred repeatedly in different zoogeographic regions. Among the remaining clades, Rutales-feeding occurs exclusively in clade 15 and may also bethe ancestral host of clade 48. Rutales-feeding therefore occurs commonly in disjunct cladesthroughout the cladogram, and application of the parsimony criterion suggests that the ancestralPsylloidea are associated with plants of the order Rutales or a direct ancestral group to theRutales. There is evidence, based on a belief that the Rutaceae and Anacardiaceae appeared earlyenough for direct migration to Australia, that the Rutales evolved at least 95 million years ago(Raven & Axelrod, 1974). If angiosperm-feeding in psyllids evolved only once then initially thiswas most likely to have been in conjunction with primitive Rutales, possibly prior to differentia-tion of the host-plant families Aceraceae, Anacardiaceae, Burseraceae, Meliaceae andRutaceae. Much of the primary differentiation of the Rutales seems to have taken place inAfrica-South America, with long standing connections to Eurasia (Raven & Axelrod, 1974). The clades which are restricted to a single plant taxon other than Rutales are as follows: clade3 (e.g. Carsidard) on Malvales and clade 6 (e.g. Homotoma) on Ficus. Examples of large cladeson a diverse range of plants are: clade 2 (e.g. Triozd), clade 13 (e.g. Aphalard) and clade 17 (e.g.Paurocephala, Spondyliaspis and Psylld). Zoogeographic evidence A vicariance approach (Platnick & Nelson, 1978) was applied to fit zoogeographic evidence tothe cladogram. The model was restricted to the time period since the earliest appearance of theangiosperms (125 million years before present or 125 m.y.B.P. ; Raven & Axelrod, 1974), and itwas assumed that the modern psyllids have evolved since the splitting of Pangea into Laurasiaand Gondwanaland (180 m.y.B.P.). Therefore any track or distribution which includes areas ofboth Laurasia and Gondwanaland is assumed to have been caused by a dispersal event.Furthermore, dispersal is also assumed to account for the presence of psyllids on oceanic islandsas this is a more tenable explanation of such tracks than the assumption of an as yet unknownvicariance event (Cracraft, 1975). Additionally some Nearctic-Palaearctic tracks may be better NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 259 crLU Q_ co I Q 5 | CO cr LUo 260 I. M. WHITE AND I. D. HODKINSON explained by dispersal across Beringia than by vicariance of Laurasia (Hodkinson, 1980). Thevicariance/dispersal tracks are shown as a reduced geological area cladogram, based on Rosen(1978) (Fig. 197). Each section of the psyllid cladogram (Figs 185-196) was compared to the geological areacladogram (Fig. 197). As an example of the reasoning applied, consider a hypothetical cladeoccurring in the Afrotropical, Malagasy, Oriental, Austro-Oriental and Palaearctic regions.The Palaearctic is not joined by a vicariance track in Fig. 197 to any of these other regions.Dispersal to the Palaearctic is proposed. The Austro-Oriental region must also be reached bydispersal. The Afrotropical, Malagasy and Oriental regions can be seen to have originated fromone biota (Fig. 197). The ancestral species of this hypothetical clade is, therefore, assumed tohave been distributed in the Afrotropical-Malagasy-Oriental biota, i.e. this is the ancestral trackand distribution. Other solutions are less parsimonious. Extinctions must occasionally be proposed to explain disjunct patterns of distribution.However, 'not yet discovered' may be the correct interpretation in many cases. The full analysisfor each major clade is given by White (1980). The general conclusions to be drawn from this analysis are as follows: 1. Most major clades probably had a Gondwanaland origin, i.e. clade 2 (Fig. 186) (e.g. Trioza),clade 4 (Fig. 187) (e.g. Paracarsidara), clade 6 (Fig. 188) (e.g. Homotoma), clade 8 (Fig. 189)(e.g. Phacopterori) , clade 10 (Fig. 190) (e.g. Calophya), clade 13 (Fig. 191) (e.g. Gyropsylla),clade 15 (Fig. 192) (e.g. Tainarys) and clade 17 (Fig. 193) (e.g. Paurocephala and Euphalerus). 2. The ancestor of the Psylloidea probably had a Gondwanaland track. This is consistent withthe host-plant evidence, i.e. that the ancestral host was a species of Rutales and that this group ofplants must have been distributed throughout Gondwanaland prior to its breakup. In the geological area cladogram no allowance was made for the possibility of dispersal acrossthe South Atlantic between the Afrotropical and Neotropical regions. Raven & Axelrod (1974)review the evidence for such a dispersal between the two continents, as they are thought to haveremained in near contact until at least 90 m.y.B.P. This dispersal route existed before thebreakup of the Australian- Antarctic-South American continent (45 m.y.B.P.). Any taxonpresent in both the Afro-Oriental biota and the Neotropical region could be explained bydispersal across the Atlantic or by vicariance of a Gondwanaland track followed by extinction inNew Zealand and Australia. Because of this choice of explanations the vicariance model wasadhered to purely as a convention. Australia was probably quite humid at a time when it was stilljoined to South America (Frakes & Kemp, 1974), and the subsequent lowering of humidity mayaccount for many extinctions. As the distribution patterns of most clades can be explained largely by vicariance eventsoccurring during the breakup of Gondwanaland, the minimum, and sometimes maximum, ageof many of the major clades can be determined from the estimated dates of vicariance events. Clade 4 (Fig. 187) (e.g. Paracarsidara) had a Gondwanaland ancestor (90-1 80 m.y.B.P.). Theinitial host was a species of Rutales. Ancestors 30, 31 and 32 also appeared to have evolvedduring this time in association with Mai vales. The origin of the Malvales seems uncertain. Raven& Axelrod (1974) state that the primary radiation of Malvales probably took place in Africa andSouth America in Maastricktian time (65-70 m.y.B.P.) or earlier. There are, however, somedoubtful Upper Cretaceous (65-110 m.y.B.P.) macrofossils of Malvaceae (Raven & Axelrod,1974). The origin of clade 32 remains uncertain. Clade 6 (Fig. 188) (e.g. Homotoma) also appears to have originated in Gondwanaland andthese taxa are all associated with Ficus (Moraceae). Raven & Axelrod (1974) say that Moraceaewere probably in existence early enough to have been dispersed more or less directly betweenAfrica and South America. Ancestor 6 may have had an Afro-Indian range and been dispersedto South America (Synoza) and later from India to Australia (Mycopsylla) by island hopping.Whichever route was taken ancestor 6 must have existed at least 90 m.y.B.P. Clades 8, 10 and 15 (Figs 189, 190, 192) are all associated with the Rutales and all appearto have a Gondwanaland distributed ancestor. These groups must each have been distinct by 90m.y.B.P. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 261 Ancestors 13 and 17 (Figs 191, 193) are of unknown host relations and each probably had aGondwanaland distribution. These groups must also have been distinct by 90 m.y.B.P. In clade17 the major host preference is for Resales, especially Fabaceae (clade 53). However, Raven &Axelrod (1974) imply that the Fabaceae were of later origin (c. 65 m.y.B.P.) although theRosales probably existed much earlier. Despite several problems a number of major conclusions can be drawn. Firstly that mostmajor clades have a distribution which is consistent with a southern ancestry, probably prior tothe breakup of Gondwanaland (90 m.y.B.P.). Furthermore, the flowering plants, typified byAnnonales, probably evolved about 125 m.y.B.P. (as indicated by data reviewed by Raven &Axelrod, 1974). The modern psyllids probably evolved with the Rutales (p. 258) and, therefore,later than 125 m.y.B.P. but earlier than 90 m.y.B.P. (although Togepsylla, a morphologicallyvery primitive psyllid associated with Annonales, may be a relic member of a group antedatingancestor 1). Conversely, the following major groups probably evolved from ancestors in the northern landmass of Laurasia: Livia, Strophingia in the Palaearctic, clade 40 (e.g. Aphalara), clade 81 (e.g.Psylla) and clade 73 (e.g. Pachypsylld). This is assuming that their ancestors dispersed toLaurasia from Gondwanaland. An alternative hypothesis is available if a pure vicariance modelis adopted, i.e. that the ancestral psyllid was distributed throughout Pangea. If this were the casethen we must accept that modern psyllids diversified before angiosperms evolved and psyllidstherefore moved onto angiosperm hosts on several separate occasions, remarkably, often ontothe same group of plants, namely the Rutales. On balance, our initial hypothesis that modernpsyllids only evolved since the appearance of the angiosperms, and therefore since the splittingof Pangea into the southern Gondwanaland and northern Laurasia, seems more tenable. Classification and phytogenyNymphal phenetic classification Several phenetic classifications have been presented which provide different summaries of theresemblances between taxa. It now becomes necessary to identify the common factors andproduce a SUMMARY classification. Analyses based upon character resemblance, such as principal component and SUMRATinformation statistic, indicated the characters of greatest importance in forming a pheneticclassification of nymphs. The species groups defined by the presence of each of these 'important'characters coincide with the major groups formed in the minimum spanning networks, pheno-grams and principal component analyses. Therefore the listing of these species groups forms asummary phenetic classification. The relationships of the phenetic groups, as defined by the 'important' characters, are shownin Fig. 198 and the approximate positions of the groups and subgroups, of the summaryclassification, are shown relative to principal components I and II (Fig. 199). This provides avisual representation of the between group relationships. A list of the species in each group,together with the initial letter of the family to which the taxon belongs in the classification ofBecker-Migdisova (1973): Aphalaridae (A), Carsidaridae (C), Liviidae (L), Psyllidae (P),Spondyliaspididae (S) and Triozidae (T), is given in Table 15. Previous phenetic classifications of psyllid nymphs (Ferris, 1925; Rahman, 1932) were basedon wing-pad shape. The only similarity between these and the above classification is that the'truncate sectasetae' subgroup roughly corresponds to the 'triozine' group of Ferris andRahman. The percentage of species examined from each of the Becker-Migdisova (1973) families,relative to the new summary classification, is shown in Table 16. The most highly congruentgroup is 3 (capitate setae) plus subgroups l.ii (truncate sectasetae) and 4.i (petiolate arolium).These groups are defined by gain characters and correspond to some Psyllidae, Triozidae andthe remaining Psyllidae respectively. By contrast, group 2 (lanceolate setae) and subgroup 4.ii(remainder) are characterised by loss characters. Such characters would be expected to form 262 I. M. WHITE AND I. D. HODKINSON LU LUCO LU < 2 crLU I SUBGROUP GROUP Fig. 198 Phenogram of groups and subgroups of species defined by the summary phenetic classification,based on nymphal characters. evolutionary convergent clusters. Group 2 and subgroups 4.ii and l.i (pointed sectasetae) areleast congruent with the families of Becker-Migdisova. Pointed sectasetae are probably theancestral character state and, therefore, subgroup l.i contains those species which have retainedan ancestral feature. Empirical taxonomic studies of psyllid classification have failed to find stable positions forcertain genera; for instance, the suggested relationships between Paurocephala and Pauropsyllaare radically different in the classifications of Crawford (1914), Becker-Migdisova (1973) andLoginova (1972). In this study, these problematical groups again tended to cluster in differentpositions in different analyses. The most stable groups in each analysis, group 3 (species withcapitate setae) and subgroup l.ii (species with truncate sectasetae), are highly congruent withthe most stable families recognised by empirical taxonomy, that is the Psyllidae and Triozidaerespectively. Numerical phenetic methods were, therefore, of little direct value in the placementof problem groups in a new general classification. However, such methods did indicate nymphalgroupings which might not be predicted from the existing empirical adult classifications.Furthermore, numerical phenetics were particularly relevant to ground plan construction forcladistic analysis and for the recognition of characters with the greatest classificatory power suchas the form of the nymphal tarsal arolium. This was found to have far greater power than hadbeen empirically expected and upon re-examination was found to be one of the most usefulcharacters for later cladistic analysis. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 263 PC II PC I Fig. 199 Groups and subgroups in the summary classification, placed relative to principal components Iand II. General classification The most convincing argument in favour of cladistic analysis as a basis for a general referenceclassification is provided by Mickevich (1978), who compared congruence of phenetic versuscladistic classifications. The latter were found to have greater stability, probably because theywere less sensitive to the adverse effects of heterogeneity in the evolutionary rates of charactersand, furthermore, they were the most predictive (Platnick, 1978). However, a truly predictiveclassification, based upon a cladogram, may be impractical if the taxon (clade) to which a speciesbelongs can only be determined from a life-cycle stage which may be unavailable. Therefore, asthe nymphs of most species are unknown and many clades are defined by attributes of onelife-cycle stage only, it is necessary to produce a compromise 'practical-predictive' classification.For example, clade 13 (Fig. 191, p. 224) (e.g. Aphalara) and clade 15 (Fig. 192) (e.g. Rhinocold)are defined by an adult and a nymphal attribute and are impracticable in a nymphal and an adultclassification respectively. If predictability is to be retained some impractical groups have to betolerated. However, for practical reasons more emphasis has occasionally been placed on adultrather than nymphal characters in finally deciding the position of a taxon. In deriving the general classification monophyletic groups were preferred. No polyphyleticgroups, in the sense of Hennig (1966) or Farris (1974), were formed and paraphyletic groups, ofHennig (1966) and Farris (1974), were allowed if they increased practicality in identifying adults. 264 I. M. WHITE AND I. D. HODKINSON This combination of both cladistic and phenetic information is, in general principle, the'evolutionary' method of Mayr (1969). The suggested classification (Table 17) includes eight families of which three are new. Theprobable positions of some taxa not examined are included and these are marked by an asterisk.A major source of information for the inclusion of these additional taxa was Loginova (1964b).The probable family or subfamily to which the residual genera belong are listed in Table 18. Aphalaridae This is probably a paraphyletic group, in the sense of Hennig (1966) and Farris (1974), butpolyphyletic according to Nelson (1971). It comprises species which are phenetically close to theground plan of the Psylloidea: clade 1 (Fig. 185) minus clades 9 (Fig. 185) and 54 (Fig. 193). Itwould be impractical to make each whole clade a separate monophyletic family because most aredefined only by derived nymphal attributes and adults could only be assigned to such familieswhen accompanied by nymphs of the same species. This is still a problem at the subfamily leveland only polyphyletic subfamily groupings within Aphalaridae would form a practical classifica-tion. The content of the family is similar to the Aphalaridae of Becker-Migdisova (1973) but withthe addition of the Diaphorininae (from Psyllidae of Becker-Migdisova), Ctenarytainini (fromSpondyliaspididae) and Liviinae (formerly Liviidae). The Diaphorininae and Ctenarytainini areincluded in the Aphalaridae largely on the basis of nymphal features. Phenetically the Liviinaeare distant from other Aphalaridae but the relationship of the single genus Livia is bestillustrated by placing it within the family. Spondyliaspididae This family is probably a paraphyletic group (in all senses) (clade 54 (Fig. 193) minus clade 57).However, it is only the genus Arepuna which falls outside of a probable monophyletic grouping(clade 55, Fig. 195). Arepuna spp. have a 'Euphalerus' adult facies and it would, therefore, beimpractical to place this genus in a separate family to the genus Euphalerus. Psyllidae This is a probable monophyletic group (clade 57, Fig. 196). Five subfamilies are tentativelyproposed, based upon clades which could only be defined by loss characters. This family is thePsyllidae of Becker-Migdisova (1973) minus the Diaphorinini and Psyllopseini which are nowplaced in the Aphalaridae: Diaphorininae, and the Euphalerini which are now in the Spondy-liaspididae. Loginova (19760, 1977) proposed a tribe Cyamophilini, which includes Amorphicola.Nymphs of Cyamophila, the type-genus, were not examined and it is possible that Cyamophilabelongs close to Amorphicola in the cladogram. Therefore, no name is proposed for a tribecontaining Amorphicola. Calophyidae This family is a possible monophyletic group (clade 10, Fig. 190), although Apsylla is onlytentatively included. All other genera belong to the subfamily Calophyinae (clade 39). Manyspecies at present referred to the genus Pauropsylla probably belong to the Calophyinae (p. 241). Phacopteronidae A family which is a possible monophyletic group (clade 8, Fig. 189). Bharatiana is only provisionally included and all other genera belong to the subfamily Phacopteroninae (clade 37). Homotomidae This is the subfamily Homotominae of Becker-Migdisova (1973) and a probable monophyletic group (clade 6, Fig. 188). Carsidaridae This family is not the Carsidaridae of Becker-Migdisova (1973) as the following genera havebeen placed elsewhere in the present classification: Diclidophlebia and Togepsylla (Aphalar-idae), Calophya, Microceropsylla and Pelmatobrachia (Calophyidae), Homotoma, Macro- NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 265 homotoma, Mycopsylla, Pseudoeriopsylla and Synoza (Homotomidae), Bharatiana, Epicarsa,Phacopteron and Pseudophacopteron (Phacopteronidae) plus Leptynoptera and Pauropsylla(Triozidae). As now defined, the Carsidaridae is a probable monophyletic group (clade 4, Fig.187). The genus Mastigimas, however, is only tentatively included and all other genera form theCarsidarinae (clade 32). Unfortunately, nymphs of the type-genus, Carsidara, were notincluded in the study but the predictive properties of the cladogram suggest that the nymph ofCarsidara should be of the type found in clade 4 which is thus referred to the existing familyCarsidaridae. Triozidae This is a probable monophyletic group (clade 2, Fig. 186) which is the Triozidae of Becker-Migdisova (1973) plus Leptynoptera and Pauropsylla from her Carsidaridae. The tribe Trioziniis probably a paraphyletic group (clade 21 minus clade 25) which requires much further study ofgeneric limits before the classification can be improved. Possible phytogeny It is conventional to illustrate a phylogeny as a lateral view of a tree diagram (Fig. 200). Theancestral group is extinct and in the absence of fossil evidence the information required to makemore than a tentative estimate of the branching sequence is unavailable. It is more informative to illustrate a terminal cross section of the phyletic tree as an unresolvedBUSH PHYLOGENY (Thorne, 1976) (Fig. 201). Such a phylogeny is said to be unresolved, as noattempt is made to show the exact sequence of branching. The Ancestral Group comprises extinct species which probably had a Gondwanalanddistribution, fed on Rutales and evolved 90-125 million years before present. A southernancestry for the psyllids has also been suggested by Eastop (1978) and Hodkinson (1980).Klimaszewski (1964), however, believed that psyllids evolved in South East Asia which,according to Takhtajan (1969), is the 'cradle of the angiosperms'. South East Asia (theAustro-Oriental region) only came into existence in the Miocene with the arrival of theAustralian plate in the vicnity of Asia, and angiosperms could not have originated there (Raven& Axelrod, 1974) and, by the same logic, neither could the psyllids. Unfortunately most of thefossils of insects resembling psyllids, as reviewed by Szelegiewicz (1971), antedate the angio-sperms. Furthermore, their morphology suggests that they were not on the direct line of descentto the modern Psylloidea. However, there are a few genera which may be as closely related tothe ancestral group as they are to any other extant taxa. They include Apsylla (Calophyidae),Bharatiana (Phacopteronidae), Mastigimas (Carsidaridae) and Strophingia (Aphalaridae) and,with the exception of Strophingia (on Ericales), they are all Rutales-feeders. The most primitive(Cronquist, 1968; Takhtajan, 1969; Thorne, 1976) and thereby the probable ancestral group ofangiosperms (Takhtajan, 1969) are thought to be the Annonales. Togepsylla (Aphalaridae),which feeds on Lauraceae (Annonales) , may possibly be a relic genus of a psyllid group whichantedates the Rutales-feeders. CALOPHYIDAEPHACOPTERONIDAEHOMOTOMIDAECARSIDARIDAE TRIOZIDAE SPONDYLIASPIDIDAEPSYLLIDAE Fig. 200 Suggested phylogenetic relationships of the families of Psylloidea; as a tree with more than oneline leading to the paraphyletic families Aphalaridae and Spondyliaspididae. 266 I. M. WHITE AND I. D. HODKINSON Fig. 201 Suggested phylogenetic relationships of the families of Psylloidea; a cross section of theunresolved bush phylogeny, in which each family (solid border) and subfamily (broken border) is showncovering an area roughly in proportion to the number of species it contains. The Aphalaridae is a collection of five phyletic lines, Strophingiinae, Liviinae, Aphalarinae,Rhinocolinae and Paurocephalinae which originate from close to the probable ancestor. TheStrophingiinae are only separated from the ground plan by loss characters. The Liviinaeprobably had a Laurasian ancestor associated with the Commelinales but further evidencesuggesting the origin of the group is unavailable. The Aphalarinae, by contrast, were most likelyto have had a Gondwanaland ancestor, although the greatest diversification of species andgenera has occurred in northern regions. However, their host-plants are diverse and do notinclude Rutales. It is conceivable that if Annonales-feeding antedates Rutales-feeding, theancestor of Aphalarinae was an Annonales-feeder. At least one extant member of the family,Gyropsylla cannela (Crawford) on Lauraceae, is associated with Annonales. The Rhinocolinae probably had an ancestor with a Gondwanaland distribution. All membersof the subfamily examined feed on Rutales. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 267Table 15 Nymphal summary classification 1. Sectasetae group. Taxa with pointed or truncate sectasetae (characters N26 to N33, Table 4, p. 207). The forewing-pad generally has a well-formed humeral lobe (Nl) and the tarsal arolia are very rarely petiolate (N4).There are two subgroups. i. Pointed sectasetae subgroup. Taxa with pointed sectasetae (characters N26 to N33) Calophya (most spp . ) C Camarotoscena unicolor A Crawforda triopsyllina T Dididophlebia eastopi C Egeirotriozaspp. T Homotomaspp. C Leptynoptera sulfurea C Leuronota michoacana T Moraniella calodendri A Paraphalaroidafremontiae A Paurocephala spp. A Synozapulchra C Trioza alacris T Triozoida silvestris T ii. Truncate sectasetae subgroup. Taxa with truncate sectasetae (characters N26 to N33). Acanthocnema casuarinae T Ceropsylla martorelli T Paratriozaspp. T Pauropsylla trichaeta C Togepsylla matsumurana C Trichochermes walkeri T Trioza (most spp . ) T 2. Lanceolate setae group. Taxa with lanceolate setae (characters N23 to N25). There is often a humeral lobe (Nl) and the tarsalarolia are rarely petiolate (N4). Agonoscenaspp. A Aphalaraspp. A Bharatiana octopsinosa C Camarotoscena speciosa A Colposceniasp. A Craspedoleptaspp. A Crastina linavuorii A Ctenarytaina eucalypti S Diaphorina spp. P Epicarsa sp. C Eucalyptolymasp. S Euphylluraspp. A Leurolophus vittatus A Livia crefeldensis L L. v emails L Neophylluraspp. A Pennavenafabulosa P Phacopteron lentiginosum C Phellopsyllasp. S Phytolyma (most spp.) A Pseudoeriopsylla nyasae C Pseudophacopteronfloccosa C Psyllopsisspp. P 268 I. M. WHITE AND I. D. HODKINSON Rhinocola aceris A Strophingiaspp. A Tainarys schini A 3. Capitate setae group. Taxa with capitate setae (Nil to N19). All of these species also have a petiolate tarsal arolium (N4). Acizzia hakeae P A. russellae P A. uncatoides P Amorphicola amorphae P Arytaina genistae P Arytainillaspp. P Ceanothiaspp. P Ciriacremum spp. P Euceropsyllaspp. P Euphalerus tantillus P E. sp. (B). P Floria variegata P Freysuila sp. P Heteropsylla spp. P Insnesia glabruscuta P Isogonoceraia divergipennis P Mitrapsylla deserata P Ps_y//fl (most spp.) P Purshivora spp. P Trigonon longicornis P 4. Other taxa. Taxa which lack sectasetae (characters N26 to N33), lanceolate setae (N23 to N25) and capitate setae(Nil to N19). Two subgroups may, however, be recognised by the presence or absence of the petiolatetarsal arolium, a character which received high eigenvector and SUMRAT values,i. Petiolate arolium subgroup. The following taxa have a petiolate tarsal arolium. Acizzia acaciae P A. acaciaebaileyanae P Anomoneura mori P Aphalaroidaspp. A Arepunasp. P Colophorina cassiae P Epipsyllaspp. P Euglyptoneura spp. P Euphalerus (most spp. ) P Neopsy Ilia spp. P Pexopsylla cercocarpi P Platycorpha princeps P Psylla betulaenanae P P. carpinicola P P. floccosa P P. galeaformis P P. mali P P. phoradendrae P P. ribesiae P P. sto'ata P P. trimaculata P Retroacizzia antennata P Spanioneurafonscolombii P ii. Remainder. Apsylla cistellata A NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 269 Calophya rhois C C. rotundipennis C Cardiaspina spp. S Ceropsylla sideroxyli T Creiissp. S Euphalerus gallicolus P Glycaspis spp. S Gyropsyllaspp. A Hevaheva swezeyi T Kuwayamapisonia T Livia (most spp.) L Macrohomotoma spp. C Mastigimas spp. C Mesohomotomaspp. C Microceropsyllasp. C Mycopsyllasp. C Neolithussp. T Pachypsyllaspp. S Paracarsidaraspp. C Pauropsylla (most spp . ) C Pelmatobrachiasp. C Phytolyma lata A Protyora sterculiae C Pseudophacopteron (most spp.) C Spondyliaspissp. S Swezeyana elongagena T Synozosp. C Tenaphalara spp. C Tetragonocephalasp. S Trioza anceps T 7". hirsuta T 7. palmicola T Triozamia lamborni T There are four remaining subfamilies of Aphalaridae; Paurocephalinae, Euphyllurinae,Diaphorininae and Aphalaroidinae. The Paurocephalinae are morphologically most primitiveand, hence, this subfamily is illustrated (Fig. 201) as deriving from the ancestral group and givingrise to the other three subfamilies. The Paurocephalinae, Euphyllurinae and Diaphorininaeprobably had Gondwanaland origins associated with unknown hosts. The ancestor of the extant Aphalaroidinae was most likely to have had a Gondwanalanddistribution, associated with Fabaceae. From such an ancestor the present day Nearctic genusAphalaroida evolved together with the ancestor of the Spondyliaspididae. Table 16 Percentage of species in each family of Becker-Migdisova (1973) in each group or subgroup ofthe nymphal summary classification (rows total 100%). Group or subgroupl.i. l.ii. 2. 3. 4.i. 4.ii. 270 I. M. WHITE AND I. D. HODKINSON Table 17 A general classification of the Psylloidea. The following classification includes eight families. An asterisk indicates the probable position in the classification of some taxa not examined. Psylloidea Low Aphalaridae LowTogepsyllinae Becker-Migdisova Togepsylla Kuwayama*Strophingiinae subfam. n. (type-genus: Strophingia Enderlein) Strophingia EnderleinLiviinae Low Livia LatreilleAphalarinae LowPhytolymini Becker-Migdisova Phytolyma ScottGyropsyllini trib. n. (type-genus: Gyropsylla Brethes) Gyropsylla BrethesColposceniini Becker-Migdisova Colposcenia Enderlein Crastina LoginovaAphalarini Low Aphalara Forster Craspedolepta Enderlein Brachystetha Loginova* Epheloscyta Loginova* Xanioptera Enderlein*Caillardiini Loginova* Caillardia Bergevin* Eumetoecus Loginova* Rhodochlanis Loginova* Rhombaphalara Loginova*Xenaphalarini Loginova* Eurotica Loginova* Xenaphalara Loginova*Rhinocolinae Becker-MigdisovaRhinocolini Becker-Migdisova Tainarys Brethes Leurolophus Tuthill Moraniella Loginova Rhinocola Forster Agonoscena Enderlein Aphorma Hodkinson* Lisronia Loginova* Rhachistoneura Hodkinson & Hollis*Pachypsylloidini Loginova*- Acaerus Loginova* Eremopsylloides Loginova* Pachypsylloides Bergevin*Paurocephalinae Becker-Migdisova Camarotoscena Haupt Paurocephala CrawfordEuphyllurinae Becker-MigdisovaDiclidophlebiini Becker-Migdisova Diclidophlebia Crawford Paraphalaroida Loginova Haplaphalara Uichanco*Euphyllurini Becker-Migdisova Euphyllura Forster NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 271 Neophyllura Loginova Katecephala Crawford* Ligustrinia Loginova* Syntomoza Enderlein* Syringilla Loginova*Ctenarytainini trib. n. (type-genus: Ctenarytaina Ferris & Klyver) Ctenarytaina Ferris & Klyver (some spp. referred to Eucalyptolyma} Eurhinocola Crawford* Syncarpiolyma Froggatt*Diaphorininae VondracekDiaphorinini Vondracek Diaphorina Low Pennavena Capener Eudiaphorina Loginova*Psyllopseini Vondracek Psyllopsis LowAphalaroidinae Loginova Aphalaroida CrawfordSpondyliaspididae SchwarzArepuniinae subfam. n. (type-genus: Arepuna Tuthill) Arepuna TuthillEuphalerinae Becker-Migdisova Euphalerus Schwarz Retroadzzia Heslop-Harrison TPachyparia Loginova* Phellopsylla Taylor Colophorina Capener Cometopsylla Froggatt*Pachypsyllinae Becker-Migdisova Pachypsylla Riley Tetragonocephala CrawfordSpondyliaspidinae Spondyliaspis Signoret Creiis Scott Cardiaspina Crawford Glycaspis Taylor Australopsylla Tuthill & Taylor* Eucalyptolyma Froggatt* (not including species examined in this study) Hyalinaspis Taylor* Lasiopsylla Froggatt*Psyllidae Low Acizziinae subfam. n. (type-genus: Acizzia Heslop-Harrison)Acizziini trib. n. (type-genus: Acizzia Heslop-Harrison) Trigonon Crawford Mitrapsylla Crawford Acizzia Heslop-Harrison Platycorypha Tuthill Neopsyllia Caldwell Freysuila AlemanMacrocorsini Becker-Migdisova* Auchmerina Enderlein* Caradocia Laing* Geijerolyma Froggatt* Macrocorsa Vondracek*Anomoneurinae Becker-MigdisovaAnomoneurini Becker-Migdisova Anomoneura Schwarz Epipsylla Kuwayama 272 I. M. WHITE AND I. D. HODKINSON Tribe - unnamed (may be Cyamophilini Loginova) Amorphicola Heslop-Harrison (many species referred to Euphalerus) ICyamophila Loginova*Ciriacreminae EnderleinCiriacremini Enderlein Euceropsylla Boselli Heteropsylla Crawford Insnesia Tuthill Isogonoceraia Tuthill Ciriacremwn Enderlein Aremica Tuthill* Delina Blanchard* Kleiniella Aulmann* Palmapenna Hollis* Panisopelma Enderlein* Russelliana Tuthill*Arytaininae Crawford Arytaina Forster Floria Low Arytainilla Loginova Alloeoneura Low* Amblyrhina Low* Livilla Curtis*Psyllinae Low Psylla Geoffroy sensu lato (including all subgenera) Spanioneura Forster Ceanothia Heslop-Harrison Euglyptoneura Heslop-Harrison Purshivora Heslop-HarrisonCalophyidae Vondracek stat. n.Apsyllinae Becker-Migdisova Apsylla CrawfordCalophyinae Vondracek Pelmatobrachia Enderlein Microceropsylla Boselli Calophya Low Holotrioza Brethes* Paracalophya Tuthill*Phacopteronidae Becker-Migdisova stat. n.Bharatianinae subfam. n. (type-genus: Bharatiana Mathur) Bharatiana MathurPhacopteroninae Becker-Migdisova Phacopteron Buckton Pseudophacopteron Enderlein Epicarsa Crawford Chineura Tuthill* Phacosemoid.es Lima & Guitton*Homotomidae Heslop-Harrison stat. n.Homotominae Heslop-Harrison Homotoma Guerin-Meneville Synoza Enderlein Macrohomotominae subfam. n. (type-genus: Macrohomotoma Kuwayama)Mycopsyllini trib. n. (type-genus: Mycopsylla Froggatt) Mycopsylla FroggattMacrohomotomini trib. n. (type-genus: Macrohomotoma Kuwayama) Macrohomotoma Kuwayama Pseudoeriopsylla Newstead NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 273 Carsidaridae CrawfordMastigimatinae Becker-Migdisova Mastigimas EnderleinCarsidarinae CrawfordTenaphalarini Heslop-Harrison Tenaphalara KuwayamaCarsidarini Crawford Protyora Kieffer Mesohomotoma Kuwayama Paracarsidara Heslop-Harrison Carsidara Walker*Triozidae LowNeolithinae subfam. n. (type-genus: Neolithus Scott) Neolithus Scott Schedoneolithus Tuthill*Triozamiinae Becker-Migdisova Triozamia VondracekTriozinae LowTriozini Low Leuronota Crawford Trichochermes Kirkaldy Egeirotrioza Boselli Aacanthocnema Tuthill & Taylor Triozoida Crawford Trio -z a Forster Paratrioza Crawford. Kuwayama Crawford Ceropsylla Riley Swezeyana Caldwell Crawforda Caldwell Hevaheva Kirkaldy Anomocephala Tuthill* Bactericera Puton* Calinda Blanchard* Epitrioza Kuwayama* Eutrioza Loginova* Hemischizocranium Tuthill* Hemitrioza Crawford* Izpania Klimaszewski* Metatrioza Tuthill* Myrmecephala Tuthill* Neotrioza Kieffer* Neotriozella Crawford* Ozotrioza Kieffer* Paracomeca Laing* Pseudotrioza Miyatake* Rhegmoza Enderlein* Schedotrioza Tuthill & Taylor* Stenopsylla Kuwayama*Pauropsyllini Crawford Pauropsylla Riibsaamen Leptynoptera Crawford Sympauropsylla Enderlein* 274 I. M. WHITE AND I. D. HODKINSON Table 18 Genera not examined in this study, and the familyor subfamily to which they probably belong. Aconopsylla Tuthill & TaylorAnomalopsylla TuthillAstragilita LoginovaAtmetocranium TuthillBrachypsylla FroggattCarsidaroida CrawfordCecidopsylla KiefferCeddotrioza KiefferCerotrioza CrawfordDiceraopsylla CrawfordDynopsylla CrawfordEngytatoneura LoginovaEriopsylla FroggattJenseniella TuthillLabicria EnderleinLanthanaphalara TuthillLeptotrioza MiyatakeLevidea Tuthill Lindbergiella Heslop-HarrisonMegadicrania LoginovaMetapsylla KuwayamaNesiope KirkaldyOptomopsylla CaldwellPaurotriozana CaldwellPexopsylla JensenPseudacanthopsylla SamyRhinopsylla RileySphingocladia EnderleinSphinia BlanchardTyora Walker Carsidarinae Aphalaridae Psyllidae Aphalaridae Psyllidae Carsidarinae Phacopteronidae Triozidae Triozidae Aphalaridae Homotomidae Triozidae Psyllidae Aphalaridae Psyllidae Aphalaridae Triozidae Triozidae Psyllidae Aphalaridae Spondyliaspididae Carsidarinae Triozidae Triozidae Psyllidae Psyllidae Triozidae Homotomidae Aphalaridae Carsidarinae The Spondyliaspididae probably derived from a Gondwanaland ancestor associated withFabaceae. However, the largest group, the Spondyliaspidinae, is associated with the genusEucalyptus (Myrtaceae). It is most likely that the Psyllidae shared a common ancestor with the Spondyliaspididae(minus Arepuniinae) in Gondwanaland in association with the Fabaceae. Present day psyllidsare largely associated with Fabaceae and Rosaceae (Rosales). The morphologically mostprimitive, and probably oldest, subfamilies of Psyllidae are the Acizziinae and Anomoneurinae,most species of which retain the habit of Fabaceae-f ceding. The Ciriacreminae have a Gond-wanaland distribution suggesting that they are also an old group. The Arytaininae (as heredefined) are probably a more recent group, restricted to the Palaearctic, but like mostCiriacreminae, retaining the habit of Fabaceae-feeding. The Psyllinae live on a variety ofhost-plants, particularly the Rosaceae, a north temperate family (Good, 1974). They are largelyHolarctic and may have had a Laurasian ancestor. The Calophyidae and Phacopteronidae have retained the habit of Rutales-feeding andprobably had Gondwanaland origins. However, the families of Rutales with which most speciesof these families are associated differ; Calophyidae feed on Anacardiaceae and Phacopteroni-dae feed on Meliaceae. The family Homotomidae probably had a Gondwanaland origin in association with Ficus(Moraceae). With the exception of a few Indian species on Santalaceae (Mathur, 1975) they feedon Ficus. Once again, the most probable origin of the Carsidaridae was in Gondwanaland. The genusMastigimas, which is only tentatively assigned to this family, has retained the habit of NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 275 Rutales-f ceding, the remaining genera are placed in the subfamily Carsidarinae, which feed onMalvales. The Triozidae is a cosmopolitan group for which a Gondwanaland origin appears most likely.There is no indication of the ancestral host relationship and, unlike other families, host-plantdiversity appears to exceed morphological diversity (p. 256). Keys Key construction These keys are intended to place most psyllid nymphs in the correct subfamily, tribe or in somecases genus. The keys are an application of the predictive properties of the new classificationand, hence, they should work for the majority of species not examined by this study as well as the15% of species which were. To maintain practicality it was sometimes necessary to artificiallysplit some polythetic taxa. For example, the family Triozidae is keyed out in 11 sections in thekey to families. In many polythetic taxa no morphological features of the nymphs could be usedto separate subordinate taxa. In such cases host-plant differences were often of more practicalvalue than morphological attributes and, hence, some key couplets contain host-plant charac-ters. Confirmatory data are given after most couplets. Nomomeristic and metric (Figs 5-7)characters are also given as confirmatory data; with the following abbreviations. Nomomeristic characters A = Number of antenna divisions (i.e. apparent segments).R = Antennal divisions upon which rhinaria occur. Metric characters (lengths) AL = Antenna length (Figs. 5,6). ARE = Circum-anal pore ring breadth (Fig. 7). BL = Body length (Figs 5,6). WL = Forewing-pad length (Figs 5,6). Metric characters (ratios) AWL = Antenna length to forewing-pad length ratio.BBBL = Body breadth (Figs 5, 6) to body length ratio. Scale lines are given on drawings of structures whose dimensions are not indicated in the text. Artificial key to families The phenetic groups which are usually monothetic, that are most congruent with the classifica-tion, are separated first. In this way, the key should have maximum reliability for species which'key-out' prior to couplet 39. Many of the characters used in the key were shown to have highclassificatory power, as indicated by high values of the SUMRAT information statistic andprincipal component eigenvectors. The key to families of White & Hodkinson (1982) should beused as an additional check on identity when Holarctic material is being examined. Artificial key to families of Psylloidea 1 Truncate sectasetae present on margin of forewing-pad (Fig. 50). [Sectasetae not present on antenna. Forewing-pad usually anteriorly produced as a humeral lobe] TRIOZIDAE (most species) (p. 289) - Sectasetae usually absent from margin of forewing-pad; if present they are pointed (except Togepsylla matsumurana Kuwayama which has two rows of truncate sectasetae on theantenna) 2 2 Scales present on body margin (Figs 173-176) TRIOZIDAE (many Hawaiian and tropical New World species) (p. 289)Scales not present on body margin 3 3 Sectasetae (pointed or truncate) present on abdomen margin and numbering more than 4 + 4.. 4 - Sectasetae usually absent from abdomen margin ; if present then numbering at most 4-1-4 12 276 I. M. WHITE AND I. D. HODKINSON 4 Tarsal arolium without a visible unguitractor (Fig. 95). [Palaearctic and Oriental. On Populus spp.] TRIOZIDAE (Egeirotrioza)(p. 289) Tarsal arolium with a distinctly visible unguitractor (Figs 65 , 66, 69-71 ,74) 5 5 Clavate setae present on dorsal surface of abdomen (Fig. 172). [Forewing-pad anteriorly produced as a humeral lobe which extends anterior to eye. Hawaii. On Tetraplasandra] TRIOZIDAE (Crawforda)(p. 289)Clavate setae absent from dorsal surface of abdomen 6 6 Sectasetae present on antenna (Figs 40, 43, 45) 7 Sectasetae absent from antenna 8 7 Antenna length to forewing-pad length ratio 0- 18-0-47. On Rutales, especially Anacardiaceae. Antennal sectasetae arranged in one row, opposite rhinaria (Fig. 40). CALOPHYIDAE (most New World spp.) (p. 288) Antenna length to forewing-pad length ratio 0-52-1-62. Usually not on Rutales, not known onAnacardiaceae. Antennal sectasetae usually in more than one row (Fig. 45). [On Malvales,Melastomataceae, Moraceae and Rutaceae.] APHALARIDAE (Paurocephala and other genera often confused with Paurocephala) (p. 278) 8 Apex of abdomen inwardly emarginate (Fig. 153). [Neotropical. On Ficus.] HOMOTOMIDAE (Synoza)(p. 288)Apex of abdomen not inwardly emarginate 9 9 Tarsal arolium with an unguitractor which forms a petiole (Fig. 69). [Palaearctic. On Populus.] APHALARIDAE (Camarotoscena) (p. 278)Tarsal arolium with a short unguitractor and no petiole 10 10 Hindwing-pad very reduced, its apex interior to margin of abdomen (Fig. 48). [Austro- Oriental and Pacific. On Calophyllum.} TRIOZIDAE (Leptynoptera) (p. 289) Hindwing-pad of normal proportions , its apex exterior to margin of abdomen 11 11 General body form very broad; body breadth more than 0-75 times body length. [Old World. On Ficus.] HOMOTOMIDAE (p. 288) General body form elongate; body breadth less than 0-70 times body length TRIOZIDAE (some spp.) (p. 289) 12 Lanceolate setae present on abdomen margin and numbering 3 + 3 or 4 -I- 4. Capitate setae (sometimes modified into tubular structures, Fig. 133; Mitrapsylla deserata Caldwell)present on abdomen margin and/or dorsal surface. [Tropical and warm temperate New World. On Fabaceae.] PSYLLIDAE (Heteropsylla and Mitrapsylla) (p. 285) Lanceolate setae usually absent from abdomen margin; if present, then capitate setae absentfrom body and wing-pads 13 13 Lanceolate setae present on abdomen margin and/or forewing-pad margin (Figs 162, 165, 167, 168) 14 Lanceolate setae absent from abdomen and forewing-pad margins 22 14 Circum-anal pore ring reduced to a few large pores (Fig. 146). [Oriental. On Burseraceae.] PHACOPTERONIDAE (Phacopteron)(p. 288)Circum-anal pore ring not reduced to a few large pores 15 15 Tibia each with a row of stout setae on outer edge (Figs 51 , 54) 16 Tibia without a row of stout setae on outer edge 17 16 Tarsal arolium with a long unguitractor which forms a petiole (Fig. 69). [Palaearctic. On Populus.] APHALARIDAE (Camarotoscena) (p. 278) Tarsal arolium very reduced (not or hardly visible). [Tropical Old World. On Meliaceae.] PHACOPTERONIDAE (Chineura)(p. 288) 17 Anal pore-field arranged as bands (similar to Fig. 155). Antenna with 10 divisions. [Neo- tropical.] PHACOPTERONIDAE (Epicarsd) (p. 288) Anal pore-field usually not arranged as bands; or if arranged as bands then antenna with atmost 8 divisions 18 18 Anal pore-field arranged as 2 rings which are each placed to one side of the anus (Figs 150-152) . [Tropical Old World. On Ficus.] HOMOTOMIDAE (some Macrohomotominae) (p. 288) Anal pore-field not arranged as 2 rings which are each placed to one side of the anus 19 19 On Anacardiaceae. Antenna with 3 divisions. Small (BL = 0-93-1-20 mm). CALOPHYIDAE (Calophya rhois)(p. 288) Usually not on Anacardiaceae; or if on Anacardiaceae, antenna with 7 or 8 divisions (someRhinocolinae) or larger (some Diaphorina, BL = 1 -34-2-13 mm) 20 20 Apical margin of abdomen truncate-acuminate (Fig. 127). Anal pore-field not arranged as a NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 277 circum-anal ring, pores concentrated at antero-lateral angle of caudal plate (Fig. 127); poresnot visible on anal plate. [Australia. On Eucalyptus.] SPONDYLIASPIDIDAE (Phellopsylla)(p. 284) Apical margin of abdomen not truncate-acuminate. Anal pore-field usually comprised of acircum-anal ring only; if pores present on caudal plate then they also occur on the anal plate. . 21 21 Circum-anal pore ring partly on caudal plate, broad and convoluted (Fig. 145). [Oriental. On Toona.] PHACOPTERONIDAE (Bharatiana)(p. 288) Circum-anal pore ring usually confined to anal plate; if partly on caudal plate then usually notconvoluted (Fig. 108), or if convoluted then pores in a narrow row (Fig. 112) APHALARIDAE (most species) (p. 278) 22 Anal pore-field (excluding circum-anal ring) arranged as 1 + 1 (Fig. 121) or 2 + 2 (Figs 120, 122) rings, which are slightly convoluted. [Tarsal arolium usually with a long unguitractorwhich forms a petiole (Fig. 77); except Euphalerus gallicolus. On Fabaceae and Rham- naceae.] SPONDYLIASPIDIDAE (Euphalerus) (p. 284) Anal pore-field (excluding circum-anal ring) usually not arranged as 1 + 1 or 2 + 2 rings; ifarranged as 1 + 1 rings then rings not convoluted (Figs 1 10, 149) 23 23 Apical margin of abdomen serrate-acuminate (Figs 125, 126, 128). Circum-anal pore ring absent. Anus posterior. [Anal pore-field, if present, comprised of small groups of pores,most of which occur on the caudal plate. On Celtis and Colophospermum.] SPONDYLIASPIDIDAE (Pachypsyllinae and Retroacizzid) (p. 284)Apical margin of abdomen usually not serrate-acuminate; or if serrate-acuminate (Figs 149, 157) circum-anal pore ring present and anus ventral 24 24 Caudal plate pointed (Figs 118, 124). [Abdomen segments usually laterally bulging (Fig. 27). Anal pore-field, if present, comprised of small groups of pores placed ventrally (Figs 118,124, 129). Anus posterior. Tarsal apical setae usually strongly capitate (Fig. 38). Australia. On Eucalyptus.} SPONDYLIASPIDIDAE (Spondyliaspidinae) (p. 284) Caudal plate (if developed) not pointed 25 25 Abdomen margin with 1 + 1, 2 + 2, 3 + 3 or 4 + 4 sectasetae .... PSYLLIDAE (many spp.) (p. 285)Abdomen margin without sectasetae 26 26 Abdomen margin with capitate setae PSYLLIDAE (many spp.) (p. 285) Abdomen margin without capitate setae 27 27 Tarsal arolium with a petiole. Arolium pad usually large relative to tarsal claws (Figs 59, 80-93) ; if pad small (Fig. 179) then host-plant is a species of Fabaceae (some Acizzia spp.) .... 28Tarsal arolium usually without a petiole; if with a petiole then arolium pad small and host-plantis a species of Meliaceae (some Pseudophacopteron spp., Fig. 94) or Terminalia (Triozahirsuta,ig.96) 31 28 Abdomen margin with rod setae (Fig. 161) 29 Abdomen margin without rod setae 30 29 Tarsal arolium with a very long petiole (Fig. 59). Antenna with 7 divisions. New World. On Fabaceae APHALARIDAE (Aphalaroida pithecolobia) (p. 278) Tarsal arolium with a short petiole (similar to Fig. 92). Antenna with 7 or 8 divisions.Palaearctic. On Ulmus PSYLLIDAE (Psylla ulmi)(p.285) 30 Tarsal arolium with a very long petiole (Fig. 59). [New World. On Fabaceae.] APHALARIDAE (Aphalaroida inermis)(p. 278)Tarsal arolium with a short petiole (Figs 79-93) PSYLLIDAE (many spp.) (p. 285) 31 Abdomen margin with clavate setae (Fig. 169). [Neotropical. On Solanaceae.] SPONDYLIASPIDIDAE (Arepuna)(p. 284)Abdomen margin without clavate setae 32 32 Anal pore-field with 1 + 1 incomplete rings in addition to circum-anal pore rings (Fig. 143). Host-plant is Mangifera indica. [Oriental.] CALOPHYIDAE (Apsylld) (p. 288) Anal pore-field usually comprised of circum-anal rings only; if 1 + 1 additional rings presentthey are complete and the host-plant is Juncus (some Livia spp., Fig. 110) or Ficus(Macrohomotoma striata, Fig. 149) 33 33 Anal pore-field comprised of circum-anal rings plus bands of pores which cover a more extensive area of the anal plate than of the caudal plate (Fig. 158). [Aftrotropical. On Antiaria.] TRIOZIDAE (Triozamid)(p. 289) Anal pore-field usually comprised of circum-anal pore rings only; if pore bands present thenthey are more extensive on the caudal plate than on the anal plate (Fig. 155) 34 34 Anal pore-field (excluding circum-anal ring which may be present or absent) comprised of pore 278 I. M. WHITE AND I. D. HODKINSON bands (Fig. 155) or a single band plus numerous ovoid pore areas (Fig. 154). [On Meliaceae andMalvales.] CARSIDARIDAE(p. 289) Anal pore-field (excluding circum-anal pore ring which is present) usually absent; if presentthen comprised of 1 -I- 1 rings (Figs 110, 149) 35 35 Apical margin of abdomen notched and with 1 + 1 stout setae (Fig. 157). [Neotropical. On Euphorbiaceae,MyrtaceaeandSolanaceae.] TRIOZIDAE (Neolithus)(p. 289) Apical margin of abdomen not notched and without 1 + 1 stout setae 36 36 Anal pore-field comprised of a circum-anal ring plus 1 + 1 additional rings each of which is separated from the circum-anal ring (Fig. 110). [Holarctic and northern Oriental. On Juncus.] APHALARIDAE (some Livia spp.)(p. 278) Anal pore-field usually comprised of circum-anal pore rings only; if 1 + 1 additional ringspresent then they are adjacent to the circum-anal pore ring (Fig. 149) 37 37 Anal pore-field comprised of circum-anal pore rings plus 1 + 1 additional rings (Fig. 149). [Austro-Oriental and Oriental. On Ficus.] HOMOTOMIDAE (Macrohomotoma)(p. 288) Anal pore-field comprised of circum-anal pore rings only (which may occasionally be incom-plete, Figs 150, 151; Mycopsylla) 38 38 Circum-anal pore rings broken and with a convoluted inner margin to the outer ring (Figs 150, 151). [BBBL more than 0-83. Australasian, Austro-Oriental and Oriental. On Ficus.] HOMOTOMIDAE (Mycopsylla) (p. 288) Circum-anal pore rings usually not broken; if broken (Fig. 147) then inner margin of outer ringnot convoluted 39 39 Antenna with 1 division. [On Rutales.] CALOPHYIDAE (Calophya rotundipennis and Microceropsylla)(p. 288)Antenna with more than 1 division 40 40 Antenna with 2 divisions . [Oriental . On Ficus .] TRIOZIDAE (Pauropsylla depressd) (p . 289) Antenna with more than 2 divisions 41 41 Antenna with 3 divisions 42 Antenna with more than 3 divisions 43 42 Tarsus with 2 segments (Fig. 52). [Oriental. On Anacardiaceae.] CALOPHYIDAE (Pelmatobrachid)(p. 288) Tarsus with 1 segment separate from the tibiotarsus (Fig. 53). [Afrotropical and Palaearctic.On Moraceae and Tamarix.] .... APHALARIDAE (Phytolyma and some Colposceniini) (p. 278) 43 General form broad (BBBL more than 0-90). [Austro-Oriental and Oriental. On Ficus.] HOMOTOMIDAE (Macrohomotomd)(p. 288)General form elongate (BBBL less than 0-85) 44 44 Antenna with 5 or 6 divisions. [On Lauraceae.] TRIOZIDAE (Pauropsylla beesoni and Trioza anceps) (p. 289)Antenna with 8, 9 or 10 divisions 45 45 General form broad (BBBL more than 0-77). Antenna with 6 rhinaria (Fig. 41 ; often difficult to see). [Tarsal arolium without a visible unguitractor (Fig. 58). New Zealand and New World. OnllexandNectandra.] APHALARIDAE(G;yr0/wy//fl)(p. 278) General form elongate (BBBL less than 0-76). Antenna with 4 rhinaria 46 46 Tarsal arolium usually not visible ; if visible then very small relative to claws , with a short petiole and a well-developed pad (Fig. 94). [Old World tropics. On Meliaceae.] PHACOPTERONIDAE (Pseudophacopteron)(p. 288)Tarsal arolium large relative to claws, with a stout petiole and a very reduced pad (Fig. 96). [Oriental. On Terminalia.] TRIOZIDAE (Trioza hirsutd) (p. 289) Keys to subfamilies and genera of Aphalaridae Many subfamilies are largely characterised by the form of the tarsal arolium, which is oftendifficult to observe. Because of this two keys to subfamilies are provided: (1) a key which followsthe classification as closely as possible and (2) a much simplified artificial key. Material ofTogepsylla sp. was not available for the analyses described earlier in this paper. However,material has since become available and the genus is tentatively included in the following keys(material in British Museum (Natural History); from New Guinea). NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 279 Key to subfamilies of Aphalaridae 1 Tarsus without an arolium, but with a pair of pulvilli (each situated beneath a claw). [Austro-Oriental, Oriental and Palaearctic. On Lauraceae.] TOGEPSYLLINAE (p. 281) Tarsus usually with a well-developed arolium; or if without a visible arolium, pulvilli alsoabsent 2 2 Tarsal arolium not visible 3 Tarsal arolium well developed and visible 4 3 Abdomen margin with short lanceolate setae (similar in proportion to those in Fig. 164). [Anal pore-field usually comprised of groups of pores on the caudal plate in addition to thecircum-anal ring (Fig. 105). Australasian, Oriental, Pacific, New Zealand and introduced toother areas on cultivated Eucalyptus. On Myrtaceae, Onagraceae and Rutaceae.] EUPHYLLURINAE (Ctenarytainini)(p. 282) - Abdomen margin usually with long simple setae; exceptionally slightly lanceolate (Fig. 162; Phytolyma minutd). [Anal pore-field comprised of circum-anal pore rings only (Fig. 162).Afrotropical. On Moraceae.] APHALARINAE (Phytolyma) (p. 281) 4 Tarsal arolium without a visible unguitractor (Figs 56-58). APHALARINAE (minus Phytolyma)(p. 281) - Tarsal arolium with a well-developed unguitractor (Figs 59-76) 5 5 Anal pore-field comprised of circum-anal pore rings plus 1 + i incomplete rings (Figs 106, 107, 112,113) EUPHYLLURINAE (minus Ctenarytainini) (p. 282) - Anal pore-field usually comprised of circum-anal rings only; or if additional pore areas present they form ovoid areas (Figs 97, 99, 100) or complete rings (Fig. 110) 6 6 Unguitractor less than half as long as whole arolium and not forming a petiole (Figs 67, 68, 71-76) 7 Unguitractor usually more than half as long as whole arolium (Figs 59-61, 69, 70); if less thanhalf as long as whole arolium then arolium petiolate (Fig. 69) 9 7 Tarsal arolium pad longer than broad (Fig. 71). [Palaearctic. On Ericaceae.] STROPHINGIINAE(p. 281) - Tarsal arolium pad broader than long (Figs 67, 68, 72-76) 8 8 Tarsal arolium pad more than 2-0 times as broad as long and with a pair of sclerotized areas (Figs 72-76). [On Rutales.] RHINOCOLINAE (Rhinocolini) (p. 282) - Tarsal arolium pad less than 1-5 times as broad as long and without sclerotized areas (Figs 67, 68). [Holarctic and Oriental. On CarexandJuncus.} LJVHNAE (p. 281) 9 Abdomen margin with lanceolate setae. Anal pore-field comprised of circum-anal rings only. Tarsal arolium as in Figs 60, 61 DIAPHORININAE(p. 283) - Abdomen margin usually without lanceolate setae; or if with lanceolate setae anal pore-field comprised of circum-anal rings plus ovoid pore areas (Figs 97, 99, 100). Tarsal arolium petiolate (Figs 59, 69, 70) 10 10 Abdomen margin with sectasetae or lanceolate setae. Petiole of tarsal arolium short relative toextension of claws (Figs 69, 70). [On Populus, Malvales and Urticales.] PAUROCEPHALINAE(p. 283) - Abdomen margin without sectasetae or lanceolate setae. Petiole of tarsal arolium long relative to extension of claws (Fig. 59). [New World. OnFabaceae.] APHALAROIDINAE(p. 283) Simplified key to subfamilies of Aphalaridae 1 Abdomen margin with rod setae. [Tarsal arolium with a very long petiole (Fig. 59). New World. On Fabaceae.] APHALAROIDINAE (Aphalaroida pithecolobia) (p. 283) - Abdomen margin without rod setae 2 On Moraceae (Clorophora and Ficus). [Abdomen margin with long setae, which may be slightly lanceolate (Fig. 162). Anus posterior. Tarsal arolium not visible. Afrotropical.] APHALARINAE (Phytolyma} (p. 281) Not on Moraceae 3 Abdomen margin with sectasetae - Abdomen margin without sectasetae 4 Antenna without sectasetae - Antenna with sectasetae . . 6 280 I. M. WHITE AND I. D. HODKINSON 5 Anal pore-field comprised of circum-anal rings plus ovoid pore areas (Figs 99, 100). Tarsal arolium petiolate (Fig. 69). On Populus [Palaearctic.] PAUROCEPHALINAE (Camarotoscena) (p. 283) Anal pore-field comprised of circum-anal rings only. Tarsal arolium not petiolate (Fig. 71). OnEricaceae (Calluna and Erica). [Palaearctic.] STROPHINGIINAE (p. 281) 6 Anal pore-field comprised of circum-anal rings plus additional broken rings (Figs 106, 113) EUPHYLLURINAE (Diclidophlebiini) (p. 282)Anal pore-field comprised of circum-anal rings only 7 7 Abdomen margin sectasetae based upon large clustered tubercles (Fig. 114). Tarsal arolium petiolate (Fig. 70). [On Malvales and possibly Urticales.] PAUROCEPHALINAE (Paurocephala)(p. 283) Abdomen margin sectasetae not based upon clustered tubercles . Tarsal arolium , if present , notpetiolate 8 8 Abdomen, wing-pads and antennal sectasetae truncate. On Lauraceae (Lindera and Litsed). Tarsal arolium absent. Paired pulvilli present, one under each claw. [Austro-Oriental, Oriental & Palaearctic.] TOGEPSYLLINAE(p. 281) Abdomen, wing-pads and antennal sectasetae pointed. On Rutaceae (Calodendrum). Tarsalarolium present (Fig. 74). Pulvilli absent. [Afrotropical.] RHINOCOLINAE (Moraniella)(p. 282) 9 Abdomen margin usually with lanceolate setae; if abdomen margin without lanceolate setae then forewing-pad margin with lanceolate setae 10 Abdomen margin and forewing-pad margin without lanceolate setae 21 10 On Carex. [Circum-anal pore rings as in Fig. 111. Tarsal arolium as in Fig. 68. Holarctic.] LIVIINAE (some spp.)(p. 281)On Dicotyledoneae 11 11 On Tamaricaceae. Lanceolate setae on abdomen and/or forewing-pad margin more than 6 times as long as broad (Fig. 163). Tarsal arolium without a visible unguitractor (Fig. 57). [Afrotropical , Oriental and Palaearctic. ] APHALARINAE (Colposceniini) (p. 281) Uusually not on Tamaricaceae; if on Tamaricaceae then lanceolate setae less than 4 times aslong as broad and tarsal arolium with a distinct unguitractor (Fig. 71) 12 12 On Calluna or Erica. [Anal pore-field comprised of circum-anal rings only (Fig. 116). Tarsal arolium as in Fig. 71. Palaearctic.] STROPHINGIINAE (p. 281) Not on Calluna or Erica 13 13 On Acer. [Tarsal arolium as in Fig. 75 . Palaearctic ] RHINOCOLINAE (Rhinocola) (p. 282) Not on Acer 14 14 Anal pore-field comprised of circum-anal rings (which may be very faint, Fig. 112) plus additional broken rings (Figs 107, 112. On Arbutus, Arctostaphylos, Olea and Phillyrea. [Tarsal arolium as in Figs 62-64. Holarctic.] EUPHYLLURINAE (Euphyllurini) (p. 282) Anal pore-field usually comprised of cirum-anal rings only; if additional pore areaspresent then they are in ovoid groups (Figs 97, 99, 100, 105) and the hosts are Populus(Camarotoscena), Rutales (Agonoscena and Ctenarytaina) , Myrtaceae or Onagraceae(Ctenarytaina) 15 15 Anal pore-field comprised of circum-anal rings plus adjacent ovoid groups of pores (Figs 97, 99, 100) 16 Anal pore-field usually comprised of circum-anal rings only; or if with ovoid groups of poresthen these are situated some distance from the outer circum-anal ring (Fig. 105) 17 16 On Populus. Tarsal arolium narrow (Fig. 69). [Palaearctic.] PAUROCEPHALINAE (Camarotoscena) (p. 283)On Ruta or Pistacia. Tarsal arolium broad (Fig. 72) RHINOCOLINAE (Agonoscena) (p. 282) 17 On Anacardiaceae in the New World. [Tarsal arolium as in Figs 73, 76.] RHINOCOLINAE (Leurolophus and Tainarys) (p. 282)Usually not on Anacardiaceae; if on Anacardiaceae then Old World 18 18 On Fraxinus. Tarsal arolium with a very long petiole (Fig. 61) DIAPHORININAE (Psyllopsis)(p. 283)Not on Fraxinus. Tarsal arolium , if visible, without a long petiole 19 19 Antenna with 8 or 9 divisions. Tarsal arolium not visible. Abdomen margin usually sinuate before apex (Fig. 105). Anal pore-field sometimes with groups of pores in addition tocircum-anal pore rings (Fig. 105). [On Myrtaceae, Onagraceae and Rutaceae.] EUPHYLLURINAE (Ctenarytaina) (p. 282)- Antenna usually with less than 8 divisions ; if with 8 divisions then tarsal arolium clearly visible . NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 281 Abdomen margin not sinuate before apex and anal pore-field comprised of circum-anal rings only 20 20 Abdomen margin lanceolate setae at least 4 times as long as broad (Fig. 164). Tarsal arolium without a visible unguitractor (Fig. 56). Antenna usually with 5 or 6 rhinaria, sometimes 4.Humeral lobe of forewing-pad not usually extended anterior to the posterior margin of theeye. [Cool temperate Holarctic and Oriental. On Asteraceae, Brassicaceae, Chenopo-diaceae, Onagraceae, Polygonaceae and Ranunculaceae.] APHALARINAE (Aphalarini) (p. 281)Abdomen margin lanceolate setae less than 4 times as long as broad (Fig. 165). Tarsal aroliumwith a long unguitractor (Fig. 60). Antenna with 4 rhinaria. Humeral lobe of forewing-padusually extended anterior to the posterior margin of the eye. [Warm temperate and tropicalAfrotropical, Oriental and Palaearctic. On numerous hosts.] DIAPHORININAE (Diaphorinini)(p. 283) 21 Anal pore-field comprised of a circum-anal ring plus 1 + 1 additional rings (Fig. 110). Tarsal arolium with a distinct unguitractor but not petiolate (Fig. 67). [Holarctic. OnJuncus.] LIVIINAE (some spp.)(p. 281) - Anal pore-field comprised of circum-anal rings only. Tarsal arolium without a distinct unguitractor (Figs 57, 58) or, if with an unguitractor then petiolate (Fig. 59) 22 22 Tarsal arolium with a long petiole (Fig . 59) . On Fabaceae . [New World . ] APHALAROIDINAE (Aphalaroida inermis) (p. 283)Tarsal arolium without a long petiole (Figs 57, 58). On Ilex, Nectandra or Tamarix APHALARINAE (Colposceniini and Gyropsylla) (p. 281) Confirmatory characters of Togepsyllinae Body, antenna and wing-pads with truncate sectasetae. Tarsus with paired pulvilli, one undereach claw. BL = 0-93-1-27 mm, WL = 0-35-0-46 mm, ARE = 0-08 mm, AWL = 0-89-1-12,BBBL = 0-40-0-51, A = 7, R = 3456. Austro-oriental, Oriental and eastern Palaearctic. OnLauraceae TOGEPSYLLA Key to species groups of Liviinae One genus only: Livia. Species formerly referred to Diraphia Waga can be separated from other species ofLivia, as follows. 1 Anal pore-field comprised of circum-anal pore rings plus 1 + 1 additional rings (Fig. 110).Tarsal arolium pad with angles acutely rounded (Fig. 67). OnJuncus. [BL = 1-44-2-45 mm,WL = 0-65-0-83 mm, ARE = 0-06-0-08 mm, AWL = 0-49-0-75, BBBL = 0-48-0-71, A = 3or 7, R = 3333 or 3577.] LIVIA (minus species formerly referred to Diraphia) - Anal pore-field comprised of circum-anal rings only (Fig. 111). Tarsal arolium pad with angles broadly rounded (Fig. 68). On Carex. [BL = 1-75-2-63 mm, WL = 0-80-0-92 mm, ARE =0-47-0-60 mm, AWL = 0-55-0-76, BBBL = 0-51-0-75, A = 7 or 10, R = 4677 or 4689.] LIVIA (formerly Diraphia spp.) Confirmatory characters of Strophingiinae Abdomen and wing-pad margins with lanceolate setae or pointed sectasetae. Tarsal arolium asin Fig. 71. BL = 1-11-1-58 mm, WL = 0-46-0-60 mm, ARE = 0-09-0-13 mm, AWL =0-41-0-56, BBBL = 0-72-0-86, A = 3, R = 3333. Palaearctic. On Calluna and Erica STROPHINGIA Key to genera of Aphalarinae 1 Each tarsus without a visible arolium . On Moraceae . [Longest seta on abdomen margin simple , others at most only slightly lanceolate (Fig. 162). Anus posterior, Fig. 162. BL = 1-72-2-47 mm, WL = 0-72-1-11 mm, ARE = 0-31-0-69 mm, AWL = 0-60-0-78, BBBL = 0-82- 1-11, A = 3, R = 3333. Afrotropical. On ClorophoraandFicus.] PHYTOLYMA Each tarsus with a visible and well-developed arolium (Figs 56-58) . Not on Moraceae 2 2 Anus posterior. Circum-anal pore ring more than 0-35 mm broad and without sharp angles (Fig. 108). [Body and wing-pads without lanceolate setae. Antenna with 6 rhinaria (may bevery difficult to see) (Fig. 41). BL = 1-52-1-87 mm, WL = 0-74-0-79 mm, ARE = 0-36-0-39 mm, AWL = 0-65-0-87, BBBL = 0-78-0-81, A = 8 or 10, R = 456788 or 456789. NewWorld and New Zealand. On Ilex and Nectandra. ] GYROPSYLLA 282 I. M. WHITE AND I. D. HODKINSON Anus ventral. Circum-anal pore ring usually less than 0-35 mm broad; if more than 0-35 mmbroad (some Craspedolepta spp.) then ring with some sharp angles (Figs 102, 103) 3 3 On Tamaricaceae. Lanceolate setae often absent; or if present they are more than 6 times as long as broad (Fig. 163) . Forewing-pad margin sometimes with a deep oblique notch 4 Not on Tamaricaceae. Lanceolate setae present and less than 6 times as long as broad(Fig. 164) . Forewing-pad margin without a deep oblique notch 5 4 General form broad (BBBL more than 0-83). [Forewing-pad usually with a humeral lobe. BL = 0-95-1-07 mm, WL = 0-37-0-43 mm, ARE = 0-09-0-11 mm, AWL = 0-38-0-43, BBBL =0-91-0-95,A = 3,R = 3333. Afrotropical, Oriental and Palaearctic. On Tamarix.] COLPOSCENIA - General form elongate (BBBL less than 0-83). [Forewing-pad without a humeral lobe. BL = 1-84 mm, WL = 0-64 mm, ARE = 0-14 mm, AWL = 0-47, BBBL = 0-75, A = 3, R =333333. Afrotropical and Palaearctic. On Tamarix and Myricaria.] CRASTINA 5 On Caltha, Polygonum, Rumex or Sisymbrium. [Antenna usually with 7 divisions, rarely 3 or 8. BL = 1-65-2-46 mm, WL = 0-70-0-93 mm, ARB = 0-16-0-31 mm, AWL = 0-40-0-62,BBBL = 0-55-0-80, R = 333333, 3577, 34577, 345677 or 456788. Holarctic and Mexico.] APHALARA - On Asteraceae or Onagraceae. [Antenna usually with 3 divisions, rarely 7. BL = 1-58-2-80 mm, WL = 0-60-1-15 mm, ARB = 0-10-0-38 mm, AWL = 0-34-0-66, BBBL = 0-52-0-91, R= 3333, 33333, 333333, 3577, 35677 or 345677. Holarctic and Mexico.] CRASPEDOLEPTA Key to genera of Rhinocolinae: Rhinocolini 1 Forewing-pad and abdomen margins with sectasetae (Fig. 166). [Tarsal arolium as in Fig. 74. BL = 1-41-1-59 mm, WL = 0-60-0-68 mm, ARB = 0-14-0-16 mm, AWL = 0-52-0-58, BBBL = 0-74-0-79, A = 3, R = 3333. Afrotropical. On Calodendrum.} . . MORANIELLA Forewing-pad and abdomen margins without sectasetae 2 Anal pore-field comprised of circum-anal rings plus adjacent ovoid groups of pores (Fig. 97). [Tarsal arolium as in Fig. 72. BL = 0-91-1-30 mm, WL = 0-34-0-42 mm, ARB = 0-12-0-18 mm, AWL = 0-76-1-17, BBBL = 0-71-0-78, A = 7, R = 3577. Afrotropical, Oriental and Palaearctic. On Pistacia and Ruta.] AGONOSCENA Anal pore-field comprised of circum-anal rings only 3 3 Anus posterior or nearly so. Outer circum-anal pore ring comprised of multiple rows of pores (Fig. 109). Antenna with 8 divisions. [Tarsal arolium as in Fig. 73. BL = 1-05-1-16 mm, WL= 0-42-0-50 mm, ARB = 0-19 mm, AWL = 0-70-0-76, BBBL = 0-68-0-76, R = 3577. Nearctic.On/Mws.] LEUROLOPHUS Anus ventral. Outer circum-anal pore ring comprised of a single row of pores (Fig. 168).Antenna with 7 divisions 4 4 Lanceolate setae on the head, wing-pads and abdomen truncate (Fig. 168). Antenna with lanceolate setae. [Tarsal arolium as in Fig. 75. BL = 1-56-2-20 mm, WL = 0-53-0-68 mm,ARB = 0-13-0-19 mm, AWL = 0-61-0-91, BBBL = 0-55-0-70, R = 3577. Palaearctic. On Acer.] RHINOCOLA Lanceolate setae on the head, wing-pads and abdomen pointed (Fig. 167). Antenna with-out lanceolate setae. [Tarsal arolium as in Fig. 76. BL = 1-61 mm, WL = 0-58 mm, ARB =0-09 mm, AWL = 0-57, BBBL = 0-78, R = 3577. Neotropical. OnSchinus.] TAINARYS Key to genera and subgenera of Euphyliurinae 1 Abdomen margin with sectasetae Abdomen margin with lanceolate or stout simple setae 3 2 Anal pore-field (other than circum-anal rings) comprised of unbroken bands (Fig. 106) . [Tarsal arolium as in Fig. 65. BL = 2-25-2-75 mm, WL = 0-68-0-72 mm, ARB = 0-08-0-09 mm,AWL = 1-24-1-37, BBBL = 0-58-0-60, A = 9, R = 3578. Old World tropics. OnSterculiaceae.] DICLIDOPHLEBIA - Anal pore-field (other than circum-anal rings) comprised of broken bands (Fig. 113). [Tarsal arolium as in Fig. 66. BL = 1-34-1-53 mm, WL = 0-59-0-68 mm, ARB = 0-07-0-10 mm,AWL = 1-44-1-62, BBBL = 0-79-0-85, A = 9, R = 3577. Tropical. On Melastomataceae,Sterculiaceae and Tiliaceae.] PARAPHALAROIDA 3 Tarsal arolium visible and well developed (Figs 62-64) 4 NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 283 Tarsal arolium not visible 6 4 Anal pore-field comprised of circum-anal pore rings, pore bands and ovoid pore groups (Fig. 107). [Tarsal arolium as in Fig. 62. BL = 1-22-2-00 mm, WL = 0-51-0-75 mm,ARB = 0-08-0-12 mm, AWL = 0-72-0-89, BBBL = 0-71-0-92, A = 8-9, R = 3578. Afrotropical, Oriental and Palaearctic. On Olea and Phillyrea.} EUPHYLLURA Anal pore-field comprised of convoluted rings (Fig. 112) 5 5 Tarsal arolium not petiolate (Fig. 64). On Arctostaphylos . [BL = 1-90-2-37 mm, WL = 0-74-1-10 mm, AWL = 0-75-1-10 mm, BBBL = 0-71-0-76, A = 7, R = 3577. Nearctic.] NEOPHYLLURA (NEOPHYLLURA) Tarsal arolium petiolate (Fig. 63). On Arbutus. [BL = 2-15-2-65 mm, WL = 0-53-0-80 mm,AWL = 0-80-1-02, BBBL = 0-66-0-68, A = 7, R = 3577. Nearctic.] NEOPHYLLURA (ARBUTOPHILA) 6 Anal pore-field comprised of circum-anal rings only, or circum-anal rings plus ovoid groups of pores which are confined to the lateral areas of the abdomen (Fig. 105). [BL = 1-18-1-38 mm, WL = 0-48-0-60 mm, ARB = 0-07-0-09 mm, AWL = 0-68-0-78, BBBL = 0-58-0-70, A = 8, R = 3577. Australia, New Zealand and Pacific. On Eucalyptus, Fuchsia andBoronia. Introduced to Afrotropical and Palaearctic on cultivated Eucalyptus.} CTENARYTAINA - Anal pore-field comprised of circum-anal rings plus broken bands of pore groups which are predominantly dorsal in position (Fig. 119). [BL = 1-51-2-00 mm, WL = 0-55-0-64 mm,AWL = 0-67-0-73, BBBL = 0-50-0-56, A = 9, R = 3578. Australia and Oriental. On Eucalyptus and other Myrtaceae . ] 'EUCAL YPTOL YMA ', ( EUPHYLLURA ' (some species currently referred, but excluding the type-species) Key to genera of Paurocephalinae 1 Abdomen margin with sectasetae based upon large clustered tubercles (Fig. 114). Antennawith sectasetae (Fig. 45). Anal pore-field comprised of circum-anal rings only. On Malvalesand Moraceae. [Tarsal arolium as in Fig. 70. BL = 0-97-1-41 mm, WL = 0-40-0-53 mm,ARB = 0-13-0-16 mm, AWL = 0-81-1-00, BBBL = 0-74-0-92, A = 3, R = 3333. OldWorld.] PAUROCEPHALA - Abdomen margin usually with lanceolate setae ; if with sectasetae , then they are not based upon large clustered tubercles. Antenna without sectasetae. Anal pore-field comprised of circum-anal rings plus adjacent crescent-shaped (Fig. 99) or ovoid (Fig. 100) pore areas. OnPopulus. [Tarsal arolium as in Fig. 69. BL = 1-04-1-98 mm, WL = 0-44-0-71 mm, ARB =0-05-0-23 mm, AWL = 0-70-0-84, BBBL = 0-76-0-85, A = 7, R = 3577. Palaearctic.] CAMAROTOSCENA Key to genera of Diaphorininae 1 Tarsal arolium with a long petiole (Fig. 61). Antenna with 8 divisions and very long (AL = 0-79-0-87 mm, AWL = 0-91-1-23). On Fraxinus. Fore wing-pad at most slightly extendedanteriorly as a humeral lobe, which does not extend anterior to procoxa. [BL = 1-52-2-57 mm, WL = 0-64-0-89 mm, ARB = 0-16-0-27 mm, BBBL = 0-64-0-81, R = 3578.Palaearctic and Oriental. Introduced to Nearctic.] PSYLLOPSIS - Tarsal arolium without a long petiole (Fig. 60). Antenna with 3 divisions and short (AL = 0-29-0-45 mm, AWL = 0-26-0-54). Not on Fraxinus. Forewing-pad usually extendedanteriorly as a humeral lobe , which extends anterior to procoxa 2 2 Body and wing-pads broad (BBBL = 0-78-0-93). [BL = 1-34-2-13 mm, WL = 0-75-1-24 mm, ARB = 0-11-0-24 mm, AWL = 0-26-0-54, R = 3333. Afrotropical, Oriental and Palaearc-tic. Introduced to Neotropical. On a wide variety of host-plants.] DIAPHORINA Body and wing-pads narrow (BBBL = 0-73-0-75). [BL = 1-91-2-06 mm, WL = 0-87-0-88 mm,ARB = 0-28 mm, AWL = 0-38-0-43, R = 3333. Afrotropical. OnStrychnos.] PENNAVENA Confirmatory characters of Aphalaroidinae Tarsal arolium with a very long petiole (Fig. 59). BL = 1-51-1-75 mm, WL = 0-59-0-66 mm,ARB = 0-07-0-09 mm, AWL = 0-63-0-77, BBBL = 0-71-0-76, A = 7, R = 3577. NewWorld . On Fabaceae . . . APHALAROIDA 284 I. M. WHITE AND I. D. HODKINSON Key to genera and species groups of Spondyliaspididae 1 Caudal plate pointed (Figs 118, 124), without apical'teeth' 2 Caudal plate not pointed (Fig. 121), with apical 'teeth' (Figs 122, 125, 127, 128) 5 2 Margin of abdomen with 'teeth' placed 1 + 1 either side of caudal plate area (Fig. 118). [BL = 3-35-3-91 mm, WL = 1-03-1-18 mm, AWL = 0-83-0-90, BBBL = 0-53-0-30, A = 10, R = 4689. Australia. On Eucalyptus.] CREIIS Margin of abdomen without 'teeth' placed 1 + 1 either side of caudal plate area (Figs 124, 129) 3 3 Anal pore-field absent. Antenna short relative to forewing-pad (AWL = 0-57-0-79). [BL = 1-61-2-89 mm, WL = 0-68-0-92 mm, BBBL = 0-54-0-78, A = 9 or 10, R = 3578 or 4689. Australia. On Eucalyptus.] CARDIASPINA Anal pore-field comprised of scattered pore groups (Figs 124-129) 4 4 Antenna with 10 divisions and about twice as long as forewing-pad (AWL = 2-02). [BL = 2-44 mm, WL = 0-61 mm, BBBL = 0-55, R = 4689. Australia. On Eucalyptus.] SPONDYLIASPIS - Antenna with 9 divisions and about as long as forewing-pad (AWL = 0-87-1-14). [BL = 1-41-2-15 mm, WL = 0-66-0-89 mm, BBBL = 0-67-0-84, R = 3578 or 34578. Australia. OnEucalyptus.] GLYCASPIS 5 Apical margin of abdomen with 'tooth-like' processes (Figs 120, 122, 125) 6 Apical margin of abdomen without 'tooth-like' processes 13 6 Anal pore-field comprised of 1 + 1 ventral and 1 + 1 dorsal rings (Figs 120 , 122) Anal pore-field usually comprised of scattered pore groups (Fig. 126) , or absent 8 7 Abdomen with large apical 'teeth' arranged 1 + 1 (Fig. 120). Forming galls on Rhamnaceae. [BL = 2-78-2-88 mm, WL = 0-73-0-80 mm, AWL = 1-56-1-68, BBBL = 0-55-0-68, A = 7,R = 3567. New World tropics.] EUPHALERUS (E. gallicolus) - Abdomen with about 4 small apical 'teeth' (Fig. 122). Forming lerps on Fabaceae. [BL = 1-15-1-17 mm, WL = 0-44-0-58 mm, AWL = 1-00-1-36, BBBL = 0-85-0-94, A = 9, R =3578. New World tropics.] EUPHALERUS (E. nidifex} 8 Antenna with 10 divisions. On Celtis 9 Antenna with 8 or 9 divisions. Usually not on Celtis , or if on Celtis antenna with 8 divisions 11 9 Abdomen apical 'teeth' without medial 'tooth' or 'teeth' enlarged (Fig. 126). Lerp forming. [BL = 3-08 mm, WL = 0-84 mm, AWL = 0-99, BBBL = 0-51, R = 4689. Eastern Palaearctic.] PACHYPSYLLA (P. japonicd) Abdomen apical 'teeth' with median 'tooth' or 'teeth' enlarged (Fig. 125) . Gall forming 10 10 Large, body length more than 4-5 mm. Forming galls on stems and leaf petioles. [BL = 4-91 mm, WL = 1-22-1-53 mm, AWL = 0-65-0-75, BBBL = 0-61-0-63, R = 4689. Nearctic.] PACHYPSYLLA (P. venusta} - Small, body length less than 4-5 mm. Forming galls on leaves. [BL = 2-03-4-25 mm, WL = 0-69-1-03 mm, AWL = 0-73-1-00, BBBL = 0-48-0-75, R = 4689. Nearctic and Mexico.] PACHYPSYLLA (minus P. japonica and P. venusta) 11 Anal pore-field absent. Tarsal arolium petiolate (Fig. 78). [Apical 'teeth' of abdomen as in Fig. 128. BL = 2-20-2-61 mm, WL = 0-76-0-87 mm, AWL = 1-25-1-43, BBBL = 0-67-0-69, A = 8, R = 4688. Afrotropical. Forming lerps on Colophospermum.] RETROACIZZIA Anal pore-field comprised of scattered pore groups (Fig. 1 27) . Tarsal arolium not visible 12 12 Abdomen margin with lanceolate setae. Antenna with 9 divisions. [BL = 2-18-2-66 mm, WL = 0-70-0-71 mm, AWL = 1-08-1-21, BBBL = 0-59-0-66, R = 3578. Australia. In discarded lerps of Spondyliaspidinae on Eucalyptus.] PHELLOPSYLLA Abdomen margin without lanceolate setae. Antenna with 8 divisions. [BL = 3 mm, WL =1-1mm, AWL = 0-9, BBBL = 0-7, R = 4688, all estimated from Ferris (1926). Nearctic andMexico. Forming lerps on Celtis.] TETRAGONOCEPHALA 13 Anal pore-field comprised of circum-anal rings only. Abdomen and wing-pad margins with clavate setae (Fig. 169). [BL = 1-44 mm, WL = 0-57 mm, ARE = 0-08 mm, AWL = 0-81, BBBL = 0-72, A = 7, R = 3577. Neotropical. On Solanaceae.] AREPUNA Anal pore-field comprised of circum-anal rings plus pore bands (Fig. 121) or pore groups (Figs117, 123) arranged as rings. Abdomen and wing-pad margins without clavate setae 14 14 Anal pore-field comprised of circum-anal rings plus pore bands (Fig. 121). [BL = 1-56-1-77 mm, WL = 0-53-0-66 mm, AWL = 1-06-1-45, BBBL = 0-60-0-80, A = 7, R = 3577 or 4677. Nearctic. On Rhamnaceae.] EUPHALERUS (E. jugovenosus, E. rugipennis and E. vermiculosus, but not the type-species of Euphalerus) NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 285 Anal pore-field comprised of circum-anal rings plus pore groups (Figs 117, 123) 15 15 Abdomen margin with 3 + 3 sectasetae (Fig. 123). [Anal pore-field as in Fig. 123. BL =1-44-1-72 mm, WL = 0-48-0-53 mm, AWL = 1-13-1-15, BBBL = 0-65-0-66, A = 8, R =3578. Afrotropical. On Fabaceae.] EUPHALERUS (some species assigned to the genus, but not the type species) Abdomen margin without sectasetae. [Anal pore-field as in Fig. 117. BL = 1-69 mm, WL = 0-64 mm, ARE = 0-12 mm, AWL = 1-11, BBBL = 0-65, A = 8, R = 3578. Afrotropical. On Fabaceae.] COLOPHORINA Keys to subfamilies and genera of Psyllidae Key to subfamilies of Psyllidae 1 Hindwing-pad margin with a pointed sectaseta. [Neotropical. On Fabaceae.] ACIZZIINAE (Neopsyllia and Platycorypha)(p. 285)Hindwing-pad margin without sectasetae 2 2 Dorsal surface of abdomen and thorax with lanceolate setae . [Neotropical . On Fabaceae . ] ACIZZIINAE (Mitrapsylla) (p. 285)Dorsal surface of abdomen and thorax without lanceolate setae 3 3 Abdomen margin with 3 + 3 or 4 + 4 tubular sectasetae (Fig. 37 [ts]) or lanceolate setae (Fig. 37 [1]). [Tropical. On Fabaceae and Euphorbiaceae.] CIRIACREMINAE (p. 286) Abdomen margin without tubular sectasetae or lanceolate setae 4 4 Tarsal arolium pad with two separate spinule-covered areas (Figs 81 , 83 , 84) . Circum-anal pore rings partly on the dorsal surface of the abdomen (Fig. 130). [Tropical Old World and eastern Palaearctic. On Fabaceae and Moraceae.] ANOMONEURINAE (Anomoneurini) (p. 286) Tarsal arolium pad with one spinule-covered area (Figs 79, 80, 86, 92, 93). Circum-anal porerings usually confined to the ventral surface of the abdomen (the exceptions being somePsyllinae, Figs 132, 134, 136, 142) 5 5 Antenna with 9 divisions 6 Antenna with less than 9 divisions 7 6 On Cercocarpus. [Nearctic.] PSYLLINAE (Psylla magnd) (p. 287) On Fabaceae or Proteaceae ACIZZIINAE (Acizzia)(p. 285) 7 Tarsal arolium pad only slightly broader at apex than at base (Fig. 93) ACIZZIINAE (Trigonon)(p. 285)Tarsal arolium pad very much broader at apex than at base (Figs 86, 92) 8 8 Antenna with 5 divisions ANOMONEURINAE (some species assigned to Euphalerus but excluding the type-species) (p. 286)Antenna with 7 or 8 divisions 9 9 On Cercocarpus. Circum-anal pore ring breadth to antenna length ratio more than 0-3 (0-36-0-40) ANOMONEURINAE (Euphalerus tantillus but not the type-species of Euphalerus) (p. 286)Usually not on Cercocarpus; or if on Cercocarpus then circum-anal pore ring breadth toantenna length ratio less than 0-3 (0-1 1-0-25) 10 10 Not on Fabaceae or Solanaceae PSYLLINAE (most species) (p. 287) On Fabaceae or Solanaceae 11 11 Circum-anal pore rings convoluted (Fig. 138) PSYLLINAE (Psylla pulchella) (p. 287) Circum-anal pore rings not convoluted 12 12 OnBauhinia PSYLLINAE (Psylla simlae) (p. 287) Not on Bauhlnia 13 13 OnGenisteae ARYTAININAE(p. 287) Not on Genisteae 14 14 Ventral surface of abdomen with capitate setae ACIZZIINAE (Freysuild) (p. 285) Ventral surface of abdomen without capitate setae ANOMONEURINAE (some species assigned to Euphalerus but excluding the type-species) (p. 286) Key to genera of Acizziinae 1 Dorsal surface of thorax and abdomen with lanceolate setae. Abdomen margin with 3 + 3lanceolate setae and 'funnel' shaped setae (Fig. 133). [Tarsal arolium as in Fig. 89. BL = 1-25 286 I. M. WHITE AND I. D. HODKINSON mm, WL = 0-45 mm, ARE = 0-09 mm, AWL = 1-67, BBBL = 0-67, A = 7, R = 3577. Neotropical. On Mimosoideae.] MITRAPSYLLA Dorsal surface of thorax and abdomen without lanceolate setae. Abdomen margin withoutlanceolate setae and 'funnel'-shaped setae 2 2 Hindwing-pad margin with a pointed sectaseta. Abdomen margin with 3 + 3 pointed sectasetae 3Hindwing-pad margin and abdomen margin without sectasetae 4 3 Antenna with 9 divisions. [Tarsal arolium as in Fig. 91 . BL = 2-41 mm, WL = 0-82 mm, ARB = 0-14 mm, AWL = 2-68, BBBL = 0-58, R = 3578. Cuba. OnMyroxylon.] PLATYCORYPHA - Antenna with 10 divisions. [Tarsal arolium as in Fig. 90. BL = 2-11-2-31 mm, WL = 0-84-0-90mm, ARB = 0-22-0-24 mm, AWL = 1-22-2-11, BBBL = 0-66-0-69, R = 4689. SouthAmerica. On Erythrina and Tipuana.] NEOPSYLLIA 4 Antenna with 7 divisions . Ventral surface of abdomen with capitate setae . [Tarsal arolium as in Fig. 86. BL = 1-54 mm, WL = 0-61 mm, ARB = 0-11 mm, AWL = 1-75, BBBL = 0-71, R = 3577. South America. On Caesalpinoideae and Solanaceae.] FREYSUILA Antenna with 8 or 9 divisions . Ventral surface of abdomen without capitate setae 5 5 Antenna with 8 divisions. Tarsal arolium pad only slightly broader at apex than at base (Fig. 93). [BL = 2-23 mm, WL = 0-64 mm, ARB = 0-25 mm, AWL = 2-77, BBBL = 0-64, R = 3578. Austro-Oriental and Pacific. Introduced to Hawaii.] TRIGONON Antenna with 9 divisions. Tarsal arolium pad much broader at apex than at base (Fig. 80) or very reduced (Fig. 79). [BL = 0-95-1-66 mm, WL = 0-41-0-61 mm, ARB = 0-07-0-11 mm,AWL = 0-91-1-57, BBBL = 0-63-0-85, R = 3578. An almost cosmopolitan tropical andwarm temperate genus. On Mimosoideae and Proteaceae. ] ACIZZIA Key to genera of Anomoneurinae The 'unnamed tribe' includes the genus Amorphicola and several species which are currentlyreferred to the genus Euphalerus (the type-species of which, E. nidifex, is here placed in theSpondyliaspididae). It is possible that this tribe should include Cyamophila Loginova. However,this is not Cyamophilini in the sense in which Loginova (1976o; 1977) defined it. Because of theuncertain status of this tribe it is not further divided in the following key. 1 Circum-anal pore rings confined to ventral surface of abdomen. [BL = 1-13-1-98 mm, WL = 0-45-0-68 mm, ARB = 0-11-0-13 mm, AWL = 0-72-0-89, BBBL = 0-60-0-81, A = 5 or 7, R = 3455 or 3577. On Fabaceae and Rhamnaceae.] 'unnamed tribe' Circum-anal pore rings partly on the dorsal surface of the abdomen (Fig. 130) 2 2 Abdomen margin with 3 + 3 sectasetae. Antenna with 9 divisions. [Tarsal arolium as in Fig. 81. BL = 2-49-3-11 mm, WL = 1-02-1-03 mm, ARB = 0-55-0-65 mm, AWL = 1-63-1-71, BBBL = 0-69-0-75, R = 4689. Oriental and Palaearctic. On Morus.} ANOMONEURA Abdomen margin without sectasetae. Antenna with 7 divisions. [Tarsal arolium as in Figs 83,84. BL = 2-09-3-38 mm, WL = 1-02-1-03 mm, ARB = 0-33-0-91 mm, AWL = 2-33-2-45,BBBL = 0-56-0-65, R = 3577. Tropical Old World. On Fabaceae.] EPIPSYLLA Key to genera of Ciriacreminae 1 Abdomen margin with 3 + 3 or 4 + 4 lanceolate setae, without tubular sectasetae (Fig. 37 [1]). [BL = 1-12-1-53 mm, WL = 0-41-0-55 mm, ARB = 0-10-0-18 mm, AWL = 1-59-2-17,BBBL = 0-58-0-74, A = 7-8, R = 3577 or 3578. Tropical and warm temperature New World. On Mimosoideae.] HETEROPSYLLA Abdomen margin without lanceolate setae but with 3 + 3 or 4 + 4 tubular sectasetae which areusually placed on slightly raised tubercles (Fig. 37 [ts]) . [Tropical . Anus posterior.] 2 2 Antenna with 9 divisions and abdomen margin with 3 + 3 sectasetae. [Tarsal arolium as in Fig. 88. BL = 1-46 mm, WL = 0-49 mm, ARB = 0-11 mm, AWL = 1-71, BBBL = 0-55, R = 3578. Neotropical and Pacific. On Mimosoideae.] ISOGONOCERAIA Antenna usually with 7 divisions. If antenna with 9 divisions (Ciriacremum harteni and C.julbernardioides) then abdomen margin with 4 + 4 sectasetae 3 3 Abdomen margin with 4 + 4 tubular sectasetae. Antenna with 7 divisions. [Tarsal arolium as in Fig. 85. BL = 1-57-2-58 mm, WL = 0-62-0-79 mm, ARB = 0-24-0-26 mm, AWL =1-97-2-53, BBBL = 0-57-0-72, R = 3577. Neotropical. On Mimosoideae.] EUCEROPSYLLA NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 287 Abdomen margin usually with 3 + 3 tubular sectasetae. If abdomen margin with 4 + 4 tubular sectasetae antenna with 9 divisions 4 4 Forewing-pad dorsal surface without capitate setae. Antenna with 7 divisions. [Tarsal aroliumas in Fig. 87. Abdomen margin with 3 + 3 tubular sectasetae. BL = 1-80-2-43 mm, WL =0-70-0-75 mm, ARE = 0-36-0-40 mm, AWL = 1-90-1-91, BBBL = 0-59-0-69, R = 3577.Pacific. On Caesalpinoideae and Euphorbiaceae.] INSNESIA - Forewing-pad dorsal surface usually with capitate setae. If fore wing-pad dorsal surface without capitate setae then antenna with 9 divisions. [Tarsal arolium as in Fig. 82. Abdomen marginwith 3 + 3 or 4 + 4 tubular sectasetae. BL = 1-33-2-25 mm, WL = 0-51-0-74 mm, ARE =0-12-0-34 mm, AWL = 1-54-2-46, BBBL = 0-60-0-71, R = 3577 or 3578.) CIRIACREMUM Key to genera of Arytaininae 1 Antenna without capitate setae. [Forewing-pad dorsal surface without capitate setae. Abdo- men margin with 3 + 3 or 4 + 4 pointed sectasetae. BL = 1-37-2-09 mm, WL = 0-57-0-72mm, ARE = 0-14-0-28 mm, AWL = 0-93-1-89, BBBL = 0-59-0-98, A = 7, R = 3577.Palaearctic. OnGenisteae.] ARYTAINILLA - Antenna with a capitate seta positioned close to rhinarium IV 2 2 Dorsal surface of forewing-pad with capitate setae. Abdomen margin with up to 3 + 3 (sometimes none) pointed sectasetae. [BL = 1-81-2-62 mm, WL = 0-62-0-73 mm, ARE =0-19-0-23 mm, AWL = 1-53-1-61, BBBL = 0-53-0-71, A = 7, R = 3577. Palaearctic and Oriental. OnGenisteae.] ARYTAINA Dorsal surface of forewing-pad without capitate setae. Abdomen margin with 4 + 4 pointedsectasetae. [BL = 1-91-2-25 mm, WL = 0-62-0-79 mm, ARE = 0-17^0-22 mm, AWL =1-14-1-37, BBBL = 0-66-0-72, A = 7, R = 3577. Palaearctic and Afrotropical. OnGenisteae.] FLORIA Key to genera of Psyllinae Keys for the separation of Psylla into subgenera have been provided by Ossiannilsson (1970),Loginova (1978) and White & Hodkinson (1982). In the following key to genera no tenable methodcould be found to distinguish Purshivora pubescens (Crawford) from some members of Psyllasubgenus Hepatopsylla Ossiannilsson. 1 Circum-anal pore rings extending onto dorsal surface of abdomen and of a convoluted shape (Fig. 142). Antenna with 7 divisions. [BL = 1-34-1-98 mm, WL = 0-68-0-75 mm, ARE =0-51-0-57 mm, AWL = 0-87-1-18, BBBL = 0-66-0-77, R = 3577. Palaearctic. On Buxus.] SPANIONEURA - Circum-anal pore rings usually not extending onto dorsal surface of abdomen (Figs 135-141); or if rings convoluted and extending onto dorsal surface of abdomen (Fig. 134), antenna with 9 divisions 2 2 Dorsal surface of forewing-pad with capitate or clavate setae. Abdomen margin with 3 + 3 pointed sectasetae. [BL = 1-25-1-62 mm, WL = 0-45-0-52 mm, ARE = 0-11-0-13 mm,AWL = 0-45-1-09, BBBL = 0-60-0-72, A = 7, R = 3577. Nearctic. On Ceanothus and Cercocarpus.} CEANOTHIA Dorsal surface of forewing-pad usually without capitate and clavate setae; if dorsalsurface of forewing-pad with capitate setae, abdomen with 4 + 4 pointed sectasetae (Psyllasimlae) 3 3 On Ceanothus. [Outer circum-anal pore ring multiple (Fig. 132). BL = 1-25-2-16 mm, WL = 0-56-0-75 mm, ARE = 0-11-0-37 mm, AWL = 0-75-1-27, BBBL = 0-69-0-85, A = 7, R =3577. Nearctic.] EUGLYPTONEURA - Not on Ceanothus 4 4 On Purshia. Outer circum-anal pore ring comprised of a multiple row of pores. [Abdomen margin without sectasetae. BL = 1-27-1-56 mm, WL = 0-43-0-53 mm, ARE = 0-13-0-20mm, AWL = 1-08-1-27, BBBL = 0-69-0-79, A = 7, R = 3577. Nearctic.] PVRSHIVORA (P. chelifera) - Usually not on Purshia; or if on Purshia outer circum-anal pore ring comprised of a single row of pores (similar to Figs 137-139) 5 5 On Purshia. [Confirmatory characters included with Purshivora chelifera and Psylla confirma- tory descriptions.] Purshivora pubescens, Psylla coryli, P. hirsuta, P. minuta 288 I. M. WHITE AND I. D. HODKINSON Not on Purshia. [Circum-anal pore rings of various forms (Figs 134-141). BL = 1-31-3-08 mm,WL = 0-46-1-15 mm, ARB = 0-06-0-85 mm, AWL = 0-61-2-20, BBBL = 0-50-1-08, A = 7or 8, R = 3577 or 3578. Holarctic and northern Oriental. On a wide variety of hosts,especially Betulaceae, Rhamnaceae, Rosaceae and Salicaceae.] PSYLLA (most species) Key to genera of Calophyidae 1 Anal pore-field comprised of circum-anal pore rings plus 1 + 1 pairs of partial rings (Fig. 143). Antenna with 10 divisions. [BL = 2-22 mm, WL = 0-81 mm, ARB = 0-13 mm, AWL = 1-05, BBBL = 0-80, R = 4689. Oriental. On Mangifera.] APSYLLA Anal pore-field comprised of circum-anal pore rings only, or not discernible . Antenna with 1 to3 divisions 2 2 Antenna length to forewing-pad length ratio 0-65-0-73. Humeral lobe of forewing-pad not extended as far forward as eye. [BL = 1-47-1-75 mm, WL = 0-60-0-68 mm, ARB = 0-08 mm, BBBL = 0-83-0-96, R = 3333. Oriental. On Buchanania.} PELMATOBRACHIA Antenna length to forewing-pad length ratio 0-18-0-55. Humeral lobe of forewing-padextended forward beyond the posterior margin of the eye 3 3 Antenna with one rhinarium. Body breadth with wing-pads to body length ratio 1-23-1-25. Margin of head, wing-pads and abdomen without sectasetae. [BL = 1-15-1-34 mm, WL =0-64-0-75 mm, ARB = 0-03-0-04 mm, AWL = 0-49-0-53, A = 1. Oriental. OnMangifera.] MICROCEROPSYLLA Antenna with 3 or 4 rhinaria. Body breadth with wing-pads to body length ratio 0-83-1-16.Margin of head, wing-pads and abdomen usually with sectasetae. [BL = 0-93-1-52 mm, WL= 0-49-0-84 mm, ARB = 0-07-0-15 mm, AWL = 0-18-0-56, A = 1, 2 or 3, R = 111, 1111,2222 or 3333. Tropical and warm temperate areas. On Anacardiaceae, Burseraceae andRutaceae.] CALOPHYA Key to genera of Phacopteronidae 1 Anal pore-field arranged as bands (similar to Fig. 155). Antenna with 10 divisions. [Abdomen margin with lanceolate setae. BL = 2-20 mm, WL = 0-70 mm, AWL = 2-14, BBBL = 0-68, R = 4689. Neotropical.] 1EPICARSA Anal pore-field arranged as circum-anal rings only (Figs 145-147), often very reduced (Fig. 146) . Antenna with 3-9 divisions 2 2 Outer circum-anal pore ring convoluted and extending onto the dorsal surface of the abdomen (Fig. 145). Antenna longer than forewing-pad length (AWL = 1-20). [Abdomen margin withsmall lanceolate setae. BL = 3-06 mm, WL = 1-25 mm, ARB = 0-91 mm, BBBL = 0-71, A = 8, R = 3578. Oriental. On Toona.} BHARATIANA Outer circum-anal pore ring not convoluted and confined to ventral surface of abdomen.Antenna shorter than forewing-pad length (AWL = 0-50-0-64) 3 3 Antenna with 3 divisions, the last of which is covered in prominent hairs (Fig. 47). On Burseraceae. [Abdomen margin with short lanceolate setae. Circum-anal pore ring veryreduced (Fig. 146). BL = 2-90-4-10 mm, WL = 1-15 mm, AWL = 0-50, BBBL = 0-64, R = 3333. Oriental.] PHACOPTERON Antenna with more than 3 divisions. On Meliaceae 4 4 Antenna with 5 divisions. Tibia each with a row of stout setae on outer edge (Fig. 54). Abdomen margin with lanceolate setae. [BL = 1-03 mm, WL = 0-37 mm, ARB = 0-29 mm, AWL = 0-62, BBBL = 0-66, R = 3355. Pacific. OnAglaia.} tCHINEURA Antenna with 8 or 9 divisions. Tibia without stout setae. Abdomen margin without lanceolatesetae. [Tarsal arolium often with a short petiole (Fig. 94). Circum-anal pore rings often withseparated pores (Fig. 147). BL = 1-78-2-31 mm, WL = 0-61-0-71 mm, ARB = 0-17-0-21mm, AWL = 0-50-0-64, BBBL = 0-60-0-75, R = 3578 or 4689. Afrotropical. On Khaya.] PSEUDOPHACOPTERON Key to genera of Homotomidae 1 Abdomen margin with sectasetae Abdomen margin without sectasetae 3 2 Dorsal surface of abdomen with pointed sectasetae. Apical margin of abdomen inwardly NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 289 marginate (Fig. 153). General form elongate (BBBL = 0-65-0-81). [BL = 1-97-3-17 mm,WL = 0-75-1-13 mm, ARE = 0-61-0-76 mm, AWL = 1-05-1-48, A = 3, R = 3333. Neotropical. On Ficus.} SYNOZA Dorsal surface of abdomen without pointed sectasetae. Apical margin of abdomen evenlyrounded. General form broad (BBBL = 0-83-1-04). [BL = 1-59-2-68 mm, WL = 0-81-1-45mm, ARE = 0-34-0-46 mm, AWL = 0-48-0-69, A = 2-3, R = 2222 or 3333. Tropical andwarm temperate Old World. On Ficus.] HOMOTOMA 3 Inner margin of outer circum-anal pore ring convoluted and pore rings confined to the ventral surface of the abdomen (Figs 150, 151). Antenna with 2 divisions. [BL = 2-33-3-09 mm, WL= 1-13-1-40 mm, AWL = 0-43-0-53, BBBL = 0-87-1-10, R = 2222. Australasian, Austro-Oriental and Oriental. On Ficus.] MYCOPSYLLA Inner margin of outer circum-anal pore ring usually not convoluted (Figs 148, 149); ifconvoluted then pore rings extending onto the dorsal surface of the abdomen (Fig. 152).Antenna with more than 2 divisions 4 4 Circum-anal pore rings extending onto dorsal surface of abdomen (Fig. 152). Antenna with 3 divisions. [BL = 2-31-2-50 mm, WL = 1-20-1-40 mm, AWL = 0-46-0-53, BBBL = 1-61-1-25, R = 3333. Afrotropical. On Ficus.] PSEUDOERIOPSYLLA Circum-anal pore rings confined to ventral surface of abdomen (Figs 148, 149). Antenna with 9or 10 divisions. [BL = 1-84-2-84 mm, WL = 0-87-1-09 mm, ARE = 0-72-0-93 mm, AWL =0-70-0-82, BBBL = 0-84-0-98, R = 4578 or 4689. Austro-Oriental and Oriental. On Ficus.] MACROHOMOTOMA Key to subfamilies of Carsidaridae 1 Anal pore-field comprised of numerous pore groups (Fig. 154). On Cedrela (Meliaceae). [BL= 1-87-2-71 mm, WL = 0-72-0-82 mm, AWL = 1-80-2-43, BBBL = 0-60-0-77, A = 9-10, R = 3578 or 4689 . Neotropical . ] M ASTIGIM ATIN AE (Mastigimas) Anal pore-field comprised of pore bands (Fig. 155). On Malvales. [BL = 1-46-3-06 mm, WL =0-59-0-87 mm, AWL = 1-53-2-73, BBBL = 0-50-0-78, A = 10, R = 4689. Tropical.] CARSIDARINAE Key to genera of Triozidae 1 Apical margin of abdomen with 'tooth-like' processes (Fig. 157). [Head, thorax, abdomen and wing-pads without secta-setae. BL = 2-80-3-02 mm, WL = 1 -03-1-13 mm, ARE = 0-24-0-26mm, AWL = 0-70-0-77, BBBL = 0-58-0-66, A = 6, R = 3355. Neotropical. On Euphor- biaceae, Myrtaceae and Solanaceae.] NEOLITHUS Apical margin of abdomen without 'tooth-like' processes 2 2 Anal pore-field comprised of circum-anal pore rings plus small groups of pores arranged as bands (Fig. 158). [Head, thorax, abdomen and wing-pads without sectasetae. BL =2-50-3-02 mm, WL = 1-15-1-16 mm, ARE = 0-08-0-09 mm, AWL = 1-40-1-46, BBBL = 0-91 , A = 10, R = 4689. Afrotropical. OnAntiaria.] TRIOZAMIA Anal pore-field comprised of circum-anal pore rings only 3 3 Head, abdomen and wing-pad margins with scales (Figs 173-177) 4 Head , abdomen and wing-pad margins without scales , usually with sectasetae 8 4 Marginal scales more than 3 times as long as broad (Fig. 173). [Forewing-pad with a well-developed humeral lobe. Marginal sectasetae absent. BL = 146 mm, WL = 0-62 mm, AWL = 0-65, BBBL = 0-76, A = 7, R = 3467. Hawaiian. On Pelea.] HEVAHEVA Marginal scales less than 2 times as long as broad 5 5 Dorsal surface of body and wing-pads with clavate setae (similar to Fig. 172). [Forewing-pad with a well-developed humeral lobe. Marginal sectasetae absent. BL = 2-06-2-31 mm, WL= 1-19-1-35 mm, ARE = 0-27-0-36 mm, AWL = 0-18-0-19, BBBL = 0-84-0-88, A = 2, R = 2222. Tropical New World. On Sideroxylon.} CEROPSYLLA (C. sideroxyli) Dorsal surface of body and wing-pads without clavate setae 6 6 Abdomen margin with sectasetae near abdominal apex, scales only present on basal three- quarters of abdomen margin (Fig. 177). [Forewing-pad with a well-developed humeral lobe.BL = 3-78 mm, WL = 2-18 mm, ARE = 0-43 mm, AWL = 0-16, BBBL = 0-74, A = 2, R = 1222. Hawaiian. On Pritchardia.] TRIOZA (T. palmicola) Abdomen margin without sectasetae 7 290 I. M. WHITE AND I. D. HODKINSON 7 Marginal scales ridged (Fig. 176). [Forewing-pad with a well-developed humeral lobe. BL = 3-11 mm, WL = 1-30 mm, ARB = 0-26 mm, AWL = 0-15, BBBL = 0-65, A = 2, R = 2222.Hawaiian. On Sideroxylon.] SWEZEYANA - Marginal scales not ridged (Fig. 175). [Forewing-pad with a well-developed humeral lobe. BL = 1-70 mm, WL = 0-95 mm, ARB = 0-21 mm, AWL = 0-20, A = 3, R = 1133. Hawaiian. OnPisonia.] KUWAYAMA (K. pisonia) 8 Forewing-pad margin with truncate sectasetae 9 Forewing-pad margin without truncate sectasetae 12 9 General form elongate (BBBL = 0-37-0-46). [BL = 2-59-2-69 mm, WL = 1-00-1-11 mm, ARB = 0-15-0-16mm, AWL = 0-18-0-24, A = 1, R = 1111. Australasian. On Casuarina.] AACANTHOCNEMAGeneral form broader (BBBL = 0-48-0-87) 10 10 Antenna with 8 or 9 divisions. Outer circum-anal pore ring comprised of a multiple row of pores. [BL = 2-59-3-12 mm, WL = 1-26-1-48 mm, ARB = 0-33-0-36 mm, AWL =0-37-0-40, BBBL = 0-52-0-56, R = 4688 or 4689. Afrotropical, Oriental and Palaearctic. On Rhamnus.] TRICHOCHERMES Antenna with 3-7 divisions . Outer circum-anal pore ring comprised of a single row of pores 11 11 On Ficus. Forewing-pad hind margin broadly rounded (Fig. 49). [BL = 1-41 mm, WL = 0-64-0-65 mm, ARB = 0-18-0-20 mm, AWL = 0-39-0-40, BBBL = 0-78, A = 4, R = 1244.Tropical Old World.] PAUROPSYLLA (P. trichaetd) - Not on Ficus. Forewing-pad hind margin usually rounded (Fig. 50). [BL = 1-15-2-95 mm, WL = 0-59-1-67 mm, ARB = 0-16-0-67 mm, AWL = 0-17-1-01, BBBL = 0-58-0-85, A = 3-7.]TRIOZA (most species of Trioza plus Paratrioza and some Ceropsylla species) 12 Abdomen margin without sectasetae [Several species which form enclosed galls and which cannot be separated by any tenable characters.]Abdomen margin with pointed sectasetae 13 13 Tarsal arolium without a visible unguitractor (Fig. 95). [Abdomen margin usually with more than a single row of sectasetae. Usually forming enclosed galls. BL = 1-98-3-05 mm, WL =1-10-1-54 mm, ARB = 0-21-0-43 mm, AWL = 0-21-0-60, BBBL = 0-51-0-79, A = 8 or 10,R = 3578 or 4689. Oriental and Palaearctic. On Populus, especially P. euphratica.] EGEIROTRIOZATarsal arolium with a clearly visible unguitractor 14 14 Dorsal surface of abdomen with clavate setae (Fig. 172). [Humeral lobe of forewing-pad extending anterior to eye. BL = 2-77 mm, WL = 1-54 mm, AWL = 0-18, BBBL = 0-87, A = 3, R = 3333. Hawaiian. On Tetraplasandra.] CRAWFORDA Dorsal surface of abdomen without clavate setae 15 15 Hindwing-pad very reduced (Fig. 48). [BL = 1-44-1-66 mm, WL = 0-76-0-85 mm, ARB = 0-32-0-35 mm, AWL = 0-46-0-53, BBBL = 0-59-0-67, A = 6, R = 3466. Austro-Oriental and Pacific. On Calophyllum.] LEPTYNOPTERA Hindwing-pad of normal proportions 16 16 Dorsal surface of abdomen with pointed sectasetae (Fig. 159). [BL = 1-74-1-98 mm, WL = 0-89-0-97 mm, ARB = 0-34-0-37 mm, AWL = 0-40-0-44, BBBL = 0-62-0-65, A = 6, R =3466. Palaearctic, introduced to New World. On Laurus and Persea.] TRIOZA (T. alacris)Dorsal surface of abdomen without sectasetae 17 17 Circum-anal pore rings convoluted (Fig. 156). [BL = 1-88 mm, WL = 0-73 mm, AWL = 0-51, BBBL = 0-48, A = 7, R = 3577. Tropical New World and Oriental. On Celtis and possibly FabaceaeandS/zoraz.] LEVRONOTA Circum-anal pore rings not convoluted (Fig. 160). [BL = 1-59 mm, WL = 0-77 mm, ARB =0-29 mm, AWL = 0-44, BBBL = 0-56, A = 1, R = 1111. Neotropical. OnPsidium.} TRIOZOIDA This study formed the basis of a Ph.D. project by the senior author, supervised by I. D. Hodkinson,Liverpool Polytechnic, and D. Hollis, British Museum (Natural History). Special thanks go to thefollowing colleagues at Liverpool Polytechnic: B. Duncan, M. Forster, J. Higgins, B. Midgeley, J. T.Rasmussen and A. J. K. Walley. I also thank all of the individuals and institutions who donated or lentspecimens (listed in Table 1), and the following for reasons too numerous to mention: V. K. Brown andR. G. 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Results from a study of the genus Glycaspis (Homoptera: Psyllidae). Australian Zoologist 15: 343-376.Nelson, G. 1971. Paraphyly and polyphyly: redefinitions. Systematic Zoology 20: 471-472.Ossiannilsson, F. 1970. Contributions to the knowledge of Swedish psyllids (Hem., Psylloidea) 1-4. Entomologica Scandinavica 1: 135-144.Pettey, F. W. 1924. South African Psyllids. Entomology Memoirs. Department of Agriculture, Union of South Africa 2: 21-30. 1925. New South African psyllids. South African Journal of Natural History 5: 125-142. 1933. New species of South African psyllids. Entomology Memoirs. Department of Agriculture, Union of South Africa 8:3-23.Pflugfelder, O. 1941. Psyllina. In Bronns, H. G. (ed.), Klassen und Ordnungen des Tierreichs. Volume 5, part 3, book 8. 95 pp. Leipzig. Platnick, N. 1. 1978. Gaps and prediction in classification. Systematic Zoology 27: 472-474.Platnick, N. I. & Nelson, G. 1978. A method of analysis for historical biogeography. Systematic Zoology 27: 1-16.Rahman, K. A. 1932. Observations on the immature stages of some Indian Psyllidae (Homoptera: Rhynchota). Indian Journalof Agricultural Science 2: 358-377.Raven, P. H. & Axelrod, D. 1. 1974. Angiosperm biogeography and past continental movements. Annals of Missouri Botanical Garden 61: 539-673. NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 295 Rohlf, F. J. 1972. An empirical comparison of three ordination techniques in numerical taxonomy. Systematic Zoology 21: 271-280.Rosen, D. E. 1978. Vicariant patterns and historical explanation in biogeography. Systematic Zoology 27: 159-188.Schaefer, H. A. 1949. Beitrage zur Kenntnis der Psylliden der Schweiz. Mitteilungen der Schweizerischen entomologischen Gesellschaft 22: 1-96.Schlee, D. 1969. Sperma-Ubertragung in ihrer Bedeutung fur das phylogenetische System der Sternorrhyn- cha. Phylogenetische Studien an Hemiptera I. Psylliformes (Psyllina + Aleyrodina) als mono- phyletische Gruppe. Zeitschrift fur Morphologic und Okologie der Tiere 64: 95-138.Schwarz, E. A. 1898. Notes on the lerp-insects (Psyllidae) of Australia. Proceedings of the Entomological Society of Washington 4: 66-75.Scott, J. 1882. On certain genera and species of the group of Psyllidae in the collection of the British Museum. Transactions of the Entomological Society of London 1882: 449-^473.1886a. Description of the nymph of Psylla mali, Schmidtberger. Entomologist's Monthly Magazine 22:281.18866. Description of the nymph of Psyllopsis (Psylla, Foerst.)fraxinicola. Entomologist's Monthly Magazine 22: 281-282.1886c. Note on Trioza remota, Foerster, together with a description of the nymph. Entomologist's Monthly Magazine 22: 282. Siegel, S. 1956. Nonparametric statistics for the behavioural sciences xvii + 312 pp. New York.Sneath, P. H. A. & Sokal, R. R. 1973. Numerical taxonomy xv + 573 pp. San Francisco.Szelegiewicz, H. 1971. Autapomorphous wing characters in the recent subgroups of Sternorrhyncha (Hemiptera) and their significance in the interpretation of the Paleozoic members of the group. Annales zoologici 29: 15-81. [In Polish, English translation - U.S. Dept. of Commerce, National Technical Information Service, Springfield, Virginia.] Takhtajan, A. 1969. Flowering plants - origin and dispersal, x + 310 pp. Edinburgh.Thome, R. F. 1976. A phylogenetic classification of the Angiospermae. Evolutionary Biology 9: 35-106.Tut hill, L. D. 1943. The psyllids of America north of Mexico (Psyllidae: Homoptera) (subfamilies Psyllinae andTriozinae). Iowa State College Journal of Science 17: 443-660. 1944. Contributions to the knowledge of the Psyllidae of Mexico. Journal of the Kansas Entomologi-cal Society 17 ': 143-159 . 1945. Contributions to the knowledge of the Psyllidae of Mexico. Journal of the Kansas Entomologi-cal Society IS: 1-29.1950. Contributions to the knowledge of the Psyllidae of Mexico. (Part II). Journal of the Kansas Entomological Society 23: 52-63. 1952. On the Psyllidae of New Zealand (Homoptera). Pacific Science 6: 83-125. 1959. Los Psyllidae del Peru Central (Insecta: Homoptera). Revista peruana de entomologia agricola 2: 1-27.1964a. Conociemientos adicionales sobre los Psyllidae (Homoptera) del Peru. Revista peruana de entomologia agricola 7: 25-32. 19646. Homoptera: Psyllidae. Insects of Micronesia 6(6): 353-376. Tuthill, L. D. & Taylor, K. L. 1955. Australian genera of the family Psyllidae (Hemiptera: Homoptera). Australian Journal of Zoology 3: 227-257.Uichanco, L. B. 1921 . New records and species of Psyllidae from the Philippine Islands, with descriptions of some preadult stages and habits. Philippine Journal of Science 18: 259-286.Vondracek, K. 1957. Mery Psylloidea. Fauna CSR 9: 1-431.1963. Jumping plant-lice (Psylloidea-Homoptera) of Central Africa. Part I (Congo). Sbornik Entomologickeho oddeleni Ndrodniho musea v Praze 35: 263-290.Watrous, L. E. & Wheeler, Q. D. 1981 . The outgroup comparison method of character analysis. Systematic Zoology 30: 1-11.White, I. M. 1980. Nymphal taxonomy and systematics of the Psylloidea (Insecta: Homoptera). xiv + 340, 12, 8, 4, 20, 8, 1 pp. Unpublished C.N.A. A. Ph.D. thesis, Liverpool Polytechnic.White, I. M. & Hodkinson, I. D. 1982. Psylloidea (Homoptera), nymphal stages, Handbooks for the Identification of British Insects (II)5b: 1-50. Yang, C-T. 1984. Psyllidae of Taiwan. Taiwan Museum Special Publication Series no. 3: 1-305.Young, D. J. & Watson, L. 1970. The classification of dicotyledons: a study of the upper levels of the hierarchy. Australian Journal of Botany 18: 387-433.Zimmerman, E. C. 1948. Insects of Hawaii 5: 12-38. Honolulu. 296 I. M. WHITE AND I. D. HODKINSON Index to host plants Acer 280, 282Aceraceae 258Aglaia 288Aquifoliaceae 256Anacardiaceae 256, 258, 274, 276, 278, 280, 288 Annonales 256, 261, 265, 266Antiaria 277, 289Arbutus 280, 283Arctostaphylos 280, 283Asteraceae257,281,282 Bauhinia 285Betulaceae 257, 288Boronia 283Brassicaceae 256, 281Buchanania 288Burseraceae 256, 258, 276, 288Buxaceae 258Buxus 287 Caesalpinoideae 286, 287Calluna 280, 281Calodendrum 280, 282Calophyllum 276, 290Caltha 282 Carex 176, 255, 279, 280, 281Casuarina 290Ceanothus 287Cedrela 289Celtis 277, 284, 290Cercocarpus285,287Chenopodiaceae 257, 281Clorophora 279, 281Colophospermum 277, 284Commelinales 256, 266Cyperaceae 256 Dicotyledoneae 255, 280 Erica 280, 281 Ericaceae 256, 257, 279, 280Ericales 265Erythrina 286 Eucalyptus 274, 277, 279, 283, 284Euphorbiaceae 257, 278, 285, 287,289 Fabaceae 256, 257, 258, 261, 274, 276, 277, 279, 281, 283, 284, 285, 286, 290Ficus 256, 274, 276, 277, 278, 279, 281,289,290Fraxinus 280, 283Fuchsia 283 Genisteae285,287Ilex 256, 278, 281 Juncaceae 256 Juncus 176, 255, 277, 278, 279, 281 Khaya 288 Lauraceae 256, 265, 266, 278, 279, 280, 281Laurus 290Lindera 280Litsea 280Loganiaceae 257 Malvaceae 257, 260 Malvales 256, 257, 258, 260, 275, 276, 278, 279, 280, 283, 289Mangifera 288Mangifera indica 277Melastomataceae 257, 276, 282Meliaceae 256, 258, 274, 276, 277, 278, 288, 289Mimosoideae 286Monocotyledoneae 255Moraceae 256, 257, 260, 274, 276, 278,279,281,283,285Moms 286Myricaria 256, 282Myroxylon 286Myrtaceae 257, 258, 274, 278, 279, 280, 283, 289 Nectandra 256, 278, 281 Olea 280, 283Oleaceae 257Onagraceae 257, 279, 280, 281, 282 Pelea 289Persea 290 Phillyrea 280, 283 Pisonia 290 Pistacia 280, 282 Polygonaceae 256, 281 Polygonum 282 Populus 276, 279, 280, 283, 290 Populus euphratica 290 Pritchardia 289 Proteaceae 257, 285, 286 Psidium 290 Purshia 287, 288 Ranunculaceae 256, 281Rhamnaceae 257, 258, 277, 284, 286, 288Rhamnus 290Rhus 282 Rosaceae 258, 274, 288Resales 256, 257, 261, 274Rumex 282Ruta 280, 282Rutaceae 256, 257, 258, 276, 279, 280, 288Rutales 256, 257, 258, 260, 261, 265, 266, 274, 275, 276, 278, 279, 280 Salicaceae 256, 257, 288 Salix 256 Santalaceae 274 Saxifragaceae 258 Schinus 282 Shorea 290 Sideroxylon 289, 290 Sisymbrium 282 Solanaceae 257, 258, 277, 278, 284, 285,286,289Sterculiaceae 257, 282Strychnos 283 Tamaricaceae 256, 280, 282Tamarix256,278,281,282Terminaiia 277, 278Tetraplasandra 276, 290Tiliaceae 282Tipuana 286Toona 277, 288Ulmaceae 257Ulmus 277Urticales 279, 280 Index Invalid and misspelt names are in italics; keys (pp. 275-290), place in general classification (table 17,p. 270), and other principal references are in bold. Names listed in tables 1,6, 11, 15, 17 and 18 are included. Aacanthocnema 167, 171 , 209, 212, Acaerus 270 213, 235, 267, 273, 290 aceris 159, 211, 236, 268 acaciae 162, 209, 213, 227, 234, 249, Acizzia 162, 209, 21 1 , 212, 213, 227,268 228,234,235,247,249,251,257, acaciaebaileyanae 162, 209, 213, 268, 271, 277, 285, 286 234,268 Acizziinae271,274, 285 Acizziini 271Aconopsylla 274aculeata!63,211acutipennis!61,210, 235aethiopica 159aggregata 165 NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 297 Agonoscena 159, 209, 211, 212, 213, 234, 267, 270, 280, 282 alacris 166, 211, 212, 213, 236, 267, 290 alba 164, 210, 235albagena 164, 21 1,236albifrons 167, 210, 235albitextura 165, 210, 234albiventris 166, 210, 235albomaculata 162, 211, 236Aleyrodoidea 238Alloeoneural54,251,272alni 154, 164, 168, 210, 235ambigua!64,211,236Amblyrhina 154, 25 1,272americana 164, 211, 236amorphae 163, 211, 235, 268Amorphicola 163, 211, 212, 213, 235, 248, 250, 251, 264, 268, 272,286 anceps 167, 211, 236, 269, 278angustipennis 160, 169, 210, 234annulata 164, 21 1,213, 236Anomalopsylla 155, 274Anomalopsyllinae 156, 160, 241, 242 Anomalopsyllini 160Anomocephala 273Anomoneura 162, 211, 212, 213, 235, 248, 249, 250, 257, 268, 271, 286 Anomoneurinae 271, 274, 285, 286Anomoneurini 162, 271, 285antennata 165, 211, 236, 268Aphalara 154, 155, 156, 160, 168, 170, 171, 210, 211, 212, 213, 234, 235, 237, 238, 247, 248, 256, 258, 261,263,267,270,282Aphalaridae 155, 156, 157, 159, 169, 171, 176, 179, 212, 213, 216, 219, 222, 226, 228, 237, 238, 242, 246, 247, 261, 264, 265, 266, 269, 270, 274, 276, 277, 278, 279Aphalarinae 154, 155, 160, 237, 266, 270, 279, 280, 281Aphalarini 160, 243, 270, 281Aphalaroida 154, 159, 179, 210, 211, 212, 213, 228, 235, 237, 247, 268, 269, 271, 277, 279, 281, 283Aphalaroidinae 269, 271, 279, 281, 283 Aphorma 270Apsylla 160, 209, 212, 213, 235, 238, 264, 265, 268, 272, 277, 288Apsyllinae 272Apsyllini 160, 241arbolensis 166, 210,236arbuti 159, 210, 236Arbutophila 159, 283arctostaphyli 159, 210, 235Aremica 272Arepuna 164, 176, 211, 212, 213, 235, 247, 257, 264, 268, 271, 277, 284 Arepuniinae 271, 274artemisiae 160, 169, 210, 234 Arytaina 154, 162, 211, 212, 213,235, 248, 249, 251, 268, 272, 287 Arytainilla 163, 210, 211, 212, 213,235, 248, 249, 251, 268, 272, 287 Arytaininae 162, 247, 251, 272, 274,285, 287 Arytainini 162, 251 Asphagidella 163, 213 Astragilita 274 atkasookensis 166, 210, 235 Atmetocranium 274 Auchmerina 271 Australopsylla 271 aylmeriae 166, 211,236 Bactericera 154, 155, 166, 273Bactericerinae 156, 166Baeopelma 163baileyi 165, 210, 234bakeri 167, 21 1,235beamed 167, 210, 235beesom'161,213,224,278bengalensis 247 betulaenanae 164, 210, 235, 268Bharatiana 161, 210, 213, 235, 238,241, 264, 265, 267, 272, 277, 288Bharatianinae 272bicolor 159, 21 1,235Brachypsylla 274Brachystetha 270brevistigmata 164, 210, 235brunneipennis 164, 210, 235buxi 163, 210, 235 Cacopsylla 163 Caillardia 270 Caillardiini 270 californical60,211,235 Calinda 273 calodendri 159, 179, 211, 236, 267 Calophya 154, 155, 156, 160, 168, 169,171,179,209,211,212,213, 226, 234, 235, 238, 241, 242, 260, 264, 267, 269, 272, 276, 278, 288Calophyidae 264, 265, 272, 274, 276, 278, 288 Calophyinae 160, 264, 272calthae 168Camarotoscena 159, 211, 212, 213, 235, 237, 246, 257, 267, 270, 276, 280, 283cannela 266capeneri 162, 211capense 162, 210Caradocia 271cardiac 162, 209, 234Cardiaspina 154, 165, 210, 212, 213, 234,235,269,271,284Cardiaspis 154 carpinicola 164, 210, 235, 268Carsidara 154, 155, 156, 238, 258, 265,273Carsidaridae 155, 156, 157, 160, 168,169,171,174,176,179,212, 213, 216, 219, 224, 226, 227, 237, 238, 241, 243, 261, 264, 265, 273,274, 278, 289Carsidarinae 154, 161, 240, 265, 273, 274, 275, 289Carsidarini 273Carsidaroida 274cassiae 163, 209, 235, 268casuarinae 167, 171, 209, 235, 267caudata 245 cayeyensis 163, 211, 236 ceanothi 163, 209 Ceanothia 163, 209, 211, 212, 213, 235,251,268,272,287ceardi 166Cecidopsylla 274Cecidotrioza 274cedrelae 16 1,209, 235celtidisgemma 165, 174celtidismamma 165celtidisvesiculum 165cercocarpi 165, 179, 210, 236, 247, 268 Ceriacreminae 154Ceriacremum 154Ceropsylla 167, 209, 21 1 , 212, 213, 228, 235, 267, 269, 273, 289, 290Cerotrioza 274chelifera 163, 210, 287chenopodii 166, 211, 235Chermes 154 Chineura 241, 272, 276,288chobauti 162, 209, 234cinereae 159, 209, 234cinnamomi 166, 211, 235Ciriacremidae 155Ciriacreminae 154, 155, 162, 272, 274, 285, 286 Ciriacremini 156, 162, 272Ciriacremum 154, 155, 156, 162, 176,210,211,212,213,228,235, 249,250,268,272,286,287cistellata 160, 209, 235, 268citri 162, 168, 210, 234clutiae 162, 210, 234cockerelli 166, 210, 235Colophorina 163, 209, 212, 213, 235,268,271,285coloradensis 162, 209, 235Colposcenia 160, 209, 212, 213, 235, 267, 270, 282Colposceniini 160, 243, 270, 278, 280, 281 Cometopsylla 271concolor 245conflecta 165conserta 165constricta 160, 210, 236coryli 164, 210, 235, 287Craspedolepta 160, 169, 209, 210, 211, 212, 213, 228, 234, 236, 256, 267, 270, 282Crastina 160, 21 1 , 212, 213, 236, 267, 270, 282Crawforda 167, 211, 212, 213, 236, 267, 273, 276, 290crefeldensis 162, 211, 236, 267 298 I. M. WHITE AND I. D. HODKINSON Creiis 154, 165, 209, 212, 213, 234, 247,269,271,284crithmi 166, 210, 235Ctenarytaina 165, 179, 210, 213, 227, 236, 244, 245, 267, 271, 280, 283 Ctenarytainini 264, 271, 279curta 160, 210, 234curvatinervis 166, 210, 235Cyamophila 264, 272, 286Cyamophilini 251, 264, 272, 286cyanoreios 165cytisi 163, 210 Delina 250, 272densitexta 165, 210, 235depressa 161, 168, 213, 224, 278deserata 165, 210, 236, 249, 268, 276Diaphorina 154, 155, 156, 162, 168, 170, 171, 176, 209, 210, 211, 212, 213, 216, 234, 236, 247, 256, 257, 267,271,276,283Diaphorini 162, 179, 219, 226, 237 (see also Diaphorinini)Diaphorininae 264, 269, 271, 279, 280,281,283Diaphorinini 247, 264, 271, 281 (see also Diaphorini: Becker- Migdisova)Diceraopsylla 274Diclidophlebia 161, 179, 211, 212, 213, 236, 237, 243, 244, 264, 267, 270, 282 Diclidophlebiini 161, 243, 270, 280diospyri 166, 174,211,235Diraphia 281divergipennis 165, 176, 211, 236, 268 dubia 160, 209, 235dugesii 161Dynopsylla 274Dynopsyllini 162 eastopi 161, 179, 21 1,236, 267Egeirotrioza 166, 170, 211, 212, 213, 228, 236, 237, 267, 273, 276, 290elongagena 167, 210, 236, 269Engytatoneura 274Epheloscyta 270Epicarsa 154, 161, 179, 211, 213, 219, 236, 238, 241, 256, 265, 267, 272, 276, 288Epipsylla 164, 209, 211, 212, 213, 236, 248, 249, 250, 268, 271, 286Epitrioza 273Eremopsylloides 270ericae 159, 210, 235Eriopsylla 274erythrinae!65,210,235erytreae 167, 212, 235eucalypti 165, 179, 210, 236, 244, 267Eucalyptolyma 165, 176, 179, 211, 212, 213, 227, 236, 244, 245, 267, 271,283Euceropsylla 163, 176, 210, 211, 212, 213, 234, 236, 250, 251, 268, 272, 286 Eudiaphorina 271Euglyptoneura 163, 179, 210, 211, 212, 213, 227, 236, 247, 251, 268, 272, 287Eumetoecus 270Euphalerinae 271Euphalerini 163, 264Euphalerus 154, 163, 170, 174, 176, 209, 210, 211, 212, 213, 224, 228, 234, 236, 247, 248, 250, 257, 258, 260, 264, 268, 269, 271, 272, 277, 284, 285, 286euphratica 166Euphyllura 154, 159, 211, 212, 213, 236,245,257,267,270,283Euphyllurinae 269, 270, 279, 280, 282 Euphyllurini 159, 237, 244, 270, 280Eurhinocola 271Eurotica 270Eutrioza 273Eutriozini 166exilis 160, 21 1,235 fabulosa 162, 210, 236, 267 falcata 167, 212, 236 fici 162, 179, 210 ficus 162, 179, 213, 258, 260 flava 164 flavida 160, 209, 234 floccosa (Pseudophacopteron) 161, 179,213,241,267floccosa (Psylla) 164, 210, 235, 268floccosa (Synoza) 162, 213florea 162, 210, 234Fiona 154, 163, 211, 212, 213, 236, 248,249,251,268,272,287foersteri 163, 21 1,236fonscolombii 164, 210, 235, 268forcipata 250fraxini 163, 210fraxinicola 163, 210, 235fremontiae 160, 174, 179, 211, 236, 243,267Freysuila Aleman 165, 211, 212, 213, 236, 249, 257, 268, 271, 285, 286 Freysuila sensu Schwarz 154frontalis 166, 21 1,235furcata 160, 210, 234fusca 160fuscinodulus 250fuscipennis (Eucalyptolyma) 245fuscipennis (Euglyptoneura) 163, 211 galeaformis 164, 210, 235, 268gallicolus 163, 174, 176, 211, 212, 213, 224, 236, 247, 257, 269, 277, 284 gardenensis 162, 210Geijerolyma 271genistae!62,211,235,268gigantea!61,168,210,235 glabruscuta 163, 176, 210, 236, 268gladiatum 162, 176, 210Glycaspis 156, 165, 210, 212, 213, 227,234,269,271,284gossypii 159, 210, 235gressitti 241Gyropsylla 160, 210, 212, 213, 234, 256, 260, 266, 269, 270, 278, 281Gyropsyllini 270 hakani 163, 210 hakeae 163, 211, 213, 235, 257, 268hamata 164, 210, 235Haplaphalara 270harteni 162, 21 1,286Hemischizocranium 273Hemitrioza 273Hepatopsylla 164, 287Heteropsylla 154, 165, 179, 211, 212, 213, 236, 249, 250, 268, 272, 276, 286 Heterotrioza 166Hevaheva 167, 211, 212, 213, 236, 269, 273, 289hibisci!61,176,210,235hirsuta (Psylla) 164, 210, 235, 287hirsuta (Trioza) 166, 212, 213, 236, 237, 269, 277, 278Holotrioza 242, 272Homoptera 154Homotoma 154, 156, 162, 170, 179, 211, 213, 236, 238, 241, 258, 260, 264, 267, 272, 289Homotomidae 264, 265, 272, 274, 276, 278, 288Homotominae 162, 226, 241, 264, 272 Homotomini 162hyalina 250Hyalinaspis 271 ilicis 160, 210, 234 imponens 165 incidata 166, 211,235 incisa 165 indica 162, 213 inermis 159, 21 1,212, 277, 281 ingae 250 Insnesia 163, 176, 210, 212, 213, 236,251,257,268,272,287Isogonoceraia 165, 176, 211, 212, 213,236,251,268,272,286Izpania 273 japonica 165, 247, 284 Jenseniella 274 jugovenosus 163, 210, 213, 224, 234, 257, 284 julbernardioides 162, 211, 286juncorum 162, 211, 236 Katecephala 271Kleiniella 155, 250, 272Kuwayama 167, 211, 212, 213, 236,269, 273, 290 NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 299 Labicria 274 lamborni 166, 212, 236, 269 Lanthanaphalara 274 Lasiopsylla 271 lata 160, 269 lavaterae 166, 210, 236 lentiginosum 161,211, 236, 267 Leptotrioza 274 Leptynoptera 161, 174, 211, 213, 236, 238, 265, 267, 273, 276, 290Leptynopterinae 161, 238Leurolophus 159, 211, 212, 213, 236, 267, 270, 280, 282Leuronota 166, 211, 212, 213, 236, 267, 273, 290Levidea 274Ligustrinia 271linavuorii 160, 21 1,236, 267Lindbergiella 274Lisronia 270litseae 166, 21 1,235Livia 154, 156, 162, 176, 209, 211, 213, 235, 236, 238, 242, 255, 256, 261, 264, 267, 269, 270, 277, 278, 281Liviidae 155, 157, 158, 162, 212, 213,226,261,264Liviinae 154, 155, 264, 266, 270, 279, 280, 281Livilla 25 1,272Livillinae 154lobata 166, 212, 236longicornis 165, 211, 236, 268longispiculata 242 Macrocorsa 271Macrocorsini 271Macrohomotoma 162, 176, 210, 211, 213, 236, 264, 269, 272, 277, 278, 289 Macrohomotominae 272, 276Macrohomotomini 272maculata 242 maculipennis (Livia) 162, 209, 235maculipennis (Paratrioza) 166, 210, 235 magna 164, 21 1,236, 285magnicauda 164, 210, 235magnoliae 166, 212, 236maideni 245 malayensis!61,209,235mali 163, 210, 213, 235, 268marginepunctata 166, 211, 235martorelli 167, 209, 267Mastigimas 154, 161, 209, 210, 213, 235, 238, 256, 265, 269, 273, 274, 289 Mastigimatinae 273, 289Mastigimatini 161matsumurana 157, 161, 174, 179, 211,236,267,275media 164, 210, 236Megadicrania 274Megatrioza 166melanoneura 164, 210, 235menoni 244 Mesohomotoma 161, 176, 209, 210, 213, 235, 269, 273Mesohomotomini 161Metapsylla 274Metatrioza 273 michoacana 166, 211, 236, 267Microceropsylla 161, 169, 211, 212, 213, 236, 240, 241, 264, 269, 272, 278, 288 Microceropsyllini 161, 241minor 164, 210, 235minuta (Craspedolepta) 160, 209, 234 minuta (Phytolyma) 160, 279minuta (Psylla) 164, 210, 235, 287minuta (Trioza) 166, 210, 235minuticona 163, 210, 234minutissima 160, 209, 234Mitrapsylla 165, 210, 212, 213, 236, 248, 249, 268, 271, 276, 285, 286monticola 160, 21 1,235Moraniella 159, 179, 211, 212, 213, 236, 243, 267, 270, 280, 282mori 162, 21 1,235, 250, 268moscovita 164, 210, 235Mycopsylla 154, 162, 179, 210, 213, 235, 260, 265, 269, 272, 278, 289Mycopsyllini 272Myrmecephala 273myrtilli 164, 210, 235 nebulosa 160, 21 1,236 negundinis 164, 211, 236 Neocalophya 242 Neolithinae 273 Neolithus 166, 21 1,212, 213, 236, 269, 273, 278, 289Neophyllura 159, 210, 211, 212, 213, 235,236,267,271,283Neopsyllia 165, 176, 210, 212, 213, 235,249,268,271,285,286Neotrioza 273Neotriozella 273nervosa 160, 210, 234Nesiope 274nidifex 163, 211, 212, 213, 224, 234, 247, 284, 286nigricornis 166, 212, 236nigripennis 160, 209, 234nigripes 249nigrita 164, 210, 235nubifera 160, 210, 234nyasae 162, 21 1,236, 267 oblonga 250obsoleta 166, 212, 236obtusa 166, 212, 236octospinosa 161, 210, 235, 267olivina 159Optomopsylla 274orientalis 165Osmopsylla 164Ozotrioza 273 Pachyparia 271 Pachypsylla 154, 165, 170, 174, 210, 212, 213, 235, 247, 261, 269, 271, 284 Pachypsyllinae 165, 247, 271, 277Pachypsylloides 270Pachypsylloidini 270Palmapenna 250, 272palmeni 164, 21 1,236palmicola 166, 210, 236, 255, 269, 289panacis 167, 212, 235Panisopelma 250, 272Paracalophya 242, 272Paracarsidara 161, 168, 210, 213, 235, 238, 256, 260, 269, 273Paracomeca 273Paracomecini 166parallela 164, 210, 235Paraphalaroida 160, 174, 179, 211, 212, 213, 226, 236, 237, 243, 267,270, 282 Paratrioza 166, 210, 212, 213, 228, 235, 236, 267, 273, 290paucivena 241Paurocephala 155, 159, 170, 174, 179, 210, 212, 213, 219, 235, 237,238, 243, 244, 246, 250, 257, 258,260, 262, 267, 270, 276, 280, 283 Paurocephalinae 159, 246, 266, 269,270, 279, 280, 283 Paurocephalini 159, 246 Pauropsylla 154, 155, 161, 168, 211, 213, 219, 224, 228, 236, 238, 242,246, 262, 264, 265, 267, 269, 273,278, 290 Pauropsyllinae 161, 241, 246, 250Pauropsyllini 161, 238, 241, 242, 273Paurotriozana 274Pelmatobrachia 161, 174, 211, 213, 236, 241 , 264, 269, 272, 278, 288Pennavena 162, 210, 212, 213, 236, 247,257,267,271,283peregrina 163, 21 1,236persicaria!60,211,235,237Pexopsylla 165, 179, 210, 212, 213, 236, 247, 268, 274Phacopterinae 161, 226, 241 (see also Phacopteroninae)Phacopterini 161Phacopteron 154, 161, 211, 213, 236, 238, 260, 265, 267, 272, 276, 288Phacopteronidae 264, 265, 272, 274, 276, 277, 278, 288Phacopteroninae 264, 272 (see also Phacopterinae: Becker- Migdisova)Phacopteronini (see Phacopterini: Becker-Migdisova)Phacosemoides 241, 272Phellopsylla 165, 179, 211, 212, 213, 227, 236, 247, 257, 267, 271, 277, 284 phillyreae 159, 212phoradendrae (Psylla) 164, 209, 213, 236, 268phoradendrae (Trioza) 167, 210, 236 300 I. M. WHITE AND I. D. HODKINSON Phytolyma 154, 155, 156, 160, 211, 212, 213, 236, 242, 256, 267, 269,270, 278, 279, 281 Phytolymini 160, 270 pisonia 167, 211, 213, 236, 269, 290 pithecolobia 159, 179, 210, 212, 247, 277, 279Platycorypha 165, 176, 210, 212, 213, 235, 249, 268, 271, 285, 286polygoni 160, 210, 234princeps 165, 176, 210, 235, 268Prionocnemidae 154 Protyora 161, 211, 213, 236, 269, 273 pruni 164, 21 1,236Pseudacanthopsylla 274Pseudoeriopsylla 162, 211, 213, 236, 265, 267, 272, 289Pseudophacopterini 161Pseudophacopteron 161, 179, 211, 213, 228, 236, 238, 241, 265, 267, 269, 272, 277, 278, 288Pseudophacopteronini (see Pseudophacopterini: Becker- Migdisova)Pseudotrioza 273Psylla 154, 155, 156, 157, 163, 164, 168, 174, 179, 209, 210, 211, 212, 213, 227, 235, 236, 238, 247, 249, 250, 251, 255, 256, 258, 261, 268, 272, 277, 285, 287, 288Psyllidae 154, 155, 156, 157, 162, 169, 170, 174, 176, 179, 212, 213, 216, 219, 222, 224, 226, 227, 237, 238, 241, 247, 261, 262, 264, 271, 274, 276, 277, 285Psyllinae 154, 155, 163, 228, 272, 274, 285, 287Psylloidea 154, 156, 157, 168, 179, 228, 238, 255, 256, 258, 260, 264, 265, 270, 275 Psyllopseini 163, 247, 264, 271Psyllopsis 154, 155, 163, 179, 210, 212, 213, 216, 219, 226, 235, 237, 247,257,267,271,280,283pubescens 163, 21 1,287pulchella 164, 21 1,236, 285pulchra 164, 210, 235, 267punctulata 162, 210, 234Purshivora 163, 210, 21 1 , 212, 213, 236,251,268,272,287putonii 162, 210, 234pyri 164, 21 1,235pyricola 164, 210, 235pyrisuga 164, 210, 235 quadripunctata 167, 210, 235 remota 166, 212, 236 Retroacizzia 165, 211 , 212, 213, 236, 247,268,271,277,284Rhachistoneura 270rhamnicola 164, 211, 236Rhegmoza 273Rhinocola 154, 159, 21 1 , 212, 213, 236, 242, 263, 268, 270, 280, 282 Rhinocolinae 266, 270, 276, 279, 280, 282 Rhinocolini 159, 270, 279, 282Rhinopsylla 154, 274Rhodochlanis 270rhododendri 164, 211,236rhois 160, 21 1,235, 269, 276Rhombaphalara 270ribesiae 164, 210, 213, 235, 268rivalis 165 robusta 163, 179, 210, 247rotundipennis 160, 169, 211, 212, 213, 235, 269, 278rugipennis 163, 210, 213, 224, 234, 284 rumicisl60,211,235russellae 163, 211, 213, 235, 268Russelliana 272russoi 163, 210, 234 salebrosa 165 saliceti 164, 210, 235 salicivora 166, 210, 235 Schedoneolithus 273 Schedotrioza 273 schini (Calophya) 160, 211, 212, 213, 235schini (Tainarys) 160, 210, 235, 268 sicki 241 sideroxyli 167, 21 1,269, 289silvestris 167, 212, 236, 267simila 160, 210, 234simlae 164, 209, 236, 285, 287sinuata 164, 210, 235solani 162, 21 1,236sonchi 160, 210, 234sorbi 163, 21 1,235Spanioneura 154, 164, 210, 212, 235,251,268,272,287spartiicola 163, 211spartiophila 163, 211speciosa 159, 212, 213, 267spegazziniana 160, 210Sphingocladia 274Sphinia 274Spondyliaspidae 155 (see also Spondyliaspididae)Spondyliaspididae 155, 157, 165, 168, 170, 179, 212, 213, 227, 237, 242, 244, 247, 261, 264, 269, 271, 274, 277, 284, 286 (see also Spondyliaspidae:auctt . )Spondyliaspidinae 165, 174, 176, 222, 244, 271, 274, 277, 284 (see also Spondyliaspinae:auctt.)Spondyliaspinae 154, 155 (see also Spondyliaspidinae)Spondyliaspis 154, 156, 165,210, 212, 213, 236, 238, 258, 269, 271, 284 squamula 165,210,234Stenopsylla 273sterculiae 161, 21 1,236, 269Sternorrhyncha 154Stigmaphalarini 243 striata (Macrohomotoma) 162, 176, 211,277 striata (Psylla) 164, 210, 235, 268stricklandil63,211,236Strophingia 156, 157, 159, 209, 210, 212, 213, 234, 235, 238, 242, 247, 256,261,265,268,270,281Strophingiinae 266, 270, 279, 280, 281 suaedae 160, 210, 234subpunctata 160, 210, 236subspiculata 164, 211, 236sulfurea 161, 174, 211, 236, 267Swezeyana 167, 210, 212, 213, 236, 269, 273, 290swezeyi 167, 21 1,236, 269Sympauropsylla 273Syncarpiolyma 271Syndesmophlebia 155Synoza 162, 211, 213, 236, 241, 260, 265, 267, 269, 272, 276, 289Synoziini 162Syntomoza 271Syringilla 271 Tainarys 155, 156, 160, 210, 212, 213, 235, 243, 260, 268, 270, 280, 282tantillus 163, 209, 213, 224, 236, 258, 268, 285Tenaphalara 156, 161, 209, 210, 211, 213, 227, 235, 236, 237, 240, 269, 273 Tenaphalarinae 161, 240, 243Tenaphalarini 161, 273tessmanni 161, 209Tetragonocephala 157, 165, 210, 212,213,236,269,271,284texana 165Thamnopsylla 164thysanura 245Togepsylla 157, 161, 174, 179, 211, 213, 236, 244, 261, 264, 265, 267, 270, 275, 278, 281Togepsyllinae 270, 279, 280, 281Togepsyllini 161torifrons251trichaeta 161, 213, 219, 224, 267, 290Trichochermes 166, 210, 212, 213, 236, 240, 267, 273, 290Trigonon 165, 211, 212, 213, 236, 248,257,268,271,285,286trimaculata 164, 210, 235, 268triopsyllina 167, 21 1,236, 267Trioza 154, 156, 157, 166, 168, 174, 210, 211, 212, 213, 228, 235, 236, 237, 238, 240, 255, 256, 258, 260,267, 269, 273, 277, 278, 289, 290 Triozamia 155, 156, 166, 212, 236,269, 273, 277, 289 Triozamiinae 166, 273 Triozidae 155, 156, 157, 166, 168,169, 170, 171, 174, 176, 179, 212,213, 216, 219, 222, 224, 227, 237, NYMPHAL TAXONOMY AND SYSTEMATICS OF THE PSYLLOIDEA (HOMOPTERA) 301 238, 261, 262, 265, 273, 274, 275, 276, 277, 278, 289Triozinae 154, 155, 166, 273Triozinil66,265,273Triozoida 167, 212, 213, 236, 267, 273, 290 triozomima 160, 209, 235tripunctata 166, 21 1,235tuxtlaensis 244Tyora 154, 274 ulmi 163, 179,211,235,277 uncatoides 163, 211, 213, 235, 268unicolor 159, 212, 213, 267urenae 159, 210, 235urticae 166, 168, 212, 236 vancouverensis 160, 210, 234variegata 163, 21 1,236, 268veaziei 160, 210, 234venusta 165, 210, 235,284vermiculosus 163, 210, 213, 224, 234, 284vernalis 162, 211,236,267 verrucosa 242verucifica 166visci 164, 210, 235vitiensis 166, 21 1,235vitreoradiata 167, 212, 236vittatus 159, 21 1,236, 267 walked 166, 210, 236, 267 Xanioptera 270Xenaphalara 270Xenaphalarini 270 British Museum (Natural History) Milkweed butterflies: their cladistics and biology P. R. Ackery & R. I. Vane- Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects oftheir biology have stimulated far more attention than can be justified by species numbersalone. In recent years, an expansive literature has grown, considering aspects of theircourtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry andgenetics. The popularity of danaines among biologists can certainly be attributed to thiscombination, within one small group, of so many of the factors that make butterflies such aninteresting group to study. The obvious need to place this wealth of biological data within anacceptable systematic framework provided the impetus for this volume. Started eight years ago within the conventions of evolution by natural selection andHennig's phylogenetic systematics, the book is now largely about natural history (what theanimals have and do, where they live and how they develop) and natural groups - as revealedby a form of analysis approaching that practised by the new school of 'transformed cladistics' .The authors have prepared a handbook that will appeal to a wide range of biologists, frommuseum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography.ISBN 565 00893 5. 1984. Price 50. Titles to be published in Volume 50 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera) By Peter S. Broomfield Nymphal taxonomy and systematics of the Psylloidea (Homoptera). By I. M. White & I. D. Hodkinson The whitefly of New Guinea (Homoptera: Aleyrodidae) ByJ. H.Martin Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural History) The whitefly of New Guinea(Homoptera: Aleyrodidae) J. H. Martin BRITISH MUSEUM (NAVtflUL HIS i- - 3 .. , ^5 .TED ^'.'XL I. Entomology seriesVol50 No 3 30 May 1985 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in fourscientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique andever-growing collections of the Museum, both by the scientific staff of the Museum and byspecialists from elsewhere who make use of the Museum's resources. Many of the papers areworks of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself,available separately, and individually priced. Volumes contain about 300 pages and severalvolumes may appear within a calendar year. Subscriptions may be placed for one or more ofthe series on either an Annual or Per Volume basis. Prices vary according to the contents ofthe individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History),Cromwell Road, London SW75BD,England. World List abbreviation: Bull. Br. Mas. nat. Hist. (Ent.) Trustees of the British Museum (Natural History), 1985 The Entomology series is produced under the general editorship of the Keeper of Entomology: Laurence A. Mound Assistant Editor: W. Gerald Tremewan ISBN 565 06010 4ISSN 0524-6431 British Museum (Natural History)Cromwell RoadLondon SW7 5BD Entomology seriesVol50No3pp303-351 Issued 30 May 1985 The whitefly of New Guinea (Homoptera:Aleyrodidae) J. H. Martin BRITISH (NATURAL HIS. Department of Entomology, British Museum (Natural History), Cromwell Road, LondonSW7 5BD Contents Synopsis .................................................................................................... 303 Introduction .............................................................................................. 303 Depositories .............................................................................................. 304 Acknowledgements ..................................................................................... 304 Checklist of New Guinea Aleyrodidae, their hosts in New Guinea, and their depositories ............................................................................................ 304 Key to whitefly subfamilies and genera of New Guinea ......................................... 309 Aleurodicinae ............................................................................................ 311 Aleyrodinae ............................................................................................... 313 References ................................................................................................ 337 Index ........................................................................................................ 351 Synopsis The whitefly fauna of New Guinea, including Irian Jaya (Indonesian New Guinea) and the smaller islandprovinces of Papua New Guinea, is reviewed. Fourteen species are described as new, six are newlysynonymized, and five new combinations are established. The checklist of whitefly from New Guineaincludes host data and depositories and those whitefly not determined to species. Keys to the subfamiliesand genera occurring in New Guinea are provided, to the named species of Aleurocanthus from NewGuinea, and to the described species of Parabemisia, Xenaleyrodes and tropical Asian Aleurodicus. Introduction In their systematic review of the whitefly of the world, Mound & Halsey (1978) listed 1156described species; only five were recorded from New Guinea, based upon material in the BritishMuseum (Natural History). Four of these species, Aleurodicus destructor, Bemisia tabaci,Neomaskellia bergii and Trialeurodes vaporariorum, are widely distributed and well-knownpests which had been submitted for identification by agricultural organisations. Previouslypublished records of three species from Papua New Guinea and Irian Jaya (Dumbleton, 1954)were not included in Mound & Halsey 's review. In an account of crop pests of Irian Jaya (WestIrian), Thomas (1962) lists nine species of whitefly, but careful scrutiny of the introduction to thework indicates that only one (an undetermined Aleurocanthus) had actually been recorded fromthere; the remaining species were from elsewhere in Indonesia, and from Territory of Papuaand New Guinea' (Dumbleton, 1954). Dumbleton, (196 la; 19616) specifically stated that thewhitefly fauna of New Guinea was practically unknown, and no publication has appeared sinceto change the situation. In 1979 the author joined the expedition 'Operation Drake' at its scientific base camp at Buso(7 25 'S, 147 15'E) on the Morobe Province coast of Papua New Guinea, some 80 km south-eastof Lae. In the vicinity of Buso, whitefly specimens were collected from angiosperm hostsrepresenting over 60 genera in 35 families, as well as from a number of unidentified hosts; the 780slide-mounted specimens almost certainly comprise the first significant collection of the group tohave been made in New Guinea, over 80 whitefly species being represented as a result of threemonths collecting in the area. This figure may be compared with 30 records of described speciesfrom Australia, 31 from New Caledonia and 11 from New Zealand, the whitefly faunas of these Bull. Br. Mus. nat. Hist. (Ent.) 50 (3): 303-351 Issued 30 May 1985 304 J. H. MARTIN areas being relatively well studied (Mound & Halsey, 1978). The Buso collection forms the basisof this paper, which also includes other New Guinea records. The taxonomy of whitefly is based upon the exuviae of the final (fourth) instar larvae, usuallyreferred to as the 'pupal cases'. Mound (1963) has demonstrated by host-transfer experimentswith Bemisia tabaci (Gennadius) that some whitefly species are polyphagous and morphological-ly variable. As in B. tabaci, this has undoubtedly led to different names being applied to variantswithin many species, and consequently only 14 of the more distinctive species are described hereas new, bringing the total of named species to 37, although over 90 are known to occur. Thechecklist includes partially identified material examined from New Guinea, their host data anddepositories; of the 54 undetermined species, it is probable that the majority are undescribed. The descriptions which follow use the standard whitefly terminology, which is here illustratedwith a schematic diagram (Fig. 48), and is similarly visually explained by Cohic (1966), David &Subramaniam (1976) and Bink-Moenen (1983); Russell (1943) also clarifies a number of termsused in whitefly descriptions. Depositories BMNH British Museum (Natural History), London BPBM Bernice P. Bishop Museum, Honolulu CSIRO Central Scientific & Industrial Research Organisation, Canberra DPIQ Department of Primary Industry, Indooroopilly, Queensland DSIR Department of Scientific & Industrial Research, Auckland HDA Department of Agriculture, Honolulu TARI Taiwan Agricultural Research Institute, Taipei UMO University Museum, Oxford USNM United States National Museum of Natural History, Washington DC Acknowledgments The author is particularly indebted to the organisers of 'Operation Drake', and to all those involved withthe Buso scientific camp, without whose support the collection of most of the material referred to in thispaper would not have been possible. Thanks, too, are due to Ted Henty and his staff at the Lae Herbarium(Papua New Guinea Department of Primary Industry), and to those botanists who worked at Buso, fortheir help in establishing the identities of many of the host plants. The help of CSIRO, Canberra, theBernice P. Bishop Museum, Honolulu and DSIR, Auckland (who all loaned material for study) is alsoacknowledged with thanks. Checklist of New Guinea Aleyrodidae, their hosts in New Guinea, and their depositories * denotes discussion of undetermined species in main text ALEYRODIDAE HOST FAMILY HOST GENUS/SPECIES DEPOSITORY ALEURODICINAEALEURODICUS Douglas,1892destructor Mackie, 1912 /io/mes//(Maskell, 1895) ALEYRODINAEALEUROCANTHUS Quaintance & Baker, 1914cocoisCorbett, 1927esaMTakahashi, 1936 LauraceaeMoraceae Palmae ?Xanthophyllaceae Sapindaceae SterculiaceaeSapindaceae Cinnamomum BMNH Ficus microcarpa var. BMNH naumannii Cocos nudfera BMNH IXanthophyllum BMNH Guioa BMNH indet. BMNH Pometia pinnata BMNH THE WHITEFLY OF NEW GUINEA * denotes discussion of undetermined species in main text 305 ALEUROTUBERCULA TVS Takahashi, 1932 306 J. H. MARTIN * denotes discussion of undetermined species in main text ALEUROTULUS Quaintance & Baker, 1914anindJJiacea Singh, 1931ASIALEYRODES Corbett,1935asp. 1* ? sp. 2* BEMISIA Quaintance &Baker, 1914a/er (Priesner & Hosny, 1934) leakii (Peal, 1903)pongamiae Takahashi, 1931tefeac/(Gennadius, 1889) sp. 1 CRENIDORSUM Russell ,1945 lasangensissp.n.nwrohensissp. n. sp. 1*DIALEURODES (Cockerell, 1902)decaspermi sp. n.kirkaldyi (Kolinsky, 1907)psidii Corbett, 1935a Gramineae EuphorbiaceaeCelastraceae Leguminosae AraceaeLeguminosaeConvolvulaceaeEuphorbiaceae LeguminosaeLeguminosae MusaceaeMyrtaceae ?Araliaceae MyrtaceaeOleaceae CelastraceaeDipterocarpaceae Ebenaceae Euphorbiaceae Gyrocarpaceae Moraceae Myrtaceae NaucleaceaeRutaceae indet. bamboo PimelodendronemboinicumSalacia Pterocarpus ?indicus Colocasia c.v.Denis Ipomoea batatasManihot esculenta c.v. & utilissimaSophora tomentosaDalbergia Musa DecaspermumMyrtellatSchefflera DecaspermumJasminum LophopetalumAnisoptera thurifera polyandraDiospyrosMacarangaGyrocarpusFicus DecaspermumMyrtellaNeonaucleaEuodea BMNH BMNHBMNH BMNH BMNHBMNHBMNHBMNH BMNHBMNH BMNHBMNHBMNHBMNH BMNH;USNMBMNH;BPBM BMNHBMNH BMNH BMNH BMNH BMNH BMNH BMNH BMNH;BPBM BMNH THE WHITEFLY OF NEW GUINEA denotes discussion of undetermined species in main text 307 ORCHAMOPLA TVS Russell ,1958niuginiisp. n. ?niuginiisp. n.PARABEMISIA Takahashi,1952 Jauan/sp. n.myrmecophtia sp. n. Guttiferae Linaceae ?Celastraceae indet.Dipterocarpaceae Saccharum officinarum BMNH;BPBM Calophyllum inophyllum BMNH;USNMDurandea BMNH ILophopetalum BMNH Anisoptera BMNHBMNH 308 J. H. MARTIN * denotes discussion of undetermined species in main text THE WHITEFLY OF NEW GUINEA 309 * denotes discussion of undetermined species in main textALEYRODIDAE HOST FAMILY HOST GENUS/SPECIES DEPOSITORY genus indet. 4, Myrtaceae Decaspermum BMNHIndoaleyrodeslDialeurodesgroup Key to whitefly subfamilies and genera of New Guinea The taxonomy of whitefly at the generic level is in such a poor state that keys like those of Sampson &Drews (1956) and David & Subramaniam (1976) frequently give misleading results when species somewhatatypical of their allotted genera are encountered. The tendency for many such species to be placed in new,monobasic genera has done nothing to clarify generic concepts. The key below will enable genera knownfrom New Guinea to be identified, but, in cases where the described species within a particular genus are sovariable as to make the genus difficult to define, some are keyed out at species level. Pupal cases 1 Subdorsum with large, ornate, compound pores, often with central spatulate processes, 4-6 pairs on abdomen and 1 cephalic pair; lingula large and excluded from vasiform orifice,bearing 4 hairs (ALEURODICINAE) ALEURODICUS(p. 311) - Subdorsum without large, ornate, compound pores (although it may possess 5 pairs of large, simple pores) ; lingula not as above (ALEYRODINAE) 2 2 Vasiform orifice transversely eliptical, elevated, usually appearing wider than long in slide- mounted specimens.Outer submargin with a single row of about 16 pairs of long hairs (Fig. 38); cuticle pale to brown. Dense, ant-attended colonies on blades of Gramineae NEOMASKELLIA(p. 329) If vasiform orifice elevated, then at least as long as wide 3 3 Submargin with a row of dentate glands (Fig. 22) . Margin weakly crenulate to finely toothed, but thoracic and caudal tracheal teeth alwaysstrongly differentiated from remainder of margin (Fig .23) ORCHAMOPLA TVS (p . 329) - Submargin without dentate glands 4 4 Inner submargin, and often also much of dorsal disc, with long, stout spines which have pointed, laciniate or rounded apices.Vasiform orifice often elevated 5 - Inner submargin without row of long, stout spines, although short, lanceolate setae sometimes present in outer submargin 6 5 Stout spines restricted to a single submarginal ring only; spines tubiform, much thicker in basal two-thirds and narrowing rather suddenly before apical third; spines may be angled sharplyat constriction (Fig. 35); true margin curled downwards, usually concealed by dorsum inslide-mounted specimens XENALEYRODES(p. 334) - In addition to submarginal ring, stout spines often occurring on dorsal disc (Figs 1 , 41 , 44, 45) , even if only one or two such additional pairs of spines present (as in Fig. 3); true margin often not curled downwards ALEUROCANTHUS(p.313) 6 Margin rather irregular, somewhat crenulate; without, or with only slight differentiation of margin at thoracic tracheal openings. Vasiform orifice triangular, sometimes sinuate lateral-ly, with apex leading into a pronounced caudal furrow; operculum occupying basal half oforifice, with triangular to elongate-oval head of lingula occupying most of remainder.Posterior margin of case normally indented, but often without obvious differentiation. Outersubmargin without a row of hairs. Pupal cases pale, not noticeably waxy in life BEMISIA (Fig. 10) (p. 321) - Combination of characters not as above 7 7 Margin differentiated at caudal and/or thoracic tracheal areas: margin may be indented as a 'pore'; or a 'comb' of teeth distinct from remainder of margin may be present, this combstanding proud, or indented. Tracheal folds often marked on venter, running mesad frommargin (as in Figs 8, 20) 13 - Margin undifferentiated at areas of tracheal openings (as in Figs 7,14) 8 8 Dorsal disc completely separated from rather wide submarginal area by a distinct line or fold concentric with margin.Pupal cases black TETRALEURODES-group (p. 333) 310 J. H. MARTIN - Dorsal disc not so separated from submarginal area 9 9 A longitudinal furrow present on each side of submedian area of dorsal disc in thoracic, and sometimes also anterior abdominal, area (as in Figs 12, 15) 10 - A longitudinal furrow not present on each side of submedian area of dorsal disc 11 10 Margin complex: cuticular markings in outer submargin usually give margin the appearance of having a double row of teeth. Pupal cases usually dark brown or black ALEUROTRACHELUS(p. 320) - Marginal teeth simple (Figs 13, 16). Pupal cases of New Guinea species pale CRENIDORSUM(p. 323) 11 Submargin either with long setae (minimally 3 pairs), or with a row of short, stout, lanceolate setae; margin weakly crenulate, not coarsely toothed 12 - Submargin without long, simple setae and without short, lanceolate setae; margin (high power detail) apparently double: one row of shallow crenulations and one row of more obviousregular teeth (Figs 7, 8) ALEUROMARGINATUS(p. 318) 12 Pupal case elongate, parallel-sided and slightly squared anteriorly and posteriorly; rather variable number of long submarginal setae present, minimally 1 anterior and 2 posteriorpairs; lingula excluded from vasiform orifice; on bamboo ALEUROTULUS (arundinacea Singh) (p. 321) - Pupal case elongate-oval; submargin with even row of 16 pairs of short, broad, lanceolate setae (Fig. 39); lingula exposed but included within vasiform orifice; on blades of Gramineae,usually in dense, ant- attended colonies ALEUROCYBOTUS (setiferus Quaintance & Baker) (p. 317) 13 Dorsal disc separated from rather wide submarginal area by a distinct line or fold concentric with margin 14 - Dorsal disc not so separated from submarginal area 15 14 Vasiform orifice triangular, longer than wide, lingula included and often obscured by opercu- lum, caudal furrow pronounced (Fig. 46); differentiation of margin at thoracic and caudaltracheal openings rather variable between species, often most apparent caudally. Pupalcases black, often with a white waxy fringe in life ALEUROLOBUS (p. 317) - Vasiform orifice small and subcircular, hardly longer than wide; thoracic and caudal tracheal openings at margin marked by small indentations ('pores'). Pupal cases pale to dusky : ASIALEYRODES(p. 321) 15 Almost always with 5 pairs of large, simple, subdorsal pores (Fig. 40); if pores are absent then outer submargin with a row of about 12 pairs of short, somewhat lanceolate setae. Margin irregular. Pupal cases in life often surrounded by small patches of secretediridescent blue waxy filaments DIALEUROPORA(p. 326) - Without 5 pairs of large, simple pores in subdorsum; if submarginal setae present, then not lanceolate 16 16 Inner submargin with a row of stout papillae. Dorsal disc often with a few papillae. Knob of lingula about as broad as long, lobulate (Fig. 29) TRIALEVRODES(p.333) Inner submargin without a row of such papillae 17 17 Thoracic and/or caudal tracheal openings at margin either in the form of notches ('pores'), which may themselves be deeply invaginated from the main outline of the margin, or in theform of thickened, smoother breaks in an otherwise toothed margin 18 - Thoracic and/or caudal tracheal openings in the form of differentiated marginal teeth, sometimes only 2 or 3 such modified teeth present, and sometimes several, forming a 'comb';differentiation often best developed caudally 23 18 Submedian area of dorsal disc somewhat elevated above remainder of pupal case, forming a rhachis with a pronounced furrow or crease running into subdorsum from submedian part ofeach abdominal segment (as Fig. 38) RHACHISPHORA(p. 333) - Rhachis not developed and submedian area not elevated above remainder of pupal case, but segmentation normally marked medially 19 19 Tracheal pores invaginated from margin, inset from margin by several times diameter of pore; outer submargin without a ring of hairs which extend beyond margin; vasiform orificetriangular, lingula exposed but included (Fig. 20) INDOALEYRODES(p. 327) - If tracheal pores invaginated from margin, then either outer submargin with a row of hairs which extend beyond margin, or vasiform orifice not both triangular and with lingulaexposed 20 THE WHITEFLY OF NEW GUINEA 311 20 Vasiform orifice about as wide as long, with operculum occupying most of area of orifice; lingula concealed by operculum 21 - Vasiform orifice normally longer than wide ; operculum covering only basal half to two-thirds of orifice; lingula exposed but included.Outer submargin with a row of hairs which extend beyond the margin 22 21 Vasiform orifice relatively small with respect to size of pupal case: inset from posterior margin by 3 or more times its own length. Caudal furrow often rather ornate (Fig. 18) DIALEURODES(p. 325) - Vasiform orifice relatively large with respect to size of pupal case: maximally inset from posterior margin by about twice its own length (Fig. 42). Caudal furrow well-marked butoften rather plain ALEUROTUBERCULATUS(p.32Q) 22 Vasiform orifice with a subcircular to trapezoidal anterior section which contains both the operculum and the lingula which has a rather wide D-shaped head: orifice continuedposteriorly as a cordate to triangular extension which is sculptured on the floor of thedepression (Fig. 28) PEALIUS-group(p. 332) - Vasiform orifice normally triangular and without a false posterior edge; lingula exposed, included, its head with a pair of lateral basal tubercles, not so short and D-shaped (Figs 25, 27) PARABEMISIA(p. 330) 23 Pupal case elongate-oval, 2-0-2-5 times longer than wide, asymmetric in outline (Fig. 8) ALEUROMARGINATUS (littoralis sp. n.) (p. 319) - Outline of pupal case symmetrical and round to oval, not elongate 24 24 Without a row of submarginal hairs; caudal furrow (dorsal) usually little-marked. Vasiform orifice cordate to rounded-triangular; thoracic and caudal folds (ventral) oftenclearly indicated as finely stippled bands running from margin into subdorsal area. Pupalcases black or pale ALEUROPLATUS(p. 320) - Normally with a row of submarginal hairs which extend beyond the margin, and with caudal furrow (dorsal) well-developed 22 ALEURODICINAEALEURODICUS Douglas Aleurodicus Douglas, 1892: 32. Type-species: Aleurodicus anonae Morgan, by subsequent designation,Quaintance & Baker, 1908: 8. Key to tropical Asian species of AleurodicusPupal cases 1 With 4 pairs of abdominal compound pores - With 5 or 6 pairs of abdominal compound pores 3 2 Margin with regular, coarse, teeth - about 60 on each side of case; central spines of compound pores stout and seta-like, reaching margin of case; case without long submarginal setae. (SriLanka) antidesmae Corbett - Margin not regularly and coarsely toothed; central spines of compound pores short and spatulate, dagger-shaped; case with 11 pairs of long, fine submarginal setae, in addition tocaudal and posterior marginal pairs.Dorsum with a dense pattern of small wax pores . * dispersus Russell 3 With 6 pairs of abdominal compound pores With 5 pairs of abdominal compound pores 5 4 All abdominal compound pores subequal in size, usually with central processes not evident, resembling rings with transversely-sculptured rims; submargin with about 12 pairs of long,fine setae (Fig. 47). (Australia, Brunei, Papua New Guinea, Philippines, Sarawak, SolomonIslands, Sulawesi, West Malaysia) destructor Mackie (p. 312) - Posterior two pairs of abdominal compound pores larger than remaining four pairs, each of posterior two pairs with a conspicuous central spine; submargin without long, fine setae. (Fiji,Java, Papua New Guinea, [Sarawak], Sri Lanka, Thailand, West Malaysia) Ao/mesi/(Maskell)(p.312) - Russell (1965) recorded A. dispersus from the Caribbean, Central and South America, southern U.S.A. andthe Canary Islands. It has since been introduced to the Philippines (Martin & Lucas, 1984), Guam, MarianasIslands and Hawaii (Russell, pers. comm.). 312 J. H. MARTIN 5 Cephalic and posterior four pairs of abdominal compound pores subequal in size, with centralspines short, not reaching margin of case; only anteriormost pair of abdominal pores smallerthan remainder. (Hong Kong, Taiwan) machili Takahashi - Cephalic and posterior two pairs of abdominal compound pores subequal in size, with centralspines usually reaching margin of case; anterior three pairs of abdominal pores subequal insize, distinctly smaller than remainder. (West Malaysia) cinnumomi Takahashi Aleurodicus destructor Mackie (Fig. 47)Aleurodicus destructor Mackie, 1912: 142. Syntype pupal cases, PHILIPPINES (USNM). DISTRIBUTION. Australia (New South Wales), Brunei, Papua New Guinea, Philippines, Sara-wak, Solomon Islands, Sulawesi, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Port Moresby, on Cocos nucifera (Palmae) and Ficus microcarpa var. naumannii(Moraceae); Wewak, on Cocos nucifera; Wau, on Cinnamomum sp. (Lauraceae); Buso, on IXanthophyl-lum sp. (Xanthophyllaceae); New Britain Province, on Cocos nucifera (all BMNH). Aleurodicus holmesii (Maskell) Aleurodes holmesii Maskell, 1895: 435, fig. xxxi-2. Syntype pupal cases, FIJI (DSIR) [examined].Aleurodicus holmesii (Maskell) Cockerell, 1903: 664. Aleurodicus malayensis Takahashi, 1951: 2, fig. 2. Syntype pupal cases, WEST MALAYSIA (BMNH)[examined]. Syn. n. Takahashi separated malayensis from other Asian species of Aleurodicus because it had pupalcases which were 'narrow and much narrowed cephalad', with the cephalic compound poresappearing very close to the margin. The syntypes of malayensis are in a very thick balsam mount,and there is considerable downward curling of the margins of the specimens, particularly in thecephalothoracic region. There is nothing else unusual about the outline of the pupal case, or inthe position of the cephalic compound pores with respect to the margin of the case. The onlyother difference between malayensis and holmesii was given as the relatively long spines issuingfrom the cephalic and last two abdominal pairs of compound pores in malayensis. The syntypesof holmesii have been compared with the syntypes of malayensis, as well as with material fromPapua New Guinea, Java, Thailand and Sri Lanka. In the syntypes of holmesii, all the compoundpore central spines are broken, and additional slides were therefore prepared from duplicate drymaterial from the Maskell collection. From specimens with unbroken spines, it is evident thatthe compound pore spines of holmesii from Maskell's original Fiji material are indeed veryshort, and do not extend beyond the pupal case margin, in contrast to the syntypes of malayensiswhich have rather longer spines. However, there is variation in the relative lengths of thesespines with respect to the pore-to-margin distance in other material present in the BMNHcollection. A. malayensis is therefore considered a junior synonym of holmesii. DISTRIBUTION. Fiji, Java, Papua New Guinea, (Sarawak), Sri Lanka, Thailand, West Malaysia. MATERIAL EXAMINED Fgi: Syntype pupal cases (of holmesii) and duplicate material bearing syntype data, on Psidium sp.(Myrtaceae) (BMNH; DSIR). West Malaysia: 8 pupal cases (syntypes of malayensis) , Kuala Lumpur, onundetermined host (BMNH). Java: 5 pupal cases (BMNH). Papua New Guinea: 3 pupal cases, 2 adult $ , Buso, on undetermined tree;7 pupal cases, Buso, on Guioa sp. (Sapindaceae) (BMNH). Sarawak: 1 pupal case, Gunung Mulu NationalPark, on Annonaceae, tentatively identified as holmesii (BMNH). Sri Lanka: 1 pupal case, Kandy (G. H.Corbett det.) (BMNH). Thailand: 4 pupal cases, Pah Meeung Mts (BMNH). THE WHITEFLY OF NEW GUINEA 313 ALEYRODINAEALEUROCANTHUS Quaintance & Baker Aleurocanthus Quaintance & Baker, 1914: 102. Type-species: Aleurodes spinifera Quaintance, by originaldesignation. More species of whitefly seen from New Guinea belong to Aleurocanthus than to any othergenus: five have been identified as previously described species, two are here described as new,and a further seven remain undetermined (p. 305). Of those which are undetermined, species 1, 2, 3 and 5 resemble regis Mound in possessingfew, if any, stout spines additional to the submarginal ring (see comments concerningpapuanus). Aleurocanthus sp. 6 is discussed with esakii which it closely resembles. Key to named New Guinea species of AleurocanthusPupal cases 1 Pupal cases pale to slightly dusky; ventral submargin punctuated by band of shallow subcircular tubercles (Fig. 1) 2 - Pupal cases brown to black ; without ventral submarginal band of shallow subcircular tubercles ... 3 2 Stout dorsal spines varying markedly in length, longest 0-25-0.30 mm long; submarginal stout spines tend to occur in groups of 3 or 4 in thoracic and anterior abdominal regions (Fig. 1) /uteussp. n.(p. 314) - Stout dorsal spines varying less extremely in length, maximally about 0-2 mm long; submarginal stout spines not occurring in groups of 3 or 4 (Fig. 41) cocois Corbett (p. 313) 3 Stout dorsal spines in a single ring of 12 submarginal pairs with the only additional pair situated on edge of rhachis on abdominal segment V (Fig. 3). Apices of spines expanded, laciniate papuanus sp. n. (p. 315) - Several pairs of stout dorsal spines present in subdorsal and submedian areas in addition to submarginal ring. Spines with acute apices 4 4 Marginal teeth very large, only 4-5 occupying 0-1 mm of margin, teeth at least as long as wide at base, blunt. Distribution of stout dorsal spines as in Fig. 44 woglumi Ashby (p. 316) - Marginal teeth smaller, more than 7 teeth occupying 0-1 mm of margin 5 5 Submargin with 11 or 12 pairs of stout spines evenly spaced around case, but with posterior 3 pairs double , the submarginal ring thus totalling about 30 spines (as in Fig .45) 6 - Submargin normally with 11 pairs of stout spines, the posterior 3 pairs not being double, submarginal ring thus totalling about 22 spines only. Arrangement of abdominal stout spines: submedian pairs on segments I-III & VIII, innersubdorsal pairs on segments II-VII spiniferus (Quaintance) (p. 316) 6 Marginal teeth acute-triangular, often somewhat uneven ; cephalothorax with 3 submedian pairs of short stout spines (Fig. 45) pendleburyi Corbett (p. 316) - Marginal teeth blunt, rather castellate and somewhat spiky; cephalothorax with only 1 pair of submedian short stout spines esalci/Takahashi(p. 314) Aleurocanthus cocois Corbett(Fig. 41) Aleurocanthus cocois Corbett, 1927: 24. Syntype pupal case, WEST MALAYSIA (BMNH) [examined].Aleurocanthus canangae Corbett, 1935a: 790. Syntype pupal cases, WEST MALAYSIA (presumed lost).Syn. n. As stated by Corbett (1935a), canangae and cocois are only separable by the different lengths ofthe dorsal spines. Examination of the material listed below, including a syntype of cocois andseven specimens identified as canangae by Corbett, indicates that the spines vary somewhat inlength, but that the pattern of spines does not. Thus canangae is here synonymised with cocois. DISTRIBUTION. Burma, Cambodia, India, Papua New Guinea, Solomon Islands, Thailand, WestMalaysia. 314 J. H. MARTIN MATERIAL EXAMINED West Malaysia: 1 syntype pupal case, Batu Gajah, on Cocos nucifera (Palmae) (BMNH); 7 pupal cases(identified as canangae by Corbett), Kuala Lumpur, on Kananga [sic] odorata (Annonaceae) (BMNH). Papua New Guinea: numerous pupal cases and larvae, Buso, on undetermined Sterculiaceae; 14 pupalcases, Wasu, on undetermined tree (all BMNH). Solomon Islands: numerous pupal cases, Honiara, on Cocos nucifera; 1 pupal cases, Russell Island; 13pupal cases, Guadalcanal, on Cocos (all BMNH). Aleurocanthus esakii TakahashiAleurocanthus esakii Takahashi, 1936: 111, fig. 1. Syntype pupal cases, PALAU ISLANDS (TARI). This species is very similar topendleburyi, from which it differs in possessing blunt to castellatemarginal teeth, and in having two pairs fewer of short cephalothoracic dorsal spines. Several pupal cases of Aleurocanthus sp. 7 (see p. 305), very similar to esakii, were collectedon Cinnamomum sp. at Wau Ecology Institute, Morobe uplands. These possess marginal teethidentical to those of esakii, but have one extra pair of short cephalothoracic dorsal spines; theyalso differ in the number and form of the fine submarginal capitate setae. These specimens thushave one pair of dorsal spines fewer thanpendleburyi, and differ further from that species in notpossessing acute triangular marginal teeth. DISTRIBUTION. Palau Islands (Caroline group), Papua New Guinea. MATERIAL EXAMINED Papua New Guinea: Buso, on single leaf of tree-crown foliage of Pometia pinnata (Sapindaceae)(BMNH). Aleurocanthus luteussp. n. (Figs 1,2) PUPAL CASE. Elongate-oval, 0-80-0-90 mm long, 0-47-0-55 mm wide, widest at abdominal segment III.Cuticle pale to yellowish dusky. Margin very slightly flattened at thoracic tracheal openings, but not atanterior or posterior edges of case. Marginal teeth well developed, even, about 9 rounded-castellate teethoccupying 0-1 mm of margin (Fig. 2). Fine anterior and posterior marginal setae present. Dorswn. Dorsal cuticle smooth, not punctuated by pores or sculpturing. Eye spots not marked. Dorsaldisc and submargin with about 40 pairs of stout spines, of varying lengths, as shown in Fig. 1; spinessmooth, pointed, with the longest (usually 1 cephalothoracic pair, subdorsal pair on abdominal segment IIIand submedian pair on abdominal segment V) up to 0-35 mm long. Each dorsal spine with a small circularpore near base. About half-way between margin and outermost stout spines is a row of about 8 pairs ofsubmarginal setae about 35 urn long, arranged as in Fig. 1. Pair of cephalic setae not evident in specimensseen, but eighth abdominal and caudal setae long: eighth abdominal setae rather stout, up to 0- 12 mm long;caudal setae very fine, up to 0-25 mm long. Base of each marginal tooth marked on dorsum by atubercle-like marking on cuticle. Dorsal disc rhachisform, with cephalic and posterior abdominal medianline raised into a pronounced keel, with the submedian spines on either side of keel. Transverse moultingsutures becoming indistinct in submargin, not apparently reaching margin of case; longitudinal suturereaching anterior edge of case. Meso-/metathoracic suture well marked, but remainder of cephalothoracicsegmentation not evident. Vasiform orifice elevated, with vertical lateral 'flanges' which fold down onslide-mounted specimens to partially obscure vasiform orifice characters (Fig. 2). Vasiform orifice openingabout 0-14 mm long, 0-13 mm wide; operculum occupying whole of orifice and obscuring setose head oflingula; vasiform orifice almost always distorted in slide-mounted specimens. Caudal furrow not evident. Venter. Submargin punctuated by shallow, subcircular tubercles which run mesad from margin in regularlines (Fig. 1), about 4-5 tubercles in each line, and about 2 marginal teeth to each line of tubercles.Thoracic and caudal tracheal folds only marked as breaks in submarginal band of tubercles. Mesad ofsubmarginal band of tubercles are many rather sparsely distributed dots in outer subdorsum and centralpart of thoracic area. A minute conical spine present at base of each middle and hind leg. Ventralabdominal setae long, fine. Holotype pupal case, Papua New Guinea: Morobe Province coast, Lasanga Island, on Macaranga sp.(Euphorbiaceae), 21.X.1979 (/. H. Martin 2746) (BMNH). Paratypes. 9 pupal cases, same data as holotype (BMNH). THE WHITEFLY OF NEW GUINEA 315 COMMENTS. A. luteus belongs to a group of species which are characterised by their pale pupalcase cuticle and the possession of a ventral submarginal band of shallow tubercles. Species in thisgroup include strychnosicola Cohic, trispina Mound, zizyphi Priesner & Hosny and mackenzieiCohic from Africa; rugosa Singh from India and West Malaysia; and cocois Corbett from theIndian Region, Pacific Region and New Guinea. With its large number of long dorsal spines,luteus most closely resembles trispina and mackenziei, but differs from both species in thedistribution and the markedly varying length and thickness of its dorsal spines; luteus furtherdiffers from trispina in having smooth dorsal spines which do not possess tiny lateral spinules. A.luteus resembles cocois in possessing about 8 pairs of submarginal setae, but the dorsal discspines are much stouter and longer, and the tendency for submarginal stout spines to occur ingroups of 3 or 4 does not occur in cocois. Aleurocanthus papuanus sp. n. (Figs 3-5) PUPAL CASE. Dark brown to black, usually requiring only a little bleaching for microscopical examination.Sexually dimorphic, $ 1-30-1 -45 mm long, 1-30-1-40 times as long as wide, cf 1-00-1-07 mm long,1-40-1-50 times as long as wide; maximum width at abdominal segments II and III. Outline rathercharacteristic, flattened to shallowly concave posteriorly, slightly indented towards thoracic tracheal areasand faintly pointed anteriorly. Marginal teeth well developed, even, rounded-triangular and distinctlypaler than remainder of case, about 6-8 teeth occupying 0-1 mm of margin; teeth modified at thoracic andcaudal tracheal openings (Fig. 4), where the line marking the tooth bases is more markedly indented thanthe margin itself. Fine anterior and posterior marginal setae present. Dorsum. Cuticle of submedian area fairly smooth, evenly pigmented, but subdorsum and innersubmargin punctuated by rather regular darker blotches (Fig. 4). Tiny porettes scattered over dorsum. Eyespots not visible. Submargin with a ring of 12 pairs of stout spines which are paler than dorsal cuticle, up to0-33 mm long in 9 (proportionately shorter in cT), with expanded laciniate apices (Fig. 4); fifthcephalothoracic pair further inset from margin than remainder, and one further pair of similar spinespresent in inner subdorsum on abdominal segment V (Fig. 3). First abdominal setae short and ratherspatulate; cephalic and eighth abdominal setae fine and dark, with caudal hairs similar but up to twice aslong. A line of tiny submarginal pores present between marginal tooth bases and line of spinal bases (Fig.4). Transverse moulting sutures only reach outer subdorsum, but longitudinal suture reaches anteriormargin. Abdominal segmentation very well marked by thin, pale, suture-like folds which are immediatelybordered by slightly darker pigmentation than on remainder of dorsum; these curious folds are continuedinto outer subdorsum almost perfectly out of phase with the segmentation on the submedian area, forminga rhachis (Fig. 3). Meso-/metathoracic suture well defined, very close to median part of transverse moultingsuture, but remaining cephalothoracic segmentation not discernible. Submedian area defined in cepha-lothorax by suture-like folds similar to those on abdomen, with one lateral branch extending as far ascephalothoracic submargin between spine pairs 3 and 4. Vasiform orifice (Fig. 5) subcircular, outer (upper)internal margin toothed, inner part slightly reticulate as shown. Operculum trapezoidal, unpigmented,almost filling internal area of orifice. Lingula included and covered by operculum, though somewhat visiblethrough operculum. Caudal furrow marked by slight darkening of cuticle (Fig. 4). Venter. Thoracic and caudal tracheal folds each marked only by a pair of fine lines extending mesad toouter subdorsal area, not spinulose or sculptured. Ventral abdominal setae well developed, fine, situatedanterior to vasiform orifice. Holotype pupal case ($), Papua New Guinea: Morobe Province coast, Buso river bank, on Xanthophyl-lum papuanum (Xanthophyllaceae), 13.ix.1979 (/. H. Martin 2557) (BMNH). Paratype pupal cases. Papua New Guinea: 35 9, 22 d", same data as holotype; 1 $, Lasanga Island, onGuioa sp. (Sapindaceae), 19.ix.1979 (/. H. Martin 2590) (BMNH; USNM). COMMENTS. A. papuanus resembles regis Mound (1965, Nigeria) in possessing spines which arevirtually all in one submarginal ring, and in the form of these spines, but differs in many othercharacters. In its general outline, shape of the transverse moulting sutures, and tendencytowards a dorsal rhachis, papuanus appears similar to hirsutus (Maskell) and T-signatus(Maskell) from Australia (Dumbleton, 1956), and to brevispinosus Dumbleton and spinithoraxDumbleton from New Caledonia (Dumbleton, 1961a). The combination of 12 pairs of submar-ginal spines with expanded apices, only one similar submedian pair, and the marked cepha- 316 J. H. MARTIN lothoracic and abdominal folds separate papuanus from other described species ofAleurocan-thus. It is worth noting that Aleurocanthus species 1,2,3 and 5 (p. 305) also display a lack of dorsaldisc spines, but seem to have closer affinities with regis than with papuanus and the Australianand Pacific species. The colony on Xanthophyllum papuanum was very numerous, but a single specimen wascollected from a species of Guioa. Pupae were not attended by ants. Aleurocanthus pendleburyi Corbett (Fig. 45)Aleurocanthus pendleburyi Corbett, 1935a: 795. Syntype pupal cases, WEST MALAYSIA (presumed lost). This species differs from esakii in possessing pointed marginal teeth and two additional pairs ofshort cephalothoracic dorsal spines. DISTRIBUTION. Papua New Guinea, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Aiyura, on Piper sp. (Piperaceae); Goroka, on Persea americana (Lauraceae)(BMNH); Nondugl, on undetermined host (BMNH; BPBM). West Malaysia: Genting Highlands, onundetermined tree (BMNH). Aleurocanthus spiniferus (Quaintance) Aleurodes spinifera Quaintance, 1903: 63; Quaintance & Baker, 1917: pi. 38, figs 1-6. Syntype pupal cases, JAVA (USNM; TARI).Aleurocanthus spiniferus (Quaintance) Quaintance & Baker, 1914: 102. This species is widespread and has hosts recorded from 15 plant families. The material listedbelow extends its known range into Java, New Guinea and Australia. DISTRIBUTION. Papua New Guinea, Java, Australia (Queensland), also widely distributed in theOld World tropics; the existence of spiniferus in the Neotropical Region is doubtful (Mound &Halsey, 1978). MATERIAL EXAMINED Papua New Guinea: Port Moresby, on Plumeria rubra c.v. (Apocynaceae) and Hibiscus sp. (Malvaceae)(BMNH). Java: Jakarta, on Citrus llimon (Rutaceae) (BMNH). Australia: Queensland, Cairns, oncustard apple (Annonaceae) (BMNH; DPIQ). Aleurocanthus woglumi Ashby(Fig. 44) Aleurocanthus woglumi Ashby, 1915: 321. Syntype pupal cases, JAMAICA (7USNM).Aleurocanthus husaini Corbett, 1939: 69. Syntype pupal cases, INDIA (depository unknown). Syn. n. Corbett (1939) distinguished husaini from woglumi on the evidence of certain longer dorsal discspines in the pupal cases of husaini, combined with an apparent difference between theforewings of adults of the two species. The type-material of husaini has not been traced, but inview of the variation in spine lengths displayed within some species of Aleurocanthus (seediscussion of cocois), it is considered that this character has little significance. Further, theillustrated forewings appear to be from a male ('husaini') and a female ('woglumi'), whichprobably accounts for the difference in shape. The pupal case of husaini as described andillustrated by Corbett is considered a variation of woglumi. Mound & Halsey (1978) did not record woglumi from New Guinea. The samples (see below)from Irian Jaya and Papua New Guinea (Wewak) were collected in 1959 and 1968 respectively,and were among undetermined material in BPBM, Honolulu. THE WHITEFLY OF NEW GUINEA 317 DISTRIBUTION. Borneo, Irian Jay a, Java, Papua New Guinea, Philippines, Singapore, Sumatra,West Malaysia in Austro-Oriental Region; Hawaii; also widely distributed elsewhere in warmerparts of the world. MATERIAL EXAMINED Irian Jaya: Cyclops Mts, on Citrus sp. (Rutaceae) (BMNH; BPBM). Papua New Guinea: Wewak, onCitrus sp. (BMNH; BPBM); Zifaseng, on Citrus sp. (BMNH); North Solomon Islands Province, 'nearArawa', on Citrus sp. (BMNH). ALEUROCYBOTUS Quaintance & Baker Aleurocybotus Quaintance & Baker, 1914: 101. Type-species: Aleurodes graminicola Quaintance, bymonotypy. Aleurocybotus setiferus Quaintance & Baker (Fig. 39)Aleurocybotus setiferus Quaintance & Baker, 1917: 357. Syntype pupal cases, JAVA, SRI LANKA (USNM). Two colonies of this species were seen at Buso, feeding on blades of the grass Imperata cylindricagrowing in sand among other strand-line vegetation. Both colonies were vigorously attended byants: Iridomyrmex sp. in one, and Polyrhachis laciniata in the other. DISTRIBUTION. Australia (Queensland), Hong Kong, Java, Papua New Guinea, Philippines, SriLanka, Taiwan, Thailand, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Buso, on Imperata cylindrica (Gramineae). Hong Kong: New Territories, on 'grass'.Sri Lanka: Peradeniya, on Imperata arundinacea. Australia: Queensland, on Imperata cylindrica (allBMNH). ALEUROLOBUS Quaintance & Baker Aleurolobus Quaintance & Baker, 1914: 108. Type-species: Aleurodes marlatti Quaintance, by originaldesignation. Aleurolobus niloticus Priesner & HosnyAleurolobus niloticus Priesner & Hosny, 1934: 1, (pi. 3). Syntype pupal cases, EGYPT (USNM). The material from New Guinea and Sarawak, detailed below, extends the distribution given byMound & Halsey (1978) into the Austro-Oriental Region. DISTRIBUTION. The Middle East, East and North Africa, India, Pakistan, Sarawak, Papua NewGuinea. MATERIAL EXAMINED Papua New Guinea: Buso, on Gmelina sp. (Verbenaceae). Sarawak: Gunung Mulu National Park, onundetermined host (all BMNH). Aleurolobus selangorensis Corbett (Fig. 46)Aleurolobus selangorensis Corbett, 1935a: 819. Syntype pupal cases, WEST MALAYSIA (presumed lost). A. selangorensis is very similar to niloticus but can be distinguished by the presence ofconspicuous dots lining the folds which run mesad from the marginal teeth into the submargin. DISTRIBUTION. Papua New Guinea, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Buso, on Erythroxylum sp. (Erythroxylaceae), Dalbergia sp. (Leguminosae),Rhizophora cf. stylosa (Rhizophoraceae); Lasanga Island, on Cerbera Imanghas (Apocynaceae); KratkeMts, on Neonauclea sp. (Naucleaceae) (all BMNH). 318 J. H. MARTIN ALEUROMARGINA TVS CorbettAleuromarginatus Corbett, 19356: 246. Type-species: Aleuromarginatus tephrosiae Corbett, by monotypy. Aleuromarginatus corbettiaformis sp. n. (Figs 6, 7) PUPAL CASE. Outline asymmetrically elongate-oval, often rather narrowed cephalad and usually widest atabdominal segments III-IV. Margin sometimes flattened at anterior and posterior ends of case, but rarelyconcave. Sexually dimorphic: $ 1-40-1-55 mm long, 0-65-0-70 mm wide; cf 1-10-1 -20 mm long, 0-47-0-50mm wide; on average 2-3 times as long as wide. Margin (Fig. 7) with regular double row of teeth; primaryteeth rather coarse and conspicuous, triangular, with only 7-9 occupying 0-1 mm of abdominal margin;secondary row in form of shallow crenulations. Margin not modified at thoracic and caudal tracheal areas.A pair of fine marginal setae present posteriorly but not anteriorly. Dorsum. Cuticle pale, but many specimens with a brown median stripe. Dorsum densely patterned;submargin with an elongate-oval fold at base of each marginal tooth, and immediately mesad is a row ofsubcircular papillae; lateral margins of submedian area of dorsal disc marked by longitudinal lines ofsimilar papillae (Fig. 6) , these lines similar in appearance to those in species of Corbettia Dozier ; remainderof dorsal disc densely punctuated by subcircular to polygonal markings which are less sharply defined thanthe papillae (Fig. 7). Dorsum bearing 21 pairs of blunt hairs which are up to 35 /Lim long and usually muchcurved; distribution as shown in Fig. 6, caudal pair on abdominal segment VIII situated submarginally.Longitudinal and transverse moulting sutures both reach margin. Pro-/meso- and meso-/metathoracicsutures well defined, unlike cephalic/prothoracic junction. Vasiform orifice subcordate, a little longer thanwide, inset from posterior margin by about 3 times its own length in $ (Fig. 7) and 2-3-2-6 times in cf ;lateral margins almost straight, anterior and posterior margins rounded; inner edges of lateral margins witha few coarse teeth. Operculum and lingula as shown (Fig. 7), similar to those of A. littoralis (following).Dorsal disc with scattered double pores, submargin with line of similar pores just mesad of marginal teeth.An apparent caudal furrow is created by differentiation of some of the dorsal markings. Venter. Thoracic tracheal folds only occasionally indicated, and then only by a pair of faint lines runningmesad from margin. Caudal tracheal fold indicated by faint longitudinal folds and scattered spinules.Ventral abdominal setae fine, long, more than half length of vasiform orifice. Anterior abdominal spiracleshook-shaped. A minute conical spine at base of each middle and hind leg. Antennae rather thick, distallyroughened and apically pointed, apex reaching to halfway between bases of fore and middle legs. Holotype pupal case $ , Papua New Guinea: Morobe Province coast, Buso, on Desmodium umbellatum(Leguminosae), 10.x. 1979 (/. H. Martin 2680) (BMNH). Paratype pupal cases. 13 $, 9 cf , same data as holotype (BMNH; USNM); 10 dry specimens on leaf,same data as holotype (BMNH). COMMENTS. The species is known from only one colony, a large, scattered population on thelower surfaces of mature leaves of Desmodium umbellatum (Leguminosae) . The pupae were notattended by ants. Each individual has a marginal fringe of waxy filaments, apparently each onecorresponding to a marginal tooth: the filaments are not dense and not very obvious to the nakedeye, although they are about as long as the subdorsum is wide. The dorsal surface is covered bydiscrete grains of whitish or almost colourless waxy secretion, each grain corresponding to acuticular marking: this secretion is developed into longitudinal ridges at the edges of thesubmedian area, corresponding to the lines of tubercles, and the secretion is similarly developedabove the line of submarginal tubercles. Each of the dorsal hairs also has an accretion of waxattached to it. This species has a marked preference for feeding sites alongside major leaf veins,which is the usual explanation of asymmetry of outline. The host plant, Desmodium umbella-tum, is widespread in strand-line vegetation from East Africa and Madagascar through tropicalAsia to Taiwan and northern Australia (Verdcourt, 1979). The density and form of cuticular patterning in corbettiaformis is quite unlike that of any of thedescribed species. Despite the differentiated longitudinal line of tubercles at the edge of thedorsal disc, the species displays all the characters of Aleuromarginatus, in particular theapparently double row of marginal teeth and 21 pairs of tiny dorsal hairs in a characteristicpattern. In the characteristics of the vasiform orifice, dorsal segmentation and caudal trachealfold, corbettiaformis resembles littoralis (see below), but the dorsal patterning, margin of caseand appearance in life are quite different. THE WHITEFLY OF NEW GUINEA 319 Two ventrally incomplete pupal cases of Aleuromarginatus (sp. 1, p. 305) were collected ona vine, possibly Dalbergia densa or D. candenatatus . These also have lines of papillae differenti-ated from the remainder of the dorsal markings, but in a more complex pattern involvingcomplete delineation of the submedian area and 8 pairs of radial 'spokes' running into thesubmargin; the specimens are symmetrical and not narrowed cephalad. Aleuromarginatus littoralis sp. n. (Figs 8, 9) PUPAL CASE. Shape generally elongate-oval, widest at abdominal segment III, with margin slightly indentedanteriorly and posteriorly. Most individuals are asymmetrical, with many being more convex on one sidethan on the other, and with precise outline being rather variable. Sexually dimorphic: $ 1-80-2-00 mmlong, 0-75-0-90 mm wide; C? 1-30-1-45 mm long, 0-55-0-65 mm wide. Pupal cases of both sexes 2-0-2-5times as long as wide, on average 2-3 times. Margin (Fig. 8, inset) with regular double row of teeth: primaryteeth conspicuous, with 12-13 occupying 0-1 mm of abdominal margin; secondary row in form of shallowcrenulations; short parallel lines run mesad from bases of teeth into submargin. Margin shallowly indentedin thoracic tracheal region where teeth are smaller and closer-set; posterior marginal teeth at apex ofcaudal fold similarly differentiated from remainder of marginal teeth. A pair of fine posterior marginalsetae present, but anterior pair apparently always absent. Dorsum. Cuticle pale, but with all segmental sutures sharply defined and often emphasised by brownishpigment. Abdominal intersegmental sclerotisation expands at lateral ends of segmental sutures and tendsto merge with that of adjacent segments; the abdominal submedian area is thus quite well defined, althoughnot delineated by a suture-like fold or rhachis. Paired submedian depressions present in anterior half ofeach of abdominal segments I- VI, and on thorax. Dorsum sculptured by a dense but irregular pattern ofpolygonal to subcircular markings which are darker and better defined nearer submargin, and somewhatvariable between specimens (Fig. 8, inset). Dorsum bearing 21 pairs of very short, often much-curvedhairs; distribution as in Fig. 8; abdomen with the pairs on segments I and III- VI just inside submedian area,pair on segment II in subdorsum, the remaining 4 pairs on segment VIII, including the subdorsally placedcaudal pair which are similar to the remainder. Transverse and longitudinal moulting sutures reach margin.Pro-/meso- and meso-/metathoracic sutures well defined; cephalic/prothoracic junction no more than aslight fold which is angled abruptly anteriad in its distal half; lengths of thoracic segments subequal. Medianlength of abdominal segment VII about half that of each of segments I-VI. Median line on abdominalsegments I-IV or V somewhat sclerotic, although not normally pigmented. Vasiform orifice (Fig. 9)elongate-cordate, 65-90 pm long, 55-75 /mi wide, inset from posterior margin by about 3 times its ownlength; lateral margins straight to slightly concave, anterior and posterior margins rounded; inner edges oflateral margins toothed. Operculum trapezoidal with rounded lateral margins, occupying about half lengthof vasiform orifice; lingula with large spinulose head, occupying most of remaining volume of orifice,exposed but included. Dorsal disc with scattered double disc pores; submargin with a line of similar poresjust mesad of marginal teeth, one double pore at base of each 4th to 5th tooth. Venter. Caudal tracheal fold rather broad, marked by slight longitudinal folding, and further punctuatedby groups of small spinules along two-thirds of its length from margin to vasiform orifice (Fig. 8). Thoracictracheal folds marked in subdorsum by scattered, very fine stipples, margins of folds often marked by faintlines. Ventral abdominal setae fine, about half as long as vasiform orifice, with bases situated at about halflength of, and a little lateral to, orifice. Anterior abdominal spiracles appear hook-like, situated just lateralto submedian area of abdomen which is very faintly stippled. A minute conical spine present at base of eachmiddle and hind leg. Antennae rather long, slender, directed posteriorly, distally roughened, apicallypointed, apex reaching half way between bases of fore and middle legs. Holotype pupal case 9, Papua New Guinea: Morobe Province coast, Lasanga Island, on 1 Denistrifoliata (Leguminosae), 7.xi.l979 (J. H. Martin 2819) (BMNH). Paratype pupal cases. Papua New Guinea: 31 $,22 cf , same data as holotype; 23 9, 21 cf, MorobeProvince coast, Buso, on same host, 10.ix.1979 (JHM 2529); 13 $, 6 d", Buso, on same host, 25.x. 1979(JHM 2760) (BMNH; USNM). A number of dry specimens on leaf, with same data as holotype (BMNH). COMMENTS. This species is known from three large, but not dense, colonies. Pupae were notattended by ants. Each pupa is devoid of obvious secretion dorsally, but is surroundedmarginally by a broad fringe of translucent waxy strands, apparently one to each marginal tooth.The marginal wax strands corresponding to the thoracic and caudal tracheal areas are denser andmore opaque, white. Pupae are scattered, apparently randomly, over the lower surfaces of thesmooth mature leaves of the host, with nothing in their positioning to indicate a possible 320 J. H. MARTIN explanation for their asymmetry. The host, tentatively identified as Denis trifoliata, is ashoreline woody climber which in the Buso area was found in sandy, beach-top situations. D.trifoliata is widespread in strand-line vegetation in East Africa, Madagascar, tropical andsubtropical Asia and Australia (Verdcourt, 1979). A. littoralis is similar to kallarensis David & Subramaniam (1976) and dalbergiae Cohic (1969)in its marked asymmetry and general appearance. It is also similar to millettiae Cohic (1968),differing in the following respects: meso- and metathoracic segments subequal in medianlengths; margin flattened to concave anteriorly, posteriorly and at thoracic tracheal areas; mostindividuals conspicuously asymmetrical. It differs from kallarensis and dalbergiae as follows:vasiform orifice longer than wide, with lateral margins straight to slightly concave; submedianarea of dorsal disc not defined by longitudinal suture-like folds; thoracic tracheal fold normallymarked on venter, punctuated by fine stippling in the subdorsal area; thoracic tracheal teethsmaller than remainder of marginal teeth, with margin indented at that point. A. littoralis furtherdiffers from dalbergiae in possessing a pair of dorsal disc hairs on abdominal segment VI, butwith no pair on segment VII; dalbergiae possesses the pair on segment VII, but not that on VI.Indeed, with kallarensis having the same pattern of dorsal hairs on the abdomen as littoralis, theonly apparent difference between kallarensis and dalbergiae is this transposition of one pair ofhairs (Figs 11 a, b, c). ALEUROPLATUS Quaintance & Baker Aleuroplatus Quaintance & Baker, 1914: 98. Type-species: Aleurodes quercusaquaticae Quaintance, byoriginal designation. Five species from the Buso area are regarded as belonging to Aleuroplatus (p. 305), althoughnone has been identified to species level. Aleuroplatus spp. 1 and 5 are evenly dark and similar ingeneral shape to bossi Takahashi; the remaining three species are less typical of the genus. ALEUROTRACHELUS Quaintance & Baker Aleurotrachelus Quaintance & Baker, 1914: 103. Type-species: Aleurodes tracheifer Quaintance, byoriginal designation. A single species is assigned to Aleurotrachelus, collected at Buso from Celtis philippinensis(Ulmaceae), Durandea sp. (Linaceae) and undetermined hosts. This species is tentativelyassigned to Aleurotrachelus and bears similarities in its outline and suture lines to dryandraeSolomon from Western Australia. ALEUROTUBERCULA TVS Takahashi Aleurotuberculatus Takahashi, 1932: 20. Type-species: Aleurotuberculatus gordoniae Takahashi, byoriginal designation. Five species of Aleurotuberculatus have been examined from New Guinea, but it has only beenpossible to name one; of the undetermined species (p. 306), sp. 2 is similar to melastomaeTakahashi, and sp. 4 is similar to siamensis Takahashi and bauhiniae Corbett. Aleurotuberculatus neolitseae Takahashi (Fig. 42)Aleurotuberculatus neolitseae Takahashi, 1934: 55. Syntype pupal cases, TAIWAN (TARI). A. neolitseae is a very distinctive species, and is well illustrated by Takahashi (1934) and Corbett(1935a). DISTRIBUTION. Papua New Guinea, Sarawak, Taiwan, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Buso area, on Anisoptera thurifera polyandra (Dipterocarpaceae), Diospyros sp.(Ebenaceae), Calophyllum sp. (Guttiferae), Elmerrillea papuana (Magnoliaceae), Gymnacranthera sp. THE WHITEFLY OF NEW GUINEA 321 (Myristicaceae), Myrtella sp. (Myrtaceae), Syzygium [= Eugenia] sp. (Myrtaceae), Schuurmansia hen-ningsii (Ochnaceae), Xanthophyllum papuanum (Xanthophyllaceae). Sarawak: Gunung Mulu NationalPark, on undetermined host. West Malaysia: Taman Negara National Park, Kuala Tahan, on undeter-mined woody vine (all BMNH). ALEUROTULUS Quaintance & Baker Aleurotulus Quaintance & Baker, 1914: 101. Type-species: Aleurodes nephrolepidis Quaintance, byoriginal designation. Aleurotulus arundinacea SinghAleurotulus arundinacea Singh, 1931: 88, pi. 35. Syntype pupal cases, INDIA (depository unknown). The eight pupal cases examined from New Guinea are all from a single bamboo clump, yet theyexhibit marked variation in the number and distribution of the long submarginal setae. Singhdescribed the species as having just 4 pairs of long submarginal setae, all on the cephalothorax,in addition to the caudal setae. The New Guinea specimens range from those with just onecephalothoracic pair and the caudal pair, to one with 4 cephalothoracic and 5 abdominal pairs.The other characters are as detailed by Singh, although the posterior marginal setae are ratherlonger. DISTRIBUTION. India, Papua New Guinea. MATERIAL EXAMINEDPapua New Guinea: Buso, on unidentified bamboo (Gramineae) (BMNH). ASIALEYRODES Corbett Asialeyrodes Corbett, 1935a: 841. Type-species: Asialeyrodes lumpurensis Corbett, by original designa-tion. Asialeyrodes sp. 1 (p. 306) is represented by a damaged pupal case; it is very flat and typicalof the species described by Corbett and Takahashi, and was collected from Pimelodendronemboinicum (Euphorbiaceae) at Buso. A second species (1 'Asialeyrodes, sp. 2) was collected from a forest-canopy vine, identified asSalacia sp. (Celastraceae); it has markedly convex pupae, unlike the described species ofAsialeyrodes, but with thoracic and caudal tracheal pores present, and with the submarginseparated from the subdorsum by a suture-like fold, it is tentatively assigned to Asialeyrodes. BEMISIA Quaintance & Baker Bemisia Quaintance & Baker, 1914: 99. Type-species: Aleurodes inconspicua Quaintance, by originaldesignation. Bemisia afer/hancocki-group Dialeurodoides afer Priesner & Hosny, 1934: 6, pi. 4. Syntype pupal cases, EGYPT (USNM; BMNH) [examined]. Bemisia afer (Priesner & Hosny) Habib & Farag, 1970: 8.Bemisia hancocki Corbett, 1936: 20, fig. 5. Syntype pupal case, UGANDA (BMNH) [examined]. [Synony- misedbyBink-Moenen, 1983.] Bemisia hancocki was described from cotton in Uganda, and variability of African material wasdiscussed by Mound (1965). The species was subsequently synonymised with afer by Bink-Moenen (1983). In the BMNH collection, there is a syntype of each of these taxa, that of aferbeing a slide-mount of a badly damaged pupal case which does not display many characters.Furthermore, a second slide preparation of 'hancocki', which was previously erroneouslylabelled as the 'type', and bears the same acquisition number as the syntype, displays differencesfrom the syntype. The true situation, even within the African material, is thus by no means clear.Specimens in the BMNH collection from the Pacific Region and from New Guinea clearly 322 J. H. MARTIN belong to the afer/hancocki-group, but vary much in size, the position of the vasiform orifice withrespect to the posterior margin of the pupal case, and the extent of dorsal tubercular sculpturing. DISTRIBUTION, 'afer' - Egypt; 'hancocki' - Mediterranean area, Africa, Madagascar, India,Pakistan; afer/hancocki-group - Papua New Guinea, Fiji, Tonga. MATERIAL EXAMINED Egypt: 1 pupal case (syntype of afer), Ibreem, on Ficus sycamorus (Moraceae) (BMNH). Uganda: 1pupal case (syntype of hancocki), on cotton (Gossypium, Malvaceae), coll. Hancock (BMNH); 1 pupalcase, on Vigna cajanus (Leguminosae), bearing same acquisition number as syntype of hancocki (BMNH).Papua New Guinea: 8 pupal cases, Lasanga Island, on Pterocarpus lindicus (Leguminosae) (BMNH). (?) Bemisia leakii (Peal) Aleyrodes leakii Peal, 1903: 87. Syntype pupal cases, INDIA (depository unknown).Bemisia leakii (Peal) Quaintance & Baker, 1914: 100. Dumbleton (19616) lists B. leakii on Colocasia sp. from Tahiti, quoting Cohic (1955), andillustrates the vasiform orifice characters while not indicating the source of the illustratedmaterial. Cohic simply listed the species as one found colonising Colocasia ('taro'), and gave nodescriptive data apart from a short observation about its feeding habits. Cohic's observation(translated) was This aleyrodid lives on the lower surface of the leaves of taro, but never indense colonies; the whitish nymphs are isolated from each other. Damage is rarely important.'These observations are also true of the specimens found on taro in New Guinea. Two slidesidentified as '? leakii' were loaned to the author by DSIR for comparison. This material wascollected by Cohic in Tahiti, and became part of the Dumbleton collection. It is undoubtedly thematerial cited in the text of Dumbleton's paper, and appears to be the source of the figure also.The vasiform orifice of the New Guinea specimens from taro matches that of Cohic's materialand also matches Dumbleton's figure, but none of these matches Peal's (admittedly poor)original figure. The author has not seen any further material identified as Bemisia leakii, so theTahiti and New Guinea records of this species must remain uncertain. DISTRIBUTION. Fiji, India, (Papua New Guinea), (Tahiti). MATERIAL EXAMINED Papua New Guinea: Buso, on Colocasia variety ('taro', Araceae) (BMNH). Tahiti: on Colocasia sp. andErythrina sp. (Leguminosae), F. Cohic coll. (DSIR). Bemisia pongamiae Takahashi Bemisia pongamiae Takahashi, 1931: 223, fig. 5. Syntype pupal cases, TAIWAN (TARI).DISTRIBUTION. Papua New Guinea, Taiwan, West Malaysia. MATERIAL EXAMINED Papua New Guinea: Buso strand-line, on Denis sp., small tree (Leguminosae). West Malaysia: TiomanIsland, Kampung Tekek, on ?Leguminosae (all BMNH). Bemisia tabaci (Gennadius)(Fig. 10) Aleurodes tabaci Gennadius, 1889: 1. Syntype pupal cases, GREECE (USNM).Bemisia tabaci (Gennadius) Takahashi, 1936: 110. Mound & Halsey (1978) list hosts of B. tabaci belonging to 63 families, and this whitefly is knownfrom most warmer parts of the world. The records from New Guinea, Sarawak and Java (seebelow) extend the recorded distribution. The New Guinea record quoted by Mound & Halseyrefers to the New Britain Province sample. DISTRIBUTION. Papua New Guinea (including New Britain), Java, Sarawak; also very widelydistributed in warmer parts of the world. THE WHITEFLY OF NEW GUINEA 323 MATERIAL EXAMINED Papua New Guinea: Lae, on Manihot esculenta c.v. (Euphorbiaceae); Lasanga Island, on Manihotutilissima and M. esculenta c.v.; Buso, on Sophora tomentosa (Leguminosae) ; New Britain Province, onIpomoea batatas (Convolvulaceae). Sarawak: Gunung Mulu National Park, on Manihot c.v. Java: Jakarta,on Manihot c.v. and undetermined shrub. (All BMNH.) CRENIDORSUM RussellCrenidorsum Russell, 1945: 55. Type-species: Crenidorsum tuberculatum Russell, by original designation. Crenidorsum was erected by Russell to accommodate 12 species from the Caribbean area;hitherto no further species have been assigned to the genus. Russell considered Crenidorsum,with the following group of characters, to be most closely allied toAleuroplatus, Aleurotrachelusand Aleurotulus: longitudinal differentiated fold/furrow in inner subdorsum on each side ofpupal case; vasiform orifice subcordate to broadly elliptical; operculum nearly filling vasiformorifice; lingula often folded into vasiform orifice, but when extended the appearance is as in Fig.14; submedian cephalic setae present; submargin not separated from dorsal disc by a completefold. The two species described below seem, in general appearance, to be closest to marginalefrom the Dominican Republic. A third New Guinea species (Crenidorsum sp. 1 - p. 306) is represented by a damagedspecimen from an undetermined host at Buso; it is unique in having the longitudinal subdorsalfurrows continued from abdominal segments IV- VII in the form of separate, segmental,comma-shaped creases. Crenidorsum lasangensis sp. n. (Figs 12-14) PUPAL CASE. Rather small, 0-74-0-92 mm long, 0-50-0-61 mm wide, about 1 -45-1-50 times as long as broad,widest at metathorax. Cuticle pale, colourless. Outline oval, but with margin almost straight for a shortdistance at thoracic and caudal tracheal areas, although not indented. Margin (Figs 13, 14) with singleregular row of teeth, about 13-16 occupying 0-1 mm of abdominal margin. Usual pairs of anterior andposterior marginal setae present, each arising from near apex of a marginal tooth, fine. Dorsum. Whole of cuticle between submedian area of dorsal disc and margin finely rugose (Fig. 13), therugae running almost parallel to margin, each sculptured by transverse pale markings, so that the rugaeresemble banded chromosomes in appearance. Cuticle of submedian area smoother, with well-markedpairs of submedian depressions on abdomen and thorax and very fine spinules medially on abdominalsegments. Pair of longitudinal subdorsal furrows present, though not as suture-like as in some othermembers of the genus; the furrows well-defined on thorax but becoming indistinct towards cephalic andabdominal areas. Median lengths of abdominal segments I-VI subequal, each about twice that of segmentVII. Large dorsal disc pores present, each with an adjacent tiny porette, distributed as in Fig. 12;additionally about 32 much smaller pore/porette adjacent pairs in an inner submarginal row (Fig. 13), withothers distributed in subdorsum. Longitudinal moulting suture reaching margin of case, transversemoulting sutures terminating in subdorsum. Vasiform orifice (Fig. 14) subcordate, a little wider than long,inset from posterior margin of case by about twice its own length; on average 36 /urn long, 45 /xm wide and70/um from margin. Operculum laterally-rounded trapezoidal, almost filling vasiform orifice. Lingula (Fig.14) longer than vasiform orifice, expanded apically into a spinulose club with a pair of fine setae, excludedfrom vasiform orifice if not folded. Thoracic and caudal tracheal areas not differentiated, caudal furrow notevident. Single pairs of cephalic, metathoracic, eighth abdominal and caudal setae present; cephalic andmetathoracic pairs up to 0-3 mm long (although often broken), eighth abdominal and caudal pairs abouthalf as long. Bases of caudal setae inset from margin of case by about twice height of marginal teeth. Venter. Caudal tracheal fold not defined. Thoracic tracheal folds defined only by a nebulous patch of finestipples located between forelegs and margin of case. A minute conical spine present at base of each middleand hind leg, not longer than broad at base. Median area of abdomen minutely roughened in transverse,segmental bands. Holotype pupal case, Papua New Guinea: Morobe Province coast, Lasanga Island, on Musa sp.(Musaceae), 18.ix.1979 (J. H. Martin 2586) (BMNH). Paratype pupal cases, Papua New Guinea: 12, same data as holotype (BMNH; USNM). 324 J. H. MARTIN COMMENTS. In the sample collected the pupae were sparsely scattered over the lower surface of alarge leaf of Musa, the plant growing in a 'garden' clearing. The pupae were not attended byants, and were not noticeably protected by waxy secretions. No adults were seen. Althoughnothing further is known about the biology of this insect, the importance of bananas andplantains throughout the tropics makes the description of this species important. C. lasangensis displays many characters associated with Aleurotulus species, particularly withregard to the often-excluded lingula and the long dorsal setae. However, the presence oflongitudinal subdorsal furrows and the form of the lingula indicate inclusion in Crenidorsum. C. lasangensis differs from differens Russell in the absence of a submedian pair of mesothor-acic setae, and in the absence of median tubercles on abdominal segments II-V; it differs frommarginale Russell in possessing remarkably long submedian setae, particularly the cephalic andmetathoracic pairs, and in the transverse moulting suture extending well beyond the longitudin-al subdorsal furrow; the dorsal disc pores in lasangensis are much more prominent than in otherspecies of Crenidorsum. Crenidorsum morobensis sp. n. (Figs 15, 16) PUPAL CASE. Pale, surrounded by broad mealy border, but without dense dorsal waxy covering. Rathersmall, 0-64-0-86 mm long, 0-45A)-63 mm wide. Shape oval, about 1-4 times as long as broad, widest atmetathorax, margin faintly indented at thoracic and caudal tracheal openings. Margin (Fig. 16) punctuatedby well-developed and evenly spaced teeth, about 11-12 occupying 0-1 mm of margin. Cuticle pale, butsome specimens dusky brownish in submedian area and in a narrow marginal ring at bases of teeth. Dorsum. Cuticle slightly wrinkled, with irregular folds running mesad from bases of marginal teeth intosubdorsum (Fig. 16); median parts of abdominal segments very finely spinulose. A pair of longitudinalsubdorsal furrows present, immediately lateral to submedian area and legs, these furrows running fromlevel of cephalic setae to approximately abdominal segment IV, remaining about parallel to margin of case(Fig. 15). Submedian area of dorsal disc somewhat raised and developed into a rhachis, delineated incephalothoracic and anterior abdominal region by a pair of rather sinuous and less sharply definedlongitudinal lines lying mesad of the subdorsal furrows. Posterior abdominal segments with raised lateralfolds extending posterolaterad into subdorsum. Median lengths of abdominal segments I-VII subequal.Submargin with single line of about 48 simple pores, inset about a marginal tooth-length from tooth bases;a little further mesad is a line of 9 pairs of minute hairs. Remainder of dorsal disc with evenly scatteredpores. Longitudinal moulting suture reaches anterior margin of pupal case, transverse moulting suturesterminate at the subdorsal furrows. Vasiform orifice (Fig. 16) subcordate, slightly elevated, a little widerthan long, inset a little more than its own width from posterior margin of pupal case; on average 32 /xmlong, 37 fjan wide and 40 /xm from posterior margin. Operculum roundly trapezoidal, almost exactly fillingvasiform orifice. Lingula longer than vasiform orifice, but normally folded into orifice and included.Thoracic and caudal tracheal openings at margin only marked by slight marginal indentations, marginalteeth not differentiated, caudal furrow hardly evident. Single pairs of short, pointed setae present on head,metathorax and abdominal segment VIII, shorter than vasiform orifice; caudal setae longer, length aboutequal to distance from vasiform orifice to margin, setal bases situated halfway between orifice and margin. Venter. Caudal tracheal fold not defined; thoracic tracheal folds sometimes marked in submargin by afaint pair of lines running mesad from the slightly indented margin. A minute conical spine present at baseof each middle and hind leg, each hardly longer than its own basal width. Holotype pupal case, Papua New Guinea: Morobe Province coast, Buso, on Myrtella sp. (Myrtaceae),3.X.1979 (/. H. Martin 2655) (BMNH). Paratype pupal cases. Papua New Guinea: 9, same data as holotype; 2, Buso riverbank, on IDecasper-mum sp. (Myrtaceae), ll.ix.1979 (JHM 2531); 1 (third instar larva), Buso, on Decaspermum sp.,14.ix.1979 (JHM 2565); 1, Buso riverbank, on undetermined sapling, 12.ix.1979 (JHM 2547); 1, Buso, onundetermined forest-canopy vine, 11. xi. 1979 (JHM 2841) (all BMNH). COMMENTS. Most specimens were found sparsely distributed on leaves of a Myrtella speciesgrowing on beach-top sand between mangroves and the sea. Further specimens were taken fromother hosts (see paratype data), but there is insufficient material generally to enable anyconclusions to be drawn on the likely host range of the species. The pupae were not attended byants. THE WHITEFLY OF NEW GUINEA 325 C. morobensis differs markedly from the described species of Crenidorsum in possessing adorsal rhachis. DIALEURODES Cockerell Aleyrodes (Dialeurodes) Cockerell, 1902: 283. Type-species: Aleyrodes citri Riley & Howard [= Aleyrodes citri Ashmead], by original designation.Dialeurodes Cockerell; Quaintance & Baker, 1914: 97 [raised to genus]. In addition to two named species and one here described as new, eight undetermined species ofDialeurodes from New Guinea have been examined (p. 307). Dialeurodes sp. 1 resembles ixoraeSingh (1931, figured) in having the submedian area delineated by a line of small papillae, but itdiffers in other respects; sp. 2 belongs to a group for which Quaintance & Baker (1917) used thesubgenus Rabdostigma; sp. 6 resembles subrotunda Takahashi. Dialeurodes decaspermi sp. n. (Figs 17-19) PUPAL CASE. Large, of striking appearance, conspicuous against the rather pale leaf underside of the host.Length 1-70-2-55 mm, width 1-40-2-35 mm, broadly oval to almost circular. Margin slightly irregular, butentire, not crenulate or castellate. Anterior and posterior pairs of marginal setae present, fine. Dorsum. Possessing a most remarkable sclerotic pattern (Fig. 17): only the extreme marginal area,cephalothoracic subdorsum, median part of abdomen and a pair of subdorsal abdominal patches pale, theremainder dark brown to black, the resultant pattern resembling a pale anchor on a dark background.Paler marginal area with very fine lines running mesad as far as the darkly pigmented submargin.Pigmented cuticle finely granular in appearance, granulations not apparent in paler areas. Whole ofdorsum except paler marginal band bearing many evenly spaced disc pores and many subcircular markingswhich give the dorsum a 'cobbled' appearance. Longitudinal moulting suture only reaches anteriorly as faras the subdorsal pale zone, transverse moulting sutures terminate above outer edges of hind legs. Vasiformorifice (Fig. 19) subcordate, 50-70 pm long, a little wider than long, situated 5-5-8-0 times its own lengthfrom posterior margin of pupal case; posterolateral margin of orifice smooth, dark, much thickened.Operculum trapezoidal, almost filling vasiform orifice. Lingula (Fig. 19) apically setose, with 4 lateralprocesses, longer than vasiform orifice but apical section usually recurved to appear shorter than orifice.Caudal and thoracic tracheal pores well marked, situated at points of slight marginal indentation. Caudalfurrow (Fig. 18) marked in posterior part by longitudinal rugae extending from caudal tracheal pore, andalong remainder of its length by denser cuticular markings. Median lengths of abdominal segments I-VIIsubequal. Minute pairs of cephalic and first and eighth abdominal setae present, capitate, resembling tinymatch sticks. A similar pair, the caudal setae, present about half-way between vasiform orifice andposterior margin of case (Fig. 18). Venter. Thoracic and caudal tracheal folds defined by bands of fine stipples. Antennae rather long,almost reaching articulation of middle legs. A fine seta present at base of each middle and hind leg, similarto ventral abdominal setae, although a little shorter. Holotype pupal case, Papua New Guinea: Morobe Province coast, Buso riverbank, on Decospermum sp.(Myrtaceae), 16.x. 1979 (/. H. Martin 2713) (BMNH). Paratype pupal cases. 5, same data as holotype; 48, same locality and host, ix-x.1979 (JHM 2528, 2621,2690, 2755) (BMNH; USNM). COMMENTS. The striking pupae of this species were found only on small bushy plants identified asa Decaspermum sp. (Myrtaceae) growing in situations alongside the river and on the beach-topat Buso. They were invariably seen on the young leaves near the growing points of the plants.The apparent absence of the species from other plants in the area throughout a three-monthperiod suggests strong host specificity. The pupae were found in shallow concavities on the leafundersides, thus remaining flush with the leaf surface. There appears to be sexual dimorphism inthis species, with two distinct size ranges within the overall size range described above, but noadults were obtained to confirm this. The pupae were not protected by any visible waxy orwoolly secretions, and were not attended by ants. Individuals were fairly evenly scattered overthe affected leaves. D. decaspermi may be distinguished from other Dialeurodes species by its most unusualsclerotic patterning, combined with its large size and distinctively sculptured dorsum. 326 J. H. MARTIN Dialeurodes kirkaldyi (Kolinsky) Aleyrodes kirkaldyi Kolinsky, 1907: 95, fig. 2. Synlype pupal cases, HAWAII (HDA; USNM).Dialeurodes kirkaldyi (Kolinsky) Quainlance & Baker, 1914: 98. DISTRIBUTION. Irian Jay a, and several counlries in each zoogeographical region excepling IheMalagasy Region. MATERIAL EXAMINEDIrian Jaya: Sukarnapura, onJasminum sp. (Oleaceae) (BMNH; BPBM). Dialeurodes psidii Corbell(Fig. 43) Dialeurodes psidii Corbell, 1935a: 734. Synlype pupal cases, WEST MALAYSIA (presumed losl).Dialeurodes lumpurensis Corbell, 1935a: 739. Synlype pupal cases, WEST MALAYSIA (presumed losl).Syn. n. Corbett (1935a) described seven species of Dialeurodes in which the longitudinal and transversemoulting sutures are joined by a cephalothoracic suture, giving rise to distinct 'trapdoors' whichcan become detached as the adult emerges. Corbett's material on which he based his 1935publication is thought to be destroyed, but specimens from New Guinea vary between samplesin the degree of cuticular marking, and even within samples to some degree; it is considered thatpsidii and lumpurensis are synonymous, with psidii having page priority. Certainly, fromCorbett's observations, and from the study of the material seen from New Guinea, it seems thatpsidii has a wide range of hosts, and it may be that other species in this group will prove to beconspecific also. DISTRIBUTION. Irian Jaya, Papua New Guinea, Sarawak, Thailand, West Malaysia. MATERIAL EXAMINED Irian Jaya: Biak, on Ficus sp. (Moraceae) (BMNH). Papua New Guinea: Buso, onAnisoptera thuriferapolyandra tree crown (Diplerocarpaceae), Celtis sp. (Ulmaceae), Decaspermum sp. (Myrlaceae), Dios-pyros sp. (Ebenaceae), Euodea sp. (Rulaceae), Gyrocarpus sp. (Gyrocarpaceae), Lophopetalum sp.(Celaslraceae), Macaranga sp. (Euphorbiaceae), Myrtella sp. (Myrlaceae), Premna sp. (Verbenaceae),Xanthophyllum papuanum (Xanlhophyllaceae), and undelermined hosls; Lasanga Island, on Euodea sp.(Rulaceae); Wau Ecology Inslilule, on Ficus sp. (Moraceae) (all BMNH); Kralke Mis, on Neonauclea sp.(Naucleaceae) (BMNH; BPBM). Sarawak: Gunung Mulu National Park, on undetermined hosl(BMNH). West Malaysia: Taman Negara Nalional Park, on Melaslomalaceae; Genling Highlands, onundelermined hosl. (All BMNH.) DIALEUROPORA Quaintance & Baker Dialeurodes (Dialeuropora) Quaintance & Baker, 1917: 434. Type-species: Dialeurodes (Dialeuropora) decempuncta Quaintance & Baker, by monotypy.Dialeuropora Quainlance & Baker; Takahashi, 1934: 46 [raised lo genus]. Most specimens from New Guinea have been identified as the very common and widespreadspecies decempuncta, but one sample (sp. 1, p. 307) from Elmerrillea sp. (Magnoliaceae) containsa species resembling brideliae (Takahashi), with the vasiform orifice rather small relative to itsdistance from the posterior margin of the pupal case and an apparent absence of short lanceolatesetae around the submargin. Dialeuropora decempuncta (Quaintance & Baker)(Fig. 40) Dialeurodes (Dialeuropora) decempuncta Quaintance & Baker, 1917: 434. Synlype pupal cases, SRI LANKA, PAKISTAN (USNM). Dialeuropora decempuncta (Quainlance & Baker) Takahashi, 1934: 46.Dialeuropora perseae (Corbell), 1935a: 749. Synlype pupal cases, WEST MALAYSIA (presumed losl). Syn. n. THE WHITEFLY OF NEW GUINEA 327 Mound & Halsey (1978) concluded that D. decempuncta varies considerably, particularly in theprecise form and size of the submarginal setae, and accordingly synonymised setigerus (Taka-hashi, 1934) and dothioensis (Dumbleton, 1961a) with decempuncta. Dialeuropora perseae(Corbett, 19350) was described as differing from setigerus only in the presence of 'a ring of smallsubmarginal pores around the case, and of similar-sized pores distributed throughout thedorsum'. The BMNH paratype of dothioensis possesses these pores, in common with manyother specimens of decempuncta. Importantly, several samples contain both individuals withobvious small pores and those with such small pores not evident, and D. perseae is regarded as ajunior synonym of decempuncta. The samples of decempuncta from Breynia in New Guinea exhibit a further variation, which isvery confusing when encountered in its most extreme form - a tendency to lose the 5 pairs oflarge simple subdorsal pores which are the principal diagnostic feature of the genus. One sampleeach from Lasanga Island and Buso contains individuals which vary from those lacking the 5pairs of pores to those with the pores all present but small, and individuals from two further Busosamples all lack pores. The remainder of the morphological characters, particularly the shortlanceolate setae in the submargin and the shape and size of the vasiform orifice and lingula, aretypical. The samples from ^.Breynia (Wau) and Glochidion (Buso) have normally developed largepores and submarginal setae, but have unusually long first and eighth abdominal, caudal andcephalic setae - longer than the width of the vasiform orifice. Material from Tonga (BMNH), identified as decempuncta and listed in Mound & Halsey,does not match even the variants described here and should be regarded as belonging to anunidentified species of Dialeuropora. DISTRIBUTION. Austro-Oriental Region: Java, Papua New Guinea, Sarawak, Singapore, WestMalaysia. Oriental Region: Cambodia, India, Pakistan, Sri Lanka, Taiwan, Thailand. PacificRegion: New Caledonia. Australasian Region: Northern Territory. MATERIAL EXAMINED Papua New Guinea: Buso, on Alphitonia sp. (Rhamnaceae), Breynia sp. (Euphorbiaceae), Colocasiac.v. (Araceae), Glochidion (Euphorbiaceae), Macaranga sp. (Euphorbiaceae), Pometia pinnata treecrown (Sapindaceae), Tetracera sp. (Dilleniaceae) and undetermined host; Lasanga Island, on Breyniasp., Macaranga sp., Pterocarpus tindicus (Leguminosae) and ?Leguminosae; Wau Ecology Institute, on? Breynia sp. (all BMNH). Java: Jakarta, on Musa sp. (Musaceae) and Psidium guajava (Myrtaceae)(BMNH). Sarawak: Gunung Mulu National Park, on IMillettia sp. (Leguminosae) and undeterminedhosts (all BMNH). Singapore: on fEugenia sp. (Myrtaceae) (BMNH). West Malaysia: Taman NegaraNational Park, Kuala Tahan, on Flemingia macrophylla (Leguminosae) and undetermined hosts; TiomanIsland, on Bauhinia sp. vine (Leguminosae) and ?Leguminosae (all BMNH). New Caledonia: DothioRiver bridge (paratypes of Dialeurodes dothioensis Dumbleton, F. Cohic. coll. , synonymised by Mound &Halsey, 1978), on undetermined host (BMNH). Australia: Northern Territory, on Eucalyptus sp.(Myrtaceae) (BMNH). INDOALEYRODES David & Subramaniam Indoaleyrodes David & Subramaniam, in Krishnamurthy, Raman & David, 1973: 75. Type-species: Indoaleyrodes pustulatus David & Subramaniam, by monotypy.Indoaleyrodes David & Subramaniam, 1976: 199. The name Indoaleyrodes pustulatus was first published in an account of the feeding damage toleaves ofMorinda tinctoria (Rubiaceae) (Krishnamurthy, Raman & David, 1973, quoting David& Subramaniam, 1972). The 1972 reference was given as 'Studies on some Indian Aleyrodidae(in press), Mem. zool. Surv. India, Calcutta' which in fact appeared, in a different journal, in1976. Although a description of the work of an animal constitutes an 'indication' for thepurposes of Article 25 of the Code, the definitive description of /. pustulatus appears in David &Subramaniam (1976), and the 'holotype' and 'paratypes' must properly be regarded as syntypes(3 in BMNH [examined]). /. pustulatus is considered a junior synonym (syn. n.) of Dialeurodes laos Takahashi (1942),and laos is here included in Indoaleyrodes (comb. n.). 328 J. H. MARTIN Indoaleyrodes differs from Dialeurodes primarily in its deeply indented thoracic and caudaltracheal pores, combined with a triangular vasiform orifice in which the operculum does notoccupy most of its area as in Dialeurodes. In addition to laos and pseudoculatus (see below), it is clear that Parabemisia reticulataDumbleton (19610) should also be included in Indoaleyrodes (comb. n.). Indoaleyrodes pseudoculatus sp. n. (Figs 20, 21) PUPAL CASE. Outline almost circular, only 1-15-1-25 times longer than wide, widest opposite hind legs,margin indented slightly at thoracic and caudal tracheal openings, but with actual tracheal pores inset frommain marginal outline by about 3 times pore diameter (Fig. 20). Apparently sexually dimorphic, withdimensions in the ranges 1-34-1-41 mm by 1-15-1-21 mm and 1-03-1-09 mm by 0-86-0-90 mm. Marginsmooth and slightly irregular, with single pairs of short, fine anterior and posterior marginal setae. Cuticlepale, transparent. Dorsum. Closely set parallel lines run mesad from margin into outer subdorsum, length of these lines alittle greater than distance from main marginal outline to tracheal pores (Fig. 20, inset). Thorax bearing apair of very prominent oval glandular areas with polygonal reticulate pattern, resembling compound eyes;each patch situated about mid-way between thoracic tracheal pore and legs, and a little longer thanvasiform orifice. A pair of comma-shaped patches of similar appearance adjacent to vasiform orifice,similar in length to orifice (Fig. 21). Remainder of dorsal surface smooth, with a few scattered tiny discpores. Moulting sutures rather faint, not apparently reaching margin. Median length of abdominalsegment VII about half that of segment VI . Dorsal disc not defined . Only 3 pairs of dorsal setae present , allfine and rather short, cephalic and caudal pairs hardly longer than diameter of tracheal pore: cephalic setaeplaced at level of apices of front legs; eighth abdominal setae lateral to anterior edge of vasiform orifice;caudal setae slightly anterolaterad of caudal tracheal pore. Vasiform orifice (Fig. 21) large, triangular,1-15-1-25 times longer than wide, inset from posterior margin of case by about 1-5-2-0 times its own length;posterior part of internal margin of vasiform orifice produced into a squared process with width similar todiameter of head of lingula. Caudal furrow not marked. Operculum rounded-trapezoidal, occupying basalhalf of vasiform orifice. Lingula long and stout, head developed into a spinulose club with a pair of basallateral lobes and an apical pair of long spines which overlap apex of vasiform orifice; lingula exposed butincluded within vasiform orifice. Venter. Caudal and thoracic tracheal folds marked by small groups of coarse spinules running mesad inwidening bands from the pores (Fig. 20) ; caudal fold achieving width of vasiform orifice at level of posteriorabdominal spiracle, and thoracic folds achieving similar maximum width. A short spine present at base ofeach middle and hind leg. Mouthparts well developed, with distinct aphid-like ultimate rostral segmentof similar length (longitudinal axis) to operculum. Ventral abdominal setae similar to dorsal eighthabdominal setae. Holotype pupal case, Papua New Guinea: Morobe Province coast, Buso, on tree-crown foliage ofSyzygium sp. (Myrtaceae), S.x.1979 (/. H. Martin 2674) (BMNH). Paratypes. 13 pupal cases, 1 larva, same data as holotype (BMNH). COMMENTS. This species is known from a single collection from young crown leaves of a 26-metrehigh tree of a Syzygium sp., reached from a walkway in the forest canopy. The pupae were notattended by ants, and no adults were found. The genus Indoaleyrodes is very little known, so nospeculation may be made about the likely host range of this remarkable new species. At least twoof the specimens were parasitized. /. pseudoculatus is at once separated from the other described species by the remarkableglandular areas on the cephalothorax, which resemble compound eyes, and by the comma-shaped glandular areas on each side of the vasiform orifice. Two pupal cases which closely resemble pseudoculatus are present in the BMNH collection(Indoaleyrodes sp. 2, p. 307). One is incomplete and the second evidently damaged by a fungusbefore it was collected. They differ from pseudoculatus in having the abdominal glandular areasfused to form a U-shape, which starts anterior to the vasiform orifice and crosses the caudalfurrow half-way between the orifice and the caudal pore; they also differ in the shape andposition of the vasiform orifice. They were collected from an undetermined host at 6,000 ft in theKampere Barola Divide. THE WHITEFLY OF NEW GUINEA 329 Another species of Indoaleyrodes (sp. 1, p. 307) is represented by a single pupal case fromCeltis philippinensis (Ulmaceae), but this possesses neither cephalothoracic nor abdominalglandular areas. NEOMASKELLIA Quaintance & BakerNeomaskellia Quaintance & Baker, 1913: 91. Type-species: Aleurodes comata Maskell, by monotypy. Neomaskellia bergii (Signoret)(Fig. 38) Aleurodes bergii Signoret, 1868: 395. Syntype pupal cases, MAURITIUS (depository unknown).Neomaskellia bergii (Signoret) Quaintance & Baker, 1914: 104. DISTRIBUTION. Papua New Guinea; also widely distributed throughout Africa, the Orient, SouthEast Asia and the Pacific Region. MATERIAL EXAMINED Papua New Guinea: Lasanga Island, on Saccharum ? officinarum (Gramineae), attended by ants(Oecophylla smaragdina); Bubia (Lae), on Saccharum officinarum and Cenchrus ciliaris (Gramineae); PigIsland (Madang), on Saccharum officinarum; Amax mining camp (Hessen Bay, Morobe Province), onundetermined grass (all BMNH); Wau, on undetermined grass (BMNH; BPBM); Laloki, on undeter-mined grass (BMNH; BPBM); 'Papua', on Saccharum sp. (BPBM). ORCHAMOPLA TVS Russell Aleuroplatus (Orchamus) Quaintance & Baker, 1917: 400. Type-species: Aleuroplatus (Orchamus)mammaeferus Quaintance & Baker, by monotypy. [Homonym of Orchamus Stal, 1876: 30 (Orthop-tera).] Orchamus Quaintance & Baker; Dumbleton, 1956: 13 [raised to genus]. Orchamoplatus Russell, 1958: 390. [Replacement name for Orchamus Quaintance & Baker.] Since Russell (1958) revised the genus, with 10 included species, a further three, dumbletoni(Cohic, 1959), perdentatus Dumbleton (19610) and sudaniensis Gameel (1968), have beendescribed. As sudaniensis does not possess submarginal glands of the dentate type characteristicof Orchamoplatus, it should be included in Neoaleurotrachelus Takahashi & Mamet [synony-mous v/ithJeannelaleyrodes Cohic (Bink-Moenen, 1983)] (comb, n.); this opinion was expressedto Gameel by Mound (pers. comm. , 1967) prior to publication of the description. Orchamoplatus niuginiisp. n. (Figs 22, 23) PUPAL CASE. Completely pale to slightly dusky, ovoid, rather large: 0-85-1-25 mm long, 0-59-0-87 mmwide, mostly 1-4-1-5 times as long as wide, widest at abdominal segment II. Margin irregularly crenulate,about 13-15 small, rounded crenulations to 0- 1 mm of abdominal margin, with fine folds running mesad fora short distance from the bases of the crenulations (Fig. 23). Margin much indented at regions of thoracicand caudal tracheal openings, marginal crenulations very strongly differentiated to form thoracic andcaudal tracheal combs. Usual pairs of anterior and posterior marginal setae present. Dorsum. Submargin with single row of evenly spaced dentate glands (the term 'gland' used by Russell,1958), 50-70 pairs in total, with 16-20 pairs anterior to thoracic combs and 38-50 pairs posteriorly (Fig. 22);structure of glands as in Fig. 23. Row of submarginal dentate glands mostly inset from margin by 3-4 timeslength of gland crown, although row of glands and margin converge at tracheal regions. Dentate glandsadjacent to thoracic and caudal combs not different to remaining glands, not reaching margin. Thoracicand caudal tracheal combs well developed, set in deep marginal concavities, with fluted areas at the toothbases extending mesad of the line of dentate glands; thoracic combs normally with 10 or 11 teeth, caudalcomb with 8-10 teeth, combs evidently arched dorsally. Seven pairs of tiny submarginal setae present, 3cephalic, 2 thoracic and 2 mid-abdominal; the line of these setae completed by minute, evenly spacedporettes. Most specimens with pronounced cephalothoracic fold parallel to margin between submargin andsubdorsum, fold extending posteriorly half-way to thoracic combs and then fading gradually. Longitudinalmoulting suture not reaching margin of pupal case, terminating at cephalothoracic fold; transverse 330 J. H. MARTIN moulting sutures terminating in outer subdorsum. Dorsal cuticle mesad of dentate glands virtually smooth,punctuated only by very fine transverse lines of spinulose sculpturing on median part of abdominalsegments II- VII, by coarser spinulose corrugations on median part of abdominal segment VIII (Fig. 23),and by an evenly spinulose patch on abdominal segment I (Fig. 22). Single pairs of cephalic, first and eighthabdominal setae present; cephalic and eighth abdominal pairs fine, cephalic pair the shorter; pair of setaeon abdominal segment I in form of thick, chitinous spatulas which are often lost in preparation forslide-mounting. Disc pores well developed, about 12 pairs distributed as shown in Fig. 22. Vasiform orifice(Fig. 23) elevated and thus liable to distortion in slide-mounted specimens, subcircular, inner walls verticalwith parallel vertical ridges; orifice about 60 ^im in diameter, usually appearing slightly wider than long,and inset a little more than its own length from posterior margin of pupal case (measured from apex ofcaudal tracheal comb). Operculum subtrapezoidal, almost filling cross-sectional area of vasiform orifice.Lingula shorter than vasiform orifice, head obscured by operculum, apparently rather square and denselyspinulose. Caudal setae very long and fine, up to 0-21 mm long, bases situated just mesad of line of dentateglands, on inner border of caudal comb area. Caudal furrow not marked. Venter. Thoracic and caudal tracheal folds virtually unmarked. A minute conical spine present at base ofeach middle and hind leg. Median parts of abdominal segments II-VIII very finely spinulose. Bases ofventral abdominal setae near anterior edge of vasiform orifice. Rostral base setae tiny, smaller thanleg-base spines. Holotype pupal case, Papua New Guinea: Morobe Province coast, Lasanga Island, on Calophylluminophyllum (Guttiferae), 18.ix.1979 (J. H. Martin 2581) (BMNH). Paratypes. Papua New Guinea: 47 pupal cases, 1 larva, same data as holotype; 30 pupal cases, MorobeProvince coast, Buso, same host, 30. ix. 1979 (JHM 2643); 19 pupal cases, 3 larvae, Lasanga Island, samehost, 7.xi.l979 (JHM 2816); 3 pupal cases, 1 larva, Buso, on Durandea sp. (Linaceae) at canopy level,8.X.1979 (JHM 2676) (BMNH; USNM). COMMENTS. The species occurs in very dense colonies on the underside of mature leaves oflittoral Calophyllum inophyllum (Guttiferae), and is not attended by ants. These denseaggregations of pupae and larvae are covered with a glassy, translucent secretion which becomeshard when specimens are stored dry; folding the leaf then causes flakes of secretion to peel awaywith the insects embedded in it. The earlier larval exuviae often remain attached to the dorsa ofthe later stages, and sometimes all instars are represented in a stack. No adults were found andthere was no sign of empty cases from which adults had emerged. A few specimens were alsotaken from a leaf of a species of Durandea vine in the forest canopy, but this provides the onlyfirm evidence of oligophagy. O. niuginii is similar to mammaeferus (Quaintance & Baker) and montanus (Dumbleton), butdiffers from both in the following characteristics: submarginal dentate glands further inset frommargin (3-4 times length of gland crown, compared with up to twice); dentate glands adjacent toboth thoracic and caudal combs not larger than remainder of glands, not reaching margin ofcase; generally a larger species, pupal cases mostly over 1-0 mm long. O. niuginii further differsfrom mammaeferus in the possession of a pair of cephalic setae. O. niuginii bears no closeresemblance to calophylli Russell, which was described from a species of Calophyllum in Tonga. A further colony of Orchamoplatus at Buso was taken from a tree-crown leaf, possiblybelonging to a species of Lophopetalum (Celastraceae). The mounted specimens agree withniuginii in most respects, but differ in the apparent shape of the vasiform orifice, and inpossessing less-indented thoracic and caudal tracheal areas (apices of centre teeth in combsstand proud of margin of case). A field note states that all were parasitised, appearing black, andthe venters failed to detach from the leaf. The mounted specimens, with parasites removed, arepale but ventrally incomplete; they are possibly morphologically modified by the parasites andare tentatively determined as niuginii. PARABEMISIA TakahashiParabemisiaTakahashi, 1952: 21. Type-species: Parabemisia maculata Takahashi, by original designation. Takahashi (1952) erected Parabemisia to accommodate his new species maculata, Bemisia acerisTakahashi and Bemisia myricae Kuwana. Dumbleton (1961a) assigned his species reticulata to Parabemisia on the basis of its laterally THE WHITEFLY OF NEW GUINEA 331 bilobed lingula head, while noting that other characters were not typical of the genus; reticulatais here transferred to Indoaleyrodes David & Subramaniam (1973) (comb. n.). The five species currently placed in Parabemisia are keyed below, in a modified version ofTakahashi's original key. Key to species of ParabemisiaPupal cases 1 Submarginal setae very short, sometimes hardly recognisable, numbering 11 pairs (excluding anterior and posterior marginal pairs and caudal pair). Head of lingula rather triangular,evenly tapering from near its base 2 - Submarginal setae long and conspicuous , subequal in length to vasiform orifice and caudal setae , normally numbering 11 or 13 pairs (excluding anterior and posterior marginal pairs and caudalpair) . Head of lingula rather ovoid , not tapering evenly from near its base 3 2 Head of lingula slender, over twice as long as wide; vasiform orifice elongated, caudal furrow distinct and slender ; tracheal folds (ventral) with fine dots aceris (Takahashi) - Head of lingula not slender, rather abruptly tapering; vasiform orifice not elongated, caudal furrow not well defined , but wider ; tracheal folds without fine dots maculata Takahashi 3 Submargin normally with 1 1 pairs of setae (excluding caudal pair) . Dorsum bearing many large , tubercular pores, each of which is about half as wide as operculum jawani sp. n. (p. 331) - Submargin normally with 13 pairs of setae (excluding caudal pair). Dorsum not bearing large, tubercular pores 4 4 Usually entirely pale, very occasionally with slight duskiness. Vasiform orifice with lateral margins straight or slightly concave myricae (Kuwana) - Never entirely pale , always with submargin and subdorsum brown . Vasiform orifice subcordate , lateral margins markedly convex, thickened myrmecophtia sp. n. (p. 332) Parabemisia jawani sp. n. (Figs 24, 25) PUPAL CASE. Pale, oval, 0-8-1-1 mm long, broadest at abdominal segment III, 1-2-1-4 times as long as wide.Posterior margin flattened but hardly indented, without marginal indentation towards thoracic trachealareas. Margin bluntly and rather unevenly crenulate, with about 12 teeth occupying 0-1 mm of margin.Thoracic and caudal tracheal openings each marked by a simple notch which is hardly wider than onemarginal tooth (Fig. 25). Posterior marginal setae present, but anterior pair apparently lacking. Dorsum. Twelve pairs of Submarginal setae present, 6 on cephalothorax and 6 on abdomen, includingcaudal pair; setae fine, a little longer than vasiform orifice, their bases situated just mesad of marginalteeth. A pair of tiny hairs present on each of abdominal segments I and VIII, but cephalic pair absent.Dorsum punctuated by evenly distributed, large, circular tubercular pores, each about half of opercularwidth in diameter, and with a paler central opening in stained specimens (Fig. 24). Median line of abdomendevoid of these pores, with evenly staining cuticle. Remainder of dorsal cuticle smooth, but staining picksout variations in sclerotisation which appear as irregular polygonal plates delineated by paler lines. Medianlengths of abdominal segments I- VI subequal, but that of segment VII much reduced. Median lengths ofmeso- and metathoracic segments also subequal. Both transverse and longitudinal moulting suturesreaching margin, the dorsal halves of cephalothorax easily becoming detached. Vasiform orifice (Fig. 25)rounded-triangular to trapezoidal, with posterior margin abruptly truncated although not marked by asharp line; orifice inset from posterior margin by about 3 times its own length; internal walls of orificesmooth, not notched. Operculum laterally-rounded trapezoidal, occupying about two-thirds of vasiformorifice. Lingula exposed, only just included in orifice; head dark, finely spinulose, with a pronounced pairof lateral processes ; apical setae not apparent. Caudal furrow well marked by a line of darkly staining spots. Venter. Caudal and thoracic tracheal folds marked by bands of fine dots running mesad from margin,reaching vasiform orifice and outer edges of legs respectively (Fig. 25). A minute conical spine present atbase of each middle and hind leg. Ventral abdominal setae long, fine, about as long as vasiform orifice.Abdominal spiracles close to ventral abdominal setae, appearing claw-like. Holotype pupal case, Papua New Guinea: Morobe Province coast, Jawani Island, on undeterminedwoody host, 2.xi.l979 (J. H. Martin 2789) (BMNH). Paratype pupal cases. 16, same data as holotype (BMNH). 332 J. H. MARTIN COMMENTS. Nothing is known about the biology of this insect, which was encountered only oncefeeding on an undetermined woody host alongside mangroves on a small offshore island. Thepupae were not attended by ants. P. jawani is very distinctive with its large dorsal tubercular pores, and can be separated by thecharacters given in the key. Parabemisia myrmecophila sp. n. (Figs 26, 27) PUPAL CASE. Outline rather pear-shaped, 0-65-0-90 mm long, 0-45-0-62 mm wide, broadest at abdominalsegment III, mostly 1-4-1-5 times as long as wide. Brown, with variable median area pale, ranging fromindividuals which are almost evenly brown to those with whole of submedian area of dorsal disc pale; palearea widest on cephalothorax (Fig. 26). Margin (Fig. 27) evenly crenulate, with about 20 rounded-triangular teeth occupying 0-1 mm of margin. Margin very gently indented towards thoracic area at pointwhere a tracheal comb is slightly differentiated from remainder of marginal teeth; posterior margin of caserounded or flattened, but not indented. Dorsum. A row of 14 pairs of submarginal setae present, including caudal pair, setal bases situated justmesad of marginal teeth; submarginal setae fine, similar in length to the single pairs of cephalic and first andeighth abdominal setae, subequal to length of vasiform orifice. Dorsal disc pores similar in size to setalbases, scattered evenly, with 4-5 pairs on each of abdominal segments III- VIII, and about 30 pairs oncephalothorax. Tiny disc porettes also present. Cuticle slightly wrinkled, with a few darker granularmarkings on the brown areas. Median length of abdominal segment VII much less than half that of segmentVI, abdomen appearing 7-segmented along median line. Longitudinal and transverse moulting suturesreaching margin of case. Vasiform orifice (Fig. 27) rather large, nearly 0-1 times length of case, subcordate,posterolaterally much thickened with toothed inner margin, posterior margin rather flattened; orifice60-80 /u,m long and wide, situated 65-100 /um from posterior margin of case. Operculum trapezoidal, onlyabout half filling orifice. Lingula shorter than vasiform orifice, exposed but included; lingula headspinulose, with an apical pair of setae which are longer than lingula head, bearing a pair of small laterallobes at base of head, each lobe with a short seta. Caudal furrow discernible in most specimens, althoughmarked to a rather variable degree. Venter. Caudal and thoracic tracheal folds not defined. A minute conical spine at base of each middle andhind leg, hardly longer than its own basal width. Ventral abdominal setae fine, similar to posterior marginalsetae, but situated under vasiform orifice and thus not easy to see. Rostral base setae relatively large, overhalf length of anterior marginal setae, fine. Holotype pupal case, Papua New Guinea: Morobe Province coast, Buso, on Cryptocarya sp.(Lauraceae), 27.ix.1979 (/. H. Martin 2629) (BMNH). Paratypes. Papua New Guinea: 48 pupal cases, same data as holotype; 11 pupal cases, 7 larvae, Buso, onAnisoptera sp. (Dipterocarpaceae), 27. ix. 1979 (JHM 2626); 44 pupal cases, 4 larvae, Buso, on Macarangasp. (Euphorbiaceae), 27. ix. 1979 (JHM 2627); 44 pupal cases, 1 larva, Buso, on Prunus sp. (Rosaceae),26.ix.1979 (JHM 2622) (BMNH; USNM). COMMENTS. This species was always found in very dense colonies on undersides of young leavesof small woody saplings growing on the forest floor. In each case, the colony was vigorouslyattended by ants, Rhoptromyrmex melleus (Emery) . In one case, adults were observed emergingand then congregating in groups on the very youngest leaves of the same plant, Cryptocarya sp.It seems likely that in this way the colony can keep pace with the growth of the plant while itremains suitable for colonisation. P. myrmecophila may be distinguished from other described species of Parabemisia bycharacters given in the key. PEAL/l/SQuaintance & BakerPealius Quaintance & Baker, 1914: 99. Type-species: Aleyrodes maskelli Bemis, by original designation. There are currently 28 species assigned to Pealius, and the genus is particularly difficult todefine; typically, the vasiform orifice has a 'false' posterior margin, continuing posteriorly as ashallow depression which is distinct from the caudal furrow (Fig. 28). Five species from New Guinea are assigned to the Pealius-group, and are listed with host dataon p. 308. THE WHITEFLY OF NEW GUINEA 333 RHACHISPHORA Quaintance & Baker Dialeurodes (Rhachisphora) Quaintance & Baker, 1917: 430. Type-species: Dialeurodes (Rhachisphora) trilobitoides Quaintance & Baker, by original designation.Rhachisphora Quaintance & Baker; Takahashi, 1952: 22 [raised to genus]. In addition to the specimen tentatively identified as R. ardisiae (see below), two further sampleshave been studied from New Guinea (p. 308). Rhachisphora sp. 1 has an oval pupal case similarto ardisiae and some other species, differing markedly from the anteriorly flattened species ofthe trilobitoides-group; this sample was collected from tree-crown leaves of a Schefflera sp.(Araliaceae) in Buso forest. The second sample (Rhachisphora^., sp. 2) contains pupal caseswhich are almost circular and markedly convex: the rather wide submedian area is smooth andbrownish; the submargin and outer subdorsum are pale; the inner subdorsal area comprises 2lateral, arcuate, imbricate zones which stain very deeply and which have about 20 radial 'spokes'leading into the submargin, forming a pronounced though highly unusual rhachis. (?) Rhachisphora ardisiae (Takahashi) comb. n.Dialeurodes ardisiae Takahashi, 1935: 50, fig. 35. Syntype pupal cases, TAIWAN (TARI). This record is based upon a single specimen which has not been compared with the type-material; accordingly, the record should be regarded as tentative, but from the description it isclear that ardisiae should be placed in Rhachisphora, rather than in Dialeurodes. DISTRIBUTION. (Papua New Guinea), Taiwan. MATERIAL EXAMINEDPapua New Guinea: Buso, on undetermined host (BMNH). TETRALEURODES Cockerell Aleyrodes (Tetraleurodes) Cockerell, 1902: 283. Type-species: Aleyrodes perileuca Cockerell, by original designation.Tetraleurodes Cockerell; Quaintance & Baker, 1914: 107 [raised to genus]. Fifty-seven species of whitefly are currently placed in Tetraleurodes, which is a difficult genus todefine. Those from New Guinea that are assigned to the Tetraleurodes-group all have darkbrown to black pupal cases, lack thoracic and caudal tracheal differentiation, and have thesubmarginal and subdorsal regions separated by a suture-like fold.The 9 species from New Guinea are listed with host data on p. 308. TRIALEURODES Cockerell Aleyrodes (Trialeurodes) Cockerell, 1902: 283. Type-species: Aleurodes pergandei Quaintance, by original designation.Trialeurodes Cockerell; Quaitance & Baker, 1915: xi [raised to genus]. Trialeurodes vaporariorum (Westwood)(Fig. 29) Aleyrodes vaporariorum Westwood, 1856: 852. Syntype pupal cases, adults, GREAT BRITAIN: England (Westwood collection, thought to be part of type series, BMNH; UMO) [examined].Trialeurodes vaporariorum (Westwood) Quaintance & Baker, 1914: 105. DISTRIBUTION. Papua New Guinea; also very widely distributed throughout the world, althoughOriental, Austro-Oriental and Australasian records are rather sparse.MATERIAL EXAMINED Papua New Guinea: Goroka, on 'squash' (Cucurbitaceae) (BMNH; BPBM); Aiyura, on Solanumlycopersicon (Solanaceae) (BMNH); Chimbu, on English potato (Solanaceae) (BMNH). Great Britain:no data, part of Westwood series (BMNH). 334 J. H. MARTIN XENALEYRODES TakahashiXenaleyrodes Takahashi, 1936: 113. Type-species: Xenaleyrodes artocarpi Takahashi, by monotypy. Takahashi (1936) erected Xenaleyrodes for artocarpi from the Palau Islands (Caroline group).The genus is distinguished from Aleurocanthus by the characteristic tubiform submarginalspines which are much thicker in the basal two-thirds, by the absence of similar spines elsewhereon the pupal case and by the deflexed margin (see generic key, p. 309). From a study of theoriginal description, and of material in the BMNH, it is clear that Neomaskellia eucalyptiDumbleton (1956) from Australia should also be placed in Xenaleyrodes (comb. n.).A key to the pupal cases of the five species currently placed in Xenaleyrodes is given below. Key to species of Xenaleyrodes Pupal cases 1 Submarginal tubiform spines distinctly curved in apical third, often abruptly angled through 90 (Fig. 35). Comb of differentiated teeth present at position of thoracic tracheal openings near margin (Fig. 36) 2 Submarginal tubiform spines much narrowed in apical third , but generally straight (Figs 3 1 , 33) .No differentiated teeth in thoracic tracheal area.Posteriorly with a single pair of very long, stout setae 0-3-0-4 mm long 4 2 Second cephalothoracic pair of submarginal tubiform spines set much closer to 1st pair than to 3rd pair. Thoracic tracheal combs normally with about 8 teeth 3 All cephalothoracic submarginal tubiform spines evenly spaced. Thoracic tracheal combsnormally with 4 teeth timonii sp. n. (p. 336) 3 Dorsal cuticle rather rough, reticulate-granular; posterior margin of vasiform orifice produced into a pointed process with a median notch; all abdominal submarginal tubiform spines evenly spaced artocarpi Takahashi (p. 334) - Dorsal cuticle smooth; posterior margin of vasiform orifice smoothly rounded, not producedinto a pointed process; posterior two pairs of abdominal submarginal tubiform spines setcloser together than remaining 4 pairs. (Australia, Victoria) eucalypti (Dumbleton) 4 Submargin with single row of about 30 pairs of shorter (90 ju,m) tubiform spines, interspersed with about 9 pairs of fine hairs up to 0-25 mm long (Fig. 30) broughae sp. n. (p. 334) Submargin with single row of 11 pairs of longer (up to 0-12 mm) tubiform spines, interspersedwith 7 pairs of short, lanciform, spines 25-40 /A long (Fig. 33) irianicus sp. n. (p. 335) Xenaleyrodes artocarpi Takahashi Xenaleyrodes artocarpi Takahashi, 1936: 113, fig. 2. Syntype pupal cases, PALAU ISLANDS (TARI).DISTRIBUTION. Palau Islands (Caroline group), Papua New Guinea. MATERIAL EXAMINEDPapua New Guinea: Buso, on Premna sp. (Verbenaceae) and IDecaspermum sp. (Myrtaceae) (BMNH). Xenaleyrodes broughae sp. n. (Figs 30-32) PUPAL CASE. Jet black, opaque, with an agglomeration of white wax (see comments). Evenly oval,1-40-1-50 mm long, 1-00-1-10 mm wide, widest at abdominal segments I & II. Cases rather deep, withmargin mostly deflexed under dorsum in slide-mounted specimens: apparent margin is thus merely part ofsubmarginal dorsum and is smooth. Region of true margin complex: true margin serrate-toothed, withventer having another 'false margin' of rounded crenulations of about 3 times width of true marginal teeth(Fig. 32). Dorsum. Dorsal surface almost smooth, and in evenly bleached specimens only slight mottling evident,even median abdominal segmentation little marked. Immediately mesad of (dorsal to) true margin is a lineof small tubercles which appear paler than rest of cuticle; inset a little further is a ring of about 30 pairs ofshort, stout, tubiform spines with expanded apices, interspersed with about 9 pairs of very long fine hairs,including caudal pair (Fig. 30). Tubiform spines about 90 /xm long, with basal two-thirds markedly swollen;fine submarginal hairs up to 0-25 mm long, the ring of spines and hairs appearing marginal owing todeflexing of true margin. Adjacent to caudal hairs, with bases a little further inset than submarginal ring of THE WHITEFLY OF NEW GUINEA 335 tubiform spines, is a pair of remarkably stout setae, up to 0-4 mm long and much darker than duskytubiform spines: it seems likely that these are modified tubiform spines. Single pairs of cephalic, first andeighth abdominal setae also present, as detailed in Fig. 30. Thoracic and abdominal tracheal openings notdiscernible. Small eyespots present, best seen in untreated specimens. Transverse moulting suturesreaching apparent margin, joined distally to longitudinal suture by a variably defined cephalothoracic foldrunning concentric to margin. Each thoracic segment with a single pair of subcircular depressions in cuticle.Vasiform orifice elevated, oval, with operculum occupying whole area of orifice and obscuring lingula;orifice inset from apparent margin by a little more than its own length; eighth abdominal setae arising fromlateral extensions of vasiform orifice elevation. Venter. Thoracic and caudal tracheal folds not marked. Entire venter smooth, not stippled or otherwisepunctuated. Usual ventral spiracles, setae, legs and antennae present. Holotype pupal case, Papua New Guinea: Southern Highlands Province, Eraue, on Citrus sp.(Rutaceae), 21.vii.1983 (E. J. Brough E630) (BMNH). Paratype pupal cases. Papua New Guinea: 40, same data as holotype (BMNH; USNM). Spirit-storedmaterial with same data as holotype (BMNH). COMMENTS. Known from a single collection from the Southern Highlands Province of PapuaNew Guinea, where it occurred as a densely populated colony on a few leaves of a mature orangetree growing in a coffee nursery. Each pupa has a thin plate of wax covering the median andsubdorsal areas, the wax thickening into dense white curls which extend from the submarginalarea to well beyond the edge of the pupa, obscuring the ring of tubiform spines and hairs. Thepupae are apparently attached to the leaf by the part of the venter inside the crenulate 'falsemargin', with a ring of white wax separating the outer ring of the venter from the leaf lamella.The colony was not apparently attended by ants. The pupal cases were exceptionally resistant to bleaching, and could only be bleachedsuccessfully by using a mixture of 880-volume ammonia and approximately 20-volume hydrogenperoxide. With this reagent, bleaching to an acceptable level occurs in only a few minutes,without use of heat. X. broughae can be distinguished from the other described species of Xenaleyrodes by thecharacters given in the key. Xenaleyrodes irianicus sp. n. (Fig. 33) PUPAL CASE. Jet black, opaque, in groups under leaves of host. Oval, up to 1-05 mm long, 0-85 mm wide,widest at abdominal segment I. Cases deep, margin deflexed, obscured by dorsum in slide-mountedspecimens. True margin crenulate, with about 12 teeth to 0-1 mm of margin; a row of tubercles of similarsize immediately mesad of true margin gives margin the appearance of being double in some prepar-ations. Dorsum. Smooth, unicolourous to slightly mottled in slide-mounted bleached specimens; abdominaland thoracic segmentation only faintly marked. Submarginal (apparently marginal, owing to deflexion oftrue margin) ring of about 11 pairs of tubiform spines up to 0-12 mm long, interspersed with 6 or 7 pairs oflanceolate spines about 25-40 /im long, distributed as shown in Fig. 33. Caudally a pair of very long, stoutspines (probably modified tubiform spines) up to 0-3 mm long, and a pair of fine caudal hairs about 0-15 mmlong. Single pairs of cephalic, first and eighth abdominal setae present, as detailed in figure. Thoracic andabdominal tracheal openings not discernible. Eyespots present. Transverse moulting sutures reachingapparent margin, longitudinal suture not so. Vasiform orifice elevated, oval, with operculum completelyobscuring lingula. Eighth abdominal setae arising from lateral extensions of vasiform orifice elevation.Whole of caudal and vasiform orifice area strikingly similar to that of A', broughae. Venter. Ventral characters entirely as in X. broughae, with the exception of characteristics of true margin. Holotype pupal case, Irian Jaya: Biak, on undetermined tree, 23. v. 1959 (T. C. Mad) (BPBM). Paratype pupal cases. 14, same data as holotype (BMNH; BPBM); dry material on leaf, same data asholotype (BPBM). COMMENTS. Each pupa of X. irianicus is surrounded and covered by a secretion which, in driedspecimens at least, is translucent and glassy, not the more usual white wax. Exuviae of the earlierinstars do not appear to remain attached to the dorsum of the pupa.X. irianicus is similar to broughae in having the submarginal ring of tubiform spines inter- 336 J. H. MARTIN spersed with a smaller number of simple hairs/spines. However, the tubiform spines are fewerand longer, and the simple spines are fewer and shorter in irianicus. A further species, Xenaleyrod.es sp. 1 (p. 308), is represented by three damaged pupal casesfrom an unidentified host from near Lake Trist, Kuper Range, Papua New Guinea -this appearsto be most similar to irianicus, but the submarginal tubiform spines do not appear to beinterspersed with lanceolate spines, and the posteriormost pair are not as long and stout as inirianicus, although longer than the remainder. Xenaleyrodes timonii sp. n. (Figs 34-37) PUPAL CASE. Jet black, opaque, with little waxy secretion evident. Pupal cases oval, up to 0-95 mm long,0-65 mm wide, widest at thoraco-abdominal suture. Cases rather deep, with submargin deflexed, foldedunder remainder of dorsum in slide-mounted specimens; the apparent margin is thus merely a part of thesubmarginal dorsum, and is smooth to irregular. True margin, at junction of dorsum and venter, smooth:apparent crenulate to serrate appearance in most specimens is due to an adjacent line of tiny tubercles (Fig.36), precise appearance dependent upon angle of tubercles to viewing axis. Dorsum. Cuticle almost completely smooth, punctuated only by one row of tiny tubercles just mesad of(i.e. vertically above, in unmounted specimens) true margin, and by disc porettes, a row just mesad ofsubmarginal tubiform spines, a few subdorsally, and also scattered between tubiform spines and truemargin. Submargin with single row of 12 pairs of evenly spaced, very prominent tubiform spines (Figs34-37); tubiform spines of specimens in situ on leaf with basal half to two-thirds (about 70 jim) horizontal,the apical part much narrower and angled downwards (see comments) through 90, apices expanded,laciniate. Single pairs of cephalic and first and eighth abdominal setae present; first abdominal pair rathervariable, with the extremes shown in Fig. 34. Caudal setae very long, fine, longer than other dorsal setae,sometimes as long as 0-2 mm. Eye spots not marked. Thoracic and abdominal tracheal furrows not marked,but combs distinct; thoracic combs (Fig. 36) normally with 4 rounded teeth, occasionally with 3 or 5, setdistinctly mesad of (above) true margin, each tooth about half as wide as base of a submarginal tubiformspine; caudal comb similar, with about 6 teeth. Longitudinal and transverse moulting sutures not reachingtrue margin; transverse sutures reaching apparent margin, transverse and longitudinal sutures joineddistally by a cephalothoracic suture. Vasiform orifice (Fig. 37) oval, elevated, produced posteriorly into ablunt process; orifice toothed posterolaterally on internal walls, a little longer than wide, inset fromposterior margin of case by about half its own length, about 65-80 /AHI long, 60-70 /im wide, about 30-50fj,m from margin. Width of dorsal opening of orifice dependent upon degree of flattening in slide-mountedspecimens, but evidently less than maximum internal width of orifice (Fig. 37), and often measuring lessthan width of operculum. Operculum rather small, trapezoidal, occupying only about half of internalcross-sectional area of vasiform orifice. Lingula short, finger-shaped, finely spinulose, exposed butincluded within vasiform orifice. Venter. Thoracic and caudal tracheal folds not marked. Most of venter densely stippled, stipples mostmarked in submargin, only absent from median area of thorax. Abdominal segmentation evident. Minuteconical seta present at base of each middle and hind leg. A group of 5-7 short, 2- or 3-pointed anchor spinespresent near anterior edge of venter, presumably to aid adhesion to leaf. Holotype pupal case, Papua New Guinea: Morobe Province coast, Buso, on Timonius sp. (Rubiaceae),10.ix.1979 (/. H. Martin 2527) (BMNH). Paratype pupal cases. Papua New Guinea: (all from same general locality and same host species) 26,same data as holotype; 19, riverside, 13.ix.1979 (JHM 2552, 2556); 28, riverside, 13.X.1979 (JHM 2691,2696); 5, beach-top, 12.X.1979 (/HM2687A); 25, beach-top, 6.ix.l979 (JHM25QQ) (BMNH; USNM). COMMENTS. Known from seven samples, always from leaves of a species of Timonius growing ongravel bars between meanders of the fast-flowing Buso river, or in beach-top sand. The pupaeoccur in dense groups and are not attended by ants. The larval exuviae often remain attached tothe dorsa of later stages, sometimes with all stages present in a stack. The species was neverencountered on other hosts, and the name timonii appears appropriate. No adults were seen. X. timonii may be distinguished from other Xenaleyrodes species by characters given in thekey. In Takahashi's original description of X. artocarpi (1936), the submarginal tubular spines aredescribed as being 'eminently curved upward on the distal part'. 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Transactions and Proceedings of the New Zealand Institute 28: 411-449.Mound, L. A. 1963. Host-correlated variation in Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae). Proceedings of the Royal Entomological Society of London (A) 38: 171-180.1965. An introduction to the Aleyrodidae of western Africa. Bulletin of the British Museum (Natural History) (Entomology) 17: 113-160. 1966. A revision of the British Aleyrodidae. Bulletin of the British Museum (Natural History) (Entomology) 17: 397-428. 338 J. H. MARTIN Mound, L. A. & Halsey, S. H. 1978. Whitefly of the World. 340 pp. Chichester. Peal, H. W. 1903. Contribution towards a monograph of the oriental Aleurodidae. Journal of the Asiatic Society of Bengal 72: 61-98.Priesner, H. & Hosny, M. 1934. Contributions to a knowledge of the whiteflies (Aleurodidae) of Egypt (III). Bulletin. Ministry of Agriculture, Egypt. Technical and Scientific Service 145: 1-11.Quaintance, A. L. 1903. New oriental Aleurodidae. Canadian Entomologist 35: 61-64. - 1908. Homoptera, Family Aleyrodidae. In Wytsman, P., Genera Insectorum 87: 1-11.Quaintance, A. L. & Baker, A. C. 1913. Classification of the Aleyrodidae Part I. Technical Series. Bureau of Entomology, United States Department of Agriculture 27: 1-93. 1914. Classification of the Aleyrodidae Part II. Technical Series. Bureau of Entomology, United States Department of Agriculture 27: 95-109. 1915. Classification of the Aleyrodidae - Contents and Index. Technical Series. Bureau of Ento- mology, United States Department of Agriculture 27: i-xi, 111-114. 1917. A contribution to our knowledge of the whiteflies of the subfamily Aleurodinae. Proceedings of the United States National Museum 51: 335-445.Russell, L. M. 1943. A new genus and four new species of whiteflies from the West Indies (Homoptera,Aleyrodidae). Proceedings of the Entomological Society of Washington 45: 131-141. - 1945. A new genus and twelve new species of Neotropical whiteflies. Journal of the WashingtonAcademy of Sciences 35: 55-65. - 1958. Orchamoplatus, an Australasian genus (Homoptera: Aleyrodidae). Proceedings of theHawaiian Entomological Society 16: 389-410. - 1965. A new species of Aleurodicus Douglas and two close relatives (Homoptera: Aleyrodidae).Florida Entomologist 48: 47-55. Sampson, W. W. & Drews, E. A. 1956. Keys to the genera of the Aleyrodidae and notes on certain genera. Annals and Magazine of Natural History (12) 9: 689-697.Signoret, V. 1868. Essai monographique sur les aleurodes. Annales de la Societe Entomologique de France (4) 8: 369-402.Singh, K. 1931. A contribution towards our knowledge of the Aleyrodidae (whitefly) of India. Memoirs of the Department of Agriculture in India 12 (1): 1-98.Stal, C. 1876. Observations Orthopterologiques 2. Les genres des Acridoidees de la faune europeenne. Bihang till Kongl. Svenska Vetenskaps Akademiens Handlingar, Stockholm 4 (5): 30.Takahashi, R. 1931. Some Formosan whiteflies. Journal of the Society of Tropical Agriculture, Taiwan 3: 218-223. - 1932. Aleyrodidae of Formosa Part I. Report. Department of Agriculture. Government ResearchInstitute, Formosa 59: 1-57. - 1934. Aleyrodidae of Formosa Part III. Report. Department of Agriculture. Government ResearchInstitute, Formosa 63: 39-71. - 1935. Aleyrodidae of Formosa Part IV. Report. Department of Agriculture. Government ResearchInstitute, Formosa 66: 39-65. 1936. Some Aleyrodidae, Aphididae, Coccidae and Thysanoptera from Micronesia. Tenthredo. ActaEntomologica 1: 109-120. - 1942. Some foreign Aleyrodidae IX. Species from Thailand and French Indo-China. Transactions ofthe Natural History Society of Formosa 32: 327-335. - 1951. Descriptions of six interesting species of Aleyrodidae from Malaya. Kontyu 19: 1-8. - 1952. Aleurotuberculatus and Parabemisia of Japan (Aleyrodidae, Homoptera). MiscellaneousReports of the Research Institute for Natural Resources, Tokyo 25: 17-24. Thomas, R. T. S. 1962. Checklist of pests on some crops in West Irian. Mededelingen van Economische Zaken, LandbouwkundigeSerieno. 1: 99.Verdcourt, B. 1979. A manual of New Guinea legumes. Botany Bulletin no. 11, Office of Forests, Division of Botany, Lae, Papua New Guinea.Westwood, J. 0. 1856. The new Aleyrodes of the greenhouse. Gardeners' Chronicle 1856: 852. THE WHITEFLY OF NEW GUINEA 339 Figs 1-5 Aleurocanthus species. 1,2, A. luteus: (1) pupal case; (2) posterior dorsal detail of pupal case.3-5, A. papuanus: (3) pupal case; (4) posterior dorsal detail of pupal case; (5) vasiform orifice detail withoperculum removed for clarity. 340 J. H. MARTIN 10 Figs 6-10 6 , 7 , A leuromarginatus corbettiaformis : (6) pupal case ; (7) posterior dorsal detail of pupal case .8, 9, Aleuromarginatus littoralis: (8) pupal case, with inset showing fine detail of dorsal cuticle; (9)vasiform orifice. 10, Bemisia tabaci, pupal case. THE WHITEFLY OF NEW GUINEA 341 lla 13 lib lie 12 Figs 11-14 11, Aleuromarginatus species, posterior dorsal detail of: (Ha) A. littoralis; (lib) A.kallarensis; (lie) A. dalbergiae - transposition of abdominal hairs arrowed. 12-14, Crenidorswnlasangensis: (12) pupal case; (13) dorsal submarginal detail of thoracic region; (14) posterior dorsaldetail of pupal case. 342 J. H. MARTIN 16 17 Figs 15-19 15, 16, Crenidorsum morobensis: (15) pupal case; (16) posterior dorsal detail of pupal case.17-19, Dialeurodes decaspermi: (17) pupal case, with detail of pigmentation; (18) vasiform orifice andcaudal furrow; (19) detail of vasiform orifice, operculum folded upwards. THE WHITEFLY OF NEW GUINEA 343 20 21 Figs 20, 21 Indoaleyrodes pseudoculatus: 20, pupal case, extremity of longitudinal moulting suturearrowed; 21, vasiform orifice and one lateral dorsal abdominal glandular patch. 344 J. H. MARTIN 25 Figs 22-25 22, 23, Orchamoplatus niuginii: (22) pupal case; (23) posterior dorsal detail of pupal case,operculum removed. 24, 25, Parabemisia jawani: (24) pupal case; (25) posterior dorsal detail of pupal case. THE WHITEFLY OF NEW GUINEA 345 27 28 Figs 26-29 26, 27, Parabemisia myrmecophila: (26) pupal case, with detail of pigmentation; (27)posterior dorsal detail of pupal case. 28, stylised vasiform orifice typical of the Pealius genus-group. 29,Trialeurodes vaporariorum, pupal case, with enlargement of vasiform orifice. 346 J. H. MARTIN 30 igs 30-32 Xenaleyrodes broughae: 30, pupal case; 31, dorsal submarginal detail and true margin; 32,true margin, ventral 'false margin', and dorsal submarginal detail. THE WHITEFLY OF NEW GUINEA 347 35 Figs 33-37 Xenaleyrodes species. 33, X. irianicus, pupal case. 34-37, X. timonii: (34) pupal case, dorsaldetail; (35) submarginal tubiform spine; (36) detail of margin at thoracic tracheal area; (37) vasiformorifice, with stylised transverse section. J. H. MARTIN 40 Figs 38-41 38, Neomaskellia bergii, pupal case ex Saccharum, New Guinea. 39, Aleurocybotus setiferus,pupal case ex Imperata, New Guinea; dotted line indicates border of submedian pigmentation. 40,Dialeuropora decempuncta, pupal case ex Colocasia, New Guinea. 41,Aleurocanthus cocois, pattern ofdorsal setae and stout spines. THE WHITEFLY OF NEW GUINEA 349 45 Figs 42-47 (after Corbett, 1935a) 42, Aleurotuberculatus neolitseae, pupal case. 43, Dialeurodes psidii,pupal case. 44, Aleurocanthus woglumi, pupal case. 45, Aleurocanthus pendleburyi, pupal case withmarginal detail. 46, Aleurolobus selangorensis, pupal case with vasiform orifice detail. 47, Aleurodicusdestructor, pupal case. 350 J. H. MARTIN / Subdorsal papill: Anterior marginal seta._ Cephalothoracic fold/suture. Longitudinal moulting suture Rostral base seta Ant enna . / i Submarginal p laciniate spinSubmarginal ,tubiform spine Fig. 48 Annotated schematic diagram of whitefly pupal case. Addendum While this manuscript was in press four samples of Orchamoplatus mammaeferus (Quaintance & Baker,1917: 400), collected in New Guinea, were received. O. mammaeferus is similar to niuginii, and thedifferences between the two species are discussed on p. 330. MATERIAL EXAMINED Papua New Guinea: near Laiagam and near Wabag, on Phaseolus vulgaris (Leguminosae) ; Wau, onCroton sp. (Euphorbiaceae); Mt Hagen, on sweet potato (Convolvulaceae) (all BMNH).This brings the total of named species (p. 304) to 38. THE WHITEFLY OF NEW GUINEA Index 351 Principal page references are in bold; page references to figures are in italics; invalid names are in italics. aceris 330, 331afer 306, 321-322Aleurocanthus 303, 304, 309, 313-17,334,339,348,349Aleurocybotus 305, 310, 317, 348Aleurodes 312, 316, 317, 320, 321, 322, 329, 333 Aleurodicinae 304, 309, 311-12Aleurodicus 303, 304, 309, 311-12, 349 Aleurolobus 305, 310, 317, 349Aleuromarginatus 305, 310, 311, 318-320,340,347 Aleuroplatus 305-311, 320, 323, 329Aleurotrachelus 305, 310, 320, 323Aleurotuberculatus 305, 311, 320-321,349 Aleurotulus 306, 310, 321, 323, 324Aleyrodes 322, 325, 326, 332, 333Aleyrodinae 304, 309, 313-337anonae311antidesmaeSllardisiae 308, 333artocarpi 308, 334, 336arundinacea 306, 310, 321Asialeyrodes 306, 310, 321 bauhiniae 320 Bemisia 303, 304, 306, 309, 321-323,330,340bergii 303, 307, 329, 345bossi 320brevispinosus 315brideliae 326broughae 308, 334-335, 346 calophylli 330canangae313, 314cinnamomi 312citri Ashmead 325citri Riley & Howard 325cocois 304, 313-314, 315, 316, 348comata 329 corbettiaformis 305, 318-319, 340Crenidorsum 306, 310, 323-325,347,342 dalbergiae 320, 347decaspermi 306, 325, 342decempuncta 307, 326-327, 348destructor 303, 304, 311, 312,349Dialeurodes 306, 311, 325-326, 327, 328,333,342,349Dialeurodoides 321Dialeuropora 307, 310, 326-327, 348differens 324 dispersus311dothioensis 327dryandrae 320dumbletoni 329 esakii 304, 313, 314, 316eucalypti 334 gordoniae 320graminicola 317 hancocki 321-322hirsutus 315holmesii304,311,312Husaini3l6 inconspicua 321 Indoaleyrodes 307, 310, 327-329, 331,343 irianicus 308, 334, 335-336, 347ixorae 325 jawani 307, 331-332, 344Jeannelaleyrodes 329 kallarensis 320, 347kirkaldyi 306, 326 laos 327, 328 lasangensis 306, 323-324, 347 leakii 306, 322 littoralis 305, 311, 318, 319-320, 340, 347 lumpurensis, Asialeyrodes 321lumpurensis, Dialeurodes 326luteus 305, 313, 314-315, 339 machili 312 mackenziei 315 maculata 330, 331 malayensis 312 mammaeferus 329, 330, 350 marginale 323, 324 marlatti317 maskelli 332 melastomae 320 millettiae 320 montanus 330 morobensis 306, 324-325, 342 myricae 330, 331 myrmecophila 307, 331, 332, 345 Neoaleurotrachelus 329neolitseae 306, 320-321, 349Neomaskellia 303, 307, 309, 329, 334,348 nephrolepidis 321niloticus305,317 niuginii 307, 329-330, 344 Orchamoplatus 307, 309, 329-330, 344, 350Orchamus 329 papuanus 305, 313, 315-316, 339Parabemisia 307, 311, 328, 330-332, 344, 345 Pealius308,311,332,345pendleburyi 305, 313, 314, 316, 349perdentatus 329pergandei 333perileuca 333perseae 326, 327pongamiae 306, 322pseudoculatus 307, 328, 343psidii 306, 326, 349pustulatus 327 quercusaquaticae 320 Rabdostigma 325regis313,315,316reticulata 328, 330, 331Rhachisphora 308, 310, 333rugosa 315 selangorensis 305, 317, 349setiferus 305, 310, 317, 348setigerus 327siamensis 320spinifera 316spiniferus305,313,316spinithorax 315strychnosicola 315subrotunda 325sudaniensis 329 tabaci 303, 304, 306, 322-323, 340 tephrosiae 318 Tetraleurodes 308, 309, 333 timonii 308, 334, 336-337, 347 tracheifer 320 Trialeurodes 303, 308, 310, 333, 345 trilobitoides 333 trispina315 T-signatus 315 tuberculatum 323 vaporariorum 303, 308, 333, 345woglumi 305, 313, 316-317, 349 Xenaleyrodes 308, 309, 334-337,346, 347 zizyphi 315 British Museum (Natural History) Catalogue of the Diptera of the Afrotropical Region Editor: R. W. Crosskey. Assistant editors: B. H. Cogan, P. Freeman, A. C. Pont, K. G. V. Smith and the late H. Oldroyd. British Museum (Natural History). 1980, 1437 pp. 55.00 The Diptera or two-winged flies are probably the most important insects that affect man.Although most flies are harmless, some have become transmitters of dangerous diseases toman and his domestic animals, and others are important pests of agricultural crops. Some fliesare beneficial because they destroy large numbers of plant-feeding insects through theirparasitic or predacious habits. Nowhere is their socio-economic and medical impact more sharply felt than in tropical Africa,where fly-borne diseases are not only a direct health hazard but can prevent or hinderdevelopment of the land. The control of such diseases as sleeping sickness and onchocerciasisdepends in great measure upon controlling the flies that carry them. This in turn requires athorough appreciation of all that is known about the insect vectors, including their basictaxonomy, so that they can be correctly identified and their geographical ranges accuratelyestablished. This catalogue synthesizes the scattered basic taxonomic work on the Diptera of tropicalAfrica and its islands by listing the known 16,500 species with their synonyms and knowngeographical ranges within a comprehensive classification. A short introduction is given toeach family and a bibliography of 4,700 titles provides references to the primary literature.Such a task has never before been attempted for the region and its completion should greatlystimulate taxonomic research. The Catalogue represents ten years' careful work by a team offorty specialists, under the editorship of six dipterists on the staff of the Natural HistoryMuseum, themselves contributors with considerable expertise in the African fauna. The Catalogue should serve for a long time as an indispensable tool to the taxonomist and anessential source-work to anyone concerned with African flies in the fields of medical,agricultural and veterinary science. Titles to be published in Volume 50 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera) By Peter S. Broomfield Nymphal taxonomy and systematics of the Psylloidea (Homoptera) By I. M. White & I. D. Hodkinson The whitefly of New Guinea (Homoptera: Aleyrodidae) By J.H.Martin Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, SuffolkPrinted in Great Britain by Henry Ling Ltd, Dorchester