Bulletin of the British Museum (Natural History). British Museum (Natural History) Converted as part of the ABLE project by Dauvit King London : BM(NH) Set of bird parts only held at TOS 112. General Library missing Vol 55-56, 1989-90 Vol.1 (1950)- 4 5 6 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.) '/ -> 1 I . BULLETIN OF /& j G£tt£RAL THE BRITISH MUSEUM^ i6juNI976(NATURAL HISTORY) ZOOLOGY Vol. 27 1974 BRITISH MUSEUM (NATURAL HISTORY)LONDON: 1976 DATES OF PUBLICATION OF THE PARTS No. i . . . . . .13 June 1974 No. 2 -"i . . . . .18 July 1974 No. 3 19 July 1974 No. 4 . . . . .20 September 1974 No. 5 . . . . .23 December 1974 No. 6 . . . . .23 December 1974 No. 7 . . . . .24 December 1974 ISSN 0007-1498 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU CONTENTS ZOOLOGY VOLUME 27 PAGE No. I. Anatomy of head and neck in the Huia (Heteralocha acutirostris)with comparative notes on other Callaeidae. By P. J. K. BURTON(i PI.) . . i No. 2. Miscellanea .......... 49 A new skink (Reptilia : Scincidae : genus Emoia) from the NewHebrides. By LORD MEDWAY ...... 53 Scutocyamus parvus, a new genus and species of whale-louse(Amphipoda : Cyamidae) ectoparasitic on the North Atlanticwhite-beaked dolphin. By R. J. LINCOLN and D. E. HURLEY (i PL) . • . • 59 Catalogue of the whale-lice (Crustacea : Amphipoda : Cyamidae)in the collections of the British Museum (Natural History). By R. J. LINCOLN and D. E. HURLEY 65 A review of Laephotis Thomas, 1901 (Chiroptera : Vespertilioni- dae). By J. E. HILL 73 The genus Finmarchinella Swain, 1963 (Crustacea : Ostracoda)and its species. By J. W. NEALE (2 Pis) .... 83 Euplotes rariseta sp. n. (Protozoa: Ciliatea). A new smallmarine hypotrich. By C. R. CURDS, B. J. WEST and J. E. DORAHY (i PL) 95 Observations on clonal cultures of Eugfypha acanthophora andEuglypha strigosa (Testacea: Protozoa). By R. H. HEDLEY, C. G. OGDEN and J. I. KRAFFT (2 Pis) 103 Descriptions of three species of Euplotes (Protozoa : Ciliatea). By C. R. CURDS (i PL) 113 New records of bats from Southeastern Asia, with taxonomic notes. By J. E. HILL 127 No. 3. The Haplochromis species (Pisces : Cichlidae) of Lake Rudolf, East Africa. By P. H. GREENWOOD 139 No. 4. A review of Scotoecus Thomas, 1901 (Chiroptera : Vespertilionidae). By J. E. HILL 167 No. 5. Catalogue of the types of terrestrial isopods (Oniscoidea) in thecollections of the British Museum (Natural History) I. SuperfamilyPseudotracheata. By R. J. LINCOLN and J. P. ELLIS . . 189 No. 6. Campylaspis species (Crustacea : Cumacea) from the deep Atlantic. By N. S. JONES -247 No. 7. A new family, genus and species of bat (Mammalia : Chiroptera) from Thailand. By J. E. HILL 301 Index to Volume 27. . . . . . • • • 337 INDEX TO VOLUME 27The page numbers of the principal references and the new taxonomic names are printed in bold type. abbreviatus, Cyamusabnormis, Gelsana .abyssinicus, Microcercus .Acaeroplastes .Acanthoniscusacanthophora, Euglypha .ActoeciaActoeciidae aculeata, Campylaspisacuminatus, Rhinolophus. . 65 . 214. 215 • 193. 219 . 103-112, PI. I . 218 . 191, 218250, 253, 268-270 . 131 acuminatus, Rhinolophus acuminatus 131-132, 137acuminatus acuminatus, Rhinolophus 131-132, 137 acuminatus audax, Rhinolophus . . 132acuminatus calypso, Rhinolophus . -132acuminatus circe, Rhinolophus . . .132acuminatus sumatranus, Rhinolophus . 132acustiserra, Porcellio .... 197 acutirostris, Heteralocha . . 1-48, i PI. Adinda ....... 244 aediculatus, Euplotes . . . .124 aegaeum, Armadillidium .... 208 aegaeus, Porcellio . . . . . 197 aenigma, Diploexochus .... 223 aerarius, Diploexochus .... 224 affinis, Armadillo . . . . .219 affinis, Campylaspis. . 250, 254, 263, 277affinis, Cubaris . . . . .219 affinis, Euplotes 113, 114, 116, 117-119, 120, 123affinis tricirratus, Euplotes . 113, 119, 123Agabiforius . . . . . -237 Agabiformius ...... 193 Agnara ....... 205 Akermania . . . . . .219 alassiense, Armadillidium . . 208, 212 alatus, Euplotes .... 97, 101 Alauda . . . . . . -41 alba, Campylaspis . . 250, 251, 256, 258albanicum, Armadillidium . . . 208 alberti, Diploexochus . . . .224 albertianus, Haplochromis . . 150, 159albescens, Diploexochus .... 224 albicornis, Lucasius .... 191, 197 albicornis, Mesarmadillo . . . -215albicornis, Porcellio . . . . 197 albifrons, Armadillidium .... 208 albigula, Scotoecus . . 180, 181, 182, 183albigula, Scotoecus hindei 180, 181, 182-i83albinus, Porcellio . . . . .198 albipennis, Tura ..... 204 albirostris, Lagenorhynchus . . 59, 71 albofuscus, Scotoecus 169, 171, 172, 174-175, 176, 177, 186 albofuscus, Scotoecus albofuscus . 175-176albofuscus, Scotophilus . . . 169, 175albofuscus albofuscus, Scotoecus . 175-176albofuscus cinnamomeus, Scotoecus . . 175albofuscus woodi, Scotoecus . 175, 176-177albomaculatus, Buddelundia . . .222albomarginatus, Armadillo . . .219albomarginatus, Porcellio . . . 198 albospinosus, Armadillo . . . .219 album, Armadillidium .... 208 alloniscus ...... 205 alluaudi, Porcellio . . . . .198 alticola, Diploexochus . . . .224 altimontis, Saidjahus .... 229 alveolata, Euglypha .... 104 ambitiosus, Armadillo . . . .229 ambitiosus, Spherillo .... 229 amblyoda, Campylaspis . . . 250, 251Amblyrhamphus . . . . -37 americanum, Ethelum . . . -213americanus, Mesarmadillo . . .213 amieti, Euplotes . . . . .124 amoenus, Porcellio ..... 198 Amphipoda . . . 59-64, I PI.; 65-72ampullatus, Hyperoodon . . . -71Anchicubaris . . . . . .219 anconanum, Armadillidium . . . 208anconanus, Metoponorthus pruinosus . 197 aneityumensis, Emoia . . . 53-55, 56Angara ....... 193 angolensis, Laephotis 73, 76-77, 78- 79, 80, 81angolensis, Laephotis wintoni . . -73angularis, Campylaspis . . . 250, 251angulata, Barentsovia . . 86, 87, 88-89, 91, 92, Pis. 1-2angulata, Cythere .... 87, 88 angulata, Cythereis . . .83, 88 angulata, Finmarchinella . . 86, 87, 88-89, 90, 91, 92, Pis. 1-2 angulata, Hemicythere . . 86, 87, 88, 89angulata, "Hemicytherinae" . . .88angulata, Nereina . . 86, 87, 88, 90, 91angusta, Tura . .204 angustulus, Porcellio .... 198 anna, Ciridops . • 25, 28 annectans, Myotis . . • 3°3 annulicornis, Cylisticus .... 192 antarctica, Campylaspis . . 250, 256, 299Anthochaera . • 34 apenninorum, Armadillidium . . . 208 338 INDEX apenninorum, Trachelipus . . . 207 apenninorum, Tracheoniscus . . . 207aperta, Campylaspis . . . 250, 253 apfelbecki, Armadillidium . . . 208 approximates, Metoponorthus . . .196apulicus, Trachelipus .... 207 apulicus, Tracheoniscus .... 207 Arachnothera . . . . . .42 arcuata, Campylaspis . 250, 253, 265-266arcuatus, Porcellio ..... 202 argentarium, Armadillidium . . . 208argentarius, Acaeroplastes . . . 193argentarius, Metoponorthus . . 193 Armadillidae . . 197, 219-232, 244-245 Armadillidiidae . 191, 208-213, 240-242 Armadillidium 208-212, 213, 221, 240, 241, 242Armadillo . 208, 219-221, 222, 224, 225, 226, 227, 228, 229, 230, 231, 232, 244, 245armatus, Benechinus . . . .213 artinii, Scotoecus . . . 180, 181, 182artinii, Scotoecus hindei . . . .182 asifensis, Metoponorthus .... 197 asifensis, Metoponorthus sexfasciatus . 197 aspera, Campylaspis . . . 250, 255 aspheronicus, Euplotes . . . .124 assimile, Armadillidium .... 209 astriger, Armadillidium .... 208 astriger, Armadillo . . . .208 ater, Porcellio . . . . .198 aternanus, Trachelipus larii . . . 207aternanus, Tracheoniscus larii . . . 207aterrimus, Phoeniculus . . . 37, 45Atlantidiidae . . . . .191 atrox, Phoniscus . . . 135-136, 137audax, Rhinolophus aciminatus . . 132 aulacoeis, Campylaspis . . . 250, 252Aulonocara . . . . . .160 auritus, Porcellio ..... 198 ausseli, Armadillo . . . . .219 australis, Eubalaena glaciolis . 67, 68, 69, 71avium, Haplachromis . . . .159 badium, Armadillidium .... 208 badium siculorum, Armadillidium . . 208Balaena . . . . . 67, 71 Balaenoptera . . . . 65, 66, 71 balaenopterae, Cyamus . . . 65-66, 71Balantiopteryx . . . . -3*7 Balionycteris . . . . .128 Balloniscidae .... 191, 205 Balloniscus ...... 205 balstoni, Scoteinus . . . . . 173 balteatus, Euplotes ..... 101 balticus, Euplotes . . . . . 101 balticus, Porcellio ..... 207 balticus, Trachelipus .... 207 balticus burzenlandicus, Porcellio . . 207balticus burzenlandicus, Trachelipus . . 207barbertoni, Bethalus . . . .221 Barentsovia . .83, 84, 86, 87-92, Pis. 1-2 barentzovoensis, Barentsovia . 83, 86-88, 89, 90, 91, Pis. 1-2 barentzovoensis, Finmarchinella 83, 86-88, 89, 90, 91, Pis. 1-2 barentzovoensis, Nereina . . . 83, 86barnardi, Bethalus ..... 222 barnardi, Cubaris . . . . .222 barroisi, Porcellio ..... 198 batesoni, Porcellio . . . . .198 benaci, Metoponorthus . . . .196 Benechinus . . . . . .213 bergomatius, Cylisticus plumbeus . .192besi, Periscyphis . . . . .216 Bethalus .... 221-222, 245 bicarinata, Campylaspis . 250, 254, 287-28gbicarinata, Merulana .... 228 bicarinata, Spherillo .... 228 bicolor, Exzaes . . . . .212 bicoloratus, Armadillo .... 229 bicoloratus, Periscyphis . . . .216 bicoloratus, Spherillo .... 229 bicornis, Mahehia . . . . . 195 biellensis, Cylisticus. .... 192 bifrons, Armadillo ..... 229 bifrons, Spherillo . . . . .229 bimaculatus, Hemichromis . . .141binotatus, Buddelundia . . . .223 bipartitus, Buddelundia .... 223 bistriatus, Porcellio . . . .198 bisulcatus, Euplotes . . . 99, 101 bituberculatus, Armadillo . . .219 bituberculatus, Diploexochus . . .224bizonatus, Periscyphops . . . .217 blattarius, Porcellio .... 198 bloyeti, Haplochromis . . 145, 162, 163blythi, Rhinolophus . . . 131, 137 blythi, Rhinolophus blythi . . -131blythi blythi, Rhinolophus . . -131blythi calidus, Rhinolophus . . .131blythi szechwanus, Rhinolophus . -131bocki, Nesodillo ..... 228 bodenheimeri, Hemilepistus . . -194bonetti, Campylaspis . . 250, 251, 261boopis, Cyamus .... 66-67, 71 borelli, Plataoniscus .... 205 borellii, Alloniscus ..... 205 botswanae, Laephotis . 73, 76, 77-79, 80, 81brachycephalus, Spherillo. . . .229 brachypterus, Philetor . . . . 134 brachypterus, Philetor brachypterus . . 135brachypterus, Vespertilio . . -134 brachypterus brachypterus, Philetor . .135brachypterus rohui, Philetor . . . 135brachypterus verecundus, Philetor 134-135, 137bredanensis, Steno .... 70, 71 brentanus, Trachelipus .... 207 brentanus, Tracheoniscus .... 207 brevicornis, Armadillo . . . .219 brevicornis, Campylaspis 250, 254, 270-271brevipennis, Porcellio . . . .198 INDEX 339 brevis, Ignambabrevis, Phalababuddelundi, PorcellioBuddelundia .bulbosa, Campylaspis . 215 . 206 . 198 222-223 250, 254, 273-274, 289 bulgaricus, Trachelipus .... 207bulgaricus, Tracheoniscus. . . . 207bullatus, Haplochromis . . 159, 160, 161Burton, P. J. K. . . . l-48, i PI. burzenlandicus, Porcellio balticus . . 207burzenlandicus, Trochelipus balticus . . 207 Cacicus ..... 36, 37 cadornae, Pipistrellus .... 303 caelatum, Armadillidium .... 208 Caeroplastes . . . . . . 193 calidus, Rhinolophus blythi . . -131caligans, Spherillo . . . . .229 Callaeas . . 3, 4, 6, 7, n, 12, 14, 22, 23. 26, 35-36, 44Callaeidae .... 1-48, i PI. callosus, Buddelundia .... 223 calmani, Porcellio ..... 198 calypso, Rhinolophus acuminatus . .132Campy laspides . . . . .270 Campylaspis ..... 247-3 oo Campylorhamphus . . . . -42 canaliculata, Campylaspis . 250, 252, 263 canaliculata, Merulana . . . .228 canaliculatus, Spherillo .... 228 canariensis, Armadillo .... 224 canariensis, Diploexochus. . . .224 canariensis, Porcellio . . . .198 caperata, Campylaspis . . . 250, 254, 279-280, 282carinatus, Armadillo .... 222 carinatus, Bethalus ..... 222 carinatus, Hemiporcellio . . . -195carniolense, Armadillidium . . . 208Carolliinae . . . . . .319 carpathicus, Protracheoniscus saxonicus . 207carthaginensis, Porcellio . . . .198 carunculata, Anthochaera . . -34 carunculatus, Creadion . 3, 4, 5, 6, 10, n, 34cassida, Armadillo ..... 222 cassida, Bethalus . . . . .222 castor, Diploexochus . . . .224 catodon, Physeter . . . 66, 67, 70, 71catodontis, Cyamus . . . 67, 71 caucasius, Cylisticus . . . .192 caudatus, Ourachaerus .... 232 cavernicola, Periscyphis . . .216 cayennensis, Porcellio . . . . 199 cela, Cacicus . . . . . -37 celsicauda, Diploexochus .... 224 ceti, Cyamus ..... 67, 71 charon, Euplotes . . 97, 113, 115, 123chindeensis, Periscyphis . . .216 Chironax .... 127-129, 137 Chiroptera 73-82; 128-138; 167-188; 301-336Cichlidae ..... 139-165 ciliata, Euglypha . . . . .104 Ciliatea . . 95-102, i PI.; 113-126, i PI.cilicius, Metoponorthus . . . .196 cinctus, Armadillo ..... 220 cinctutus, Pyrgoniscus . . . .232 cinerascens, Buddelundia . . . 223 cinerea, Callaeas . . . 3, 4, 6, n, 12, 22, 23, 26, 35-36 cinerea, Creatophora . . . -37cingendum, Armadillidium . . . 208cingendum, Armadillidium maculatum . 208cingulatus, Diploexochus .... 224 cinnamomomeus, Scotoecus albofuscus . 175circe, Rhinolophus acuminatus . . . 132Circoniscus . . . . . .218 circumdatus, Pipistrellus . . 132-134, *37Ciridops ..... 25, 28 civilis, Periscyphis . . . . .216 clathrata nuda, Cythere . . . 86, 87, 88clausus, Synarmadillo . . . .231 clavata, Campylaspis . . 250, 255, 292Coelops ...... 320 cognata, Campylaspis 250, 254, 275, 277-278cognatus, Porcellio . . . .199 Coleura ....... 323 collicolus, Porcellidium .... 206 collicolus, Porcellio ..... 206 collicolus, Porcellium .... 206 collinus, Armadillo . . . .220 coloratus, Diploexochus .... 224 compressa, Euglypha . . . .104 concinella, Normanicythere . . -85concinnus, Scleropactes . . . .218 congener, Armadillo . . . .224 congener, Diploexochus . . . .224 conglobator, Armadillo . . . ^ . 220conisaleus, Diploexochus . . . 224 contra, Sturnus . . . . -37 Contractus, Porcellio . . . . 199 Corcorax . . . . . .29 corcyraeum, Armadillidium . . . 209cornutus, Rhinolophus . . . -131cornutus pumilus, Rhinolophus. . -131Corvus ....... 38 costata, Campylaspis . . 250, 253, 266costata speciosa, Campylaspis . . . 266coxalis, Metoponorthus .... 196 Graseonycteridae .... 301-336 Craseonycteris .... 301-336 crassus, Armadillo . . . . .224 crassus, Diploexochus .... 224 Creadion . . . 3, 4, 5, 6, 7, 10, n, 14. 34- 35. 36. 38. 44 Creatophora . . . . . -37 crenosus, Euplotes ..... 101 creper, Saidjahus ..... 229 crispa, Campylaspis. . . . 250, 253 cristatus, Euplotes . . . 99, 101, 121 340 INDEX cristatus, Hemilepistus .... 194 cruentatus, Porcellio . . . . 199 Crustacea . 59-64, i PL; 65-72; 83-94, 2 Pis.; 189-246; 247-300 Cubaris . 219, 220, 221, 222, 225, 226, 231 Cumacea ..... 247-300 Curds, C. R. . 95-102, i PL; 113-126, i PL curvicosta, Barentsoviacurvicosta, Finmarchinella Cushmanidea Cyamidae Cyamus .... cyanomelas, Rhinopomastus Cylisticidae Cylisticus Cythere .... Cythereis . 83, 86, 87, 88, 89, 90-91, Pis. 1-2 . 83, 86, 87, 88, 89, 90-91, Pis. 1-2 . 86 59-64, i PL; 65-72 • 59, 65-70, 71 . 42 191, 192 . 192 86, 87, 88 . 83, 84, 88 dalmatinus frascatensis, Orthometopon . 206danae, Armadillo ..... 230 danae, Spherillo . . . . .220 davidi, Armadillidium .... 209 debueni, Porcellio ..... 199 decoratus, Spherillo . . . .229 decumanus, Psarocolius . . . -37delphini, Isocyamus . . . 70, 71 dentifrons, Cylisticus . . . .192 depressus, Armadillo .... 222 depressus, Bethalus .... 222 Desertoniscus . . . . . .193 desertorum, Protracheoniscus . . . 206 Desmodontidaediadaleos, Euplotes .Diclidurinae .Diclidurus dimorphus, PorcellionidesDiploexochus .disjunctus, Diploexochus .dispar, Porcelliodispersus, Spherillo . .dollfusi, EubelumDolphin, white-beakedDorahy, J. E.dugesi, Armadillodugesi, Diploexochusdumorum, Armadillodumorum, Venezillo ecaudatus, Diploexochusechinata, Campylaspisegens, Armadilloelbanum, ArmadillidiumEllis, J. P. elongatus, HemilepistusEluma .emarginatus, Bethalus 319, 320, 322, 330. 124 • 319323, 325. 326, 333 . 204 223-227, 245. 224. 199. 229. 214 59-64, i PL. 95-102, i PL • 225 • 225 • 232 • 232 • 225250, 253, 270 . 220 . 209 . 189-246 • 194 212, 242. 222 emarginatus, Scotophilus . Emballonuridae . 317,322, 323,328, 329, Emballonuroidea Emoa .... Emoia .... emunitus, Armadillo emunitus, Bethalus . Ennurensis episimus, Hekelus . epops, Upupa Eptesicus erinaceus, Armadillo erinaceus, Spherillo . erraticus, Cyamus . erythroleucus, Armadillo . Eschrichtius . esterelanum, Armadillidium esterelanus, Cylisticus Ethelum Eubalaena Eubelidae . . 191 Eubelum Euglypha Euglyphidae . Euplotes . 95-102, i Euporcellio eurystomus, Euplotes 118, evansi, Porcellio exarata, Campylaspis 250, Exzaes . • 173318, 319, 320, 321, 324. 325, 326, 327,330. 332, 333, 334 • 329, 334 . 5653-58 . 222 . 222 . . . 193 213 37. 42 • 74. 134 . 229. 229. 67-68, 71 . 220 70. 71. 2O9. 192213-214 67, 68, 69, 71, 213-218, 242-244214, 215, 242, 243103-112, 2 Pis. . IO3-II2, 2 Pis. PL; 113-126, i PL. 207 119, I2O, 121, 124. 199 254, 266, 284-285 . 212 falabae, Scotoecus . . 169, 178, 183, 184falabae, Scotoecus hindei . . . 183-184Falculea. . . . . . -42 famosa, Nectarinia . . . .42 feae, Armadillo . . . . .228 feae, Pericephalus ..... 228 festivus, Armadillo ..... 225 festivus, Diploexochus .... 225 ficorum, Porcellio . . . . .199 ficulneus, Porcellio . . . . . 199 finmarchica, Cythereis . . . 83, 84finmarchica, Finmarchinella . 83, 84-85, 86, Pis. 1-2 finmarchica, Hemicythere . . .84 Finmarchinella . . . 83-94, 2 Pis.fissifrons, Pagana . . . . .206 flavocinctus, Porcellio . . . . 199 flavoscutatum, Armadillidium . . . 209flavovittata, Porcellio . . . . 199 flavus, Armadillo ..... 220 fongosiensis, Anchicuboris . . .219 fontium, Protracheoniscus . . . 206formosana, Nagara .... 205 formosanus, Nagurus .... 205 fragilis, Agnara . . . . .205 frascatensis, Orthometopon dalmatinus . 206 INDEX frigida, Campylaspis . . . 250, 255 fritschei, Reductoniscus .... 229frontalis, Sphaeroniscus . . . .218frontalis, Spherillo . . . . .230frontexcavatum, Armadillidium . . 209 frontirostre, Armadillidium . . . 209frontosus, Buddelundia .... 223frontosus, Metoponorthus . . . 196 furcatum, Armadillidium .... 209furcatus, Diploexochus . . . .225Furipteridae . 318, 319, 320, 326, 327, 328, 330fuscomarmoratus, Metoponorthus . .196fusiformis, Campylaspis . . . 250, 252 grisescens, Spherillo.grossus, Armadillo .grossus, Spherilloguttata, CampylaspisGymnastinops • 230 • 230. 230 250, 251, 258-259 • 39 galapagoensis, Cubans . galapagoensis, Diploexochus galeatus, Armadillo . galeatus, Pericephalus . gallicus, Porcellio . . Gelsana . . . . gemmulatus, Porcellio . Gerufa .... Gerutha ... gestroi, Armadillidium . gibbosus, Eschrichtius . gigas, Armadillidium simoni gigas, Cubaris .. gigas, Diploexochus . glabra, Campylaspis . glacialis, Eubolaena. . glacialis, Eubolaena glacialis glacialis australis, Eubalaena glacialis glacialis, Eubalaena glacialis japonica, Eubalaena glebulosa, Campylaspis . Globicephala ... globosa, Campylaspis globus, Synarmadillo . Glomerulus ... glomus, Armadillo . . glomus, Diploexochus . Glossophaga . . . gordoniensis, Diploexochus gracilior, Merulana translucida gracilipennis, Cylisticus . gracilis, Cyamus . . grandinatum, Armadillidium graniger, Porcellio . . granulata, Campylaspis . granulatum, Armadillidium . . granulatum peloponnesiaca, Armadillidium granuliferus, Porcellio ... gravei, Metoponorthus . . . ... 225 . . . 225... 228... 228. . . 199. . .214... 199... 194.. 214-215. . . 209. . 70, 71. . .211... 225. . .225. 250, 252, 261. . .69. 68, 69, 71. 67, 68, 69, 71. 68, 69, 71. . 69, 71. 250, 255, 293-295... 70, 71250, 255, 291, 295, 297. . . .231.... 228. . . .225.... 225. . . -325. 225. 228.19268, 71. 209. 199250, 252.210209200196 Greenwood, P. Hgrenadensis, Armadillogrenadensis, Venezillogreyii, Scoteinus .griseoalbus, Armadillogriseoalbus, Bethalus 139-i65 . 232.232. 173.222.222 Haloporcellio .. Haplarmadillo . . Haplochromis ...harpa, Euplotes . .harsadiensis, Armadillo .hartae, Campylaspis . hebridarum, Spherillo .Hedley, R. H. . . Hekelus . . . Hemichromis . . Hemicythere ..."Hemicytherinae" . .Hemignathus . . . .Hemilepistus ....Hemiporcellio . . . hermagorensis, ProtracheoniscusHermanites . . . .herzegovinensis, Porcellio . .herzegovinensis, Porcellium .Heteralocha ....Hiallum . . . . hilgendorfi, Eubelum . .hilgendorfi, Hiallum . . ... 194. . -215.. 139-I&5 . . 101, 119... 220. . 250, 253... 230103-1 12, 2 Pis.. . .213. . .14184, 86, 87, 88, 89. . .88. . 5, 42 .. 194, 237. . .19520686206206i PI.215215215 1- Hill, J. E. 73-82; 127-138; 167-188; 301-336hindei, Nycticeius hirundo . . .182hindei, Scotoecus . 169, 171, 172, 175, 178, 179-i8o, 181, 182, 183, 184, 185, 186hindei, Scotoecus hindei 179, 180-i82, 183, 184hindei, Scotoecus hirundo . 181, 182, 183 hindei albigula, Scotoecus 180, 181, 182-i83hindei artinii, Scotoecus . . . .182hindei falabae, Scotoecus . . . 183-1 84hindei hindei, Scotoecus 179, 180-i82, 183, 184Hipposideridae 317, 318, 319, 320, 322, 323, 325, 326, 327, 328, 329, 330, 332, 333Hipposideros ...... 303 hirsutulus, Protracheoniscus . . . 206hirticornis, Gerufa . . . . . 194 hirtum, Armadillidium .... 209 hirtus, Agabiformius . . . . 193 hirtus, Lucasius ..... 193 hirundo, Scotoecus . . 169, 171, 172, 175, 177-179, 181, 183, 184, 185, 186 hirundo, Scotophilus . . . . 177 hirundo hindei, Nycticeius . . .182hirundo hindei, Scotoecus . 181, 182, 183 hispida, Metoponorthus . . . .196 hispida, Porcellio . . . . .196 hispida, Porcellionides . . . 196 hispidus, Ennurensis . . . . 193 holosericeus, Amblyrhamphus . . -37horrida, Campylaspis . . 250, 255, 292 342 INDEX horridoides, Campylaspis .horsfieldii, Myotis .horsfieldii, Myotis horsfieldiihorsfieldii horsfieldii, Myotishorsfieldii lepidus, Myotis.hughscotti, PeriscyphisHuia ....Hurley, D. E.hybridum, Armadillidiumhybridum, SchizidiumHyperoodon .hypotoreus, SpherilloHypotrichida .hypselos, Diploexochushypsinephes, Diploexochus 250, 255, 292-293. 132 . 132, 137. 132, 137 • 132. 216 1-48, i PI. 59-64, i PI.; 65-72 . 213 . 213 . 71 • 23095-102, i PI.; 113 . 225. 225 Ignamba . . . . . .215 ignavum, Eubelum . . . . .214 imbutus, Porcellio ..... 200 imbutus pellegrinensis, Porcellio . . 200immotus, Armadillo .... 220 immsi, Hemiporcellio . . . . 195 immsi, Porcellio . . . . .195 inaequalis, Buddelundia .... 223 incanus, Porcellio ..... 200 Inchanga ...... 195 incicus, Scleropactes . . . .218 incisa, Nagara ..... 205 incisus, Nagurus ..... 205 inconspicuus, Armadillo .... 220 indentatus, Euplotes .... 101 ingens, Spherillo ..... 230 iniqua, Merulana ..... 228 iniquus, Spherillo ..... 228 inkystans, Euplotes . . . . 101 inornata, Campylaspis . . . 250, 251instrenuum, Eubelum . . . .214 integer, Armadillo .... 220 intercalarius, Porcellio .... 200 intermedia, Campylaspis 250, 255, 277, 289, 297intermixtus, Armadillo .... 220 interpolator, Porcellio .... 200 Isocyamus ..... 59, 70, 71 japonica, Barentsovia . . . 91-92 japonica, Eubalaena glacialis . . 69, 71japonica, Finmarchinella . . . 91-92japonica, Nereina . . . . 83, 91jelskii, Porcellio ..... 200johnstoni, Campylaspis . . . 250, 253johorensis, Tadarida . . . 136-137 Jones, N. S 247-30O jonesi, Campylaspis. . . 250, 253, 263 kaokoensis, Diploexochus . . . 225 kiiensis, Campylaspis . . . 250, 252 KerivoulaKerivoulinae .Kisuma . kogmani, DiploexochusKrafft, J. I. .kuhlii, Scotophilus .kunenensis, Periscyphiskunenensis, Periscyphops . 328319, 320, 325 • 232 • 225 103-112, 2 PIS. 174 . 216. 216 labiatus, Buddelundia .... 223 lacteolus, Metoponorthus .... 196 Laephotis ..... 73-82 laevigata, Campylaspis 250, 251, 258, 260-26ilaevigatus, Buddelundia .... 223 laevis, Gerutha . . . . .214 laevis, Porcellio . . . . .201 Lagenorhynchus .... 59, 71 lamellatus, Porcellio . . 191, 194, 201 laminigerum, Armadillidium . . .210larii, Trachelipus ..... 207 larii, Tracheoniscus ..... 207 larii aternanus, Trachelipus . . . 207larii aternanus, Tracheoniscus . . . 207Lasiurus ...... 328 lateralis, Buddelundia .... 223 laticarpa, Campylaspis . . . 250, 254laticauda, Mahehia .... 195 laticauda, Tura ..... 204 latidactyla, Campylaspis . . . 250, 253Latidens . . . . . -303 latifrons, Armadillo . . . .222 latifrons, Bethalus . . . . .222 latissimus, Periscyphis . . . .216 latissimus, Porcellio. .... 200 latus, Euplotes patella .... 124 latus, Protracheoniscus .... 206 legendrei, Campylaspis . . . 250, 253lekaguli, Hipposideros .... 303 lentus, Spherillo ..... 230 lepidus, Myotis . . . . .132 lepidus, Myotis horsfieldii. . . .132 lepidus group, Rhinolophus . . .129lepidus subgroup, Rhinolophus . .129 lepineyi, Porcellio ..... 200 Leptotrichus . . . . . 195, 237 leucocephalus, Microcercus . . -215leucocephalus, Periscyphis . . -215leucorhamphus, Cacicus . . . -37leucostictus, Sarotherodon . . .163lifuensis, Armadillo ..... 220 lifuensis, Cubaris . . . . .220 limbata, Periscyphis . . . .216 limbatus, Bethalus . . . .222 limenites, Diploexochus . . . .225 Lincoln, R. J.. 59-64, I pl-.' 66-72; 189-246lineare, Trinema . . . . -103 Liponycteris . . . . . -323 littoralis, Metoponorthus . . . .196 INDEX 343 loati, Haplochromis. . . . .162 Lonchorhina ...... 326 longicauda, Porcellio .... 200 longipennis, Porcellio .... 200 longipes, Armadillo .... 225 longipes, Diploexochus .... 225 loyaltiensis, Emoia samoensis . . 53-58lubricum, Eubelum . . . . .214 Lucasius . 191, 193, 195, 197, 201, 237 lugubris orarum, Porcellio . . .201 lugubris vizzavonensis, Porcellio . .201lusitanus, Porcellio .... 202 lusitanus, Porcellio scaber . . . 202 lutshniki, Trachelipus .... 207 lutshniki, Tracheoniscus .... 207 Lygosoma . . . . . -56 Lyprobius ...... 205 macconneli, Haplochromis . 141, 149, 153, 154-i6i, 162, 164macmahoni, Armadillo .... 220 macrocneme, Miniopterus medius . 135, 137macrodens, Bethalus .... 222 macrophthalma, Campylaspis . 250, 253, 264macrops, Gerufa ..... 194 maculata, Balloniscus .... 205 maculata, Campylaspis . . . 250, 255maculata, Mahehia . . . . 195 maculata, Philoscia .... 205 maculatum cingendum, Armadillidium . 208maculipes, Porcellio. .... 200 maculosa, Pagana ..... 206 maculosus, Spherillo .... 230 madagascariensis, Agnara . . . 205 madagascariensis, Metoponorthus . . 205magna, Sturnella . . . . -37 magnicirratus, Euplotes . . . . 101 magnificus, Porcellio .... 200 mahagiensis, Haplochromis . . .162Mahehia. ...... 195 makuae, Diploexochus .... 226 Mammalia . 127-138; 167-188; 301-336 mammillata, Sunniva . . . -213mansa, Campylaspis 250, 254, 283, 286-287marcidus, Armadillo . . . .228 marcidus, Pericephalus .... 228 mareoticum, Armadillidium . . .210marginalis, Uramba. .... 205 marginatus, Spherillo . . . -230marginenotatus, Porcellio. . . . 200 marginepilosa, Gerutha . . . -215marinensium, Armadillidium . . .210marmorata, Gerufa . . . . .194 marmoratus, Synarmadillo . . .231marquesarum, Spherillo .... 230 marquesarum, Xestodillo .... 230 marshalli, Rhinolophus .... 303 mauritanica, Campylaspis . 250, 251, 261 mayeti, Armadillo . . . . .220medius, Miniopterus . . . -135 medius macrocneme, Miniopterus . 135, 137Medway, Lord .... 53-58 Megadermatidae . . 318, 319, 320, 322, 325, 326, 327, 329, 330, 332Megaptera . . . .66, 67, 70, 71 mehelyi, Protracheoniscus . . . 206 mehelyi, Protracheoniscus politus . . 206meiringi, Diploexochus . . . .226 melaena, Globicephala . . . 70, 71melanocephalus, Chironax . 127-129, 137 melanurus, Acaeroplastes . . -193 melanurus, Metoponorthus . . . 193meleagris, Metoponorthus . . . 197 meleagris, Metoponorthus pruinosus . 197 Merulana ...... 228 Mesarmadillo . . 213, 214, 215, 243 Metoponorthus . . 193, 196-197, 199, 200, 204, 205, 206, 237, 238 197 • 215, 243318, 322, 324, 325 325 228 . 228 . 215 . 201 319, 327135. 137. 328 250, 253. 129, 131 . 213. 129 193. IOI 213 193. 22O. 2I4. 214 113, 114, 115-117,Il8, 119, I2O, 121, 123. 201 moebiusi quadricirratus, Euplotes . 113, 123 mohamedanicum, Armadillidium . .210 Molossidae . . 318, 319, 320, 322, 324, 325, 326, 327, 328, 329, 330 monocellatus, Haplarmadillo . . -215monoceros, Monodon . . 68, 69, 71 Monodon .... 68, 69, 71 monodontis, Cyamus . . . 68, 71montagui, Diploexochus .... 226montana, Gerufa . . . . . 194montanus, Armadillo .... 220montanus, Porcellio. .... 203montanus, Porcellio spinipennis . . 203 monticola, Buddelundia .... 223montivagus, Armadillo .... 230montivagus, Spherillo .... 230 Mica MicrocercusMicrochiropteraMicronycteris .microps, Armadillo .microps, Glomerulusmicrops, Ignamba .mildei, PorcellioMiniopterinae .Miniopterusminor, Campylaspisminor, Rhinolophus.minor, Sunnivaminor subgroup, Rhinolophusminuta, Angaraminuta, Euplotesminuta, Pareluma .minutus, Agabiformiusmiser, Armadillomodestum, Ethelummodestus, Mesarmadillo .moebiusi, Euplotes moebiusi, Porcellio 344 INDEX Mormoopidae . 319. 320, 323. 325. 326.327, 328, 329, 330, 332, 333 • 325 . 222 250, 256, 297-2QQ . 2IO . 320 205 . 20565, 71 100, 117 • 132,137. 303 Mormoops tnucidus, Bethalus . multinodosa, Campylaspis muricatum, Armadillidium Murininae mus, Lyprobius mus, Uramba musculus, Balaenoptera . mutabilis, Euplotes Myotis .... myrmecophilus, Metoponorthus myrmecophilus 196myrmecophilus, Myrmeconiscus myrmecophilus 196myrmecophilus myrmecophilus, Metoponorthus 196myrmecophilus myrmecophilus, Myrmeconiscus 196myrmicidarum, Metoponorthus . .196 Mystacinidae . ' . mystica, SunnivaMysticeti mysticetus, BalaenaMyzopodidae . 318, 319, 320, 324, 325. 327. 329, 33° . 213 . 71 67, 71 318, 320, 326, 327, 330 Nagara .Nagurus. namibensis, Laephotisnana, Nagara .Nannastacidaenanus, Diploexochusnanus, Nagurusnarentanus, Porcellionasatum, Armadillidium . 205 . 205, 23973, 75-76, 77, 79, 80. 205. 249. 226• 205. 201 . 212 nasatum sorrentinum, Armadillidium . 210 nasatus, Cylisticus ..... 192natalensis, Inchanga .... 195 Natalidae . 318, 320, 326, 327, 328, 330 naupliensis, PorcellioNeale, J. W. .nebulosus, DiploexochusNectariniaNeocyamusNereina .Nesodillo .nicklesi, Porcellionicobaricus, Armadillonicobaricus, Spherillo .nigra, Emoia .nigra, Porcellionigricans, Periscyphisnigripes, Buddelundia .nigrobrunneus, Metoponorthusnigromarginatus, Armadillonigropunctatus, Periscyphisnigropunctatus, Synarmadillo . 2OI 83-94, 2 Pis. 226. 4259, 70, 71 83, 86, 87, 88, 90, 91. 228f . . 201 • 230 • 230 • 53 . 201 . 217 . 223 . 196 . 221 231. 231 niloticus, Sarotherodonnitens, Campylaspis.nitidulus, Armadillidium .nobilis, Spherillo . .Noctilio . Noctilionidae . . 318,325, 326 noctulinus, Nycticejusnoctulinus, Vespertilionodosus, Cyamusnodosus, Hemilepistusnodulosa, Campylaspisnormani, Lucasius .normani, Porcellio .Normanicytherenovaeangliae, Megaptera .nuda, Campylaspis .nuda, Cythere clathrata .Nyctalus Nycteridae . 318, 319,325. 326, 327Nycticeius Nyctophilinae 140, 149 250, 251, 256, 258. 210 • 230 • 329 319, 320, 323, 324,327, 329, 330, 332 • i?4. 172, 174 68-69, 7i • 194. 250, 255 . 2OI . • . 201 85 66, 67, 70, 71 250, 251, 256-258 86, 87, 88 . 171 320, 322, 323, 324,329, 330, 332, 333169, 170, 171, 173, 174, 175, 182, 186• 317, 322 obliquidens, Diploexochus . . . 226 obliquipes, Spherillo . . . -230 obsoletus, Porcellio ..... 201obtusa, Angara . . . . . 193obtusifrons, Porcellio . . . .201obtusiserra, Porcellio .... 201obtusus, Agabiformius .... 193occidentals, Protracheoniscus . . .196ocellatus, Porcellio . . . . .201octocarinatus, Euplotes . . . .124octocirratus, Euplotes . . . 100, 101odherni, Armadillidium . . . .210odherni, Armadillidium peraccai . .210Odontoceti . . . . . -71oertzeni, Armadillidium . . . .213oertzeni, Schizidium . . . .213 officinalis syriaca, Armadillo . . .221 Ogden, C. G 103-112, 2 Pis. oliveti, Armadillidium . . . .210olivieri, Porcellio . . . . .198Oniscoidea ..... 189-246opacus, Buddelundia .... 223opihensis, Actoecia . . . .218 orarum, Porcellio lugubris . . . 201 orarum, Porcellio orarum . . .201 orarum orarum, Porcellio . . .201 orarum vizzavonensis, Porcellio . . 201 orientalis, Campylaspis . . . 250, 251ormeanum, Armadillidium . . .210ornatus, Scotomanes . . . . 173 Orodillo 228 orphanus, Diploexochus .... 226Orthometopon ..... 206 Ostracoda .... 83-94, 2 Pis. INDEX 345 Ourachaerus .ovalis, Campylaspis.ovalis, Cyamusovampoensis, Cubarisovampoensis, Diploexochusoxyzomus, Armadillooxyzomus, Cubaris • 232 250, 253 69, 71 . 226. 226 . 221. 221 pachytos, Diploexochus pacifica, Campylaspis pacificus, Cyamus paeneglabra, Campylaspis Pagana . palaestinus, Hemilepistus pallasi, Armadillidium palliata, Falculea pallidus, Cylisticus . pallidus, Lucasius pallidus, Porcellio pallidus, Scotoecus . pallidus, Scotophilus Pandanus panurus, Armadillo . panurus, Bethalus . papillata, Campylaspis papillosa, Kisuma Paracubaris paradoxolophus, Rhinolophus Paraperiscyphis Pareluma parietinus, Porcellio parkei, Euplotes parvus, Armadillo parvus, Porcellio parvus, Scutocyamus . parvus, Spherillo patella, Euplotes patella latus, Euplotes paucinodosa, Campylaspis paucispina, Campylaspis. pauper, Porcellio pauperculus, Diploexochus pellegrinense, Armadillidium pellegrinensis, Porcellio imbutus . . 200 Pelmatochromis . . . . .141 peloponnesiaca, Armadillidium granulatum 209peltatus, Spherillo . . . . -230 penicilliger, Haloporcellio . . . -194peraccae, Armadillidium . . . .210 peraccai, odherni, Armadillidium . .210Pericephalus ...... 228 Periscyphis . . 215,216-217,218,231,243 Periscyphops . . .216,217-218,243,244perkinsi, Armadillo . . . . .231 perkinsi, Spherillo . . . . .231 petraeum, Armadillidium . . . .210 petronius, Haplochromis . . . 161, 163phaeacorum, Porcellio rathkei . . . 207 . 226 . 250,25170, 71 250, 251, 258, 261. 206 • 194 . 2IO . 42 . 192 . 195 195 171,172-174, 175, 186169, 170, 172, 173 • 195 . 222. 222250, 255, 296 232. 218 303 . 216 • 213 . 2OI 119-121, 122, 123, 124, PI. I. 230. 201 59-64, i PL; 71 230 97. 99. IO°. 117. 124. 124 25°. 255, 291-292250,254, 275-277 . 201 . 226 . 210 phaeacorum, Trachelipus .Phalaba .... phaleronensis, Metoponorthus .phaleronensis, Orthometoponphaleronensis, Porcelliopharyngalis, Haplochromis Philetor Philoscia .... philoscoides, MetoponorthusPhoeniculus ....Phoniscus ....Phyllostomatidae 317, 319, 320, 327. Phyllostomatoidea .physalus, BalaenopteraPhyseter .... physeteris, Neocyamuspictus, Rotunguspiger, Armadillopila, Gerutha ....pileus, Campylaspis .pilosa, Campylaspis . 250, pilosa, Geruthapilularis, Armadillo .pilum, Eubelum Pipistrellus . . . 132-Pisces .....pisum, Armadillopisum, Diploexochusplanarius, Porcellio .Plataoniscus ....Platycyamus ....platyuropus, Campylaspis.plicata, Campylaspis 250, 254, plicata, Tadaridaplumbeus, Cylisticusplumbeus bergomatius, Cylisticusplumbeus umbricus, Cylisticus .plumipes, Euplotes .politus, Leptotrichuspolitus mehelyi, Protracheoniscuspollex, DiploexochusPolyacanthus ....polythele, Diploexochus .pontremolensis, Cylisticus.porcata, Campylaspis . 250, 252,Porcellidium ....Porcellio 191, 194, 195, 196, . 207 206, 239, 240. 206. 206. 206161, 163 134-135. 137 • 205. 19637-45 135-136, 137322, 325, 326, 329, 330. 332 • 329. 65, 66, 71 66, 67, 70, 7170, 71. 218 . 221. 215250, 252254,271-272 215. 221. 215 134. 137. 303. 139-165. 226. 226. 202. 205 59.70-71250, 252, 259280-282, 289 • 136, 137 . 192. 192. 192117, 119, 124 • 195. 206. 226. 229. 226 PorcellionidaePorcellionides .Porcellium .porphyrivagus, Caeroplastesporphyrivagus, Metoponorthus .portofinense, Armadillidiumpraeustus, Porcellio .pretoriensis, Armadillopretoriensis, BethalusPrionoplus .procerus, Hemignathus . 192 261-2&3, 265. 206 197-204, 206,207, 238, 239I9L 193-205, 237-239196, 204. 206 193193 2102O2222222 442 346 INDEX Protozoa 95-102, i PI.; 103-112, 2 Pis.; 113-126, i PI. Protracheoniscus . . . 196,206-207provincialis, Porcellio .... 202 proximatus, Armadillo . . . .221 pruinosus, Metoponorthus . 199, 200,204 pruinosus anconanus, Metoponorthus . . 197pruinosus meleagris, Metoponorthus . . 197Psarocolius ..... 37,39 pseudoratzeburgi, Euporcellio . . . 207pseudoratzeburgi, Porcellio . . . 207pseudoratzeburgi, Trachelipus . . . 207pseudorcae, Syncyamus . . . .62 Pseudotracheata .... 189-246 Pteropodidae . . . . . -325 pujetanum, Armadillidium . . .211pujetanus, Porcellio .... 202 pulchella, Campylaspis . . . 250,251pulcher, Periscyphis. . . ..217 pumila, Campylaspis . . 250,252,263 pumilus, Armadillo ..... 232 pumilus, Rhinolophus cornutus . . -131pumilus, Venezillo ..... 232 purpurascens, Eluma . . . .212 purpurascens, Spherillo . . . .231 purpureus, Phoeniculus . . . -37purpureus, Porcellio..... 202 pusillus, Rhinolophus . . 129-131, 137pusillus group, Rhinolophus . . 129,131pustulosa, Campylaspis . . . 250,255pygmaeus, Armadillo . . . .221 pygmaeus, Periscyphis . . . -231pygmaeus, Synarmadillo . . . -231pyrenaeus, Porcellio..... 202 Pyrgoniscus ...... 232 pyriformis, Tetrahymena . . . -97Pyrrhocorax . . . . . -37 pyrrhocorax, Pyrrhocorax . . -37 quadricirratus, Euplotes moebiusi . 113,123 quadrimaculatus, Diploexochus. . .226 quadrimaculatus, Mesarmadillo . -215 quadrimaculatus, Periscyphis . . .217 quadriplicata, Campylaspis . . 250,254 quadriserriatum, Armadillidium . .211 quadritracheata, Buddelundia . . . 223 quietum, Eubelum . . . . .214 quinquecarinatus, Euplotes . . 97, 101 quinquepustulatum, Armadillidium . .211 ragusae, Porcellio ..... 202raikovi, Euplotes . . .99, 100, 101, 123rariseta, Euplotes . 95-IO2, i PL; 115, 118rathkei phaeacorum, Porcellio . . . 207 razae, Alauda .reaumuri, Hemilepistusredacta, Campylaspis . 250,Reductoniscusreflexum, Ethelum .reflexus, Mesarmadillorehobotense, ArmadillidiumReptilia ....reticularis, Prionoplusreticulata, CampylaspisRhinolophidae 318,319,320,327. 328,RhinolophoideaRhinolophus . Rhinopoma . . 304,316,RhinopomastusRhinopomatidae3i6, 317, 318,323, 324, 325, rhodesiae, Bethalus .rhodesiensis, Diploexochusribauti, Sphaerobathytropariparium, Armadillidium .robusta, Arachnotherarohui, Philetarrohui, Philetar brachypterusrosai, Armadillidiumroscida, Campylaspisroscoffensis, Euplotesrostellata, Campylaspis 250rostrata, Campylaspisrotunda, Euglypha .RotungusRousettus rubicunda, Campylaspis . 250rubromaculata, Campylaspis,rudolfianus, Haplochromis 153, rufa, Campylaspisrufescens, Diploexochusruficauda, Periscyphisrufobrunneus, Agabiformiusrufomarginatus, Spherillo .rugifrons, Buddelundiarugulosus, Cubaris .rugulosus, Spherillo .rupta, Campylospis . sabuleti, Metoponorthussabulifer, Porcellio .Saccolaimus .Saccopteryx .sagamiensis, CampylaspisSaidjahus saldanhae, Diploexochussalimalii, Latidens . . 41 • 194254,274-275, 278 . 229. 214. 214 . 211 53-58 4 . 250,253323,324, 325, 326,329, 33°, 332, 333 • 332129-132, 137, 303 ,321,323,332,333 . 38,42,45 319, 320, 321, 322, 326, 328, 329, 330, 332,333. 334 . 222 . 227 . 218 . 211 . 42 134.135 135. 211 . 250, 254. IOI ,253,267-268, 270 250, 255, 268, 289 103, 108 . 218 . 129 ,251,259-26o, 261 • 250,255141,142-150, 152, 155. 156, 157. 158. 159, 160, 161, 162 . 250,252 . 227. 217 • 193. 231. 223. 231 • 231. 250, 252 . 197 . 202 323 323250, 255 . 229 . 227 303 INDEX 347 salisburyensis, Diploexochus . . .227saltuum, Porcellio ..... 202 samoense, Emoa . . . . .56 samoense, Lygosoma . . . -56samoensis, Emoia .... 54, 55, 56 samoensis loyaltiensis, Emoia . . 53-58sanctum, Armadillidium . . . .211 sanfordi, Emoia .... 55,56 sarajevensis, Porcellio .... 202 sarasini, Nesodillo . . . . .228 sarculatus, Porcellio..... 202 Sarotherodon .... 148, 149, 163 sauteri, Orodillo . . . . .228 savonense, Armadillidium. . . .211 saxicola, Haplochromis . . . -157saxonicus, Protracheoniscus . . . 207saxonicus carpathicus, Protracheoniscus . 207scaber, Porcellio .... 199, 201 scaber lusitanus, Porcellio . . . 202 scaberrimum, Armadillidium . . .211scabrum, Armadillidium . . . .211 scammoni, Cyamus .... 70, 71 schellenbergi, Nesodillo .... 228 schirasi, Hemilepistus .... 194 Schizidium . . . . . -213 schlieffenii, Nycticeius . . . . 175 schlieffenii, Scoteinus . . 173, 181, 182Scincidae ..... 53-58 Scleropactes . . . . .218, 244 Scoteinus . . 170, 171, 173, 181, 182, 186Scotoecus ..... 167-i88 Scotomanes . . . . . . 173 Scotophilus . . . 169, 170, 171, 172, 173,174,175, 177Scotozous . . . . . .169 Scutocyamus . . 59-64, l Pl-» 71 serratipes, Campylaspis . . . 250, 254serratum, Armadillidium . . . .211 setaceus, Spherillo . . . . .231 sexfasciatus, Metoponorthus . . . 197sexfasciatus asifensis, Metoponorthus . . 197sharpi, Armadillo . . . . .221 siculorum, Armadillidium badium . . 208silvanus, Periscyphops . . . .217 silvarum, Armadillo .... 227 silvarum, Diploexochus . . . .227 similis, Campylaspis. . . . 250,252 simoni, Armadillidium . . . .211 simoni gigas, Armadillidium . . .211simplex, Synarmadillo .... 232 simulator, Porcellio ..... 202 sinuosa, Campylaspis . . . 250, 253Skink . .... 53-58 sociablis, Porcellio ..... 203 societatis, Pipistrellus . . . 133, 134solitarius, Cacicus .... 36, 37 sollers, Spherillo . . . . .231 sordidus, Porcellio ..... 203 sorrentinum, Armadillidium nasatum . 210 spatulata, Porcellio . . . . .203 spatulatus, Porcellio .... 200 speciosa, Campylaspis . . . 250, 253speciosa, Campylaspis costata . . .266speiseri, Emoia .... 54,55 Sperillo ....... 245 speyeri, Armadillidium . . . .212 Sphaerobathytropa . . . . .218 Sphaeroniscidae . . . 191,218,244Sphaeroniscus. .... 218,244 Spherillo . . .220,228,229-231,245 sphinx, Haloporcellio . . . 191, 194spicatus, Spherillo . . . . .231 spinicornis, Porcellio .... 203 spinifera, Campylaspides . . . .270 spiniger, Acanthoniscus . . . .219 spinipennis, Porcellio .... 203 spinipennis, Porcellio spinipennis . 202, 203spinipennis montanus, Porcellio. . . 203 spinipennis spinipennis, Porcellio . 202, 203spinipes, Porcellio ..... 203 spinosa, Akermania . . . . .219 spinosa, Campylaspis . 250, 253, 267, 270spinosus, Circoniscus . . . .218 spinosus, Paracubaris . . . .218 spretus, Porcellio ..... 203 squamatus, Periscyphops . . . .217 squamifera, Campylaspis . 250, 255, 293, 295, 296squamosus, Periscyphops . . . .217 stebbingi, Paraperiscyphis . . .216 steenbrasi, Diploexochus . . . .227 Steno . . . . . . 70,71 Stenoderminae . . . . .319 sticta, Campylaspis. . 250,253,264-265stipulatum, Eubelum . . . .214 stolikanum, Armadillidium . . .212straeleni, Haplochromis . . . .162 strelkovi, Euplotes ..... 101 striata, Campylaspis . . . 250,252 stricticauda, Armadillo .... 222 stricticauda, Bethalus . . . .222 strigosa, Euglypha . . . 103-112, PI. 2Sturnella . . . . . -37 Sturnus ....... 37 subbadius subgroup, Rhinolophus . .129subdentatum, Armadillidium . . .212suberorum, Cylisticus .... 192 subjaponica, Cushmanidea . . .86submersa, Campylaspis . 250, 255, 290-291subterraneus, Desertoniscus . . . 193subtransversus, Periscyphis . . .217succinctus, Porcellio .... 203 sulcata, Campylaspis . . 250,253,275sulcatus, Buddelundia .... 223 sumatranus, Rhinolophus acuminatus . 132 Sunniva ...... 213 sylvatica, Exzaes . . . . .212 Synarmadillo ..... 231-232 Syncyamus ..... 59,62 syriaca, Armadillo omcinalis . . .221szechwanus, Rhinolophus blythi . -131 INDEX tabularis, Diploexochus . . . .227 Tadarida . . . 136-137 tamei, Periscyphis . . . . .217 Taphozous . . 323,326,328,333 tarangensis, Spherillo . . . .231 tardus, Mica ...... 197 tardus, Porcellio ..... 197 taschkententis, Protracheoniscus . . 207tegulatus, Euplotes .... 101, 124 temminckii, Scotophilus . . . . 174 tendanum, Armadillidium . . .212 tenuipunctatus, Armadillo . . .221 Testacea .... 103-112, 2 Pis. testacea, Tura ..... 204 Tetrahymena . . . . . -97 thetidis, Campylaspis . . . 250,255thompsoni, Campylaspis . . . 250,252thompsoni, Platycyamus . . . 70-71thomsoni, Cyamus . . . . .70 thonglongyai, Craseonycteris . 301-336Thyropteridae . 318,319,320,326,327,330 Tura ....turkanae, Haplochromis . . 204 141, 149, 150-153,155.i60 tigris, Armadillidium Tilapia . tirolense, Armadillidium tomentosus, Buddelundia Toradjia torulosa, Campylaspis tosaensis, Hermanites trachealis, Porcellio . Trachelipidae . Trachelipus .... Tracheoniscus tradouwi, Bethalus . translucida, Merulana translucida gracilior, Merulana . translucidus. Armadillo transmutatus, Porcellio transsilvaticus, Cylisticus . transvaalensis, Polyacanthus travancoria, Nagara. travancorius, Nagurus Trematocranus triangulifera, Uramba triarticulatus, Periscyphis. triarticulatus, Periscyphops tricirratus, Euplotes affinis trifolium, Armadillo. trilobatus, Porcellio . Trinema. .... triplicata, Campylaspis trisulcatus, Euplotes trivialis, Periscyphis trochilirostris, Campylorhamphus truncorum, Armadillo truncorum, Venezillo . 212148, 149. 212• 223. 242250,255, 295-296 • 203191,205-207, 239-240 . 207. 207 . 222 . 228 . 228 . 228 203. 192. 229 205 205 . 160 205 . 218. 218113,119,123 . 221 2O3. 103 • 250,253. IOI . 217 . 42 232 232 tuberculatus, Mesarmadillotubulata, Campylaspistuffraui, Euplotestugelae, Diploexochustunetanum, Armadillidium • 215250,252101, 124 . 227 . 212 umbensis, Campylaspis . . . 250, 253umbricus, Cylisticus plumbeus . . .192undata, Campylaspis .250, 253, 283, 287, 289undulata, Periscyphis . . . .217uniformis, Sunniva . . . . .213uniplicata, Campylaspis . . . 250, 252unisulcata, Campylaspis . . 250,252,263Upupa . 37, 42 Uramba ..... 205,239 uraniponnica, Finmarchinella . . 83,86 valleculata, Campylaspis. 250, 254, 282-284vallombrosae, Armadillidium . . .212vanderhorsti, Haplochromis . . .162vannus, Euplotes . . . . . 101 variabilis, Euplotes . . . . .124 variabilis, Porcellionides .... 204 velifer, Haplochromis . . . -143venetus, Protracheoniscus . . . 207 Venezillo ...... 232 venustus, Armadillo. . . . .232 venustus, Venezillo .... 232 verecundus, Eptesicus . . . -134verecundus, Philetar brachypterus 134-135, T37verrucosa, Campylaspis . . 250,255,297verrucosus, Armadillo . . . .227 verrucosus, Diploexochus . . . .227 versicolor, Armadillidium . . . .212 Vespertilio . . . . 134, 172, 174 vespertilio, Porcellio .... 203 Vespertilionidae 73-82, 167-188, 317, 318, 319, 320, 322, 324, 325, 326, 327, 328,329, 330 violaceus, Porcellio .... 198, 204 virgatus, Metoponorthus .... 197 viridis, Metoponorthus .... 197 viticola, Armadillo . . . . .221 vitrea, Campylaspis . . 250, 252, 263, 285vittatus, Periscyphis . . . .217 vizzavonensis, Porcellio lugubris . . 201 vizzavonensis, Porcellio orarum . .201 vulcani, Tilapia .... 148, 149 vulgare, Armadillidium . . 210,212,221vulgaris, Sturnus . . . . -37 wagleri, Psarocolius .weberi, Armadilloweberi, SpherilloWest, B. J. .Whale-liceWhite-beaked dolphin • 37. 231 • 23195-102, i PI. 59-64, i PI.; 65-7259-64, i PI. INDEX wilsmorei, Armadillowilsmorei, Cubaris .wingatii, Haplochromiswintoni, Laephotis . wintoni, Laephotis wintoniwintoni angolensis, Laephotiswintoni wintoni, Laephotiswoodi, Scotoecuswoodi, Scotoecus albofuscus Xestadillo . 221 . 221 162, 163 73. 74-75, 76, 77,78, 79, 80, 81 • 73 • 73 • 73 176,182, 183175, 176-177 230 yemenensis, Porcellio zachvatkini, Hemilepistuszealandicus, Porcelliozebricolor, Armadillozebricolor, Cubaris .zenkewitchi, Euploteszigzag, Armadillozigzag, Diploexochuszwartbergensis, Diploexochus 349204 194204 221221 I24227227227 17 ANATOMY OF HEAD AND NECKIN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITHCOMPARATIVE NOTES ON OTHER CALLAEIDAE P. J. K. BURTON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. i LONDON: 1974 FRONTISPIECE. Male and female Huias (Heteralocha acutirostris) showing probable feedingmethods. The male is depicted excavating decayed wood by the 'gaping' technique described in the text. 17 JULI9I ANATOMY OF HEAD AND NECK IN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITH COMPARATIVE NOTES ON OTHER CALLAEIDAE BY PHILIP JOHN KENNEDY BURTON Pp 1-48 ; i Plate, 26 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. I LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. i, of the Zoological series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 13 June, 1974 Price £2-30 ANATOMY OF HEAD AND NECK IN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITH COMPARATIVE NOTES ON OTHER CALLAEIDAE By PHILIP J. K. BURTON CONTENTS PageI. INTRODUCTION ......... 3 II. METHODS AND MATERIAL ....... 5 III. SKULL AND LIGAMENTS ........ 6 IV. JAW MUSCULATURE ........ l6 V. BUCCAL CAVITY AND GLANDS ....... 24 VI. TONGUE APPARATUS ........ 24 VII. NECK AND NECK MUSCULATURE ...... 28 VIII. OTHER CALLAEIDAE ........ 34 IX. FUNCTIONAL ASPECTS ........ 36 X. SEXUAL DIMORPHISM ........ 40 XI. CONCLUDING REMARKS ........ 44 XII. ACKNOWLEDGEMENTS ........ 46 XIII. REFERENCES ......... 46 SYNOPSIS Spirit specimens and skeletons of the Huia (Heteralocha acutirostris) have been used in adetailed study of feeding adaptations in this extinct species. The Huia showed remarkablesexual dimorphism in bill form, and particular attention is paid throughout to differencesbetween male and female. Comparison is made with the other members of the Callaeidae -Creadion carunculatus and Callaeas cinerea. The features studied indicate that the Huia washighly specialized for feeding by 'gaping' in decayed timber, in order to gain access to beetlegrubs. The full extent of sexual dimorphism revealed by this study is reviewed, and its sig-nificance is discussed. I. INTRODUCTION THE extinct Huia (Heteralocha acutirostris) of New Zealand is frequently quoted asan example of extreme sexual dimorphism among birds. This dimorphism is shownmost conspicuously by the bill, which in the male was typically fairly straight,moderately long, and tapered evenly to a point; while in the female it was muchlonger, more slender, and strongly downcurved. There is little information on theextent of dimorphism beyond this. Phillipps (1963) mentions plumage differencesand a tendency towards stronger claws in older males, and Selander (1966) givesbill, wing and tarsus measurements for a small sample of each sex. The anatomicalstudy of the Huia by Garrod (1872) concentrated principally on establishing theaffinities of the species, and did not inquire closely into its sexual dimorphism. Theproblem of its relationships is still not fully resolved beyond the fact that the Huiais closely related to two other New Zealand species-the Saddleback (Creadion 4 P. J. K. BURTON camnculatus) and the Kokako (Callaeas cinerea) (Stonor, 1942). The three togetherform the family Callaeidae, placed between the Dicruridae and Grallinidae by Mayrand Green way (1956) ; close affinity with the Sturnidae has been suggested in thepast (Garrod, 1872 ; Gray, 1870). The present investigation has been concerned with the anatomical basis of theHuia's feeding specializations. In particular, I have attempted to discover thefull extent of sexual dimorphism in feeding structures, and to correlate this wherepossible with bill form and feeding methods. It is thus a study of adaptation,unusual in that the principal forms to be compared are the two sexes of a singlespecies. The material on which the investigation is based is contained in the aviananatomical collections of the British Museum (Natural History). These consist ofsix spirit and one osteological specimens of the Huia, one spirit specimen of theSaddleback and four spirit specimens of the Kokako. It is thought that the Huiaspirit specimens may be the only fluid-preserved examples of this species in existence,although there is at least one skeleton in New Zealand (Phillipps, 1963) and anotherin the U.S.A. (Bock, igGoa.). The history of the Huia's disappearance, and many aspects of its life and habits,are summarized by Phillipps. Last reported alive in 1907, it is probable that somesurvived after this date, and Phillipps entertains the possibility that a few may stillexist. Its range was restricted to the mountainous Wellington Province of NorthIsland, New Zealand. Here, the combination of a mild climate and heavy rainfallhas produced densely forested country with a high incidence of timber decay andepiphytic growth. The Huia fed principally on insects, varied with some vegetablematter, but by far the most important prey appears to have been the larvae of theHuhu beetle, Prionoplus reticularis (Cerambycidae). This large beetle is commonin many parts of New Zealand, and its larvae, found in decaying timber, may reacha length of 70 mm and a width of 20 mm. The most detailed account of the Huia'smanner of extracting these is provided by Buller (1888). This account has beenoften quoted, but because of its relevance to the present study, it seems essentialto do so again. Buller's description refers to a pair of captive Huias which at firsthe fed on individual huhu grubs : 'On offering one of these to the Huia he would seize it in the middle, and, at once transferringit to his perch and placing one foot firmly upon it, he would tear off the hard parts, and thenthrowing the grub upwards to secure it lengthwise in his bill, would swallow it whole . . . Theyseemed never to tire of probing and chiselling with their beaks. Having discovered that thecanvas lining of the room was pervious, they were incessantly piercing it, and tearing off largestrips of paper, till, in the course of a few days, the walls were completely defaced. But what interested me most of all was the manner in which the birds assisted each other intheir search for food, because it appeared to explain the use, in the economy of nature, of thedifferently formed bills in the two sexes. To divert the birds, I introduced a log of decayed woodinfested with the huhu grub. They at once attacked it, carefully probing the softer parts withtheir bills, and then vigorously assailing them, scooping out the decayed wood till the larva orpupa was visible, when it was carefully drawn from its cell, treated in the way described above,and then swallowed. The very different development of the mandibles in the two sexes enabledthem to perform separate offices. The male always attacked the more decayed portions of thewood, chiselling out his prey after the manner of some Woodpeckers, while the female probedwith her long pliant bill the other cells, where hardness of the surrounding parts resisted the ANATOMY OF HEAD AND NECK IN THE HUIA 5 chisel of her mate. Sometimes I observed the male remove the decayed portion without beingable to reach the grub, when the female would at once come to his aid, and accomplish with herlong slender bill what he had failed to do. I noticed, however, that the female always appro-priated to her own use the morsels thus obtained.' An additional reference to the different feeding techniques of male and female isthe note by J. M. Wright quoted by Oliver (1955): 'I have watched them in pairs hunting for wetas.* The male would tear away at the outerpart of a green sapling. The female then tried to retrieve the weta with her long slender bill.If not successful she would stand back while the male tried to enlarge the hole. Unless thetree was a maire the birds would generally succeed and would then fly away with the weta.1 Phillipps (1963) also provides the most detailed information available on theextent of dimorphism ascertainable from skins, and on the range of variation inthe two sexes, based on 119 specimens in New Zealand museums. Young birdsshow little difference in size and shape of the bill. Some male bills were more curvedthan usual, but were normally deeper than those of females. The male bills reachedan extreme length of 59-60 mm (from feathers), and females up to 104 mm. Phillippsmentions two doubtful cases of birds with bills 63 mm long which he has treated asmales, though they may be females. The skin collection of the British Museum(Natural History) includes two doubtful individuals ; one, with a bill length of76-4 mm is labelled as a male ; the other, labelled as a female, has a bill of 71-8 mm.Excluding these, bill length data for other specimens in the collection may besummarized as follows : Males : Mean 59-7 ; Min. 54-7 ; Max. 65-0 (16 specimens).Females : Mean 96-3 ; Min. 87-1 ; Max. 108-0 (7 specimens). Buller (1878) also gives details on the range of bill form in the Huia and figuresseveral specimens, including a female with the upper jaw much longer than the lower,recalling some species of Hemignathus (Drepanididae). II. METHODS AND MATERIAL Most of the observations reported here were obtained simply by dissection usinga stereomicroscope at powers mainly in the range X2| to X20. The main com-plicating factor was the irreplaceable nature of the Huia specimens, and becauseof this, it was thought prudent to keep dissections of this species to a minimum. Ingeneral, dissections have been limited to the left side, and as little structure aspossible has been removed. Because of its history, the single specimen of Creadioncarunculatus was regarded as of similar value, and no dissections were attemptedwhich would have required removing or severing structures. For convenient reference, the Huia specimens were given a simple individualcoding in addition to their British Museum (Natural History) Registration numbers. * Orthoptera, Gryllacridoidea. 6 P. J. K. BURTON Details of the specimens of Callaeidae used, with their conditions at commencementof the study, are as follows : Heteralocha acutirostris $ i. Reg. No. A 1973.1.3. Intact spirit specimen. <J 2. Reg. No. A 1973.1.2. Intact spirit specimen. (£3. Reg. No. 1940.12.8.107. Intact spirit specimen. $4. Reg. No. 81.1.17.91. Complete skeleton specimen. $ A. Reg. No. A 1973.1.4. Spirit specimen with skull almost completely skeletonized. Rhamphotheca of upper jaw retained (but detached), tongue, hyoid muscles, M. depressor mandibulae and all neck muscles intact.$ B. Reg. No. A 1973.1.1. Spirit specimen with cranium and upper jaw completely skeletonized, removed and maintained as dried specimen. Upper jaw broken and rhamphotheca missing. Lower jaw with rham- photheca intact, detached, but retained in spirit. Tongue, hyoid muscles and all neck muscles intact.$ C. Reg. No. 1940.12.8.108. Intact spirit specimen. Creadion carunculatus Reg. No. 1845.2.21.392. Spirit specimen with skull detached and skele-tonized except for M. depressor mandibulae which is intact. Rhampho-theca of both jaws missing. Tongue, hyoid muscles and all neck musclesintact. This specimen is believed to have been collected on CaptainCook's third voyage (Burton, 1969). It is probably the oldest survivingspirit specimen of any bird, but is in good condition. Callaeas cinerea Reg. Nos. A 1973.1.5 and A 1973.1.6. 2 intact spirit specimens. Various spirit, skeleton or skin specimens of birds in other families were alsoexamined for comparative purposes. The species involved are mentioned in thetext or in the tables. III. SKULL AND LIGAMENTS The skull of a male Huia was described and figured by Garrod (1872), who calledattention to some of its conspicuous features. A short description of the skull isalso given by Oliver as an appendix to the book by Phillipps (1963). The skulls ofmale and female specimens are depicted in Figs. 1-3. It will be noted that with therhamphotheca removed, the disparity between the sexes in jaw form appears muchless than in skin specimens ; this is due to the much greater development of therhamphotheca in the female. Striking features of the skull in both sexes (thoughmore marked in the male) are the prominent occipital crest, well-developed exoc-cipital process and very long retroarticular process. Less obvious, but equally ANATOMY OF HEAD AND NECK IN THE HUIA FIG. i. Heteralocha acutirostris, skull in lateral view. A. Male (<$ 4). B. Female (? A). For abbreviations see p. 47. unusual, is the position and form of the nasal-frontal hinge. The nasals extendbackward in the form of a rounded spur, reaching a point roughly level with themiddle of the orbits. The nasal-frontal hinge is consequently situated much furtherposteriorly than in typical passerines ; the drawings of Creadion and Callaeas (Figs.4-9) show the normal condition. Quantitative data for the specimens studied are summarized in Table i. Relativevalues for the various measurements taken have been assigned by giving for each itsratio to length of the jugal bar. Several other ratios are also included. In theabsence of weight data, sternum length is used as a crude index of body size(Amadon, 1943 ; Zusi, 1962) ; measurements of wing, tail and tarsus are also given(R to U). P. J. K. BURTON Pi 20mm FIG. 2. Heteralocha acutirostris, skull in dorsal view.A. Male (<$ 4). B. Female ($ A). For abbreviations see p. 47. ANATOMY OF HEAD AND NECK IN THE HUIA Pi 20mm FIG. 3. Heteralocha acutirostris, skull in ventral view.A. Male (^ 4). B. Female ($ A). For abbreviations see p. 47. P. J. K. BURTON 10mm FIG. 4. Creadion carunculatus, skull in lateral view. 10mm FIG. 5. Creadion carunculatus, skull in dorsal view. ANATOMY OF HEAD AND NECK IN THE HUIA ii 10mm FIG. 6. Creadion carunculatus, skull in ventral view. 20mm FIG. 7. Callaeas cinerea, skull in lateral view. 12 P. J. K. BURTON Oor EEo CM J ANATOMY OF HEAD AND NECK IN THE HUIA FIG. 10. Skull of Heteralocha acutirostris showing the method of taking variousmeasurements given in Table i. For key see below. The methods by which skeleton measurements were taken are as follows :A Upper jaw length : Fig. 10. B Upper jaw with rhamphotheca removed : Fig. 10.C, D Lower jaw : tip of mandible to posterior tip of retroarticular process, with and without rhamphotheca.E Retroarticular process : measured from the posterior edge of the lateral condyle of the quadrate.F Bill depth : measured at right angles to the tomia, level with the junction of nasal and jugal bars. Only given for specimens with rhamphotheca intact on both jaws. G Bill width : measured at the same point as F, with or without rhamphotheca.H Cranium length : Fig. 10./ Cranium breadth : the greatest breadth of the cranium (just posterior to the orbits).K Anterior cranium breadth : the maximum breadth of the cranium anterior to the orbits.L Orbit length : Fig. 10.M Jugal bar : Fig. 10.N Quadrate : from the lateral extremity of the articulation with the jugal bar, to the medial tip of the orbital process.0 Pterygoid : from the anterior to the posterior extremities visible in antero- lateral view.P Tongue length : from the anterior tip to the tips of the postero-lateral papillae.Q Sternum length : from the midpoint of the posterior margin to the antero- ventral tip of the keel. The postorbital, internal jugo-mandibular and occipito-mandibular ligaments arepresent. Strong connective tissue running from the postorbital process to theectethmoid process across the ventral lateral side of the orbit amounts to an ill-definedsuborbital ligament. There is no external jugo-mandibular ligament. The occipito-mandibular ligament is extremely stout, and much of its central region is ossified,more so in male specimens. The internal jugo-mandibular ligament contains two 14 P. ]. K. BURTON 6org-cx-o+6i H SF y> o N O P H y> b Jo s , «wt?77»3 (M °,w a b >o «!t c- p 'ro M -^ p ^o" &ot it t j'81 1 ro ro l l Tj- 10 cb ^ UOlpVUQ N •2- J^ ^2- SU9UII09ds £ JO UB9J\[ O b 00 «o in ^t- v ^N (N 'fO>O M V1{OOlV49iaH ^3 ^~ >ti o o IH ^^ ^ro H ^ 0 X T3 ! SU9UIID9ds £ JO tTE9J^ b iO ^ yi 00 ro oo" O Tf b oo 04 ro 0 IHM co" cob 9 M "t^ P ^ 00 M — ^ "0? ^ 01 IN {H O o ot Q\ P b OI f^ ro JM »o M VilOOJV^SJSpl 00 ^ro "^ ^ O M ^^ ^ro M ^. O 1 P! ^oo 'b" iO "ro 1) a. 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J. K. BURTON sesamoids, one placed laterally and the other posterior to the quadrate. In commonwith all passerines so far studied, the lower jaw has no medial brace (see Bock,I96ob). IV. JAW MUSCULATURE A work of major importance for studies of avian jaw musculature is Lakjer's(1926) review of the trigeminal musculature. For passerines in particular, thepaper by Fiedler (1951) provides valuable clarification. Investigations by Bock(i96oa and MS) have resolved many remaining problems of structure and ter-minology within the order, and his nomenclature for the subdivision of complexmuscles is the one followed here. The jaw musculature of the Callaeidae remainsundescribed beyond the comments of Garrod (1872) and Lowe (1938) on the largesize of the 'digastric muscle' (= M. depressor mandibulae). In describing the structure of complex muscles it is often useful to designate theirvarious aponeuroses by letters or numbers. It is far from clear to what extent theseaponeuroses can be homologized between different groups, although the work ofStarck and Barnikol (1954) has made progress in this respect for M. adductormandibulae externus. In the present case, a simple numbering system has been usedwhere necessary, but is not intended to imply homology with any numbered apo-neuroses described from other species, with the exception of Aps. i, 2 and 3 in M.adductor mandibulae externus. M. adductor mandibulae externus This important and complex muscle acts to raise the lower jaw and to maintaina grip on objects held in the bill. Its architecture is intricate, and the relationshipsof its aponeuroses can be better understood by the examination of cross sections(Fig. 18). The abbreviation 'M.a.m.e.' is used in parts of the description to shortenthe otherwise unwieldy names of some subdivisions. The three major aponeurosesdescribed by Starck and Barnikol (1954) in a wide range of birds of several orderscan be recognized with little difficulty in Heteralocha, and their numbering here(Aps. i, 2 and 3) is the same as Starck and Barnikol's. M. adductor mandibulae externus rostralis The most dorsal part of M. adductor mandibulae externus, with an extensivefleshy origin on the cranium. Insertion is made principally via Ap. i, which nar-rows anteriorly to form a flat tendon attached to a crista on the surangular. Follow-ing Bock, three sections are recognized : M.a.m.e. rostralis medialis. Origin is from the postero-lateral border of the orbit,medial to the postorbital process, and from a vertically oriented aponeurosis (Ap. 4)attached to the orbit along the medial border of the muscle. Insertion is made viaAp. i. Medially, a group of fibres from the ventral part of the muscle overlaps partof Ap. i as a more or less distinct slip. M.a.m.e. rostralis temporalis. Origin is fleshy from the border of the temporalfossa ; a short aponeurosis (Ap. 5) runs forwards from the tip of the postorbital ANATOMY OF HEAD AND NECK IN THE HUIA Mdt Mdm L po FIG. ii. Heteralocha acutirostris, male (<$ i), jaw muscles and ligaments.For abbreviations see p. 47. process across the dorsal lateral surface, and in some specimens extends onto thedorsal surface of M.a.m.e. rostralis medialis. The fibres of this section convergeanteriorly, falling into an ill-defined bipinnate arrangement, and are attached to theventro-lateral surface of Ap. i ; a few fibres overlap its dorsal surface posteriorly, butit is exposed for some 8 mm from the insertion. M.a.m.e. rostralis lateralis. This section lies superficial to the dorsal part ofM.a.m.e. ventralis. Its origin is from the lateral edge of the zygomatic process, andfrom the lateral edge of Ap. 2 in its narrow posterior region. Fibres run upwardand medially from the origin and insert on the lateral surface of the mandible overa narrow zone dorsal to the insertion of M.a.m.e. ventralis. This insertion is fleshy,and via a superficial aponeurosis (Ap. 6) which merges medially with Ap. i. Theanterior ventral part of this section lies medial to the jugal bar and is in consequenceflattened into a relatively thin sheet of fibres. M. adductor mandibulae externus ventralis Origin is from the zygomatic process, by a strong aponeurosis (Ap. 2) whichtraverses the gap between the cranium and mandible as a fairly narrow tendon, thenfans out over the lateral surface of the muscle. Fibres arising from this aponeurosismake an extensive fleshy insertion on the lateral surface of the mandible, spanningthe fenestra in the posterior part of the mandible. M. adductor mandibulae externus caudalis The shortest section of M. adductor mandibulae externus. Origin is from theotic process of the quadrate and from the ventral surface of the posterior half ofAp. 2. The origin is fleshy, and by a weak aponeurosis (Ap. 7). Insertion is on thedorsal lateral edge of the surangular, posterior and ventral to that of Ap. i. Theinsertion is made by a short, narrow aponeurosis (Ap. 3), which bifurcates within i8 P. J. K. BURTON the body of the muscle to form a dorsal (Ap. 3a) and a ventral (Ap. 3b) branch.Ap. 7 passes between 3a and 3b before fading out. The muscle shows multipinnatestructure, clearest in its middle third, and most conspicuous about Ap. 3a. Fibresrun forwards from Ap. 2 to Ap. 3a, back from Ap. 3a to Ap. 7, and forwards againfrom Ap. 7 to Ap. 3b. M. pseudotemporalis superficialis A muscle of relatively simple structure, whose action is adduction of the lower jaw.The origin is a broad fleshy one on the posterior wall of the orbit, immediatelymedial to M.a.m.e. rostralis medialis. There is also a weak dorsal aponeurosisattached to the orbit. Insertion is made via a strong tendon attached to the medialside of the surangular, ventral to the ramus mandibularis of the trigeminal nerve. Mdt Md m L po FIG. 12. Heteralocha acutirostris, female ($ C), jaw muscles and ligaments.For abbreviations see p. 47. M. pseudotemporalis profundus This muscle is an adductor of the lower jaw, and by its action on the quadrate italso functions to lower the upper jaw. It takes its origin from the dorso-lateralsurface of the orbital process of the quadrate, and from a strong aponeurosis attachedto the expanded tip of the orbital process. It makes an extensive fleshy insertionon the medial surface of the mandible, overlapping slightly onto its dorsal edge,adjacent to Ap. i of M.a.m.e. rostralis. M. adductor mandibulae posterior The actions of this muscle are similar to those of M. pseudotemporalis profundus.Due to its situation closer to the mandible-quadrate articulation, it has a lowermechanical advantage, but its upward component may be of value in holding themandible against the quadrate. M. adductor posterior is contiguous with M. ANATOMY OF HEAD AND NECK IN THE HUIAMptvm Mptdm Mpsp Mpq1 Mamerm Mamert M a m ev M amec FIG. 13. Heteralocha acutirostris (<$ i), jaw muscles in lateral view.For abbreviations see p. 47. pseudotemporalis profundus at its origin, which is also a fleshy one on the quadrate,occupying the base of the orbital process and the dorsal surface of the quadrate body.The two muscles are separated (following the criterion used by Lakjer, 1926) byN. pterygoideus. The muscle inserts on the expanded region of the dorsal surfaceof the mandible just anterior to the base of the internal process, and posterior to theinsertion of M.a.m.e. caudalis. The insertion is fleshy and via a weak dorsal apo-neurosis. M. pterygoideus This important muscle complex acts both to lower the upper jaw and to raise thelower. Its attachments to the palatine and pterygoid are arbitrarily regarded as theorigin, and attachment to the lower jaw as the insertion. M. pterygoideus dorsalis lateralis This section has an extensive fleshy origin on the dorsal surface of the palatine.It inserts on the medial surface of the mandible, posterior to M. pseudotemporalisprofundus, fleshily, and by a superficial aponeurosis (Ap. i), which is strongestventrally. M. pterygoideus ventralis lateralis Origin is from an aponeurosis (Ap. 2) which is attached to the postero-lateral tipof the palatine, and extends for some distance across the ventral surface of the muscle.The medial edge of this aponeurosis serves to demarcate it from M. pterygoideus 20 P. J. K. BURTON ventralis medialis seen in ventral view. M. pterygoideus ventralis lateralis is at-tached at the base of the internal process of the mandible, ventral to the insertionof M. pterygoideus dorsalis lateralis. There is no overlap (Venter externus') ontothe ventral edge or lateral surface of the mandible as in many birds, since the ventralmedial edge of the mandible in this region is occupied by a forward extension of M.depressor mandibulae. M pt v m M pt d m aM pt d m pMpt r Mptdl Aplame Mpsp Mamerm Mamert5mm FIG. 14. Heteralocha acutirostris (£ i), jaw muscles in dorsal view.For abbreviations see p. 47. M. pterygoideus dorsalis medialis In dorsal view, this muscle is seen to lie immediately adjacent and posterior toM. pterygoideus dorsalis lateralis, separated from it by a narrow groove which widensmedially, exposing a small area of the dorsal surface of M. pterygoideus ventralismedialis. Its origin is confined to the pterygoid, and occupies much of its surface.The muscle falls into anterior and posterior portions, whose fibres diverge in aposterior direction roughly along the lines of the pterygoid, so that it appears bi-pinnate in dorsal view. M. pterygoideus dorsalis medialis anterior originates more laterally on the ptery-goid, and inserts narrowly on the mandible, immediately caudal to M. pterygoideusdorsalis lateralis. The insertion is fleshy and aponeurotic, the aponeurosis (Ap. 3)being strongest ventrally. M. pterygoideus dorsalis medialis posterior originates medially on the pterygoid,and fans out to a wide fleshy insertion on the dorsal part of the internal process ofthe mandible, dorsal to that of M. pterygoideus ventralis medialis. ANATOMY OF HEAD AND NECK IN THE HUIA M pt v m Eus Mpt r Ap 3a d m M d m 5mm FIG. 15. Heteralocha acutirostris (<$ i). M. pterygoideus and M. depressor in ventral view.Left side of palate left intact to show papillae. For abbreviations see p. 47. M. pterygoideus ventralis medialis There is an extensive fleshy origin on the ventral surface of the palatine andinsertion is made on the internal process of the mandible, fleshily, and by a strongdorsal aponeurosis (Ap. 4). The fibres of this section are orientated more nearlyparallel to the skull axis than those of any other part of M. pterygoideus. On themedial side of the muscle is a slip (M. pterygoideus retractor) exclusively specializedfor retracting the palatal apparatus, attached at one end to the medial caudal tipof the palatine, and at the other to the basitemporal plate of the skull. A few of itsmost medial fibres are attached to the edges of the opening of the Eustachian tube. M. protractor quadrati et pterygoidei This muscle acts to raise the upper jaw by moving the palatal framework andjugals forwards. It has a wide fleshy origin from the posterior part of the interorbitalseptum and the adjacent region of the posterior orbital wall. The lateral borderof the muscle lies immediately adjacent to M. pseudotemporalis superficialis. 22 P. J- K. BURTON Two parts can be distinguished. The more anterior and medial part (M. protractori), arising mainly on the interorbital septum, shows a bipinnate fibre arrangement.Its raphe is a strong aponeurosis attached to a spur on the posterior dorsal tip of thepterygoid, immediately adjacent to the quadrate. This raphe serves as the principalsite of insertion for the fibres of M. protractor i. The lateral part (M. protractor 2) originating from the posterior wall of the orbitinserts fleshily, and by a weak dorsal aponeurosis on the caudal edge of the basal halfof the orbital process. Mptdl MptdmaMpss Mpql MamermM pt d m p / Ma mert M d m Mamev Ap2ame Mamp Mamec 5mm FIG. 16. Callaeas cinerea, jaw muscles in lateral view. For abbreviations see p. 47. M. depressor mandibulae This muscle, which depresses the lower jaw, is of enormous bulk (and, presumably,fibre number) in Heteralocha relative to the other jaw muscles. There is a widefleshy origin on the squamosal, parietal and exoccipital. Fibres originating high onthe cranium converge to insert fleshily near the extremity of the long retroarticularprocess, mainly on its medial side. Those originating lower, on the exoccipital, fanout to insert fleshily on both surfaces of the retroarticular process. Fibres insertingmedially extend far forward beyond the retroarticular process along the ventral halfof the mandible, ventral to the insertion of M. pseudotemporalis profundus. Thisextension is here termed the pars anterior of M. depressor mandibulae. ANATOMY OF HEAD AND NECK IN THE HUIAMptdl Mptdma Mptdmp Mpq1 Mpss M a merm M a m e v M a m e rt Mdm 5mm FIG. 17. Callaeas cinerea, jaw muscles in dorsal view. For abbreviations see p. 47. M.pseudotemporalis profundus is concealed by M. pseudotemporalis superficialis. 2mm FIG. 18. M. adductor mandibulae externus, TS at about the midpoint of the otic process ofthe quadrate, and at right angles to it. Numbering of aponeuroses as explained in text.A - Callaeas cinerea. B - Heteralocha acutirostris (<$ i). The left side is lateral in bothdiagrams. The following aponeuroses can be distinguished : Ap. i. This is attached to the cranium around the dorsal edge of the origin on thesquamosal and parietal, and extends some way across the lateral surfaceof the muscle. Ap. 2. A strong aponeurosis with a narrow attachment at about the middle of theposterior border of the auditory meatus, fanning out across the lateralsurface of the muscle, and acting as a surface of origin for fibres insertingon the anterior half of the lateral surface of the retroarticular process. 24 P. J- K. BURTON Ap. 3. A strong aponeurosis attached to the occipital crest, which marks thepostero-ventral limit of the origin. Ap. 3 extends some way across themedial surface of the muscle, and becomes stronger on the exoccipitalprocess, whence it is produced as a strong raphe (Ap. 3a) across theventral region of the muscle, and on into the pars anterior. Fibresarising from its medial side insert mainly on the medial surface of theinternal process of the mandible. Those running from its lateral sideinsert on the medial surface of the basal half of the long retroarticularprocess, on the posterior face of the internal process at its base, and onthe ventral medial surface of the mandible. Ap. 4. An aponeurosis attached to the dorsal edge of the internal process of themandible, extending across much of the dorsal surface of the short partof the muscle between this and the exoccipital. This aponeurosis isbranched and infolded within the muscle. Ap. 5. An aponeurosis attached to the dorsal edge of the retroarticular process,but concealed from lateral view by the thin sheet of fibres originatingfrom Ap. 2, and inserted on the lateral surface of the process. V. BUCCAL CAVITY AND GLANDS Backwardly directed horny papillae are abundant on the surface of the palate.They are longest where they fringe the choanae, and in a densely clustered transverserow posterior to the internal opening of the Eustachian tubes. The surface of thelarynx is also papillate, the papillae being longest at its posterior margin. Three pairs of salivary glands are present. The Gl. angularis oris lies on the sideof the head immediately below the skin and just ventral to the jugal bar ; its ductopens at the angle of the gape. A second gland, which may be termed a Gl. palatinae(see Antony, 1920), lies between M. depressor mandibulae, pars anterior, and M.pterygoideus ventralis lateralis, immediately below the mucosa of the palate. Thethird gland is a sublingual one. It lies immediately lateral to M. genioglossus, andextends from the level of the anterior border of M. mylohyoideus to the region ofthe tongue base. Its duct runs alongside M. genioglossus, but diverges from itnear the mandibular symphysis to open in a lateral position. VI. TONGUE APPARATUS The tongue is narrow, tapering evenly towards its anterior tip which is somewhatfrayed and brush like (Fig. 19). The lateral edge bears a few papillae posteriorly,and the posterior edge is crowded with pointed papillae, longest laterally. Tonguelength for the specimens examined is shown in Table i. The tongue skeleton is similar to that of many passerines. The paraglossaliawhich provide support for the tongue itself have long posterior processes, andanteriorly they meet and run side by side to a point about one-third of the way fromthe tip of the tongue. The remainder of the tongue is purely corneous. The basihyal ANATOMY OF HEAD AND NECK IN THE HUIA 5mm FIG. 19. Heteralocha acutirostris (<$ i), tongue and larynx in dorsal view. is a slender rod of roughly triangular section, with the apex of the triangle upper-most. The flattened urohyal, expanded at its posterior tip, and the hyoid horns(each consisting of the ceratobranchial posteriorly and epibranchial distally) are ofsimilar form to those of many passerines. The tongue muscles in Heteralocha show few unusual features, and no significantsexual dimorphism was encountered in the dissections. The descriptions given aretherefore brief ones ; illustrations are given in Figs. 21 and 22. More detailedaccounts and background information are given in the general review by George andBerger (1966) and the paper on passerine tongue muscles by Engels (1938). A veryfull description of the tongue muscles of a single passerine species is that by Bock(1972) for the extinct Ciridops anna (Drepanididae). For discussions on function,reference may also be made to the account of wader tongue muscles by Burton (I974)- M. mylohyoideus A thin muscular sheet with a long narrow origin on the medial side of the man-dibular ramus, inserting on a median raphe. M. mylohyoideus lies ventral to thetongue and all the muscles attached to it. M. serpihyoideus Origin is on the occipital plate, medial to the exoccipital process (Bock, igGob,p. 38) and insertion is on a median raphe continuous with that of M. mylohyoideus. 26 P. J. K. BURTON M. genioglossus A narrow, strap-like muscle whose origin is from the posterior edge of the mandi-bular symphysis. The left and right muscles lie side by side near the origin and thendiverge, running along the ventral side of the mucosa of the floor of the buccalcavity, on either side of the position of the tongue. In the region of the basihyal,the fibres of the muscle fan out to insert on the connective tissue and mucosa over-lying the basihyal and its musculature, and, anteriorly, on the posterior process ofthe paraglossa. FIG. 20. Callaeas cinerea, tongue and larynx in dorsal view. M. stylohyoideus A long, narrow muscle originating on the ventral edge of the base of the retro-articular process, and running along the anterior edge of M. serpihyoideus, to inserton the lateral surface of the basihyal, just anterior to M. thyreohyoideus. M. branchiomandibularis (= M. geniohyoideus of many authors) A bulky muscle whose origin is on the medial surface of the mandible, ventralto that of M. mylohyoideus. A broad anterior and narrow posterior position can bedistinguished. The two run parallel to insert on the hyoid horn. The anteriorportion meets the horn from the ventral side, and is twisted around it for some dis-tance before inserting on the epibranchial. The posterior portion meets the anterioron the dorso-medial side, and merges with it. M. ceratohyoideus A thin, weakly developed muscle which was found only in <$ i. It originates onthe hyoid horn, on the ventral lateral surface of the distal tip of the ceratobranchial, ANATOMY OF HEAD AND NECK IN THE HUIA Apm Mthh Mseh FIG. 21. Heteralocha acutirostris ($ B), tongue muscles in ventral view.For abbreviations see p. 47. deep to M. ceratoglossus. The insertion is on a median raphe continuous with thatof M. serpihyoideus and M. mylohyoideus, near the posterior end of the urohyal. M. ceratoglossus A unipinnate muscle whose fibres arise on the surface of the ceratobranchial, andthe anterior end of the epibranchial, and insert on a long lateral tendon which isitself inserted on the ventral surface of the paraglossal, just level with the anteriortip of the basihyal. No fibres insert on the tendon over the region lying alongsidethe basihyal, but just anterior to its attachment to the paraglossal, a small fleshyslip arises. It merges with that from the other side, and together they insert on astrong medial aponeurosis which runs along the ventral side of the paraglossalia, andinserts on the corneous anterior part of the tongue. This slip is referred to by Bock(1972 and MS) as M. hypoglossus anterior. A similar slip in shorebirds was describedby Burton (1974) as M. ceratoglossus anterior ; the term 'M. hypoglossus anterior' isused by Burton (1974) to refer to a quite distinct muscle, apparently absent from 28 P. J. K. BURTON passerines, arising on the posterior tip of the paraglossals and also inserting on themedian aponeurosis. M. hypoglossus obliquus Origin is on the postero-lateral process of the paraglossa. The right and leftmuscles are merged, the whole forming a bulky loop passing ventral to the anteriorthird of the basihyal. Mthh Mtrh M th h M tr I FIG. 22. Heteralocha acutirostris (? C), ventro-lateral view of tongue muscles attached to thecricoid cartilage. Cricoid and urohyal shown stippled. Both left and right Mm.tracheolateralis are visible. For abbreviations see p. 47. M. tracheohyoideus Origin is from the skin of the neck, and insertion on the lateral surface of thecricoid, just below the dorsal origin of M. thyreohyoideus. M. tracheolateralis The muscle originates on the syrinx, and passes along the side of the trachea,broadening anteriorly to insert by two heads on the lateral surface of the cricoid. M. thyreohyoideus This muscle originates from the lateral surface of the cricoid, by a dorsal and aventral slip. The dorsal one is slender, and is attached just below the dorsal edge ofthe cricoid. The ventral slip is broader, and arises between the two heads of insertionof M. tracheolateralis. These slips unite anterior to the larynx and insert on theanterior lateral surface of the basihyal posterior to the insertion of the more slenderM. stylohyoideus. M. ceratoglossus superior (Bock, 1972) was not found in Heteralocha. The glottalmuscles (M. thyreoarytenoideus and M. constrictor glottidis) conform to Bock'sdescription for Ciridops anna. VII. NECK AND NECK MUSCULATURE The thorough review by Boas (1929) has provided the basis for most subsequentstudies of the avian neck and its musculature. Boas concentrated on non-passerines,principally large species, but detailed information on several small passerines isgiven by Palmgren (1949). There is, however, little information on the cervical ANATOMY OF HEAD AND NECK IN THE HU1A Mrcl M co M s co M sp Mil Mrcv Mfcb Mas6-3 Mas7-3 Mas8-5 10mm FIG. 23. Heteralocha acutirostris ($ C), lateral view of superficial neck muscles.For abbreviations see p. 47. muscles of larger passerines comparable in size with the Callaeidae. The majorityof neck muscles perform several actions which interact in a complex way, and theirindividual functions are omitted from the present account in the interest of brevity.Reference may be made to Boas and Palmgren, and to the excellent discussions onneck muscle function by Zusi (1962, 1969). There are 14 cervical vertebrae in Heteralocha, as in most passerines. This totalincludes the two cervico-dorsal vertebrae (13 and 14) which bear movable ribs notarticulating with the sternum, but from a functional standpoint are best treatedwith the neck. The rib on 13 is extremely short, while that on 14 is long, almostreaching the sternum, but lacking an uncinate process. Six pairs of ribs actuallyarticulate with the sternum, as in Cor cor ax, but unlike most passerines which onlyhave 5 ribs attached to the sternum (Beddard, 1898). Boas showed that the neck of birds consists of three sections, distinguishable byboth functional and morphological differences. Section I, the most anterior, canonly be flexed downward, and Section II only upward. Section III can be flexeddownward, and also upward at its anterior end. In Heteralocha, the constitution ofthe segments agrees with Palmgren's division for smaller passerines (although Palm-gren omitted the cervico-dorsal vertebrae from his count). The division is asfollows : Section I, vertebrae i to 4 ; Section II, vertebrae 5 to 9 ; Section III,vertebrae 10 to 14. The vertebrae of Section I (except the atlas) have strong neuralspines and hypapophyses. Those in Section II lack neural spines (except for a weakone on 5) and have no hypapophyses ; they are more elongated than the vertebraeof Sections I and III. The vertebrae of Section III have hypapophyses ; that on 10is weak, but they increase in size posteriorly. The last, and largest, hypapophysisis on 15, the first dorsal vertebra. Vertebra 14 has a strong neural spine. The account below follows the same order and terminology as Palmgren, withmodification in the case of Mm. splenii accessorii and Mm. intercristales. 30 P. J. K. BURTON M. biv enter This muscle arises from the dorsal surface of the aponeurosis of origin of M. spinalisin the region of 13, and inserts on the dorsomedial edge of the occipital deep to M.complexus. The muscle consists of two fleshy bellies linked by a flat tendon ex-tending approximately from 8 to 5. Little variation was encountered. M. spinalis The muscle consists of a series of fleshy slips from the ventral surface of anaponeurosis attached to the neural spines of vertebrae 14 to 18. These insert on theanapophyses of 2 and of 5 or 6 to 13 ; the most posterior of these is feeble and in-distinct in most of the specimens. Variations in the specimens of Heteralocha wereas follows : Slips to both 5 and 6 present: $ A, $ B. Slip to 6 present, slip to 5 absent : ^ i, ^3, $ C. Slip to 5 present, slip to 6 absent: $2. Mm. splenii colli These muscles are a series of slips arising from the lateral surfaces of the neuralspines of 3 or 4 successive vertebrae, and joining the most anterior slip of M. spinaliswhich inserts on 2. The vertebrae of origin in the Heteralocha specimens are : 4 to 7 : <$ i, $'s A, B and C. 4 to 6 : $ 2, ^ 3. Mm. splenii accessorii and Mm. intercristales In order to clarify the relationships between these two rather similar groups ofmuscles it seems desirable to deal with them together. The muscles included underthese headings in Heteralocha are as follows : a. A muscle running from the anterior surface of the neural spine of 14 to thetransverse-oblique crest of 13, and a similar muscle connecting 13 and 12. b. A series of muscles connecting the transverse-oblique crests of successive verte-brae from 13-12 to 6-5. c. Muscles arising on the ventro-lateral surfaces of the neural spines of 5 to 3 andinserting on the transverse-oblique crests of 4 to 2. d. A muscle arising on the dorso-lateral surface of the neural spine of 3 and insert-ing on the anapophysis of 2, and a similar but weaker muscle arising on the neuralspine and medial part of the neural arch of 2 and inserting on the anapophysis of i.That arising on 3 closely resembles Mm. splenii colli. e. A narrow, flat slip arising from the ventral part of the neural spine of 5 andinserting on the anapophysis of 3, somewhat resembling Mm. dorsales pygmaei. According to Palmgren's criteria, groups a and b only should be regarded as com-prising the Mm. intercristales, and groups c, d and e should be treated as Mm. spleniiaccessorii. (In the small species examined by Palmgren the two muscles runningantero-laterally from the neural spine of 3 are inseparable.) However, it wouldseem more consistent with Boas's work to include group c also in Mm. intercristales,and to reserve the term Mm. splenii accessorii for groups d and e. ANATOMY OF HEAD AND NECK IN THE HUIA 31 No noteworthy variations between individuals were found. M. splenius capitis This muscle originates from the neural spine of 2 and inserts on the posteriorsurface of the skull deep to M. complexus and M. biventer cervicis. No variations insiting were found, and there is little indication of cruciform structure (Burton,i97ia). 1 2 3 4 5 6 7 To 15 FIG. 24. Diagram to show arrangement of slips of some ventral muscles in the anteriorpart of the neck in Heteralocha acutirostris (? B). Heavy lines = M. flexor colli brevis.Broken lines = M. flexor colli profundus. Fine lines = anterior slips of M. longus colliventralis. Mm. Pygmaei Origin is from the medial region of the neural arches of 12 to 8. Each muscleinserts on the lateral edges of the transverse-oblique crest of the second vertebraanterior to it except that from 12 which inserts on n. The latter muscle is weak inmost of the specimens and absent in $ i. Mm. interspinales These three muscles connect the well-developed neural spines of 2, 3, 4 and 5, andwere present in all specimens examined. Mm. ascendentes cervicis These muscles arise on the diapophyses of cervical vertebrae up to and including 6.Most consist of two slips inserting on the anapophyses of the second and thirdvertebrae anterior. That from 7 consists of three slips inserting on 3, 4 and 5 in allspecimens, and that from 8 sends an additional slip to 4 in ^'s i and 2. The shortslips arising on 12 and posteriorly are relatively weak, and also showed some minorvariations in siting. The series is continued posteriorly as Mm. ascendentes thoracicis,but those arising posterior to 14 were not dissected. Mm. intertransversarii These laterally situated muscles connect successive vertebrae. The most anteriorare those from 3 to 2, and the most posterior those from 13 to 12. Each musclearises from the anterior surface of the transverse process and inserts on the posteriorsurface of the transverse process of the vertebra in front ; that from 4 to 3 inserts on 32 P. J. K. BURTON the medial surface of the rib of 3, and that from 3 to 2 inserts on the lateral surfaceof the centrum of 2. As far as 6-5, the Mm. inclusi lie deep to Mm. intertransversarii, and closelyassociated with them. Palmgren regards the muscles anterior to this as continuingthe series of Mm. inclusi; Boas and Zusi treat them with Mm. intertransversarii asis done here. The Mm. intertransversarii are multipinnate muscles, traversed by interdigitatingraphes from origin and insertion. The muscles are bulkiest and the number ofraphes greatest in the region from 7 to 10. Up to 9 raphes have been detected in9-8 or in 10-9, the number and situation of raphes showing small individual varia-tions. The muscle connecting 13 and 12 is reduced to a small dorsal slip, whilethose anterior to 5 are also of small size. Mm. inclusi These muscles are concealed by Mm. intertransversarii, and can only be separatedfrom them with difficulty. Each one arises on the anterior surface of the transverseprocess medial to M. intertransversarius, and inserts on the lateral surfaces of theneural arch and centrum of the next vertebra in front. Most show division intodorsal and ventral bellies (Mm. inclusi superiores and inferiores). In the mostposterior two (12 to n and n to 10), only inferiores can be distinguished. Themost anterior are those connecting 5 and 6. M. longus colli ventralis This complex muscle consists of a series of fleshy slips arising on the sublateralprocesses, hypapophyses and anterior part of the centra of vertebrae 15 to 6. Themain part of the muscle inserts by a series of 7 tendons on the ribs of n to 5. Eachvertebra sends a slip to join each of the tendons traversing it; there are thus 7 suchslips from each of vertebrae 15 to 12 after which the number of slips decreases byone for each vertebra anterior to this. In the region 15 to 12 the slips are denselycrowded and difficult to separate and some of the deepest fibres appear to attach toadjacent vertebrae. A smaller group of slips situated anteriorly is also included with M. longus colli.Three of these arise from a tendon attached to the sublateral process of 7, whichalso provides origin for part of M. flexor colli profundus. They insert on ribs 3 and4 by short aponeuroses and on the long tendon attached to rib 5. Slips also ariseon the ventral anterior surfaces of the lateral processes of 7, 6 and 5, the two formerin close association with the Mm. intertransversarii, and immediately below them.These attach on rib 3, and those from 6 and 7 also on rib 4 ; minor variations occur.Two slips arise from the sublateral process of 6 and insert on ribs 3 and 4, and thereis also a longer slip arising on 9 (8 in $ C) which inserts by a weak tendon on rib 4.This tendon also appears to receive some fibres from the posterior part of the muscle. M. flexor colli brevis Lateral and medial parts may be distinguished in this muscle. The lateral partconstitutes the greater bulk of the muscle and originates from the ventral surface ANATOMY OF HEAD AND NECK IN THE HUIA 33 of the lateral strut of 3 ; and from the lateral processes of 4 and 5 in the $'s and 6 alsoin the g's. The medial portion is separated from the lateral by the anterior part ofM. longus colli; it takes origin from the sublateral processes of 3, 4 and 5. The lateral and medial portions join anterior to rib 3 and insert by a tendon onthe postero-ventral processes of the centrum of the atlas. 10 12 13 14 15 FIG. 25. Diagram to show arrangements of slips of some neck muscles in Heteralocha acu-tirostris ($ A or $ B). Heavy lines = M. splenius capitis and Mm. splenii colli. Finelines = M. spinalis (dorsal) and M. longus colli ventralis (ventral). Broken lines = Mm.splenii accessorii (as denned in text). Dotted lines = Mm. pygmaei. Tendons of M.spinalis and M. longus colli ventralis are represented by double lines. M. flexor colli profundus Origin is from the sublateral processes of 4, 5, 6 and 7. The slips from 4 and 5arise immediately deep to the medial portion of M. flexor colli brevis and insert onthe hypapophysis of 2. From 6, slips run to insert on the hypapophyses of 2, 3 and4 (c? 2> $ 3» $ B) or of 3 and 4 only ($ i, $ A, $ C). From 7 a long slip runs to thehypapophysis of 2 ; this slip shares a tendinous origin with the medial slip of M.longus colli from 7 to 3. In $ 2, $ 3 and $ B the slip bifurcates and a branch con-tiunes forwards to fuse with M. flexor colli brevis near its insertion. The slips arisingon 6 show some fusion with M. rectus capitis ventralis at the origin. M. complexus Origin is from the lateral strut of 4, the diapophysis of 5, and from an aponeurosisattached to the diapophysis of 6. Insertion is on the dorsal edge of the occipitals. M. rectus capitis lateralis Origin is from the hypapophyses of 2, 3 and 4, and insertion on the lateral dorsaledge of the exoccipital. M. rectus capitis superior This muscle lies immediately superficial to M. flexor colli brevis. Origin is fromthe lateral surface of neural arch i, from the anterior surfaces of anapophyses 2 and 34 P. J- K. BURTON 3, from the lateral strut of 4, and from the transverse processes of 5 and 6. Originfrom 6 is absent in all the female specimens and in <£ 2 ; origin from 5 is also absentin $ C. Insertion is aponeurotic and fleshy on a ridge at the posterior edge of thebasitemporal plate. M. rectus capitis ventralis Origin is from the ventral surface of i, from the hyapophyses of 2, 3, 4 and 5, andfrom the sublateral process of 6 where there is some fusion with M. flexor colliprofundus. The right and left Mm. recti capiti ventrales are fused in the midlineand insert together on the basitemporal plate, anterior to M. rectus capitis superior. VIII. OTHER CALLAEIDAECreadion carunculatus The description of the Huia's skull by Oliver (in Phillipps, 1963) refers to thegeneral similarity of the skull of Creadion. Oliver notes the greater extent of ossi-fication in Heteralocha, manifested especially at the front of the orbit. The skulland skeleton of this species are figured by Shufeldt (1913), who includes some briefnotes on its skull for comparison with Anthochaera carunculata (Meliphagidae). The following additional points may be noted. The retroarticular process ishighly developed in Creadion, but is nevertheless relatively shorter than that of theHuia ; there is no raised occipital crest, and little development of an exoccipitalprocess. Otherwise, skull proportions are similar to Heteralocha, particularlyfemale specimens, though the quadrate and pterygoid are relatively smaller. Theskull is apparently larger relative to body size in the Saddleback if a rough index(sternum length) from a single specimen can be relied upon. Ligaments have beenremoved from the skull of the available specimen, and could not be studied. With the exception of M. depressor mandibulae, jaw muscles have also beenremoved from the specimen of Creadion. This remaining muscle is, however, ofconsiderable interest, since it possesses a pars anterior as in Heteralocha - a featurewhich otherwise appears to be unique among birds so far studied. The rest of themuscle is also well developed, and similar in structure to that of the Huia, thoughrelatively less massive. The tongue, like the jaws, is smaller relative to the skull inCreadion, but the hyoid musculature is closely similar in both Saddleback and Huia. Due to the historic interest of the specimen, only limited dissection of the neckmuscles could be undertaken for Creadion. Five of the muscles exhibiting variationin the Huia or among other passerines were examined. These showed an overallreduction in the number of sites of attachment. M. spinalis lacks the slip insertingon 5 which is present in three of the Huia specimens, though a slip to 6 (absent inHuia <$ 2) is present. M. splenius colli lacks the slip to 7 found in four of the Huiaspecimens. M. flexor colli brevis and M. rectus capitis superior lack attachment to 6,a feature found in three and two of the male Huia specimens respectively. M. corn-plexus also lacks a slip to 6, although this is present in all six Huias dissected. ANATOMY OF HEAD AND NECK IN THE HUIA 35 Callaeas cinerea The skull and other features of the skeleton in the Kokako have been described insome detail by Stonor (1942). It is sufficient here to draw attention to the extensivedifferences in skull proportions and geometry between this bird, on the one hand,and Heteralocha and Creadion, on the other. The bill is shorter, but much deeper,and the jugal bar meets it at a greater angle. The quadrate is relatively much larger,and is rotated backwards by comparison with the other Callaeidae, so that its oticprocess meets the lower jaw more nearly at right angles. The cranium is shortened,and smaller relative to the orbits ; the jugal is very long relative to the skull. Thelower jaw is deep, and bears strong cristae for the attachment of the adductormusculature. There is no retroarticular process, and the articular is extremelyshallow at its articulation with the quadrate. The prominent sesamoids at thisarticulation were noted by Stonor, and their origin and functions are discussed indetail by Burton (1973). As in the Huia, the external jugomandibular ligament isabsent. The occipitomandibular ligament is not ossified at all. The jaw musculature is notable for the highly developed adductor musculature,and the very small M. depressor by comparison with the other Callaeidae. M.adductor mandibulae externus is considerably more bulky in actual (as well as rela-tive) size than in Heteralocha, with increased number of fibres and greater com-plexity. Its aponeuroses are generally stouter, and more branched and subdividedinternally (Fig. 18), providing extra surface for fibre attachment, and consequentlya greater use of pinnate structure. M. pseudotemporalis superficialis is also muchlarger than in the Huia and Saddleback, and in dorsal view completely conceals M.pseudotemporalis profundus, which is much reduced - a consequence of the back-ward displacement of the quadrate. M. pterygoideus is bulky and the retractorpalatini slip is especially well developed, and prominent in dorsal view. M. depressormandibulae is not only smaller than in the other Callaeidae, but structurally simpler,and entirely lacks a pars anterior. It is unique, however, for the pulley arrangementbetween its aponeurosis and the internal jugo-mandibular ligament (Burton, 1973). The tongue is much broader relative to its length than in the Huia and Saddleback ;its shape is almost rectangular, and the tip brush-like. The paraglossa which sup-port it are widely separated. There is no median aponeurosis and M. ceratoglossusanterior is absent ; otherwise, the hyoid musculature resembles that of the othertwo genera of Callaeidae. The palate is provided with horny papillae similar tothose of the other Callaeidae, but the horny lining of both jaws is developed near thebill tip into raised, papillate bosses. There is a pair of Gil. angularis oris, but a Gl.palatinae has not been found. Neck muscles showing variation in their points of attachment in Heteralocha orother passerines were examined in Callaeas. The two specimens dissected had thesame number and sites of attachment points for the muscles examined. Slips ormuscle components absent in some Huia specimens, but present in the Kokakospecimens, were the insertions of M. spinalis cervicis on 5 and 6 ; M. splenius colliarising from 7 ; M. intercristalis from 5 to 3 ; the slip of M. ascendens from 8 to 4 ;and the slip of M. flexor colli brevis to 6. No Mm. pygmaei arise from 12, thoughthis site is occupied in all but one male Huia specimen. However, an additional M. 36 P. J. K. BURTON pygmaeus from 7 to 6 is present in the Kokako specimens, though absent in all theHuias dissected. The Kokakos lacked the slip of M. flexor colli profundus from 6to 2, present in three of the Huias, and the slip of M. complexus to 6, present in allthe Huias. M. flexor colli brevis in the Callaeas specimens lacks the slip to 6 presentin two male Huias, but has a slip to 5 (unlike Creadion and one female Huia specimen). IX. FUNCTIONAL ASPECTS Jaw mechanism Next to the form and sexual dimorphism of the bill, the most striking features of theHuia's cranial morphology are the huge M. depressor mandibulae and associatedskull modifications - the prominent occipital crest, providing extra surface for itsorigin, and the very long retroarticular process providing increased leverage for themuscle. Clearly these adaptations must permit the lower jaw to be depressed withgreat force. This can only be necessary if it is to be opened against considerableresistance by external forces, and it seems certain, therefore, that the Huia washighly specialized for feeding by 'gaping' or 'prying'. This feeding technique con-sists basically of thrusting the bill into a potentially food-bearing medium (earth,wrood, fruit, etc.) and opening it, to widen the hole and so facilitate exploration orprey extraction. Gaping is a technique described from birds of several families. Among passerines,good examples are furnished by some Sturnidae and many Icteridae. Gapingbehaviour and related modifications of M. depressor mandibulae have been studiedin the Icteridae by Beecher (1951). It is interesting to compare the Huia with otherbirds specialized for gaping, and some figures for relative length of the retroarticularprocess in several examples, mainly of passerines, are given in Table 2. The nearestapproach is shown by Cacicus solitarius in which the retroarticular process is slightlylonger relative to the lower jaw than in the Huia. However, Cacicus solitarius is ashorter billed bird ; relative to skull length, its retroarticular process is shorter thanin Heteralocha, a difference which would be even greater but for the Huia's elongatedskull with enlarged occipital crest. It seems clear that the Huia was very highlyspecialized for feeding by gaping, perhaps more so than any bird now living. Buller's account of the Huia's feeding behaviour is an excellent one, but clearlygives an incomplete picture of the male's excavation methods. Buller remarks thatthe captive male used its bill to 'chisel out' pieces of decayed wood in a woodpecker-like manner. Almost certainly its precise technique must usually have been todrive the bill into the wood and 'gape' to split portions off; this is actually quitedifferent in principle from the methods of woodpeckers which depend purely onblows. Buller states that the female he observed fed in a quite different manner,by probing into relatively hard wood. However, M. depressor mandibulae and theretroarticular process, though somewhat smaller in the female, are still very large bycomparison with other birds, and her capacity for forceful depression of the lowerjaw must also have been very high. Due to the greater length of the female's bill,less force could be exerted at its tip than in the male ; and the more flexible natureof its anterior, purely rhamphothecal, portion, appears rather inefficient for gaping. ANATOMY OF HEAD AND NECK IN THE HUIATABLE 2 Relative length of the retroarticular process in single specimens of various birds showinggaping adaptations, and in all intact Huia specimens used in this study 37 f S I (Si o .a K Upupa epopsPhoeniculus purpureusPhoeniculus aterrimusPsarocolius decumanusPsarocolius wagleriCacicus celaCacicus leucorhamphusCacicus solitariusSturnella magnaAmblyrhamphus holosericeusCreatophora cinereaSturnus vulgarisSturnus contraHeteralocha acutirostris, $ (mean of 3)Heteralocha acutirostris, $ (mean of 2)Pyrrhocorax pyrrhocorax 81-060-4 45'465-272-6 43'i 48-7 53'544'2 45'441-7 43'949-589-4 109-577-0 6-2 4'4 7-87.94-05'210-3 5'58-02-85'0 4'716-4 I3-55'4 25'426-320-5 36-9 37'029-129-831-1 31-3 28-028-829-427-846-8 40-9 0-07o-io O'lOO'I2O'llO'O9O'll 0-19 0*12 0-180-07 O'll 0-090-18 0-07 0'220'24O'2I0'2I0'2I0-140-17 o'330-170-29 O'lO 0-17 0-17 0-310-13 Nevertheless, gaping must also have been an important part of her repertoire offeeding techniques in addition to exploratory probing (as described by Buller) forwhich the decurved bill shape is evidently adapted. In probing timber tunnelledby beetles, it seems feasible that the bill might occasionally be inserted through acrack into a larger cavity, so that gaping could take place with contact only in themore rigid basal part of the bill. However, the possibility remains that the female'sapparent gaping adaptations reflect principally the shared genotype of the two sexes(see under Sexual Dimorphism). In addition to its large size, M. depressor mandibulae in the Huia also showsinteresting structural complexities. A major contribution to the force of depressionis obviously provided by the large mass arising posteriorly and dorsally on thecranium, since this is not only bulky, but has the longest moment arm due to itsinsertion near the posterior extremity of the retroarticular process. However, the 38 P. J. K. BURTON anterior parts of the muscle also exhibit various modifications. Several aponeurosesare present, serving as the basis for pinnate fibre arrangements ; this may be relatedto their short working distance, for which pinnate structure should provide moreforceful contraction than parallel fibred muscles of the same physiological crosssection (Gans and Bock, 1965). The pars anterior of the muscle (present also inCreaction) is of particular interest, since it appears not to have been described pre-viously in any other bird, and was certainly absent from gaping species of otherfamilies dissected during this study. Its functions are hard to surmise, and asatisfactory explanation will probably require a much more detailed knowledge ofthe jaw mechanics involved in gaping than is available at present. Except for fibres inserting on the internal process, the greater bulk of M. depressormandibulae has a medial component in contraction which is greatest for the mostposterior parts of the muscle originating near the midline of the cranium. Theunusual prominence of the medial condyle of the quadrate may in part serve to resistthe stress which this places on the quadrate/mandible articulation. Probably M. depressor also aids protraction of the upper jaw by the action of itsupward force component on the quadrate (see Bock, 1964, 1968, and Zusi, 1962, 1967,for discussions of this mechanism), but M. protractor quadrati et pterygoidei isitself well developed. It is certainly evident that upper jaw action contributessubstantially to 'gaping'. The unusual situation of the nasal-frontal hinge may berelevant in this respect. Its position, more dorsal and posterior than in otherpasserines, places it further from the line of action of the protractor force, actingthrough the palatines and jugals ; the moment arm of this force is consequentlyincreased. The remainder of the jaw musculature, concerned with adduction of the lower jawand depression of the upper jaw, calls for less comment. It is well developed, asseems necessary to provide sufficient grip while extracting huhu grubs, but notremarkably so ; in general features, the muscles other than M. depressor and M.protractor resemble those of Corvus described by Bock (MS). Skull architecture There is a need for detailed and comparative mechanical analyses of the skull in'gaping' passerines of a comparable depth to those available for biting and seedcrushing forms, e.g. Bock (i96oa, 1964^ 1966) and Bowman (1961). Here, somefeatures of possible significance in the skull of Heteralocha will be briefly noted. The wedge-shaped bill of the male Huia is typical of the majority of 'gapers' andresembles that of many Icterids. The straight-sided culmen and gonys are moreefficient than the blunter, arched bill of most passerines for the task of forcing thebill into the substrate prior to gaping. Decurved bills, such as those of the female's,or some wood hoopoes (Rhinopomastus spp.), depend for this on the presence ofexisting crevices or tunnels. In the case of the wedge-shaped bill, the narrower andmore acutely conical its form, the more easily it will penetrate the substrate ; alimit is imposed by the danger of damage by forces directed across the bill axis,since this hazard is less in a more broadly based cone (see Bowman, 1961, p. 222-224).Evidently this danger was not exceptionally high in the Huia, since the male's bill ANATOMY OF HEAD AND NECK IN THE HUIA 39 is narrower and more acute than in many Icterids, including the wood-prying caci-ques ; and in both sexes, the angle between the jugal bars and mandibular rami(giving lateral support to the bill) is no greater than in many unspecialized passerines. The culmen of the bill and frontal bridge of the skull are virtually in line, againa feature of many 'gapers', but particularly well marked in the Huia, with the nasal-frontal hinge shifted to an unusual position nearly halfway across the orbital region.This profile eliminates the difficulties that might arise with a more conventionalskull possessing a marked 'forehead' at the nasal-frontal hinge if the bill were thrustinto the feeding substrate up to or past its base. Many Icterids in fact have theculmen raised above the level of the frontal bridge ; the horny plate extending ontothe forehead of some members of this family (e.g. Gymnostinops, Psarocolius) isprobably adaptive primarily to gaping in succulent fruits (Beecher, 1951). The stoutness of the quadrate-jugal articulation in Heteralocha is probably relatedto the great development of M. depressor mandibulae. The prominence of themedial condyle of the quadrate and its orientation are also unusual; the long axisof the condyle is directed more nearly at right angles to the skull axis than in mostpasserines, a feature which may be adaptive to resisting backward disarticulation bystrong external forces. The nasal-frontal hinge appears to be unusually ill defined in Heteralocha, andmanipulation of skulls relaxed by boiling suggests that the bending zone is con-siderably longer and stiffer than in most holorhinal birds. This may have somesignificance as a safety factor, since a wood 'gaper' could be exposed to some dangerof over-protraction in the event of timber splitting suddenly while a strong force wasstill being exerted by the upper jaw. However, there are no bony protraction stopscomparable to the overhanging frontal 'brow' of woodpeckers - part of a completelydistinct adaptive complex based on the use of M. protractor to distribute compressionforces while hammering (Spring, 1965, Bock, 1966). Buccal cavity, tongue and hyoid musculature Although the Huia's manner of gaining access to its food is specialized, the fooditself (Cerambycid larvae) and the process of swallowing it pose no unusual problems.It is consequently no surprise that the papillae of the upper jaw, tongue and larynxresemble those of many other passerines which feed on invertebrate prey. Thehyoid musculature is similarly unspecialized. Cerambycid larvae are smoothskinned, and would not seem to require a great deal of lubrication. The limitedcomplement of salivary glands is consistent with this supposition. Neck musculature By comparison with the other two species of Callaeidae, the neck musculature ofHeteralocha appears better developed inasmuch as several of the muscles or musclegroups occupy additional sites of origin or insertion. It is difficult, however, todraw general inferences from this, owing to the scarcity of information on otherpasserines. Palmgren's (1949) study, though detailed, concentrated on small speciesof passerines ; Boas (1929) confined his work mainly to non-passerines. In theabsence of such information for larger passerines, it is not possible to know to what 4o P. J. K. BURTON extent this difference is due to the Huia's larger size. For the present it is onlypossible to remark that greater development of the neck musculature would beexpected in a bird so highly adapted to the extraction of prey from timber byvigorous techniques. Other Callaeidae The Saddleback is known to excavate for insect food in decayed timber, bark,epiphytes, etc., and also takes some huhu grubs. It is clear from its skull that inthis species, too, gaping must play a large part in its feeding activities. However, itis less specialized than the Huia, even apart from the lack of pronounced sexualdimorphism. Correlated with its smaller body size, it includes a larger proportionof small insects and other items in its diet, and has never reached the high degree ofdependence on a single prey species seen in the Huia. The Kokako is primarily a fruit and leaf eater. In many features of its cranialmorphology it shows convergence with finches or with parrots. These featuresare centred around the need for powerful adduction for biting hard or tough vegetablefoods. In the skull, the jaws are shortened and deepened, and the quadrate/mandiblearticulation shifted posteriorly. The curious structure of the mandible/quadratearticulation is evidently a consequence of these trends (Burton, J-973a). There is noretroarticular process, and it is doubtful if gaping adaptations were ever present inits ancestry. The adductor musculature is more bulky and complex than in theHuia and Saddleback, and the tongue is specialized, probably for dealing with fruits. X. SEXUAL DIMORPHISM Bill The great difference in size and shape of the bill between male and female Huias isof course well known. It does not appear to have been realized, however, to what alarge extent this is due to differing development of the rhamphotheca. With therhamphotheca removed, the bill length difference is reduced and the skulls of thetwo sexes resemble each other much more closely (Fig. 26). The greatly elongatedrhamphotheca of the female is remarkable not only by comparison with the male,but also when compared with other birds of similar bill form. Table 3 gives figuresfor upper jaw length with and without rhamphotheca in specimens of a variety ofbirds (mainly passerines) with long decurved bills. It can be seen that all of thesefall well short of the female Huia in relative development of the rhamphotheca, andsome bills of very extreme form are bony right to the tip. It is difficult to assess thesignificance of this feature. From a functional viewpoint, it might be suggestedthat the relatively flexible material of the rhamphotheca would be in less danger ofbreakage than bone when used for vigorous exploration of timber. However, developmental and genetic factors should also be taken into considera-tion. The greater development of the rhamphotheca in the female may be a specialcase of allometric growth ; this might be investigated by soft x-ray photography ofthe fairly extensive series of Huia skins in the world's museums. It is also possible ANATOMY OF HEAD AND NECK IN THE HUIA 20mm FIG. 26. Comparative development of the rhamphotheca in male and female of Heteralochaacutirostris (<$ i and $ C). Solid black areas represent the profile of the rhamphotheca. that the answer may lie in the differences between the genetic control of developmentof rhamphotheca and of bone ; it may simply be 'easier', genetically, to evolveextreme sexual dimorphism of bill length in this way. Other examples of thisphenomenon may exist. For example, the lark Alauda razae shows marked sexualdimorphism in bill length (Burton, igyib) and soft x-ray photographs of threespecimens of each sex show proportionately greater rhamphothecal development inthe longer-billed males. Investigation of longer series of this and other sexuallydimorphic species may shed further light on this problem. Close examination of Huia specimens showing features intermediate between maleand female (e.g. some specimens mentioned by Phillipps, 1963) also seems desirable.It may be mentioned in passing that the male specimen figured by Garrod (1872)seems to have been unusual in this respect. Garrod mentions that its tongue was athird the length of the bill. Assuming the tongue was of normal length, this wouldmean a bill some 90 mm long, a length normally found only in females. The figuresshow other unusual features ; the jaw tips are attenuated and slightly decurved,rather as in female specimens, and the occipital crest is less prominent than in themales examined in this study. On the other hand, bill depth and stoutness of thejugal, palatines and pterygoids are characteristically male. The specimen wasacquired by the Zoological Society of London nearly two years before its death 42 P. J. K. BURTON TABLE 3 Extension of rhamphotheca beyond upper jaw tip in single specimens of various birdswith long downcurved bills, and in male and female Huia specimens used in this study Upupa epops Rhinopomastus cyanomelasCampy lor hamphus trochilirostrisFalculea palliataNectarinia famosaArachnothera robustaHemignathus procerusHeteralocha acutirostris, <$ (mean of 3)Heteralocha acutirostris, $ (mean of 2) C Si) -SO -E, 03 63-537-762-066-7 35'254'960-650-2 90-5 o n3 -i-> g i*i o 3 _^ oj — •3 -g a s?-s 2 — 5 **-• T3 <U 03 lu O ri-a I-R-S h-> O V-, 52-3 33'0 55'558-530-8 53'946-8 43'5 49'7 •M ns 5 0-18 O'I2O'lOO'I20-13O'O20'23 o-io 0-45 (Zoological Society of London, 1870) ; possibly it was young when acquired, anddeveloped abnormally in captivity. Jaw mechanism M. depressor mandibulae is of essentially similar structure in both sexes, butdistinctly less massive in the female. Differences in skull features associated withthis are the less prominent occipital crest (and consequently shorter skull), and shorterexoccipital and retroarticular processes of the female. Comparisons between male and female for the other jaw muscles are of verylimited value, since these remain in only one female specimen ($ C). In this speci-men, M. adductor mandibulae externus appears distinctly more bulky than in thethree males, although the medial slip of M.a.m.e. rostralis median's is shorter. Greaterdevelopment of this muscle would seem reasonable in view of the female's greaterbill length, a consequence of which would be to reduce the mechanical advantage ofthe muscle for bill tip adduction - an action which might require considerable forcewhen gripping and extracting huhu grubs. Skull Features related to differences in bill form and in development of M. depressormandibulae have already been mentioned. Apart from these, the proportions of the ANATOMY OF HEAD AND NECK IN THE HUIA 43 skull are very similar in both sexes, as can be seen from Table i. The only significantdifference in proportion is in the region of the bill base. In the female, the bill isrelatively narrower and less deep at the base, and the skull is narrower anterior to theorbits. Probably her mode of feeling involved less exposure to forces directed acrossthe bill axis than the more vigorous techniques of the male. Ratios of jugal lengthto sternum length give no indication of any difference between the sexes in relativehead size. Buccal cavity, tongue and tongue musculature Despite the much greater length of the female's bill, the tongue is of similar sizein both sexes. It would have been of little use for the extraction of prey by thefemale, and in both sexes was probably concerned simply with manoeuvring foodduring the process of swallowing. It is therefore not surprising that the hyoidmusculature shows no obvious sign of sexual dimorphism. The same is true also ofthe buccal papillae and salivary glands, whose functions would have been similarin both sexes. Neck musculature Several of the neck muscles and muscle groups showed variations in the numberand disposition of components. Table 4 compares the distribution of these variablecomponents in the three spirit specimens of each sex, and a fairly clear patternemerges. Variable slips and components of the dorsal muscles (concerned mainly TABLE 4 Distribution of neck muscle components present in some, but not all, of the Huia specimens Number of specimens in which present Dorsal components M. spinalis insertion on 5M. spinalis insertion on 6M. splenius colli origin on 7M. pygmaeu? from 12 to IIM. ascendens from 8 to 4 TOTALVentral components M. flexor colli profundus from 6 to 2M. flexor colli profundus slip joining M. flexor colli brevisM. flexor colli brevis origin from 6M. rectus capitis superior origin from 5M. rectus capitis superior origin from 6 Male i 2 I22 Female 2333o iii io 2O TOTAL 3545 219 3 33 5 2 TOTAL 12 16 44 P. J- K. BURTON with raising the neck) are slightly better represented in the female specimens.However, a much greater disparity exists among the variable components of theventral muscles (concerned with lowering the head and neck), which are much betterrepresented in the male specimens. This discrepancy is entirely consistent with the difference between the feedingmethods of male and female. Forceful downward movements of head and neckmust have been essential to enable the male to insert his bill into timber before'gaping' to split it ; this would have been much less important in the female whorelied on the presence of existing tunnels and crevices into which to insert her bill.The preponderance of variable components of the dorsal musculature in the femaleis less extreme, but may also correspond to a difference in use. Extracting huhugrubs must have entailed exertion in lifting the head and neck for both sexes ; butthis exertion may often have been greater for the female, since a larger proportionof her prey would have to be removed from more or less intact tunnels due to theharder wood in which she fed and her lower capacity for forceful 'gaping'. XI. CONCLUDING REMARKS Several studies of the Huia's morphology have been undertaken in attempts toclarify its systematic position. I would be reluctant, however, to draw any con-clusions about the Huia's affinities from the results of the present investigation. Mostof the foregoing descriptions and discussions have been centred around the extremesof sexual dimorphism and adaptation for gaping shown by the Huia, but neitherphenomenon is unique among passerines except in degree. Indeed several of thefeatures to which Garrod (1872) gave greatest weight in allying Heteralocha with theSturnidae were gaping adaptations and the genera of 'Sturnidae' which he examinedincluded several New World forms now assigned to the quite distinct family Icteridae.Firstly, the size of M. depressor mandibulae (the 'digastric' muscle) which impressedGarrod and also Lowe (1938) with its similarity to various Sturnidae is an unsatisfac-tory character unless considered in conjunction with the structure of the muscle.In this respect, Heteralocha and Creadion both show a feature which is apparentlyunique - a pars anterior, extending forwards from the articulation with the quadrate.Secondly, Callaeas, an undoubted relative (Stonor, 1942) of these two genera, showsno gaping adaptations, and quite possibly did not evolve from a gaping ancestor.It seems reasonable to suggest that the gaping adaptations of the Huia and Saddle-back evolved independently of other families and in isolation, and if this is correct,they are poor evidence for affinity with other gaping forms. The details revealed by this study do not, either, affect the general conclusion ofSelander (1966) that extreme sexual dimorphism in feeding structures is particularlyfrequent in insular bird species, since it provides a means of utilizing a wider rangeof food resources in conditions of reduced competition. However, the dimorphismof the bill in the Huia is the most extreme example known of this phenomenon, andit is natural to wonder how such a large difference between the sexes evolved. Thegaping adaptations may well have played a crucial part. Gaping is an unusual means ANATOMY OF HEAD AND NECK IN THE HUIA 45 of exploiting the food resources of timber, and among passerines in general, probingis a much more common technique. Nevertheless, gaping was almost certainlythe primary adaptation in the stock from which the Huia arose. The existence ofsimilar adaptations in both sexes of the Saddleback lends strong support to thispresumption. If so, the female Huia's bill form appears to be a secondary adaptationfor probing, probably evolved from what was initially a slightly more slender(and very likely shorter) bill than that of the male. An early appearance of a sexualdifference in feeding technique would be essential for this to take place. It shouldbe stressed here that Buller's account gives no evidence for cooperation between thetwo sexes in feeding, although the terms in which it is couched appear to suggest thisand have often been interpreted in this way. Almost certainly each simply fedindependently in the manner for which it was adapted. The female probablyderived occasional benefit from the excavations of the male, but the reasons for theHuia's regular association in pairs may primarily have been social or sexual. Assuming the Huia to be extinct, a full understanding of the significance of itsdimorphism may never be achieved. However, there are still various lines of in-vestigation which might profitably be pursued. Further use could be made of theexisting material of Heteralocha. In addition to the 119 New Zealand specimensexamined by Phillipps (1963), several museums in other parts of the world holdseries of skins. Examination of all this scattered material may prove difficult, butit would seem desirable to have the fullest available quantitative data on the extentof dimorphism and of variation within each sex. The data so far available suggestthat the female's bill may have been more variable than the male's. If true, thismight indicate that females obtained food in a greater variety of situations or bymore versatile techniques than males (although Phillipps suggests that the male atea greater proportion of insects other than huhu grubs). It certainly appears that thepair bond was very strong in the Huia, and that both sexes participated in feedingthe young. Studies of the Saddleback may provide information from which inferences aboutthe Huia's behaviour can be drawn, and investigations on its feeding ecology andfunctional anatomy are currently in progress (Jenkins, pers. comm.). Finally,studies on other timber-feeding birds may shed further light on the Huia's adapta-tions. The closest parallels to these appear to be shown not by passerines, but bythe Wood Hoopoes (Order Coraciiformes, family Phoeniculidae). Within this family,the genus Phoeniculus (particularly P. aterrimus) appears to parallel the male Huia,while Rhinopomastus spp. resemble the female, both genera showing well-markedgaping adaptations. A study of feeding behaviour and anatomy in this family, aswell as being worth while in its own right, would make an interesting comparisonwith the Huia, in which similar bill forms and feeding methods have evolved in thetwo sexes of a single species. Yet, finally, it must be admitted that these indirect methods of investigation area very poor substitute for observations on the living bird. A study of this species inlife might have provided much information of general importance for evolutionarybiology, and would certainly have been of intense interest ; its loss is a matter forthe greatest regret. 46 P. J. K. BURTON XII. ACKNOWLEDGEMENTS I am much indebted to Dr Walter J. Bock for reading the first draft of this paper,and offering valuable criticism and advice. The excellent drawings of skulls (Figs.1-9) are the work of Sharon Chambers, to whom I am deeply grateful. Mr P. F.Jenkins, of the University of Auckland, kindly let me have news of his work on theSaddleback. Thanks are due to the British Trans-Americas Expedition (1971-2)for enabling me to collect several of the specimens used for comparison with theCallaeidae. XIII. REFERENCES AMADON, D. 1943. Bird weights as an aid in taxonomy. Wilson Bull. 55 : 164-177.ANTONY, M. 1920. Uber die Speicheldriisen der Vogel. Zool. Jb. (Abt.Anat.) 41 : 545-660.BEECHER, W. J. 1951. Adaptations for food getting in the American Blackbirds. Auk 68 : 411-440.BOAS, J. E. V. 1929. Biologisch-anatomische Studien iiber den Hals der Vogel. K. dansk. Vidensk. Selsk. Skr. (Naturvidenskab. Math. Afdel.) Ser. 9, 1 : 102-222.BOCK, W. J. I96oa. The palatine process of the premaxilla in the Passeres. Bull. Mus. comp. Zool. Harv. 122 : 361-488. ig6ob. Secondary articulation of the avian mandible. Auk 77 : 19-55. Kinetics of the avian skull. /. Morph. 114 : 1-41. Bill shape as a generic character in the cardinals. Wilson Bull. 76 : 50-61.— 1966. An approach to the functional analysis of bill shape. Auk 83 : 10-51. 1972. Morphology of the tongue apparatus of Ciridops anna (Drepanididae). Ibis 114:61-78. MS : Drawings of jaw and tongue musculature in passerine birds. BOWMAN, R. I. 1961. Morphological differentiation and adaptations in the Galapagos finches. Univ. Calif. Publs Zool. 58 : vii + 326 pp.BULLER, W. L. 1878. Further descriptive notes of the Huia (Heteralocha acutirostris). Trans. N.Z. Inst. 10: 211. 1888. A History of the Birds of New Zealand. 2nd edition. Vol. i. London. BURTON, P. J. K. 1969. Two bird specimens probably from Cook's voyages. Ibis 111 : 388- 390. I97ia. Some observations on the splenius capitis muscle of birds. Ibis 113 : 19-28. i97ib. Sexual size dimorphism in Alauda razae. Bull. Brit. Orn. Club 91 : 108-109. 1973- Structure of the depressor mandibulae muscle in the Kokako Callaeas cinerea. Ibis 115 : 138-140. 1974- Feeding and the Feeding Apparatus in Waders. Brit. Mus. (Nat. Hist.). ENGELS, W. L. 1938. Tongue musculature of passerine birds. Auk 55 : 642-650.FIEDLER, W. 1951. Beitrage zur Morphologic der Kiefermuskulatur der Oscines. Zool. Jb. (Abt.Anat.) 71 : 235-288. CANS, C. & BOCK, W. J. 1965. The functional significance of muscle architecture - a theoreti-cal analysis. Ergebn. Anat. EntwGesch. 38 : 115-142.GARROD, A. H. 1872. Notes on the anatomy of the huia bird (Heteralocha gouldi). Proc. Zool. Soc. Lond. : 643-647. GEORGE, J. C. & BERGER, A. J. 1966. Avian Myology. London and New York.GRAY, G. R. 1870. Hand-list of Genera and Species of Birds distinguishing those contained in the British Museum. Part 2. London.LAKJER, T. 1926. Studien iiber die trigeminus-versorgte Kaumuskulatur der Sauropsiden. Copenhagen.LOWE, P. R. 1938. Some anatomical and other notes on the systematic position of the genus Picathartes, together with some remarks on the families Sturnidae and Eulabetidae. Ibis i4th Ser. 2 : 254-269. ANATOMY OF HEAD AND NECK IN THE HUIA 47 MAYR, E. & GREENWAY, J. C. 1956. Sequence of passerine families (Aves). Breviora No. 58 : i-n.OLIVER, W. R. B. 1955. New Zealand Birds. 2nd edition. Wellington. 1963. The skull of the Huia. In The Book of the Huia by W. J. Phillipps. Christchurch. PALMGREN, P. 1949- Zur biologischen Anatomic der Halsmuskulatur der Singvogel. In Ornithologie als biologische Wissenschaft, pp. 192-203. Heidelberg.PHILLIPPS, W. J. 1963. The Book of the Huia. Christchurch.SELANDER, R. K. 1966. Sexual dimorphism and differential niche utilization in birds. Condor 68 : 113-151.SHUFELDT, R. W. 1913. On the osteology of the Red Wattle-Bird (Anthochaera carunculata). Emu 13 : 1-14.STARCK, D. & BARNIKOL, A. 1954- Beitrage zur Morphologic der Trigeminusmuskulatur der Vogel. Morphol. Jahrb. 94 : 1-64.STONOR, C. R. 1942. Anatomical notes on the New Zealand Wattled Crow (Callaeas) with especial reference to its powers of flight. Ibis nth Ser., 6 : 1-18.ZOOLOGICAL SOCIETY OF LONDON, 1870. Report on additions to the Society's Menagerie during the month of May. Proc. Zool. Soc. Lond. : 380-384.Zusi, R. L. 1962. Structural adaptations of the head and neck in the Black Skimmer Rynchops nigra Linnaeus. Publs Nuttall orn. Club No. 3 : vii + ioi pp. 1969. Osteology and myology of the head and neck of the Pied-billed Grebes (Podilvmbus). Misc. Publs Mus. Zool. Univ. Mich. No. 139 : 1-49. KEY TO ABBREVIATIONS IN FIGURES Ap i a m e Aponeurosis i, M. adductor mandibulae externus Ap 2 a m e Aponeurosis 2, M. adductor mandibulae externus Ap 3a d m Aponeurosis 3a, M. depressor mandibulae Ap m Median aponeurosis of tongue C o Occipital crest E u s Opening of Eustachian tube G a o Gl. angularis oris L i j m Internal jugomandibular ligament L p o Postorbital ligament M a m e c M. adductor mandibulae externus caudalis M a m e r 1 M. adductor mandibulae externus rostralis lateralis M a m e r m M. adductor mandibulae externus rostralis medialis M a m e r t M. adductor mandibulae externus rostralis temporalis M a m e v M. adductor mandibulae externus ventralis M a m p M. adductor mandibulae posterior M as M. ascendens cervicis. M b m M. branchiomandibularis Meg M. ceratoglossus M c g t Tendon of M. ceratoglossus M co M. complexus M d m M. depressor mandibulae M d m a M. depressor mandibulae pars anterior M dt M. dermotemporalis M f c b M. flexor colli brevis M hg a M. hypoglossus anterior M hg o M. hypoglossus obliquus M i t M. intertransversarius M p q i M. protractor quadrati et pterygoidei i M p q 2 M. protractor quadrati et pterygoidei 2 M ps p M. pseudotemporalis profundus 48 P. J. K. BURTON M ps s M. pseudotemporalis superficialis M pt d 1 M. pterygoideus dorsalis lateralis M pt d m a M. pterygoideus dorsalis medialis anterior M pt d m p M. pterygoideus dorsalis medialis posterior M pt r M. pterygoideus retractor M pt v 1 M. pterygoideus ventralis lateralis M pt v m M. pterygoideus ventralis medialis M r c 1 M. rectus capitis lateralis M r c v M. rectus capitis ventralis M s co M. splenius colli M se h M. serpihyoideus M sp M. spinalis M st h M. stylohyoideus M th h M. thyreohyoideus M tr h M. tracheohyoideus M tr 1 M. tracheolateralis N f h Nasal-frontal hinge P ex Exoccipital process P i Internal process of mandible P r Retroarticular process of mandible Dr P. J. K. BURTON Sub-department of OrnithologyBRITISH MUSEUM (NATURAL HISTORY)TRINGHERTFORDSHIRE A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.1965. (Out of Print.) £3.75. 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogyof the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates,77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur]1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : theBiology and Evolution of a Species Flock. Pp. 1-134 > T Plate, 77 Text-figures.1974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU MISCELLANEA 19 AUG1974 •v * .c^V BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 2 LONDON: 1974 MISCELLANEA Pp 49-138 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 2 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 2, of the Zoological series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 18 July, 1974 Price £4.70 CONTENTS Page A new skink from the New Hebrides. By LORD MEDWAY ... 53 New genus of whale-louse. By R. J. LINCOLN & D. E. HURLEY . . 59 Catalogue of whale-lice. By R. J. LINCOLN & D. E. HURLEY ... 65 Review of Laephotis Thomas 1901. By J. E. HILL .... 73 The genus Finmarchinella Swain. By J. W. NEALE .... 83 Euplotes rariseta sp. n. By G. R. CURDS, B. J. WEST & J. E. DORAHY . 95 Clonal cultures of Euglypha (Protozoa). By R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT .......... 103 Three species of Euplotes. By C. R. CURDS . . . . . . 113 Bats from Southeastern Asia. By J. E. HILL . . . . . 127 A NEW SKINK (REPTILIA: SCINCIDAE: GENUSEMOIA) FROM THE NEW HEBRIDES WITH COMMENTS ON THE STATUS OF EMOIA SAMOENSISLOYALTIENSIS (ROUX) By LORD MEDWAY DURING the Royal Society/Percy Sladen expedition to the New Hebrides, 1971,three immature skinks of an undescribed taxon were caught on Aneityum island.Subsequently three further specimens, taken on the same island by Miss EvelynCheesman, were found among the collections of the British Museum (NaturalHistory) where they had been catalogued as Emoia nigra. Apart from a note thatthe period of collecting was 1954-55, Miss Cheesman's material lacks field data. Itis likely, but not certain, that the skinks were caught near Red Crest, the campthree miles from the coast (at Anelgauhat) at about 1200 ft (366 m) elevation,occupied by Miss Cheesman from mid-March through July 1955 (see Cheesman,1957 : 312-327). Of this site, Miss Cheesman has written : 'Plenty of lizards darted about all over the clearing on sunny days, overcast daysmade them inactive and then one found them under logs, in holes, or hollow trees . . .Individuals were inclined to be tame, a smooth blue-grey skink with a green matewould appear whenever I had meals outside on the clearing. She accepted cheese,bread, biscuit, boiled rice and banana but refused cooked taro, beating a retreat asif insulted. To catch lizards was another matter, but I did succeed in getting agood series.' During the 1971 expedition skinks were also collected on all other islands visited,i.e. in order northwards from Aneityum, Tanna, Erromanga, Efate, Malekula, Malo,Aore and Espiritu Santo. Previous collections from these islands, and from othersin the group, have been reported by Boulenger (1887), Roux (1913), Schmidt & Hurt(1930), Burt & Burt (1932) and Angel (1935). Additional unpublished materialfrom the New Hebrides, examined in the course of preparation of this note, is pre-served in the Museum National d'Histoire Naturelle, Paris (Frangois collection),the Natur-Museum Senckenberg, Frankfurt a. Main (Bregulla collection), and theBritish Museum (Natural History) (mainly collected by J. R. Baker and associatesor by E. Cheesman). Other than the six examples from Aneityum, no other repre-sentatives of the undescribed skink have been found among any collections. Sinceit is evidently confined to the island of Aneityum, it is appropriate that the newtaxon be named : Emoia aneityumensis n. sp. - the Aneityum skink HOLOTYPE. BM 1956.1.3.65, adult male, collected by Miss E. Cheesman onAneityum island, New Hebrides, South West Pacific, 1954-55 ; for likely place andperiod of collection, see above. Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 54 LORD MEDWAY PARATYPES. BM 1956.1.3.63, adult male, and 1956.1.3.64, subadult female, alsocollected on Aneityum by Miss Cheesman during 1954-55 ; BM 1973.1534 (col-lectors' number RSNH 14-01), juvenile, apparently female, caught on 19 July1971, BM 1973.1535 (RSNH 18-01), immature female, 21 July 1971, and BM 1973.1536 (RSNH 19-03), juvenile male, 23 July 1971, all taken in disturbed and partiallyregenerated forest about i mile northeast of Anelgauhat, Aneityum, by Lord Medwayand A. G. Marshall. DIAGNOSIS. In size and colour, and in the scalation of the head, similar to Emoiasamoensis (Dumeril & Dumeril), differing in the greater number of longitudinalscale rows and the smaller number of subdigital lamellae. Similarly coloured toEmoia speiseri (Roux), but again differing in details of scalation, larger as a matureadult, and evidently producing a greater number of eggs at one time. DESCRIPTION. An Emoia, possessing the characters of that genus as defined byGray (1845 : 95) and elaborated by Smith (1937). On the head, the prefrontalsform a short medial suture, thus excluding the nasals from contact with the frontals.The interparietal is a separate shield, not fused to the parietals. The fifth or sixthupper labial is large, underlying the eye. The coloration of the holotype, after 18 years in alcohol, is dark brown on theupperparts, with a broken row of irregularly shaped black spots or flecks extendingas an interrupted dorsolateral line from the ear orifice to the base of the tail, passingabove the origins of the hindlimbs. The flanks are brown, barred and flecked byinvasions of the buffy white ventral colour. In the three young animals, whenfreshly caught, the upperparts were grey-brown or olive-brown, marked with abroken row of irregular spots, flecks or blotches of black, intermixed with flecks ofeau-de-nil, running from the ear to the base of the tail. The flanks were olive-brown,barred and flecked with the colour of the underparts, which were dirty white. Theunbroken tail was the same brown as the ground colour of the upperparts, butregenerated portions were a lighter shade of brown, more or less unmarked. Thespecimen BM 1956.1.3.63 (again, after 18 years in preservative) is uniformly greyishbrown above and buff below, without markings ; in my opinion, in life it could wellhave appeared 'smooth blue-grey' (cf. Cheesman 1957 : 325, quoted above). Details of scalation and measurements are given in Table i. By comparison,specimens of Emoia samoensis (two syntypes nos. 2764 & 7070 in the Paris museum,two from Samoa nos. 4632 & 4633 in the Basel museum, and two from Erromanganos. 1860.3.18.8 & 1860.3.18.11 in the British Museum) have 32-36 scale rows atmid-body, 44-51 subdigital lamellae under the fourth toe, and measure snout-ventlength 92-102 mm, total length 250-290 per cent. Twelve specimens of Emoiaspeiseri from all parts of this species' range in the New Hebrides have 28-34 scalerows at mid-body, 38-48 subdigital lamellae under the fourth toe, and measure60-72 mm snouth-vent, total length about 270 per cent (cf. Roux, 1913). The female E. aneityumensis BM 1956.1.3.64 has four enlarged ova in the leftovary and at least one in the right, and BM 1973.1535 has four enlarging ova dis-cernible in the right ovary (left ovary not recognized). Of six undoubted femaleE. speiseri examined, two only had enlarging ova ; one had a single large follicle in A NEW SKINK FROM THE NEW HEBRIDES 55 the left ovary only, and the other had one large follicle in each ovary. Among E.samoensis, Roux (1913), has reported a gravid female containing four large ova oreggs ('gros oeufs'). There is no reason to believe that E. aneityumensis is confined to the vicinity ofAnelgauhat on Aneityum. Retrospectively I attribute to this species a grey skinkjudged to be at least the size of the holotype (Table i), seen (but not caught) climbingthe cliff above the sea-shore at Aegiptzav, on the north coast of the island, in July1971. TABLE i Scale counts and measurements of the type series of Emoia aneityumensis Reg. no. Longitudinal Lamellae under Snout-vent Total length1 BM scale rows fourth toe length (as % snout-vent) at mid-body (mm) Left Right 1956.1.3.652 40 42 42 92 (250) 1956.1.3.63 40 38 38 89 274 1956.1.3.64 40 36 36 83 (240)I973-I534 42 39 38 49 257I973-I535 4° 41 41 70 (203)I973-I536 41 38 39 66 (221) 1 Figures are given in parentheses if the tail showed signs of loss and subsequent regeneration.* Holotype. DISCUSSION The genus Emoia is represented in the New Hebrides by ten taxa (Medway &Marshall, in prep.), of which E. sanfordi, E. speiseri and E. aneityumensis areendemic to the archipelago. Only E. aneityumensis is confined, as far as known, toa single island. Of the two taxa most closely resembling E. aneityumensis, E. speiseri is sympatricwith it on Aneityum island. Differences between the two, described above, establishthat they are distinct species. E. samoensis, on the other hand, has been found inthe New Hebrides only on Erromanga, some 180 km north-northwest of Aneityumand the next large island but one. The specimens (BM 1860.3.18.8 & 1860.3.18.11)were collected more than a century ago by Mr Cuming (Boulenger, 1887). In 1971we did not find the species, but our stay on Erromanga was brief and collecting notintensive ; there are no grounds to doubt the record. The third of Cuming's speci-mens allocated to this species by Boulenger (BM 1860.3.18.12) is in fact E. sanfordi,an identification which confirms at least that the collection derived from the NewHebrides. Samples of E. samoensis from all parts of its range, from the Loyalty Islands toFiji and the Samoan group, provide no evidence of geographical variation. Separa-tion of the Loyalty Islands population, described as a subspecies by Roux (1913), isnot justified. The distinguishing character - adult size invariably less than that ofexamples from neighbouring archipelagoes ('archipels voisins') - was established by 56 LORD MEDWAY comparison with specimens from the New Hebrides (Roux 1913 : no). This material(in the Basel Museum of Natural History) was re-examined in the course of prepara-tion of this note. All New Hebrides specimens identified by Roux (1913 : 155) asLygosoma (Emoa) samoense, a synonym of E. samoensis, prove to be E. sanfordi andthus provide a fallacious basis for comparison. In the scalation of the head, thenumber of longitudinal scale rows and of subdigital lamellae, specimens from theLoyalty Islands fall within the range of variation of E. samoensis from other partsof the South West Pacific. The absence of demonstrable geographical variation in E. samoensis over thiswide area makes it unlikely that the species would differentiate markedly betweenErromanga and Aneityum islands. E. samoensis and E. aneityumensis may replaceeach other ecologically on their respective islands in the New Hebrides, but there areno grounds for treating the two taxa as geographical races of one species. Thecharacters that distinguish Emoia aneityumensis set the taxon apart from all othermembers of the genus, and it must be treated as a distinct species. ACKNOWLEDGEMENTS The Royal Society/Percy Sladen expedition was led by Dr K. E. Lee, on whom fellthe chief burden of organization and administration. During field work, Dr A. G.Marshall was jointly responsible for the collection and curation of specimens ; wewere assisted on Aneityum by Fred Boe, John Wycliffe and Hugo Tamata. TheRoyal Society subsequently provided a travel grant which enabled me to visitEuropean museums containing collections of New Hebridean vertebrates ; for theirkindness in making available specimens in their care and for other acts of hospitality,I am grateful to Professor J. Guibe, Professor U. Rahm and Dr K. Klemmer. Allother work for this paper has been carried out at the British Museum (NaturalHistory), and I am particularly grateful to the Curator of Herpetology, Miss A. G. C.Grandison, and staff for the facilities provided. Miss Grandison and Dr Marshallkindly read and criticized this paper in draft. REFERENCES ANGEL, F. 1935. Liste des reptiles recoltes par la Mission Aubert de la Rue aux Nouvelles Hebrides ou dans les iles voisines. Bull. Mus. Hist, nat., Paris (2) 7 : 54-56.BOULENGER, G. A. 1887. Catalogue of Lizards in the Collection of the British Museum. 2nd ed. London : British Museum.BURT, C. E. & BURT, M. D. 1932. Herpetological results of the Whitney South Sea expedition VI. Pacific island amphibians and reptiles in the collection of the American Museum of Natural Nistory. Bull. Am. Mus. nat. Hist. 63 : 462-597.CHEESMAN, E. 1957. Things Worth While. London : Hutchinson & Co.GRAY, J. E. 1845. Catalogue of Lizards in the Collection of the British Museum. London : British Museum.MEDWAY, LORD & MARSHALL, A. G. (in prep.). Observations on the vertebrate fauna of the New Hebrides.Roux, J. 1913. Les reptiles de la Nouvelle-CalMonie et des iles Loyalty. Appendice : Note sur quelques reptiles des Nouvelles Hebrides, des lies Banks et Santa Cruz. In: F. Sarasin & J. Roux (eds), Nova Caledonia, Zoologie 1 : 80-160, pis IV, V. A NEW SKINK FROM THE NEW HEBRIDES 57 SCHMIDT, K. P. & BURT, C. E. 1930. Herpetological results of the Whitney South Sea expedi-tion V. Description of Emoia sanfordi, a new lizard from the islands of the Western Pacific(Scincidae). Am. Mus. Novit. 436 : 1-3. SMITH, M. A. 1937. Review of the genus Lygosoma (Scincidae : Reptilia) and its allies. Rec.Indian Mus. 39 : 213-234. LORD MEDWAYGREAT GLEMHAM HOUSESAXMUNDHAMSUFFOLK IPiy iLP SCUTOCYAMUS PARVUS, A NEW GENUS AND SPECIES OF WHALE-LOUSE (AMPHIPODA: CYAMIDAE) ECTOPARASITIC ON THE NORTH ATLANTIC WHITE-BEAKED DOLPHIN By ROGER J. LINCOLN & D. E. HURLEY 'The sea-lowse is an insect that is an enemy of all kinde of Whales, which bybiting and tickling it puts into such a rage, that they are forced to run uponthe sand, and hasten to dry land : I know nothing concerning the use ofthese creatures ; but I seriously exhort posterity to search out the use ofthem.' From : The Theater of Insects or Lesser Living Creatures Book n Chapter 38p. 1126 by Tho. Mouffet, Doctor inPhysick, London, 1658. INTRODUCTION It is rare to find a new species of cyamid, rarer still to discover a new genus. Sincethe first figure and description in the scientific literature of a 'Walfisches Lauss' byMartens in 1675, nineteen accepted species of cyamid have been described, onlythree of these in this century (Leung, 1967). Fifteen species are ascribed to thegenus Cyamus and one each to Neocyamus, Platycyamus, Syncyamus and Isocyamus.Although the last two were established in 1955, only Syncyamus is based on a recentlydescribed species. Their rarity is not unexpected. Cyamids are host-specific to cetaceans which area well-studied and numerically limited group. It is even more surprising, then, todiscover in the collections of the British Museum (Natural History), unrecognizedsince their collection in 1933, specimens of a distinct and new species of cyamid froma common North Atlantic dolphin from which cyamids have never, to our knowledge,been recorded. These cyamids stood out for two reasons ; their generally small size in comparisonwith other species in the collection, and their distinctively rounded, saucer-shaped,low-profile outline which immediately suggested their novelty. These featuresindicated that their host would prove to be a fast-swimming species, which is indeedthe case, and the subsequent examination has revealed further adaptation to lifein an area of turbulent flow in the shape of the dactyls and cramp-iron teeth on thebody and appendages. The cyamids were collected by Dr F. C. Fraser from a white-beaked dolphin,Lagenorhynchus albirostris Gray, and later presented to the Crustacea Section. The Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 60 R. J. LINCOLN & D. E. HURLEY dolphin was caught in the North Sea, off Peterhead in Aberdeenshire, on 21 July1933. It was a young female measuring 5 ft 7! in. in length with a pronounceddeformity in the curvature of the spine behind the dorsal fin (Fraser, pers. comm.). SCUTOCYAMUS gen. nov. DIAGNOSIS. Pereon segments 3-4 and 6-7 fused. Antenna i, 2-articulate ;antenna 2, i-articulate. Maxilla 2 with outer lobes absent. Maxillipeds fused andmarkedly reduced to form a small cleft flap. Pereopods i and 2 strongly dissimilarin shape and with i very much smaller than 2 ; pereopod 2 only 3-articulate. Malewithout accessory gills. Type species, Scutocyamus parvus sp. nov. Scutocyamus parvus sp. nov.(Text figs. la-g, 2a-h ; PI. la-e) DIAGNOSIS. With the characters of the genus given above. Body of small size,robust, strongly flattened dorsoventrally, head weakly immersed into anterior pereonsegment. Pereopod i (Fig. ic) lacking unguis, dactylus armed with numerous smallteeth (PI. le) ; pereopod 2 and pereopods 5-7 powerfully developed and held incharacteristic posture giving a general oval symmetry to the shape of the animal(Fig. la ; PI. la). Gills simple, single. Pereon segments 5-7 each with a pair oflarge spines on the ventral surface. DESCRIPTION. Length of body from apex of head to end of pereon 1-7-2-4 mmin male and 2-6-3-1 mm in ovigerous female. Maximum width of body at the levelof tergite 5 from 0-9 to 1-3 mm in male and from 1-4 to 1-8 mm in ovigerous female.No trace of pigmentation remaining in the alcohol preserved material. Pereon(Fig. la) somewhat oval in outline (pereon segment i fused with head), male rathermore slender than female ; tergites 3-4 in female slightly shorter than other tergites,in male very much shorter and also much narrower than other tergites (Fig. ib) ;tergite 7 weakly immersed into pereon segment 6 (Fig. la, b) ; a pair of strong spineson the ventral side of segments 5-7. Head with sides convex, immersed slightlyinto anterior pereon segment, anterolateral angles expanded into large lobes giving awide frontal margin to head; eyes small, oval, mid-dorsal. Antenna i (Fig. 2c)small, 2-articulate, article 2 longer than i, apex with prominent group ofsensory setae. Antenna 2 (Fig. 2d) extremely small, i-articulate, but constrictionnear tip gives impression of separate terminal article, apex with several largesensory setae. Upper lip (Fig. 2e) large, outer margin weakly concave and denselyfringed with fine setae. Mandibles (Fig. 2g) with 2 incisor processes, the anteriorof 1-2 teeth, the posterior incisor of several teeth ; right mandible with 2 penicils,left with only one ; molar process a rounded protuberance covered with very finesetae. Lower lip (Fig. 21) with inner lobes fused into a single elongate lobe, outerlobes slightly broader than inner, distal margins fringed with long setae. Maxilla i(Fig. 2h), apex with 3 pairs of curved spines each with 2-4 small teeth on innermargin ; palp i-articulate, reaching to about the apex of the outer lobe and with NEW GENUS AND SPECIES OF WHALE-LOUSE 61 g FIG. i. Scutocyamus parvus sp. nov. a, dorsal entire, female ; b, dorsal entire, male ; c,pereopod i ; d, pereopod 2, ventral; e, pereopod 5, ventral; f, pereopod 6, ventral;g, pereopod 7, ventral; bar scale a-b, i-o mm ; c, 0-2 mm ; d -g, 0-4 mm. 62 R. J. LINCOLN & D. E. HURLEY small group of sensory setae at tip. Maxilla 2 (Fig. 2b) a single elongate lobe witha small group of sensory setae at tip ; right and left maxilla 2 fused along mid-line.Maxillipeds (Fig. 2b) fused and reduced to a small cleft flap. Pereopod i (Fig. ic)extremely small, simple, 5-articulate, propodus elongate and about twice as long aswide ; dactylus broad with numerous combs of small teeth towards apex (PL le),unguis absent. Pereopod 2 (Fig. id) powerfully developed, only 3-articulate,proximal article large and robust with small marginal spine ; article 2 very broadand flattened, the outer margin with a deep indentation which probably marks thepoint of fusion of two articles ; article 2 with 2 small marginal spines ; dactylusrobust strongly angled at its mid-point, and with a small but sharply pointed unguis.These cyamids have a quite characteristic posture with the expansive secondpereopods held across the front of the head to form an effective shield. Pereopods5-7 (Figs. le, f, g) powerful, 5-articulate, basal article (basis + ischium) short andstout with 3 well-developed ventral spines ; merus broad and flat with a singlespine on distal margin ; carpus about equal to size of merus with a small spine ondistal margin and an extremely large mid-ventral spine, the inner posterior angleproduced into a small triangular process overlapping the dorsal surface of merus ;propodus elongate and extremely robust; dactylus acutely angled and with sharplypointed unguis. Gills single, quite short, tapering somewhat, and held acrossventral surface of pereon in forward direction ; accessory gills absent in male. Broodpouch rounded (Fig. 2a), containing only 7-10 eggs or young ; margins of oostegitesfringed with many short setae ; genital valves well developed with inner marginalso fringed with small setae. Pleon a small bilobed structure. MATERIAL EXAMINED. n$$ (ovigerous), 2-6-3-1 mm length, 1-4-1-8 mm width:21$$ (immature), 2-0-2-5 mm length, 1-1-1-4 mm width : 28^, 1-7-2-4 mm length,0-9-1-3 mm width : 37 juveniles. Holotype $ registration no. 1973 : 105 ; paratypesregistration no. 1973 :106, deposited in the collections of the British Museum (NaturalHistory). REMARKS. Scutocyamus parvus can be immediately recognized by the small bodysize, general symmetry given to the body posture by the expansive second pereopods,and by the fusion of pereon tergites 3-4 and tergites 6-7. Only one other species,Syncyamus pseudorcae Bowman, has a similar fusion of the pereon segments but it isrestricted to tergites 6-7. While the amalgamation of tergites 3-4 is complete inScutocyamus (PI. id) the posterior tergites retain a weak demarcation line whichcan be mistaken for a suture under a light microscope. Confirmation that pereontergites 6-7 are in fact fused was obtained by examining material with a scanningelectron microscope. The demarcation line was found to be a shallow depressionwhich follows the line of fusion of the two segments. Of the five recognized genera of cyamids, Scutocyamus seems to be most closelyallied to the monotypic Syncyamus. In addition to the fusion of the pereon tergitesboth have a similar reduction of the mouthparts, especially the reduction of themaxillipeds to a small flap, the fusion of the second maxillae, and the fusion of theinner lobes of the lower lip. Also, the absence of an unguis on the small pereopod Iis a character shared by the genera. However, Scutocyamus retains a number of NEW GENUS AND SPECIES OF WHALE-LOUSE FIG. 2. Scutocyamus parvus sp. nov. a, ventral entire, female ; b, mouthparts, ventral; c,antenna i ; d, antenna 2 ; e, upper lip ; f, lowerlip ; g, mandible with upper lip ; h,maxilla i ; bar scale a, i-o mm ; b-h, o-i mm. 64 R. J. LINCOLN & D. E. HURLEY unique characters such as the fusion of pereon tergites 3-4, extreme reduction of theantennae and pereopod i, a 3-articulate pereopod 2, and an absence of accessorygills in the male. The armature of comb-like teeth on the propodus of pereopod ihas not previously been reported and appears to be an adaption to scrape the surfaceof the host. ACKNOWLEDGEMENTS The authors wish to thank Dr F. C. Fraser for his assistance with original data andfor drawing attention to the opening quotation. The junior author wishes tothank the Trustees of the British Museum (Natural History) for facilities madeavailable during Official Leave from New Zealand. REFERENCES BOWMAN, T. E. 1955. A new genus and species of whale-louse (Amphipoda, Cyamidae) from the false killer whale. Bull. mar. Sci. Gulf Caribb. 5 (4) : 315-320.LEUNG, Y.-M. 1967. An illustrated key to the species of whale-lice (Amphipoda, Cyamidae), ectoparasites of Cetacea, with a guide to the literature. Crustaceana 12 (3) : 279-291.MARTENS, F. 1675. F. Martens. . . . Spitzbergische oder Groenlandische Reise Beschreibung gethan im Jahr 1671, etc. pp [viii] 132, 16 pis. Hamburg. Dr ROGER J. LINCOLN Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 Dr D. E. HURLEY NEW ZEALAND OCEANOGRAPHIC INSTITUTE P.O. Box 8009 WELLINGTON NEW ZEALAND It PLATE i Scutocyamus parvus sp. nov. a, entire ventral, male (X 30) ; b, mouthparts, male (xi8o) ;c, pleon ventral, male ( X 120) ; d, fusion pereon tergites 3-4, dorsal female ( x 120) ; e, pereopodi, apex of propodus, male (x i-i k). Scanning electron microscope ; gold coating ; 20 kv. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i CATALOGUE OF THE WHALE-LICE (CRUSTACEA : AMPHIPODA : CYAMIDAE) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) By ROGER J. LINCOLN & D. E. HURLEY INTRODUCTION THE collections of the British Museum (Natural History), containing as they domuch of the material taken during the Discovery Investigations (1925-27) on thebiology of the southern whales and much incidentally collected Northern Hemispherematerial, represent one of the more significant collections of cyamid material.Numerically this collection is not large but it contains some types and materialwhich has been little studied. Furthermore the group is usually very poorly repre-sented in museum collections. The discovery of a new genus and species of cyamid (Lincoln & Hurley, 1974) hasbeen one of the benefits resulting from a re-examination of the material in thecollections of the Crustacea Section. Since all identifications have now been verifiedand labelling updated following the nomenclature given in the recent cyamid synopsisof Leung (1967), this is a suitable opportunity to present a published catalogue ofholdings for the benefit of other workers. The nomenclature for the whales followsthat used by Hershkovitz (1966). The assistance of Miss J. Ellis in collating the information is gratefully acknow-ledged. CYAMUS Latreille, 1796 Cyamus abbreviatus Say, 1818(i) Type ? i specimen (dry collection). North America. Leach Collection. Cyamus balaenopterae Barnard, 1931 (1) Syntypes, 6^, 4$$ ; Ann. Mag. nat. Hist. (10) 7 : 425-430. Reg. no. 1936:11:2:3495-3501. Saldanha Bay, South Africa ; on Balaenoptera physalis (L.)[blue whale according to label] whale no. W 1096. Discovery Collection. (2) 2&J, 8$$, i juv. Reg. no. 1936:11:2:3502-3531. Saldanha Bay, South Africa ;on Balaenoptera musculus (L.) whale no. W 867. Discovery Collection. (3) 9&?> 6$$. Reg. no. 1936:11:2:3502-3531. Durban, South Africa; onBalaenoptera physalus (L.) whale no. W Dig. Discovery Collection. (4) *(?, 5?? (immature). Reg. no. 1936:11:2:3502-3531. Saldanha Bay, SouthAfrica, 18.8.1926 ; on Balaenoptera musculus (L.) whale no. W 961. DiscoveryCollection. Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 66 R. J. LINCOLN & D. E. HURLEY (5) 3<£?> 6$$, i? ovig., i juv. Reg. no. 1936:11:2:3502-3531. Saldanha Bay,South Africa, 15.8.1926 ; on Balaenoptera physalus (L.) whale no. W 948.Discovery Collection. (6) 20(3$, 15??, 10 juvs. Reg. no. 1973:104:45. No locality data, 24.10.1948 ; onBalaenoptera physalus (L.). Discovery Collection. Cyamus boopis Lutken, 1870 (1) i$. Reg. no. 1853:68. Greenland. Collected by Holboll. (2) 10(3$, 10 juvs. Reg. no. 1910:8:9:1-6. Bransfield Straits, South Shetlands ;on Megaptera novaeangliae (Borowski). Collected by J. A. Morch. (3) 2c?$> J? (immature). Reg. no. 1911:11:8:23064-066. No locality data. A. M.Norman Collection. (4) 3$$. Reg. no. 1920:7:5:61-62. Deception Island, South Shetlands, 1914 ;on Megaptera novaeangliae (Borowski). Collected by A. G. Bennett. (5) iJc&J- Reg- no- 1924:7:10:1-5. Olna Firth, Shetland Islands, 19.6.1924 ; onMegaptera novaeangliae (Borowski). Collected by F. V. Morley. (6) 3(3$, 3?$ (immature), 2 juvs. Reg. no. 1928:12:1:2966-71. Durban, SouthAfrica, 23.7.1908 ; on Megaptera novaeangliae (Borowski). T. R. R. StebbingCollection. (7) i$, 2?? (all immature). Reg. no. 1936:11:2:3542-3581. South Georgia ; onMegaptera novaeangliae (Borowski) whale no. 355. Discovery Collection. (8) 6(3$, i$. Reg. no. 1936:11:2:3542-3581. South Georgia; on Megapteranovaeangliae (Borowski), whale no. 373, genital region. Discovery Collection. (9) 2(&J, 2$$, 5 juvs. Reg. no. 1936:11:2:3542-3581. South Georgia; onMegaptera novaeangliae (Borowski), whale no. 387. Discovery Collection. (10) 3(3$, 3??, 10 juvs. Reg. no. 1936:11:2:3542-3581. Deception Island, SouthShetlands ; on Megaptera novaeangliae (Borowski). Discovery Collection. (n) 6$$, 4$?, 10 juvs. Reg. no. 1936:11:2:3542-3581. Saldanha Bay, SouthAfrica, 29.9.1926 ; on Megaptera novaeangliae (Borowski), whale no. 1125.Discovery Collection. (12) ii(3$, i?. Reg. no. 1936:11:2:3542-3581. Durban, South Africa; onMegaptera novaeangliae (Borowski), whale no. 03. Discovery Collection. (13) 2$$, 4??. Reg. no. 1936:11:2:3542-3581. Durban, South Africa, on Physetercatodon L., whale no. Dai. Discovery Collection. (14) 10$$, 7$$, 2 juvs. Reg. no. 1936:11:2:3542-3581. Saldanha Bay, SouthAfrica, 6.8.1926; on Megaptera novaeangliae (Borowski), whale no. 918.Discovery Collection. (15) i6(3$, 9$$, 10 juvs. Reg. no. 1965:9:8:1-15. Paita, Peru, 05^5' S, 8i°io' W,19.10.1960, on Megaptera novaeangliae (Borowski), $ 13-3 m, whale no. PAi.Collected by Robert Clarke. (16) 2<&J. Reg. no. 1972:556:2. Varanger Fiord, Norway ; on Megaptera novae-angliae (Borowski). G. O. Sars Collection. (17) 50(3$, io$$ (immature). Reg. no. 1972:568:60. Off Russell Bay, Bay ofIslands, New Zealand, August 1912 ; on Megaptera novaeangliae (Borowski).Terra Nova Collection. CATALOGUE OF WHALE-LICE 67 (18) 150(2(2??. Reg. no. 1972:569:150. Off Russell Bay, Bay of Islands, NewZealand, August 1912 ; on Megaptera novaeangliae (Borowski). Terra NovaCollection. (19) 14^, 6??. Reg. no. 1972:577:20. No locality data. No collection data. (20) 9(2(2, 6?? (immature), i juv. Reg. no. 1972:579:16. St. Vicente. Collected byR. T. Lowe. Cyamus catodontis Margolis, 1954 (1) 300(2(2?? (mature and immature). Reg. no. 1965:9:8:16-45. Paita, Peru05°05' S, 8i°io' W, 30.9.1960 ; on Physeter catodon L., <$ 13-2 m, whale no-Pa870. Collected by R. Clarke. (2) 100(2(2??. Reg. no. 1965:9:8:16-45. Pisco, Peru, I3°47' S, 76°i5' W, 17.10.1960 ; on Physeter catodon L., <$ 13-6 m, whale no. Pi59- Collected by R. Clarke. (3) I5°^?? (mature and immature). Reg. no. 1965:9:8:16-45. Pisco, Peru,I3°47' S, 76°i5' W, 17.1.1961 ; on Physeter catodon L., ^ 15-2 m, whale no.Pii6o. Collected by R. Clarke. (4) 150(2(2, 150$$. Reg. no. 1973:103:300. No locality data ; on Physeter catodonL., (2 54 ft, whale no. 20. Discovery Collection, 1951-52. Cyamus ceti (L., 1758) Lamark, 1801 (1) i(2, i$. Reg. no. 1911:11:8:23075-75. No locality data; A. M. NormanCollection. (2) 3<2c?, 8??- Reg. no. 1928:12:1:2978-2984. Greenland ; on Balaena mysticetusL. T. R. R. Stebbing Collection. (3) 6^(2, 3??- Reg. no. 1956:10:10:340-343. Arctic Sea, 1893. UniversityCollege, Dundee Collection (S.S. Eclipse). (4) 300(2(2, 200$$, 50?$ (immature), 50 juvs. Reg. no. 1972:556-600. NorthEurope ? No collection data. (5) 2(2c2, 4??. Reg. no. 1972:588:6. Murray Firth, Scotland. Leach Collection. (6) 6(2(2, *?, 6 juvs. Reg. no. 1972:559:13. No locality data. No collection data. (7) 9cft2> 10??. Reg. no. 1972:567:19. No locality data. No collection data. (8) 10(2(2, io$$, 20 juvs. Reg. no. 1972:578:40. Baffin Bay, 20.7.1894. UniversityCollege, Dundee Collection (S.S. Eclipse}. (9) Reg. no. 6o4a, b, d, c, dry collection (Leach cab : 31). Britain. LeachCollection. Cyamus erraticus Roussel de Vauzeme, 1834 (x) 5c&2, 4?? (immature). Reg. no. 1912:10:2:1-7. 6 miles N.E. of Flugga,Shetland. Collected by Lovett. (2) 4(2(2, i$. Reg. no. 1928:12:1:2966-71. Durban, South Africa, 23.7.1908 ; onEubalaena. T. R. R. Stebbing Collection. (3) 8(2(2, 10??. Reg. no. 1936:11:2:3532-3541. Saldanha Bay, South Africa,23.7.1908 ; on Eubalaena glacialis australis Desmoulins, whale no. 1020.Discovery Collection. 68 R. J. LINCOLN & D. E. HURLEY (4) 2c&?» 5??i i? (immature). Reg. no. 1972:560:8. Rockall, North Atlantic,1.7.1914 ; on flipper of Eubalaena glacialis glacialis Miiller. No collection data. (5) I4c£c?, 3$9. Reg. no. 1972:570:17. No locality data. Albany Museum,Grahamstown Collection. (6) !<£. Reg. no. 1972:575:1. No locality data ; on Eubalaena glacialis glacialisMiiller. Collected by Professor Goldberg. Cyamus gracilis (Roussel de Vauzeme, 1834) (1) i<£. Reg. no. 1907:12:2:218. Pacific Ocean. University College, DundeeCollection. (2) i$, i$ (immature). Reg. no. 1911:11:8:23069-70. No locality data. A. M.Norman Collection. (3) 8&J (immature), 4$$. Reg. no. 1928:12:1:2972-74. False Bay, Cape, SouthAfrica. T. R. R. Stebbing Collection. (4) i$, 6 juvs. Reg. no. 1936:11:2:3582-3584. South Georgia ; on Eubalaenaglacialis australis Desmoulins, whale no. 503. Discovery Collection. (5) Reg. no. 6o5a (dry collection), Leach cab : 31. British Sea. LeachCollection. Cyamus monodontis Liitken, 1873 (1) i specimen (slide collection) . No locality data ; on Monodon monoceros L. Nocollection data. (2) 3$$, 10 juvs. Reg. no. 1928:12:1:2975-77. No locality data ; on Monodonmonoceros L. T. R. R. Stebbing Collection. (3) 2<£<£, 2$$. Reg. no. 1911:11:8:23071-74. No locality data ; on Monodonmonoceros L. A. M. Norman Collection. (4) 5^, 5$$, 10 juvs. Reg. no. 1956:10:10:344-346. Davis Straits, Greenland,Canada ; on Monodon monoceros L. University College, Dundee Collection. (5) i$. Reg. no. 1972:561:1. No locality data. University College, DundeeCollection. (6) i$. Reg. no. 1972:562:1. No locality data ; on Monodon monoceros L.Collected by Nuttall. (7) ioo<^$$. Reg. no. 1972:563:100. Upernavik, Greenland, 1892. UniversityCollege, Dundee Collection. Cyamus nodosus Liitken, 1860 (1) 4$$, 2$$ (immature), 14 juvs. Reg. no. 1907:12:2:212-217. Davis Straits,Greenland, Canada. University College, Dundee Collection. (2) 200^$$. Reg. no. 1879:26. Omenak, Greenland ; on Monodon monoceros L.Collected by E. Whymper. (3) 2(J(J, i?, i juv. Reg. no. 1911:11:8:23031-33. Omenak, Greenland. A. M.Norman Collection. (4) 2(££, 4$?, 8$$ (immature). Reg. no. 1972:564:12. No locality data ; onMonodon monoceros L. University College, Dundee Collection. CATALOGUE OF WHALE-LICE 69 (5) i$. Reg. no. 1972:565:1. No locality data ; on Monodon monoceros L. Col-lected by Nuttall. (6) 5cJcJ, 3?$ (immature), 9 juvs. Reg. no. 1972:566:17. No locality data.University College, Dundee Collection. (7) 3 specimens (slide collection). No locality data ; on Monodon monoceros L.No collection data. Cyamus ovalis Roussel de Vauzeme, 1834 (1) 3°c?c??$> attached to section of skin. Reg. no. 1911:10:16:1-10. SouthGeorgia, 2.2.1911 ; on Eubalaena glacialis Muller. Collected by G. Cruick-shank. (2) I50&J??, attached to section of skin from lower maxilla. Reg. no. 1911:10:16:11-20. South Georgia, 15.3.1911 ; on Eubalaena glacialis Muller. Collectedby G. Cruikshank. (3) I0c££, 20 juvs. Reg. no. 1911:11:8:23034-53. Iceland; on Eubalaenaglacialis glacialis Muller. A. M. Norman Collection. (4) 2<3cT, 3?$, 30 juvs. Reg. no. 1911:11:8:23054-23063. Pacific Ocean; onEubalaena glacialis japonica Lacepede. A. M. Norman Collection. (5) 1503$$?, attached to section of skin. Reg. no. 1912:6:15:1-20. SouthGeorgia; on Eubalaena glacialis australis Desmoulins. Collected by G.Cruikshank. (6) 6<^, 8$$, 30 juvs. Reg. no. 1912:10:2:8-27. 6 miles N.E. of Flugga, Shetland.Collected by Lovett. (7) 300dc?$$> attached to three separate sections of skin. Reg. no. 1920:10:21:23-42. Stromness Whaling Station, South Georgia ; on right whale (upper jaw).Collected by J. Rasmussen and Alex Lange. (8) 7dtf, 7?$, 6 juvs. Reg. no. 1936:11:2:3483-3494. South Georgia; onEubalaena glacialis australis Desmoulins, whale no. 503. Discovery Collection. (9) i$, 2$$, i juv. Reg. no. 1936:11:2:3483-3494. Saldanha Bay, South Africa,26.8.1926 ; on Eubalaena glacialis australis Desmoulins, whale no. 1020.Discovery Collection. (10) 10^, 30$$, 100 juvs. Reg. no. 1972:571:140. Port Elizabeth. Collected byI. L. Drege. (11) 3<3<$> 20 juvs. Reg. no. 1972:572:23. Faeroerne Island, North Atlantic,62° N, 07° W, June 1898 ; on Eubalaena glacialis glacialis Muller, $. Nocollection data. (12) 2<&£, 2$$. Reg. no. 1972:573:4. No locality data; on Eubalaena glacialisjaponica Lacepede. Collected by W. N. Lockington. (13) 7c&£> IO??» 3° juvs. Reg. no. 1972:574:47. South Georgia ; from 'bonnet' ofright whale. Collected by P. Stammwitz. (14) 3^, 9$9, 10 juvs. Reg. no. 1972:576:22. No locality data ; on Eubalaenaglacialis glacialis Muller. Collected by Goldberg. (15) 20 specimens (dry collection). Port Jackson and South Africa. LeachCollection. yo R. J. LINCOLN & D. E. HURLEY Cyamus pacificus Liitken, 1873 (J) 5cTc?- Reg- n°- I&77-3' No locality data ; on Megaptera novaeangliae (Borow-ski). Collected by W. N. Lockington. Cyamus scammoni Ball, 1872 (J) 3&?» !?• Reg- n°- X877:3. No locality data; on Eschrichtius gibbosus (Erxleben). Collected by W. N. Lockington.(2) 30^$?. attached to section of skin. Reg. no. 1891:2:10:1-12. Wladiwostock, Victoria Bay, Sea of Japan. Collected by Fridolf Hook. Cyamus thomsoni Gosse, 1855 see Platycyamus ISOCYAMUS Gervais & Van Beneden, 1859 Isocyamus delphini (Guerin-Meneville, 1836) (1) 2<3c?. Reg. no. 1911:11:8:23067-68. No locality data; on Globicephalamelaena Traill (pilot whale). A. M. Norman Collection. (2) 4&?> 5??, 6 juvs. Reg. no. 1936:11:2:3585-3600. i4°45' N, i8°34'W, 27.10.1925 ; on Steno bredanensis Lesson (rough-toothed dolphin), Dolphin A.Discovery Collection. (3) i(J, i$, ii juvs. Reg. no. 1936:11:2:3583-3600. 14^5' N, i8°34' W, 27.10.1925 ; on Steno bredanensis Lesson, Dolphin B. Discovery Collection. (4) 9c?c£??- Reg. no. 1972:547:9. Tonnybwlch Beach, Aberystwyth, Cardigan-shire ; on stranded porpoise. Collected by P. Miles. NEOCYAMUS Margolis, 1955 Neocyamus physeteris (Pouchet, 1888) (1) 4$$. Reg. no. 1965:9:8:46-65. Talcahuano, Chile, 36°45'S, 73°i2' W,23.3.1961, on Physeter catodon L., $ 10-1 m, whale no. T6. Collected by R.Clarke. (2) i<J, 2?$. Reg. no. 1965:9:8:46-65. Paita, Peru, 05^5' S, 8i°io' W, 23.6.1959,on Physeter catodon L., $ 9-6 m, whale no. Pa2i. Collected by R. Clarke. (3) i$. Reg. no. 1965:9:8:46-65. Paita, Peru, 05°05'S, 8i°io'W, 23.4.1960,on Physeter catodon L., <$ 13-7 m, whale no. Pa6n. Collected by R. Clarke. (4) i?. Reg. no. 1965:9:8:46-65. Paita, Peru, 05°O5' S, 8i°io' W, 21.4.1959, onPhyseter catodon L., <$ 13-3 m, whale no. Pan. Collected by R. Clarke. (5) i<J, io$$, i juv. Reg. no. 1965:9:8:46-65. Paita, Peru, 05^5' S, 8i°io' W,25.1.1960, on Physeter catodon L., $ 10-2 m, whale no. 417. Collected by R.Clarke. PLATYCYAMUS Liitken, 1870Platycyamus thomj>soni (Gosse, 1855) (i) i<$. Reg. no. 1907:12:2:219. No locality data. University College, DundeeCollection. CATALOGUE OF WHALE-LICE 71 (2) i(£, i$. Reg. no. 1911:11:8:23077-78. No locality data; on Hyperoodonampullatus Forster. A. M. Norman Collection. (3) i$. Reg. no. 1956:10:10:347. No locality data. Collected by UniversityCollege, Dundee. (4) Holotype. Ann. Mag. not. Hist. (2) 16 : 30-31. Reg. no. 1856:131 dry coll.,Leach cab : 31. Portland Roads S. England ; taken from beneath the eye ofstranded Bottlenose whale, Hyperoodon ampullatus Forster. Leach Collection. SCUTOCYAMUS Lincoln & Hurley, 1974 Scutocyamus parvus Lincoln & Hurley, 1974 (i) Holotype ?, io$$ ovig., 21$$ immature, 28$$, 37 juvs. Reg. no. 1973:105.Off Peterhead, Aberdeenshire, North Sea, 21.7.1933 ; on Lagenorhynchusalbirostris Gray (white-beaked dolphin). Collected by Dr F. C. Fraser. Summary of hosts of whale-lice in the collections Host Mysticeti (whalebone whales) Gray whale, Eschrichtius gibbosus (Erxleben) Blue whale, Balaenoptera musculus (L.) Fin whale, Balaenoptera physalis (L.) Humpback whale, Megaptera novaeangliae (Borowski)North Pacific right whale, Eubalaena glacialis japonica LacepedeSouthern right whale, Eubalaena glacialis australis Desmoulins North Atlantic right whale, Eubalaena glacialis glacialis MiillerGreenland right whale, Balaena mysticeti L.Odontoceti (toothed whales)Sperm whale, Physeter catodon L. Bottlenose whale, Hyperoodon ampullatus Forster Pilot whale, Globicephala melaena TraillNarwhal, Monodon monoceros L. Whale-lice Cyamus scammoni DallCyamus balaenopterae BarnardCyamus balaenopterae BarnardCyamus boopis LiitkenCyamus pacificus LutkenCyamus ovalis R. de Vauzeme Cyamus erraticus R. de VauzemeCyamus gracilis (R. de Vauzeme)Cyamus ovalis R. de VauzemeCyamus erraticus R. de VauzemeCyamus ovalis R. de VauzemeCyamus ceti (L.) Cyamus boopis LutkenCyamus catodontis MargolisNeocyamus physeteris (Pouchet)Platycyamus thompsoni (Gosse) Isocyamus delphini (Guerin-Meneville)Cyamus monodontis LutkenCyamus nodosus L.Scutocyamus parvus Lincoln & Hurley White-beaked dolphin, Lagenorhynchus al-birostris Gray Rough-toothed dolphin, Steno bredanensis Isocyamus delphini (Guerin-Me'neville) Lesson REFERENCES HERSHKOVITZ, P. 1966. Catalog of living whales. Bull. U.S. natn. Mus. 246 : 1-259.LEUNG, Y.-K. 1967. An illustrated key to the species of whale-lice (Amphipoda, Cyamidae),ecto-parasites of Cetacea, with a guide to the literature. Crustaceana 12 (3) : 279-291. 72 R. J. LINCOLN & D. E. HURLEY LINCOLN, R. J. & HURLEY, D. E. 1974. Scutocyamus parvus, a new genus and species of whale-louse (Amphipoda : Cyamidae) ectoparasitic on the North Atlantic white-beaked dolphin.Bull. Br. Mus. nat. Hist. (Zool.) 26 : 59-64. Dr ROGER J. LINCOLN Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD Dr D. E. HURLEY NEW ZEALAND OCEANOGRAPHIC INSTITUTE P.O. Box 8009 WELLINGTON NEW ZEALAND A REVIEW OF LAEPHOTIS THOMAS, 1901(CHIROPTERA: VESPERTILIONIDAE) By J. E. HILL INTRODUCTION IN recent years the African genus Laephotis Thomas, 1901 has been consideredmonotypic (e.g. Ellerman, Morrison-Scott & Hayman, 1953 : 78) with one subspecies,L. wintoni wintoni Thomas, 1901 in Kenya and a second, L. w. angolensis Monard,1935 in Angola, Zambia, Botswana and the southern part of Zaire. However,Peterson (1971) while reporting a further example from Kenya reviewed much of thelimited material of Laephotis available in collections and was led to suggest thatwintoni and angolensis might represent distinct species, with the possibility that athird undescribed taxon might occur in Zambia. Setzer (1971) treated wintoni andangolensis as specifically distinct (as had Monard when describing the latter) anddescribed two further species, namibensis from the Namib Desert, South WestAfrica, and botswanae from Botswana, Zambia and southern Zaire. Although thisauthor referred to botswanae a number of the specimens from Zambia and southernZaire hitherto allocated to angolensis, he did not have available all of the material soreported : in particular he did not examine the specimen (in the Harrison ZoologicalMuseum, Sevenoaks, England) from Ndola, Zambia, which Peterson (1971 : 888)then thought could represent either the then unknown female of wintoni or possiblyan unnamed taxon. More recently, Peterson (1973 : 602), in describing the firstknown female of wintoni has suggested that the specimen from Ndola may be moreclosely related to botswanae than to other specimens from Zambia or Zaire. Current accessions to the collections of the British Museum (Natural History)have included three female specimens of L. wintoni from Ethiopia, whence the genushas been hitherto unreported. These have led to a further examination of the sevenexamples of Laephotis already in the collections in London, together with six fromZaire in the Musee Royal de 1'Afrique Centrale, Tervuren, Belgium, and of twospecimens in the Harrison Zoological Museum, one the apparently enigmatic examplefrom Ndola. The four species recognized by Setzer (1971) are retained : wintoniand namibensis are apparently very similar but differ much more sharply fromangolensis and botswanae than do these from each other. SYSTEMATIC SECTION LAEPHO TIS Thomas, 1901Laephotis Thomas, 1901 : 460. TYPE SPECIES. Laephotis wintoni Thomas, 1901. Head low, flat-crowned ; muzzle broad, flat, the nostrils opening sublaterally ;ears moderate to large, when laid forward reaching to or beyond the tip of the Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 74 J- E. HILL muzzle, more or less triangular in outline ; anterior margin of the ear originatingnear the centre of the forehead, lacking any posteriorly directed basal lobe but witha distinct basal fold ; anterior margin convex proximally then less so or nearlystraight; tip rounded ; posterior margin of ear slightly emarginated just below tip,otherwise convex, with moderate rounded or semi-circular antitragal lobe, terminat-ing just behind the angle of the mouth. Tragus (Fig. i) as a rule rather wide, aboutone-third the length of the ear, its anterior margin slightly concave, tip bluntlypointed, anteriorly directed ; posterior margin of tragus convex or angular, a smalltriangular lobe at its base. Calcar extending along about two-thirds of the uropata-gial margin, with small, rounded post-calcarial lobe. Seven palate ridges in L.wintoni from Ethiopia, the first and second uninterrupted, the last reduced but alsocomplete (Fig. 2) : however, Peterson (1973 : 602, fig. i) reported six ridges only in asubadult female of L. wintoni from Kenya. Skull with slightly flattened braincase ; slight lambdoid but no sagittal crest;rostrum rather narrow, unexpanded, with little supraorbital inflation; a shallow,median rostral depression ; zygomata slender ; narial emargination narrow, broadlyU-shaped but slightly angular, extending posteriorly to or a little beyond a linejoining the anterior margins of the anteorbital foramina ; pre-palatal emarginationextending laterally a little beyond the inner faces of i2~2 and posteriorly almost to aline joining the posterior faces of c1"1; palate long, rather narrow, strongly domed ;maxillary toothrows usually almost parallel; post-palatal extension moderate tolong, its length from a line across the rear faces of m3~3 to the tips of the pterygoidhamulars approaching or equal to the length of the palate from the same line to theback of the pre-palatal emargination (Table 2) ; short, broad post-palatal spine ;pterygoid hamulars strong, sharply deflected inwards ; bullae inflated, large. 21 I ^ Dentition i -, c -, pm -, m - = 32. Inner upper incisor (i2) long, with secondary 3i 23 postero-internal cusp extending almost to its tip ; i3 small, a little wider than long,its main cusp barely extending above the cingulum of i2, with small internal basalcusp, in toothrow, touching i2, separated from c1 by a narrow space ; c1 normal;pm4 a little wider than long, in contact with c1; m3 not reduced, with meta-cone and three commissures ; ij_3 tricuspid, imbricated, i3 slightly the largest ;ct reduced, weak, little higher than pm4 ; pm2 reduced, about one-half the heightand one-quarter or a little more the crown area of pm4, tightly compressed in tooth-row ; m3 little reduced, the hypoconid and entoconid well developed. The genus occurs in Ethiopia and Kenya, in southern Zaire, Zambia, Botswana,South West Africa and in Angola. It appears to be related to Eptesicus. Laephotis wintoni Thomas, 1901 Laephotis wintoni Thomas, 1901 : 460. Kitui, Kenya, c. 3500 ft. Size large for the genus (length of forearm 36-41 mm, condylobasal length15-2-15-8 mm) with very large ears (length from meatus 21 mm or more) ; tragusbroad, at its widest at about one-third of its height from its base, its posterior marginstrongly convex, sometimes slightly angular (Fig. la). Dorsal surface a shade of REVIEW OF LAEPHOTIS THOMAS, 1901 75 mid-brown, the pelage dark based with brown, slightly coppery tips ; ventral surfacesimilar but paler, the hairs similarly dark based but with paler brown or, especiallyposteriorly, with buffy white tips. Skull elongate with long narrow rostrum ;palate long, the maxillary toothrows nearly parallel; post-palatal extension verylong, its length from a line across the rear faces of m3"3 to the tips of the pterygoidhamulars equal or nearly equal to the length of the palate from the same line to theback of the pre-palatal emargination ; bony post-palate long, its length from a lineacross the rear faces of m3~3 to the anterior edge of the mesopterygoid fossa exceed-ing the distance from the anterior edge of the mesopterygoid fossa to the tips ofthe pterygoid hamulars (Table 2). The female of wintoni has remained unknown until recently. Peterson (1971 : 888)speculated that a female specimen (1.2533) m the Harrison Zoological Museum,Sevenoaks, from Ndola, Zambia, considerably smaller than the male examples thathe had examined, might represent it. Setzer (1971 : 264) noted that data presentedby Peterson (p. 886) for specimens of Laephotis in the British Museum (NaturalHistory) from Zaire reveal a slight sexual variation, females being on the average alittle larger than males in external and cranial measurements. However, the skullhad not then been extracted from the sole male specimen (B.M. 57.435, in alcohol)in this series (B.M. 57.435-438) which apparently represents two species (videinfra). The measurements given by Peterson show a female (B.M. 55.1135) fromZambia to be externally a little larger on the whole than a male (B.M. 55.1134)obtained at the same locality but so only in a limited number of cranial dimensions.Peterson (1973 : 601) has reported a subadult female of wintoni from Kenya which iscomparable to male specimens in most dimensions. The three female specimens ofwintoni now available from Ethiopia are consistently larger in most respects than twomales from Kenya (Table i) or than the male and subadult female specimensreported from Kenya by Peterson (1971 : 885, 886 ; 1973 : 601). The number ofspecimens available is too small to establish any firm evidence of sexual variation insize, and the possibility remains that the population of wintoni in Ethiopia consistsof larger individuals than does the population in Kenya. The species occurs (Fig. 3) in KENYA (Thomas, 1901 : 460 ; Harrison, 1961 : 292 ;Hayman & Hill, 1971 : 49 ; Peterson, 1971 : 885, 887, fig. i (head), pi. i (fig. 2)(skull) ; Setzer, 1971 : 262, fig. ic (tragus)) ; Peterson, 1973 : 601, fig. i (soft palate),and in ETHIOPIA (B.M. 72.4397-4399, from Koka, Shoa Province, 8°27' N, 39°o6' E,at 1700 m). Laephotis namibensis Setzer, 1971 Laephotis namibensis Setzer, 1971 : 259, 263, fig. id (tragus). Kuiseb River, near NamibDesert Research Station, Gobabeb, South West Africa. No specimens are available for examination, the species being represented atpresent only by the female holotype and paratype in the collections of the UnitedStates National Museum of Natural History at the Smithsonian Institution, Washing-ton. From the description namibensis is characterized by its very large ears ; well-developed tragus and antitragus ; pale coloration ; large, relatively long and narrow 76 J. E. HILL skull; relatively long, narrow palate ; and by great inflation of the bullae. It isevidently very similar to wintoni but has larger ears (length 24-25 mm) which arebroader at the base, and a larger tragus. It is markedly paler in colour than wintoni,the dorsal surface pale drab, the ventral surface paler, the hairs tipped with white.Cranially, the braincase is more domed than in wintoni, the postorbital region moreconstricted, the zygomatic arches more arcuate in the vertical plane, the maxillarytoothrows more nearly parallel and the bullae more inflated. Although the skull israther longer than in male specimens of wintoni from Kenya, it is only marginallylonger than in female specimens from Ethiopia (Table i). Laephotis angolensis Monard, 1935 Laephotis angolensis Monard, 1935 : 45- Tyihumbwe" (Chiumbwe River, a tributary of theKasai, 15 km west of Dala), Angola. Smaller than wintoni or namibensis (length of forearm 32-35 mm, condylobasallength c. 12-9 mm) with smaller, narrower ears (length from meatus less than 16 mm) ;tragus (Fig. ib) smaller than in wintoni or namibensis (vide Setzer, 1971 : 263, fig. i),less broadened and rather more spatulate. Colour apparently similar to that ofwintoni but the only available specimens are in alcohol and have been so for someyears. According to Setzer (1971 : 260) namibensis is paler than angolensis. Skullvery much smaller than in wintoni or namibensis (Table i), the bony part of the post-palatal extension much shorter than in wintoni, its length from a line across the rearfaces of m3~3 to the anterior edge of the mesopterygoid fossa less than the distancefrom the anterior edge of the mesopterygoid fossa to the tips of the pterygoidhamulars, not exceeding it as in that species (Table 2). A number of specimens have been allocated hitherto (Hayman & Hill, 1971 : 49)to angolensis. These include a further example from Angola, from a locality 35miles east of Dande, in the collections of the American Museum of Natural History,New York (Hill & Carter, 1941 : 49) which is accepted as representative of angolensisby Setzer (1971 : 260 et seq.j. Specimens from Zambia, Zaire and Botswana wereregarded previously as angolensis but Setzer (1971 : 260) took the sole example fromBotswana as the holotype of a new species, botswanae, to which he referred such ofthe material from Zambia and Zaire as he was able to examine. Two (B.M. 55.1134-1135) of the three specimens so far reported from Zambia were seen by Setzer : thethird is the example (Harrison Zoological Museum 1.2533) discussed by Peterson(1971 : 885, 888 ; 1973 : 602) which apparently also represents botswanae (vide infra). The specimens (B.M. 57.436, 57.438) from Zaire seen by Setzer are from a series often collected at Mumene, 70 km east of Lumbumbashi ( = Elizabethville), Katanga,which, with a further three from the nearby locality of Musonge, 2 km to the west,were originally reported by Hayman (1957 : 43). This author examined ten of these ;four (B.M. 57.436-438 from Mumene, B.M. 57.435 from Musonge) are in the collec-tions of the British Museum (Natural History) and a further six (M.R.A.C. 26.402-406 from Mumene, M.R.A.C. 26.407 from Musonge) in the Musee Royale de 1'AfriqueCentrale, Tervuren. Hayman, Misonne & Verheyen (1966 : 50) list nine specimens REVIEW OF LAEPHOTIS THOMAS, 1901 77 at Tervuren but no more than six can be found (Thys van den Audenaerde, in litt.),corresponding to the total examined in London by Hayman. Of these, eight must be referred to botswanae but two, B.M. 57.435 and B.M. 57.437apparently represent not botswanae but angolensis as it is understood by Setzer(1971 : 260 et seq.). They agree closely with the descriptions of this species byMonard (1935 : 45) and Hill & Carter (1941 : 49) : their measurements (Table i) aresimilar to those of the holotype and to those of the second Angolan example as theyare reported by Hill & Carter (p. 176) and Setzer (1971 : 261). The specimens differfrom the others in the series from Zaire in smaller size, narrower tragus with generallya less angular posterior margin, and in slightly shorter, narrower rostrum and palate.The tragus of B.M. 57.435 is a little wider than that of B.M. 57.437 while amongbotswanae the tragus of M.R.A.C. 26.404 is rather narrow, with its posterior margina little less angular than is usual in that series. Setzer (1971 : 262) notes thatbotswanae has a more massive rostrum than angolensis but the rostrum in B.M. 57.435and B.M. 57.437 is no more than slightly shorter and not less massive when com-pared with the narrowest of botswanae. The rearward extension of the occipitalcrests, said by Setzer to be less in botswanae, does not differ consistently in the speci-mens examined. The toothrows of B.M. 57.435 are more divergent posteriorly thanthose of B.M. 57.437 to approach specimens referred to botswanae although, as Setzernotes of this species, its toothrows are generally more divergent posteriorly than inangolensis. The specimens available do not confirm the statement by Setzer that thebullae in botswanae are relatively as well as actually smaller than in angolensis : ifB.M. 57.435 and B.M. 57.437 correctly represent angolensis then the bullae of bots-wanae are proportionately about the same in size and in actual terms a little larger.That angolensis and botswanae might occur together was suggested by Setzer (1971 :262) : records (Fig. 3) of angolensis are thus restricted to ANGOLA (Monard, 1935 : 45 ;Hill & Carter, 1941 : 49, 176 ; Setzer, 1971 : 260, 263, fig. la (tragus)) and probablyZAIRE (Hayman, 1957 : 43 (in part) ; Hayman, Misonne & Verheyen, 1966 : 50 (inpart) ; Peterson, 1971 : 885 (in part)). Laephotis botswanae Setzer, 1971 Laephotis botswanae Setzer, 1971 : 260, 263, fig. ib (tragus). 50 miles west, 12 miles south ofShakawe, Botswana. Size between angolensis and the large species wintoni and namibensis (length offorearm 34-38 mm, condylobasal length (13-5-14-3 mm) ; ears and tragus generallya little larger than in angolensis (length of ear from meatus more than 16 but lessthan 18 mm) but markedly smaller than in either of the large species. Tragus(Fig. ic) usually with a distinct angularity in its posterior margin at its widest point.Dorsal surface similar in colour to wintoni ; ventral surface paler than in that species,the hairs more liberally tipped with buffy white rather than pale brown. Rostrumrelatively long, narrow ; maxillary toothrows slightly divergent posteriorly ; post-palatal region as in angolensis, the length of its bony part from a line across the rearfaces of m3~3 to the anterior edge of the mesopterygoid fossa less than the distance 78 J. E. HILL from the anterior edge of the mesopterygoid fossa to the tips of the pterygoidhamulars (Table 2). The specimen (Harrison Zoological Museum 1.2533) from Ndola, Zambia, whichPeterson (1971 : 888) thought might represent the female of wintoni or possibly an 5 mm FIG. i. Tragus of (a) Laephotis wintoni; (b) L. angolensis ; (c) L. botswanae. FIG. 2. Palate ridges of Laephotis wintoni. REVIEW OF LAEPHOTIS THOMAS, 1901T 79 FIG. 3. Distribution of • Laephotis wintoni ; A L. namibensis ; T L. angolensis L. botswanae. undescribed taxon and later (1973 : 602) closely related to botswanae proves in factreferable to this species. Although larger than specimens from Zaire or than those(B.M. 55.1134-1135) from Solwezi Boma, Zambia (i2°io'S, 26°3o' E) its tragus(from the dry specimen) is angular posteriorly and the specimen agrees precisely incoloration with the other examples from Zambia, differing in this respect from theholotype of wintoni in its distinctly paler, more buffy white underparts, especiallyposteriorly. Specimens referable to this species have been recorded (Fig. 3) from BOTSWANA(Smithers, 1968 : 48, 49 (map), fig. a (head) (as angolensis) ; Setzer, 1971 : 260, 263,fig. ib (tragus) ; ZAMBIA (Ellerman, Morrison-Scott & Hayman, 1953 :78 (asangolensis} ; Ansell, 1957 : 538 (as angolensis) ; Hayman, 1957 : 43 (as angolensis} ;Ansell, 1960 : 21 (as angolensis) ; Peterson, 1971 : 885, pi. i (fig. 2) (skulls, includingspecimen from Ndola) (as angolensis and Laephotis sp.) ; Setzer, 1971 : 260) ; ZAIRE(Hayman, 1957 : 43 (in part, as angolensis) ; Hayman, Misonne & Verheyen, 1966 :50 (in part, as angolensis) ; Peterson, 1971 : 885 (in part, as angolensis) ; Setzer,1971 : 260). 1 •«! i rt rt .0 _U oj o3 ri rf PH &t OH ^ ^ Aweooq e c a e '3 13 13 t^ > rt C rf ^ .r-t .^H .^H nO 4)0) 5l)<l)DiUiU4)a>(U'13'13^-1 *-HHHhHI-MWWhH C/) (/) - eiuo v"> 7*" T ?*" "9 !P Y"> ^< NN WNNNNNNNNNNN OOt^ MOONOiOO MONO* 10 »o u"i *o 10 *o »oui oiQiduioo o »o co i^ • • • • 1 jo q^Sugq ° 2 2 2 eui-o 9 9 r 91 N N 9 9^9 •^•M- inin^^inin 10^1-10 OOON ^-MION NON0000 'IONONONON (ONON •0 Cx ON -00 ON m ON NO roOt^^^^N .«OOO rt- io!n inioio^w^inin in*n O O0t^ Tt-ONQOOOOO .Oro V roro T}-ro4j.Tt-rj-rj-ro 'fTj-•^ ^f ^ .(ONVOOOOO roiO1* ^BJAL ob ob ob ob ob ob ob 9S'BDUl'Bjq 00 OO O 00 CO t^ 00 t^ ON ^^ tn ro c^> . | vo vo vo r^* in in vo 9S'BOurejq ^ ro n- 10 o «o o vO 0000 OOO|Tf-rOMN OM^J- O' M OO O P q^PTAY ^ ^ ^ ^ ^ «^ ^ vo oo t^o r^t^c^c^ t>-t^t^. f^ w ro i CO oo o M- ^ " " VO .TM ro N t^ t^ •• in TAV Ol^moSXz 1 1 ON ob 00 ON ON ON ' 0 ob1 ob11111 ''ob 4) • « ON M ^JOja^UI ^.S'69'T rororororororo roroiqOBI SSOJOB r>i -r 10 •* ro ro roro roro^rorororo rororoO0t^ ONO>-IMON Oroin ,2 o> fe " ^ ° iqjo jo q^piAV 10 10 in in in^'B^iqjoa^u'B in \o •<*• t^ vo •^•M* *n ***> in if) *n ~*t~ ininin SSOJOB q^piA\ * •*• •* -^- Tj- q^§U9'T ro ro ro ro ro UIUBOOlApUOQ 2" IT 1? 1? 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IBS^qofApuoQ >o 10 10 >n 10 10 10 roN I'rorororo rororj- G -a 5 jmilS JO OOOOMOrOrOM tO«O t^ OOC^ in inrOOro rOO o 2 £- •g n 3 ^ 3u9j ^sa^Bajf) minovovoovo oo ro roro ^•I'^t-Tf^J-Tf- Irt-io Jo fc i^ i^gui taoij _ * H U-N ^ ^_ OO> MiOinrocoOiMTt' •^ S , *8 S "i fe||6ffi 1^9 jo q^Sugq ^ N N o3 N N 1 oiin f^vbooot^^-^111 « OT fl 8 «a U3 - O <n ™ ( •^- inro rONOoOOOO-^-Mt^NrOO s § i | i^ 10 U1SU9T t^ovof--666 obob ( X3S ^O'b'XDCH-OfOfCH- OfO N "con ** « ""> ^ _ •jfr^-'ooinr^oonN MM[ UOI^BJ;ST§9^I m^O-?O^ONON N N,n2.ror^rororo •^-•'t-"•^vOvO^-Th-^- "ororo •ONCOVO^J.J.^ i^ i_( i_, .J2 ^•s* « s a si ^N in^i- f^r^c^t^io^ooo^-io^ ro roro rococororororocororororo*O *O O Of Ot- CH- CH- O O O Of Of *o Of CH- ia- * r»-. o ^-- ^Q BM British Museum (NaturHZM Harrison Zoological MuROM Royal Ontario MuseumUSNM United States NationalAMNH American Museum of J 1UT? AP M.,c/So T?™rolo /!Q 1* \trir REVIEW OF LAEPHOTIS THOMAS, 1901 81 TABLE 2 Palatal measurements (in mm) of Laephotis o fc _o I •u 2'be K wintoMZ BM 1.5.6.5*HZM 2.2030BM 72.4397BM 72.4398BM 72.4399 (?) angolensisBM 57.435BM 57.437 botswanaeBM 57.436BM 57.438MRAC 26.402MRAC 26.403MRAC 26.404MRAC 26.405BM 55.1134BM 55.1135HZM 1.2533 $ ,-H <U O 0 _O 03 bo 13 'C ko *4H o rt <u ~y w 43 w O 'O •+•1 <- o ft -M 13 VH 13 g 00 O *-" <D •*•* 0 ^•2 o3 ft b „ ft. 2 M bC o ft^ P s a <u o3 i 03 hi ^ rO -4-* C/3 O ^ yj C/3 A c £ £.!"s w 9j X8 S -2 " " S CO o oS 0 03 ^ •g SJ1 O '•£ 03 ctf Sio <u C >, * .2 £ 111 ^ O o3 <u -2 o JT! *-M rt o; P c^ a; a^ d o 3.§ * I 3 m r^ "bo S -S -e 2 1 & I'S * 2 c o 1 a ft t; HH o 1 £"* P 53 -a tH -LJ O § S^ 2^'S 13 fit I •*-! 'a'C s a| fl O en r! 00 SJ±3 !D H •« PU WJ A O >> is en 1-1 fl o be tS (U *-i £ 43 £ "§> S HH fli be ^ ^ be « ^g S ft «> S -^ J"ft 1 a 7-9 6-5 4-4 2-1 1-9 4-0 Kenya - - - - - - Kenya 8-1 6-8 4-4 2-4 1-8 4-2 Ethiopia 8-2 6-4 4'2 2-2 2-0 4-2 Ethiopia 8-5 6-8 4'3 2-5 1-9 4-4 Ethiopia 6-2 4-8 3-6 1-2 2-O 3-2 Zaire 6-1 4-8 3'5 i-3 1-9 3-2 Zaire _ 5'4 4-0 1*4 _ _ Zaire 6-7 5-2 3-8 1-4 2-0 3'4 Zaire 6-6 5'2 3'7 i '5 2-O 3'5 Zaire 6-6 5'2 3'7 I-5 1-9 3'4 Zaire 6-4 5'2 3-8 1-4 1-9 3'3 Zaire 6-5 5-3 3-8 1-5 1-9 3'4 Zaire 6-5 5'2 3'7 I-5 - - Zambia 6-4 5'2 3'7 I-5 1-9 3'4 Zambia 7-1 5-6 4-1 I-5 2-O 3'5 Zambia BM British Museum (Natural History), London.HZM Harrison Zoological Museum, Sevenoaks.MRAC Mus6e Royale de 1'Afrique Central, Tervuren.* Holotype. SUMMARY The majority of specimens of Laephotis hitherto reported in the literature areallocated to the appropriate one of the four species recognized in the most recentstudy of this genus, and these are briefly reviewed. Female specimens of L. wintoniare recorded from Ethiopia, whence until now the genus has been unreported. ACKNOWLEDGEMENTS My thanks are due to Dr M. J. Largen, of the Natural History Museum, HaileSelassie I University, Addis Ababa, who obtained the specimens from Ethiopia that 82 J. E. HILL led to this study and which have been donated to the British Museum (NaturalHistory) : to Drs M. Poll and D. Thys van den Audenaerde of the Musee Royale de1'Afrique Centrale, Tervuren, for the loan of the material of Laephotis in their care,and to Dr D. L. Harrison, who made available the specimens in his collections atSevenoaks. REFERENCES ANSELL, W. F. H. 1957. Some mammals from Northern Rhodesia. Ann. Mag. nat. Hist. (12), 10 : 529-551. 3 tabs. 1960. Mammals of Northern Rhodesia. Lusaka: Government Printer. EIXERMAN, J. R., MORRISON-SCOTT, T. C. S. & HAYMAN, R. W. 1953- Southern African mammals 1758 to 1951: a reclassification. London : British Museum (Natural History).HARRISON, D. L. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. Jl E. Africa nat. Hist. Soc. 23 : 286-295. HAYMAN, R. W. 1957. Further notes on African bats. Rev. Zool. Bot. afr. 56 : 41-45. & HILL, J. EDWARDS. 1971. In: Meester, J. & Setzer, H. W. (Eds). The Mammals of Africa. An identification manual, i. Chiroptera. Washington: Smithsonian Institution. MISONNE, X. & VERHEYEN, W. 1966. The bats of the Congo and of Rwanda and Burundi. Annls Mus. r. Afr. cent. Ser. 8°, Sci. Zool. No. 154, i-viii, 1-105, 2O P^8-HILL, J. ERIC & CARTER, T. D. 1941. The mammals of Angola, Africa. Bull. Am. Mus. nat. Hist. 78 : 1-211, 36 figs., 17 pis.MONARD, A. 1935. Contribution a la mammalogie d'Angola et Prodrome d'une faune d'Angola. Archos Mus. Bocage, 6 : 1-314, 47 figs.PETERSON, R. L. 1971. Notes on the African long-eared bats of the genus Laephotis (family Vespertilionidae). Can. J. Zool. 49 : 885-888, i fig., i pi., i tab. 1973- The first known female of the African long-eared bat Laephotis wintoni (Vesperti-lionidae : Chiroptera). Can. J. Zool. 51 : 601-603, i fig.SETZER, H. W. 1971. New bats of the genus Laephotis from Africa (Mammalia : Chiroptera). Proc. biol. Soc. Wash. 84 : 259-264, i fig., i tab.SMITHERS, R. H. N. 1968. A checklist and atlas of the mammals of Botswana (Africa). ? Salisbury : Trustees of the National Museums of Rhodesia.THOMAS, O. On a new genus and species of Vespertilionine bat from East Africa. Ann. Mag. nat. Hist. (7), 7 : 460-462. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD THE GENUS FINMARCHINELLA SWAIN 1963(CRUSTACEA : OSTRACODA) AND ITS SPECIES By JOHN W. NEALE SYNOPSIS Six species of the genus Finmarchinella from the Northern Hemisphere are found to fall intotwo groups for one of which the new subgenus Barentsovia is instituted. The confusion surround-ing the four North Atlantic and Arctic species is analysed and the new species Finmarchinella(Barentsovia} curvicosta is described. INTRODUCTION IN 1963 Swain established the genus Finmarchinella with Cythereis finmarchica Sars1866 as the type species. In 1957, however, Mandelstam had described the newspecies Nereina barentzovoensis from Russian Harbour, Novaya Zemlya, as thegenotype of his new genus Nereina. In 1966 Ishizaki pointed out that Nereina wasused for a gastropod described by Cristofori and Jan in 1832 and that Mandelstam'sgenus required a new name. The following year, Hazel (i967a, p. 18) examined theclassification of Recent Hemicytheridae from off North-eastern North America andregarded Finmarchinella finmarchica and Nereina barentzovoensis as congenericplacing them both in Nereina and considering Finmarchinella to be a junior synonym.Subsequently Hazel (igGyb) noted that Nereina was preoccupied and since Fin-marchinella was available he validated Swain's genus which thus contained these twospecies and Cythereis angulata Sars 1866. In addition Nereina japonica Ishizaki1966 from the Miocene and Pliocene of Japan and Finmarchinella uraniponnicaIshizaki 1969 from the Recent seas of western Honshu also belong here. These fivespecies, together with a new Arctic species fall into two distinct groups. F. finmar-chica and F. uraniponnica are regarded as falling within the strict interpretation ofFinmarchinella ; the remaining four species are separated and placed in the newsubgenus Barentsovia based on F. (B.) barentzovoensis (Mandelstam). The principal aim of this paper, however, is to sort out the confusion that has arisenbetween the occidental species of the subgenus Barentsovia and place their taxonomyon a sound footing. This has been done on the basis of carapace characters sincesoft parts have not been available for study in most species. The uncontroversialFinmarchinella s.s. group is dealt with first. LOCATION OF MATERIAL Specimens preceded by 'BM' are in the collections of the British Museum (NaturalHistory), those preceded by 'HU' are in the collections of Hull University. SYSTEMATIC DESCRIPTIONS Genus FINMARCHINELLA Swain 1963TYPE SPECIES. Cythereis finmarchica Sars 1866. Bull. Br. Mus. nat. Hist. (Zool.) 27, 23* 84 JOHN W. NEALE Subgenus FIN MARC BIN ELL A Swain 1963 Finmarchinella (F.) finmarchica (Sars 1866)(PI. i, figs. 6, 7 ; PL 2. figs, i, 5 & n) Cythereis finmarchica Sars, 1866 : 41. Hemicythere finmarchica, G. O. Sars, Sars, 1925 : 185-6, pi. LXXXV, fig. 3. DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Male left valve B.M. No. M aaSyA PI. i, fig. 6 0-688 0-331 Female left valve B.M. No. M 32876 PI. i, fig. 7 0-720 0-392 PI. 2, fig. ii Male carapace B.M. No. M 32876 PI. 2, fig. i 0-701 0-328 0-318 Female carapace B.M. No. M 32870 PI. 2, fig. 5 0-746 0-405 0-389 DISCUSSION. This species presents no difficulties in interpretation and only theoriginal reference and the original author's later figure are given here in the synonymy.The characteristic features are the rather 'smoothed-out' appearance with well-developed reticulation and pitting but a lack of strong costation. In dorsal view theoutline is smooth and more oval than in species of the subgenus Barentsovia (cf.PL 2, figs, i, 5 and PL 2, figs. 2-4). As in other species of the genus, sexual dimor-phism is very marked with the males much slimmer and lower in proportion to thelength than the females (PL i, figs. 6, 7 ; PL 2, figs, i, 5). DISTRIBUTION. Since this species has not hitherto been confused with any otherit is possible to use records in the literature to plot its distribution with some con-fidence (Fig. i). Sars originally described F. finmarchica from some empty shellsfrom shell sand in Oxfjord, Finmark, and subsequently found it at Korshavn on theNorwegian coast further south. Besides Sars localities it has been found in Vadsoand Lang Fjords (Brady & Norman, 1896, appendix). In the Norwegian andBarents Sea areas the author has found it in samples from Ernest Holt Station 2(70°29' N, I7°27' E) where it accounted for 2-18 per cent of the specimens, Station 6(69°54' N, I7°oo' E, 1-62 per cent of the fauna), H.M.S. Vidal Station 29 (c. 68°44' N,4i°23'5' E) and Station 46 on the Spitzbergen Shelf (75°n-2' N, 22°i4' E, 16-47 Percent). Widely distributed in high latitudes, it was recorded from three localities inthe Davis Strait and has been found in material from the Hunde Islands (68°52' N,53°07' W, 1-58 per cent), Holsteinsborg Harbour, Greenland (66°55' N, 53°25' W,7-73 per cent), off Cape Flora, Franz Joseph Land (79°57' N, 50°oi' E, 1-25 per cent)and Russian Harbour, Novaya Zemlya (76°i3' N, 62°4o' E, 0-97 per cent). Hazel(i967a) found it at Ungava Bay (6o°o8' N, 67°47' W) and gives a number of localitiesin the Gulf of Maine and on the Atlantic Shelf (Hazel, I967a, 1970). It was fairlywidely recorded round Scotland and England by Brady (1868) and Brady and Nor-man (1889), many of whose records can be confirmed by reference to material in theHancock Museum, Newcastle-upon-Tyne ; Ostenfeld and Wesenberg-Lund (1909) THE GENUS F IN M ARCH IN ELLA SWAIN Finmarchinella (F.) finmarchica (Sars 1866)f=fossil ZENITHAL EQUAL AREA FIG. i note it from the North Sea and the author has found it in the Celtic Sea in core sample5652 (49°38' N, 9°i9' W) and from both top and bottom of the 88 cm core sample5674 (49°5o-9' N, 9°o8' W). Vos (1957) added the Chateau du Taureau (entranceto the River Morlaix Estuary). It has been widely encountered in the Bay of Biscay(Peypouquet, I97ib, gives 22 localities) but it is always rare and Peypouquet (i97ia)and Moyes & Peypouquet (1971) regard it as representing a palaeothanatocoenose ofHolocene and Pleistocene age. This could explain the record from St Vincent, CapeVerd, by Brady & Norman (1889) but on balance one is inclined to agree with Hazel(i967a) who thought that it was most probably a mis-identification. It is wellknown as a fossil in the post-Tertiary deposits of Britain (Brady, Crosskey & Robert-son, 1874), Hazel (1968) has found it in the Pleistocene of a number of submarinecanyons off the eastern seaboard of the United States and Lev (1969) has found it inhis Normanicythere concinella community in the Kazantsevian deposits of the rightbank of the River Yenesei and Cheshkaya Bay. Thus, F. finmarchica is essentiallyan Arctic and boreal-Arctic species whose southern limit appears to be about Ushant(Vos figures soft parts from the Chateau du Taureau locality). 86 JOHN W. NEALE Finmarchinella (Finmarchinella) uraniponnica Ishizaki 1969 Finmarchinella uraniponnica Ishizaki, 1969 : 217-218, pi. 26, figs. 12, 13 ; pi. 24, fig. 4 ; Ishizaki,1971 : 83, pi. 3, fig. 3. This species is most easily differentiated from F. finmarchica by its more asym-metrical anterior and more sloping posterior margins in lateral view. The slightconcavity in the anterior half of the ventral margin seems a little more accentuatedalso. Not so far recorded outside Honshu, it appears to be rather rare in WesternHonshu where it was only found at one station in the Nakanoumi Estuary. Here itwas accompanied by Cushmanidea subjaponica and Hermanites tosaensis which werealso confined to this station, and seventeen other species which had a much widerdistribution. In N.E. Honshu in Aomori Bay, however, Ishizaki (1971) has shownthat it occurs much more commonly. Subgenus BARENTSOVIA subgen. nov. TYPE SPECIES. Finmarchinella barentzovoensis (Mandelstam, 1957). DIAGNOSIS. A subgenus of Finmarchinella which differs from Finmarchinella s.s.in the development of marked costation, particularly in the posterodorsal part of thevalve and in the prominent development of the subcentral and eye tubercles. As shown below, the three occidental species of Barentsovia have been the subjectof confusion amongst themselves and it is impossible to use the records in the litera-ture to establish their distribution unless there are accompanying illustrations whichshow the distinctive features. In consequence the distributions on the map (Fig. 2)use only the latter and material actually verified by the author. Finmarchinella (Barentsovia) barentzovoensis (Mandelstam, 1957)(PI. i, figs. 4, 5, 9 ; PI. 2, fig. 12) Cythere clathrata, Reuss var. nuda Brady, 1866 : 376-377, pi. 59, figs. 9, 10. Hemicythere angulata (Sars) Akatova, 1946 : 227, fig. 5. Nereina angulata (Sars) Hazel, 1967 (pars) : 19, pi. i, figs. 7, n. (non pi. i, figs. 8, 9, 10 = F. (B.) curvicosta.)Nereina barentzovoensis Mandelstam, 1957 : 180, pi. 3, figs. 7, 8. DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height (mm) (mm) Male left valve HU. 8o.R.23 PI. i, fig. 4 0-766 0-392 Female left valve HU. 8o.R.24 PI. i, fig. 5 0-831 0-457 Female left valve HU. 8o.R.25 PI. i, fig. 9 0-821 °'447 PI. 2, fig. 12 DISCUSSION. This species has been confused with F. (B.) angulata (Sars). Theearliest record of this species is that of Brady (1866) who figured it as a variety of THE GENUS FINMARCHINELLA SWAIN Finmarchinella (B.) angulata (Sars 1866)Finmarchinella (B.) barentzovoensis (Mande!stam1957)Finmarchinella (BJ curvicosta sp.nov. IHM. EQUAL AREA FIG. 2 Cythere clathrata Reuss which he called nuda, giving as localities the Hunde Islandsand Norway. His figures show the characteristic ornamentation and this is con-firmed by a slide from the Hunde Islands in the Brady Collection in the HancockMuseum labelled 'Cythere angulata ? Sars = var : nuda Brady'. This contains fivespecimens, male and female right valves and two female left valves of F. (B.)barentzovoensis and a female right valve of F. (B.) angulata (Sars). Later Brady &Norman (1889) placed Cythere clathrata var. nuda in synonymy with Sars' speciesand the confusion has persisted. Akatova (1946) figured a specimen as Hemicythereangulata (G. O. Sars). More recently Hazel (i967a), commenting on the difficulty ofinterpreting Mandelstam's species from the published figures and lacking comparativematerial, figured two specimens of this species as Nereina angulata (Sars). The dif-ferences, however, are clear and well seen by comparing PI. i, figs. 4, 5 with PI. 2,fig. 8, and PI. 2, figs. 10, 12. The species is most easily recognized in lateral view bythe strong rib which follows the anterior margin in its dorsal half and then curves into the body of the valve at about mid-height. The posterodorsal quarter of the valve 88 JOHN W. NEALE surface is also particularly distinctive with two oblique ribs which run postero-dorsally from the subcentral tubercle and are linked by a very well-developed in-flation or tubercle at their posterodorsal termination. The lower of these two ribs isthe stronger. DISTRIBUTION. Mandelstam's original material came from Russian Harbour,Novaya Zemlya, and material from this locality collected by Professor H. V. Howein 1937 yielded 146 excellent specimens (46 per cent of them adults) of which threeare figured here. Hazel's specimens came from Frobisher Bay (63°io' N, 67°45' W)and the Gulf of Maine (44°o8' N, 68°i3'W). Akatova figured a specimen fromstation 80 on the Novosiberian Shelf (j6°$2' N, I33°23' E). The present author hasfound it in British Museum (N.H.) samples collected by the 'Ernest Holt' at Stationsi (70°32' N, i8°23' E) and 6 (69^4' N, i7°oo' E) and by H.M.S. Vidal at Station 46(75°n-2' N, 22°i4' E) on the Spitzbergen Shelf, and in material from the HundeIslands (68°52' N, 53°O7' W) in the Hancock Museum. The distribution of F. (B.)barentzovoensis is essentially Arctic (Fig. 2), all the records (except for the somewhatanomalous specimen from the Gulf of Maine figured by Hazel) being from latitudeshigher than 63° N and the maximum depth at which it has been recorded is 143fathoms (Ernest Holt Station 6), the bulk of the occurrences being between 10-40fathoms. It may thus be regarded as a relatively shallow-water Arctic form and itis possible that Hazel's Gulf of Maine right valve is a sub-Recent specimen. Finmarchinella (Barentsovia) angulata (Sars 1866)(PL i, fig. 8 ; PL 2, figs. 2,4, 6 & 10) Cythereis angulata G. O. Sars, 1866 : 46. Cythere angulata (Sars), Brady, 1868 : 409, pi. 26, figs. 39-42 ; Brady, Crosskey & Robertson, 1874 : 162-163, pi. 4, figs. 17-24, pi. 10, fig. 22 ; Brady & Norman, 1889 (pars) : 165 (non Cythere dathrata, var. nuda, Brady = F. (B.) barentzovoensis). Hemicythere angulata, G. O. Sars, Sars, 1925 : 187-188, pi. LXXXVI, fig. 2. 'Hemicythermae' (genre ?) angulata (Sars), Wagner, 1957 : 61-62, pi. XXXVIII, figs, i, 2 ;Yassini, 1969 : 42-43, pi. XXXIX, fig. 17. Non Hemicythere angulata (G. O. Sars), Akatova, 1946 : 227, fig. 5 (= F. (B.) barentzovoensis). Non Finmarchinella angulata (Sars), Swain, 1963 : 813 (pi. 97, fig. 22 = juv. indet. ; pi. 99, fig.9 & text-fig, nc. = F. (B.) curvicosta). Non Nereina angulata (Sars, 1865), Hazel, 1967 : 19 (pi. i, figs. 7, n = F. (B.) barentzovoensis ;pi. i, figs. 8, 9, 10 = F. (B.) curvicosta). DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Female left valve Oslo Museum F 1447.4 PL l> fig- 8 0-704 0-380 PI. 2, fig. 10 Female right valve Oslo Museum F 1447.6 PL 2, fig. 6 0-668 °'357 Male carapace B.M. No. 1973.331 PL 2, fig. 2 0-701 0-367 o*325 Female carapace B.M. No. 1973.332 PI. 2, fig. 4 0-740 °'4I5 °'377 THE GENUS FINMARCHINELLA SWAIN 89 DISCUSSION. This species has been confused with both F. (B.) barentzovoensis andF. (B.) curvicosta in the past. Sars' illustration in the Crustacea of Norway (1925,pi. LXXXVI, fig. 2) shows a rib which slightly affects the lateral outline postero-dorsally. Sars (1925, p. 188) notes that he had taken this species 'in several placeson the Norwegian coast from Christiana Fjord to Finmark, in the laminaria zone,though nowhere in any abundance'. The type material could not be traced but theZoologisk Museum, Oslo, kindly searched its collections and provided a phial ofspirit material labelled in Sars' own hand 'Hemicythere angulata G.O.S.' Thisyielded 13 specimens of F. (B.) angulata including four juveniles which showed thatSars' figure emphasized the essential features. Dorsally there is a rib which runsfrom approximately the mid-length of the valve towards the posterodorsal cardinalangle where it becomes accentuated, often affecting the outline posterodorsally inlateral view. The rib then turns downward at almost a right angle, or even moreacutely, before disappearing at about mid-height (PI. i, fig. 8a ; PL 2, fig. 10). Theposterior part of the dorsal margin is slightly concave in lateral view. F. (B.) angulata is most easily differentiated from F. (B.) barentzovoensis and F. (B.)curvicosta by the lack of any rib connecting the ventral end of this posterodorsalaccentuation with the subcentral tubercle. Sars' material also shows the presenceof a marginal rib anteriorly (PI. i, fig. 8a) which coincides with the outline at theanterior cardinal angle and ventrally, but lies some distance inside anteriorly whereit follows the general curve of the outline but with the dorsal two-thirds forming amuch flatter arc. This species also lacks the incurved part of the anterior rib atabout mid-height which is characteristic of F. (B.) barentzovoensis. DISTRIBUTION. Sars found this species living in shallow water along the length ofthe Norwegian coast. It occurs at Russian Harbour, Novaya Zemlya, and ErnestHolt Station 6 in the Barents and Norwegian Sea areas and the author has also foundjuveniles in Cockburn Bank samples 5668 (49°5o' N, 9°i8' W) and 5673 (49°5o' N,9°09' W) although these Celtic Sea specimens are possibly not indigenous. Slidesin the British Museum (N.H.) show that in the Greenland area it was found at Hol-steinsborg Harbour (where it occurred with almost equal numbers of F. (B.) curvicosta)and at Station 3 (69°3i' N, 56° 01' W) of H.M.S. Valorous in 1875 ; in the Norwegianarea at Hollingpollen, Drobak and Lungegaards-vandet, Bergen ; and round the Irishand Scottish coasts at Clew Bay, County Mayo, between the Cumbraes in the Firthof Clyde, off Tarbert, Loch Fyne and in the Minch. From the Brady Collection inthe Hancock Museum, Newcastle-upon-Tyne, it has been possible to verify the pre-sence of this species in the Hunde Islands in the Greenland region ; in Stromness Bay,Scapa Flow, Rothesay Bay, Cumbrae, Rosneath, and Bo'ness in the Firth of Forthin the Scottish area and in Westport Bay in Ireland. From the same collection onecan also confirm that it occurs fossil at Loch Gilp, Dalmuir, Colintraive, Cumbraeand in raised beaches at Oban and Tarbert. It also occurs fossil in the Quaternaryof the Netherlands (Wagner, 1957) and Yassini (1969) found it at eight localities inthe Bay of Biscay although he did not regard it as living in that area at the presentday. Most records of this species are from less than 50 fathoms and only rarely hasit been found at depths of more than 100 fathoms. Its known occurrence is shownin Fig. 2. go JOHN W. NEALE Finmarchinella (Barentsovia) curvicosta sp. nov.(PL i, figs. 1-3 ; PI. 2, figs. 3, 7-9, 13) Finmarchinella angulata (Sars) Swain, 1963 : 813-814, pi. 99, fig. 9 & text-fig, nc. non pi. 97,fig. 22 (= juv. indet.) Nereina angulata (Sars) Hazel, 1967 (pars) : 19, pi. i, figs. 8, 9, 10. non pi. i, figs. 7, n ( =F. (B.) barentzovoensis]. HOLOTYPE. A female carapace (mounted as separate valves) from HolsteinsborgHarbour, Greenland, B.M. No. 1973.310. PARATYPES. Nineteen specimens from Holsteinsborg Harbour, Greenland,including males, females and juveniles, B.M. Nos. 1973.311-329. OTHER MATERIAL. Fifty-two specimens in the collections of Hull University andLouisiana State University. DIMENSIONS OF FIGURED MATERIAL. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Female left valve, holotype B.M. No. 1973.310 PI. i, fig. 3 0-808 0-451 PI. 2, fig. 13 Male left valve, paratype B.M. No. 1973.311 PI. i, fig. 2 0-743 0-383 Female right valve, paratype B.M. No. 1973.312 PI. i, fig. i 0-844 °'45° PI. 2, fig. 8Female left valve, paratype B.M. No. 1973.313 PI. 2, fig. 9 0-844 °'454 Female carapace, paratype B.M. No. 1973.314 PI. 2, fig. 3 0-811 0-441 0-431 Female left valve HU. 8o.R.26 PI. 2, fig. 7 0-876 0-506 DIAGNOSIS. A species of Finmarchinella (Barentsovia} with gently convex postero-dorsal margin, well-developed costation and overall reticulation. An almosthorizontal rib runs posteriorly from the subcentral tubercle to about five-sixths of thelength where it joins a strong curved rib at an acute angle. The curved rib passesinto a weaker oblique rib which runs back to the subcentral tubercle. The elongateloop formed by these costae usually contains two rows of polygonal fossae. Dorsaland ventral ribs complete the principal costation. DESCRIPTION. Valves in lateral view rounded-rectangular of typical Finmar-chinella shape. Posterodorsal margin gently convex, posterior margin concave. Indorsal view rounded subhexagonal. Eye tubercles and subcentral tubercle well-developed. Ornamentation of overall reticulation consisting of finely pitted poly-gonal fossae and costation. There are four principal costae, the most prominent ofwhich is an almost horizontal rib which runs posteriorly from the subcentral tubercle.At about five-sixths the length this joins at an acute angle a short, curved, accen-tuated rib which runs towards the middle of the dorsal margin and generally formsthe most prominent feature of the ornamentation. Dorsally, this strong, curved THE GENUS FINMARCHINELLA SWAIN 91 rib runs into a relatively weak oblique rib which runs anteroventrally back to thesubcentral tubercle. This straight section of the curved rib is the most weaklydeveloped of the principal costae. As a whole the costate pattern in this area formsan elongated loop with its apex at the subcentral tubercle. The loop normallycontains two rows of polygonal fossae. A third principal rib starts just below theposterior part of the eye tubercle and runs obliquely in a posterodorsal direction tojoin the dorsal margin at about three-quarters of the length (PI. i, fig. 2a; PI. 2,fig. 3). A fourth, slightly flexuous costa delimits the junction between the lateraland ventral surfaces and is very prominent at its posterior termination. Hingeantimerodont, anterior and posterior toothplates with 5-6 (usually 5) teeth andmedian locellate groove in the right valve. Left valve with complementary struc-tures. Normal pore canals large, simple and scattered. Marginal pore canalssimple and straight ; approximately 36 anteriorly and 10 posteriorly. Anterior andposteroventral vestibules exceedingly narrow. Muscle scar pattern typical of thegenus consisting of a vertical row of four adductor scars with three scars anteriorly.The long axes of the adductor scars lie obliquely to the length of the valve, declininganteriorly. The middle scars are very clearly divided. Sexual dimorphism marked,the males being lower in proportion to their length than the females. AFFINITIES AND DIFFERENCES. This species is most closely related to the twopreceding species. It is most easily differentiated by the posterodorsal margin whichis convex in lateral view compared with the concavity in this region seen in F. (B.)barentzovoensis (PI. i, figs. 4, 5) and F. (B.) angulata (PI. i, fig. 8a). Besides otherdifferences in ornamentation F. (B.) curvicosta lacks the prominent incurving of theanterior rib of F. (B.) barentzovoensis and the curve of the posterodorsal rib is muchflatter and lies well below the valve margin compared with F. (B.} angulata (cf. PL 2,figs. 13 and 10). DISTRIBUTION. F. (B.) curvicosta is essentially Arctic in distribution (Fig. 2).Slides in the British Museum (N.H.) show that it occurs in 10 fathoms at Holsteins-borg Harbour and 5-25 fathoms at Godhavn Harbour, Disco in the Greenland areaand was dredged from 100 fathoms at Station 3 (69°3i' N, 56°oi' W) by H.M.S.Valorous in 1875. The Brady Collection in the Hancock Museum, Newcastle-upon-Tyne, contains a slide of this species from the Hunde Islands (68°52'N, 53°O7' W),also in the Greenland area. It occurs at Kneeland Bay (62°5g' N, 67°28' W) andUngava Bay (6o°o8' N, 67°47' W) where it was recorded by Hazel (ig67a) as Nereinaangulata (Sars) and Swain (1963) figured a female left valve from the PleistoceneGubik Formation of Alaska, also as Finmarchinella angulata (Sars). In the easternhemisphere it was found quite commonly in 8 fathoms at Russian Harbour, NovayaZemlya. Finmarchinella (Barentsovia) japonica (Ishizaki, 1966) Nereina japonica Ishizaki, 1966 : 143-144, pi. 19, figs. 1-4, text-fig, i, figs. 3, 4. DISCUSSION. This species shows the well-developed submedian and eye tuberclesof Barentsovia. The costation is difficult to ascertain from the figure of the holotype 92 JOHN W. NEALE but in his description Ishizaki notes that the dorsal rib extends to the posterior endwhere it turns downward and disappears at mid-height. This, coupled with thegeneral reticulation, is much more reminiscent of F. (B.) angulata (Sars) than of theother species of Barentsovia. It differs from F. angulata in the more prominentventral rib. Ishizaki's species is exclusively fossil and has not so far been found outside theMiocene and Pliocene of Japan. CONCLUSIONS The genus Finmarchinella is essentially a cold water genus which includes non-costate (s.g. Finmarchinella} and costate (s.g. Barentsovia} groups. The confusionbetween species of the latter is easily resolved by reference to the shape and ornamen-tation of the valves. ACKNOWLEDGEMENTS It is a pleasure to thank the authorities of the British Museum (Natural History),London, the Hancock Museum, Newcastle-upon-Tyne, and the Zoologisk Museum,Oslo, who kindly arranged the loan of the material on which this work was based. REFERENCES AKATOVA, N. A. 1946. On the fauna of Ostracoda from the Novosiberian Shelf waters. In: Volume III. Results of the drifting expedition in the main northern seaways in the ice-breaker steamship 'Sedov' 1937-1940. Isdat. Glavsevmorputi, Moscow, Leningrad, 224-230, 8 figs. (In Russian.)BRADY, G. S. 1866. On New or imperfectly known Species of Marine Ostracoda. Trans. zool. Soc. London, 5 : 359-393, pis. 57-62. 1868. A Monograph of the Recent British Ostracoda. Trans. Linnean Soc. London, 26 : 353-495. pis. 23-41.— CROSSKEY, H. W. & ROBERTSON, D. 1874. A Monograph of the post-Tertiary Entomo- straca of Scotland (including species from England and Ireland). Palaeont. Soc. Monogr. London, 1-232, pis. 1-16. & NORMAN, A. M. 1889. A Monograph of the Marine and Freshwater Ostracoda of the North Atlantic and of North-western Europe. Section I. Podocopa. Sci. Trans. Royal Dublin Soc. 4 (Series II) : 63-270, pis. 8-23. & NORMAN, A. M. 1896. A Monograph of the Marine and Freshwater Ostracoda of the North Atlantic and of North-western Europe. Part II. Sections II-IV. Myodocopa, Cladocopa, and Platycopa. Sci. Trans. Royal Dublin Soc. 5 (Series II) : 621-746, pis. 50-68.CRISTOFORI, J. DE & JAN, G. 1832. Conspectus methodicus testaceorum in collectione mea exstantium. Scientiae Naturalis Cultoribus : 8 pp.HAZEL, J. E. ig67a. Classification and Distribution of the Recent Hemicytheridae and Trachyleberididae (Ostracoda) off Northeastern North America. U.S. Geol. Surv. Prof. Paper 564 : 1-49, pis. i-n.— ig67b. Corrections : Classification and Distribution of the Recent Hemicytheridae and Trachyleberididae (Ostracoda) off Northeastern North America. /. Paleont. 41 : 1284- 1285. THE GENUS FINMARCHINELLA SWAIN 93 HAZEL, J. E. 1968. Pleistocene ostracode zoogeography in Atlantic Coast submarine canyons. /. Paleont. 42 : 1264-1271, 3 text-figs. 1970. Atlantic Continental Shelf and Slope of the United States. Ostracode Zoogeography in the Southern Nova Scotian and Northern Virginian Faunal Provinces. U.S. Geol. Surv. Prof. Paper 529-E : i-v, Ei-E2i, 69 maps, n text-figs., 3 tables.ISHIZAKI, K. 1966. Miocene and Pliocene Ostracodes from the Sendai Area, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 37 : 131-163, pis. 16-19, I fig.—— 1969. Ostracodes from Shinjiko and Nakanoumi, Shimane Prefecture, Western Honshu, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 41 : 197-224, pis. 24-26, 16 figs., 2 tables. 1971. Ostracodes from Aomori Bay, Aomori Prefecture, Northeast Honshu, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 43 : 59-97, pis. 1-7, 8 figs., 4 tables.LEV, O. M. 1969. Analysis and Comparison of Ostracod Communities from Late Cenozoic Deposits in the Lower Reaches of the Yenesei, Ob', Pechora, Mezen and N. Dvina Rivers. In: Uchen. Zap. Paleont. Biostrat., nauch-issled. Inst. geol. Arktiki, 28 : 25-34, 5 P^s- (In Russian.)MANDELSTAM, M. I., SCHNEIDER, G. F., KUZNETSOVA, Z. V. & KATZ, F. I. 1957. New Genera in the Families Cypridae and Cytheridae. Ezheg. vses. paleont. obshch. 14 : 166-193, P^3- I-IV. (In Russian.) MOVES, J. & PEYPOUQUET, J. P. 1971. Les Ostracodes indicateurs d'un paleorivage pleisto-cene en bordure du plateau continental du Golfe de Gascoyne. C. R. Somm. Stances Soc. Geol. France. Fasc. 44 : 219-220.OSTENFELD, C. H. & WESENBERG-LuND, C. 1909. Catalogue des especes de plantes et d'ani- maux observees dans le plankton recueilli pendant les expeditions periodiques depuis le mois d'Aout 1905 jusqu'au mois de Mai 1908. Publs Circonst. Cons. perm. int. Explor. Mer. No. 48.PEYPOUQUET, J. P. 1971 a. La distinction des biocenoses, thanatocenoses, paleothanato- cenoses: probleme fondamental sur une plateforme continentale. Bull. Inst. Geol. Aquitaine, 11 : 191-208, i map. i97ib. Inventaire de la microfaune d'Ostracodes de la region de Cap-Breton. Butt. Inst. Geol. Aquitaine, 11 : 209-217, i map, i table.SARS, G. O. 1866. Oversigt af Norges Marine Ostracoder, Norske Vidensk.-Akad. forhandlingar (1865), 1866, pp. 1-130. 1922-28. An Account of the Crustacea of Norway, 9 : 1-277, P^8- I-H9- Bergen Museum. SWAIN, F. M. 1963. Pleistocene Ostracoda from the Gubik Formation, Arctic Coastal Plain, Alaska. /. Paleont. 37 : 798-834, pis. 95-99, 13 figs.Vos, A. P. C. DE. 1957. Liste annot^e des Ostracodes marins des environs de Roscoff. Archs Zool. exp. gen. 95 : 1-74, 29 pis., i table.WAGNER, C. W. 1957. ^ur ^es Ostracodes du Quaternaire recent des Pays-Bas et leur Utilisation dans VEtude geologique des Depots holocenes. Mouton & Co., The Hague, pp. 1-259, 50 pis., 21 figs.YASSINI, I. 1969. Ecologie des Associations d'Ostracodes du Bassin d'Arcachon et du Littoral Atlantique. Application a 1'Interpretation de quelques Populations du Tertiaire Aquitain. Bull. Inst Geol. Aquitaine, I-XXIV, 1-288, 39 pis. Dr JOHN W. NEALEDepartment of GeologyTHE UNIVERSITYHULL HU6 7RX 3t PLATE iFinmarchinella Magnification in all cases X 72 ± 4 FIG. i. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female right valve, B.M. No1973.312. a. Internal lateral view, b. Oblique dorsal internal view. Holsteinsborg Harbour,Greenland. FIG. 2. Finmarchinella (B.) curvicosta sp. nov. Paratype. Male left valve, B.M. No.1973.311. a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour,Greenland. FIG. 3. Finmarchinella (B.) curvicosta sp. nov. Holotype. Female left valve of carapace,B.M. No. 1973.310. a. External lateral view. b. Oblique anterior view. HolsteinsborgHarbour, Greenland. FIG. 4. Finmarchinella (B.) barentzovoensis (Mandelstam). Male left valve, HU. 8o.R.23.External lateral view. Russian Harbour, Novaya Zemlya. FIG. 5. Finmarchinella (B.) barentzovoensis (Mandelstam). Female left valve, HU. 8o.R.24.External lateral view. Russian Harbour, Novaya Zemlya. FIG. 6. Finmarchinella (F.) finmarchica (Sars). Male left valve, B.M. No. M.3287 A. a.External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 7. Finmarchinella (F.) finmarchica (Sars). Female left valve, B.M. No. M.3287 B.a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 8. Finmarchinella (B.) angulata (Sars). Female left valve, Oslo Museum F 1447.4.a. External lateral view. b. Oblique anterior view. Locality unknown. FIG. 9. Finmarchinella (B.) barentzovoensis (Mandelstam). Female left valve, HU. 8o.R.25.Oblique anterior view. Russian Harbour, Novaya Zemlya. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i PLATE 2Finmarchinella FIG. i. Finmarchinella (F.) finmarchica (Sars). Male carapace, B.M. No. M.3287 C. Dorsalview, x 74. Holsteinsborg Harbour, Greenland. FIG. 2. Finmarchinella (B.) angulata (Sars). Male carapace, B.M. No. 1973.331. Dorsalview x 74. Holsteinsborg Harbour, Greenland. FIG. 3. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female carapace, B.M. No.1973.314. Dorsal view X73. Holsteinsborg Harbour, Greenland. FIG. 4. Finmarchinella (B.) angulata (Sars). Female carapace, B.M. No. 1973.332. Dorsalview x 72. Holsteinsborg Harbour, Greenland. FIG. 5. Finmarchinella (F.) finmarchica (Sars). Female carapace, B.M. No. M.3287 D.Dorsal view x 74. Holsteinsborg Harbour, Greenland. FIG. 6. Finmarchinella (B.) angulata (Sars). Female right valve, Oslo Museum No. F 1447.6.Internal view showing muscle scar pattern x 335. Locality unknown. FIG. 7. Finmarchinella (B.) curvicosta sp. nov. Female left valve, HU. 8o.R.26. Externallateral view x 60. Russian Harbour, Novaya Zemlya. FIG. 8. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female right valve, B.M. No. 1973.312. a. Anterior part of hinge x 160. b. Posterior part of hinge x 160. HolsteinsborgHarbour, Greenland. FIG. 9. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female left valve, B.M. No. 1973.313. Oblique ventral view x62. Holsteinsborg Harbour, Greenland. FIG. 10,. Finmarchinella (B.) angulata (Sars). Female left valve, Oslo Museum No. F 1447.4.Lateral view of posterodorsal part of valve x 160. Locality unknown. FIG. ii. Finmarchinella (F.) finmarchica (Sars). Female left valve, B.M. No. 3287 B.Lateral view of posterodorsal part of valve x 144. Holsteinsborg Harbour, Greenland. FIG. 12. Finmarchinella (B.) barentzovoensis (Mandelstam). Female left valve, HU. 8o.R.25.Lateral view of posterodorsal part of valve x 144. Russian Harbour, Novaya Zemlya. FIG. 13. Finmarchinella (B.) curvicosta sp. nov. Holotype. Female left valve (of carapace),B.M. No. 1973.310. Lateral view of posterodorsal part of valve X 135. Holsteinsborg Harbour,Greenland. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE 2 EUPLOTES RARISETA SP. N. (PROTOZOA:CILIATEA) A NEW SMALL MARINE HYPOTRICH By C. R. CURDS, B. JEAN WEST & JEAN E. DORAHY INTRODUCTION TUFFRAU (1960) listed 55 specimens of the genus Euplotes Ehrenberg, 1830 that hadbeen described to that date, and in his extensive revision of the genus eliminatedmany of the species. In doing so, Tuffrau (1960) suggested that there were fourcharacters of fundamental importance which should be used for taxonomic purposesat the species level. These characters were : the number of laterodorsal kinetics,the arrangement of the dorsal silver-line network, the number of frontoventral cirriand the shape of the non-dividing macronucleus. Borror (1968) added the appear-ance of the pellicular or cortical sculpturing that is often a feature of some Euplotesspp. as a further criterion. Using these characters, in addition to some of the moretraditional features, Borror (1972) listed 43 species of Euplotes in his revision of theorder Hypotrichida Stein, 1859. When both traditional and modern criteria are considered, the species of Euplotesdescribed in the present paper does not conform to any of those listed by Borror(1972) nor Carter (1972). MATERIALS AND METHODS (a) Source and cultivation The hypotrich was originally contained in a sample of seawater collected fromAberystwyth, Wales. Some seawater was inoculated into a flask of Erdschreibersolution ('Medium i', Committee on Cultures, Society of Protozoologists, 1958)which was then incubated in the dark at room temperature (about 20 °C) for severalweeks. Under these conditions, Euplotes was the dominant ciliate which grew.Some of the cells so obtained were used by Miss Sheila Andrews to initiate a cultureat the Cambridge Culture Centre of Algae and Protozoa (Euplotes sp. LB i624/2a).Later, a clonal culture was established (Euplotes sp. LB i624/2b) and most of thework described herein relates to that clone. The hypotrich was maintained in the dark at room temperature in conical flasksor test tubes containing Erdschreiber solution. Cultures were fed weekly with a fewdrops of a thick suspension of freeze-dried baker's yeast and subcultured at monthlyintervals. The ciliate also grew well in 150 ml conical flasks containing 100 ml ofseawater and 10-12 rice grains, but the maximum populations reached in thesecultures were lower than those achieved by the former culture method. In order toestablish the possible ecological range of this species, some experimental work wascarried out with media consisting of various dilutions of Erdschreiber solution sup-plemented with baker's yeast and seawater supplemented with rice grains. In these Bull. Br. Mus. nat. Hist. (Zool.) 27, 24 96 C. R. CURDS, B. J. WEST & J. E. DORAHY experiments, duplicate cultures in each medium were subcultured serially into flaskscontaining media of progressively lower salinity at 10 per cent (v/v) intervals. Theinoculum size was always 4 ml per 100 ml of medium and a period of 1-5 weeks atroom temperature was allowed between each transfer. In this way it was possibleto establish the approximate lower salinity limit for this species. (b) Light microscopy Observations and measurements were made using both living and fixed material.Living organisms were slowed by immersion in methyl cellulose (5 per cent w/v inseawater), and osmium tetroxide vapour (2 per cent w/v) was found to be a suitablefixative for cells which were to be drawn or measured. Large numbers of Euploteswere fixed in bulk and their nuclei were stained using Dippell and Chao's modificationof De Lamater's basic fuchsin stain (Sonneborn, 1950). The silver-line system wasdisplayed using the 'wet' method modified by Chatton & Lwoff (1930) and Corliss(1953). (c) Scanning-electron microscopy The preparation of these marine organisms for the scanning-electron microscopeproved to be rather more complex than the method described by Small and Marszalek(1969). It was found necessary to wash the cells 5 or 6 times prior to fixation inorder to separate the cells from bacteria, yeast and debris. The first two washeswere carried out by slow centrifugation (approx. 400 g) for a few seconds. In thiscase the majority of the Euplotes were retained in the supernatant. The cells werethen repeatedly washed in membrane-filtered seawater and centrifuged at approxi-mately 1750 g. The cells in the final pellet were rapidly killed in osmium tetroxidevapour to prevent the cirri curling during fixation. Considerable effort was made tofind a good method of fixation for these marine ciliates which was suitable for scan-ning-electron microscopy. The fixative suggested by Small & Marszalek (1969) wasthat used by Parducz (1967) and although this was acceptable in some respects, ithad the disadvantage of disrupting the structure of the cirri and the adoral zone ofmembranelles (AZM). The fixative finally chosen, although it is still under review,consisted of equal parts of 2 per cent (w/v in distilled water) osmium tetroxide and asaturated aqueous solution of mercuric chloride. Sufficient sodium chloride wasthen added to give a final concentration of 3-3 per cent (w/v) ; this prevented thecells swelling during fixation. The three component parts were mixed immediatelybefore use and then membrane filtered ; cells were fixed in the filtered solution over-night. Following fixation, the cells were washed repeatedly (at least 8 times) inmembrane-filtered triple glass-distilled water and centrifuged at approximately400 g between each wash. The washed cells were then frozen by being dropped ontopieces of aluminium floating on liquid nitrogen and were dried in a tissue dryeressentially as described by Small & Marszalek (1969). The pieces of aluminiumcovered with organisms were glued to stubs and coated with approximately 100 A ofgold as described by Harris, Martin & Ogden (1972), specimens were examined witha Cambridge 'Stereoscan' Mk II scanning-electron microscope (Cambridge Instru-ments Ltd, Cambridge, England). EUPLOTES RARISETA SP. N. 97 Euplotes rariseta sp. n. Diagnosis Small (30-45 /xm long, 20-31 /mi wide), ovoid, marine hypotrich with 10 fronto-ventral, 5 transversal and 3 caudal cirri; caudal cirrus below AZM stout. Ventralsurface heavily sculptured with 6 posteriorly projecting ridges. Dorsal surface with6 double-edged longitudinal ridges. Dorsal bristles sparse ; 6 kinetics with a maxi-mum of 6 bristles in the central kinetics. Dorsal silver-line system double of thepatella type. Macronucleus S-shaped. Micronucleus small, situated anteriorly. Type slides showing silver-line systems, nuclear apparatus and whole mounts havebeen deposited in the slide collection of the Protozoa Section, BM (NH) : holotypeReg. No. 1972:11:1:11 ; paratype Reg. Nos. 1972:11:1:12-18. Body size and shape The size distribution data of Euplotes cells at the stationary and logarithmicphases of growth are given in Fig. i. It should be noted that Euplotes cells from thelogarithmic phase are considerably larger (42 ± 3 /mi long by 28 ± 3 /mi wide) thanare cells from the stationary phase (34-6 ± 3 /mi long by 24-2 ± 2-3 /mi wide). Thisis in agreement with the observations of Curds & Cockburn (1971) who noted thatthe size of the ciliate Tetrahymena pyriformis was related to its rate of growth. Thedata on size show that this species of Euplotes is amongst the smallest recorded forthe genus. The dorsal surface is conspicuously sculptured with 6 double-edged longitudinalridges (PL i, figs, a and c). To the left of each ridge is a parallel row of pits fromwhich the short (2 /u.m long) dorsal cilia or bristles emerge. The ventral surface isalso strongly sculptured (Fig. 2 ; PI. i, figs, a and c), particularly in the posteriorregion between the transverse cirri. There are 6 ventral ridges, 3 of which areprominent and consist of an oblique midventral ridge, and 2 ridges which runlongitudinally flanking the 2 outermost transverse cirri. The other 3 ridges are lessprominent and are more or less restricted to the area immediately between thetransverse cirri. All 6 ventral ridges project slightly posteriorly between the trans-verse cirri and form fin-like structures (Fig. 2 ; Plate i, figs, b and c). The rightbuccal overture has a slight anterior evagination as in Euplotes alatus Kahl, 1932(see Borror, 1968) but in the species under description it extends only just past theAZM. Ciliary organdies This Euplotes has 10 frontoventral cirri which are arranged as shown in PI. i,fig. b and in Figs. 2 and 3b. The arrangement closely resembles that of bothEuplotes charon Ehrenberg, 1830 and Euplotes quinquecarinatus Gelei, 1950 (seeBorror, 1968). There is the normal complement of 5 transverse cirri which arisebetween the ventral ridges and these, particularly cirri III i and IV i (following thenumeration of Wallengren, 1900), are frequently seen to be frayed at the distal end(Fig. 2). The tendency for the transverse cirri to disrupt in this way is greatlyincreased during fixation. There are only 3 caudal cirri, and in this species the one 98 C. R. CURDS, B. J. WEST & J. E. DORAHY 99- £95- >80- E ° 5- 2'0 2'4 28 32 36 SIZE (microns) 40 44 48 FIG. i. Size distribution data of Euplotes rariseta. Curve A. Breadth of cells at stationaryphase. Curve B. Breadth of cells during logarithmic growth. Curve C. Length of cellsat stationary phase. Curve D. Length of cells during logarithmic growth. Stationary-phase cells were fixed in osmic acid vapour. Logarithmic-phase cells were measuredalive immersed in methyl cellulose solution. 10/jm FIG. 2. Ventral aspect of Euplotes rariseta showing cirri and ventral ridges (camera lucida drawing). EUPLOTES RARISETA SP. N. 99 a b FIG. 3. Silver-line system of Euplotes rariseta, (a) dorsal surface, (b) ventral surface. below and nearest to the AZM is characteristically held stiffly out to the left side andis much stouter than the other two caudal cirri which arise beneath the centraltransverse cirri (Fig. 2). The presence of this large stiff caudal cirrus appears to bea feature unique to this species ; in the living cell, the cirrus seems to act as a rudder-like structure. The dorsal bristles are sparse in this species ; there are 6 rows andthe mid-dorsal rows have only 6 bristles, in which respect this species resemblesEuplotes bisulcatus Kahl, 1932 (see Borror, 1968). Nuclei The macronucleus of this species is an irregular S-shape (Fig. 4). In this respectit appears to be unique in the genus Euplotes. The posterior part of the macro-nucleus points anteriorly and ventrally towards the position of the cytostome. Themicronucleus is situated close to the macronucleus on the left anterior edge of thebody (Fig. 4). Silver-line system The geometry of the dorsal silver-line system is of the double or patella-type andclosely resembles that of Euplotes raikovi Agamaliev, 1966 with a series of alternatelongitudinal rows of narrow long polygons and wide short polygons (Fig. 3a). Theventral argyrome (Fig. 3b) consists of an irregular network of polygons in similarnumbers to that of E. cristatus Kahl, 1932 (see Tuffrau, 1960). The dorsal silver-linenetwork has been seen on some scanning-electron micrographs (PL i, fig. d). C. R. CURDS, B. J. WEST & J. E. DORAHY FIG. 4. Ventral view of the nuclei of Euplotes rariseta from a stained preparation.(M = macronucleus, m = micronucleus.) Ecological data Cells inoculated into Erdschreiber solution supplemented with yeast grew well innine different dilutions of this medium from 100 down to 20 per cent (v/v). Cellstransferred from 20 to 10 per cent Erdschreiber solution showed no growth duringthe 6 weeks following inoculation. Growth in seawater supplemented with rice grainswas good in seven dilutions from 100 down to 40 per cent (v/v), but cells transferredfrom 40 to 30 per cent seawater showed good growth only after 3 weeks in culture.Subsequent transfer of cells from 30 to 20 per cent seawater led to poor growth after5 weeks, and no growth was observed in 10 per cent seawater. Contractile vacuoleactivity was not observed in organisms from any of the tested dilutions of seawateror Erdschreiber solution. This is in agreement with the findings of Yocum (1934),who found that the activity of the contractile vacuole in Euplotes patella Ehrenberg,1831 diminished and became imperceptible at concentrations of seawater above10 per cent. Yocum (1934) also found that E. patella would survive well and dividein 66 per cent seawater. In view of the finding that the species of Eiiplotes describedin the present paper will survive and divide in dilutions of seawater or Erdschreibersolution between 20 and 100 per cent, it seems likely that, in nature, this organismcould be found in both marine and brackish environments. However, for thelower dilutions tested, organisms were not as abundant in seawater medium as theywere in Erdschreiber medium, and it is possible that in these experiments the amountsof substances other than sodium chloride were limiting growth. DISCUSSION AND CONCLUSIONS In the past, some authors have noted that the number of caudal cirri is not asconstant as the others, for example, Euplotes mutabilis Tuffrau, 1960 has 4-5caudals, E. raikovi has 2-3 caudals and there are other similar examples. In viewof these observations perhaps the use of the number of caudal cirri as a charactershould be regarded with caution. Although E. octocirratus Agamaliev, 1967 is theonly other species of Euplotes which has precisely the same numbers of cirri on theventral surface, that is 10 frontoventrals, 5 transversals and 3 caudals, one should EUPLOTES RARISETA SP. N. 101 consider those species with a 10 : 5 : 4 cirri complement. These include the follow-ing : E. alatus, E. balteatus Kahl, 1932, E. crenosus Tuffrau, 1960, E. cristatus, E.harpa Stein, 1859, E. indentatus Carter, 1972, E. inkystans Chatton in Tuffrau, 1960,E. minuta Yocum, 1930, E. magnicirratus Carter, 1972, E. quinquecarinatus, E.roscoffensis Dragesco, 1966, E. trisulcatus Kahl, 1932, E. tuffraui Berger, 1965 andE. vannus Minkjewicz, 1901. In all these species the shapes of the macronuclei arequite unlike that of the species described in this paper and furthermore the silver-line systems of these species are different. It is unfortunate that the dorsal and ventral silver-line systems have not beendescribed for all the known species of Euplotes ; however, of those listed above nonehave 6 dorsal kinetics as found in E. rariseta ; only E. octocirratus and E. trisulcatushave 7 kinetics, the remainder have 8 or more. There are four other species with6 dorsal kinetics and these are E. raikovi, E. strelkovi Agamaliev, 1967, E. tegulatusTuffrau, 1960 and E. balticus Kahl, 1932. None of these four species have the cor-rect numbers of cirri and furthermore all possess far too many dorsal kinetosomes. E. bisulcatus is the only other species with a maximum of 6 dorsal kinetosomes perkinety but there are 8 dorsal kinetics in that species. Euplotes rariseta differs from all previously described species of Euplotes in pos-sessing an S-shaped macronucleus and a very stout caudal cirrus. In addition,although the remaining characters have been observed amongst other species ofEuplotes, it is evident from the information given above that none of these specieshas the same combination of characters. For these reasons, it is considered that thespecies of Euplotes described in this paper is sufficiently distinct from all others tobe designated as a separate species, and because of the paucity of dorsal bristleshas been named Euplotes rariseta. Following the revised classification of the Com-mittee on Taxonomic Problems of the Society of Protozoology (Honigberg et al.1964) Euplotes rariseta is placed into class Ciliatea Perty, 1852, order HypotrichidaStein, 1859, family Euplotidae Ehrenberg, 1838. ACKNOWLEDGEMENTS The authors wish to thank Mr R. Harris, Department of Zoology, British Museum(Natural History), for his considerable aid and advice on the treatment of specimensfor scanning-electron microscopy. We are indebted to Dr R. H. Hedley and MrC. G. Ogden, British Museum (Natural History), and to Mr E. A. George and Dr F. C. Page, Culture Centre for Algae and Protozoa, who read the original manu-script and made several constructive suggestions. Part of this work was carriedout while B. J. W. was employed by the Natural Environment Research Council,Culture Centre for Algae and Protozoa, Cambridge. REFERENCES BORROR, A. C. 1968. Systematics of Euplotes (Ciliophora, Hypotrichida) ; towards union of the old and the new. /. Protozool. 15 (4) : 802-808. — 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Protozool. 19 (i) :1-23. 102 C. R. CURDS, B. J. WEST & J. E. DORAHY CARTER, H. P. 1972. Infraciliature of eleven species of the genus Euplotes. Trans. Amer. Microsc. Soc. 91 (4) : 466-492.CHATTON, E. & LWOFF, A. 1930. Impregnation, par diffusion argentique, de 1'infraciliature des cilies marins et d'eau douce, apres fixation cytologique et sans dessication. C. r. Seanc. Soc. Biol. 104 : 834-836.COMMITTEE ON CULTURES, SOCIETY OF PROTOZOOLOGISTS. 1958. A Catalogue of Laboratory Strains of Free-living and Parasitic Protozoa. /. Protozool. 5 (i) : 1-38.CORLISS, J. O. 1953. Silver impregnation of ciliated protozoa by the Chatton-Lwoff technic. Stain Tech. 28 : 97-100.CURDS, C. R. & COCKBURN, A. 1971. Continuous monoxenic culture of Tetrahymena pyriformis. J. gen. Microbiol. 66 : 95-108.HARRIS, R. H., MARTIN, B. S. & OGDEN, C. G. 1972. Notes on the preparation of natural history specimens for scanning electron microscopy. Bull. Br. Mus. nat. Hist. (Zool). 24 (3) : 223-228.HONIGBERG, B. M., BALAMUTH, W., BOVEE, E. C., CORLISS, J. O., GOJDICS, M., HALL, R. P., KUDO, R. R., LEVINE, N. D., LOEBLICH, A. R., WEISER, J. & WENRICH, D. H. 1964. A revised classification of the Phylum Protozoa. /. Protozool. 11 pp. 7-20.PARDUCZ, B. 1967. Ciliary movement and co-ordination in ciliates. Int. Rev. Cytol. 21 : 91- 128.SMALL, E. B. & MARSZALEK, D. S. 1969. Scanning electron microscopy of fixed, frozen and dried protozoa. Science, N.Y. 163 : 1064-1065.SONNEBORN, T. M. 1950. Methods in general biology and genetics of Paramecium aurelia. J. exp. Zool. 113 : 84-147.TUFFRAU, M. 1960. Revision du genre Euplotes, fondle sur la comparaison des structures superficielles. Hydrobiologia, 15 : 1-77.WALLENGREN, H. 1900. Zur Kenntnis der vergleichenden Morphologic der Hypotrichen. Bih. K. svenska Vetensk A had. Handl. 26 : 1-31.YOCUM, H. B. 1934. Observations on the experimental adaptation of certain fresh-water ciliates to seawater. Biol. Bull. mar. biol. Lab. Woods Hole, 67 : 273-276. Dr COLIN R. CURDS Miss B. JEAN WEST, B.Sc.* Mrs JEAN E. DORAHY, M.Sc. CULTURE CENTRE OF ALGAE AND PROTOZOA Department of Zoology STOREY'S WAY BRITISH MUSEUM (NATURAL HISTORY) CAMBRIDGE CROMWELL ROAD LONDON SW7 5BD * Present address. Microbiology Department, School of Biological Sciences, University of Bath,England. 4t PLATE i Scanning micrographs of Euplotes rariseta a. Dorsal view showing ridges and rows of dorsal bristles ( x 2-ik) b. Ventral view showing the arrangement of cirri and ventral ridges (x i-8k) c. Posterolateral aspect showing double-edged dorsal ridges ( x 2-yk) d. Dorsal surface showing argyrome (x 6-ik) Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i OBSERVATIONS ON CLONAL CULTURES OF EUGLYPHA ACANTHOPHORA AND EUGLYPHA STRIGOSA (TESTACEA: PROTOZOA) By R. H. HEDLEY, C. G. OGDEN & JUNE I. KRAFFT INTRODUCTION MANY testate amoebae are known only from their original descriptions, based onobservations made at the magnification limits of the optical microscope. Theintroduction of scanning-electron microscopy for surface ultrastructure examinationnow allows more accurate descriptions to be made of these animals. The presentaccount describes the shell morphology and biology of two species of EuglyphaDujardin, 1841, namely Euglypha acanthophora (Ehrenberg, 1841) and Euglyphastrigosa (Ehrenberg, 1871), both of which have been recently isolated from samplesof sewage sludge and moss respectively, collected in southern England. This is thethird paper in a series dealing with testate amoebae. The previous papers (Hedleyand Ogden, 1973, 1974) are devoted to the biology and fine structure of Euglypharotunda and Trinema lineare respectively. Fine structure observations on thecytoplasm of E. acanthophora and E. strigosa will be the subject of a separatepublication. The genus Euglypha belongs to the family Euglyphidae - the classification adoptedhere is that proposed by Loeblich and Tappan (1961) : Class RHIZOPODEA Von Siebold, 1845 Subclass FILOSIA Leidy, 1879 Order GROMIDA Claparede and Lachmann, 1859 Superfamily EUGLYPHACEA Loeblich and Tappan, 1961 Family EUGLYPHIDAE Wallich, 1864 test hyaline, symmetrical, elongate, composed ofrounded siliceous scales, aperture rounded orelongate ; one nucleus. Previous work - biology Euglypha acanthophora. Most of the early biological observations of Euglyphawere made on this species. These reports (Leidy, 1877 ; Gruber, 1881 ; Blochmann,1887 ; Schewiakoff, 1888 ; Carter, 1889 ; Penard, 1890 ; Reukauf, 1912 ; Popoff,1912) dealt mainly with nuclear division and reproduction by binary fission. Carter(1865, 1889), Reukauf (1912) and Popoff (1912) also described another process ofdivision in which many individuals were produced at one time, but all the stages ofthis division were not seen. A report by Jones (1929), made from observations on an Bull. BY. Mus. nat. Hist. (Zool.) 27, 25* 104 R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT old protozoan culture, described a flagellate stage for E. acanthophora. This has notbeen confirmed and it is possible that the cultures were contaminated. Encystation was observed by Penard (1890, 1902), Leidy (1879), Popoff (1912)and Reukauf (1912). Leidy (1879) described the cyst as being enclosed in twodistinct envelopes, and Popoff (1912) illustrated the various stages in the process ofencystation. When two individuals were observed joining to form a single largespecimen and immediately formed a cyst, Reukauf (1912) suggested that this was acopulation cyst which at excystation produced many new individuals. In studies on the distribution of testaceans in relation to their habitats, Heal(1961) found E. acanthophora in small numbers associated with fen areas, whereasde Graaf (1956) considered this species to have no particular associations. Euglypha strigosa. This species has been recorded from both wet and dryhabitats, de Graaf (1956) noted that in a Netherland fen it preferred drier Sphagnum,whilst Heal (1961) described it as a eurytopic species occurring in large numbers inbog hummocks in Northern England and also noted (Heal, 1962) that it appearedto be restricted to the lower levels of the Sphagnum. Differences in shell size andshape of specimens of E. strigosa were described by Chardez and Leclercq (1963), whosuggested that these differences could be related to the type of habitats in whichthey occur. Previous work - taxonomy In a review of the British Freshwater Rhizopoda fauna Cash et al. (1915) summarizeprevious taxonomic reports of both Euglypha acanthophora and E. strigosa. Thepresent account excludes these references and is restricted to subsequent descrip-tions. Euglypha acanthophora (Ehrenberg, 1841). This species was originally describedas E. alveolata but was made a synonym of E. acanthophora by Cash et al. (1915) whogave a detailed account of their reasons, and this has since been accepted by mostauthors. They (Cash et al. 1915) redescribed E. acanthophora and also two varietieswhich differed mainly in size, and the shape and number of the elongated shell-plates. Further redescriptions and sizes were given by Hoogenraad and de Groot(1940) and Decloitre (1962, 1965), the latter author also described (Decloitre, 1962)seven varieties of E. acanthophora. Euglypha strigosa (Ehrenberg, 1872). Cash et al. (1915) described this species as'the most generally distributed and numerous of the Euglyphae', easily distinguishedfrom E. ciliata and E. compressa by its thickened apertural-plates and circular aper-ture. In a review of the genus Euglypha, Decloitre (1962) redescribed E. strigosaand three varieties. Additional sizes and descriptions were given by Bartos (1963)and Decloitre (1964). The surface ultrastructure of the shell of E. strigosa wasdescribed by Cambar et al. (1964), from metal and carbon-shadowed preparations. MATERIALS AND METHODS Euglypha acanthophora was isolated from a sample of sewage sludge from MapleLodge Works of the Colne Valley Sewage Board, Hertfordshire, in December, 1972. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 105 Euglypha strigosa was isolated from a sample of sphagnum moss collected in theNew Forest, near Christchurch, Hampshire, in April, 1970. Both species wereobtained from crude cultures made from small portions of each type of materialplaced in a shallow layer of culture liquid and kept in the laboratory at roomtemperature, 18-20 °C. Agnotobiotic cultures were kept in small plastic containers,on a thin layer of agar agar (i per cent in distilled water) with a sterilized wheat grainadded prior to setting and covered with a shallow layer of culture liquid. The cultureliquid was a 5 per cent (w/v) solution of soil extract, plus 100 mg/1"1 sodium nitrateand 15 mg/l"1 sodium dihydrogen orthophosphate, in distilled water. E. strigosahas more recently been kept on 1-5 per cent agar in amoeba saline, with a sterilizedrice grain added and covered with a shallow layer of Prescott and James solution(Prescott & James, 1955). Clonal cultures were established by isolating single active animals. One clonehas been used subsequently to produce the working cultures of each species. Theseclones have been deposited, E. strigosa (No. 1520/2), E. acanthophora (No. 1520/3),and maintained at the Culture Centre of Algae and Protozoa, The Natural Environ-ment Research Council, Cambridge, England. Subcultures made at 4-5 weekintervals are adequate to maintain active animals that readily feed and reproduce. Optical microscopy. Specimens were examined by both phase-contrast and bright-field illumination, either live or after fixation with Schaudinn's fluid. Smears offixed animals were stained with either borax carmine or iron haematoxylin. Scanning-electron microscopy. Live specimens were either cleaned by transferringthem from the culture vessels through several changes of triple-distilled water, orfixed in glutaraldehyde and then thoroughly washed. They were then manipulatedusing a single-hair brush onto a small cover-slip, previously cleaned with acetone andlint-free tissue, to which they adhere when dry. To obtain single siliceous shell-plates, an individual clean specimen was placed on a small segment of cover-slip andcovered with a small drop of concentrated sulphuric acid. This was heated gently toevaporate the acid and liberate the plates from the organic cement. These preparedcover-slips were secured to 'Stereoscan' stubs with an electrically conductive paint,'Silver Dag', prior to being coated evenly with 10-15 nm °f g°ld using the methodrecommended by Harris et al. (1972). The stubs were examined on either the Cam-bridge Stereoscan Mk II or Stereoscan 600, operating at 15 or 20 kV and the resultswere recorded on Ilford HP4 film. DISTRIBUTION E. acanthophora is usually found in damp mosses, submerged vegetation andstanding water, whilst E. strigosa is found in similar habitats and additionally in driermosses and soil. A full list of locality records and references was given by Decloitre (1962). Sincethis review several authors, namely, Bartos (1963), Bonnet (1966), Decloitre (1964,1965) and Stepanek (1963), have recorded these species from other localities andthese are included here. io6 R. H. MEDLEY, C. G. OGDEN & J. I. KRAFFT The following lists illustrate the geographically widespread distribution of bothspecies : Euglypha acanthophora EUROPE NORTH AMERICASOUTH AMERICA :AFRICA : ASIA :AUSTRALASIA : England, Wales, Scotland, Ireland, France, Holland, Belgium, Germany, Spain, Switzerland, Hungary, Czechoslovakia, Luxembourg, Iceland and Lapland. Greenland, Canada and the United States of America. Venezuela and Colombia. Congo, Cameroons, French West Africa, South Africa and Madagascar. China, Japan, Java and Sumatra. Australia and New Zealand. Euglypha strigosaEUROPE: NORTH AMERICASOUTH AMERICA ;AFRICA : ASIA : AUSTRALASIA :ANTARCTICA : England, Wales, Scotland, Ireland, France, Belgium, Holland, Spain, Germany, Switzerland, Italy, Hungary, Czechoslovakia, Bulgaria, Rumania, Spitzbergen, Finland and Iceland. Greenland, Canada and the United States of America. Central America, Colombia, Venezuela and Chile. South Africa, French West Africa, Madagascar and the Canary Islands. Krakatoa Island, Borneo and Java. Australia, Tasmania and New Zealand. Adelaide Island. MORPHOLOGY AND VARIATION Euglypha acanthophora. The shell has a mean length of 70-8 /u,m ± 9-6 /mi anda mean breadth of 39-0 /zm ±4-4 ju,m ; it is elliptical in shape through the majoraxis and circular through the minor axis (PL i, fig. A), and is made up of threedifferent types of siliceous plates- apertural-plates (PL i, fig. E), shell-plates (PL i,fig. B) and elongated shell-plates (PL i, fig. B). The aperture is terminal, circularand has a mean diameter of 18-4 /u,m ± 4-9 /u,m, it is surrounded by between 10 and13 evenly spaced apertural-plates (PL i, fig. D). Each apertural-plate is roughlycircular in shape and varies in size from 8-1 /u,m to 12-5 /u,m in length and 7-2 /u,m to11-4 ju,m in width. The dentate edge of the apertural-plate is slightly thickened andcarries a median tooth with either four or five smaller lateral teeth on each side(PL i, figs. C and E). The apertural-plates are found not only bordering the aper-ture, but are also seen in the second and third row inside the aperture (PL i, fig. E).There are approximately 200 shell-plates which range in size from 10-9 /urn to 12 ju.niin length and 7-9 /u.m to 9-1 /mi in width. They are roughly circular in shape with theanterior edge usually having three small pointed projections (Fig. la), and arearranged regularly in alternate longitudinal rows (PL i. fig. A). In the posterior CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 107 a b c FIG. i. Diagram to show the comparative shape of shell-plates (a) E. acanthophorafrom anterior region, (b) E. acanthophora from posterior region, (c) E. strigosa. region of the shell there are normally between one to six elongated shell-plates whichhave one edge tapering to a fine point (PL i, fig. B). These plates are approximatelytwice as long as a normal shell-plate, nevertheless, the projections break easily sothat they are often seen to be short and end bluntly. The pointed edge of theseelongated shell-plates is usually directed posteriorly so that they are seen to projectfrom the outline of the shell, but on occasions they follow the curvature of the shelland are difficult to see by light microscopy. Abnormally shaped shells occur occasionally, although the incidence of such formsappears to be less than three per cent. They are usually specimens with eitherdeformed anterior regions or small specimens, about half or two-thirds normal length,with wide apertures. The first type is the most frequent abnormal shell form, havinga normal arrangement of shell-plates in the posterior region but anterior to the middleregion the shell-plates are compacted together haphazardly, so that the shell onlyattains half the normal length. These deformed shells usually collapse when air-dried whilst normal shells retain their shape. The animals inside these deformedshells appear to possess the normal cytoplasmic organelles. Euglypha strigosa. The shell has a mean length of 81-3 /mi ± 3-8 /mi and a meanbreadth of 42-2 /mi ±6 /mi; it is elliptical in shape through both the major andminor axes (PI. 2, fig. A). Two distinct types of siliceous plates make up the shell,apertural-plates and shell-plates, and in addition there are numerous elongatedsiliceous spines (PI. 2, fig. A). The aperture is terminal and circular, with a meandiameter of 14-9 /mi + 0-8 /mi; it is surrounded by between 10 and 13 evenly spaced,apertural-plates - the usual number of apertural-plates is n (PI. 2, fig. E). Eachapertural-plate is roughly circular in shape and varies in size from 9-2 /mi to 10-8 /miin length and 7-8 /mi to 8-5 /mi in width. One margin is distinctly thicked and carriesa large median tooth with either three or four smaller teeth on each side (PI. 2,fig. C). The shell-plates are roughly oval in shape (Fig. ic), with a size range of from8-0 /mi to 11-3 /mi in length and 4-1 /mi to 6-4 /mi in width, and they are arranged inregularly spaced, alternate longitudinal rows. The thin, elongated spines vary inshape and size from small pear-shaped spines approximately 2 /mi long to thin io8 R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT spines 23 p,m long, and they project from any part of the shell surface (PL I, fig. A).They are found at the junction of shell-plates, where they are held in position by asimilar organic cement material to that which holds the shell-plates together inE. rotunda (Hedley and Ogden, 1973). This has been confirmed from the presentauthors unpublished observations of the fine structure of E. strigosa. The spinesare single structures, although two or more are often seen projecting from the samejunction as though they were connected. They usually project from the surface atright-angles, but they may lie close to the surface and point in any direction. Thereappears to be no regular arrangement in the distribution of the spines, althoughusually only the smaller spines are seen at the junction of the apertural-plate andshell-plate region, and the largest spines seem to lie along the lateral margins. Thespines appear to be removed easily by abrasive action (PI. 2, fig. B).No abnormal forms of this species have been seen in our cultures. REPRODUCTION Euglypha acanthophora. Growth curves produced from observations made onthree replicate cultures gave an estimated doubling time of between 2-3 and 2-8 days.Cytoplasmic division proceeds in a similar sequence to that previously described forE. rotunda by Hedley and Ogden (1973). The approximate time taken to produce thedaughter shell is 25 minutes, whereas the time taken for the completion of binaryfission is two hours. The animals are usually active after division with strong pseudo-podial movement (Fig. 2). Eight specimens with abnormal apertures were isolated and after six days had dividedto produce 27 normal individuals. Observations on individual, dividing abnormalforms, such as those with a wide aperture and others being half normal size, show thatthey often produced smaller individuals with apparently the normal shell shape.One abnormal animal at division produced an almost normal shell, a small shell andsome loose shell-plates (Fig. 3). Cytoplasm was present in the larger daughtershell, and the group did not separate after the normal time taken for division, butmoved after this time as a group. During division in a normal individual, at thetime when all the reserve shell-plates had passed into the pseudopodial trunk, asmall circle of shell-plates became detached from the posterior end of the daughtershell. Binary fission proceeded normally and the two individuals separated, leavingbehind the circle of shell-plates. Euglypha strigosa. An estimated doubling time of between 3-9 and 4-3 days wasobtained from growth curves produced from observations on three replicate cultures.Division is preceded by the retraction of the filose pseudopodia, followed by theextrusion of a large pseudopodial trunk and shortly after by the apertural-plates(PI. 2, fig. D). The shell-plates are then arranged progressively until the daughter-shell is almost complete, at which point the spines are pushed out between the shell-plates with considerable movement. This continues for a short time after the shellsare equal in size, but then the movement of both shell-plates and spines ceasesabruptly. The time taken to produce a daughter-shell is approximately 50 minutes,with complete binary fission taking about three hours. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 109 / 10 FIG. 2. Drawings of a moving specimen of E. acanthophora observed over a period of30 minutes, to show the diversity of pseudopodial movement; the arrow denotes thedirection of movement. It was noted that in preparations of dividing specimens for scanning electronmicroscopy, the daughter-shell consistently collapsed. This is possibly due to anincomplete stabilization of the organic cement lining. no R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT FIG. 3. Diagram representing the division products of an abnormal form ; A - abnormalform, B - apparently normal daughter and C - a small form without cytoplasm. ACKNOWLEDGEMENT We would like to acknowledge the technical assistance of Miss E. M. Sides, whowas responsible for collecting and isolating E. strigosa. REFERENCES EARTHS, E. 1963. Rhizopoden einiger Moosproben aus Java. Ada Universitatis Carolin. : 119-190.BLOCHMANN, F. 1887. Zur Kenntnis der Fortpflanzung von Euglypha alveolata Duj. Morph. Jb. 13:173-183.BONNET, L. 1966. Le peuplement thecamoebien de quelques sols du Chili (I). Protistologica 2 : 113-139.CAMBAR, R., LE BLANC, M., MERCIER, M. & THOMAS, R. 1964. Etude en microscopie electroni- que des microstructures superficielles de Protistes fossiles ou vivants. C. r. Acad. Sci. (Paris), 258 : 3554-3555-CARTER, H. J. 1865. On the fresh- and salt-water Rhizopoda of England and India. Ann. Mag. nat. Hist. (3) 15 : 277-293. 1889. Ramulina parasitica, a new species of fossil foraminifera infesting Orbitolites mantelli, var. theobaldi, with comparative observations on the process of reproduction in the Myce- tozoa, freshwater rhizopoda and foraminifera. Ann. Mag. nat. Hist. (6) 41 : 94-101.CASH, J., WAILES, G. H. & HOPKINSON, J. 1915. The British Freshwater Rhizopoda and Heliozoa. Vol. 3, Rhizopoda, part 3. The Ray Society London. 156 pp., 24 pis.CHARDEZ, D. & LECLERCQ, J. 1963. Variabilite des populations d'Euglypha strigosa (Ehren- berg) Leidy en fonction de 1'habitat (Rhizopoda testacea). Bull. Inst. agron. Stns Rech. Gembloux, 31 : 21-27. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) in DECLOITRE, L. 1962. Le genre Euglypha Dujardin. Arch. Protistenk. 106 : 51-100. 1964. Thecamoebiens de la Xlleme Expedition Antarctique Fran9aise. Territoire des Terres Australes et Antarctique Francaises. Expeditions Polaires Fran9aises (Missions Paul-Emile Victor). Publ. No. 259 : 1-47. 1965. Amoebida testacea (Rhizopoda). Zoology Iceland, 2 (i) : 1-58. GRAAF, FR. de 1956. Studies on Rotatoria and Rhizopoda from the Netherlands. I. Rotatoria and Rhizopoda from the 'Grote Huisven'. Biol. Jaarb. 23 : 145-217.GRUBER, A. 1881. Der Theilungsvorgang bei Euglypha alveolata. Z. wiss. Zool. 35 : 431-439.HARRIS, R. H., MARTIN, B. S. & OGDEN, C. G. 1972. Notes on the preparation of natural history specimens for scanning electron microscopy. Bull. Br. Mus. nat. Hist. (Zool.) 24 : 223-228.HEAL, O. W. 1961. The distribution of testate amoebae (Rhizopoda : Testacea) in some fens and bogs in Northern England. /. Linn. Soc. (Zool.) 44 : 369-382. 1962. The abundance and micro-distribution of testate amoebae (Rhizopoda : Testacea) in Sphagnum. Oikos, 13 : 35-47.HEDLEY, R. H. & OGDEN, C. G. 1973. Biology and fine structure of Euglypha rotunda (Testacea : Protozoa). Bull. Br. Mus. nat. Hist. (Zool.) 25 : 119-137. 1974. Observations on Trinema Hneare Penard (Testacea : Protozoa). Bull. Br. Mus. nat. Hist. (Zool.) 26 : 185-199.HOOGENRAAD, H. R. & DE GROOT, A. A. 1940. Fauna van Nederland. Vol. 9, Zoetwater- rhizopoden en Heliozoen. Leiden. 302 pp.JONES, P. M. 1929. Indications of a flagellate stage in Euglypha alveolata. Am. Nat. 63 : 565- 570. LEIDY, J. 1877. The birth of a Rhizopod. Proc. A cad. nat. Sci. Philad. 29 : 261-265. 1879. Freshwater Rhizopods of North America. In: Vol. 12, United States Geological Survey of the Territories. Washington. 324 pp.LOEBLICH, A. R. & TAPPAN, H. 1961. Suprageneric classification of the Rhizopodea. /. Paleontology, 35 : 245-330.PENARD, E. 1890. Etudes sur les Rhizopodes d'eau douce. Mem. Soc. Phys. Hist. nat. Geneve, 31 : 1-230. 1902. Faune Rhizopodique du Bassin du Leman. Geneva. 700 pp. POPOFF, M. 1912. Uber die geschlechtliche Fortpflanzung von Euglypha alveolata Duj. Arch. Protistenk. 25 : 8-26.PRESCOTT, D. M. & JAMES, T. W. 1955. Culturing of Amoeba proteus on Tetrahymena. Exp. Cell. Res. 8 : 256-258. REUKAUF, A. 1912. Zur Encystierung von Euglypha alveolata. Zool. Anz. 39 : 372-375.SCHEWIAKOFF, W. 1888. Uber die karyokinetische Kerntheilung der Euglypha alveolata. Morph. Jb. 13 : 193-258.SxfipANEK, M. 1963. Die Rhizopoden aus Katanga (Kongo-Afrika). Annls Mus. r. Afr. cent. 117 : 9-91. Dr R. H. HEDLEY C. G. OGDEN JUNE I. KRAFFT BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD 5*t PLATE i A. Lateral view of E. acanthophora showing the arrangement of the shell-plates. x 1500 B. Preparation of shell-plates to illustrate two elongated shell-plates. X 3000 C. Lateral view of single apertural-plate with characteristic teeth. X 7600 D. Aperture of E. acanthophora showing the arrangement of the apertural-plates. x 2900 E. Ventral view of apertural-plate preparation, to show thickened dentate edge and thearrangement of three rows of aperture-plates. X 3500 Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i PLATE 2 A. Lateral view of E. strigosa to show the distribution of siliceous spines. x 1200 B. Area of shell surface illustrating the holes left by removal of spines. x 5000 C. Lateral view of a single apertural-plate. X 4700 D. Apertural view of the first stage of division ; note the large pseudopodial trunk (arrowed)with some daughter-shell apertural-plates arranged on it. X 2700 E. Aperture of E. strigosa with eleven evenly spaced apertural-plates. x 3000 Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE 2 D DESCRIPTIONS OF THREE SPECIES OFEUPLOTES (PROTOZOA: CILIATEA) By C. R. CURDS INTRODUCTION IN HIS revision of the order Hypotrichida Stein, 1859, Borror (1972) listed 43 speciesof the genus Euplotes Ehrenberg, 1830 and in the same year Carter (1972) added afurther four new species. The latter author suggested the following charactersconstitute a reliable basis for separating species within the genus ; the pattern of thedorsal interkinetal silver-line network or argyrome, the number of dorsolateralkinetics, the shape of the adoral zone of membranelles and the number of membran-elles therein, the number and arrangement of the ventral cirri and the shape of thenon-dividing macronucleus. Most of these characters were initially introduced byTuffrau (1960) while Borror (1968) added the appearance of the cortical sculpturingthat is sometimes a feature of the dorsal and ventral surfaces of Euplotes spp. Three species of Euplotes are described in the present paper. The first two arefreshwater forms that were isolated from samples of activated sludge and thesewere subsequently identified as E. moebiusi forma quadricirratus and E. affinis formatricirratus respectively. The third is a euryhaline species from Austria which doesnot conform to any of those species described in the literature when both traditionaland modern criteria are taken into consideration. Euplotes moebiusi Kahl, 1932 and the variety with four caudal cirri, E. moebiusiforma quadricirratus Kahl, 1932, have not been described in the literature since theirfirst brief descriptions and illustrations by Kahl (1932). However, this species hasregularly been observed in activated-sludge plants treating sewage and industrialwaste waters over many years (Curds & Cockburn, 1970 ; Ministry of Technology1968). Photographs of E. moebiusi were published by Klein (1958) in order todemonstrate the 'dry' silver method but these were not sufficiently comprehensivefor taxonomic purposes. Euplotes affinis Dujardin, 1841 and its variety with three caudal cirri E. affinisforma tricirratus Kahl, 1932 are also examples that have not been described sincetheir originals and yet have been seen regularly in aerobic waste-treatment processes(Curds & Cockburn, 1970) and in other organically polluted situations (Bick, 1972).Tuffrau (1960) thought it likely that both E. moebuisi and E. affinis were synonymsfor the species E. charon Ehrenberg, 1830. MATERIALS AND METHODS (a) Source and cultivation A clonal culture of Euplotes moebiusi was isolated direct from an activated-sludgesample obtained from Maple Lodge Sewage Treatment Works, Rickmansworth,Hertfordshire. This species was maintained in freshwater Erdschreiber solution Bull. Br. Mus. nat. Hist. (Zool.) 27, 26* n4 C. R. CURDS ('Medium i', Committee on Cultures, Society of Protozoologists, 1958), either intest-tubes or in Petri dishes. The largest populations were obtained in Petri dishescontaining a thin layer of Musgrave and Clegg's agar (2-5 per cent agar, 0-5 per centsodium chloride and 0-5 per cent Liebig's beef extract in distilled water) which wasstreaked with the bacterium Klebsiella aerogenes (National Collection of IndustrialBacteria, NCIB 8017) as a food supply and then flooded with Erdschreiber solution.Subcultures were transferred at monthly intervals. Euplotes affinis was collected by Mr A. Cockburn from a sample of activated sludgetaken from an experimental small-scale pilot plant operated at the Water PollutionResearch Laboratory, Stevenage, Hertfordshire. It was sent to the British Museum(Natural History) as a clonal culture and was maintained in a similar manner toE. moebiusi. The third hypotrich, a small euryhaline Euplotes sp., was originally collected byProfessor E. Tschermak from a freshwater source in Schlospark Schonbrunner inVienna which is the locus classicus for the alga Ruttnera spectabilis Geitler, 1942 (seeGeitler, 1942, 1943). Samples of this alga were sent to Dr Mary Parke at the MarineBiological Station at Plymouth where the hypotrich was first noticed and cultured.Cultures of the ciliate were subsequently deposited with the NERC Culture Collectionof Algae and Protozoa where it was cultivated in saltwater Erdechreiber solution.Later, a clonal culture was established in freshwater and marine media at the BritishMuseum (Natural History) and the descriptions herein relate to organisms from thatclone. This small Euplotes sp. could be maintained equally well in test-tubes orplastic Petri dishes containing either fresh or sea water Erdschreiber solutions. Cul-tures were kept in the dark at room temperature, and were fed at weekly intervalswith a few drops of a thick suspension of baker's yeast. Cultures were transferredat monthly intervals. (b) Microscopy Light microscopy and the methods used for observations and measurements weresimilar to those described by Curds, West & Dorahy (1974). Silver-line systemswere displayed using the 'wet' method of Corliss (1953) in the cases of Euplotes affinisand the small euryhaline species. The 'dry' method of Klein (1958) was used toshow the silver-line system of E. moebiusi. The latter method proved to be farmore reliable and quicker than the conventional 'wet' method. As Klein (1958)pointed out, the success of the 'dry' method depends on the cell drying and dyingmore or less simultaneously and this was achieved by removing excess moisture withthe aid of screws of paper tissue and by flicking single cells out from drops of liquidonto dry parts of the slide by means of an eyelash mounted in a glass rod. Nucleiwere stained using Dippell and Chao's modification of De Lamater's basic fuchsinmethod described by Sonneborn (1950). The nuclei of E. moebiusi and E. affiniswere stained after fixation on the slide by air-drying with equal success as the con-ventional method of chemical fixation. The techniques used for scanning-electron microscopy of the small euryhalineEuplotes sp. were similar to those previously described by Curds et al. (1974) with theexception of the fixation methods. Here the hypotrichs were not killed in osmium THREE SPECIES OF EUPLOTES 115 tetroxide vapour, and fixation was best using the osmium-meruric chloride fixative(Parducz, 1967) which was recommended by Small & Maraszalek (1969). A com-parison of the results obtained by the fixation methods of Curds et al. (1974) and thoseof Small & Marszalek (1969) for this species is demonstrated in PI. i. Plate la showsa cell fixed in Parducz's solution following the recommendations of Small & Mars-zalek (1969) where PI. ib shows a similar cell that had been killed in osmium vapourfixed in a solution containing equal parts of 2 per cent (w/v) osmium tetroxide andsaturated mercuric chloride solutions. It is apparent that the Parducz's fixativewas far better for this species than were the methods of Curds et al. (1974),whereasthe reverse was true for the species Euplotes rariseta Curds, West & Dorahy, 1974.These results suggest therefore that the choice of fixative may vary with the speciesunder consideration. RESULTSEuplotes moebiusi Kahl, 1932 DIAGNOSIS. Medium (60 /-mi long, 40 /Am wide), ovoid freshwater hypotrich with10 frontoventral, 5 transverse and 4 caudal cirri. Ventral surface heavily sculpturedwith 7 ridges, dorsal surface with 5 longitudinal ridges. Adoral zone with 35-40membranelles which extend two-thirds the length of the cell. Dorsal silver-linesystem with 5 longitudinal rows of narrow polygons interspersed with an irregularnetwork of larger polygons ; 7 dorsolateral kinetics bearing a maximum of n dorsalcilia. Macronucleus 3-shaped, micronucleus small, situated anteriorly. Slides showing silver-line systems, ventral ridges and nuclei have been depositedin the slide collection of the B.M. (N.H.), Reg. Nos. 1973:4:14:1-5. DETAILED DESCRIPTION. It can be seen from the size distribution data given onFig. i that Euplotes moebiusi is a medium-sized species and is 62-25 ± 6'6 /u,m longand 39-45 ± 5-87 /u,m wide. The outline shape of the body is oval and there is adefinite notch at the anterior end of the body where the adoral zone of membranelles(AZM) begins. The ventral surface is heavily sculptured with 7 ridges. (Fig. 2).One flattened ridge runs along the edge of the peristome and terminates posteriorlyin a sharp point. One short ridge is restricted to the anterior half of the body andlies between the front-ventral cirri separating cirrus streak I and II from streak III(using the method of cirrus numeration of Wallengren, 1900). Three short ridgesare confined to the posterior and lie between the transverse cirri. One ridge stretchesthe entire length of the body beginning at the anterior notch (between streaks IIIand IV) and terminating between the transverse cirri III i and IV i. One mediumlength ridge is restricted to the central portion of the body and separates cirri V 3and VI 2 from cirrus V 2. The positions and shapes of these ridges are similar tothose figured by Kahl (1932). There are 5 longitudinal ridges on the dorsal surface. Euplotes moebiusi has 10 frontoventral cirri which are distributed as shown inFigs. 2 and 3b. The arrangement resembles that of E. char on. There are 5 transverseand 4 caudal cirri. No specimens were observed with 3 caudal cirri as was shown inthe original descriptions by Kahl (1932). The AZM is composed of 35-40 membran-elles and its extends two-thirds of the way down the body (Fig. 2). 6** n6 C. R. CURDS I 99-S 98~ 3. 95-^-290- CO 80- »> >- CO -4—. = ;-= 6oH 01 La <-> 03 i: -a 40- 20- 20 30 40 50 SIZE (microns) 60 70 FIG. I. Size distribution data of Euplotes moebiusi (broken lines) and Euplotes affinis(continuous lines). Triangles denote width of cells, circles denote length. FIG. 2. Ventral aspect of Euplotes moebiusi showing cirri and ridges(scale represents 10 pim). THREE SPECIES OF EUPLOTES 117 The geometry of the dorsal silver-line system of Euplotes moebiusi differs from anyof those published. It consists (Fig. 3a) of 5 ladder-like longitudinal rows of narrowpolygons with the pits of the dorsal cilia or bristles positioned on the right. In otherEuplotes spp. with a double patella-type of dorsal argyrome the dorsal pits are situatedon the left of the rows of smaller polygons. In addition, an irregular network ofpolygons, resembling that of E. mutabilis Tuffrau, 1960, is sandwiched between theladder-like rows. There are 7 dorsolateral kinetics with the central kinetics bearinga maximum of n dorsal bristles. The ventral silver-line network (Fig. 3b) is aconventional series of irregular polygons whose general pattern closely resemblesthat of E. patella Muller, 1773 (see Tuffrau, 1960). a b FIG. 3. Silver-line system of Euplotes moebiusi, (a) dorsal surface, (b) ventral surface. The macronucleus of Euplotes moebiusi is an irregular 3 -shape (Fig. 4) resemblingthat of E. plumipes Stokes, 1884 (see Tuffrau, 1960), except that the posterior tail isshorter than in that species. The micronucleus is small and lies very close to, andsometimes overlaps, the left anterior edge of the macronucleus. Euplotes affinis Dujardin, 1842Small (38 /um long, 26 /*m wide), ovoid freshwater hypotrich with DIAGNOSIS. 9 fronto ventral, 5 transverse and 3 caudal cirri. Ventral surface sculptured with 3prominent ridges and dorsal surface with 5 longitudinal ridges. AZM with 18-20 n8 C. R. CURDS FlG. 4. M Ventral view of the nuclei of Euplotes moebiusi from a stained preparation(M = macronucleus, m = micronucleus). membranelles which extends two-thirds the length of the cell. There is a smallundulating membrane. Dorsal silver-line system of the double eurystomus typewith 7 dorsolateral kinetics and a maximum of 9 dorsal cilia in the central kinetics.The macronucleus is 3-shaped and there is a small anterior micronucleus. Slides showing silver-line systems and nuclei have been deposited in the slidecollection of the B.M. (N.H.), Reg. Nos. 1973:9:26:1-10. DETAILED DESCRIPTION. Euplotes affinis is one of the smaller freshwater specieswhose dimensions are 38-4 ± 4-3 /^m long and 25-8 ± 4-0 /mi wide. The size distribu-tion data of this species are compared with those of E. moebiusi in Fig. i. Theoutline shape of E. affinis resembles that of E. moebiusi and there is a marked notchat the anterior of the cell which denotes the origin of the AZM. The ventral surfaceis heavily sculptured by 3 longitudinal ridges (Fig. 5) which travel almost the entirelength of the cell. The outer pair of ventral ridges flank the transverse cirri at theposterior end of the body and terminate anteriorly between cirrus streaks I and IIand between streaks V and VI. There are 3 other minor ridges that are restrictedto the posterior end of the cell and these lie between the transverse cirri. Thepositions and shapes of the complete ventral ridging conforms closely with thosefigured by Kahl (1932). The dorsal surface is also heavily sculptured with 5 longi-tudinal ridges. Euplotes affinis has 9 frontoventral cirri whose distribution is shown in Figs. 5and 6b. There are 5 transverse cirri and 3 caudal cirri ; one of the caudals is largerthan the others and is held out stiffly to the right in a manner similar to that of E.rariseta (see Curds et al. 1974). No specimens have been observed with 4 caudal THREE SPECIES OF EUPLOTES 119 cirri as was shown in the original descriptions of E. affinis by Dujardin (1841),although Kahl (1932) described the variety E. affinis forma tricirratus which had 3caudal cirri. The AZM of E. affinis extends two-thirds the length of the body andis composed of 18-20 membranelles which is approximately half the number foundin E. moebiusi. m FIG. 5. Ventral aspect of Euplotes affinis showing cirri, ridges and nuclei(scale represents 10 (zm, M = macronucleus, m = micronucleus). The dorsal and ventral silver-line systems are shown in Fig. 6. The geometry ofthe dorsal argyrome (Fig. 6a) is of the double type resembling that of Euploteseurystomus Wrzesniowski, 1870. There are 7 dorsolateral kinetics in E. affinis andthe central kinetics bear a maximum of 9 dorsal cilia. The ventral silver-line systemis of a common type consisting of a series of relatively few polygons. The macronucleus is 3-shaped and resembles those of Euplotes harpa Stein, 1859and E. plumipes (see Tuffrau, 1960). The micronucleus is small and is situated atthe anterior edge of the macronucleus. Euplotes parkei sp. n. DIAGNOSIS. Small (41 /zm long, 30 /mi wide) euryhaline species ; broadly oval inoutline. Dorsal surface with 6 low longitudinal ridges and ventral surface with 7minor ridges. AZM approximately two-thirds body length, composed of 18 mem-branelles. A deep pocket near the cytostome bears an undulating membrane.Usually 8, but rarely 9, frontoventral cirri ; 5 transverse and 4 caudal cirri. Thereare 8 dorsolateral kinetics with a maximum of n dorsal cilia in the central kinetics. C. R. CURDS Dorsal silver-line system of the double eurystomus-type. Macronucleus C-shapedwith anteriorly situated micronucleus. Type slides showing silver-line systems and nuclei have been deposited in the slidecollection of the B.M. (N.H.). Holotype Reg. No. 1973:4:2:1, and paratype Reg.Nos. 1973:4:2:2-5. a b FIG. 6. Silver-line system of Euplotes affinis, (a) dorsal surface, (b) ventral surface. DETAILED DESCRIPTION. This is a small euryhaline species (41-215-6 /urn long,30-615-3 /u,m wide) whose size distribution data are given in Fig. 7. It is broadlyoval in outline shape with the dorsal surface sculptured with 6 relatively low longi-tudinal ridges. To the left of each ridge is a parallel row of pits from which theshort (2 /urn) dorsal bristles emerge (PI. i, figs, c & d). The ventral surface alsobears 7 longitudinal ridges (PI. i, fig. a) but these are not as prominent as in Euplotesmoebiusi. One ventral ridge travels the complete length of the body along theextreme edge of the peristome, while the other 6 are relatively short and are confinedto the posterior half of the cell. The transverse cirri arise from between these 6minor ridges. The AZM extends two-thirds the length of the body and is composedof 18 membranelles. There is an undulating membrane situated in a deep pocketon the right of the peristome in the proximity of the cytostome. This species of Euplotes usually bears 8 frontoventral cirri which are arranged asshown in Figs. 8, gb and PI. i, fig. a ; however, a gth frontoventral cirrus is occasion-ally present within the same clone and this lies in a position V 2 (PI. i, fig. b). The9th frontoventral cirrus has been found only in animals cultured in freshwaterErdschreiber solution even though the marine cultures have been searched thoroughly THREE SPECIES OF EUPLOTES 121 £= 99- CO 98- o.o 95- •^— QJ "s 90n o COas 80- CD +- t± — 60-| cB -g 4°H Q_ 2Q_ a> 03 20- 10- 5- 2-1- 20 30 40 SIZE (microns) 50 60 FIG. 7. Size distribution data of Euplotes parkei. Triangles denote width and circles length of the cells. for this variant. It is not yet known if the salinity of the culture plays any part inpromoting this type of intraspecific polymorphism. The 5 long transverse and 4caudal cirri were constant in their numbers. There are 8 dorsolateral kinetics in this species and the central kineties bear amaximum of n dorsal cilia. The dorsal and ventral silver-line systems are shownin Fig. 9. The disposition of the dorsal argyrome (Fig. ga) is of the double typeresembling that of Euplotes eurystomus. The ventral silver-line system consists of aseries of few but large polygons and in this respect resembles that of E. cristatusKahl, 1932 (see Tuffrau, 1960). The dorsal argyrome can be seen on some scanning-electron micrographs (PI. i, figs, c & d) as a series of tiny specks. The macronucleus(Fig. 8) is C-shaped and the micronucleus is situated close to the left anterior edge ofthe macronucleus. DISCUSSION AND CONCLUSIONS Euplotes moebiusi is one of the nine species of the genus whose silver-line systemhad not been fully described, and its identity relied solely upon the brief descriptionof Kahl (1932). It is evident from the description of the silver-line system given in 122 C. R. CURDS FIG. 8. Ventral aspect of Euplotes parkei showing cirri, ridges and nuclei(scale represents 10 pun, M = macronucleus, m = micronucleus). a b FIG. 9. Silver-line system of Euplotes parkei, (a) dorsal surface, (b) ventral surface. THREE SPECIES OF EUPLOTES 123 the present paper that E. moebiusi is a distinct species and is not a synonym of E.charon as was suggested by Tuffrau (1960). Before the completion of the present studies it was the opinion of the author thatEuplotes moebiusi was likely to be a synonym of E. affinis because of the similaritiesbetween the two species which may be listed thus ; they both have the same generalshape and size with a definite anterior notch ; both have ventral ridges of similarappearance ; both E. moebiusi and E. affinis are reported to have 4 and 3 caudalcirri variants called E. moebiusi forma quadricirratus and E. affinis forma tricirratusrespectively ; both have 3-shaped macronuclei and both may be found in organicallypolluted freshwater habitats. This meant that there was only one known characterleft to separate the two species, namely the presence or absence of frontoventralcirrus V 2. In view of the findings presented in this paper concerning the intra-specific polymorphism of E. parkei due to the variability of cirrus V 2 it is evidentthat the presence or absence of this cirrus is not as reliable a character as was origin-ally believed. The results presented here however have clarified the situationconsiderably and E. affinis can now be distinguished from E. moebiusi by the 5characters listed in Table i. The most reliable character is the geometry of thedorsal silver-line system which is completely different in the two species (compareFigs. 3b & 6b). In the author's opinion E, affinis should not be regarded as asynonym of E. charon as was suggested by Tuffrau (1960), and there are suffi-cient reliable characters to regard E. affinis as a species distinct from all others. TABLE i List of characters which serve to distinguish between Euplotes moebiusi and Euplotes affinis Character Number of frontoventral cirriNumber of membranelles in AZMMaximum number of dorsal cilia in mid- dorsolateral kineticsDorsal argyromeVentral argyrome E. moebiusi 1035-40 ii Complex E. affinis 918-20 Double eurystomus Many small polygons Few large polygons Agamaliev (1967) reported intraspecific polymorphism in the number of fronto-ventral cirri in his description of Euplotes raikovi Agamaliev, 1966. In the CaspianSea strain of E. raikovi, Agamaliev (1967) noted that there were 7 or 8 frontoventralcirri and indicated that cirrus V 2 was that which did not develop in some specimens.However in a recent paper, Washburn & Borror (1972) described a strain of E.raikovi, isolated from a sand sample taken from the New Hampshire coast of theU.S.A., in which they could not find an 8th cirrus (cirrus V 2) although they didobserve a completely barren plaque in each case. Negative evidence such as thiscan never be conclusive and one must accept that Agamaliev's strain did exhibitpolymorphism of cirrus V 2 as he claimed, particularly in the light of the photo-graphic evidence presented in this paper where there can be no doubt that E. parkeimay have 8 or 9 completely normal frontoventral cirri. More work must be carried 124 C- R- CURDS out on the morphogenesis of E. parkei particularly on the fate of cirrus V 2, but theevidence so far obtained indicates that there is not even a barren plaque in the caseof E. parkei specimens with 8 frontoventral cirri. Furthermore, more work isneeded to test adequately whether or not intraspecific polymorphism such as thiscan be induced by adjusting the salinity of the culture medium or if the results sofar obtained can be attributed to pure chance. It is possible that one of the reasons why Euplotes parkei has remained unnoticeduntil now is because none of its characters, in isolation, will distinguish this speciesfrom all others. A combination of characters is required to do so, but there can belittle doubt that this small euryhaline Euplotes is a separate and distinct species.The following species have the combination of characters - a double dorsal argyromewith 8 or 9 frontoventral, 5 transverse and 4 caudal cirri - E. aediculatus Pierson,1943 ; E. amieti Dragesco, 1970 ; E. aspheronicus Agamaliev, 1966 ; E. diadaleosDiller & Kounaris, 1966 ; E. eurystomus ; E. octocarinatus Carter, 1972 ; E. patella ;E. patella forma latus Agamaliev, 1967 ; E. plumipes ; E. tegulatus Tuffrau, 1960 ;E. tuffraui Berger, 1965 ; E. variabilis Stokes, 1887 (see Carter, 1972) ; and E.zenkewitchi Burkovsky, 1970. However only four of these, E. aediculatus, E.eurystomus, E. variabilis and E. octocarinatus, have 8 dorsolateral kinetics. SinceE. octocarinatus has a patella-like double dorsal argyrome there are only three remain-ing species with which to compare and contrast E. parkei. All three of theseEuplotes spp. are large (over 100 jum long) and have at least 40 membranelles in theAZM, whereas E. parkei is small (under 50 /um long) and has less than 20 mem-branelles. Furthermore, the shapes of the macronuclei differ and all three specieshave many more dorsal cilia than E. parkei. It is evident therefore that Euplotes parkei differs from all previously describedspecies of Euplotes and the differences are considered to be sufficiently distinct forthis organism to be designated as a separate species. It is named Euplotes parkeiafter Dr Mary Parke of the Marine Biological Station Plymouth who first isolatedand cultivated this hypotrich. Following the revised classification of the Committeeon Taxonomic Problems of the Society of Protozoology (Honigberg et al., 1964),Euplotes parkei is placed into class Ciliatea Perty, 1852, order Hypotrichida Stein,1859, family Euplotidae Ehrenberg, 1838. ACKNOWLEDGEMENTS The author is indebted to Mrs J. E. Dorahy, Department of Zoology, BritishMuseum (Natural History), for her technical assistance, to Miss B. J. West, NERCCulture Centre for Algae and Protozoa, who made the original silver preparations ofEuplotes parkei, to Mr A. Cockburn, Water Pollution Research Laboratory of theDepartment of the Environment, who isolated E. affinis, to Miss M. Yeoh, MapleLodge Sewage Treatment Works, Rickmansworth, who supplied samples of activatedsludge containing E. moebiusi and to Mr R. Harris, Department of Zoology, BritishMuseum (Natural History), for his aid and advice on the preparation of ciliates forscanning-electron microscopy. THREE SPECIES OF EUPLOTES 125 REFERENCES AGAMALIEV, F. G. 1967. Faune cilies mesopsammiques de la cote ouest de la Met Caspienne. Cahiers de Biol Mar. 8 : 359-402.BICK, H. 1972. Ciliated Protozoa. An illustrated guide to the species used as biological indicators in freshwater biology. World Health Organization, Geneva.BORROR, A. C. 1968. Systematics of Euplotes (Ciliophora, Hypotrichida) ; towards union of the old and the new. /. Protozool. 15 (4) : 802-808.- 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Protozool. 19 (i) : 1-23.CARTER, H. P. 1972. Infraciliature of eleven species of the genus Euplotes. Trans. Amer. Microsc. Soc. 91 (4) : 466-492.COMMITTEE ON CULTURES, SOCIETY OF PROTOZOOLOGISTS. 1958. A Catalogue of Laboratory Strains of Free-living and Parasitic Protozoa. /. Protozool. 5 (i) : 1-38.CORLISS, J. O. 1953. Silver impregnation of ciliated protozoa by the Chatton-Lwoff technic. Stain Tech. 28 : 97-100.CURDS, C. R. & COCKBURN, A. 1970. Protozoa in biological sewage-treatment processes - I. A survey of the protozoan fauna of British percolating niters and activated-sludge plants. Wat. Res. 4 : 225-236.— WEST, B. J. & DORAHY, J. E. 1974. Euplotes rariseta sp. n. (Protozoa, Ciliatea), a new small marine hypotrich. Bull. Br. Mus. nat. Hist. (Zool), 27 (2) : 95-102.DUJARDIN, F. 1841. Histoire Naturelle des Zoophytes. Infusoires. Paris. 678 pp.GEITLER, L. 1942. Neue luftlebige Algen aus Wien. Ost. hot. Z. 41 (i) : 49-51. 1943- Eine neue atmophytische Chrysophycee, Ruttnera spectabilis, nov. gen., nov. spec. Int. Revue ges. Hydrobiol. 43 : 100-109. HONIGBERG, B. M., BURBANCK, W., BOVEE, E. C., CORLISS, J. O., GOJDICS, M., HALL, R. P., KUDO, R. R., LEVINE, N. D., LOEBLICH, A. R., WESIER, J. & WENRICH, D. H. 1964. A revised classification of the Phylum Protozoa. /. Protozool. 11 : 7-20.KAHL, A. 1932. Urtiere oder Protozoa. I : Wimpertiere oder Ciliata (Infusoria), eine Bearbeitung der freilebenden und ectocommensalen Infusorien der Erde, unter Ausschluss der marinen Tintinnidae. Un: Dahl, F. Die Tierwelt Deutschlands, Teil 25, pp. 399-650. G. Fischer, Jena. KLEIN, B. M. 1958. The 'dry' silver method and its proper use. /. Protozool. 5 (2) : 99-103.MINISTRY OF TECHNOLOGY. 1968. Protozoa in sewage-treatment processes. In: Notes on Water Pollution, No. 43. H.M.S.O., London.PARDUCZ, B. 1967. Ciliary movement and co-ordination in ciliates. Int. Rev. Cytol. 21 : 91- 128.SMALL, E. B. & MARSZALEK, D. S. 1969. Scanning electron microscopy of fixed frozen and dried Protozoa. Science, N.Y. 163 : 1064-1065.SONNEBORN, T. M. I95O. Methods in general biology and genetics of Paramecium aurelia. J. exp. Zool. 113 : 84-147.TUFFRAU, M. 1960. Revision du genre Euplotes, fondee sur la comparaison des structures superficielles. Hydrobiologia, 15 : 1-77.WALLENGREN, H. 1900. Zur Kenntnis der vergleichenden Morphologie der Hypotrichen. Bih. K. svenska Vetensk Akad. Handl. 26 : 1-31.WASHBURN, E. S. & BORROR, A. C. 1972. Euplotes raikovi Agamaliev, 1966 (Ciliophora, Hypotrichida) from New Hampshire : Description and morphogenesis. /. Protozool. 19 (4) : 604-608. Dr C. R. CURDS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD 6*f PLATE i Scanning-electron micrographs of Euplotes parkei a. Ventral view showing ridges and cirri. Cell fixed in Parducz's (1967) fixative. b. Ventral view of a 9 frontoventral cirri variant. Cell killed and fixed following the methodsof Curds et al. (1974). c. Dorsal view showing the outline of the dorsal argyrome as specks. d. Dorsal view showing ridges, dorsal cilia and outline of argyrome. Bull. Brit. Mus. nat. Hist. (Zool). 27, 2 PLATE i NEW RECORDS OF BATS FROM SOUTH-EASTERN ASIA, WITH TAXONOMIC NOTES By J. E. HILL INTRODUCTION BATS from Malaya, Java and Sulawesi (Celebes) identified in recent years at theBritish Museum (Natural History) have included specimens representing a numberof poorly known species and thereby of taxonomic interest and importance, or whichprovide further distributional records. The majority of the Malayan specimenshave come to London through the agency of Lord Medway and were collected byhim or by Mr G. C. Yong. A further interesting specimen from Malaya has beenprovided by Dr D. R. Wells of the School of Biological Sciences, the University ofMalaya. The Indonesian specimens are from a collection submitted for identifica-tion by Captain P. F. D. Van Peenen, M.C., U.S.N., Officer in Charge, U.S. NavalMedical Research Unit No. 2, Djakarta Detachment. My thanks are due also toDr G. G. Musser and Dr Karl F. Koopman of the American Museum of NaturalHistory, New York, who waived a prior claim to the Sulawesian specimens, and toDr H. W. Setzer of the United States National Museum of Natural History, theSmithsonian Institution, Washington, who arranged the loan of one of thespecimens discussed. Measurements are in millimetres : unless otherwise indicated,the specimens have been donated to the collections of the British Museum(Natural History). SYSTEMATIC SECTION ^ Chironax (?) melanocephalus (Temminck, 1825)Sulawesi: Soroako, south Sulawesi. $ (young adult) B.M. 73.1802. Chironax has been unreported hitherto from Sulawesi. This young adult specimendiffers in a number of features from Malayan material referred to C. melanocephalusand apparently also in some ways from Javan specimens, of which none is availablefor comparison. Consequently, it is referred to C. melanocephalus with considerablehesitation. This Sulawesian specimen lacks the distinctive blackish cap usually characteristicof Chironax although the nape and crown are darker brown than the back which iswarm brown, tinged with grey over the shoulders. The throat and the sides of theneck are creamy white, the belly yellowish white and the flanks brown, this colourextending across the hinder part of the ventrum. In colour the specimen agreesquite well with the original description by Temminck (1825 : 190) of specimens from Bull. Br. Mus. nat. Hist. (Zool.) 27, 27 128 J. E. HILL Java : it differs from mainland specimens in its generally paler dorsal surface andalso lacks any rufous or orange at the sides of the neck, a feature sometimes found inmainland examples. There are some differences in wing structure when a comparison is made withmainland and Javan specimens. In particular, the fifth metacarpal is relatively alittle longer, and the second phalanges of the third, fourth and fifth digits are rela-tively a little shorter, those of the fourth and fifth digits not exceeding in length thefirst phalanges of their respective digits as they do in continental and Javan examples.The relevant wing structures are summarized in Table i and in these respects itcan be seen that the Sulawesian specimen approaches the closely related genusBalionycteris. TABLE i Wing indices of Chironax and BalionycterisLength of forearm = 1000 Chironax Balionycteris Malaya Java* Sulawesi Third digit Metacarpal 677-711 640 635 705 710-780 First phalange 474~533 489 488 5°7 474~548 Second phalange 584-657 640 623 568 559-653 Fourth digit Metacarpal 617-679 617 578 641 693-739 First phalange . 368-415 373 378 373 374-43° Second phalange 396-444 418 422 360 352-415 Fifth digit Metacarpal 659-704 640 622 708 712-764 First phalange 3i8~36o 344 322 319 330-364 Second phalange 339~384 348 344 3*7 3^-354* 'Co-types' of C. melanocephalus, from Andersen (1912 : 676). Cranially the specimen from Sulawesi is a little smaller than the mainland examplesbut its cranial dimensions fall within the range of variation reported for the Javan'cotypes' by Andersen (1912 : 678). The supraorbital region is a little more swollenmedianly than in continental specimens and the postorbital processes are moremassively developed. As might be expected in a young adult, the premaxillae arenot solidly fused anteriorly. The post-canine teeth are generally rather smallerthan are those of specimens from the mainland, with pm4 more rounded, lessrectangular in outline, while pm3 lacks completely the antero-external cusp customaryin C. melanocephalus. This cusp, however, is very small in some Malayan specimens. Chironax melanocephalus has been reported from Java, Sumatra (Chasen, 1940 : 28,30), Nias Island (Thomas, 1923 : 252), Malaya (Chasen, loc. cit. ; Hill, 1961 : 640)and from southern Thailand (Hill and Thonglongya, 1972 :181), who also reportedfurther Malayan specimens. The presence of Chironax in Sulawesi is not unexpected BATS FROM SOUTHEASTERN ASIA 129 but the exact taxonomic status of the Sulawesian population must remain to someextent uncertain until further specimens from Sulawesi and Java are available. Measurements of the specimen from Sulawesi : length of forearm 44-2 ; length ofthird metacarpal 31-1 ; length of its first phalange 22-4 ; length of its secondphalange 25-1 ; length of fourth metacarpal 28-3 ; length of its first phalange 16-5 ;length of its second phalange 15-9 ; length of fifth metacarpal 31-3 ; length of itsfirst phalange 14-1 : length of its second phalange 14-0 ; greatest length of skull22-2 ; condylobasal length 21-4 ; condylocanine length 20-8 ; palatal length n-i ;palation to incisive foramina 9-3 ; palation to basion 8-2 ; length orbit-nares 5-2 ;width of braincase 9-8 ; mastoid width 10-0 ; zygomatic width 14-3 ; n^-m1(crowns) 6-3 ; p4-p4 (crowns) 6-3 ; lachrymal width 6-1 ; c1-^1 (cingula) 4-3, (alveoli)4-0 ; postorbital width 5-5 ; interorbital width 4-9 ; width of mesopterygoid fossa2-9 ; width between p4-p4 3-8 ; width between bases of canines 1-9 ; orbital dia-meter 5-6 ; length of mandible from condyle 16-0 ; length of complete mandible fromboth condyles 15-2 ; coronoid height 7-3 ; c-m1 (crowns) 6-9 ; c-m2 (crowns) 7-5(in order of Andersen, 1912 : 678, with interpolations). Measurements of teeth : length x width of pm3 1-7x1-1 ; of pm4 1-7x1-2 ; ofm1 1-3x0-9: of pnij 0-7 x 0-7 ; of pm3 1-6x1-1 ; of pm4 1-7 x 1-2 ; of nij 1-4x0-9;of m2 0-8x0-6 (notation of teeth based on Andersen, 1912 : 680). It may be noted that by error the illustration of Chironax in the editions and print-ings of Walker (1964, 1965, 1968) is not of C. melanocephalus but of a youngRousettus. ~ Rhinolophus pusillus Temminck, 1834 Malaya : Pasang Kamunting, Kg. Ginting, Penang. $ B.M. 73.608. Andersen (1905 : 121) summarized the many different forms of Rhinolophus beforethen confused under Rhinolophus minor Horsfield, 1824, envisaging a lepidus groupcomposed of lepidus, minor and subbadius subgroupings. Later, the same author(1918 : 376) in a paper issued on his behalf by Oldfield Thomas renamed this thepusillus group and added a number of briefly diagnosed new forms. Tate andArchbold (1939 : 3) listed the names allocated to the group, retaining the sub-divisions proposed by Andersen. These authors included R. minor and R. pusillusas distinct species but Andersen (1905 : 126) considered pusillus a synonym of minor :however, Rhinolophus minor Horsfield, 1824 is preoccupied (Chasen, 1940 : 38 ;Ellerman & Morrison-Scott, 1951 : 116). The species has not hitherto been recorded from the Malay Peninsula : this speci-men from Penang agrees closely with a small series from Java and with one specimenfrom Madura identified by Andersen as R. pusillus, the species that he called R.minor in his early investigation (1905 : 121) of the group. There is close agreementalso between the Penang specimen and a series in the collections of the BritishMuseum (Natural History) from the island of Tioman, off the east coast of Malaya.Rhinolophus pusillus is small, easily recognized by its upright triangular connectingprocess and small skull (Table 2). 7* 13o J. E. HILL it 41 fl rt ti C C too?*E2 o3 rt rt CH f* 3 ga g 2^ >>>>csS5ogg M ;- ;- ^ g ^ H H H Eui-o pI o O* N c^ O co •^* io vb vb vb vo o 9}.9jduioo jo l-t •*$• ON M b I b ON b IO ON O» O O ON ON O 8UI-O vO CO OO vp CO>O IO IO if) if) 8UI-8UI To-to OO OO CO CO I CO CO CO ON CO 00 ro ro ro ro -s; J"o8•»•« ft! 9S'BOUIBjq jo M op op ON 00 t^ vb vb vb I vb CO 00 IO M N p N c^ I c^ f^ cb ob ob cb HO O t*^ vO t^ 00 I CO CO CO CO | CO jo i i «-) CO CO vp M H M jo cocococo rorororororo X9g CH-CH-CH-Of'ri^CH-CH-CH- I *O H. H. M. H O M M M ON 00 00MVOOvO M <S <N BATS FROM SOUTHEASTERN ASIA 131 Andersen (1905 : 126) recorded R. minor (= pusillus) from Darjeeling, Thailandand Java. As Osgood (1932 : 215) pointed out, when Andersen wrote further on thepusillus group (1918 : 376) and named R. blythi blythi and R. b. szechwanus, evidentlyhe had concluded that the mainland forms were separable from pusillus of Java.Indeed, Andersen has re-labelled the specimens in the British Museum (NaturalHistory) on which the continental records of pusillus were based as 'szechwanus'.Osgood considered that specimens obtained in Cochin China (South Vietnam) byDelacour and Lowe (B.M. 28.7.1.18-19) and referred to pusillus by Thomas weredoubtless closely related if not identical to R. blythi calidus Allen, 1923, to whichOsgood referred a specimen from Tonkin (North Vietnam), suggesting that since thedistinctions between the various forms seemed unclear it might be preferable totreat them as races of pusillus, the earliest name. There is every probability,therefore, that Rhinolophus pusillus Temminck, 1834 must replace R. blythi Andersen,1918 if this is thought to be specifically distinct. Ellerman and Morrison-Scott(1951 : 117) considered R. blythi a subspecies of R. cornutus Temminck, 1835, towhich Van Peenen, Ryan & Light (1969 : 61) allocated specimens from South andNorth Vietnam (presumably those reported by Osgood) and from Thailand. How-ever, Wang et al. (1962 : 556, 558, 568) reported R. cornutus pumilus Andersen, 1905and R. blythi (as a species) from Kwangsi. ^ Rhinolophus acuminatus acuminatus Peters, 1871Malaya : Kuala Kenarong, Ulu Perak. £, $ B.M. 73.609-610. Rhinolophus acuminatus has remained unreported from Malaya although there is anumber of records from Thailand, Laos and Cambodia (Hill & Thonglongya, 1972 :185), and the species occurs also on Sumatra and the nearby islands of Engano andNias, on Borneo, Java and on Lombok. Like those from Thailand and Laospreviously reported, these Malayan specimens agree closely in size with R. a.acuminatus from Java. Both have been preserved in alcohol. The female is verysimilar in colour to reddish brown examples from Java, its dorsal surface brown,tinged with russet, especially posteriorly. Ventrally it is greyish brown, the hairspale based, pale grey brown for much of their length and extensively tipped withgrey. Dorsally, the male is browner, lacking russet, lightly tipped over the shoulderswith silver grey : ventrally it is paler than the female, more nearly fawn, with abrownish tinge. Measurements ($ B.M. 73.609, $ B.M. 73.610) : length of forearm 49-1, 47-8 ;width of noseleaf 8-1, 8-0 ; greatest length of skull to canine 21-5, 20-5 ; condylo-canine length 19-2, 18-2 ; width across nasal swellings 6-1, 5-8 ; least interorbitalwidth 2-5, 2-7 ; zygomatic width n-6, 11-3 ; width of braincase 9-0, 9-0 ; mastoidwidth 10-2, 9-9 ; c1-c1 5-6, 6-0 ; m3-m3 8-3, 8-3 ; c-m3 8-6, 8-0 ; length of completemandible 15-2, 13-7 ; c-m3 9-2, 8-8. The subspecies of R. acuminatus as they are defined by Andersen (1905 : 132,1906 : 657) seem separable only by relatively small features. Mainland specimenshave the lower part of the lateral margins of the sella more or less parallel as in 132 J. E. HILL acuminatus from Java and in audax from Lombok, the sella not abruptly widenedbasally as in sumatranus from Sumatra or circe from Nias, or in calypso from Enganoin which the sella, although widened, is less so than in these but which has a verywide noseleaf. There seem no consistent size differences between the subspecies :a series from Java suggests a degree of sexual dimorphism in size, with femalestending to be a little smaller than males. Myotis horsfieldii horsfieldii (Temminck, 1840)Sulawesi: Wawondula, south Sulawesi. $ B.M. 73.1804. This is the first of Myotis horsfieldii to be reported from Sulawesi, the speciesbeing known otherwise from Java, Borneo (Medway, 1965 : 60), Malaya and perhapsThailand (Hill, 1972 : 31). For the present it is referred to the nominate subspecies :cranially it corresponds to or slightly exceeds the upper limits of size for Javanspecimens as represented in London, but these derive from a single series from onelocality. It is similar in size to the larger of Bornean specimens (M. h. lepidus(Thomas, 1915)) but no more than three examples of this subspecies are available.Medway (1965 : 60) suggests that lepidus from Borneo may be distinguished fromJavan horsfieldii by slightly shorter skull and slightly narrower braincase, conditionsnot supported by the available specimens. The condylobasal length of eighteenJavan specimens ranges from 13-2 to 14-7, in three from Malaya from 13-7 to 14-3and from 14-3 to 14-7 in three from Borneo : the width of the braincase varies from6-9 to 7'4 in Javan examples, from 6-9 to 7-3 in those from Malaya and from 7-1 to7-3 in the specimens from Borneo, which include the holotype of lepidus. TheBornean subspecies appears therefore to be only weakly if at all separable. Measurements of the specimen from Sulawesi: length of forearm 37-7 ; greatestlength of skull 15-7 ; condylobasal length 14-4 ; condylocanine length 13-8 ; leastinterorbital width 3-6 ; zygomatic width — ; width of braincase 7-4 ; mastoidwidth 8-0 ; c^c1 4-2 ; m3-m3 6-0 ; c-m3 5-7 ; length of complete mandible 11-3 ;c-m3 6-2. The registration numbers of Malayan specimens measured by Hill(1972 : 31) are incorrectly assigned. The measurements cited are of B.M. 61.2133(forearm only), B.M. 65.320, B.M. 65.321, and of $ B.M. 16.4.21.16 (this numberomitted) from Batu Burong, Pahang, in that order. ^ Pipistrellus circumdatus (Temminck, 1840) Java : Cibodas ( = Tjibodas), Cianjur, west Java, o6°45' S, I07°oo' E, at 1350 m.<$. Original number 1435. Bogor Museum No. 10069 (a* present at UnitedStates National Museum of Natural History, the Smithsonian Institution,Washington). Kandang Badak, Cianjur, west Java, o6°47' S, io6°59' E, at 2425 m. <$. Originalnumber 1761. U.S. Naval Medical Research Unit No. 2, Djakarta Detachment. Situ Gunung, Sukabumi (= Soekaeboemi), west Java, o6°5o'S, I07°55'E, at1000 m. $. B.M. 73.1805. Malaya : Telecommunications Tower, Eraser's Hill, Pahang. ^ B.M. 73.618. BATS FROM SOUTHEASTERN ASIA 133 Extant specimens of the poorly known south-east Asian species Pipistrelluscircumdatus (Temminck, 1840) were briefly reviewed recently by Hill (1972 : 189),who described a closely related species, P. societatis, from Malaya. At that time, nomore than five examples of P. circumdatus were known. Although Temminck (1840 : 215) remarked that Boie and Macklot, who firstobtained the species in the Tapos district of Java, sent several individuals to theMusee des Pays-Bas (the Rijksmuseum van Natuurlijke Historic, Leiden), thecatalogues by Jentink (1887 : 277 ; 1888 : 178) of that collection report only a singlespecimen, in 1887 as 'one of the types of the species' for the skull, and in 1888 as'type of the species' for the mounted specimen to which the skull belongs. Thisspecimen remains in the Rijksmuseum van Natuurlijke Historic : a second examplefrom Java, now in the collection of the British Museum (Natural History) (B.M.7.1.1.401) is from the collection of R. F. Tomes, and since this collector obtainedspecimens from many sources, including the major museums of his day, it is possiblethat it is one of the original specimens obtained by Boie and Macklot. Known continental specimens are limited to three. One, (B.M. 61.12.10.1) inthe collection of the British Museum (Natural History), is of uncertain provenance.It came from T. C. Jerdon and is said by Blanford (1891 : 312) to have originatedfrom southern India. Another (A.M.N.H. 114850), in the American Museum ofNatural History, is from Pyepat, Upper Burma, and has been reported by Anthony(1941 : 81), Tate (1942 : 250) and by Hill (1972 : 36). The third, listed above, fromEraser's Hill, Pahang, Malaya, was briefly reported by Hill (1972 : 36, footnote). Itis therefore of particular interest to record three specimens from Java obtained bythe United States Medical Research Unit No. 2, Djakarta Detachment, and to presenta more detailed account of the newly collected Malayan specimen. The specimens from Java agree closely with the description by Temminck and withB.M. 7.1.1.401. The males are exactly similar in colour to this example, the blackishbrown dorsal pelage finely but profusely tipped with shining orange or bronze, theventral pelage dark brown, the hairs lightly tipped with grey except on the flankswhere the tipping is brownish, sometimes faintly orange. The membranes areblack and in B.M. 7.1.1.401 evidently have faded to a browner shade. The femalespecimen has dorsal pelage a little less finely tipped with orange, the individualorange hair tips slightly more extensive to give a coarser effect : its ventral surfaceresembles the ventral surface of the other specimens although the tips of the hairstend to be more yellowish buff, especially anteriorly. Cranially, the specimens con-form closely to B.M. 7.1.1.401, described, with the skull of the holotype of circum-datus, in the account already published (Hill, 1972 : 36). The example from Malaya is very similar to the Burmese specimen (A.M.N.H.114850) discussed by Hill (1972 : 36), its skull differing slightly in a number of featuresfrom the skulls of Javan specimens. It is generally a little smaller ; the cranialcrests are less prominent, the supraorbital ridges slightly less evident but in thisexample not terminating in slight tubercles ; the frontal depression is a little shal-lower but the post-palatal extension is not shorter than in the Javan skulls as is thepost-palatal extension of the Burmese example. As in the Burmese specimen, thepost-palatal spine is narrow : although broken in two of the Javan skulls examined, I34 J- E- HILL in the other two skulls from Javan specimens the spine is more broadly based. Thereis little difference in the dimensions of the teeth, although the canines of the Malayanspecimen are very slightly less massive than are those of the Javan examples. Initially, a comparison of the Burmese specimen with Javan material suggested(Hill, 1972 : 36) that the mainland and Javan populations might be subspecificallydistinct. The new material confirms that small differences exist between the twopopulations but as yet insufficient specimens are available for any definite conclusionto be drawn. The Malayan specimen is of especial interest since it demonstrates thepresence of P. circumdatus in Malaya with the newly described P. societatis, a closelysimilar species differing chiefly in smaller size, particularly of the rostrum, palateand dentition, and in its short post-palatal region (length of bony post-palate infour Javan specimens of circumdatus 1-5-1-7, in the Malayan specimen 1-7, in theholotype of societatis (B.M. 67.1605) i.i). Measurements ($ 1435, 3 1761, ? B.M. 73.1805 from Java, $ B.M. 73.618 fromMalaya, in that order) : length of forearm 41-2, 41-1, 42-1, 40-5 ; greatest length ofskull 16-0, 16-3, 16-6, 15-5 : condylobasal length 15-7, 15-7, 16-0, 14-9 ; condylo-canine length 15-4, 15-4, 15-7, 14-7 ; palatal length (excluding the post-palatal spine)8-4, 8-4, 8-6, 8-0 ; length palatal bridge (excluding the post-palatal spine) 6-5, 6-6,6-8, 6-4 ; length orbit-gnathion 4-2, 4-0, 4-2, 4-1 ; lachrimal width 7-1, 7-2, 7-3, 7-2 ;width across supraorbital tubercles or swellings 6-6, 6-6, 6-7, 6-4 ; zygomatic width11-4, —, 11-7, — ; least interorbital width 4-2, 4-4, 4-3, 4-3 ; width of braincase 8-0,8-1, 8-1, 7-9 ; mastoid width 8-7, 8-7, 9-1, 8-5 ; c1-^ 5-2, 5-3, 5-4, 5-0 ; m3-m3 —,7-6, 7-7, 7-3 ; c-m3 6-2, 6-1, 6-3, 5-9 ; length of complete mandible —, 11-9, 11-9,n-i ; c-m3 6-5, 6-4, 6-7, 6-3. Philetor brachypterus verecundus (Chasen, 1940)Malaya : Ulu Gombok, Selangor. <$ B.M. 73.597. The genus Philetor with its sole species P. brachypterus is known from New Guinea,Malaya, Sumatra and, doubtfully, from Java and Banka (Hill, 1971 : 140). Itsoccurrence in Malaya was based initially on two female specimens from Perakoriginally described as Eptesicus verecundus by Chasen (1940 : 53), one the holotypecollected on Mount Kledang by Dr R. Hanitsch (now B.M. 47.1437) and the other(at one time in the collections of the former Raffles Museum, Singapore) from anunspecified locality in that State. Medway (1969 : 35) mentions that it hasbeen collected in Selangor. Eptesicus verecundus was transferred to Philetorby Hill (1966) who considered it a subspecies of Philetor rohui Thomas, 1902 fromNew Guinea : later, the same author (1971) referred Vespertilio brachypterus Tem-minck, 1840 from Sumatra to Philetor as a third subspecies, brachypterus takingprecedence as the specific name. This further specimen of P. b. verecundus has been donated to the collections of theBritish Museum (Natural History) by Dr D. R. Wells of the School of BiologicalSciences, the University of Malaya. Its muzzle, ears and tragus agree closely withthose of the holotype : the structure of the penis, hitherto unknown in verecundus, BATS FROM SOUTHEASTERN ASIA 135 is almost exactly like that of P. b. rohui (figured by Hill, 1966 : 375, fig. i), differingonly in slightly wider and deeper fissures in the ventral part of the glans. Cranially there is close agreement with the holotype of verecundus. The braincaseis very slightly more inflated than in P. b. rohui as it is in the holotype : the narialemargination, however, rather wider at the roots of i2~2 and broadly V-shapedapically in the holotype of verecundus, is in this specimen rather narrower and morerounded at the apex, thus conforming with rohui and with the holotype of P. b.brachypterus from Sumatra (Hill, 1971 : 141). The upper canines of the specimenfrom Selangor are a little more massive than in the holotype of verecundus, and theSelangor specimen differs from this and from rohui in having a narrow rather thanwide post-palatal spine, and its basioccipital pits are a little less prominent. Measurements : length of forearm 35-9 ; greatest length of skull — ; condylobasallength 14-7 ; condylocanine length 14-3 ; lachrimal width 6-7 ; width across ante-orbital foramina 5-5 ; least interorbital width 4-6 ; zygomatic width — ; width ofbraincase 8-3 ; mastoid width 9-0 ; c^c1 5-4 ; m3-m3 6-7 ; c-m3 4-8 ; i2-m3 5-6 ;length of complete mandible 10-4 ; c-m3 4-9. Miniopterus medius macrocneme Revilliod, 1914Sulawesi: Wawondula, south Sulawesi. $, <$ B.M. 73.1806-1807. The length of the forearm and tibia, with the globose, high braincase and slendercanines refer these specimens to M. medius, hitherto unreported from Sulawesi.They display the less massive dentition characteristic of the easterly subspeciesM. m. macrocneme which until now has not been known to occur further west thanthe islands of Ambon and Ceram, in the Molucca Islands. Measurements ($ B.M. 73.1807, $ B.M. 73.1806) : length of forearm 44-4, 43-3 ;length of tibia 19-1, 18-8 ; greatest length of skull 13-7 ; 13-2 ; condylobasal length13-1, 12-9 ; condylocanine length 12-6, 12-1 ; least interorbital width 3-4, 3-5 ;zygomatic width 7-4, 7-5 ; width of braincase 7-4, 7-2 ; mastoid width 7-6, 7-5 ;c1-^1 3-9, 3-7 ; m3-m3 5-4, 5-4; c-m3 5-1, 5-0 ; length of complete mandible 9-7,9'5 ; c-m3 5-6, 5-3. Phoniscus atrox Miller, 1905Malaya : Tekom Forest Reserve, Jerantut, Pahang. <$ B.M. 71.1135. Until recently P. atrox was known in the literature from the holotype from Sumatraand from two further examples reported from southern Thailand by Kloss (1916 :12).The first record for Malaya is of a subadult from the Ulu Gombok Forest Reserve,Selangor, at 2000 ft, reported by Medway (1969 : 43). This second Malayan speci-men agrees exactly with those from southern Thailand. Measurements : length of forearm 34-2 ; greatest length of skull 15-5 ; condylo-basal length 13-7; condylocanine length 13-7 ; least interorbital width 3-8; 136 J. E. HILL zygomatic width 8-7 ; width of braincase 7-0 ; mastoid width 7-4 ; c^-c1 3-6;m3-m3 5-5 ; c-m3 6-0 ; length of complete mandible 10-5 ; c-m3 6-5. Tadarida johorensis (Dobson, 1873) Malaya : Pulai, Kelantan, 4^8' N, ioi°57' E, c. 780 ft. &J (n subadult) B.M.73-632, 636, 638-642, 647, 649-651, $$ (6 subadult) B.M. 73.633-635, 637, 643, 644,646, 648. Extant accounts of T. johorensis are based on one or other of two specimens. Thedetailed descriptions by Dobson (1876 : 180, 183, fig. a ; 1877 : 718, 726, fig. 5 ;1878 : 421, 432) refer to the holotype from Johore, Malaya. Later, Andersen(1907 : 39, 44) described a further specimen from Soekaranda, Deli, Sumatra andgave an account of the features of its skull and dentition. These remained apparentlythe only known specimens until Chasen (1940 : 59) recorded the species from Selangor,Malaya and from the island of Sri Buat, off the east coast of Johore. More recently,Medway & Yong (1969 : 33) have recorded the material from Kelantan on whichthese notes are based, but as T. plicata. Dobson and Andersen in their descriptions of this species emphasize the largeinteraural pocket which can be closed by an extension of the interaural band. Theinner or anterior margins of the ears are joined by a thick, deep band of integumentoriginating from a low transverse ridge on the muzzle, its upper margin elevatedbetween the ears in a truncately triangular flap. Posteriorly the lower part of thisband forms the anterior wall of a deep subtriangular or subquadrangular pocketbetween the ears, its lateral walls more or less parallel and its posterior wall a low,slightly curved ridge. The pocket is divided by a low median septum and containsa few long hairs at the bottom of its anterior part : otherwise the integument formingand surrounding the pocket is naked or nearly so. The free upper part of theinteraural band is convex anteriorly, hollowed posteriorly, and forms a lid or coverto the pocket. Previous descriptions of this structure refer only to the male. Theinteraural box in two adult females from Kelantan exactly resembles that of the adultmale : there is little variation among the younger examples in the series but in thesethe posterior wall of the pocket is sometimes less prominent and the pocket a littleshallower than in the older specimens. The dorsal pelage of specimens (in alcohol)from Kelantan is uniformly mid-brown, the hairs paler at the extreme base. Theventral pelage is predominantly fawn but is browner at the flanks, this shade extend-ing also to the sides of the throat. Tadarida johorensis is very similar to T. plicata from which it differs chiefly in thepresence of the interaural pocket. The small cranial differences separating the twospecies are admirably summarized by Andersen (1907 : 40) and are confirmed by anadult female from the series from Kelantan. These are the absence of markedcranial ridges in johorensis, its low, ill-defined sagittal crest terminating more pos-teriorly and not extending on to the supraorbital region, with the facial foramenplaced farther posteriorly immediately above the narrowest part of the postorbital BATS FROM SOUTHEASTERN ASIA 137 constriction ; a low rostrum in johorensis, with straight or slightly concave rostralprofile which contrasts with the more inflated, convex profile of plicata ; a shallownarial emargination in johorensis which extends posteriorly to a line joining theanterior margins of the anteorbital foramina ; the upper incisors are shorter andmore massive in johorensis which has shorter upper canines and smaller anteriorpremolars (pmf), with the principal cusp of the second upper premolar (pm4) lessdeveloped. Measurements of adult specimens from Kelantan (<$ B.M. 73.636, $ B.M. 73.635(forearms only), $ B.M. 73.637) : length of forearm 46-7, 49-2, 46-4 ; greatest lengthof skull 19-6 ; condylobasal length 17-3 ; condylocanine length 16-7 ; least inter-orbital width 4-6 ; zygomatic width 11-2 ; width of braincase 9-9 ; mastoid width10-6 ; c1-^1 4-7 ; m3-m3 8-2 ; c-m3 6-8 ; length of complete mandible 12-5 ; c-m37'3- SUMMARY New records of bats from the mainland and islands of south-eastern Asia includethe first reports of Rhinolophus pusillus Temminck, 1834 (probably conspecific withR. blythi Andersen, 1918, which it antedates) from Malaya and the island of Tioman,and of R. acuminatus acuminatus from Malaya. Chironax melanocephalus, Myotishorsfieldii horsfieldii and Miniopterus medius macrocneme are recorded for the firsttime from Sulawesi. Further specimens of Pipistrellus circumdatus are recordedfrom Java and the first Malayan example of this species is reported in detail. Thefirst male specimen of Philetor brachypterus verecundus is described from Malaya anda further specimen of Phoniscus atrox reported from that country. A series ofTadarida johorensis from Kelantan is reported and establishes the presence of theinteraural pocket, hitherto known only from the adult male, in female and youngexamples. REFERENCES ANDERSEN, K. 1905. On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty-six new forms. Proc. zool. Soc. Land. 75-145, i fig., 2 pis. 1906. On some bats of the genus Rhinolophus, collected by Dr W. L. Abbott in the islands of Nias and Engano. Proc. U.S. natn. Mus. 29 : 657-659. 1907. Chiropteran notes. Annali Mus. civ. Star. nat. Giacomo Doria (3), 3 (43) : 5-45. —— 1912. Catalogue of the Chiroptera in the collection of the British Museum. I. Megachirop- tera. London.- 1918. Diagnoses of new bats of the families Rhinolophidae and Megadermatidae. Ann. Mag. nat. Hist. (9), 2 : 374-384-ANTHONY, H. E. 1941. Mammals collected by the Vernay-Cutting Burma Expedition. Publs Field Mus. nat. Hist. (Zool. Ser.), 27 : 37-123, 4 pis., map.BLANFORD, W. T. 1891. The Fauna of British India, including Ceylon and Burma. Mammalia. Part II. London.CHASEN, F. N. 1940. A handlist of Malaysian mammals. Bull. Raffles Mus. No. 15 : i-xx, 11-29, map.DOBSON, G. E. 1876. Monograph of the Asiatic Chiroptera and Catalogue of the species of bats in the collection of the Indian Museum, Calcutta. London. 138 J. E. HILL DOBSON, G. E. 1877. A monograph of the group Molossi. Proc. zool. Soc. Lond. 701-735, 6 figs. 1878. Catalogue of the Chiroptera in the collection of the British Museum. London. ELLERMAN, J. R. & MORRISON-SCOTT, T. C. S. 1951. Checklist of Palaearctic and Indian Mammals, 1758-1946. ist ed., London.HILL, J. E. 1961. Fruit-bats from the Federation of Malaya. Proc. zool. Soc. Lond. 136 : 629- 642, i fig., 4 tabs. — 1966. A review of the genus Philetor (Chiroptera : Vespertilionidae). Bull. Br. Mus.nat. Hist. (Zool.), 14 : 373-387, 3 figs. 1971. The status of Vespertilio brachypterus Temminck, 1840 (Chiroptera : Vespertilioni-dae). Zool. Meded. Leiden 45 : 141-146. 1972. The Gunong Benom Expedition 1967 4. New records of Malayan bats, with taxonomic notes and the description of a new Pipistrellus. Bull. Br. Mus. nat. Hist. (Zool.), 23 : 21-42, 3 tabs. & THONGLONGYA, K. 1972. Bats from Thailand and Cambodia. Bull. Br. Mus. nat. Hist. (Zool.), 22 : 171-196, 4 figs., 2 tabs.JENTINK, F. A. 1887. Museum d'Histoire Naturelle des Pays-Bas. Catalogue Osteologique des Mammiferes. Tome IX. Leiden.- 1888. Museum d'Histoire Naturelle des Pays-Bas. Catalogue Systematique des Mammiferes (Rongeurs, Insectivores, Cheiropteres, Edentes et Marsupiaux). Tome XII. Leiden.KLOSS, C. B. 1916. On a collection of mammals from Siam. /. nat. Hist. Soc. Siam, 2 : 1-31.MEDWAY, LORD. 1965. Mammals of Borneo. Field keys and an annotated checklist. /. Malay. Brch R. Asiat. Soc. (1963), 36, 3 (No. 203) : i-xiv, 1-193, 9 figs., 34 pis., 5 tabs., map. — 1969. The wild mammals of Malaya and offshore Islands, including Singapore. KualaLumpur, Singapore, London. & YONG, G. C. 1969. Parasitic 'earwigs' (Arixenia esau) on the wrinkle-lipped bat. Malay. Nat. J. 23 : 33.OSGOOD, W. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic Expeditions. Publs Field Mus. nat. Hist. (Zool. Ser.), 18 : I93~339, 30 figs., 2 pis.TATE, G. H. H. 1942. Results of the Archbold Expeditions. No. 47. Review of the Vespertilionine bats, with special attention to genera and species of the Archbold Collec-tions. Bull. Am. Mus. nat. Hist. 80 : 221-297, 3 fig3- & ARCHBOLD, R. 1939. Results of the Archbold Expeditions. No. 24. Oriental Rhinolophus, with special reference to material from the Archbold Collections. Am. Mus. Novit. No. 1036 : 1-12, i tab.TEMMINCK, C. J. 1825. Monographies de Mammalogie. i. Paris. 1840. Monographies de Mammalogie. 2. Paris. THOMAS, O. 1923. On some small mammals, chiefly bats, from the East Indian Archipelago. Ann. Mag. nat. Hist. (9), 11 : 250-255.VAN PEENEN, P. F. D., RYAN, P. F. & LIGHT, R. H. 1969. Preliminary Identification Manual for mammals of South Vietnam. Washington. WALKER, E. P. 1964, 1965, 1968. Mammals of the World. Baltimore.WANG, S., Lu, C-K., YAO, Y-T. & Loo, T. C. 1962. On the mammals from southwestern Kwangsi, China. Acta zool. sin. 14 : 555-570, 2 pis. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.1965. (Out of Print.) £3.75. 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogyof the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227;72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : theBiology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.1974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol 884 sNU THE HAPLOCHROMIS SPECIES (PISCES : CICHLIDAE) OF LAKE RUDOLF, EAST AFRICA 7 P. H. GREENWOOD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 3 LONDON: 1974 THE HAPLOCHROMIS SPECIES(PISCES : CICHLIDAE) OF LAKE RUDOLF, EAST AFRICA BY PETER HUMPHRY GREENWOOD Pp 139-165 ; 10 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 3 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 3, of the Zoologicalseries. The abbreviated titles of periodicals citedfollow those of the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 19 July, 1974 Price £1.4° THE HAPLOCHROMIS SPECIES(PISCES : CICHLIDAE) OF LAKE RUDOLF, EAST AFRICA By P. H. GREENWOOD CONTENTS Page INTRODUCTION ........... 141 Haplochromis rudolfianus ... ...... 142 Haplochromis turkanae sp. nov. ........ 150 Haplochromis macconneli sp. nov. ....... 154 DISCUSSION ........... 161 ACKNOWLEDGEMENTS ......... 164 REFERENCES ........... 164 INTRODUCTION LAKE RUDOLF is outstanding amongst the Rift Valley Great Lakes of Africa for thepaucity of its endemic cichlid species, and particularly for the absence of a Haplo-chromis species flock (Trewavas, 1933). Whereas all other large Rift Valley lakeshave a well-defined flock of endemic Haplochromis species (even if, like Lake Albert,the flock comprises only a few species), Lake Rudolf was thought to possess but asingle Haplochromis species, the endemic H. rudolfianus Trewavas, 1933. The lake'sone positive ichthyological peculiarity, the presence of an endemic species belongingto the west African genus Pelmatochromis, has now been shown to stem from amisidentification. The Rudolf Pelmatochromis is, in fact, a specimen of the wide-spread taxon Hemichromis bimaculatus Gill (see Trewavas, 1973). Endemicity amongst the non-cichlid species is also at a low level (see Worthington& Ricardo, 1936 ; personal observations on collections recently made in the lake). Various ideas have been advanced to explain the absence of a Haplochromisspecies flock in Lake Rudolf (see summaries and comment in Fryer and lies, 1972).Of these, the most likely would seem to be the relative youth of the present lakefauna, an invasion from the Nile in post-Middle Pleistocene times. Coupled withthis factor are the shape and recent history of the lake basin, neither of which wouldprovide opportunities for the isolation (and subsequent speciation) of populationsliving in the lake. That until now the only Haplochromis known to inhabit thelake was of a structurally and ecologically generalized type similar to the fluviatilespecies of eastern Africa, would seem to agree with such a postulated lake history. Recently, however, a second and anatomically specialized species has been dis-covered in the deeper waters of the lake. A few specimens of a third species (similarto H. rudolfianus, see p. 150) have also been collected, and there are indications ofyet another taxon (see p. 149). These and other specimens, made available through the efforts of the Lake RudolfFishery Research Project, have provided sufficient material to describe the newdeep-water taxon, the new Haplochromis rudolfianus-like fish, and to redescribe in i* i42 P. H. GREENWOOD greater detail H. rudolfianm itself. Unfortunately the putative fourth species isrepresented by so few and distorted specimens that I consider it inadvisable todescribe it at present. Nevertheless, sufficient information has been gathered fromthese few specimens to show that this species too is like H. rudolfianus. As part of a planned review of phyletic relationships within the Haplochromis-group cichlids, some consideration is also given to the relationships of the LakeRudolf species with those of other lakes, especially Lakes Albert and Victoria. Haplochromis rudolfianus Trewavas, 1933 (Text-figs. 1-4)Haplochromis rudolfianus Trewavas, 1933, /. Linn. Soc. (Zool.), 38 : 321-322. LECTOTYPE. A specimen, 51-0 mm standard length (BMNH reg. no. 1933.2.23 :163), from a weedy lagoon on the east shore of Lake Rudolf, near Mt El Moitat(station number 285, see Worthington, 1932). The specimen has been eviscerated,but judging from its preserved coloration it is probably an adult male. PARALECTOTYPES. Three specimens (BMNH reg. nos. 1933.2.23:164-166),39-0-42-5 mm S.L., from the same locality as the lectotype (all are eviscerated),and one other specimen (BMNH reg. no. 1933.2.23 : 167), 45-0 mm S.L. fromCentral Island (Worthington's station no. 264). This latter fish is also evisceratedbut is probably an adult male. Comment on the original description of H. rudolfianus Trewavas' (1933) original description was based on 5 syntypes, but 25 otherspecimens were also examined although not included in the description. I havere-examined these fishes and would confirm their identification as H. rudolfianus.Dr Trewavas also included in this species, but with certain reservation, a largerspecimen, 61-0 mm S.L. and 80 mm total length (BMNH reg. no. 1933.2.23 : 169).The locality label for this fish reads '? Lake Rudolf, the uncertainty stemming fromthe collector's notes on the provenance of the specimen. A number of differencesbetween this fish and other specimens of H. rudolfianus were noted by Trewavas. Ican confirm these differences and would add others. The size discrepancy onceexisting between this specimen and others of H. rudolfianus is virtually obliteratedby the larger specimens of the latter species now in the Museum's collections. Thusit seems very unlikely that the various differences listed by Trewavas are, as shethen suggested, size correlated ones. The specimen in question departs from H. rudolfianus in having a deeper body(37'4 Per cent of standard length), somewhat deeper preorbital (17-7 per cent ofhead length), wider interorbital distance (25-6 per cent of head), a deeper cheek(25-6 per cent of head), a smaller eye diameter (30-0 per cent of head) and a markedlyshorter caudal peduncle (14-7 per cent of standard length, cf. 16-5-19-7, mean18-0 per cent in H. rudolfianus). The fish also differs from specimens of H. rudolfianusin having fewer gill rakers (7 cf. 8 or 9), larger scales on the nape and in the lateralline series (30), in lacking the characteristic dark vertical bars on the body and caudal LAKE RUDOLF HAPLOCHROMIS SPECIES 143 peduncle (see p. 147), in having no melanic pigment in the ovarian walls (see p. 146)and in having stouter and somewhat obliquely cuspidate outer jaw teeth. The lowerpharyngeal bone is finer than in H. rudolfianus, and the median lower pharyngealteeth are also finer (see p. 146). In my opinion, this specimen cannot be identified as a member of H. rudolfianus,nor can it be placed in either of the other Haplochromis species from Lake Rudolf.All the evidence certainly suggests that it in fact belongs to the fauna of anotherlake. On this assumption, the specimen was compared with Haplochromis speciesfrom all the lakes sampled by the Cambridge University Expedition of 1930-31. Itcannot be identified with any known Lake Albert Haplochromis species, nor does itclosely resemble any of the described (or known but undescribed) species of LakesEdward and George ; see Greenwood, 1973. It does, however, closely agree in allmorphometric and anatomical features (especially the dentition) with H. veliferTrewavas of Lake Nabugabo, Uganda (see Greenwood, 1965). Since extensivecollections were made in this lake by the Cambridge Expedition, I would identifythe fish as a specimen of H. velifer and suggest that the locality label be altered toread 'Lake Nabugabo'. The redescription of H. rudolfianus given below is based on the 5 type specimens(39'0-5i-o mm S.L.) and 20 additional fishes (30-5-58-0 mm S.L.) collected in 1972by the Lake Rudolf Fisheries Research Project team at Topi point, Allia Bay, andFerguson's Gulf. Coloration and certain anatomical details were also checked onthe other specimens from which meristic and morphometric data were not taken(BMNH reg. nos. 1973.11.13 : 151-170). All counts and measurements used in this description are those defined by Green-wood (1973). Dorsal head profile straight or gently curved, sloping at an angle of 25-30 degreeswith the horizontal; premaxillary pedicels rarely breaking the smooth outline of theprofile. Length of head 31-0-35-3 (mean, M = 33-5) per cent of standard length, depthof body 29-4-33-7 (M = 32-1) per cent. Preorbital depth 12-5-17-9 (M = 15-0) per cent of head, least interorbital width20-0-27-4 (M = 22-7) per cent, neither dimension showing allometry with standardlength. Length of snout 26-6-32-3 (M = 29-9) per cent of head, 0-7-1-1 (mode 0-8)times its breadth. Eye diameter 30-5-35-7 (M = 32-9) per cent of head (showingill-defined negative allometry), depth of cheek 17-4-25-8 (M = 21-7) per cent. Caudal peduncle 16-5-19-7 (M = 18-0) per cent of standard length, 1-2-1-7(modal range 1-3-1-5) times as long as deep. Mouth horizontal or slightly oblique. Length of upper jaw 30-8-38-0 (M = 33-7)per cent of head, length of lower jaw 35-7-48-3 (M = 39-7) per cent, 1-4-1-8 (modalrange 1-5-1-7) times as long as broad. Posterior tip of maxilla reaching a verticalthrough the anterior margin of the eye. Gill rakers. Relatively stout, with the lower i or 2 on the first gill arch reduced ;9 (less frequently 8, rarely 10) on the lower limb of that arch. Well-developed'pseudorakers' (fleshy protuberances between the inner and outer row of true rakers)are present on the first arch. P. H. GREENWOOD 5mm FIG. i. Haplochromis rudolfianus. Lectotype. Drawn by Gordon Howes. Scales. Strongly ctenoid, those on the anteroventral aspects of the thoracicregion very small, with an abrupt size transition between them and the posteriorscales of this region. Immediately anterior to the first dorsal fin spine there is asmall naked area (about i| or 2 scales in area), and often there is also a narrow nakedstrip below the ventral horizontal row of cheek scales. Lateral line with 30 (f2),31 (fio), 32 (fi2), or 33 (fi) scales, cheek with 3 (rarely 2 or 4) rows. Six or 7 scalesbetween the dorsal fin origin and the upper lateral line, 6 or 7 (rarely 5) between thepectoral and pelvic fin bases. Fins. Dorsal with 24 (fi2), 25 (fn) or 26 (f2) rays, comprising 14 (f3), 15 (121)or 16 (fi) spinous and 9 (19), 10 (fi5) or n (fi) branched rays. Anal with 3 spinousand 8 (13), 9 (f2o), 10 (fi) or n (fi) branched rays. First branched ray of the pelvic fin very slightly produced in both sexes, but rela-tively more so in adult males. Caudal subtruncate, scaled on its basal quarter to third. Teeth. The majority of teeth in the outer row of both jaws are unequally bicuspidand moderately stout (Text-fig. 2) ; the major cusp is acutely pointed and equilateralin outline. A few unicuspid teeth sometimes occur posteriorly in the upper jaw, butmore often the posterior teeth are tricuspid ; rarely are these teeth bicuspid. Anoteworthy feature of the samples examined is the pronounced wear pattern seenon the outer teeth. Worn teeth have the major cusp either obliquely truncate orthe wear may be so great that all demarcation between major and minor cusps hasdisappeared and the crown is spatulate. There are 28-48 teeth in the outer premaxillary row, the number showing somepositive correlation with the fish's length. All inner row teeth are small tricuspids, and are arranged in 2 or 3 (mode), rarely 4,rows in the upper jaw and in 2 (mode) or 3 rows, rarely a single row, in the lower jaw. LAKE RUDOLF HAPLOCHROMIS SPECIES 145 0.5 mm FIG. 2. Haplochromis rudolfianus. Outer row jaw teeth (in labial view) from leftdentary, anterolateral in position. OSTEOLOGY. The neurocranium (Text-fig. 3A) of H. rudolfianus is of the general-ized 'H. bloyeti'-type (see Greenwood, 1974), with a moderately decurved profileand a relatively short preotic skull length (ca 62-5 per cent of total neurocraniallength). None of the cephalic laterosensory canals or pores, nor any of the bonescarrying the canals, is at all hypertrophied (cf. p. 155 ; Text-fig. 36). 3 mm FIG. 3. Haplochromis rudolfianus. A : Neurocranium, left lateral view.B : Bones in the infraorbital series of the right side. The lower pharyngeal bone (Text-fig. 4) is triangular in outline, with the denti-gerous area a little broader than long (ca i-i times). In all specimens examinedthe bone is noticeably stout, especially compared with that of the other Rudolfspecies or that of the generalized species in other lakes and in the east African rivers.The degree of enlargement, however, varies between individuals. i** 146 P. H. GREENWOOD The two median tooth rows are composed of teeth clearly stouter than their lateralcongeners ; the degree of enlargement, like that of the bone itself, shows considerableindividual variability. All five specimens in the type series have the stoutest bonesand dentition of all the specimens examined ; the lectotype (51 mm S.L.) is excep-tional even amongst the type series in having submolariform teeth posteriorly inthe median rows (in all other specimens the bicuspid crown is still retained). Thelectotype also has the relatively most massive lower pharyngeal bone (Text-figs.4A and B). FIG. 4. Haplochromis rudolfianus. Lower pharyngeal bones to show variation in size ofmedian teeth. A and B : Bone from one of the paratypes, in occlusal and left lateralviews respectively. C and D : Bone from another specimen of the same size, in occlusaland left lateral views respectively. Lower pharyngeal bone and tooth enlargement does not seem to be size correlatedbecause some of the syntypes are amongst the smaller fishes examined, and a speci-men 7 mm longer than the lectotype has a less massive bone and dentition. Vertebral counts in the 30 specimens radiographed are : 27 (fi), 28 (13), 29 (fi8)or 30 (f8), comprising n (fi), 12 (14), 13 (123) or 14 (f2) abdominal and 16 (123) or17 (17) caudal vertebrae. (The fused first ural and preural centra are not includedin these figures.) Only two specimens show any sign of fusion or close apposition between elementsin the hypural series (cf. p. 158) ; in these two fishes the first and second hypuralsseem to be fused. VISCERA. The intestine is from i£ to i| times the standard length ; the ovariesare of unequal size with the right ovary noticeably larger than the left in mostspecimens or, rarely, it alone shows signs of oogenesis. A very characteristic featureis the intensely melanic tissue of the ovary wall; the testes, however, are but slightlypigmented. The entire peritoneum is also densely melanic. This extreme visceralmelanism is probably correlated with the intense sunlight to which fishes living inthe shallows of Lake Rudolf would be subjected. LAKE RUDOLF HAPLOCHROMIS SPECIES 147 COLORATION. In freshly killed specimens of adult male H. rudolftanus the groundcolour is greenish-yellow (but whitish on the chest). Dark vertical bars cross theflank and caudal peduncle (see notes on preserved colours below). Each scale onthe flanks has an opalescent centre in which yellow, blue and green colours can bedetected ; yellowish pigment predominates on the anterior body scales, especiallythose around the base of each pectoral fin. Posteriorly on the body the dominantcolour in the scale centres is electric blue, and is especially noticeable around thebase of the anal fin. The head, like the anterior part of the body, is an opalescentgreenish-yellow and blue, the operculum marked with a golden yellow area on itslower part. The branchiostegal membrane is a delicate pale yellow except for aclearly demarcated black area anteriorly. The dorsal fin membrane is dark golden-yellow, the lappets are scarlet and thesoft part of the fin has a bright yellow basal streak and light yellow dots dorsally,the intervening areas melanic. Black spots also occur on the spinous part of the fin.Two dark basal blotches occur on the spinous dorsal; the first lies between the sixthto ninth spines, the second between the last two or three spines. The anal fin isyellow but with scattered black and red chromatophores, and with two or three lightgolden-yellow ocelli. The caudal is a marbled yellowish-green, with red and blackspots, the red colour being most intense in the dorsal and ventral angles of the distalmargin. The pelvic fins are dusky. Adult females have a similar but more subdued coloration, with the whitish ventralareas more extensive and without the black anterior region to the branchiostegalmembrane. The pelvic fins are hyaline tinged with yellow, and the red lappets andmargin to the dorsal and anal fins respectively are barely discernible. On the analfin there are pale yellow spots in the same position as the ocelli of males. I am indebted to Mr and Mrs Hopson for supplying the notes on which this descrip-tion is based. Preserved coloration is virtually identical in both sexes, except that the pelvics infemales are clear and not dusky, and the ground coloration is lighter than in males. The ground coloration is a pale brown (fawn) shading to greyish-charcoal on thechest and belly. The flanks and caudal peduncle are crossed by 5-7 (rarely 8-10)dark and clearly defined vertical bars ; the bars on the flanks reach almost to theventral profile, but those on the caudal peduncle rarely extend to below the level ofthe midlateral line where they merge with a short and faint horizontal bar extendingthe length of the peduncle. This bar is of variable intensity and is barely visible insome specimens. Anteriorly there is a broad, sometimes ill-defined dark bar over-lying the cleithrum and following the outline of that bone ; dorsally the bar joins adark, saddle-shaped blotch on the nape. A fainter bar is sometimes visible along thevertical limb of the preoperculum. The snout is crossed by a pair of parallel darkbars, and there is an intense and clearly demarcated lachrymal stripe. All fins are a greyish-hyaline, the soft dorsal and the entire caudal are maculate,the dorsal with two distinct dark blotches basally (see above), and with dark lappets.Spotting on the caudal fin is most distinct proximally, the spots often arranged so asto produce two to four dark vertical bands on that part of the fin. The pelvics aredusky in males, greyish in females. 148 P. H. GREENWOOD ECOLOGY. Haplochromis rudolfianus is apparently confined to the shallow andprotected inshore areas of the lake, although there is a population inhabiting a craterlake on Central Island (see below}. No data are yet available on the feeding habitsof this species, nor whether it shows any particular substrate preferences. Males appear to reach a larger size than do females, and are adult at a standardlength of 45-47 mm. Females, however, mature at a smaller size, namely ca 30 mm.No data are available on the breeding habits or seasons of the species. THE CENTRAL ISLAND POPULATION. One of the paralectotypes (see p. 142) and 12additional specimens collected in 1965 by Dr R. L. Welcomme are from CentralIsland. The paralectotype is apparently from the shore of the island (stationno. 264, see Worthington, 1932) but the other fishes are from one of the crater lakesin that island (see Beadle, 1932, for details of the lakes). The Central Island fishes, particularly those from the crater itself, are of consider-able interest since they are apparently isolated from other populations inhabitingthe mainland shores of the lake. The apparent absence of H. rudolfianus from open-water localities suggests that it does not leave (or at least not frequently) its shallow-water inshore habitats. A population of Sarotherodon living in Crater Lake A of Central Island is sufficientlydistinctive for it to have been referred to a new species (Tilapia vulcani, Trewavas,1933, see also Trewavas, 1973, for distinguishing features of the genera Tilapia andSarotherodon). A population from the neighbouring Crater Lake C, however, did notdiffer significantly from the populations of Sarotherodon niloticus living in the mainlake (Trewavas, 1933). The explanation for this seemingly anomalous situationappears to be that Lake C is, on occasion, connected with the main lake. (Personalcommunication from Dr K. E. Banister, based on information he was given bymembers of the Fishery Research Team on Lake Rudolf ; the last interconnectionwas in 1972.) Lake A, on the other hand, is completely isolated from the mainlake by a crater wall at least 10 m above current lake level. It is the more regrettable then that the H. rudolfianus collected in 1965 bear nomore precise locality data than 'Crater Lake, Central Island', and that all are distortedand poorly preserved. Allowing for the difficulty of measuring distorted specimens,I can find no meristic or morphometric differences between the Central Island fishesand those from the main lake. Several of the Island fishes are, however, larger thanany recorded from the lake (maximum size 72 mm, cf. 58 mm for lake fishes). Thelargest Island fish has four or five unicuspid teeth situated posterolaterally in theupper jaw, but this could well be a size-correlated phenomenon (see Greenwood,1974). Most of the Island fishes are much darker than are the lake fishes andconsequently the vertical barring on the body is less obvious ; in a few paler indi-viduals, however, the bars are quite distinct. Like their main lake congeners,female Crater Lake fishes have only the right ovary well developed, but unlike thelatter populations the ovarian wall in these fishes is but faintly melanized and thenonly on the dorsal and lateral aspects. These observations were paralleled by thoseI was able to make on specimens and colour photographs obtained by Dr Banisterduring his recent visit to the lake. Once again, the specimens are not well preservedand add little anatomico-morphological information to that already available. LAKE RUDOLF HAPLOCHROMIS SPECIES 149 However, the similarity in coloration between Dr Banister's fishes from Crater LakeA and those from the Welcomme sample strongly indicate that the latter are fromthe same source. The dark coloration of Central Island H. rudolfianus seems to parallel that of'Tilapia vulcani' from Crater Lake A on the island (see Trewavas, 1933). In othercharacteristics the parallelism is not clearly apparent. That is to say, the H.rudolfianus specimens do not show the leanness, larger eyes, larger heads, longerdorsal fin spines and broader bands of inner teeth that Trewavas noted in theTilapia specimens (Trewavas, 1933). In contrast to the Central Island Crater Lake A specimens of H. rudolfianus, theparalectotype from the shore of the island does not show any marked darkening ofthe ground coloration. Unfortunately, it is eviscerated so no check can be made onthe melanization of its gonads. The peritoneum is very dark. Without more precise locality data, more material from different localities on theisland and more details on live coloration, little can be said about the taxonomicstatus of Central Island populations of H. rudolfianus. It does, however, seemprobable that, like the Crater Lake A population of Sarotherodon niloticus (with whichspecies 'T. vulcani' should now be synonymized ; Dr Trewavas, personal com-munication) , the Crater Lake A Haplochromis show some ecophenotypic response totheir peculiar environment. The relative melanism shown by Crate Lake A Haplo-chromis rudolfianus populations is of particular interest because male colorationseems to be an important species recognition character (see Greenwood, 1974).Altered male coloration apparently is one of the first morphological differences seenin recently speciated Haplochromis (Greenwood, 1965). DIAGNOSIS AND AFFINITIES OF H. rudolfianus. The distinctively barred colourpattern of this species immediately serves to distinguish it from all other Haplo-chromis species in Lake Rudolf (see footnote, below). From H. macconneli describedon p. 154, H. rudolfianus is further distinguished by the absence of hypertrophiedlaterosensory canals and pores on the head, by several morphometric characters (seep. 155) and in having few, if any tricuspid teeth in the outer series of either jaw. Inaddition to the hypertrophied canal system, the neurocranium in H. macconneli is ofa more derived type than that of H. rudolfianus. The nearest living relatives of H.rudolfianus are probably H. turkanae (see p. 153), and the fourth but as yet undescribedHaplochromis species in the lake.1 The validity of these relationships can only be tested when more material of theputative relatives is available. 1 Four specimens 37-0-64-0 mm standard length (and a fifth prepared as a skeleton) may representa fourth Haplochromis species. Morphometrically these fishes are not distinguishable from H. rudolfianus,and the dentition is similar except for there being only a single row of inner teeth in each jaw. Theprincipal 'interspecific' difference seems to be in the coloration, both that observed by Mr Hopson (inlitt.) when the fishes were alive and that remaining in the preserved material. There is also an observable(but non-quantifiable) difference in head shape, and these fishes have a colourless (not black) peritoneum.The gonads show no sign of melanization, unlike those of H. rudolfianus. All five specimens are from water between 15 and 35 m deep; that is, from somewhat greater depthsthan H. rudolfianus, but within the range of H. turkanae, from which species they differ in the samemorphometric characters as does H. rudolfianus. Since only five specimens are available and because all are in some way damaged or distorted, I wouldconsider it inadvisable to describe a new taxon on this material. P. H. GREENWOOD 10mm FIG. 5. Haplochromis turkanae. Holotype. Drawn by Gordon Howes. In its gross morphology and in the details of its pectoral and predorsal squamation,H. rudolfianus resembles H. albertianus Regan, a species endemic to Lake Albert(see Trewavas, 1938). The two species differ in their preserved colour patterns, andin adult H. albertianus (of the same size as H. rudolfianus) having unicuspid outer jawteeth and a more massive lower pharyngeal bone with a greater number of enlarged,submolariform teeth. In other words, H. albertianus shows a greater degree ofspecialization in those characters that are already somewhat specialized in H.rudolfianus. STUDY MATERIAL Register number BMNH1973.11.13 : 151-154 1973.11.13 : 155-17° Locality : Lake Rudolf Ferguson's Spit (shallow water, inshore)Topi Point, Allia Bay Haplochromis turkanae sp. nov.(Text-figs. 5 and 6) HOLOTYPE. An adult male, 73-0 mm standard length (BMNHreg.no. 1973.11.20:1),caught in a bottom trawl fished at a depth of 16 m, over a mud and rock bottom,5-6 km north-west of Porr. PARATYPES. Three adult males (BMNH reg. nos. 1973.11.20 : 2-4), 77-5-86.0 mmS.L. from the same locality and trawl as the holotype. 33-5 20-5 20-5 30-6 28-6 28-6 45-0 41-0 17-5 32-3 18-0 22-0 32-0 30-0 32-0 44-0 40-0 17-4 32-7 18-9 20-8 34'° 34*0 26-4 45'3 41-5 18-5 3I-4 18-5 22-O 29-6 27-9 27-0 44'5 40-8 16-9 LAKE RUDOLF HAPLOCHROMIS SPECIES 151 DESCRIPTION. Based on these four fishes, 73-0-86-0 mm S.L. With so fewspecimens available most of the morphometric data can be presented most con-veniently in tabular form. S.L. Depth* Head* PO % IO % Snt % Eye % Cheek % Lj % Uj % CP* 77'5 35-o81-0 34-586-0 33-2 * = per cent of standard length. % =per cent of head length. PO = preorbital depth; IO = least interorbital width; Snt = snout length; Lj= lower jaw, and Uj:upper jaw length. CP = caudal peduncle length. Caudal peduncle 1-4-1-5 times longer than deep. Dorsal profile of head straightor gently curved dorsally, but straight anteriorly, sloping at an angle of 30-35degrees with the horizontal. The ascending processes of the premaxillae barelyinterrupt the outline of the profile. Mouth slightly (but noticeably) oblique, the lips a little thickened ; posterior tipof the maxilla reaching a vertical through the anterior part of the eye. Jaws equalanteriorly, the lower 1-6-1-7 times longer than broad. Snout broader than long,its anterior profile, when viewed from above, smoothly rounded. None of thecephalic laterosensory canals (or their pores) is noticeably enlarged. Gill rakers. Relatively stout, 8 (13) or 9 (fi) on the lower part of the first arch,the lowermost one or two rakers reduced in size. Pseudorakers (see p. 143) aredeveloped between the inner and outer rows of gill rakers, but are not conspicuous. Scales. Ctenoid ; lateral line with 31 (13) or 32 (fi) scales, cheek with 4 (fi) or5 (13) imbricating rows. Six or 7 scales between the upper lateral line and thedorsal fin origin ; a naked area (about i| scales in extent) immediately before thefirst dorsal fin spine. Nine or 10 scales between the pectoral and pelvic fin bases ;the scales of the chest are also very small and grade abruptly with the larger scalesof the post-pectoral region. Fins. Dorsal with 14 (fa) or 15 (f2) spinous rays and 9 (14) branched rays, analwith 3 spines and 7 (12) or 8 (12) branched rays. The first pelvic ray is produced,very noticeably so in two specimens where it extends as far as the second branchedanal ray. The caudal fin is strongly subtruncate, almost rounded, and scaled on itsproximal quarter to half. Pectoral fin 25-6-28-7 per cent of standard length,81-5-88-0 per cent of head length. Teeth. In all four specimens the posterior three or four teeth in the outer pre-maxillary row are strong, dagger-like unicuspids. In two fishes the other teeth inthis row are a mixture of caniniform unicuspids and weakly bicuspids, while in theother two specimens distinctly bicuspid teeth predominate, although a few uni-cuspids occur anterolaterally. There are 42-52 teeth in this row. There is a similardifference in the predominant tooth type of the outer row in the lower jaw. Thefirst two specimens have mainly unicuspid teeth with a few bicuspids, the second 152 P. H. GREENWOOD FIG. 6. Haplochromis turkanae. Lower pharyngeal bone of holotype inA : occlusal view ; B : left lateral view. pair have a predominance of bicuspids. Bicuspid teeth in the lower jaw have theircrowns more distinctly incurved than do the upper jaw teeth. Bicuspid teeth in both jaws are of the typical generalized Haplochromis type, andclosely resemble the teeth of H. rudolfianus (see p. 144 ; Text-fig. 2). The inner series of teeth in both jaws are composed of small tricuspids arranged in3 rows in the upper and 2 (or irregularly 2) rows in the lower jaw. OSTEOLOGY. With only four specimens available, no skeletal preparations werepossible. Vertebral counts made from radiographs are: 28 (13) and 29 (fi),comprising 13 abdominal and 15 or 16 caudal vertebrae (the fused first preural andural centra not included). No specimen has any fused elements in the caudal finskeleton, but in two fishes hypurals I and 2 are closely apposed. COLORATION. No information is available on the live colours of this species. Thefour adult (but not sexually active) male specimens (fixed in formalin) have a lightgrey to yellowish-grey ground colour that extends ventrally to a clearly demarcatedhorizontal line on the body and caudal peduncle, at which level it becomes pearlywhite. The line is at about the horizontal level of the lowermost insertion of thepelvic fin ; thus in lateral view little of the caudal peduncle shows the white ventralcoloration. On the flanks, most dorsal, lateral and ventrolateral scales have anarrow margin of dark pigment. The lower jaw, branchiostegal membrane, sub-operculum and the lower part of the cheek are also pearly-white. About five very faint vertical dark bars are visible on the flanks, and another mayalso occur posteriorly on the caudal peduncle ; these bars are narrower dorsallythan ventrally, and do not extend onto the white ventral coloration of the body.The vertical limb of the preoperculum is faintly to clearly dusky and there is a broad,intensely black lachrymal bar that, at about the level of the posterior maxillarytip, narrows abruptly and then continues ventrally and a little medially onto thelower jaw. LAKE RUDOLF HAPLOCHROMIS SPECIES 153 The entire soft dorsal fin and the posterior half of the spinous dorsal are denselyand distinctly spotted, with the spots arranged in from four horizontal rows anteriorlyto six or more rows posteriorly on the fin. On the anterior part of the spinous dorsalthe spots are confluent and form vertically aligned, dark, interspinous streaks.The lappets of the dorsal fin are black. The entire caudal fin is covered with darkand discrete spots so arranged as to form wavy, vertical bars when the fin is notfully opened. The anal is hyaline except for three rather pale ocelli. The pel vieshave the anterior half dusky, the posterior half hyaline ; the elongate first pelvicray is dead-white. ECOLOGY. Virtually nothing is known about the biology of this species. Thefour specimens came from deeper water than is usual for H. rudolfianus, and shallowerwater than is usual for H. macconneli (see pp. 148 and 159). All four specimens have the stomach and intestine packed with fragments ofostracod shells. It is interesting to note that, unlike H. rudolfianus but like H. macconneli, there isno trace of dark pigment in the peritoneum and neither is there any on the gonads(cf. p. 146). All four specimens are adult males, but judging from the size and shape of thetestes, none is sexually active. DIAGNOSIS AND AFFINITIES. Haplochromis turkanae is immediately distinguish-able from H. macconneli because of its non-hypertrophied cephalic laterosensorycanal system. There are also morphometric differences between the species,differences in their dentition and, apparently, in the preserved coloration of adultmales (see below, pp. 156-159). In its general appearance, its dentition and in several morphometric characters,H. turkanae closely resembles H. rudolfianus. It differs principally in having morerows of scales on the cheek (4 or 5, cf. 3), smaller scales between the pectoral andpelvic fin insertions (9 or 10, cf. 6 or 7), a deeper cheek (26-4-32-0 per cent of head,cf. 17-4-25-8, M = 21-7 per cent), longer upper jaw (40-8-41-5, cf. 30-8-38-0, M = 33-7percent of head), and a somewhat longer lower jaw (44-0-45-3, cf. 35-7-48-3, M = 39-7per cent of head). Haplochromis turkanae also differs in having a more elongatefirst pelvic ray (which is distinctively coloured) and in its overall coloration. Forexample, it lacks the prominent vertical bars on the body, has more intensely anddensely maculate dorsal and caudal fins, and has the peculiar dribble-like extensionof the lachrymal bar onto the lower jaw. The lower pharyngeal bone and dentitionof H. turkanae are somewhat more massive than those of most H. rudolfianus, butare quite comparable with those in the type series of that species (see above, p. 146,and cf. figs. 4 and 6). Without more osteological information on H. turkanae it is difficult to say muchabout its affinities with H. rudolfianus. From what is known, however, the twospecies would seem to be very closely related phyletically. Indeed, the resemblanceparallels that seen between many pairs of Haplochromis species in Lake Victoria.As in many of the Victoria pairs there is also an apparent ecological replacementinvolved, with H. rudolfianus being the inshore, shallow-water species and H.turkanae the species of deeper waters (see Greenwood, 1974). 154 P. H. GREENWOOD 10mm FIG. 7. Haplochromis macconneli. Holotype. Drawn by Gordon Howes. Haplochromis macconneli sp. nov.(Text-figs. 7-10) This most distinctive species was discovered by Mr and Mrs A. J. Hopson when theLake Rudolf Research Project team began trawling operations in the deeper watersof the lake. Apparently it is confined to water more than 20 m deep and has beencollected from depths down to 75 m. Besides its various anatomical peculiarities, H. macconneli is also noteworthy forits marked sexual dimorphism in adult size. No males larger than 35-5 mm standardlength have yet been recorded. Indeed, despite intensive efforts, only two maleshave so far been recognized amongst the several hundreds of specimens examined.Females, on the other hand, are common components of deep-water trawl catches. The taxon is named for Mr R. B. McConnel, Officer in Charge of the FisheriesDepartment at Lake Rudolf, in grateful recognition of the assistance he has un-stintingly given to Mr Hopson and his research team. HOLOTYPE. A female, 77-0 mm standard length (BMNH reg. no. 1973.11.13 : 37),caught near the bottom in 50-64 m of water 3 miles north-west of Central Island. PARATYPES. (i) Twenty-one specimens (BMNH reg. nos. 1973.11.13 : 38-58),51-0-72-0 mm S.L., all females and from the same locality as the holotype ; (ii) 6juveniles (of indeterminable sex), 22-0-38-0 mm S.L., caught in a bottom trawl at adepth of 20 m 1-6 km off Ferguson's Spit (BMNH reg. nos. 1973.11.13 : 59-64 ;(iii) 2 adult males, 31-0 and 35-5 mm S.L., caught in a bottom trawl at a depth of30 m, 2-4 km east of North Island (BMNH reg. nos. 1973.11.13 : 65-66). DESCRIPTION. Based on the holotype and 29 paratypes, 22-0-77-0 mm S.L.Various characters have been checked on the 78 additional specimens (size range15-0-59-0 mm S.L.) from the Ferguson's Spit station and one other station 3-2 km LAKE RUDOLF HAPLOCHROMIS SPECIES 155 off-shore at a depth of 75 m. None of this extra material has, however, been in-cluded in the morphometric counts and measurements. Because most of the proportional measurements used in this description showallometry with the fish's size, the material has been divided into two size groups,one of fishes 51-77 mm S.L. and the other of fishes 22-38 mm S.L. Ranges andmeans for the larger specimens (N = 22) are given first, followed in parenthesesby those for the smaller fishes (N = 8). Length of head 33'3-39'9, M = 35-3 (31-6-39-4, M = 36-0) per cent of standardlength, depth of body 317-35-1. M = 33'3 (25-3-32-3. M = 297) Per cent- Dorsal profile of head clearly but gently concave above the eye, sloping at an angleof ca 35-40 degrees to the horizontal. The profile of the snout varies from straightto somewhat decurved, with the premaxillary pedicels always breaking the outline.The anterior tip of the lower jaw is usually produced into a symphysial knob, whichis most obvious in fishes over 65 mm S.L. An outstanding feature of the head (in specimens of all sizes) is the greatly enlargedopenings to all the laterosensory canals (Text-figs. 7 and 9). Particularly obviousare those on the preorbital and preopercular bones. The underlying canals arehypertrophied, with the result that the bones involved have a distinctly inflatedappearance. Laterosensory canals on the neurocrariium, especially the temporal canal of thepterotic bone, are also inflated, particularly in comparison with those in H. rudol-fianus and H. turkanae, and indeed with those of the generality of Haplochromisspecies (see below, p. 158). Canals, and their openings, in the extrascapular andsupracleithrum are equally affected by this trend, as are the nasals although thelatter are relatively the least cavernous of the cephalic laterosensory canal bones.In well-preserved specimens the canal openings are occluded by a thin mem-brane. Depth of preorbital 16-6-22-2, M = 20-1 (12-5-16-7, M = 15-4) per cent of head,least interorbital width 18-8-24-2, M = 21-8 (14-7-25-0, M = 21-2) per cent. Snoutslightly broader than long (rarely as long as broad) in specimens of all sizes, its length27-8-38-1, M = 33-8 (26-1-33-3, M = 29-6) per cent of head for the size groupsrespectively. Diameter of eye 23-7-29-3, M = 26-8 (25-0-33-3, M = 28-7) and depthof cheek 22-8-29-3, M = 26-7 (i8-i-25'-o, M = 21-2) per cent of head. Caudal peduncle 16-5-21-5, M = 18-6 (17-8-22-6, M = 20-0) per cent of standardlength, 1-5-2-2 times as long as deep in fishes of all sizes. Lower jaw with a distinct symphysial knob in fishes of all sizes, the protuberancemost marked in individuals of > 65 mm S.L. Length of lower jaw 40-8-51-0,M = 45-6 (37-6-46-2, M = 42-0) per cent of head, 1-6-2-0 (modal range 1-8-2-0)times longer than broad in specimens of all sizes. Length of upper jaw 31-8-44-0,M = 38-4 (30-6-37-6, M = 33-3) per cent of head. Mouth slightly oblique, the posterior tip of the maxilla reaching a vertical throughthe anterior margin of the orbit, or to a point slightly posterior to that line. Gill rakers. Of variable form, but usually with the upper 2 or 3 rakers on the firstgill arch flattened and branched, those on the middle section of the arch relativelyslender, and the lowermost I or 2 rakers reduced ; no pseudorakers are developed 156 P. H. GREENWOOD FIG. 8. Haplochromis macconneli. Dentition. A: Outer teeth from the premaxilla.B : Outer teeth from the dentary. In both, the teeth are from an anterolateral positionin the jaw and are viewed labially. (see p. 143). There are 7 (fi), 8 (f8) or 9 (121) rakers on the lower limb of the first gillarch. Scales. Ctenoid, those on the chest very small but grading in size with those onthe subpectoral region (that is, the demarcation between small and large scales inthis region is less abrupt than in H. rudolfianus ; see p. 154). Five and a half to 7 (mode 6) scales between the dorsal fin origin and the lateralline, 7-9 (modes 8 and 9), rarely 5, between the pectoral and pelvic fin bases. Cheekwith 3 or 4 rows of imbricating scales. In most specimens there is a small naked area(about the area of one scale) in front of the first dorsal fin spine. Fins. Dorsal with 22 (f6), 23 (fi6) or 24 (f8) rays, comprising 13 (12), 14 (fi6),15 (fn) or 16 (fi) spinous and 8 (fn), 9 (fi8) or 10 (fi) branched rays. Anal with3 spines and 7 (f2), 8 (£25) or 9 (13) branched rays. First ray of the pelvic fin slightlyproduced, more so in larger fishes. Caudal truncate, scaled on its proximal quarter(mode) to third. Pectoral fin 25-8-34-5, M = 28-6 (23-7-28-3, M = 26-1) per centof standard length, and 73-0-92-5, M = 81-1 (63-5-77-0, M = 72-3) per cent of head. Teeth. In fishes more than 50 mm S.L. the outer row of premaxillary teeth iscomposed of unequally bicuspid teeth anteriorly, but of tricuspids laterally andposteriorly (Text-fig. 8) ; the median cusp of the tricuspids, and the major cusp of thebicuspids is slightly incurved. Many specimens have some tricuspids intercalatedamongst the biscuspids anteriorly, and in a few fishes almost the entire outer row iscomposed of tricuspid teeth. When there is a mixture of bi- and tricuspids the latterpredominate. At all positions in the tooth row the bi»- and tricuspids are of equalsize. Tooth morphology and arrangement in the lower jaw are similar to those in theupper, although more individuals have only tricuspid teeth present; a predominantlybicuspid outer row is rarely encountered. Fishes in the size range 20-40 mm S.L. have mainly bicuspid teeth in the upperjaw, with those tricuspids present restricted to a posterolateral position in the row. LAKE RUDOLF HAPLOCHROMIS SPECIES 157 B FIG. g. Haplochromis macconneli. A : Neurocranium in right lateral view. B : Neuro-cranium, dorsal surface. C : Bones of the infraorbital series, right side. D : Right lowerjaw, viewed from a slightly ventrolateral position. Bicuspid teeth also predominate in the lower jaw, but some unicuspid (and slender)teeth may occur posterolaterally. Tooth form is like that of the larger fishes. There are 41-64 (M = 52) outer premaxillary teeth in fishes 50-77 mm S.L., and24-42 in specimens 22-39 mm long. Irrespective of a fish's size, the inner teeth in both jaws are all tricuspid and small,and are usually arranged in a single series but double rows are encountered occasion-ally. OSTEOLOGY. The characteristic hypertrophy in the cephalic laterosensory canalsystem has been commented upon above (see also Text-figs. 7 and 9). Canal bonesin the pectoral skeleton are also affected, and the otic region of the skull is noticeablyinflated (see Text-fig. 9). The neurocranium (Text-fig. 9) departs from the generalized Haplochromis type(as seen in H. mdolfianus, Text-fig. 3) and clearly approaches that found in H.saxicola and allied species in the Lake Victoria species flock (Greenwood, 1974). Inother words, the preotic region of the skull is slightly more elongate than in thegeneralized type, and associated with this and the shallower braincase, the preoticskull roof is straighter and slopes upwards at a smaller angle. The supraoccipital 158 P. H. GREENWOOD crest is lower than in most generalized types of skull, and the otic region is narrower. Although all the canal bones of the skull (and also those bones encasing the innerear) are noticeably inflated, the dermopterotics show the greatest hypertrophy of all(see Text-fig. gA and B). The lower pharyngeal bone (Text-fig. 10) is triangular in outline (length and breadthof the dentigerous surface almost equal), is relatively slender, and has an anteriorblade that is neither noticeably long nor noticeably short. The teeth on this boneare rather sparsely arranged in 16-20 irregular rows. Without exception, the teethare weakly bicuspid, with a low, blunt or even barely visible anterior cusp and acrown that slopes gently into the sharper and larger posterior cusp ; all are fine andcompressed but those in the two median rows are somewhat coarser. FIG. 10. Haplochromis macconneli. Lower pharyngeal bone in occlusal view. Vertebrae and the caudal fin skeleton. Vertebral counts for the 24 specimensradiographed are : 27 (fi), 28 (f8), 29 (fi4) or 30 (fi), comprising 12 (fi6) or 13 (f8)abdominal and 15 (fi), 16 (fi5) or 17 (f8) caudal centra. The fused first ural andfirst preural centra are excluded from these counts. The caudal fin skeleton is unusual in showing a high degree of fusion betweenvarious hypurals (or if not fusion, such extremely close apposition as to be in-distinguishable from fusion on radiographed specimens). The extent of fusion orapposition is generally complete, but in a few fishes there are short unfused sectionsbetween the otherwise conjoined elements. About 77 per cent of the 26 specimens radiographed showed some degree of fusionbetween various hypurals. Only 6 specimens (ca 23 per cent of the sample) had all5 hypurals completely free from each other. Most frequently (i.e. in 20 fishes)hypurals I and 2 are fused. In the upper part of the skeleton the commonestpattern of fusion is to have hypurals 3 and 4 fused, but hypural 5 free (n specimens) ;only 4 specimens had all 3 upper hypurals fused, and 5 fishes showed no fusionbetween any of the 3 upper hypurals. In H. rudolfianus, it will be recalled, only 2out of 30 specimens (i.e. about 7 per cent) had any fused hypurals ; in both thesefishes fusion was between hypurals i and 2 (see p. 146). LAKE RUDOLF HAPLOCHROMIS SPECIES 159 For comparison with the situation in Lake Rudolf, the caudal skeleton in severalendemic Haplochromis species from other lakes was examined. Of 174 specimens(representing 12 species) from Lake George, Uganda, about 8 per cent showed some(but never complete) fusion between elements in both the upper and the lower partsof the skeleton. The frequency of fusion seems equally distributed amongst thespecies examined. In Lake Victoria, too, hypural fusion is relatively rare. One hundred andseventy-eight specimens representing 22 species (with a modal sample size of 10specimens per species) were examined. Of these, 20 specimens (i.e. about 13 percent) had fused hypurals. The commonest pattern here is of fusion between hy-purals 3 and 4, less frequently it occurs between hypurals i and 2, and only once wasit recorded, with certainty, between hypurals i and 2, and 3 and 4 in the sameindividual (although two other individuals may show this pattern). The LakeVictoria species examined are from several phyletic lineages within the Haplochromisflock of that lake and cover a wide range of body forms. Far fewer specimens are available of the four endemic Haplochromis species fromLake Albert. Of these, H. bullatus Trewavas has 3 out of 18 fishes (i.e. about 17 percent) with hypurals i and 2 fused (or very closely apposed), while H. avium Regan(9 specimens) and H. albertianus Regan (15 specimens) have none. Both the latterspecies, however, have some individuals in which the hypurals are closely apposed. Thus, even allowing for the small sample sizes involved in this survey, it doesseem that the frequency of hypural fusion in H, macconneli is exceptionally high.The significance of this phenomenon remains unexplained. VISCERA. Because of poor preservation it is impossible to measure precisely thelength of the gut in H. macconneli ; I would estimate the intestine to be about i^times longer than the standard length. In strong contrast to H. rudolfianus there isno trace of melanin in the peritoneal tissue and neither are the gonadial walls pig-mented (see p. 146 above). COLORATION. In life, adult females are a pale greenish-fawn with traces of greenishiridescence on the flanks. All fins are colourless except for three conspicuous andbright yellow spots on the anal. Live colours for males have not been recorded. Preserved colours. I suspect that the material I have examined is somewhatbleached and thus the coloration is probably lighter than it might otherwise havebeen. There is apparently little sexual dichromatism. In both sexes the ground colourof the body and head is a pale yellowish-fawn with no tonal variation between dorsumand venter. All fins are hyaline with, in both sexes, dark maculations on the proxi-mal third to half of the caudal fin. The males examined have the dorsal fin somewhatdarker than that of the females, and there is a fairly dense aggregation of melano-phores on the anterior third of the pelvic fins. (These specimens may, however, beless bleached than are the females.) ECOLOGY. Judging from Mr Hopson's records, H. macconneli is confined to watermore than 20 m deep, and is probably most abundant in deeper water (i.e. at depths 160 P. H. GREENWOOD of 50-70 m). Some of the smallest specimens collected (15-20 mm S.L.) werecaught at a depth of 75 m, and none, of any size, has been recorded from littoralhabitats. The pale coloration of this species and the absence of melanic pigmentsin the peritoneum and gonads may well be correlated with this deep-water distribu-tion. No data are available yet on the feeding habits of the species. Breeding biology. The marked sexual dimorphism in adult size has been notedalready (p. 154). Despite a very thorough search, Mr and Mrs Hopson were at firstunable to find any males amongst the several hundreds of H. macconneli they ex-amined. Eventually two males, 31-0 and 35-5 mm S.L., were identified in a catchmade some 2-4 km east of North Island at a depth of 30 m. Both these fishesappear to be sexually adult; one is probably in an advanced stage of ripening, theother at a slightly earlier stage of development. Females are certainly adult at alength of ca 50 mm and it seems likely that they may mature at a smaller size(Hopson in litt.}. Be that as it may, females seem to grow to a much greater lengththan do males. In a sample of 21 adult females examined, only 2 have both ovaries equallydeveloped. Eleven fishes have the right ovary much larger than the left one (at allstages of oogenesis), and 8 have only the right ovary developed. Unlike H. rudolfi-anus, there is no trace of melanization in the ovarian wall of H. macconneli. No details are available on the breeding habits or seasons of the species. DIAGNOSIS AND AFFINITIES. Haplochromis macconneli is readily distinguishedfrom the other Lake Rudolf species by the hypertrophy of its cephalic laterosensorycanal system, its dentition (see p. 156), and in specimens > 35 mrn S.L., from H.rudolfianus by several morphometric characters (deeper cheek and preorbital,longer snout, smaller eye and longer jaws). Indeed, H. macconneli is so distinct morphologically from H. rudolfianus andH. turkanae (and the putative fourth species) that it is difficult to establish its phyleticaffinities with these other taxa. In addition to the characters listed above, H.macconneli also differs from H. rudolfianus (and from the undescribed species too)in the shape of its neurocranium, which is of a more specialized type (see above,p. 157 and Greenwood, 1974). Almost certainly the hypertrophy of the cephaliclateral line canals is an adaptation associated with the deep-water habitat of H.macconneli, and strongly suggests that the species evolved within the lake. Nofluviatile Haplochromis exhibits this specialization. It is interesting to note that the cephalic lateral line canal hypertrophy seen inH. macconneli is much greater than that found in any known Haplochromis speciesliving at comparable depths in Lake Victoria (Greenwood & Gee, 1969). AmongstHaplochromis-gToup species an equivalent hypertrophy is seen only in H. bullatusof Lake Albert, and in species of the endemic Lake Malawi genus Trematocranus(Trewavas, 1935). Species of another Malawi endemic, Aulonocara, show greaterdevelopment in certain parts of the system (the infraorbital series for example), butotherwise exhibit a level of hypertrophy comparable with that of H. macconneli. Trematocranus and Aulonocara are manifestly more closely related to each otherand to other taxa from Lake Malawi (Trewavas, 1935 ; personal observations) than LAKE RUDOLF HAPLOCHROMIS SPECIES 161 to any species occurring outside the lake. Neither need be considered further in thepossible phylogeny of H. macconneli. On purely morphological grounds H. bullatus of Lake Albert could be consideredthe nearest living relative of H. macconneli. Both species share the specializationof enlarged laterosensory canals on the head, and both share (with other species fromLakes Albert and Rudolf) certain peculiarities in the predorsal and thoracic squama-tion patterns (see above, p. 144). The latter character is, however, difficult to assesswith respect to its being a primitive or a derived one. At present all that can besaid is that the pattern is not encountered amongst the Haplochromis species ofLake Victoria nor is it seen in the fluviatile Haplochromis of Kenya, Uganda andTanzania. It is rarely encountered in the Haplochromis species flock of LakesEdward and George (where it is known from two species, H. pharyngalis Poll andH. petronius Greenwood ; see Greenwood, 1973), but it does seem to characterizethe Haplochromis of the River Nile, Lake Albert, Lake Rudolf and the River Zaire(personal observations ; also Greenwood, 1971). Haplochromis macconneli differs from H. bullatus in the shape of its neurocranium(see p. 157 above) which is like that found in the moderately specialized insectivore-piscivore radiation in Lake Victoria (see Greenwood, 1974, pp. 80-93). Thisdifference would not, of course, debar H. bullatus from consideration as the livingplesiomorph sister species of H. macconneli. But the fact that the Lake Rudolfand Lake Albert basins have never been interconnected (and if there had been someriverine connection, the probability that any presumed common ancestor of the twospecies would itself have been abyssal in habits) seems to rule out any such closephyletic relationship. Interspecific similarities in laterosensory canal hypertrophyare thus to be interpreted as the product of parallel evolution. The shared peculiari-ties in squamation patterns may well reflect a common ancestry but this is likely tobe a relatively distant one (see below, p. 162). It seems probable, therefore, that the relationships of H. macconneli should besought amongst the species of Lake Rudolf. Two interpretations seem possible.First, H. macconneli may be an immediate derivative of an H. rudolfianus-likeancestor (i.e. H. rudolfianus and H. macconneli may be true sister species). Alter-natively, H. macconneli might be the apomorph survivor of another lineage therelatively plesiomorph, that is H. rudolfianus-like members of which have becomeextinct (unlike the situation in Lake Victoria, for example, where it is possible tofollow, from species still extant, the specialization of a lineage ; see Greenwood,1974). DISCUSSION Even with the discovery of two and possibly three new Haplochromis species, thetotal fish fauna of Lake Rudolf still stands low on the scale of endemicity in Africanlakes. Its Haplochromis species flock also shows a low level of adaptive radiation,probably lower than that of Lake Albert (see Trewavas, 1938 ; Greenwood, 1971)where an anatomically specialized mollusc-eater, a specialized grazer on epiphytesand a species adapted for life in deep water have evolved. 162 P. H. GREENWOOD This comparison must, however, be interpreted with care. For one thing, H.mahagiensis David & Poll (the mollusc-crushing species) of Lake Albert may wellbelong to a different lineage from that of the other species in the lake. It could bethe local representative of a fluviatile mollusc-crushing species represented elsewhereby H. straeleni Poll and H. vanderhorsti Greenwood (see Greenwood, 1954 & iQSQa, fordiscussion). Furthermore, a temporal element is probably involved. There is littleevidence of Lake Albert having dried out at any time in its history, but Lake Rudolfprobably was severely reduced, or even completely desiccated, during the middlepart of the Pleistocene (see Fryer & lies, 1972, for review). Refilling of the Rudolfbasin appears to have been through what is now the River Sobat at some time withinthe later Pleistocene. Subsequently the connection was broken and has never beenre-established. Such an historical background has two consequences, namely thatLake Rudolf is to be considered a relatively young lake, and that its colonizers (or,perhaps more accurately, its recolonizers) were species of Nilotic origin. The relative youth of Lake Rudolf, coupled with the nature of its basin may, asFryer & lies (1972) suggest, account for the paucity of endemic species and, I wouldalso suggest, for the muted adaptive radiation seen amongst the three or fourHaplochromis species that evolved there. With two exceptions, all the Haplochromis species of Lake Albert are apparentlyconfined to the basin of that lake. The two more widely distributed species, H.wingatii (Blgr.) and H. loati Greenwood, both have dental specializations that arenot shared with any Lake Rudolf taxa (Greenwood, 1971). Thus, it seems unlikelythat an extant Albertine Haplochromis species was the original recolonizer of LakeRudolf in later Pleistocene times. Nor does it seem probable that the recolonizerclosely resembled any species from the Nile (amongst which must be considered H,wingatii and H. loati or their ancestors). In this situation the only conclusion thatcan be drawn is that some fluviatile species, now extinct, provided the founderpopulation for the Lake Rudolf microflock. It is, of course, possible that the present-day Rudolf species (or some part of them)are descendants from the relicts of a previous flock, possibly a more complex one,that inhabited the early Pleistocene lake and which survived the subsequent periodof desiccation. The neurocranial differences between H. rudolftanus and H. maccon-neli might be explained in this way. If this was the history of the present-dayspecies then it follows that their ancestors were derived not from the Nile (whichdid not then exist in its present form ; see Berry & Whiteman, 1968), but from ariver that originated in the eastern highlands and emptied into the developing Nilesystem in the region of the present River Sobat. Shared peculiarities in the squamation pattern of the thoracic and predorsalregions hint at a common ancestry for the Lake Rudolf and Lake Albert Haplochromisspecies (see above, p. 161). This character complex is not found in the H. bloyeti-likespecies group that is widespread in the rivers of Uganda, Kenya and Tanzania.Members of this species complex are thought to be close relatives of the ancestralspecies which gave rise to the sister species flocks in Lake Victoria and LakesEdward and George (see Greenwood, 1973 & 1974). Fishes in these flocks, with oneexception in Lake George and one in Lake Edward, all lack the Albert-Rudolf scale LAKE RUDOLF HAPLOCHROMIS SPECIES 163 pattern. The exceptional species, H. petronius and H. pharyngalis, resemble theNilotic H. wingatii in several features as well as the one of scale pattern, and theymay represent an exotic element amongst the otherwise H. bloyeti-like. derivativesinhabiting these lakes (Greenwood, 1974). The possible phyletic relationship ofHaplochromis species from Lakes Rudolf and Albert raises some interesting pointsof zoogeography. The lakes are several hundred kilometres apart and any form ofpast interconnection they may have had would have been of an indirect nature (seeabove, p. 162). In contrast, Lake Albert is close to Lake Edward and the lakes arein direct contact through the River Semliki. Yet, their faunas, both cichlid andnon-cichlid, are quite distinct (Greenwood, 1959^. The present barrier to faunalinterchange, principally the Semliki rapids, is clearly an effective one. I have argued elsewhere for a close phyletic relationship between the Haplochromisspecies of Lakes Edward and Victoria and for their derivation in parallel from acommon ancestor that once inhabited the westward flowing rivers of eastern Africaduring the Pleistocene (Greenwood, 1973). It seems now that perhaps this conceptshould be qualified by postulating an ancestral species that lived in some but notall of those rivers. The reasoning behind this qualification is, of course, the presenceof species in Lake Albert that would seem to be derived from a different lineage, alineage that also gave rise to the species of the Nile and, possibly, Lake Rudolf aswell (see above, p. 161). Furthermore, it is possible that the Haplochromis speciesof the River Zaire may share ancestry with these species (p. 161). The evidence upon which these postulated phylogenies are based, a shared scalepattern, is admittedly tenuous, particularly since it is not yet possible to determinewhich of the two types is to be considered the primitive condition. Further researchis planned to investigate the phylogeny of Haplochromis-group species and I wouldcertainly not consider the ideas expressed here as more than a working hypothesis. The presence of two species with Albert-Rudolf scale patterns (H. petronius andH. pharyngalis) in Lakes George and Edward demands explanation. Three pos-sible explanations can be considered. First, the ancestor of these species made itsway into the Lake Edward basin in fairly recent times and via the River Semliki.The likelihood of this, however, is reduced by the fact that no reciprocal exchange ofHaplochromis seems to have taken place (although the two lakes share anothercichlid Sarotherodon leucostictus [Trewavas]). Second, the prerift rivers each mayhave contained species of both squamation types. The absence of species with theAlbert-Rudolf pattern from most rivers in eastern Africa (and probably from LakeVictoria as well) would seem unlikely if both types had been represented therepreviously. The third possibility is one based on the assumption of there havingbeen a single basin in which the proto-Lakes Albert and Edward developed, probablyas a series of partly interconnected small and swampy lakes. The northern regionof this basin (the future Lake Albert) could have been fed by rivers in which theHaplochromis species had an Albert-Rudolf facies, while the southern end (futureLakes Edward and George) was fed by rivers with Haplochromis of a Victoria-Edwardfacies. The next assumption would be that only a limited exchange of species tookplace between the two regions before their continuity was broken. Victoria-typespecies, if any penetrated to the north, were, presumably, unsuccessful in that 164 P. H. GREENWOOD environment or in competition with the Albert-Rudolf types. Such a generalcontact between the early lakes may also account for the similarity in their Pleisto-cene fish faunas (see Greenwood, I95gb ; also new and unpublished observations),although one can equally argue that a fauna of this type was widespread in theprerift river systems. Whatever the explanation, it does seem that both Lake Albert and Lake Rudolfhave, since their inceptions, been relatively isolated from each other and from otherwater bodies in eastern Africa. ACKNOWLEDGEMENTS I am particularly grateful to Mr and Mrs A. J. Hopson for, in the first place, sendingme the specimens on which this paper is based, and then for the great pains they tookto obtain extra information and specimens. To my colleague Mr Gordon Howesgoes my gratitude for all the help he has given in the preparation of the paper, forhis draughtsmanship displayed in Text-figs. 5, 7 and 8, and for his skill in radio-graphing several hundred specimens. Finally, I thank my colleague Dr KeithBanister whose visit to Lake Rudolf (made since this paper went to press) led to hisproviding me with valuable extra information on Crater Lake species, and on thescarcity of H. macconneli males ; despite diligent searches he found none of thelatter. REFERENCES BEADLE, L. C. 1932. Scientific results of the Cambridge expedition to the East African lakes,1930-1. 4. The waters of some East African lakes in relation to their fauna and flora./. Linn. Soc. (Zool.}, 38 : 157-211. BERRY, L. & WHITEMAN, A. J. 1968. The Nile in the Sudan. Geogrl. J. 134 : 1-37. FRYER, G. & ILES, T. D. 1972. The Cichlid Fishes of the Great Lakes of Africa. Their Biologyand Evolution. Oliver and Boyd. Edinburgh. GREENWOOD, P. H. 1954. On two species of cichlid fishes from the Malagarazi River (Tangan-yika), with notes on the pharyngeal apophysis in species of the Haplochromis group. Ann.Mag. nat. Hist. (12), 7 : 401-414. I959a. The monotypic genera of cichlid fishes in Lake Victoria, Part II. Bull. Br. Mus. nat. Hist. (Zool.), 5 : 163-177. !959b. Quaternary fish fossils. Explor. Pare. natn. Albert Miss. J. de Heinzelin de Braucourt, 4 : 1-80. 1965. On the cichlid fishes of Lake Nabugabo, Uganda. Bull. Br. Mus. nat. Hist. (Zool.), 12 : 3I3-357- 1971. On the cichlid fish Haplochromis wingatii (Blgr.) and a new species from the Nile and Lake Albert. Revue Zool. Bot. afr. 84 : 344-365. 1973- A revision of the Haplochromis and related species (Pisces : Cichlidae) from Lake George, Uganda. Bull. Br. Mus. nat. Hist. (Zool.), 25 : 139-242. 1974- The cichlid fishes of Lake Victoria, east Africa : the biology and evolution of a species flock. Bull. Br. Mus. nat. Hist. (Zool.), Suppl. No. 6 : 1-134. & GEE, J. M. 1969. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), Part VII. Bull. Br. Mus. nat. Hist. (Zool.), 18 : 1-65.TREWAVAS, E. 1933. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. ii. The cichlid fishes. /. Linn. Soc. (Zool.}, 38 : 309-341. LAKE RUDOLF HAPLOCHROMIS SPECIES 165 TREWAVAS, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10), 16:65-118.— 1938. Lake Albert fishes of the genus Haplochromis. Ann. Mag. nat. Hist, (n), 1 : 435- 449- 1973- On the cichlid fishes of the genus Pelmatochromis with proposal of a new genus for P. congicus ; on the relationship between Pelmatochromis and Tilapia and the recognition of Sarotherodon as a distinct genus. Bull. Br. Mus. nat. Hist. (Zool.), 25 : 1-26.WORTHINGTON, E. B. 1932. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. I. General introduction and station list. /. Linn. Soc. (Zool.), 38 : 99-119. & RICARDO, C. K. 1936. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. 15. The fish of Lake Rudolf and Lake Baringo. /. Linn. Soc. (Zool.), 39 : 353-389. P. H. GREENWOOD, D.Sc. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.1965. (Out of Print.) £3.75. 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure andMineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ;29 Plates, 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : theBiology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.I974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol B$4 sNU /•v A REVIEW OF SCOTOECUS THOMAS, 1901(CHIROPTERA : VESPERTILIONIDAE) J. E. HILL BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 4 LONDON: 1974 /v OCTI974 A REVIEW OF SCOTOECUS THOMAS,(CHIROPTERA : VESPERTILIONIDAE) BY JOHN EDWARDS HILL Pp. 167-188 ; 4 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 4 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 4, of the Zoologicalseries. The abbreviated titles of periodicals citedfollow those of the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 20 September 1974 Price £1.05 A REVIEW OF SCOTOECUS THOMAS, 1901(CHIROPTERA : VESPERTILIONIDAE) By JOHN E. HILL INTRODUCTION CURRENT classification of the genus Scotoecus rests basically on Ellerman, Morrison-Scott & Hayman (1953 : 83) who recognized two species in southern Africa, one(albofuscus) with pale wings, the other (hirundo) with dark wings. Hayman(1963 : 104) expanded this opinion in a brief review, and allocated the named formsthen in the genus to one or other of these as subspecies. Rosevear (1965 : 300)retained faldbae from northern Nigeria as a distinct species, but Hayman & Hill(1971 : 36) recognized only albofuscus and hirundo as valid species. However,specimens examined recently at the British Museum (Natural History) have suggestedthat the dark-winged group is composed of two species, one smaller, for whichhirundo is available, the other a little larger in size, for which the earliest name ishindei. Hitherto, Scotoecus has been considered exclusively African, but a study ofthe Indian species first described as Scotophilus pallidus by Dobson (1876 : 186) andnowadays generally referred to Nycticeius shows it to be more closely allied toScotoecus, to which it is transferred. Measurements throughout are in millimetres : minimum and maximum valuesare given for series, with (in parentheses) the number of specimens measured. SYSTEMATIC SECTION SCOTOECUS Thomas, 1901Scotoecus Thomas, 1901 : 263. Type species Scotophilus albofuscus Thomas, 1890. DIAGNOSIS. Similar to Scotozous and Nycticeius but with spatulate or nearlyspatulate tragus (Fig. i) ; braincase elevated frontally to give a slightly convexfrontal profile ; rostrum greatly widened at the lachrymals ; narial and pre-palatal emarginations exceptionally deep ; zygomata tenuous, usually lost in pre-paration ; and anterior face of the upper canine flattened and grooved. DESCRIPTION. Small (length of forearm 28-38) ; muzzle broad, flattened,anteriorly sparsely haired, nares circular, opening obliquely ; ear rounded withrounded tip, its anterior margin lacking any basal lobe ; tragus spatulate or nearlyso with small, more or less triangular posterior basal lobe ; calcar extending alongtwo thirds of the uropatagial margin with small post-calcarial lobe ; penis sometimesexceptionally long. Skull low, braincase broad, with slight occipital helmet and low lambdoid andsagittal crests ; skull slightly elevated frontally to produce a faintly convex profile ;interorbital region wide, rostrum broad, much expanded at lachrymals ; anteorbitalforamen moderate or large, closed by strong, nearly vertical bar ; small supra-orbital tubercles often present; narial emargination wide, U-shaped or slightly i7o J. E. HILL V-shaped, the apex rounded, deep, extending posteriorly at least to a line joining theanterior margins of the anteorbital foramina, sometimes to a line joining the anteriormargins of the orbits ; pre-palatal emargination wide, extending posteriorly to apoint lying between a line joining the anterior faces of the large upper premolars(pm4-4) and a line joining the anterior faces of the first upper molars (m1-1) ; zygo-mata tenuous, in African species often lost in preparation ; maxillary toothrowsnot markedly convergent anteriorly ; a narrow post-palatal spine and shallowbasioccipital depressions. 1123 1113 Dental formula i-, c-, pm -, m - = 32 or i -, c-, pm -, m - = 30. Massive,3i 23 3123 unicuspid inner upper incisor (i2), in contact with the canine or nearly so ; anteriorface of upper canine flattened, grooved, sometimes prominently so ; anterior upperpremolar (pm2) minute or absent, when present compressed into a lingual recessbetween canine and the second upper premolar (pm4) which are otherwise in contact ;pm4 with antero-medial cusp or protocone low or absent ; upper molars (m1'3) withprotocones prominently developed ; the lingual shelves widely separated, m3 notreduced, its crown area exceeding one half the crown area of m1 or m2, with promi-nent metacone, metastyle and three commissures ; lower incisors (ij.g) imbricated ;anterior lower premolar in the Indian species reduced to one third to one half thecrown area of the second lower premolar (pm4) which is not compressed in thetoothrow ; in the African species pm2 a little less in crown area than pm4, the latterslightly shortened antero-posteriorly, compressed between pm2 and the first lowermolar (m^ ; third lower molar (m3) not reduced or only very slightly reduced, itsposterior triangle not at all or only a little smaller than the anterior triangle, hypo-conid and entoconid present, the latter slightly the lesser. LITERATURE. Trouessart, 1904 : 85 (spelt Scotaecus) ; Miller, 1907 : 217 (genericdiagnosis) ; Wettstein, 1918 : 654 (review, key) ; Simpson, 1945 : 59 (incorporatedinto Nycticeius) ; Ellerman, Morrison-Scott & Hayman, 1953 : 83 (status, partial key,a subgenus of Nycticeius) ; Hayman, 1963 : 104 (review, tooth formula, variability ofpm2, a subgenus of Nycticeius) ; Koopman, 1965 : 17 (status, generically valid) ;Rosevear, 1965 : 289, 297 (status, description, partial key, generically valid) ; Ansell1967 : 21 (as a genus) ; Hayman & Hill, 1971 : 36 (key, notes, distribution, subgenusof Nycticeius) ; Koopman, in litt., in Hayman & Hill, 1971 : 36 (a subgenus ofNycticeius}. DISTRIBUTION AND SPECIES. Indian and African, in India a semi-desert speciesinhabiting tropical thorn forest, in Africa occurring chiefly in open woodland inthree species with few records from the high forest. The Indian species (first de-scribed as Scotophilus pattidus by Dobson, 1876 : 186) was for many years referredto Scoteinus (see below) but is here allocated to Scotoecus on account of its aural androstral features, together with its grooved upper canines. Its features are lessextreme than are those of the African species of Scotoecus and apparently it forms alink between these and Nycticeius, with which in recent years it has been associatedThe African members of the genus may be divided into two sections, the one pale-winged, with brownish ventral surface, the other dark-winged, the ventral surface REVIEW OF SCOTOECUS THOMAS, 1901 171 pale, usually some shade of greyish. These correspond to the two species currentlyrecognized in Africa by most modern authors. The examination of Ethiopian speci-mens and of specimens from Cameroon and Uganda in the course of preparing thisstudy has suggested the further division of the dark-winged section into two species,one smaller, the other larger ; however, the genus is still known from relatively fewspecimens and the classification presented here is to that extent provisional. Thespecies may be keyed : 1 Tragus long, relatively narrow; zygomata not cartilaginous; pm2 reduced, its crown area one third to one half that of pm4 ; this tooth not compressed in toothrow ...... pallidus (p. 172) Tragus short, relatively wide ; zygomata cartilaginous ; pm2 unreduced,its crown area subequal to that of pm4 ; this tooth compressed intoothrow ........... 2 2 Tragus more or less parallel-margined, the upper part of its anterior margin not prolonged anteriorly ; outer part of wing membrane pale ;ventral surface of body brown ; braincase slightly flattened albofuscus (p. 174)Tragus slightly hatchet-shaped, the upper part of its anterior marginprojected forward ; outer part of wing membrane dark; ventralsurface of body greyish, at most tinged with brown ; braincase slightlyelevated ........... 3 3 Smaller, length of forearm generally less than 33-0, condylobasal length 13-2 or less, c-m3 5-1 or less . . . . . . hirundo (p. 177) - Larger, length of forearm generally exceeding 32-0, condylobasal length 13-5 or more, c-m3 5-2 or more ..... hindei (p. 179) HISTORY. First separated from Scotophilus at the generic level by Thomas(1901 : 263), Scotoecus was considered generically valid by Miller (1907 : 217), whoallied it with Nycticeius and Scoteinus. Allen (1939 : 98) listed Scotoecus as a distinctgenus for Africa but since that date opinions of its status have diverged. Simpson(1945 : 59) incorporated it into Nycticeius, a lead followed by Ellerman, Morrison-Scott & Hayman (1963 : 83) who treated Scotoecus as a subgenus of Nycticeius withthe comment that the characters given by Thomas and Miller seemed more subgenericthan generic. Hayman (1963 : 104) and Hayman & Hill (1971 : 36) adopted thisview but Rosevear (1965 : 289) noted that Scotoecus is readily distinguished fromNycticeius and considered that it merited generic separation, a view followed byAnsell (1967 : 21) and shared by Koopman (1965 : 17) who did not consider Scotoecuscongeneric with Nycticeius but later (in litt., in Hayman & Hill, 1971 : 36) revisedthis opinion to accord Scotoecus subgeneric rank only. REMARKS. There seem no good grounds for considering Scotoecus congenericwith Nycticeius : the form of the tragus, the elevated cranium and widened rostrumwith deep anterior emarginations, and the grooved upper canines of Scotoecusseparate it sharply from this genus. The convexity of the frontal area in Scotoecus,its broad rostrum with deep emarginations and its weak zygomata present a parallelwith Nyctalus, in which, however, no modification of the upper incisors has occurred. 172 J. E. HILL Scotoecus pallidus (Dobson, 1876) ? Vespertilio noctulinus Geoffrey, 1831 : 92, pi. 3. Bengal, India. Scotophilus pallidus Dobson, 1876 : 186, Appendix D, 208. Mian Mir, Punjab, northwesternIndia. DIAGNOSIS. Differing from the African S. albofuscus, S. hirundo and S. hindei inthicker, more fleshy ears, more swollen at the antitragus ; tragus (Fig. i) longer andrelatively narrower, its tip more expanded ; narial emargination faintly V-shaped,its lateral margins slightly convergent posteriorly, the apex rounded ; pre-palatalemargination narrower ; anteorbital foramen smaller ; zygomata tenuous but notcartilaginous ; anterior lower premolar (pm2) reduced, its crown area one third toone half that of the second lower premolar (pm4) which is not compressed in thetoothrow ; third lower molar (m3) slightly reduced, the posterior triangle a littlenarrower than the anterior triangle. Differs further from S. hirundo and S. hindeiin less inflated and less elevated braincase, in this respect resembling 5. albofuscus. DESCRIPTION. Large for the genus (length of forearm 34-38) ; lips fleshy ; earmoderate, reaching about halfway to tip of muzzle, with fleshy membrane and anti-tragal thickening ; anterior margin of tragus straight, tip rounded, anteriorly directed,posterior margin convex, penis not exceptionally developed. Dorsal surface palebrown, tinged with fawn, ventral surface paler, greyish white. Skull relatively massive, with (especially in older individuals) low occipital helmetand posterior sagittal and lambdoid crests ; slight supraorbital ridges with incipientsupraorbital tubercles ; anterior margin of orbit nearly vertical, the supraorbital andlachrymal regions swollen ; anteorbital foramen moderate, closed by narrow bar ;narial emargination wide, faintly V-shaped, slightly narrowed posteriorly, its lateralmargins slightly convergent, extending posteriorly a little more than halfway to theanterior orbital margin ; pre-palatal emargination wide, U-shaped, extending pos-teriorly to a line joining the anterior faces of the second upper premolars (pm4"4) ;zygomata narrow, tenuous but not cartilaginous ; post-palatal extension narrow,with small, delicate post-palatal spine. ii 13Dental formula i -, c -, pm -, m - = 30. Upper canine with flattened, faintly grooved anterior face ; small upper premolar (pm2) absent; large upper premolar(pm4) compressed between the canine and the first upper molar (m1), with smallantero-medial cusp or protocone ; m1'3 with strong protocones, their lingual shelveswidely separated ; anterior lower premolar (pm2) much reduced, one third to onehalf the crown area of the second lower premolar (pm4) and less than one half itsheight, compressed between the canine and pm4 ; third lower molar (m3) very slightlyreduced, posterior triangle a little narrower than the anterior triangle, hypoconid andentoconid slightly smaller than protoconid, paraconid and metaconid. MEASUREMENTS. Length of forearm (14) 34-1-38-4 ; condylobasal length (8)I4'i-i4'9 ; condylocanine length (8) 13-9-14-7 ; least interorbital width (10)4-0-4-4 ; zygomatic width (i) 10-5 ; width of braincase (9) 7-3-8-1 ; mastoid width(7) 8-9-9-7 ; ^-c1 (9) 4-9-5-5 ; m3-m3 (9) 6-5-6-9 ; c-m3 (u) 5-2-5-7 ; c-m3 (n)5-8-6-2. REVIEW OF SCOTOECUS THOMAS, 1901 173 LITERATURE. Dobson, 1876 : 186 (description), 208 (holotype listed) ; 1878 : 257,258 (in Scoteinus, a subgenus of Scotophilus), 264 (description repeated), pi. xv,fig. 3 (ear) ; Blanford, 1891 : 222 (further description) ; Trouessart, 1897 : 119,1904 : 85 (in Scoteinus, a subgenus of Scotophilus) ; Siddiqi, 1961 : 125 (Pakistanlocalities reviewed) ; Sinha & Chakraborty, 1971 : 53 (cranial features, measure-ments), figs, iA, 2A, 3A (skull). DISTRIBUTION. WEST PAKISTAN (Sind : Dobson, 1877 : 310 ; Anderson, 1881 :137 ; Wroughton, 1916 : 752 ; Siddiqi, 1961 : 125 ; Sinha & Chakraborty, 1971 : 54). INDIA (Punjab : Dobson, 1876 : 186, 208, 1877 : 310, 1878 : 264 ; Anderson,1881 : 137 ; Allen, 1908 : 48. Uttar Pradesh ; Bihar : Khajuria, 1951 : 120 ; Sinha& Chakraborty, 1971 : 54 ; specimens collected at Bahgownie, Darbhanga District,Bihar by N. A. Baptista in the collections of the British Museum (Natural History)). The record from Massowa, Eritrea, Ethiopia by Thomas & Doria (1886 : 206)in fact refers to Nycticeius schlieffenii, these authors having been misled by a com-parison with a specimen of schlieffenii with an erroneous locality, identified by Dobsonas pallidus (Thomas, 1890 : 86). HISTORY. A rarely collected species to which there are relatively few references.Described as a Scotophilus, the species was placed by Dobson (1878 : 257, 258) andby Trouessart (1897 : 119,1904 : 85) in the subgenus Scoteinus, subsequently elevatedto generic rank by Miller (1907 : 217) but diagnosed by this author without referenceeither to its type species (designated by Miller, p. 217) Scotophilus emarginatusDobson, 1871 or to Scotophilus pallidus Dobson, 1876. The type species has beenshown by Sinha & Chakraborty (1971 : 53) to be congeneric with the species knownfor many years as Scotomanes ornatus (Blyth, 1851)* and pallidus does not in anyevent conform to the diagnosis of Scoteinus by Miller : the lachrymal region is widenedas in the African species of Scotoecus and the third molars (m f) are almost un-reduced, the narrowing of the rostrum and reduction of m f being the chief diag-nostic features of Scoteinus as understood by Miller, who had seen only the Australianspecies balstoni and greyii and the African schlieffenii, all nowadays usually referredto Nycticeius. REMARKS. As might be expected from the disjunct distribution, the Indian 5.pallidus differs more widely from its African congeners than these do from each other.In particular, a number of its features are less extreme than in the African speciesand its dentition approaches more nearly to the type exemplified by Nycticeius.There seem no good grounds for referring pallidus to this genus, however, since inNycticeius the rostrum is not broadened and the upper canine is rounded anteriorly,not at all flattened or grooved : the same is true of the Australian species referredhitherto to Scoteinus and more recently to Nycticeius. In these, additionally, agreater measure of reduction of the third molars (m f) has occurred. * The generic synonymy was not discussed by Sinha and Chakraborty. Scoteinus and Scotomaneswere proposed simultaneously by Dobson (1875: 371) Scoteinus having line priority. Article 24 of theInternational Code of Zoological Nomenclature provides that in such a case the relative priority of thenames is determined by the action of the first reviser. Sinha and Chakraborty may be considered to befirst revisers and continue to use Scotomanes. In view of the varied uses of Scoteinus this action is inaccord with Recommendation 2^A of the International Code, which suggests that the name that ensuresstability and universality of nomenclature be selected. 174 J- E- HILL Vespertilio noctulinus I. Geoffrey, 1831. Tate (1942 :282) and Ellerman &Morrison-Scott (1951 : 177) suggested that this name should be associated withpallidus rather than considered a synonym of Scotophilus temminckii (= S. kuhlii) asit had been hitherto. It has a long history in the early literature : the descriptionis repeated by Temminck (1840 : 211), who, in a supplement to the same work,subsequently provided (p. 266) a further description with details of the dentition,referring specimens from Singapore in the collections of the Rijksmuseum vanNatuurlijke Historic, Leiden, to noctulinus. Cantor (1846 : 185), in a catalogue ofthe mammals of the Malay Peninsula referred the name to the synonymy of Scoto-philus temminckii (= 5. kuhlii) with the comment that Nycticeius noctulinus,Temminck referred to the young of the species, a view supported by Jentink (1888 :183) who listed a young example of 'Vespertilio noctulinus Is. Geoffroy' as Scotophilustemminckii. Wagner (1840 : 543, 1855 : 765) also provided further descriptions :Fitzinger (1870 : 367) gave a detailed synonymy to that date, with another descrip-tion (as Nycticejus noctulinus) but Dobson (1876 : 120, 1878 : 258) referred the nameto the synonymy of Scotophilus temminckii. Here it has since remained (Blanford,1891 : 320, although this author evidently had some reservations, placing a queryafter the allocation ; Trouessart, 1897 : 118) until recent years. It is of interest tonote that Oldfield Thomas has made a marginal note in his copy of Dobson's Cata-logue (1878) which reads 'Size of a pipistrelle! F.a. 36 (Not a Scotophilus}!' -thegrounds on which Tate suggested association with pallidus. Scotoecus albofuscus (Thomas, 1890) DIAGNOSIS. Similar to S. pallidus but smaller ; ears not especially fleshy ;tragus (Fig. i) smaller, spatulate, not prolonged anteriorly ; distal part of wingwhitish ; supraorbital region widely expanded ; lateral margins of narial emargina-tion not convergent posteriorly; anteorbital foramen large ; zygomata cartila-ginous ; anterior lower premolar (pm2) unreduced and third lower molar (m3) barelyreduced. DESCRIPTION. Small (length of forearm 28-5-31-0) ; ear membrane and anti-tragus relatively thin ; tragus small, spatulate, upper part of its anterior marginnot prolonged anteriorly. Dorsal surface brown, ventral surface similar but slightlypaler ; wing blackish brown from body to a line joining elbow to knee but whitishfrom forearm, the forearm and digits dark ; interfemoral membrane dark brown. Skull with broad, low, flattened braincase ; supraorbital region widely expanded ;narial emargination U-shaped, its lateral margins not convergent posteriorly;anteorbital foramen large, closed by a strong, nearly vertical bar; zygomata car-tilaginous, often lost in preparation. ii 13 Dental formula i -, c -, pm -, m - = 30. Upper canine with flattened, strongly312 3 grooved anterior face ; small upper premolar (pm2) absent in all eight specimensexamined and in one (No. 22412, Musee Royale d'Afrique Centrale, Tervuren) seenby Dr F. de Vree (in litt.) ; large upper premolar (pm4) with antero-medial cusp or REVIEW OF SCOTOECUS THOMAS, 1901 175 a i L mmi i i. j FIG. i. Tragus of (a) Scotoecus pallidus. (b) Scotoecus albofuscus. (c) Scotoecus hirundo. (d) Scotoecus hindei. protocone low or absent; upper molars (m1'3) with strongly developed protocones ;anterior lower premolar (pm2) unreduced, its crown area almost equal to that of thesecond lower premolar (pm4) which is compressed in the toothrow and is slightlyflattened antero-posteriorly ; third lower molar (ms) barely reduced, the width of itsposterior triangle nearly equal to that of the anterior triangle. DISTRIBUTION AND SUBSPECIES. Known from a limited number of specimensfrom localities in West Africa, Zaire, Tanzania and Malawi. Following Hayman(1963 : 105), Hayman & Hill (1971 : 37) recognized three subspecies, 5. a. albofuscusin West Africa, 5. a. cinnamomeus Wettstein, 1916 in the Sudan and S. a. woodiThomas, 1917 in southern and eastern Zaire, Tanzania and Malawi. However,cinnamomeus is based upon an example of Nycticeius schlieffenii*. Scotoecus albofuscus albofuscus (Thomas, 1890) Scotophilus albofuscus Thomas, 1890 : 84. Bathurst, Gambia. DIAGNOSIS. Upper canine with rounded base, its antero-internal margin notsharply angular, postero-internal margin rounded, lingual shelf narrow. MEASUREMENTS. Length of forearm (4) 29-8-31-0; condylobasal length (3)12-7-13-5 ; condylocanine length (3) 12-8-13-3 ; least interorbital width (3) 4-1-4-4 ;zygomatic width — ; width of braincase (4) 7-3-7-7 ; mastoid width (3) 8-2-9-0 ;ci-c1 (3) 4-4-5-1 ; m3-m3 (3) 6-6-6-7 ; c-m3 (4) 4'7~5'0 '> c-m3 (2) 5-1-5-3. * The status of cinnamomeus Wettstein, 1916 from Kordofan has been reviewed in some detail by Kock(1969: 188). The preliminary description by Wettstein (1916: 191) was succeeded by a more detailedstudy (1918: 653) with illustrations (p. 653, fig. 12) of the ear and tragus and more particularly (pi. 4,ngs- 5' 5a~b) of the skull and dentition. As Kock points out, these ally cinnamomeus undoubtedly withNycticeius schlieffenii. This opinion is supported by Koopman (in litt., in Hayman & Hill, 1971: 37)who has examined the holotype in Vienna. 176 J. E. HILL FIG. 2. Distribution of Scotoecus albofuscus. LITERATURE. Rosevear, 1965 : 298, 299 (description), fig. 78 (skull), 305, fig. 8oa(tragus). DISTRIBUTION. SENEGAL (Badi: Thomas, 1911 : 119, as a location in Gambia) ;GAMBIA ; SIERRA LEONE (both Rosevear, 1965 : 299, from specimens in BritishMuseum (Natural History)); IVORY COAST (Ayeremou (= Ahieremou, 6°i4'N.,4°56' W. ?), specimens in the Museum d'Histoire Naturelle, Geneva (F. de Vree,in litt.)) ; NIGERIA (Dollman, 1908 : 546). Scotoecus albofuscus woodi Thomas, 1917 Scotoecus woodi Thomas, 1917 : 280. Chiromo, southern Malawi, 200 feet. DIAGNOSIS. Upper canine with angular base, its antero-lateral margin forming aright angle, postero-internal margin angular and projecting, the lingual shelf wide. MEASUREMENTS. Length of forearm (5) 28-5-30-5 ; condylobasal length (4)12-9-13-7 ; condylocanine length (4) 12-9-13-7 ; least interorbital width (4) 4-1-4-3 ;zygomatic width — ; width of braincase (4) 7-2-7-6 ; mastoid width (3) 8-8-9-0 ; REVIEW OF SCOTOECUS THOMAS, 1901 177 c1-^ (4) 4-8-5-1 ; m3-m3 (4) 6-3-6-8 ; c-m3 (4) 4-8-5-1 ; c-m3 (4) 5-1-5-5 (speci-mens all from Chiromo, Malawi). Dr F. de Vree (in litt.) has measured a specimen(No. 22412) in the Musee Royale d'Afrique Centrale, Tervuren, from Baudouinville(07°O3' S., 2g°42' E.), Zaire : length of forearm 29-4 ; condylobasal length 12-9 ;least interorbital width 4-6 ; zygomatic width — ; width of braincase 7-3 ; mastoidwidth c. 8-4 ; m3-m3 6-7 ; c-m3 5-2 ; c-m3 (alveoli) 4-8 ; c-m3 (alveoli) 5-5. LITERATURE. Harrison, 1958 : 98, fig. i (whole animal), fig. 2 (head). DISTRIBUTION. ZAIRE (Hayman, 1957 : 44; Hayman, Misonne & Verheyen,1966 : 56) ; TANZANIA (Harrison, 1958 : 96 ; Vesey-Fitzgerald, 1964 : 64) ; MALAWI ;probably in ZAMBIA (Ansell, 1960, Appendix A, p. no, Appendix B, p. 117). REMARKS. According to Thomas (1917 : 280), woodi of southern Africa can bedistinguished from the West African albofuscus by its smaller size, proportionatelybroader skull and by the different shape of the base of the upper canine. The smallseries of albofuscus and woodi available in the collections of the British Museum(Natural History) do not support the statement that woodi is significantly smallerthan albofuscus and it appears to differ in size from albofuscus only in a generallyvery slightly wider rostrum, equalled in width by one of albofuscus. Scotoecus hirundo (de Winton, 1899) Scotophilus hirundo de Winton, 1899 : 355. Gambaja, Ghana, 1300 feet. DIAGNOSIS. Similar in most respects to 5. albofuscus but anterior margin oftragus (Fig. i) projected anteriorly in its distal part to give the tragus a slightlyhatchet-shaped outline, its upper margin nearly horizontal, the junction of the upperand posterior margins angular, slightly obtuse ; wing membranes wholly dark ;ventral surface of body some shade of grey ; braincase slightly inflated and roundeddorsally, less flattened ; pm2 usually but not always present, the dental formula .11 23 .11 13 i -, c -, pm -, m - = 32 or i -, c -, pm -, m - = 30.3123 ii 23 VARIABILITY OF PM2. The presence of a minute anterior upper premolar (pm2)sandwiched tightly between the canine and pm4 is variable in 5. hirundo, pm2 beingtotally absent from three of the nineteen specimens examined. The tooth is presentin both sides of the jaw in a specimen (in the Museum National d'Histoire Naturelle,Paris) from Saboya, Senegal. Dr F. de Vree (in litt.) has examined further specimens(in the Museum d'Histoire Naturelle, Geneva) from the Bandia Forest, Senegal: twoof these possessed the small pm2 but so far as could be seen, this was lacking from athird. It is present in both sides of the jaw in one from Sierra Leone but totallyabsent from another (Hayman, 1963 : 105). There is no trace of pm2 in the holotypefrom Ghana : it is present on both sides of the jaw in one specimen from the Cameroonbut in another present in the left hand side only. Of ten examples from Ethiopia,eight have the tooth on both sides of the jaw, in one it is present in the right hand sideonly and in one it is totally absent. Three specimens from Uganda have pm2present on both sides of the jaw. 178 J. E. HILL MEASUREMENTS. Length of forearm (18) 29-7-32-8 ; greatest length of skull(15) 12-6-13-7 ; condylobasal length (16) 12-1-13-0 ; condylocanine length (16)12-1-13-1 ; width of rostrum (18) 5-9-6-5 ; width across anteorbital foramina (18)4-6-5-2 ; least interorbital width (18) 4-2-4-6 ; zygomatic width (i) 10-3 ; width ofbraincase (16) 7-0-7-7 ; mastoid width (13) 8-0-8-8 ; c1-c1 (18) 4-3-5-0 ; m3-m3(18) 5-9-6-7 ; c-m3 (18) 4-7-5-1 ; length of complete mandible (15) 8-9-9-9 >c-m3 (18) 4-9-5-4. Dr F. de Vree (in litt.) has measured three specimens (seeabove) (one skull only prepared) from the Bandia Forest, Senegal: length offorearm 31-0-32-3 ; greatest length of skull 13-9 ; condylobasal length 12-9 ; leastinterorbital width 4-7 ; width of braincase 7-8 ; mastoid width 8-9 ; m3-m3 6-5 ;c-m3 5-3 ; c-m3 (alveoli) 4-9 ; length of mandible 10-1, c-m3 (alveoli) 5-3. LITERATURE. Rosevear, 1965 : 300 (further description), 305, fig. 8oc (tragus). DISTRIBUTION. SENEGAL (Saboya, specimen in Museum National d'HistoireNaturelle, Paris ; Bandia Forest, specimens in the Museum d'Histoire Naturelle,Geneva (F. de Vree, in litt.)) ; Sierra Leone (Hayman, 1963 : 105) ; GHANA ;CAMEROON (Yagoua, specimens obtained by F. de Vree and W. Verheyen) ; ETHIO-PIA (Gambela, 515 and 1768 metres, 8°i5' N., 34°35' E., and Didessa River, WollegaProvince, 1190 metres, 9°O2' N., 36°O9' E., specimens in British Museum (NaturalHistory)); UGANDA (Budongo, i°39' N., 3i°35' E., specimens in British Museum(Natural History)). REMARKS. In recent years (following Hayman, 1963 : 104) the named forms inthe dark-winged section of Scotoecus have been considered subspecies of hirundo,the earliest name. However, measurements of specimens in the collections of theBritish Museum (Natural History) and of specimens examined in 1971 for Dr F. deVree, Antwerp, suggest that two species are involved, basically one of smaller sizeand northern distribution for which the prior name is hirundo, and a second speciesof larger size and predominantly more southern distribution for which the earliestname is hindei. A specimen (B.M. 72.4421) from the Didessa River, Wollega Province, Ethiopia,is in close agreement with hirundo from West Africa : another (B.M. 72.4423) fromthe same locality is clearly referable to hindei. These are supported by a series ofspecimens (B.M. 70.2263-2270) and a further example (B.M. 72.4420), all fromGambela, western Ethiopia, which agree with hirundo : others (B.M. 70.2262,72.4422) from Bulcha, Lake Margherita, some 200 miles to the southeast of Gambela,are referable to hindei. Scotoecus has apparently not been collected hitherto inEthiopia. The two also occur almost sympatrically in Uganda : specimens (B.M.74.1-5) (including pregnant or lactating females) obtained at Budongo (i°39' N.,3i°35' E.) by J. F. Kingdon and clearly referable to hirundo contrast sharply in sizewith others (B.M. 63.1151, 65.3435) referable to hindei from West Madi (3°3o' N.,3i°35' E.) and Nabumali, South Bugisu (o°59' N., 34°i2' E.). Furthermore, twospecimens obtained at Yagoua, northern Cameroon, by F. de Vree and W. Verheyenagree very closely with hirundo and were collected concurrently at that locality withtwo further larger specimens referable with little doubt to falabae, here consideredto be a subspecies of hindei. No Scotoecus has been reported hitherto from the REVIEW OF SCOTOECUS THOMAS, 1901 179 Cameroon. There is evidence to suggest, therefore, that in Ethiopia a small species(hirundo) occurs with or in close proximity to a larger species (hindei) : both occurin nearby localities in Uganda and are again sympatric in the Cameroon. Measure-ments of hirundo and of hindei from the areas of sympatry or near sympatry arecontrasted in Table i. FIG. 3. Distribution of Scotoecus hirundo. Scotoecus hindei Thomas, 1901 DIAGNOSIS. Similar to S. hirundo but larger, especially cranially, with the supra-orbital and anteorbital regions wider and more greatly expanded. VARIABILITY OF PM2. The minute anterior upper premolar (pm2) is generallypresent in 5. hindei : its presence in twenty-six examples in the collection of theBritish Museum (Natural History) is variable only in those from Ethiopia and Kenya.The tooth is present on both sides of the jaw in two specimens from Ethiopia, butin a third example present only in the left-hand side of the jaw. Among Kenyanspecimens, the tooth is present on both sides in eight, present in the right-hand sidebut absent from the left in another, and totally absent from two further examples,one the holotype of S. hindei hindei. Hollister (1918 : 94) notes that pm2 is com-pletely absent from one of five Kenyan specimens in the Museum of Natural History, i8o J. E. HILL FIG. 4. Distribution of Scotoecus hindei. Records referred to 5. h. albigula areindicated by unshaded circles O- the Smithsonian Institution, Washington, but that an alveolus can be seen on oneside. The tooth is present in a further specimen reported (as albigula) by this authorfrom Kiriba, Sudan (see below). Furthermore, Dr F. de Vree notes (in litt.) thatpm2 is absent from a specimen (M.R.A.C. 22192) in the Musee Royale d'AfriqueCentrale, Tervuren, from Kapalowe, Katanga Zaire. Allen (1911 : 330) describespm2 in detail. DISTRIBUTION. Nigeria, Cameroon and Sudan to Ethiopia and Somalia ; Kenya,Uganda and Tanzania to Zambia ; Zaire ; Angola. Scotoecus hindei hindei Thomas, 1901 Scotoecus hindei Thomas, 1901 : 264. Kitui, Kenya, 1150 metres (in original description as 3500 feet).(?) Scotoecus artinii de Beaux, 1923 : 98. Archer's Post, northern Guaso Nyiro, Kenya. MEASUREMENTS. Length of forearm (15) 32-0-37-2 ; greatest length of skull (14)13-8-15-0 : condylobasal length (15) 13-4-14-4 ; condylocanine length (15) 13-3-14-4 ; width of rostrum (14) 6-6-7-8 ; width across anteorbital foramina (14) REVIEW OF SCOTOECUS THOMAS, 1901 181 5-1-6-0 ; least interorbital width (15) 4-4-4-8 ; zygomatic width (i) 10-7 ; widthof braincase (15) 7-3-8-0; mastoid width (15) 8-8-9-7 '< ^-c1 (14) 5-0-5-6;m3-m3 (14) 6-6-7-4; c-m3 (16) 5-3-5-7; length of complete mandible (13)10-0-11-2 ; c-m3 (16) 5-5-6-4. Dr F. de Vree (in litt.) has measured a specimen(see above) from Kapalowe, Katanga, Zaire : length of forearm 33-2 ; greatest lengthof skull 14-7 ; condylobasal length 13-8 ; least interorbital width 4-5 ; zygomaticwidth 10-5 ; width of braincase 7-6 ; mastoid width 9-1 ; m3-m3 7-0 ; c-m3 5-4 ;c-m3 (alveoli) 4-9 ; length of mandible 10-1 ; c-m3 (alveoli) 5-3. DISTRIBUTION. SUDAN (G. M. Allen, 1914 : 349, as Scoteinus schlieffenii, identifiedas Scotoecus by Koopman, 1965 : 16 ; Hollister, 1918 : 94, from Kiriba, 'Uganda', alocality in southern Sudan according to Kock, 1969 : 192, as albigula ; Kock, 1969 :192) ; ETHIOPIA (from Bulcha, Lake Margherita, 1800 metres, 6°n' N., 38°io' E.,and from Didessa River, Wollega Province, 1190 metres, 9°02' N., 36°09' E., speci-mens in British Museum (Natural History)) ; SOMALIA (de Beaux, 1924 : 155, asartinii ; Funaioli, 1971 : 29, 64, as hirundo) ; ZAIRE (J. A. Allen, 1916 : 447, asScoteinus schlieffenii, identified as Scotoecus hirundo hindei by Koopman, 1965 : 16 ;Hayman, 1954 : 291 ; Leleup, 1956 : 77 ; Anciaux de Faveaux, 1958 : 271 ; Hay-man, Misonne & Verheyen, 1966 : 56) ; UGANDA (Hayman & Hill, 1971 ; 36, asalbigula) ; KENYA (G. M. Allen, 1911 : 330 ; Dollman, 1914 : 309, in part Nos. 148,149, 150, as Scoteinus schlieffenii, re-identified in the present study ; Hollister, 1918 :94 ; Harrison, 1961 : 293) ; TANZANIA (a manuscript note by R. W. Hayman in aBritish Museum (Natural History) copy of Hayman & Swynnerton, 1951 (p. 294),records a specimen from Lake Mangona, but this example not seen ; Dodoma,specimen in British Museum (Natural History)). Scotoecus artinii de Beaux, 1923. The precise status of artinii is far from clear.Koopman (1965 : 17) is inclined to regard artinii, hindei and a third more southerlyform, albigula, as synonymous and notes that a specimen reported from Niangara,eastern Zaire, by J. A. Allen (1916 : 447) as Scoteinus schlieffenii is in fact a Scotoecus,referring it to S. hirundo hindei (= S. hindei hindei} with the remark that its bestagreement seems to be with artinii. Hayman & Hill (1971 : 36) listed artinii as asubspecies of hirundo (S. hindei as here understood), considering (p. 37) on the basisof specimens from the Cherangani Hills, western Kenya, that possibly it might be asmall form distinct from both hindei and albigula. Three specimens attributed to artinii by R. W. Hayman were collected in 1966 byMr A. N. Start at the Wei Wei River, Sigor, northeastern Cherangani Hills, Kenya,at about 3000 feet, the same collector subsequently obtaining in 1967 further examples(now B.M. 68.481-482) at the same locality. Mr Start also obtained three specimensattributed to albigula by R. W. Hayman from the Cherangani Hills at 6000 feet in1965. Of these eight it has been possible to examine only B.M. 68.481-482, but thecollections of the British Museum (Natural History) include three further examples(B.M. 14.7.31.15-17) of 5. hindei from the Wei Wei River, formerly identified asScoteinus schlieffenii (Nos. 148, 149, 150 of Dollman, 1914 : 309). Male specimens from the Wei Wei River agree quite closely with the male holotypeof 5. hindei hindei and with male specimens referred to this subspecies from the 182 J. E. HILL Lorian Swamp and the Guaso Nyiro, differing only in their slightly more slendercanines. In this respect they resemble the male examples B.M. 63.1151 from WestMadi, Uganda, and B.M. 65.3435 from Nabumali, South Bugisu, Uganda. How-ever, a single male specimen collected 30 miles northwest of Baringo, Kenya, bridgesthis difference. Specimens from the Wei Wei River and from Uganda are generallya little smaller than specimens from more easterly locations and are similar in sizeto the specimen from Zaire measured by J. A. Allen (1916 : 447) (as Scoteinusschlieffenii}. On the whole they are a little larger than specimens of S. hindei fromthe northern Cameroon. De Beaux gives few diagnostic characters for artinii but his measurements indicatea short skull with the palatal width and toothrow dimensions of 5. hindei : Hollister(1918 : 94) reports hindei from Archer's Post, Kenya, the type locality of artinii.For the present artinii is considered a provisional synonym of S. h. hindei but thereremains the possibility that a subspecies with slender canines is to be found in westernKenya, Uganda and eastern Zaire and it is to this that perhaps artinii refers. Ifartinii can be distinguished then three subspecies of hindei occur in Kenya, namelyS. h. hindei to the north, east and south, S. h. artinii to the west and northwest and5. h. albigula to the southwest. It is possible also that the rather larger albigula maybe found to be specifically valid, but direct sympatry with hindei (including artinii)has yet to be demonstrated. The reported occurrences of the three forms in EastAfrica display a degree of overlap. Scotoecus hindei albigula Thomas, 1909 Scotoecus albigula Thomas, 1909 : 544. Kirui, Mount Elgon, Kenya, 6000 feet. DIAGNOSTIC. Larger than S. h. hindei, with larger, more massive teeth ; canineslonger and heavier. MEASUREMENTS. Length of forearm (3) 35-0-38-5 ; greatest length of skull (3)15-3-15-5 ; condylobasal length (3) 14-7-14-8 ; condylocanine length (3) 14-8-15-0 ;width of rostrum (3) 7-2-7-7 ; width across anteorbital foramina (3) 5-7-6-0 ; leastinterorbital width (3) 4-5-4-9 ; zygomatic width — ; width of braincase (3) 8-0-8-1 ;mastoid width (3) 9-6-9-9 ; cl-cl 5-3-5-8 ; ms-m3 (3) 7-5-8-0 ; c-m3 (5) 5-9-6-0 ;length of complete mandible (3) 11-2-11-6 ; c-m3 6-5-6-6. LITERATURE. Monard, 1935 : 52 (notes, description repeated, in French) ; Hill& Carter, 1941 : 52 (notes, descriptive data), 177 (measurements of Angolan specimen,from Monard). DISTRIBUTION. KENYA (three examples collected in 1965 by A. N. Start in theCherangani Hills, east of Mount Elgon, northeast of Kitale, Kenya, at 6000 feet,examined by R. W. Hayman : see Hayman & Hill, 1971 : 36, 37) ; ANGOLA (Monard,*935 : 52 I Hill & Carter, 1941 : 52, 177 ; Hayman, 1963 : 104) ; ZAMBIA (Wrough-ton, 1907 : 4 ; Lancaster, 1953 : 18, as Scotoecus woodi, identified as Nycticeiushirundo hindei (= Scotoecus hindei hindei) by Ansell, 1960 : 23, Appendix A, p. no,Appendix B, p. 117 ; Ansell, 1967 : 21, as 5. hirundo hindei (= S. hindei hindei).Hollister (1918 : 94) recorded albigula from Kiriba, Uganda, but Kock (1969 : 192) REVIEW OF SCOTOECUS THOMAS, 1901 183 considers this to represent S. hirundo hindei (= S. hindei hindei) and notes thatKiriba is in the southern Sudan, 10 miles south of Gondokoro. REMARKS. The dimensions of B.M. 7.1.11.6 from Petauke, Zambia (Wroughton1907 : 4), are similar to those of S. h. albigula, and this specimen has the generallyheavier canines and more massive cheekteeth associated with this form, to which itis referred. Specimens in the Kaffrarian Museum (K.M. 1982, 1983) reported asScotoeciis woodi from the Fort Jameson District by Lancaster (1953 : 18) are identifiedas 5. hirundo hindei (= S. hindei hindei) by Ansell (see above). These may in factalso be referable to albigula, and provisionally are listed as such here, as is a specimenreported from Mfuwe Camp, Zambia by Ansell, 1967 : 21 as S. hirundo hindei (= S.hindei hindei} but which is larger than this subspecies. It is possible that further specimens may show albigula to be specifically distinct.It occurs as far north in Kenya as Mount Elgon and the Cherangani Hills, while S. hhindei has been obtained from more southerly localities in Katanga, Zaire and inTanzania. However, in Kenya albigula has been reported only from high elevations,5. h. hindei only from lower altitudes, and direct sympatry has yet to be shown tooccur. Scotoecus hindei falabae Thomas, 1915 Scotoecus falabae Thomas, 1915 : 447. Kabwir, northern Nigeria, 2500 feet. DIAGNOSIS. Similar in most respects to 5. h. hindei but slightly smaller and dorsallya little paler. MEASUREMENTS. Length of forearm (8) 32-0-35-2 ; greatest length of skull (7)13-8-14-4 ; condylobasal length (6) 13-2-13-5 ; condylocanine length (6) 13-5-13-7 ;width of rostrum (7) 6-5-7-0 ; width across anteorbital foramina (7) 5-0-5-3 ; leastinterorbital width (7) 4-3-4-6 ; zygomatic width — ; width of braincase (6) 7-3-7-9 ;mastoid width (6) 8-4-9-0 ; c^c1 (7) 4-7-5-0 ; m3-m3 (7) 6-6-6-9 '> c-m3 (8)5-2-5-5 ; length of complete mandible (5) 9-8-10-0 ; c-m3 (7) 5-6-5-8. LITERATURE. Rosevear, 1965 : 300 (further description, as a species), 305, fig. Sob(tragus). DISTRIBUTION. NIGERIA (Kabwir ; Yaba ; Jos, specimens in British Museum(Natural History)) ; CAMEROON (Yagoua ; Mokolo (Mayo Louti), specimens ob-tained by F. de Vree and W. Verheyen). REMARKS. Specimens from the northern Cameroon are very similar in size to5. h. falabae from Nigeria but on the whole have slightly more massive canines and,to a lesser extent, more massive cheekteeth. In this respect they approach 5. h.hindei. There is apparently no sexual dimorphism in size either in 5. hirundo or in 5.hindei : however, female specimens in both have slightly smaller, more slendercanines than do male examples. The canines of specimens attributed to 5. hirundoare generally less massive than are those of specimens referred to S. hindei, but in thecase of S. hirundo and 5. hindei falabae the canines of males of hirundo are almostexactly similar in size to those of females of falabae. 184 J. E. HILL The only localities at which specimens attributed to S. hirundo have been found sofar to occur sympatrically with others referred to 5. hindei are Yagoua, northernCameroon (Scotoecus having been hitherto unreported from the Cameroon), andDidessa River, Wollega Province, Ethiopia, with near sympatry at other Ethiopianlocalities and in Uganda. From Yagoua, F. de Vree and W. Verheyen obtained fourexamples of Scotoecus, two males and two females. The two male specimens arequite clearly referable to S. hindei falabae with which they are in close agreement.The two female specimens have considerably shorter skulls and agree closely withfemales of 5. hirundo. F. de Vree and W. Verheyen also obtained a male and femalefrom Mokolo (Mayo Louti) but these are of similar size, agreeing in this respect withfalabae to which they are referred. The females from Yagoua are smaller than thefemale of falabae from Mokolo and consequently are referred to 5. hirundo. Measure-ments of these specimens, and of others from Ethiopia and Uganda, appear in Table i. TABLE Measurements (in millimetres) of specimens from areas o Registration No. Sex Length offorearm 5. hirundo B.M. 72.4421 9 3i'7 B.M. 70.2267 6" 32-0 B.M. 70.2269 6* 31-1 B.M. 70.2263 1 70 2266 1 29-7 B.M. 70.2268 rB.M. 70.2270 J (6) B.M. 72.4420* $ 31-8 B.M. 74.4 3 32-0 B.M. 74.1 9 32-5 B.M. 74.2 9 30-4 B.M. 74.3 9 30-8 B.M. 74.5* $ 30-1 1.430 9 3i-5 I-43I ? 32-8 S. hindei hindei B.M. 72.4423 6* 34-1 B.M. 70.2262 6* 37'2 B.M. 72.4422 6* 33'4 B.M. 63.1151 6* B.M. 65.3435 $ 32-7 S. hindei falabae I-4I5 6* 32-9 1.429 6* 32-6 1.667 6* 32-0 Greatest length Condylo- Condylo- of basal canine skull length length 1.682 33-5 12-6 12-8 •13-4 (5) 13-2 13-213-5 14-214-8 14-414-2 13-813-914-0 13-9 12-412-7 I2-I •12-5 (5) 12-412-712-9 12-5 12-412-7 13-614-213-613-9 13-7 13-4I3-313-2 13-2 12-312-6 I2-I •12-5 (5)12-4 12-8 13-0 12-6 12-512-9 13-6 14-213-5 13-8 13-9 13-513-513-5 13-6 Rostralwidth 6-16-36-5 6-1 -6-4 (6) 6-0 6-56-4 6-2 6-2 6-0 7-16-7 6-66-7 6-56-87-0 6-5 Width across Least anteorbital interorbit foramina width 4-64-9 5'2 4'7-5-0 (6) 4-65'25'° 4-6 4-94-8 5'45'3 5'35'3 5-2 4-2 4'44-2 4'2 •4-5 (6) 4-24'5 4'7 4'4 4'34'3 4-84-64-64-6 4'7 4'4 4'34-5 4'4 Young adult REVIEW OF SCOTOECUS THOMAS, 1901 185 ACKNOWLEDGEMENTS My thanks are due to Dr M. J. Largen of the Haile Selassie I University, AddisAbaba, Ethiopia, whose specimens, donated to the British Museum (NaturalHistory), first suggested this study. Dr F. de Vree of the Laboratorium voorAlgemene Dierkunde, Rijksuniversitair Centrum, Antwerp, Belgium, has generouslyallowed me to make use of his important specimens (collected with Dr W. Verheyen)from the northern Cameroon, and, furthermore, has been especially helpful in pro-viding measurements and data from specimens in other European collections, drawnfrom his own records. I am indebted also to Dr V. Aellen of the Museum d'HistoireNaturelle, Geneva, Switzerland, who has given permission for the publication of datataken from specimens in the collections in his care, and to Mr J. F. Kingdon, whosespecimens from Uganda proved invaluable to the course of the work. mpatry or near sympatry between 5. hirundo and S. hindei Width gomatic of Mastoid width braincase width m3-m3 c-m3 Length of completemandible c-m3 Location 7-1 7-57-2 7'3 •7-5 (4) 7-0 7'57-6 7-1 7'37'3 8-2 8-3 8-2 -8-3 (5) 7'7 8-3 8-3 8-2 4'3 4'74-8 4-6 •4'7 (6) 4'3 4-8 4'74-6 4'54'5 6-06-66-6 6-3•6-5 (6) 5'96-36-6 6-3 6-66-7 4'74-9 4'7•4-8 (6)4'7 4-84'9 9'4 8-9.9-9 (6)9-1 9'99-6 9'4 9-09-6 4'95'35'3 4-9 •5-2(6) 5-i5'55'3 5'4 Didessa, EthiopiaGambela, EthiopiaGambela, Ethiopia Gambela, Ethiopia Gambela, EthiopiaBudongo, UgandaBudongo, UgandaBudongo, UgandaBudongo, UgandaBudongo, UgandaYagoua, CameroonYagoua, Cameroon 7-5 8-9 5-1 6-9 5-3 io-o 5-6 Didessa, Ethiopia 7-8 9-4 5-2 7-0 5-7 10-6 6-1 Bulcha, Ethiopia 7-9 8-8 — 5-4 5-8 Bulcha, Ethiopia 7-3 8-8 5-3 6-9 5-3 10-3 5-7 West Madi, Uganda 8-0 9-2 5-2 6-5 5-4 10-6 5-8 Nabumali, Uganda 7-5 8-4 4-8 6-7 5-5 io-o 5-8 Yagoua, Cameroon 7-3 8-6 5-0 6-9 5-3 io-o 5-8 Yagoua, Cameroon 7-4 8-6 4-9 6-7 5-2 9-8 5-6 Mokolo (Mayo Louti), Cameroon7-4 8-9 4-7 6-8 5-3 io-o 5-7 Mokolo (Mayo Louti), Cameroon 186 J. E. HILL SUMMARY The vespertilionid genus Scotoecus is reviewed in detail, and is considered to includefour species, one, pallidus, from India, having been referred at one time to Scoteinusand more recently to Nycticeius. The three remaining, albofuscus, hirundo and hindeiare African : evidence is brought forward to support the view that hirundo andhindei are specifically distinct, not conspecific as thought by the majority of modernauthors. Both are reported for the first time from the Cameroon and from Ethiopia. REFERENCES ALLEN, G. M. 1908. Notes on Chiroptera. Bull. Mus. comp. Zool. Harv. 52 : 25-62, i pi. 1911. Bats from British East Africa. Bull. Mus. comp. Zool. Harv. 54 : 321-331. 1914. Mammals from the Blue Nile Valley. Butt. Mus. comp. Zool. Harv. 58 : 305-355, i fig. 1939- Checklist of African mammals. Bull. Mus. comp. Zool. Harv. 83 : 1-763. ALLEN, J. A. 1917. In ALLEN, J. A., LANG, H. & CHAPIN, J. P. The American Museum Congo Expedition collection of bats. Part I. Systematic List. Butt. Am. Mus. nat. Hist. 37 : 405-478, 15 figs., i pi.ANCIAUX DE FAVEAUX, F. M. 1958. Speologica africana. Chiropteres des grottes du Haut-Katanga (Congo Beige). Bull. Inst. fr. Afr. noire Ser. A, 20 : 263-275.ANDERSON, J. 1881. Catalogue of Mammalia in the Indian Museum, Calcutta. Part I. Primates, Prosimiae, Chiroptera, and Insectivora. Calcutta.ANSELL, W. F. H. 1960. Mammals of Northern Rhodesia. Lusaka. 1967. Additional records of Zambian Chiroptera. Arnoldia, (Rhodesia), 2 (38) : 1-29. BLANFORD, W. T. 1891. The Fauna of British India, including Ceylon and Burma. Mammalia. Part II. London.CANTOR, T. 1846. Catalogue of the Mammalia inhabiting the Malayan Peninsula and islands. /. Asiat. Soc. Bengal, 15 : 171-203.DE BEAUX, O. 1924. Mammiferi della Somalia Italiana. Elenco delle specie e sottospecie finora riconsciute presenti, con annotazioni di sistematica e zoo-geografia. Atti Soc. ligust. Sci. nat. geogr. N.S. 3 : 149-168.DE WINTON, W. E. 1899. On mammals collected by Lieut.-Colonel W. Giffard in the Northern Territory of the Gold Coast. Ann. Mag. nat. Hist. (7), 4 : 353-359-DOBSON, G. E. 1876. Monograph of the Asiatic Chiroptera and Catalogue of the species of bats in the collection of the Indian Museum, Calcutta. London. 1877. Notes on a collection of Chiroptera from India and Burma, with descriptions of new species. /. Asiat. Soc. Beng. 46 : 310-313. 1878. Catalogue of the Chiroptera in the Collection of the British Museum. London. DOLLMAN, G. 1908. On a collection of bats from Yola, northern Nigeria, collected by Mr. G. M. Webster. Ann. Mag. nat. Hist. (8), 2 : 545-547. 1914. Notes on a collection of East African mammals presented to the British Museum by Mr. G. P. Cosens. Proc. zool. Soc. Lond. 307-318.ELLERMAN, J. R. & MORRISON-SCOTT, T. C. S. 1951. Checklist of Palaearctic and Indian Mammals, 1758-1946. London. - & HAYMAN, R. W. 1953. Southern African Mammals, 1758-1951 : a reclassifica- tion. London.FITZINGER, L. J. 1870. Kritische Durchsicht der Ordnung der Flatterthiere oder Handfliigler (Chiroptera). Familie der Fledermause (Vespertiliones). V. Abtheilung. Sber. Akad. Wiss. Wien. 62, (i) : 353-448. FUNAIOLI, U. 1971. Guida breve dei Mammiferi della Somalia. Oltremare.GEOFFROY, M. I. 1831. In BELANGER, M. C. Voyage aux Indes-Orientales, etc. Zool. Paris. REVIEW OF SCOTOECUS THOMAS, 1901 187 HARRISON, D. L. 1958. Two bats (Microchiropetra) new to the fauna of Tanganyika Territory. Durban Mus. Novit. 5 : 95-98, 2 figs. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. // E. Africa nat. Hist. Soc. 23 : 286-295.HAYMAN, R. W. 1954. Notes on some African bats, mainly from the Belgian Congo. Rev. Zool. Bot. afr. 50 : 277-295. 1957- Further notes on African bats. Rev. Zool. Bot. afr. 56 : 41-45. 1963. Mammals from Angola, mainly from the Lunda District. Publcoes cult. Co. Diam. Angola, No. 66 : 81-140, 15 figs. & HILL, J. EDWARDS 1971. In MEESTER, J. & SETZER, H. W. The Mammals of Africa. An Identification Manual. Part 2. Chiroptera. Washington.— MISONNE, X. & VERHEYEN, W. 1966. The bats of the Congo and of Rwanda and Burundi. Annls Mus. r. Afr. cent. ser. in 8°, Sci. zool. No. 154 : i-viii, 1-105, 2O P*s-HILL, J. ERIC & CARTER, T. D. 1941. The mammals of Angola, Africa. Bull. Am. Mus. nat. Hist. 78 : 1-211, 34 figs., 17 pis.HOLLISTER, N. 1918. East African mammals in the United States National Museum. Part i. Insectivora, Chiroptera and Carnivora. Bull. U.S. natn. Mus. No. 99 : 1-194, 55 P^s-JENTINK, F. A. 1887. Museum d'Histoire Naturelle de Pays-Bas etc. Tome XII. Cata-logue Systdmatique de Mammiferes. (Rongeurs, Insectivores, Cheiropteres, Edentes et Marsupiaux.) Leiden.KHAJURIA, H. 1951. Taxonomic studies on some Indian Chiroptera. Rec. Indian Mus. 50 : 113-128.KOCK, D. 1969. Die Fledermaus-Fauna des Sudan (Mammalia, Chiroptera). Abh. sencken. naturforsch. Ges. 521 : 1-238, 20 figs.KOOPMAN, K. F. 1965. Status of forms described or recorded by J. A. Allen in 'The American Museum Congo Expedition Collection of Bats'. Am. Mus. Novit. No. 2219 : 1-34.LANCASTER, D. G. 1953. A check list of the mammals of Northern Rhodesia. Lusaka.LELEUP, N. 1956. La faune cavernicole du Congo Beige et considerations sur les Coleopteres reliques d'Afrique intertropicale. Annls Mus. r. Congo Beige, ser. in 8°, Sci. zool. No. 46 : 1-170, 118 figs., 5 pis.MILLER, G. S. 1907. The families and genera of bats. Bull. U.S. natn. Mus. 57 : i-xvii, 1-282, 49 figs., 14 pis.MONARD, A. 1936. Contribution a la mammalogie d'Angola et Prodrome d'une faune d'Angola. Arches Mus. Bocage, 6 : 1-309. ROSEVEAR, D. R. 1965. The Bats of West Africa. London.SIDDIQI, M. S. 1961. Checklist of mammals of Pakistan with particular reference to the mammalian collection in the British Museum (Natural History), London. Biologia, Lahore 7 : 93-225.SIMPSON, G. G. 1945. The principles of classification and a classification of mammals. Bull. Am. Mus. nat. Hist. 85 : i-xvi, 1-349.SINHA, Y. P. & CHAKRABORTY, S. 1971. Taxonomic status of the vespertilionid bat, Nycti- cejus emarginatus Dobson. Proc. zool. Soc. Calcutta 24 : 53-59, 3 figs., i tab.SWYNNERTON, G. H. & HAYMAN, R. W. 1951. A checklist of the land mammals of the Tanganyika Territory and the Zanzibar Protectorate. // E. Africa nat. Hist. Soc. 20 : 274- 392.TATE, G. H. H. 1942. Results of the Archbold Expeditions. No. 47. Review of the Ves- pertilionine bats, with special attention to genera and species of the Archbold Collections. Bull. Am. Mus. nat. Hist. 80 : 221-297, 5 n§s-TEMMINCK, C. J. 1840. Monographies de Mammalogie. Paris.THOMAS, O. 1890. Description of a new Scotophilus from the Gambia, with remarks on some of the allied species. Annali Mus. civ. Stor. nat. Giacomo Doria (2), 9 : 84-88. 1901. A new Scotophiline bat from British East Africa, with the description of a new genus of the group. Ann. Mag. nat. Hist. (7), 7 : 263-265. 1909. New African mammals. Ann. Mag. nat. Hist. (8), 4 : 512-549. i88 J. E. HILL THOMAS, O. 1911. List of small mammals obtained by Mr. G. Fenwick Owen on the Upper Gambia and in Fouta Djallon. Ann. Mag. nat. Hist. (8), 8 : 117-124. 1915. A new bat from Northern Nigeria. Ann. Mag. nat. Hist. (8), 16 : 447-448. — 1917. A new bat of the genus Scotoecus. Ann. Mag. nat. Hist. (8), 19 : 280-281. & DORIA, G. 1886. Note intorno al alcuni Chirotteri appartenenti al Museo Civico di Geneva e descrizione di due nuove specie del genere Phyllorhina. Annali Mus. civ. Stor. nat. Giacomo Doria (2), 4 : 201-207.TROUESSART, E.-L. 1897. Catalogus mammalium tarn viventium quam fossilium. Tomus I. Primates, Prosimiae, Chiroptera, Insectivora, Carnivora, Rodentia, Pinnipedia. Berlin. 1904. Catalogus mammalium tarn viventium quam fossilium. Quinquennale supple- mentum. Berlin.VESEY-FITZGERALD, D. F. 1964. Mammals of the Rukwa Valley. Tanganyika Notes Rec. No. 62 : 61-72.WAGNER, J. A. 1840. Die Sdugethiere in Abbildungen nach der Natur mit Beschreibung von D. Johann Christian Daniel von Schreber. Supplementband i. Leipzig. 1855. Die Sdugethiere in Abbildungen nach der Natur mit Beschreibung von D. Johann Christian Daniel von Schreber. Supplementband 5. Leipzig.WETTSTEIN, O. v. 1916. Neue Affen und Fledermause aus Nordost-Afrika. Anz. Akad.Wiss. Wien. 53 : 189-192. 1918. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Kaiserliche Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommen zoologischen Expedition nach dem Anglo-Agyptischen Sudan (Kordofan) 1914. II. Bearbeitung der auf der Expedition gesammelten Vogel und Saugethiere. Denkschr. Akad. Wiss. Wien. (1917), 1918, 94 : 555-693, 13 figs., 4 tabs., map.WROUGHTON, R. C. 1907. On a collection of mammals made by Mr. S. A. Neave in Rhodesia, north of the Zambesi, with field notes by the collector. Manchester Memoirs, 51 (5) : 1-39. 1916. Bombay Natural History Society's Mammal Survey of India, Burma and Ceylon. Report No. 24. Sind. /. Bombay nat. Hist. Soc. 24 : 749-758. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.1965. (Out of Print.) £3.75. 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure andMineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ;29 Plates, 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa: theBiology and Evolution of a Species Flock. Pp. 134; I Plate, 77 Text-figures.1974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 sNU :••; CATALOGUE OF THE TYPESTERRESTRIAL ISOPODS (ONISCOIDEA)IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY)I. SUPERFAMILY PSEUDOTRACHEATA R. J. LINCOLN AND J. P. ELLIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 5 LONDON: 1974 CATALOGUE OF THE TYPES OF TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) I. SUPERFAMILY PSEUDOTRACHEATA BY ROGER ]. LINCOLN AND JOAN P. ELLIS Pp. 189-246 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 5 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, «issued in Jive series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 5, of the Zoologicalseries. The abbreviated titles of periodicals citedfollow those of the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 23 December, 1974 Price £3.10 CATALOGUE OF THE TYPES OF TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) I. SUPERFAMILY PSEUDOTRACHEATA By ROGER J. LINCOLN AND JOAN P. ELLIS THE superfamily Pseudotracheata comprises the following ten families of terrestrialisopods : Cylisticidae, Porcellionidae, Balloniscidae, Trachelipidae, Atlantidiidae,Armadillidiidae, Eubelidae, Sphaeroniscidae, Actoeciidae, Armadillidae, encompas-sing all those groups of woodlice which possess the characteristic pseudotrachealrespiratory apparatus (Vandel, 1962, 1963, 1968, 1970). The collections of the British Museum (Natural History) are very rich in typematerial of this group of woodlice, acquired in large part from the extensive collec-tions of Gustave Budde-Lund, A. M. Norman and Karl W. Verhoef, and to a lesserextent from C. L. Koch, Adrien Dollfus, J. Omer-Cooper, K. H. Barnard, W. E.Collinge and others. This important collection of types has now been indexed andcatalogued for the first time, and it is hoped at a future date to provide a similarcatalogue listing the rest of the holdings of woodlice types. The catalogue contains in all some 672 separate items relating to 522 species in77 genera. The genera are listed alphabetically within families, and the species aresimilarly listed within the appropriate genus. All available information, includingthe reference to the original description, number and sex of specimens, type status,locality, date, collector, method of acquisition and British Museum (Natural History)registration number, is given with each item. Unless otherwise stated the materialis preserved in spirit. Where the author's name appears in brackets it indicates that the entry is made ina combination other than the original combination, in which case the latter is givenin square brackets thus : albicornis (Dollfus) (iSg6b : 3) [Lucasius albicornis] Where a particular species is now considered a junior synonym of another speciesthe latter is also indicated in brackets after the entry : sphinx Verhoeff (i93ia : 533) [Transferred to Porcellio lamettatus Uljanin] A full index of species, genera and families is provided at the end of the catalogue. The collection also contains a large number of specimens labelled as types, mostlyfrom the Budde-Lund collection, for which no reference can be found in publishedliterature. It seems probable that these are specimens with manuscript nameswhich have been incorporated into personal collections but which have never beenpublished. A list of this material comprising 167 items is given in Appendix i, anda list of the manuscript names has been deposited in the library of the BritishMuseum (Natural History). I92 R. J. LINCOLN AND J. P. ELLIS Family GYLISTICIDAE CYLISTICUS Schnitzler annulicornis Verhoeff (igoSb : 184) SYNTYPES: three females. Reg. no. 1907:11:4:36-38. Toscana region, Italy. Purchasedfrom K. W. Verhoeff.biellensis Verhoeff (i93oa : 176) SYNTYPES : two females. Reg. no. 1930:5:26:54-55. Piemonte, Italy. Purchased fromK. W. Verhoeff. SYNTYPES: two males; six females. Reg. no. 1937:7:6:109-113. Piemonte, Italy.Purchased from K. W. Verhoeff.caucasius Verhoeff (1917 : 157) SYNTYPES: male. Reg. no. 1921:6:10:46. Gagri, Caucasus Mts. Collected by N.Lignau. Purchased from K. W. Verhoeff.dentifrons Budde-Lund (1885 : 81) HOLOTYPE : fragments. Reg. no. 1921:10:18:4212. Astrakhan, USSR. Budde-LundCollection.esterelanus Verhoeff (1917 : 155) SYNTYPE : female. Reg. no. 1921:6:10:45. Esterel Mts, France. Purchased from K. W.Verhoeff.gracilipennis Budde-Lund (1885 : 79, 229) SYNTYPE: male. Reg. no. 1911:11:8:10063. Rome, Italy. Collected by Bergs0e.Norman Collection (ex Copenhagen Museum). SYNTYPE : male. Reg. no. 1921:10:18:4213. (Redetermined as Cylisticus esterelanusVerhoeff.) Pierrefeu, France. Collected by E. Simon. Budde-Lund Collection. SYNTYPES : three males ; three females. Reg. no. 1921:10:18:2414-4218. Rome, Italy.Collected by Bergs0e. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1956:10:10:201-202. Rome, Italy. Collectedby Bergs0e. Presented by University College, Dundee.nasatus Verhoeff I93ia : 527 SYNTYPE : one juvenile. Reg. no. 1931:4:27:42. Villa Napoleon, Elba I. Collected byK. Strasser. Purchased from K. W. Verhoeff. SYNTYPE : female. Reg. no. 1930:5:26:52. Ormea, Liguria, Italy. Purchased fromK. W. Verhoeff.pallidus Verhoeff (19280 : 159) SYNTYPE : female. Reg. no. 1930:5:26:53. Liguria, Italy. Purchased from K. W.Verhoeff.plurnbeus Verhoeff (igoib : 73) SYNTYPE : female. Reg. no. 1901:9:19:41. Lake Garda, Italy, Purchased from K. W.Verhoeff.plurnbeus bergomatius Verhoeff (19280 : 162) SYNTYPES: one male ; one female. Reg. no. 1928:7:4:107-108. Bergamo, Italy. Pur-chased from K. W. Verhoeff.plurnbeus umbricus Verhoeff (19280 : 163) SYNTYPES: one male; one female. Reg. no. 1928:7:4:105-106. Umbria, Italy. Pur-chased from K. W. Verhoeff.pontremolensis Verhoeff (i936a : 112) SYNTYPES: one male; three females. Reg. no. 1937:7:6:114-117. Spezia, N. Italy.Purchased from K. W. Verhoeff.suberorum Verhoeff (193 la : 526) SYNTYPE : juvenile. Reg. no. 1931:4:27:41. Monte Massoncello, Italy. Purchasedfrom K. W. Verhoeff.transsilvaticus Verhoeff (igoSb : 185) SYNTYPE: female. Reg. no. 1907:11:4:39. Siebenbiirgens. Purchased from K. W. Verhoeff. TYPES OF TERRESTRIAL ISOPODS 193 Family PORCELLIONIDAEACAEROPLASTES Verhoeff argentarius (Verhoeff) (i93ia : 541) [Metoponorthus (A.) argentarius] SYNTYPE : female. Reg. no. 1931:4:27:62. Monte Argentario, Italy. Purchased fromK. W. Verhoeff.melanurus (Budde-Lund) (1885 : 181) [Metoponorthus melanurus] SYNTYPES : two females. Reg. no. 1921:10:18:5273-5274. Porto Calvien, Corsica.Collected by E. Simon. Budde-Lund Collection. SYNTYPES: two males; one female. Reg. no. 1956:10:10:145-146. Bona, Algeria.Collected by Meinert. Presented by University College, Dundee. AGABIFORMIUS Verhoeff hirtus (Aubert & Dollfus) (1890 : 8) [Lucasius hirtus] SYNTYPE : male. Reg. no. 1911:11:8:10532. Marseille, France. 'In soil at the foot of awall.' Collected by A. Dollfus. Norman Collection. SYNTYPE : female. Reg. no. 1921:10:18:4023. Marseille, France. Budde-Lund Collec-tion.minutus (Budde-Lund) (igoga : 8) [Angara minuta] SYNTYPES : three females. Reg. no. 1921:10:18:4028-4030. Porto Tunis, Tunisia. Col-lected by F. Silvestri. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1921:10:18:4031-4032. Sidi Bel Akaren.Collected by F. Silvestri. Budde-Lund Collection.obtusus (Budde-Lund) (igoga : 8) [Angara obtusa] SYNTYPES: one male; four females. Reg. no. 1921:10:18:4061-4065. Cairo, UAR.18.5.1901. Collected by L. A. Jagerskjold. Budde-Lund Collection.rufobrunneus Omer-Cooper (1923 : 401) SYNTYPE: fragments. Reg. no. 1922:5:18:6. Amara, Iraq. 13.2.1918. Collected byW. E. Evans. Presented by J. Omer-Cooper. SYNTYPES : micropreparations of appendages : six males, one female. Reg. no. 1922:5:18:51-57. Amara, Iraq. 13.2.1918. Collected by W. E. Evans. Presented by J. Omer-Cooper. CAEROPLASTES Verhoeff porphyrivagus (Verhoeff) (1918 : 134) [Metoponorthus (C.) porphyrivagus] SYNTYPE : female. Reg. no. 1921:6:10:31. French Riviera. Purchased from K. W.Verhoeff. DESERTONISCUS Verhoeff subterraneus Verhoeff (igsob : 120) SYNTYPE : female. Reg. no. 1930:5:26:73. Turkestan, USSR. Purchased from K. W.Verhoeff. ENNURENSIS Collinge hispidus Collinge (19153, : 143) PARATYPE : female. Reg. no. 1919:4:26:519. Ennur, near Madras, India. 'Under stoneson sand 19.10.1913.' Collected by N. Annandale. Collinge Collection. 194 R- J- LINCOLN AND J. P. ELLIS GERUFA Budde-Lund hirticornis Budde-Lund (igogb : 59) SYNTYPES : two females. Reg. no. 1921:10:18:1468-1469. Cape Flats, South Africa.Collected by L. Schultze. Budde-Lund Collection.macrops Barnard (1932 : 276) SYNTYPES : four males; nineteen females. Reg. no. 1933:1:25:208-215. Oudebosch,River Zonder End Mts, Cape Province, South Africa. Collected by K. H. Barnard. Pre-sented by the South African Museum.marmorata Barnard (1932 : 277) SYNTYPES : four males ; thirty-six females. Reg. no. 1933:1:25:216-227. George, CapeProvince, South Africa. Collected by K. H. Barnard. Presented by the South AfricanMuseum.montana Barnard (1932 : 275) SYNTYPES : five females. Reg. no. 1933:1:25:204-207. Riversdale Mts, Cape Province,South Africa. Collected by K. H. Barnard. Presented by the South African Museum. HALOPORCELLIO Verhoeff penicilliger Verhoeff (1917 : 170) HOLOTYPE : male. Reg. no. 1921:6:10:78. Yafo, Israel. Collected by Aharoni. Pur-chased from K. W. Verhoeff.sphinx Verhoeff (i93ia : 533) [Transferred to Porcellio lamellatus Uljanin] SYNTYPE : female. Reg. no. 1931:4:27:61. Populonia, Massoncella, Italy. Purchasedfrom K. W. Verhoeff. HEMILEPISTUS Budde-Lund bodenheimeri Verhoeff (i93ib : 40) [Transferred to Hemilepistusreaumuri (Audouin & Savigny)}SYNTYPE : female. Reg. no. 1931:4:27:68. Palestine. Collected by Bodenheimer.Purchased from K. W. Verhoeff. SYNTYPES: one male; three females. Reg. no. 1938:7:7:41-44. Palestine. Collectedby Bodenheimer. Purchased from K. W. Verhoeff.cristatus Budde-Lund (1885 : 153) SYNTYPES: one male ; three females. Reg. no. 1911:11:8:10443-10446. Serdscen, Iran.Norman Collection. SYNTYPES : one male; one female. Reg. no. 1956:10:10:156-157. Serdscen, Iran.Presented by University College, Dundee.elongatus Budde-Lund (1885 : 160) HOLOTYPE : female. Reg. no. 1921:10:18:4103. 'Taschburun in Transcaucasius.' June1879. Collected by A. Brandt. Budde-Lund Collection.nodosus Budde-Lund (1885 : 159) SYNTYPES: one male; one female. Reg. no. 1921:10:18:4115-4116. 'Tschinas, RossiaAsiatica.' 1878. Collected by V. Russow. Budde-Lund Collection.palaestinus Verhoeff (i93ib : 38) SYNTYPES : two males. Reg. no. 1931:4:27:65-67. Palestine. Purchased from K. W.Verhoeff. SYNTYPES: three males ; three females. Reg. no. 1938:7:7:35-40. Palestine. Purchasedfrom K. W. Verhoeff.schirasi Lincoln (1970 : 127) PARATYPE : male. Reg. no. 1970:199:1. Shiraz, Iran. Collected by Kollar. Budde-Lund Collection.zachvatkini Verhoeff (i93ob : 122) [Transferred to H. nodosus Budde-Lund] SYNTYPE : female. Reg. no. 1930:5:26:74. Turkestan, USSR. Purchased from K. W.Verhoeff. TYPES OF TERRESTRIAL ISOPODS 195 HEMIPORCELLIO Collinge carinatus Collinge (igisa : 145) PARATYPE : female. Reg. no. 1919:4:26:458. Lake Chilka, Rambha, Ganjam District,Madras, India. 'At edge of Lake, 1913.' Collinge Collection.imtnsi (Collinge) (i9i4a : 207) \Povcellio immsi] HOLOTYPE : female. Reg. no. 1914:8:26:1. Allahabad, India. 16.9.1907. Collectedand presented by A. D. Imms. PARATYPES : one male ; one juvenile female. Reg. no. 1919:4:26:456-457. Allahabad,India. 16.9.1907. Collected by A. D. Imms. Collinge Collection. INCHANGA Barnard natalensis Barnard (1932 : 278) SYNTYPES : five males ; eleven females. Reg. no. 1933:1:25:228-233. Inchanga, Natal,South Africa. 1917. Collected by K. H. Barnard. Presented by the South AfricanMuseum. LEPTOTRICHUS Budde-Lund politus Omer-Cooper (1923 : 402) SYNTYPES: three males; one female. Reg. no. 1922:5:18:7-10. Amara, Iraq. 1918.Collected by R. G. Tame. Presented by J. Omer-Cooper. SYNTYPES : micropreparations of appendages : three males ; six females. Reg. no.1922:5:18:35-43. Amara, Iraq. 1918. Collected by R. G. Tame. Presented by J.Omer-Cooper. LUCASIUS Kinahan pallidus (Budde-Lund) (1885 : 134) [Porcellio pallidus] SYNTYPE : male. Reg. no. 1921:10:18:5182. Algier. Budde-Lund Collection.SYNTYPES : three males. Reg. no. 1921:10:18:5183-5185. Tlemcen, Algeria. Collectedby E. Simon. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1921:10:18:5186. Mid-France. Collected by E. Simon.Budde-Lund Collection. MAHEHIA Budde-Lund bicornis Budde-Lund (1913 : 376) SYNTYPES : one male ; two females. Reg. no. 1913:1:8:64-66. Silhouette I., Seychelles,Indian Ocean. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. SYNTYPES : Seven males ; seven females. Reg. no. 1921:10:18:997-1008. Silhouette I.,Seychelles, Indian Ocean. Budde-Lund Collection.laticauda Budde-Lund (1913 : 376) SYNTYPES : six males ; one female. Reg. no. 1913:1:8:58-63. Cascade, Mt Alphonse,Mahe I., Seychelles, Indian Ocean. 1800 ft, 'in base of Pandanus leaf. Collected by the'Sealark' Expedition. Presented by J. S. Gardiner.tnaculata Budde-Lund (1913 : 375) SYNTYPE : female. Reg. no. 1913:1:8:50. Mt Alphonse, Mahe I., Seychelles, Indian Ocean.3.12.1905. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. SYNTYPE : female. Reg. no. 1921:10:18:1021. Top of Mt Morne, Mahe I., Seychelles,Indian Ocean. 2700 ft. Budde-Lund Collection. SYNTYPE : few fragments only. Reg. no. 1921:10:18:1022. Seychelles, Indian Ocean.Budde-Lund Collection. I96 R. J. LINCOLN AND J. P. ELLIS METOPONORTHUS Budde-Lund approximates Budde-Lund (1885 : 185) SYNTYPE : male. Reg. no. 1921:10:18:5230. Sebastopol. Budde-Lund Collection.benaci L. Koch (1901 : 56) SYNTYPES : one male; three females. Reg. no. 1925:7:22:697-700. Torbole, Italy.Koch Collection.cilicius Verhoeff (1918 : 138) SYNTYPE : female. Reg. no. 1921:6:10:35. Toros Gari Mts, Cilicia, Turkey. Purchasedfrom K. W. Verhoeff.coxalis Budde-Lund (1885 : 175) SYNTYPE: female. Reg. no. 1911:11:8:10520. 'Ouled Anteurs', Algeria. Collected byE. Simon. Norman Collection. SYNTYPES: one male; two females. Reg. no. 1921:10:18:5235-5237. 'Rochers desOuled Anteurs', Algeria. Collected by E. Simon. Budde-Lund Collection.frontosus Budde-Lund (1885 : 183) SYNTYPE : male. Reg. no. 1911:11:8:10525. Biskra, Algeria. Collected by Meinert.Norman Collection (ex E. Simon collection). SYNTYPES: one male; one female. Reg. no. 1921:10:18:5239-5240. Collected byMeinert. Budde-Lund Collection.fuscomarmoratus Budde-Lund (1885 : 189) SYNTYPE: female. Reg. no. 1911:11:8:10521. Oran, Algeria. Collected by E. Simon.Norman Collection. SYNTYPES: three females. Reg. no. 1921:10:18:5241-5243. Oran, Algeria. Collectedby E. Simon. Budde-Lund Collection.gravei Verhoeff (1918 : 135) SYNTYPE : male. Reg. no. 1921:6:10:36. Sicily. Purchased from K. W. Verhoeff.hispida (Miers) (i877b : 676) {Porcellio (Porcellionides) hispida] SYNTYPES : one male ; one female. Reg. no. 1879 : 21. Mongolia. Presented by Prof.Wrzesniowsky.lacteolus Budde-Lund (1885 : 186) SYNTYPE: female. Reg. no. 1911:11:8:10492. 'Bou-Saada', Algeria. Collected by E.Simon. Norman Collection. SYNTYPES : one male; one female. Reg. no. 1921:10:18:4400. 'Bou-Saada', Algeria.Collected by E. Simon. Budde-Lund Collection.littoralis Budde-Lund (1885 : 179) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:5247-5248. 'Kertsch', Crimea.Collected by V. N. Uljanin. Budde-Lund Collection. SYNTYPES: two males; one female. Reg. no. 1921:10:18:5249-5251. (RedeterminedProtracheoniscus occidentalis Vandel.) Montpellier, France. Collected by E. Simon. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1921:10:18:5252-5253. Crimea, Ukraine,USSR. Collected by V. N. Uljanin. Budde-Lund Collection. myrmicidarum Verhoeff (1918 : 132) [Transferred to M. (Myrmeconiscus) myrmecophilusmyrmecophilus (Stein)] SYNTYPES: one male; one female. Reg. no. 1921:6:10:37-38. Sicily. Purchased fromK. W. Verhoeff.nigrobrunneus Budde-Lund (1896 : 47) SYNTYPE : head only (both antennae and mouthparts from right-hand side missing).Reg. no. 1921:10:18:5328. Kappari, Greece. 14.6.1887. Collected by Oertzen. Budde-Lund Collection.philoscoides Budde-Lund (1885 : 175) SYNTYPES: three females. Reg. no. 1911:11:8:10506-10508. Algeria. Collected by E.Simon. A. M. Norman Collection (ex Dollfus collection). TYPES OF TERRESTRIAL ISOPODS 197 SYNTYPES : six females. Reg. no. 1921:10:18:5329-5334. Oned-Zitoun, Algeria. May1883. Collected by E. Simon. Budde-Lund Collection. SYNTYPES: two males ; one female. Reg. no. 1928:12:1:1518-1520. Algeria. Collectedby E. Simon. Stebbing Collection (ex Dollfus collection). SYNTYPES : two females. Reg. no. 1956:10:10:141-142. Oned-Zitoun, Algeria. Col-lected by E. Simon. Presented by University College, Dundee.pruinosus anconanus Verhoeff (ig28c : 144) SYNTYPE : male. Reg. no. 1928:7:4:89. Ancona, Italy. Purchased from K. W. Verhoeff.SYNTYPES : two males ; one female. Reg. no. 1973:152:3. Mid-Italy. Larwood Collec-tion (ex Verhoeff collection).pruinosus meleagris Budde-Lund (1885 : 168) [Metoponorthus meleagris] SYNTYPES: two males ; seven females. Reg. no. 1921:10:18:5279-5287. France ('GalliaMeridionalis'). Collected by E. Simon. Budde-Lund Collection.sabuleti Budde-Lund (1885 : 186) SYNTYPES : one male; one female. Reg. no. 1921:10:18:3920-3921. Biskra, Algeria.Budde-Lund Collection. SYNTYPES: two females. Reg. no. 1921:10:18:3922-3923. Algeria. Budde-Lund Collec-tion. SYNTYPE : male. Reg. no. 1921:10:18:4735. Biskra Mt, Algeria. Collected by Tat-schanowski. Budde-Lund Collection.sexfasciatus Budde-Lund (1885 : 167) SYNTYPES : five females. Reg. no. 1921:10:18:5306-5310. Algiers. Collected by E.Simon. Budde-Lund Collection.sexfasciatus asifensis Verhoeff (i938b : 63) [Metoponorthus asifensis] SYNTYPE : female. Reg. no. 1938:7:7:45. Atlas, Morocco. Purchased from K. W.Verhoeff.virgatus Budde-Lund (1885 : 182) SYNTYPES: two males; nine females. Reg. no. 1921:10:18:5514-5525. Florida, N.America. Collected by Leuckart. Budde-Lund Collection.viridis Budde-Lund (1885 : 179) SYNTYPES : three males. Reg. no. 1921:10:18:5526-5527. Bona, Algeria. Collected byE. Meinert. Budde-Lund Collection. MICA Budde-Lund tardus (Budde-Lund) (1885 : 305) [Porcellio tardus] SYNTYPE: female. Reg. no. 1911:11:8:10533. Tlemcen, Algeria. Collected by E.Simon. Norman Collection. PORCELLIO Latreille acustiserra Barnard (1940 : 358) SYNTYPES: thirteen males; twenty females. Reg. no. 1949:2:2:138-162. Lakes ofAddas, shore of Hora Harsadi, Ethiopia, 7000 ft. 3.12.1936. Collected and presented by J.Omer-Cooper.aegaeus Verhoeff (igo7a : 257) SYNTYPE : female. Reg. no. 1907:11:4:44. Naxos I., Greece. Purchased from K. W.Verhoeff.albicornis (Dollfus) (i8g6b : 3) [Lucasius albicornis] SYNTYPE: male. Reg. no. 1911:11:8:10529. Ficuzza, Palermo, Sicily. 900 m. NormanCollection (ex Dollfus collection). SYNTYPES : not separated from other (non-type) specimens. Reg. no. 1921:10:18:5163-5172 (part). Ficuzza, Palermo, Italy. Budde-Lund Collection (ex Dollfus collection). 198 R. J. LINCOLN AND J. P. ELLIS albinus Budde-Lund (1885 : 142) SYNTYPE : male. Reg. no. 1921:10:18:5163. Onargla, Sahara, Algeria. Collected by E.Simon. Budde-Lund Collection.albomarginatus Vogl (1875 : 516) SYNTYPES : four males. Reg. no. 1925:7:22:449-451. Syra I., Cyclades, Greece. Col-lected by Ebner. Koch Collection.(illuaudi Dollfus (1893 : 52) SYNTYPES : two males. Reg. no. 1921:10:18:4249-4250. Canary Is. Collected by C.Alluaud. Budde-Lund Collection.atnoenus Dollfus (i829b : 178) [Transferred to Porcellio violaceus Budde-Lund] SYNTYPES: two males; one female. Reg. no. 1911:11:8:10396-10398. Cuenca, Spain.Norman Collection.angustulus Budde-Lund (1885 : 146) SYNTYPE: female. Reg. no. 1911:11:8:10429. Bou-Saada, Algeria. Collected by ESimon. Norman Collection (ex Dollfus collection). SYNTYPES : one male; four females. Reg. no. 1921:10:18:3792-3796. Algeria. Col-lected by E. Simon. Budde-Lund Collection.ater Budde-Lund (1896 : 45) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:4284-4285. Rothenthurm Pass,Transylvania, Rumania. Budde-Lund Collection.auritus Budde-Lund (1885 : 126) SYNTYPE : female. Reg. no. 1921:10:18:4978. Seville, Spain. Collected by Fr. Meinert.Budde-Lund Collection.barroisi Dollfus (i8g2c : 127) SYNTYPE : female. Reg. no. 1911:11:8:10400. 'Ouadys de la Mer Morte.' Collected byT. Barrois. Norman Collection (ex Dollfus collection)batesoni Collinge (i9i5b : 461) PARATYPE : male. Reg. no. 1919:4:26:540. S. Spain, 1894. Collected by W. Bateson.Collinge Collection.bistriatus Budde-Lund (1885 : 88) SYNTYPE : male. Reg. no. 1921:10:18:4288. Istanbul, Turkey. Collected by Jelski.Budde-Lund Collection.blattarius Budde-Lund (1885 : 131) SYNTYPE : male. Reg. no. 1921:10:18:4286. Algeria. Budde-Lund Collection.brevipennis Budde-Lund (1885 : 97) SYNTYPES : one male; three females. Reg. no. 1921:10:18:3798-3801. Algeria. Col-lected by E. Simon. Budde-Lund Collection.btuldelundi Simon (1885 : 10) SYNTYPE : female. Reg. no. 1911:11:8:10428. Tunisia. Norman Collection (ex Dollfuscollection). SYNTYPES: two males ; one female. Reg.no. 1921:10:19:4979-4981. Tunisia. Budde-Lund Collection (ex Simon collection).calrnani Omer-Cooper (1923 : 399) SYNTYPES: two females. Reg. no. 1922:5:18:4-5. Baku, Azerbaydzhan, USSR. Col-lected by P. A. Buxton. Presented by J. Omer-Cooper.canariensis Dollfus (1893 : 5°) SYNTYPE: male. Reg. no. 1911:11:8:10409. Canary Is. Collected by C. Alluaud.Normal Collection (ex Dollfus collection). SYNTYPES : two males; one female; one juvenile. Reg. no. 1921:10:18:4289-4291.Canary Is. Collected by C. Alluaud. Budde-Lund Collection.carthaginensis Silvestri (1897 : 415) [Transferred to Porcellio olivieri Audouin] SYNTYPES: four males; seven females. Reg. no. 1921:10:18:3802-3811. Carthage(ruins), Tunisia. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 199 cayennensis Miers (iSyyb : 667) [Transferred to Porcellio scaber Latreille] SYNTYPES : two females. Reg. no. 1941:6:27:15. Cayenne, Guiana. Presented by Prof.Wrzesniowsky.cognatus L. Koch (1901 : 52) SYNTYPE : female. Reg. no. 1925:7:22:453-454 (part). Meran Molde. Koch Collection.SYNTYPE : female. Reg. no. 1925:7:22:453-454 (part). Ratzes. Koch Collection.contractus Dollfus (i8g2c : 8) SYNTYPE: male. Reg. no. 1892:12:6:1. Syria. Presented by W. D'Arcy Thompson.SYNTYPES: one male; one female. Reg.no. 1911:11:8:10384-10385. Syria. Collectedby L. Bleuse. Norman Collection. SYNTYPES: two males. Reg. no. 1928:12:1:1464-1465. Saida, Syria. Collected by L.Bleuse. Stebbing Collection.cruentatus L. Koch (1901 : 37) SYNTYPES : one male ; one female. Reg. no. 1925:7:22:479-480. Sugenheim, W. Ger-many. Koch Collection.debueni Dollfus (i892b : 176) SYNTYPES: three females. Reg. no. 1911:11:8:10393-10395. Villa Rutis, Coruna, Spain.Collected by Bolivar. Norman Collection. SYNTYPE : female. Reg. no. 1921:10:18:4360. Coruna, Spain. Collected by Bolivar.Budde-Lund Collection.dispar Verhoeff (190id : 407) SYNTYPES: one male; two females. Reg. no. 1901:9:19:86-88. Coimbra, Portugal.Collected by Moller. Purchased from K. W. Verhoeff. SYNTYPE: female. Reg. no. 1907:11:4:72. Coimbra, Portugal. Collected by Moller.Purchased from K. W. Verhoeff.evansi Omer-Cooper (1923 : 398) HOLOTYPE : male. Paratypes : one male; one female. Reg. no. 1922: 5:18:1-3. Amara,Mesopotamia, Iraq. Collected by C. L. Boulenger. Presented by J. Omer-Cooper. PARATYPES : micropreparations of appendages: two males; five females. Reg. no.1922:5:18:44-50. Amara, Iraq. Collected by C. L. Boulenger. Presented by J. Omer-Cooper.ficorutn Verhoeff (i93ia : 532) SYNTYPE : male. Reg. no. 1931:4:27:57. Monte Massoncello, Italy. Purchased fromK. W. Verhoeff.ficulneus Budde-Lund (1885 : 98) SYNTYPES : two males ; two females. Reg. no. 1921:10:18:4995-4998. Syria. Collectedby E. Simon. Budde-Lund Collection. SYNTYPES : two females. Reg. no. 1956:10:10:228-229. Syria. Collected by E. Simon.Presented by University College, Dundee.flavocinctus Budde-Lund (1885 : 109) SYNTYPES : two females. Reg. no. 1956:10:10:226-227. Spain. Collected by L. Lundand Fr. Meinert. Presented by University College, Dundee.flavovittata Miers (i877b : 669) [Transferred to Metoponorthus pruinosus (Brandt)] SYNTYPES : four females. Reg. no. 1879:21. Cayenne, Guiana. Presented by Prof.Wrzesniowsky.gallicus Dollfus (1903 : 63) SYNTYPES : two females. Reg. no. 1907:11:4:73-74. Pyr6n6es. Purchased from K. W.Verhoeff.gemtnulatus Dana (1853 : 725) [Transferred to Porcellio scaber Latreille] SYNTYPES : five males; one female. Reg. no. 1877:3. San Francisco, N. America.Presented by W. N. Lockington.graniger Miers (1876 : 226) HOLOTYPE : female. Reg. no. 1973:512:1 (original registration no. 1854:4). NewZealand. Presented by Col. Bolton. R. J. LINCOLN AND J. P. ELLIS Reg. no. 1921:10:18:3812. Spain. Collected by Reg. no. 1921:10:18:4425- Spain. Collected by Fr. Bona, Algeria. Collected by Fr. Meinert. Reg. no. 1921:10:18:4632-4633. 'Ordus.' Pre- Pre- granuliferus Budde-Lund (1885 : 128) SYNTYPES : one female ; one juvenile.L. Lund. Budde-Lund Collection.imbutus Budde-Lund (1885 : 145) SYNTYPES : not separated from other (non-type) specimens.4436 (part). Sicily. Budde-Lund Collection.imbutus pellegrinensis Verhoeff (igoSa : 366) SYNTYPE : male. Reg. no. 1921:6:10:32. Sicily. Purchased from K. W. Verhoeff.incanus Budde-Lund (1885 : 102) SYNTYPES : two females. Reg. no. 1921:10:18:4437-4438.Meinert. Budde-Lund Collection.intercalarius Budde-Lund (1885 : 104) SYNTYPE : female. Reg. no. 1921:10:18:4439.Budde-Lund Collection.interpolator Budde-Lund (1885 : 93) SYNTYPES : specimens coated with mould.Budde-Lund Collection.jelskii Miers (iSyyb : 668) [Transferred to Metoponorthus pruinosus Brandt] SYNTYPES: one male; two females. Reg. no. 1879:21. Maraynioe (?), Guiana,sented by Prof. Wrzesniowsky. SYNTYPES: three males; three females. Reg. no. 1879:21. Pumamarca, Peru,sented by Prof. Wrzesniowsky. SYNTYPES : nine males ; ten females. Reg. no. 1887:20. Caracas, Guiana. Presentedby Dr Ernst.latissimus Budde-Lund (1885 : 95) [Transferred to Porcellio spatulatus (Costa)] SYNTYPES: three males; five females. Reg. no. 1921:10:18:4988-4992. Corsica (collectedby E. Simon) and Sardinia (collected by Keitel). Budde-Lund Collection.lepineyi Verhoeff (i937b : 305) SYNTYPES: one male; three females. Reg. no. 1938:7:7:7-10. Haute Atlas, Morocco.Collected by J. de Le'piney. Purchased from K. W. Verhoeff.longicauda Budde-Lund (1885 : 112) SYNTYPE : male. Reg. no. 1911:11:8:10358. Algeria. Collected by Fr. Meinert. NormanCollection. SYNTYPES: two females. Reg. no. 1911:11:8:10359-10360. Algeria. Collected by E.Simon. Norman Collection. SYNTYPES : one male ; five females.Lund Collection. SYNTYPES : one male ; one female,by University College, Dundee.longipennis Budde-Lund (1885 : 91) SYNTYPES : two females. Reg. no. 1921:10:18:4475-4476.Collected by Nordmann. Budde-Lund Collection.maculipes Budde-Lund (1885 : 105) SYNTYPES: two males; two females. Reg. no. 1921:10:18:4488-4491.Collected by Sorensen. Budde-Lund Collection.magnificus Dollfus (i8g2b : 170) SYNTYPE: female. Reg. no. 1911:11:8:10430. Alcazaba, Almeria, Spain.R. Nickles. Norman Collection (ex Dollfus collection). SYNTYPE : female. Reg. no. 1921:10:18:4492. Alcazaba, Almeria, Spain.R. Nickles. Budde-Lund Collection (ex Dollfus collection).tnarginenotatus Budde-Lund (1885 : 145) SYNTYPE : male. Reg. no. 1921:10:18:4500. Bona, Algeria. Collected by Fr. Meinert.Budde-Lund Collection. Reg. no. 1921:10:18:4466-4471. Algeria. Budde-Reg. no. 1956:10:10:230-231. Algeria. Presented Avhasia, Caucasus Mts.Madeira I. Collected byCollected by TYPES OF TERRESTRIAL ISOPODS 201 SYNTYPE : female. Reg. no. 1956:10:10:223. Bona or Constantine, Algeria. Collectedby Fr. Meinert. Presented by University College, Dundee.mildei L. Koch (1901 : 493) HOLOTYPE : male. Reg. no. 1925:7:22:493. Meran. Collected by Milde. Koch Collection.moebiusi Verhoeff (igoia : 38) [Transferred to Porcellio lamellatus (Ulj.) Budde-Lund] SYNTYPES : two females. Reg. no. 1901:9:19:95-96. Dalmatia, Yugoslavia. Purchasedfrom K. W. Verhoeff.narentanus Verhoeff (igoya : 251) SYNTYPE : male. Reg. no. 1907:11:4:84. Herzegowina, Yugoslavia. Purchased fromK. W. Verhoeff.naupliensis Verhoeff (igoid : 403) SYNTYPES: one male; one female; one juvenile. Reg. no. 1901:9:19:60-61. Greece.Purchased from K. W. Verhoeff.nicklesi Dollfus (i8g2b : 171) SYNTYPE : male. Reg. no. 1921:10:18:4582. Rio Clariano, Valencia, Spain. Collectedby R. Nickles. Budde-Lund Collection.nigra Say (1818 : 432) [Transferred to Porcellio scaber Latr.] SYNTYPE : fragments only (dry). Reg. no. 1973:509:1. Pennsylvania, USA. Presentedby T. Say.normani (Dollfus) (1899 : 67) [Lucasius normani] SYNTYPES : one male; two females. Reg. no. 1911:11:8:10535-10537. Ilheo dosEmbarcadouros, Madeira I. 1897. Collected by A. M. Norman. Norman Collection.obsoletus Budde-Lund (1885 : 116) SYNTYPES : three males; five females. Reg. no. 1921:10:18:5002-5007. Sebastopol,Ukraine, USSR. Budde-Lund Collection.obtusifrons Haswell (1882 : 280) [Transferred to Porcellio laevis Latreille] SYNTYPES : two females. Reg. no. 1895:11:14:38-39. Sydney, Australia. Presentedby the Australian Museum, Sydney.obtusiserra Barnard (1940 : 357) SYNTYPES: six males; one female; one juvenile. Reg. no. 1949:2:2:118-137 (part).Addis Ababa, Ethiopia. 16.9.1926. Collected and presented by J. Omer-Cooper. SYNTYPES: one male; one female. Reg. no. 1949:2:2:118-137 (part). D j em Dj em Forest,Ethiopia. 25.9.1926. Collected and presented by J. Omer-Cooper. SYNTYPES: nine males ; twenty females. Reg. no. 1949:2:2:118-137 (part). Djem DjemForest, Ethiopia. Pond no. i, 8100 ft. 21.9.1926. Collected and presented by J. Omer-Cooper.ocellatus Budde-Lund (1885 : in) SYNTYPES: four males; three females. Reg. no. 1921:10:18:4600-4606. Spain ('His-pania meridionalis'). Collected by L. Lund. Budde-Lund Collection.or arum or arum Verhoeff (1910 : 139) {Porcellio lugubris or arum] SYNTYPES : one male ; four females. Reg. no. 1921:6:10:23-27. Riviera, France. Pur-chased from K. W. Verhoeff.orarum vizzavonensis Verhoeff (i928c : 123) \_Porcellio lugubris vizzavonensis] HOLOTYPE : male. Reg. no. 1928:7:4:12. Vizzavona, Corsica. Purchased from K. W.Verhoeff.parietinus L. Koch (1901 : 49) [Transferred to Porcellio scaber Latreille] SYNTYPES : one male ; two females. Reg. no. 1925:7:22:496-498. Landstuhl, W. Ger-many. Koch Collection.parvus Budde-Lund (1885 : 132) SYNTYPES : three females. Reg. no. 1921:10:18:5026-5027. Sicily. Collected by Groh-mann. Budde-Lund Collection.pauper Budde-Lund (1885 : 305) SYNTYPE : specimen in fragments. Reg. no. 1921:10:18:5845. Tlemcen, Algeria. Col-lected by E. Simon. Budde-Lund Collection. 202 R. J. LINCOLN AND J. P. ELLIS planarius Budde-Lund (1885 : 95) SYNTYPE : male. Reg. no. 1921:10:18:4993. Sicily. Collected by Grohmann. Budde-Lund Collection.praeustus Budde-Lund (1885 : 302) SYNTYPE : male. Reg. no. 1921:10:18:4994. Bon-Merzong, Algeria. Collected by E.Simon. Budde-Lund Collection.provincialis Aubert & Dollfus (1890 : 7) SYNTYPE: male. Reg. No. 1911:11:8:10402. 'Galets de la Crau', Marseilles, France.Norman Collection (ex Dollfus collection). pujetanus Verhoefif (1910 : 136) [Transferred to Porcellio spinipennis spinipennis Budde-Lund]SYNTYPE : female. Reg. no. 1921:6:10:22. French Riviera. Purchased from K. W.Verhoeff.purpureus Budde-Lund (1885 : 303) SYNTYPE : male. Reg. no. 1911:11:8:10422. Oran, Algeria. Collected by E. Simon.Norman Collection (ex Dollfus collection).pyrenaeus Dollfus (i8g2b : 175) SYNTYPES : one female; one juvenile. Reg. no. 1911:11:8:10407-10408. Vallee deSallanques, Spanish Pyrenees. 1200 m. Norman Collection (ex Dollfus collection).ragusae Dollfus (i8g6b : 2) SYNTYPES : two females. Reg. no. 1911:11:8:10391-10392. Selinunte (ruins), Sicily.Norman Collection (ex Dollfus collection). SYNTYPES: one male ; one female. Reg. no. 1921:10:18:4702-4703. Ficuzza, near Paler-mo, Italy. Budde-Lund Collection (ex Dollfus collection) SYNTYPE : female. Reg. no. 1921:10:18:4711. Ficuzza, Sicily. 15.2.1889. Budde-LundCollection.sabulifer Verhoeff (igo7a : 265) SYNTYPE : female. Reg. no. 1907:11:4:100. Attika. Purchased from K. W. Verhoeff.saltuum L. Koch (1901 : 45) [Transferred to Porcellio arcuatus Budde-Lund] SYNTYPE : female. Reg. no. 1911:11:8:10431. Tyrol. Norman Collection (ex Kochcollection). SYNTYPES: one male; one female. Reg. no. 1925:7:22:623-639 (part). Vahrn, Brixen,Austria. Koch Collection. SYNTYPES : one male ; two females. Reg. no. 1925:7:22:623-638 (part). Meran, Trentino,Italy. Koch Collection. SYNTYPES: two males; four females. Reg. no. 1925:7:22:623-638 (part). Ratzes. KochCollection. SYNTYPES : one male; two females. Reg. no. 1925:7:22:623-638 (part). Bosen, W,Germany. Koch Collection. SYNTYPE: female. Reg. no. 1925:7:22:623-638 (part). Torbole, Italy. Koch Collection.SYNTYPE : female. Reg. no. 1925:7:22:623-638 (part). Bayerische Alpen, W. Germany.Koch Collection.sarajevensis Verhoeff (igo7a : 255) SYNTYPE : male. Reg. no. 1907:11:4:101. Boznya, USSR. Purchased from K. W. Verhoeff.sarculatus Budde-Lund (1896 : 46) SYNTYPES : fragments of two males. Reg. no. 1921:10:18:4736. Ukraine Peninsula.Budde-Lund Collection.scaber japonicus Verhoeff (ig28b : 36) SYNTYPE: male. Reg. no. 1928:7:4:10. Hokkaido, Japan. 10.11.1904. Collected byH. Sauter. Purchased from K. W. Verhoeff.scaber lusitanus Verhoeff (i907a : 263) [Porcellio lusitanus] SYNTYPE: female. Reg. no. 1907:11:4:80. Portugal. Purchased from K. W. Verhoeff.simulator Budde-Lund (1885 : 147) SYNTYPES: three males; one female. Reg. no. 1911:11:8:10418-10421. Algeria.Collected by E. Simon. Norman Collection (ex Dollfus collection). TYPES OF TERRESTRIAL ISOPODS 203 SYNTYPES : one male ; three females. Reg. no. 1921:10:18:4882-4886. Algeria. Col-lected by E. Simon. Budde-Lund Collection.sociabilis L. Koch (1901 : 51) SYNTYPES : one male ; two females. Reg. no. 1911:11:8:10432-10434. Tyrol. NormanCollection (ex Koch collection). SYNTYPES: ten males; eight females. Reg. no. 1925:7:22:661-670. Vahrn, Brixen,Austria. Koch Collection.sordidus Budde-Lund (1885 : 108) SYNTYPE : female (?) (specimen in fragments). Reg. no. 1921:10:18:4888. Balearic Is.Collected by Schaufuss. Budde-Lund Collection.spatulata Barnard (1940 : 357) SYNTYPES : twenty-four males ; thirty-eight females. Reg. no. 1949:2:2:88-117. DjemDjem Forest, Ethiopia. Pond no. 2, 8000 ft. 26.9.1926. Collected and presented by J.Omer-Cooper. SYNTYPES: five males; ten females. Reg. no. 1949:2:2:88-177 (part). Djem Djem Forest,Ethiopia. Pond no. i, 8100 ft. 21.9.1926. Collected and presented by J. Omer-Cooper. SYNTYPE : female. Reg. no. 1949:2:2:88-117 (part). Djem Djem Forest, Ethiopia.22.9.1926. Collected and presented by J. Omer-Cooper. SYNTYPES: three males; twelve females. Reg. no. 1949:2:2:88-117 (part). Djem DjemForest, Ethiopia. 25.9.1926. Collected and presented by J. Omer-Cooper.spinicornis Say (1818 : 431) SYNTYPES : two males. Reg. no. 1973:508:2. North America. Presented by T. Say.spinipennis montanus Budde-Lund (1885 : 121) \Porcellio montanus] SYNTYPES : one male ; four (?) females (specimens in fragments). Reg. no. 1921:10:18:4503-4506. France. Collected by E. Simon. Budde-Lund Collection.spinipennis spinipennis Budde-Lund (1885 : 732) [Porcellio spinipennis] SYNTYPES : two females. Reg. no. 1921:10:18:4889-4890. Menton, France. Collectedby E. Simon. Budde-Lund Collection.spinipes Dollfus (1893 : 53) SYNTYPE: male. Reg. no. 1921:10:18:4892. Canary Is. Collected by C. Alluaud.Budde-Lund Collection.spretus Budde-Lund (1885 : 89) SYNTYPE: female. Reg. no. 1921:10:18:4891. Sicily. Collected by Grohmann. Budde-Lund Collection.succinctus Budde-Lund (1885 : 304) SYNTYPE: female. Reg. no. 1911:11:8:10364. Carthagena, Spain. May 1883. Col-lected by E. Simon. Norman Collection (ex Dollfus collection). SYNTYPES: one male; two females. Reg. no. 1921:10:18:4893-4895. Carthagena,Spain. May 1883. Collected by E. Simon. Budde-Lund Collection.trachealis Budde-Lund (1885 : 90) SYNTYPE: female. Reg. no. 1921:10:18:4896. Moldavia. Collected by Jelski. Budde-Lund Collection.transmutatus Budde-Lund (1885 : 122) SYNTYPES: four males; one female. Reg. no. 1911:11:8:10436-10440. Algeria. Col-lected by Fr. Meinert. Norman Collection. SYNTYPES : five males; twelve females. Reg. no. 1921:10:18:3924-3940. Algeria.Budde-Lund Collection.trilobatus Stein (1859 : 260) SYNTYPE : male. Reg. no. 1921:10:18:4947. Mehadia, Roumania. Budde-Lund Collec-tion (ex Stein collection). SYNTYPES: one male; one female. Reg. no. 1925:7:22:672-675 (part). Mehadia,Roumania. Koch Collection (ex Stein collection).vespertilio Budde-Lund (1896 : 46) SYNTYPE: female. Reg.no. 1921:10:18:4954. Dalmatia, Yugoslavia. Budde-Lund Collection. 204 R- J- LINCOLN AND J. P. ELLIS violaceus Budde-Lund (1885 : 117) SYNTYPE : female. Reg. no. 1921:10:18:4952-4953 (part). Murviedro, Spain. Collectedby Collin. Budde-Lund Collection.yemenensis Barnard (1941 : 57) SYNTYPE: juvenile. Reg. no. 1949:2:2:1-25 (part). Western Aden Protectorate : Dhala.4800 ft, in detritus under succulent plants. 14.9.1937. Collected by H. Scott. BritishMuseum Expedition to S.W. Arabia, 1937-38. SYNTYPES : two males; one female. Reg. no. 1949:2:2:1-25 (part). Yemen: JebelMasnah, S.W. of Ma'bar. c. 8400 ft. 9.3.1938. Collected by H. Scott and E. B. Britton.British Museum Expedition to S.W. Arabia, 1937-38. SYNTYPES: four males; one female. Reg. no. 1949:2:2:1-25 (part). Western AdenProtectorate: Jebel Jihaf. From summit, 7800 ft, under stones. 20.9.1937. Collected byH. Scott. British Museum Expedition to S.W. Arabia, 1937-38. SYNTYPES: one male ; two females. Reg. no. 1949:2:2:1-25 (part). Yemen: Ghaiman.13 miles S.E. of San'a, c. 8500 ft, found in valley near streams. 17.2.1938. Collected by H.Scott. British Museum Expedition to S.W. Arabia, 1937-38. SYNTYPES : five females. Reg. no. 1949:2:2:1-25 (part). Yemen : Birkat al Bu'r, nearHaz. About 16 miles N.W. of San'a, 9200 ft. 4.2.1938. Collected by E. B. Britton. BritishMuseum Expedition to S.W. Arabia, 1937-38. SYNTYPES: two males; four females; three juveniles. Reg. no. 1949:2:2:1-25 (part).Yemen: top of Al Errein, near Haz. 9300 ft. 3.2.1938. British Museum Expedition toS.W. Arabia, 1937-38. SYNTYPES: two males; two females. Reg. no. 1949:2:2:1-25 (part). Yemen: top ofpass over Jebel Sumara. 9200-9700 ft. 2-3.1.1938. Collected by E. B. Britton. BritishMuseum Expedition to S.W. Arabia, 1937-38.zealandicus Miers (1876 : 226) [Transferred to Metoponorthus pruinosus Brandt] HOLOTYPE : female (dry). Reg. no. 1845:30. New Zealand. Purchased from Mr Earl. PORCELLIONIDES Miers dimorphus Jackson (1926 : 186) SYNTYPES: three males; three females. Reg. no. 1925:10:14:28-32. Mont Estoril,near Lisbon, Portugal. Collected and presented by E. R. Speyer. SYNTYPES: two males ; five females. Reg.no. 1952:4:18:153-159. Mont Estoril, nearLisbon, Portugal. Collected by E. R. Speyer. Jackson Collection.variabilis Jackson (1926 : 187) SYNTYPES: four males ; twelve females. Reg. no. 1925:10:14:34-43. Mont Estoril, nr.Lisbon, Portugal. February 1925. Collected and presented by E. R. Speyer. SYNTYPES : two males; eleven females. Reg. no. 1952:4:18:160-169. Mont Estoril,near Lisbon, Portugal. February 1925. Collected by E. R. Speyer. Jackson Collection. TURA Budde-Lund albipennis Budde-Lund (i9i3b : 379) SYNTYPES: two males. Reg. no. 1921:10:18:3983-3984. Ethiopia. Collected by E.Wache. Budde-Lund Collection.angusta Budde-Lund (1913b : 378) SYNTYPES: one male; nine females. Reg. no. 1921:10:18:3985-3994. Aldabra I.,Indian Ocean. Collected by H. P. Thomasset. Budde-Lund Collection.laticauda Budde-Lund (i9i3b : 379) SYNTYPE : male. Reg. no. 1921:10:18:3995. Ethiopia. Budde-Lund Collection.testacea Budde-Lund (1908 : 282) SYNTYPES: six males; seventeen females. Reg. no. 1921:10:18:3996-4007. Majunga,Madagascar. 1894. Collected by A. Voeltzkow. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 205 URAMBA Budde-Lund marginalis Budde-Lund (1910 : 19) SYNTYPES : six males; three females. Reg. no. 1921:10:18:4070-4078. Tanganyika:Kibonoto, Kilimanjaro ; Ngara na nyuki, Meru, in acacia forest. January 1906. Budde-Lund Collection.mus (Budde-Lund) (1898 : 9) [Lyprobius mus] SYNTYPES : one male; one specimen in fragments. Reg. no. 1921:10:18:4079-4080.Zanzibar I., East Africa. Collected by A. Voeltzkow. Budde-Lund Collection.triangulifera Budde-Lund (1910 : 18) SYNTYPES : three males ; seven females. Reg. no. 1921:10:18:4085-4094. Kilimanjaro,Tanganyika. Budde-Lund Collection. Family BALLONISGIDAEBALLONISCUS Budde-Lund tnaculata (Budde-Lund) (1885 : 215) [Philoscia maculata] SYNTYPES : three females. Reg. no. 1921:10:18:1708-1710. Buenos Aires (probably SanNicolas), Agentina. Collected by W. S0rensen. Budde-Lund Collection. PLATAONISCUS Vandel borelli (Dollfus) (1897 : 3) [Alloniscus borellii] SYNTYPE : male. Reg. no. 1921:10:18:1701. San Lorenzo, Jujuy, Argentina. Collectedby A. Borelli. Budde-Lund Collection. SYNTYPE : male. Reg. no. 1956:10:10:151. San Lorenzo, Jujuy, Argentina. Collectedby A. Borelli. Presented by University College, Dundee. Family TRAGHELIPIDAE AGNARA Budde-Lund fragilis Budde-Lund (1908 : 287) SYNTYPES: three males; one female. Reg. no. 1921:10:18:5551-5554. Ceylon. January1904. Collected by K. Kraepelin. Budde-Lund Collection.madagascariensis (Budde-Lund) (1885 : 189) [Metoponorthus madagascariensis] SYNTYPES: two males; one female. Reg. no. 1921:10:18:5556-5558. Madagascar. Col-lected by Hildebrandt. Budde-Lund Collection. NAGURUS Holthius formosanus (Verhoeff) (ig28a : 214) [Nagara formosana] SYNTYPE : male. Reg. no. 1928:7:4:7. Taiwan I. Purchased from K. W. Verhoeff.incisus (Verhoeff) (ig28a : 215) [Nagara incisa] SYNTYPE : female. Reg. no. 1928:7:4:8. Taiwan I. Purchased from K. W. Verhoeff.nanus (Budde-Lund) (1908 : 285) [Nagara nana] SYNTYPES: six males; seven females. Reg. no. 1921:10:18:1495-1506. Madagascar.Budde-Lund Collection.travancorius (Verhoeff) (i936b : 100) [Nagara travancoria] SYNTYPES : two females. Reg. no. 1938:7:7:22-23. Deccan, India. Purchased fromK. W. Verhoeff. 206 R. J. LINCOLN AND J. P. ELLIS ORTHOMETOPON Verhoeff dalmatinus frascatensis Verhoeff (1918 : 147) SYNTYPES : one male; one female. Reg. no. 1921:6:10:33-34. S. Italy. Purchasedfrom K. W. Verhoeff.phaleronensis (Verhoeff) (190id : 407) [Porcellio (Metoponorthus) phaleronensis] SYNTYPES : two females. Reg. no. 1901:9:19:42-43. Phaleron, Greece. Purchased fromK. W. Verhoeff. PAGANA Budde-Lund fissifrons Budde-Lund (1908 : 289) SYNTYPES: one male; two females. Reg. no. 1921:10:18:5547-5548. Mauritius I.,Indian Ocean. Budde-Lund Collection.tnaculosa Budde-Lund (1908 : 288) SYNTYPES : two females. Reg. no. 1921:10:18:5549-5550. Mauritius I., Indian Ocean.Budde-Lund Collection. PHALABA Budde-Lund brevis Budde-Lund (1910 : 19) SYNTYPES: two males; twelve females. Reg. no. 1921:10:18:2091-2103. Kibonoto,Kilimanjaro, Tanganyika. Budde-Lund Collection. PORCELLIUM Dahl collicolus (Verhoeff) (i9O7a : 247) [Porcellio (Porcellidium) collicolus'] SYNTYPE : male. Reg. no. 1907:11:4:65. W.Hungary. Purchased from K. W. Verhoeff.herzegovinensis (Verhoeff) (igoib : 69) [Porcellio herzegovinensis] SYNTYPES : one male; two females. Reg. no. 1901:9:19:90-92. Herzegovina, Yugo-slavia. Purchased from K. W. Verhoeff. PROTRACHEONISCUS Verhoeff desertorum Verhoeff (19305 : 112) SYNTYPE : female. Reg. no. 1930:5:26:71. Tashkent, Uzbekistan, USSR. Purchasedfrom K. W. Verhoeff.fontiutn Verhoeff (i93ob : 112) SYNTYPES: one female ; four juveniles. Reg. no. 1930:5:26:65-69. Turkestan, USSR.Purchased from K. W. Verhoeff.hertnagorensis Verhoeff (1927 : 327) SYNTYPE : female. Reg. no. 1931:4:27:47. Karnten, Austria. Purchased from K. W.Verhoeff.hirsutulus Verhoeff (i93ob : 115) SYNTYPE : female. Reg. no. 1930:5:26:70. Tashkent, Uzbekistan, USSR. Purchasedfrom K. W. Verhoeff.latus Verhoeff (i93ob : no) SYNTYPES : two females. Reg. no. 1930:5:26:63-64. Turkestan, USSR. Purchasedfrom K. W. Verhoeff. SYNTYPES : two males; two females. Reg. no. 1938:7:7:26-29. Turkestan, USSR.Purchased from K. W. Verhoeff.politus tnehelyi Kesselyak (1930 : 250) [Protracheoniscus mehelyi] SYNTYPE : female. Reg. no. 1931:4:27:48. Monte Maggiore, Italy. Purchased fromK. W. Verhoeff. TYPES OF TERRESTRIAL ISOPODS 207 saxonicus Verhoeff (1927 : 324) SYNTYPES : one male; one female. Reg. no. 1928:7:4:50-51. Bayern, W. Germany.Purchased from K. W. Verhoeff.saxonicus carpathicus Verhoeff (19280 : 52) SYNTYPE : female. Reg. no. 1928:7:4:52. Carpathian Mts, Central Europe. Purchasedfrom K. W. Verhoeff.taschkententis Verhoeff (i93ob : 113) SYNTYPE : female. Reg. no. 1930:5:26:72. Turkestan, USSR. Purchased from K. W.Verhoeff.venetus Verhoeff (1927 : 328) SYNTYPES: one male; one female. Reg. no. 1928:7:4:53-54. Veneto region, Italy. Pur-chased from K. W. Verhoeff. TRACHELIPUS Budde-Lund apenninorutn (Verhoeff) (i93ia : 538) [Tracheoniscus apenninoruwi] SYNTYPE : female. Reg. no. 1931:4:27:57. Umbria, Italy. Purchased from K. W.Verhoeff. SYNTYPES : three males ; one female. Reg. no. 1937:7:6:35-38. Apennine Mts, Italy.Purchased from K. W. Verhoeff.apulicus (Verhoeff) (1939 : 215) [Tracheoniscus apulicus] SYNTYPES: one male ; one female. Reg. no. 1938:7:7:16-17. Apulia, Italy. Purchasedfrom K. W. Verhoeff.balticus (Verhoeff) (1907^: 258) [Porcellio balticus] SYNTYPE: female. Reg. no. 1907:11:4:63. Brandenburg, E. Germany. Purchased fromK. W. Verhoeff.balticus burzenlandicus (Verhoeff) (i907a : 258) [Porcellio balticus burzenlandicus] SYNTYPE: female. Reg. no. 1907:11:4:64. Siebenbiirgen, Germany. Purchased fromK. W. Verhoeff.brentanus (Verhoeff) (1927 : 332) [Tracheoniscus brentanus] SYNTYPE : female. Reg. no. 1928:7:4:95. Venezia province, Italy. Purchased from K.W. Verhoeff.bulgaricus (Verhoeff) (ig26a : 143) [Tracheoniscus bulgaricus] SYNTYPE : male. Reg. no. 1928:7:4:93. Bulgaria. Collected by I. Buresch. Purchasedfrom K. W. Verhoeff.larii (Verhoeff) (1927 : 331) [Tracheoniscus larii] SYNTYPE : male. Reg. no. 1928:7:4:94. Lake Como, Italy. Purchased from K. W.Verhoeff.larii aternanus (Verhoeff) (193la : 539) [Tracheoniscus larii aternanus] SYNTYPE : female. Reg. no. 1931:4:27:53. L'Aquila, Abruzzi, Italy. Purchased fromK. W. Verhoeff.lutshniki (Verhoeff) (1933 : 107) [Tracheoniscus lutshniki] SYNTYPES : two females. Reg. no. 1938:7:7:20-21. Sotshi, N. Caucasus Mts. July 1931.Collected by V. Lutshnik. Purchased from K. W. Verhoeff.phaeacorum (Verhoeff) (igoib : 71) [Porcellio rathkei phaeacorum] SYNTYPE : male. Reg. no. 1901:9:19:64. Korfu I., Greece. Purchased from K. W.Verhoeff. SYNTYPE: male. Reg. no. 1907:11:4:85. Korfu I., Greece. Purchased from K. W.Verhoeff.pseudoratzeburgi (Verhoeff) (i9O7a : 259) [Porcellio (Euporcellio) pseudoratzeburgi] SYNTYPE: female. Reg. no. 1907:11:4:87. Boznya, USSR. Purchased from K. W.Verhoeff. 208 R. J. LINCOLN AND J. P. ELLIS Family ARMADILLIDIIDAEARMADILLIDIUM Brandt aegaeum Strouhal (1929 : 102) SYNTYPE : female. Reg. no. 1931:4:27:32. Skyros I. May 1927. Collected by F.Werner. Purchased from K. W. Verhoeff.alassiense Verhoeff (1910 : 121) SYNTYPES : four females. Reg. no. 1937:7:6:6-9. Italian Riviera. Purchased fromK. W. Verhoeff.albanicum Verhoeff (i901 a : 37) SYNTYPES : two females. Reg. no. 1901:9:19:15-16. Korfu I., Greece. Purchased fromK. W. Verhoeff. SYNTYPE : male. Reg. no. 1921:10:18:2622. Aulaona, Albania. Collected by Oertzen.Budde-Lund Collection.albifrons L. Koch (1901 : 25) SYNTYPE : female. Reg. no. 1925:7:22:207. Merano, Italy. Collected by Milde. KochCollection.album Dollfus (1887 : 4) SYNTYPE : female. Reg. no. 1921:10:18:2623. Arcachon, France. Budde-Lund Collec-tion.anconanum Verhoeff (i928c : 98) SYNTYPES : two males; one female. Reg. no. 1928:7:4:97-99. Ancona, Italy. Pur-chased from K. W. Verhoeff.apenninorum Verhoeff (i928c : 109) SYNTYPE : male. Reg. no. 1928:7:4:103. Ancona, Italy. Purchased from K. W.Verhoeff.apfelbecki Dollfus (i896d : 584) SYNTYPE : female. Reg. no. 1901:9:19:31. Herzegovina, Yugoslavia. Purchased fromK. W. Verhoeff. SYNTYPES: two females. Reg.no. 1911:11:8:9956-9957. Stolac, Yugoslavia. NormanCollection.argentarium Verhoeff (1931 a : 518) SYNTYPES : two females. Reg. no. 1931:4:27:39-40. Monte Argentario, Italy. Pur-chased from K. W. Verhoeff. SYNTYPE : female. Reg. no. 1970:76:1. Monte Argentario, Italy. Larwood Collection(ex Verhoeff collection).astriger (C. L. Koch) (1841 : 4) [Armadillo astriger] SYNTYPE (?) : one female. Reg. no. 1925:7:22:208. Dalmatia, Yugoslavia. KochCollection.badium Budde-Lund (1885 : 60) SYNTYPES: two males ; one female. Reg. no. 1921:10:18:2661-2663. Egypt. Collectedby Fischer. Budde-Lund Collection.badium siculorum Verhoeff (igoSc : 491) SYNTYPES: two males; two females. Reg. no. 1921:6:10:14-17. Palermo region,Sicily. Purchased from K. W. Verhoeff.caelatum Miers (i877b : 665) SYNTYPES : two males ; one female. Reg. no. 1879:21. Cayenne, Guiana. Presented byProf. A. Wrzesniowsky.carniolense Verhoeff (igoib : 67) SYNTYPES: one male ; one female; one juvenile. Reg. no. 1928:7:4:100-102. Adelsberg,Krain (Carniola), Austria. Purchased from K. W. Verhoeff.cingendum Verhoeff (1910 : 119) [Armadillidium maculatum cingendum] SYNTYPES: one male ; two females. Reg.no. 1921:6:10:11-13. Italian Riviera. Pur-chased from K. W. Verhoeff. TYPES OF TERRESTRIAL ISOPODS 209 SYNTYPE : male. Reg. no. 1970:32:1. Italian Riviera. Larwood Collection (ex Verhoeffcollection).corcyraeum Verhoeff (igoib : 68) SYNTYPES : two males. Reg. no. 1901:9:19:18-19. Corfu I., Greece. Purchased fromK. W. Verhoeff. SYNTYPE : specimen in fragments. Reg. no. 1921:10:18:2713. Corfu I., Greece. Budde-Lund Collection.davidi Dollfus (1887 : 4) SYNTYPE: female. Reg. no. 1911:11:8:9930. Syria. Collected by Abbe David. NormanCollection.elbanum Verhoeff (i93ia : 522) [Transferred to Armadillidium assimile Budde-Lund] SYNTYPES : one male ; one juvenile ; one specimen with genitalia missing. Reg. no.1931:4:27:36-37. Elba. Purchased from K. W. Verhoeff. SYNTYPES : one male ; one female. Reg. no. 1970:77:2. Elba. Larwood Collection (exVerhoeff collection).esterelanum Dollfus (1887 : 2) SYNTYPE : female. Reg. no. 1892:12:6:4. Esterel, France. Presented by. W. D'ArcyThompson. SYNTYPE: female. Reg. no. 1911:11:8:9980. Mt Vinaigre, Esterel Mts, France. NormanCollection. SYNTYPES: two males; two females. Reg. no. 1921:10:18:2839-2841. Esterel Mts,France. Budde-Lund Collection. SYNTYPES: one male ; one female. Reg. no. 1921:12:1:1422-1423. Esterel Mts, France.Stebbing Collection. SYNTYPE: female. Reg. no. 1956:10:10:181. Esterel Mts, France. Presented by Univer-sity College, Dundee.flavoscutatum Strouhal (1927 : 28) SYNTYPE : female. Reg. no. 1928:7:4:115. Shkoder (formerly Skutari), Albania. Pur-chased from K. W. Verhoeff.frontexcavatum Verhoeff (i93ia : 517) SYNTYPE : male. Reg. no. 1931:4:27:35. Umbria, Italy. Purchased from K. W.Verhoeff. SYNTYPE : male. Reg. no. 1970:78:1. Umbria, Italy. Larwood Collection (ex Verhoeffcollection).frontirostre Budde-Lund (1885 : 52) SYNTYPES: one male; two females; one juvenile. Reg. no. 1921:10:18:2863-2866.Italy. Collected by J. Collin. Budde-Lund Collection.furcatum Budde-Lund (1885 : 72) SYNTYPES : one male; one female. Reg. no. 1921:10:18:2885-2886. Rome, Italy.Budde-Lund Collection.gestroi Tua (1900 : 5) SYNTYPES : six males; six females. Reg. no. 1921:10:18:2887-2898. Finalmarina.Collected by R. Gestro. Budde-Lund Collection.graecorum Verhoeff (igo7b : 474) SYNTYPE : male. Reg. no. 1907:11:4:2. Mid-Greece. Purchased from K. W. Verhoeff.grandinatum Budde-Lund (1885 : 58) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:2899-2900. Sicily. Collectedby Grohmann. Budde-Lund Collection.granulatum peloponnesiaca Verhoeff (igoic : 139) SYNTYPE : female. Reg. no. 1901:9:19:1. Navplion, Greece. Purchased from K. W.Verhoeff.hirtutn Budde-Lund (1885 : 64) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:2944-2945. Sicily. Collectedby Grohmann. Budde-Lund Collection. 2io R. J. LINCOLN AND J. P. ELLIS laminigerutn Verhoefif (igojb : 481) SYNTYPE : male. Reg. no. 1907:11:4:4. Middle Greece. Purchased from K. W. Verhoeff.mareoticum Budde-Lund (1885 : 59) SYNTYPE : female. Reg. no. 1921:10:18:2992. Egypt. Collected by Fischer. Budde-Lund Collection.marinensium Verhoeff (19280 : 105) SYNTYPE : female. Reg. no. 1928:7:4:116. San Marino Republic, Italy. Purchasedfrom K. W. Verhoeff.tnohatnedanicutn Verhoeff (1929 : 131) SYNTYPE : female. Reg. no. 1928:7:4:104. Kuru Dag Mt, Gelibolu, Thrace region,Turkey. Collected by I. Buresch. Purchased from K. W. Verhoeff.muricatum Budde-Lund (1885 : 297) SYNTYPES : two males ; eleven females. (This includes unregistered specimens collectedby Ehlers in 1879, which are in the same tube.) Reg. no. 1921:10:18:3038-3042. Cartagena,Spain. Collected by E. Simon. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1911:11:8:9958-9959. Cartagena, Spain. Collected byE. Simon. Norman Collection.nasatum sorrentinum Verhoeff (igoSc : 487) SYNTYPES: one male; four females. Reg. no. 1921:6:10:4-9. Southern Italy. Pur-chased from K. W. Verhoeff.nitidulits Collinge (191 $b : 463) [Transferred to Armadillidium vulgare (Latreille)] SYNTYPE : female. Reg. no. 1919:4:26:503. Madeira, Spain. Collected by J. W. Clark.Collinge Collection.odherni Verhoeff (i93oc : 45) [Armadillidium peraccai odherni] SYNTYPES: two males ; two females. Reg. no. 1930:5:26:6-11. Dalmatia. Purchasedfrom K. W. Verhoeff. SYNTYPES: three males; four females. Reg. no. 1937:7:6:1-5. Yugoslavia. Purchasedfrom K. W. Verhoeff.oliveti L. Koch (1901 : 31) [Transferred to Armadillidium vulgare (Latreille)] SYNTYPES: two males. Reg. no. 1911:11:8:9998-9999. Tirol, Austria. NormanCollection. SYNTYPES: five males; nine females. Reg. no. 1925:7:22:281-297 (part). Nago, nr.Torbole, Italy. Koch Collection. SYNTYPES: two males ; one female. Reg. no. 1925:7:22:281-297 (part). Bozen. KochCollection.ormeanum Verhoeff (1931 a : 521) SYNTYPE : female. Reg. no. 1930:5:6:24. Liguria, Italy. Purchased from K. W.Verhoeff. SYNTYPE: female. Reg. no. 1970:68:1. Liguria, Italy. Larwood Collection (ex Verhoeffcollection).pellegrinense Verhoeff (igoSc : 456) [Transferred to Armadillidium granulatum Brandt]. SYNTYPE : male. Reg. no. 1921:6:10:8. Sicily. Purchased from K. W. Verhoeff.peraccae Tua (1900 : 2) [Transferred to Armadillidium pallasi Brandt] SYNTYPES: two males; two females. Reg. no. 1911:11:9:9940-9943. Otranto, Italy.Collected by Peracca. Norman Collection.petraeum L. Koch (1901 : 26) SYNTYPES: two males. Reg. no. 1911:11:8:9996-9997. Tirol, Austria. NormanCollection. SYNTYPES: eight males; seven females. Reg. no. 1925:7:22:311-325. Torbole, Italy.Koch Collection.portofinense Verhoeff (19080 : 459) SYNTYPE: female. Reg. no. 1907:11:4:21. Portofino, Italian Riviera. Purchased fromK. W. Verhoeff. TYPES OF TERRESTRIAL ISOPODS 211 SYNTYPES : one male ; three females. Reg. no. 1970:42:6. Portofino, Italian Riviera.Larwood Collection (ex VerhoefE collection).pujetanum Verhoeff (1910 : 118). [Transferred to Armadillidium simoni Dollfus] SYNTYPE : female. Reg. no. 1921:6:10:10. Riviera. Purchased from K. W. Verhoeff.quadriserriatum Verhoeff (igoSc : 457) SYNTYPES : one male ; three females. Reg. no. 1907:11:4:23-26. Noli, Italian Riviera.Purchased from K. W. Verhoeff. SYNTYPE : male. Reg. no. 1970:43:1. Noli, Italian Riviera. Larwood Collection (exVerhoeff collection).quinquepustulatum Budde-Lund (1885 : 294) SYNTYPES : two males. Reg. no. 1921:10:18:3183-3184. Ouled Anteurs, Algeria.Collected by E. Simon. Budde-Lund Collection.rehobotense Verhoeff (1917 : 172) SYNTYPES : two males ; one female. Reg. no. 1921:6:10:19-20. Jaffa, Israel. Collectedby Aharoni. Purchased from K. W. Verhoeff.riparium L. Koch (1901 : 28) SYNTYPES : one male ; one female. Reg. no. 1925:7:22:374-375. Torbole, Italy. KochCollection.rosai Arcangeli (1913 : 184) SYNTYPE : female. Reg. no. 1931:4:27:33. Bergamask Alpen, Bergamo, Italy. Pur-chased from K. W. Verhoeff.sanctum Dollfus (18920 : 4) SYNTYPE : female. Reg. no. 1911:11:8:9979. Ramlah, Syria. Collected by Letourneux.Norman Collection. SYNTYPES: one male; three females. Reg. no. 1921:10:18:5850-5853. Ramlah, Syria.Collected by Letourneux. Budde-Lund Collection.savonense Verhoeff (i93ia : 520) SYNTYPE : female. Reg. no. 1930:5:6:23. Savona, Italy. Purchased from K. W.Verhoeff.scaberrimutn Stein (1859 : 264) SYNTYPE : male. Reg. no. 1921:10:18:3228. Dubrovnik (Ragusa), Yugoslavia. Budde-Lund Collection.scabrutn Dollfus (i8g2b : 165) SYNTYPES: one male ; one female. Reg. no. 1911:11:8:9991-9992. Constantina, Spain.Collected by Calder6n. Norman Collection.serratum Budde-Lund (1885 : 297) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:3239-3240. La Preste, centralFrance. Collected by E. Simon. Budde-Lund Collection.simoni Dollfus (1887 : 2) SYNTYPE : male. Reg. no. 1892:12:6:3. Cannes, France. Presented by W. D'ArcyThompson. SYNTYPES: three females. Reg. no. 1911:11:8:9963-9965. Cannes, France. 'Pineforest, under stones.' Norman Collection. SYNTYPES: three males ; one female. Reg. no. 1921:10:18:3249-3252. Cannes, France.Budde-Lund Collection (ex Dollfus collection). SYNTYPES : two females. Reg. no. 1928:12:1:1455-1457 (part). Cannes, France. Steb-bing Collection (ex Dollfus collection). SYNTYPE : male. Reg. no. 1956:10:10:198. Cannes, France. Presented by UniversityCollege, Dundee.simoni gigas Verhoeff (1931 a : 519) SYNTYPE : female. Reg. no. 1931:4:27:38. Toscana, Italy. Purchased from K. W.Verhoeff. SYNTYPE : female. Reg. no. 1970:69:1. Monte Pisani, Italy. Larwood Collection (exVerhoeff collection). 212 R. J. LINCOLN AND J. P. ELLIS speyeri Jackson (1923 : 224) [Transferred to Armadillidium nasatum Budde-Lund] HOLOTYPE : male. Reg. no. 1924:5:15:1. Cheshunt, Hertfordshire, England. 'In thecucumber houses of the Experimental and Research Station.' Collected and presented by E.Speyer. PARATYPES : six males ; eleven females. Reg. no. 1924:5:15:2-11. Cheshunt, Hertford-shire, England. Collected and donated by E. Speyer. PARATYPES: twenty-one juveniles. Reg. no. 1952:4:18:231-235. Cheshunt, Hertford-shire, England. Collected by E. Speyer. Jackson Collection.stolikanum Verhoeff (igoyb : 469) SYNTYPE : female. Reg. no. 1907:11:4:22. Stoliko, Middle Greece. Purchased fromK. W. Verhoeff. subdentatum Haswell (1882 : 279) [Redetermined by Budde-Lund as Armadillidium vulgare(Latreille)] SYNTYPE: male. Reg. no. 1895:11:14:87-88 (part). Tasmania, Australia. Presentedby the Australian Museum, Sydney.tendanum Verhoeff (i93ia : 522) [Transferred to Armadillidium alassiense Verhoeff] SYNTYPE : female. Reg. no. 1930:5:6:25. Tenda, Italy. Purchased from K. W. Verhoeff.SYNTYPES : one male ; two females. Reg. no. 1937:7:6:10-12. Tenda, Italy. Purchasedfrom K. W. Verhoeff. SYNTYPE : male. Reg. no. 1970:71:1. Tenda, Italy. Larwood Collection (ex Verhoeffcollection).tigris Budde-Lund (1885 : 55) SYNTYPE : male. Reg. no. 1921:10:18:3304. Madeira I. Collected by Metschnikoff.Budde-Lund Collection.tirolense Verhoeff (igoib : 67) SYNTYPES: one male ; four females. Reg. no. 1901:9:19:10-14. Southern Tirol, Austria.Purchased from K. W. Verhoeff.tunetanum Verhoeff (igo7b : 473) SYNTYPE : female. Reg. no. 1907:11:4:30. Tunis, Tunisian Republic, N. Africa. Pur-chased from K. W. Verhoeff.vallombrosae Verhoeff (i9O7b : 480) SYNTYPES: five females. Reg. no. 1937:7:6:21-25. Vallombrosa, Apennines, Italy.Purchased from K. W. Verhoeff.versicolor Stein (1859 : 265) SYNTYPE: female. Reg. no. 1921:10:18:3350. Dubrovnik (formerly Ragusa), Yugo-slavia. Budde-Lund Collection. ELUMA Budde-Lund purpurascens Budde-Lund (1885 : 48) SYNTYPES: three females; one juvenile. Reg. no. 1921:10:18:3769-3772. Madeira I.Collected by Metschnikoff. Budde-Lund Collection. EXZAES Barnard bicolor Barnard (1932 : 300) SYNTYPES : fifty females; two juveniles. Reg. no. 1933:1:25:292-301. OudeboschForest, River Zonder End Mts, Cape Province, South Africa. Collected by K. H. Barnard.Presented by the South African Museum.sylvatica Barnard (1932 : 299) SYNTYPES : two females. Reg. no. 1933:1:25:287-291. George Forest, Cape Province,South Africa. Collected by K. H. Barnard. Presented by the South African Museum. TYPES OF TERRESTRIAL ISOPODS 213 HEKELUS Barnard episimus Barnard (1932 : 298) SYNTYPES : three males ; fourteen females ; thirteen juveniles. Reg. no. 1933:1:25:278-286. Table Mts and Noordhoek Forest, Cape Peninsula, South Africa. Collected by K. H.Barnard. Presented by the South African Museum. PARELUMA Omer-Cooper minuta Omer-Cooper (1923 : 396) SYNTYPES: two females. Reg. no. 1922:5:18:11-12. Amarah, Iraq. 13.11.1918.Collected by W. E. Evans. Presented by J. Omer-Cooper. SCHIZIDIUM Verhoeff hybridum (Budde-Lund) (1896 : 44) [Armadillidium hybridum} HOLOTYPE : fragment only. Reg. no. 1921:10:18:2960. Symi, Greece. 7.6.1887. Col-lected by E. von Oertzen. Budde-Lund Collection.oertzeni (Budde-Lund) 1896 : 42 [Armardillidium oertzeni] SYNTYPES : one male ; five females. Reg. no. 1921:10:18:3086-3091. Karpathos (May1887) and Rhodes, Greece. Collected by E. von Oertzen. Budde-Lund Collection. SUNNIVA Budde-Lund tnammillata Barnard (1936 : 8) SYNTYPES : one male ; two females. Reg. no. 1936:7:13:43-44. Le Pouce, Mauritius I.,2400 ft. 10.1.1935. Collected and presented by R. F. Lawrence, Natal Museum.minor Budde-Lund (1908 : 268) SYNTYPE : few fragments only. Reg. no. 1921:10:18:937. Mauritius I. Collected by A.Voeltzkow. Budde-Lund Collection.mystica Budde-Lund (1908 : 267) SYNTYPE : few fragments only. Reg. no. 1921:10:18:938. Madagascar I. Collected byA. Voeltzkow. Budde-Lund Collection.uniformis Barnard (1936 : 9) SYNTYPES : five males ; six females. Reg. no. 1936:7:13:45-49. Le Pouce, Mauritius I.,2400 ft. 10.1.1935. Collected and presented by R. F. Lawrence, Natal Museum. Family EUBELIDAEBENECHINUS Budde-Lund arm at us Budde-Lund (1910 : 4) SYNTYPES : two males; four females. Reg. no. 1921:10:18:772-777. Meru Mt, Tan-ganyika, 3000-3500 m. Budde-Lund Collection. ETHELUM Budde-Lund americanum (Dollfus) (i8g6c : 397) [Mesarmadillo americanus] SYNTYPE : female. Reg. no. 1896:1:11:20. Leeward Is., West Indies. 'Lowland nearsea. May. Under stones.' Collected by H. H. Smith. Presented by the West IndiaCommittee. SYNTYPES: four females. Reg. no. 1896:1:11:21. St Vincent, West Indies. 'Sugarcanefield. March. Under decaying cane leaves.' Collected by H. H. Smith. Presented by theWest India Committee. 214 R. J. LINCOLN AND J. P. ELLIS SYNTYPES : four females. Reg. no. 1896:1:11:22. St Vincent, West Indies. 'Shadyplace. 259 ft. Under old boards. September.' Collected by H. H. Smith. Presented bythe West India Committee. SYNTYPES: one male ; one female ; one juvenile. Reg.no. 1896:1:11:23. Leeward Is.,West Indies. '500 ft. March. Under rubbish, shady place.' Collected by H. H. Smith.Presented by the West India Committee. SYNTYPE : female. Reg. no. 1921:10:18:771. St Vincent, West Indies. Collected byH. H. Smith. Budde-Lund Collection.modestum (Dollfus) (18960 : 397) [Mesarmadillo modestus] HOLOTYPE (?) : female. Reg. no. 1896:1:11:24. St Vincent, West Indies. 'Low groundS.E. of the Island, under rubbish.' Collected by H. H. Smith. Presented by the West IndiaCommittee.reflexum (Dollfus) (18960 : 398) [Mesarmadillo reflexus] HOLOTYPE : female. Reg. no. 1896:1:11:19. St Vincent, West Indies. 'Open swampyland, southern end of Island. 27 September. Under rubbish.' Collected by H. H. Smith.Presented by the West India Committee. EUBELUM Budde-Lund dollfusi Budde-Lund (1899 : 8) SYNTYPE : female. Reg. no. 1921:10:18:589. Antoto, Shoa district, Ethiopia. August1885. Collected by V. Ragazzi. Budde-Lund Collection.ignavum Budde-Lund (1899 : 9) SYNTYPE: fragments only. Reg. no. 1921:10:18:590. Let-Marfia, Shoa district, Ethiopia.March 1885. Collected by V. Ragazzi. Budde-Lund Collection.instrenuum Budde-Lund (19120 : 2) SYNTYPES : one male; ten females. Reg. no. 1921:10:18:597-607. Escarpment, EastAfrica. Collected by E. Lonnberg. Budde-Lund Collection.lubricum Budde-Lund (1885 : 292) SYNTYPES: four males; three females. Reg. no. 1921:10:18:619-625. Landana, Angola,Southwest Africa. Budde-Lund Collection. SYNTYPES : one male; one female. Reg. no. 1956:10:10:199-200. Landana, Angola,Southwest Africa. Presented by University College, Dundee.quietum Budde-Lund (1899 : 7) HOLOTYPE (?) : male. Reg. no. 1921:10:18:639. Antoto, Shoa district, Ethiopia. March1885. (Locality in publication: Let-Marefia.) Collected by V. Ragazzi. Budde-LundCollection.stipulatum Budde-Lund (1899 : 5) SYNTYPES: four males ; two females. Reg. no. 1921:10:18:640-645. Cameroons, WestAfrica. November 1891. Collected by Y. Sjostedt. Budde-Lund Collection. GELSANA Budde-Lund abnormis Budde-Lund (1910 : 6) SYNTYPES : one male ; two specimens in very poor condition. Reg. no. 1921:10:18:790-792. Kilimanjaro, Tanganyika. Collected by Y. Sjostedt. Budde-Lund Collection. GERUTHA Budde-Lund laevis Budde-Lund (19120 : 6) SYNTYPES: one male; one female. Reg. no. 1921:10:18:717-718. Escarpment, EastAfrica. 'Under logs in the forest at Escarpment station. 11.1.1911.' Collected by E.Lonnberg. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 215 tnarginepilosa Budde-Lund (19120 : 6) SYNTYPES : one male; four females. Reg. no. 1921:10:18:719-723. Kaffa, Ethiopia.Collected by E. Lonnberg. Budde-Lund Collection.pila (Budde-Lund) (1898 : 4) [Eubelum piluni] SYNTYPES: one male; twofemales. Reg. no. 1921:10:18:724-725. Ru-Nsororo. 2600 m.9.6.1891. Collected by Stuhlmann. Budde-Lund Collection.pilosa Budde-Lund (i9i2c : 6) SYNTYPE : female. Reg. no. 1921:10:18:726. Shoa district, Ethiopia. Collected by E.Lonnberg. Budde-Lund Collection. HAPLARMADILLO Dollfus monocellatus Dollfus (18960 : 400) HOLOTYPE : female. Reg. no. 1896:1:11:25. Richmond Valley, St Vincent, West Indies.'Under rotting leaves, nooft. i8th January.' Collected by H. H. Smith. Presented bythe West India Committee. HIALLUM Budde-Lund hilgendorfi (Budde-Lund) (1898 : 4) [Eubelum hilgendorfi] SYNTYPES: one male; one female. Reg. no. 1921:10:18:769-770. Runsororo, EastAfrica. 'Bamboo forest, 9-13.6.1891.' Collected by Stuhlmann and E. Pasha. Budde-Lund Collection. IGNAMBA Budde-Lund brevis Budde-Lund (1910 : 5) SYNTYPES : four females. Reg. no. 1921:10:18:778-781. Kilimanjaro, Tanganyika.Collected by Y. Sjostedt. Budde-Lund Collection.microps Budde-Lund (1910 : 6) SYNTYPES : seven females. Reg. no. 1921:10:18:782-789. Kiboscho, Kilimanjaro,Tanganyika. 'In clefts of mountains. 3000 m. February 1906.' Collected by Y. Sjostedt.Budde-Lund Collection. MESARMADILLO Dollfus albicornis Budde-Lund (1899 : 13) SYNTYPES: one male; one female. Reg. no. 1921:10:18:727-728. N'dian, Cameroons,West Africa. June 1891. Collected by Y. Sjostedt. Budde-Lund Collection.quadritnaculatus Budde-Lund (1899 : 14) SYNTYPES: two males. Reg. no. 1921:10:18:733. Cameroons, West Africa. Collectedby Y. Sjostedt. Budde-Lund Collection.tuberculatus Dollfus (i8g2a : 387) SYNTYPE : female. Reg. no. 1921:10:18:735. Assinie, West Africa. 1886. Collectedby C. Alluaud. Budde-Lund Collection. MICROCERCUS Budde-Lund abyssinicus Barnard (1940 : 365) SYNTYPES: one male; one female. Reg. no. 1949:2:2:231-232. Djem Djem Forest,Ethiopia. 21.9.1926 and 11.10.1926. Collected and presented by J. Omer-Cooper.leucocephalus (Budde-Lund) (1895 : 607) [Periscyphis leucocephalus] SYNTYPE : female. Reg. no. 1921:10:18:1267. Rangoon, Burma. July 1887. Collectedby L. Fea. Budde-Lund Collection. 216 R. J. LINCOLN AND J. P. ELLIS PARAPERISCYPHIS Stebbing stebbingi Collinge (19145 : 207) HOLOTYPE : male. Reg. no. 1919:4:26:455. Anamali Hills, Madras, S. India. 4000 ft.22.1.1912. Collected by T. B. Fletcher. Collinge Collection. PERISCYPHIS Gerstaecker besi Barnard (1941 : 63) SYNTYPES : four males; eight females. Reg. no. 1949:2:2:73-84 (part). Jebel Harir,Western Aden Protectorate. 5000-6000 ft. 10.11.1937. Collected by H. Scott and E. B.Britton. British Museum Expedition to S.W. Arabia, 1937-38. SYNTYPES : three females. Reg. no. 1949:2:2:73-84 (part). Yemen, 2 miles West ofSan'a. 8000 ft. 14.2.1938. Collected by H. Scott and E. B. Britton. British MuseumExpedition to S.W. Arabia, 1937-38. SYNTYPE : female. Reg. no. 1949:2:2:73-84 (part). Jebel Jihaf, Western Aden Protec-torate. 'Under stones and in damp soil at roots of plants, on steep face of mountain facingnorth-west. 7700 ft. 4.10.1937.' Collected by H. Scott and E. B. Britton. British MuseumExpedition to South West Arabia, 1937-38. SYNTYPE: female. Reg. no. 1949:2:2:73-84. Wadi Dareij a, south-west of Dhala, WesternAden Protectorate. 4600 ft. 9.11.1937. Collected by H. Scott and E. B. Britton. BritishMuseum Expedition to South West Arabia, 1937-38.bicoloratus Barnard (1941 : 65) SYNTYPES : three males. Reg. no. 1949:2:2:85-87. Dhala, Western Aden Protectorate.'From detritus under succulent plants. 4800 ft. 14.9.1937.' Collected by H. Scott. BritishMuseum Expedition to South West Arabia, 1937-38.cavernicola Omer-Cooper (1926 : 371) SYNTYPES: one male ; two females. Reg. no. 1921:10:18:1289-1291. Ethiopia. Budde-Lund Collection.chindeensis Barnard (1932 : 294) HOLOTYPE : male. Reg. no. 1933:1:25:262. Chinde, mouth of the Zambeze River,Mofambique. 1912. 'In sand dunes near the shore.' Collected by K. H. Barnard. Pre-sented by the South African Museum.civilis Budde-Lund (1908 : 279) SYNTYPES: two males; one female. Reg. no. 1921:10:18:1291-1295. Witu, Kenya.Collected by A. Voeltzkow. Budde-Lund Collection.hughscotti Barnard (1940 : 362) SYNTYPES: seventeen males; twenty-five females. Reg. no. 1949:2:2:201-230. DjemDjem Forest, Ethiopia, c. 8100 ft. 21.9.1926-10.10.1926. Collected and presented by J.Omer-Cooper. SYNTYPES: twenty-four males; seventeen females. Reg. no. 1949:2:2:201-230 (part).Addis Ababa, Ethiopia. Over 8000 ft. 10.9.1926. Collected and presented by J. Omer-Cooper.kunenensis (Barnard) (1924 : 231) [Periscyphops kunenensis] SYNTYPES: seven males ; twelve females. Reg.no. 1933:1:25:263-270. Ovambolande,Kunene River, South West Africa. Collected by K. H. Barnard. Presented by the SouthAfrican Museum.latissimus Omer-Cooper (1926 : 395) HOLOTYPE : male. Reg. no. 1921:10:18:1309. Ethiopia. Budde-Lund Collection.limbata Omer-Cooper (1926 : 384) SYNTYPES: two females. Reg. no. 1921:10:18:1310-1311. Kibwesa, Tanganyika.Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 217 nigricans Omer-Cooper (1926 : 382) SYNTYPES : three females. Reg. no. 1921:10:18:1312-1314. Locality given in publication :'Abyssinia, Schoa Galla ; Somaliland'. No locality data with specimen. 9.10.1901. Col-lected by V. Erlangen. Budde-Lund Collection.pulcher Budde-Lund (1898 : 7) HOLOTYPE : female. Reg. no. 1921:10:18:1315. Tanga, East Africa. Collected byMoller. Budde-Lund Collection.quadrimaculatus Budde-Lund (1908 : 279) [Transferred to Periscyphis trivialis Gerst] SYNTYPES: eleven males; seven females. Reg. no. 1921:10:18:1316-1327. Manda I.,Kenya. 21.2.1903. Collected by A. Voeltzkow. Budde-Lund Collection.ruficauda Budde-Lund (1908 : 278) SYNTYPES : fourteen males ; thirteen females ; one other specimen in fragments. Reg. no.1921:10:18:1328-1339. Manda I., Kenya. 21.2.1903. Collected by A. Voeltzkow. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1952:4:18:59. Manda I., Kenya. Collected by A. Voeltzkow.Jackson Collection.subtransversus Omer-Cooper (1926 : 392) SYNTYPES : one male ; two females. Reg. no. 1921:10:18:1340-1342. Egypt. Budde-Lund Collection.tamei Omer-Cooper (1923 : 394) SYNTYPE : one specimen in fragments. Reg. no. 1922:5:18:13-14 (part). Al Amarah,Iraq. 1918. Collected by R. G. Tame. Presented by J. Omer-Cooper. SYNTYPE : one female. Reg. no. 1922:5:18:13-14 (part). Ruz ('Robat' on label), N.E. ofBaghdad. 1918. Collected by W. E. Evans. Presented by J. Omer-Cooper. SYNTYPES : micropreparations of appendages : six males ; fourteen females. Reg. no.1922:5:18:15-34. Amara, Iraq. 1918. Collected by R. G. Tame. Presented by J. Omer-Cooper.trivialis Gerstaecker (1873 : 526) SYNTYPES : three females. Reg. no. 1921:10:18:1343-1345. Lake Jipe, Kenya. Budde-Lund Collection.undulata Omer-Cooper (1926 : 373) SYNTYPES : three males; five females. Reg. no. 1921:10:18:1346-1353. Bala, Shoadistrict, Ethiopia. Budde-Lund Collection.vittatus Omer-Cooper (1926 : 366) HOLOTYPE : male. Reg. no. 1921:10:18:1354-1359 (part). Obock, Somaliland. Budde-Lund Collection. PARATYPES : six females. Reg. no. 1921:10:18:1354-1359 (part). Obock, Somaliland.Budde-Lund Collection. PERISCYPHOPS Hilgendorf bizonatus Budde-Lund (1899 : 18) SYNTYPES: three males; four females. Reg. no. 1921:10:18:736-744. Cameroons,West Africa. 1891. Collected by Y. Sjostedt. Budde-Lund Collection.silvanus Budde-Lund (1899 : 16) SYNTYPES : four males ; two females. Reg. no. 1921:10:18:749-753. Cameroons, WestAfrica. 1891. Collected by Y. Sjostedt. Budde-Lund Collection.squamatus Budde-Lund (1899 : 20) SYNTYPE : small fragment only. Reg. no. 1921:10:18:755. Bibundi, Cameroons, Nigeria.August 1891. Collected by Y. Sjostedt. Budde-Lund Collection.squamosus Budde-Lund (1899 : 21) SYNTYPE : fragments only. Reg. no. 1921:10:18:756. Bibundi, Cameroons, Nigeria.August 1891. Collected by Y. Sjostedt. Budde-Lund Collection. 2i8 R. J. LINCOLN AND J. P. ELLIS triarticulatus (Hilgendorf) (1893 : 152) [Periscyphis (Periscyphops) triarticulatus] SYNTYPES : one male; three females. Reg. no. 1921:10:18:760-763. Bismarkburg, Togo,West Africa. March 1891. Collected by Buttner. Budde-Lund Collection. ROTUNGUS Collinge pictus Colh'nge (1916 : 544) PARATYPE : female. Reg. no. 1919:4:26:541. Kobo, Abor country, Assam, India, 400 ft,'under bark'. Collected by S. W. Kemp. Presented by W. E. Collinge. Family SPHAERONISGIDAECIRCONISCUS Pearse spinosus (Collinge) (1918 : 62) [Paracubaris spinosus] SYNTYPES : two males. Reg. no. 1919:4:26:542-543. Mazakuvi River, Guiana. May1916. 'In decaying wood.' Collected by G. E. Bodkin. Collinge Collection. SCLEROPACTES Budde-Lund concinnus Budde-Lund (1885 : 240) SYNTYPES : two males ; one female ; one other specimen in fragments. Reg. no. 1921:10:18:961-964. Tambillo, Peru. Collected by J. Stolzmann. Budde-Lund Collection. incicus Budde-Lund (1885 : 241) HOLOTYPE : male. Reg. no. 1921:10:18:968. Peru. Collected by J. Stolzmann. Budde-Lund Collection. SPHAEROBATHYTROPA Verhoeff ribauti Verhoeff (igoSb : 171) SYNTYPES : two females. Reg. no. 1908:6:1:15-16. St Beat, Pyrenees, France. Col-lected by Ribaut. Purchased from K. W. Verhoeff. SPHAERONISCUS Budde-Lund frontalis Richardson (1914 : 31) SYNTYPE : male. Reg. no. 1928:5:1:84-87 (part). Buenavista, nr Viota, Colombia.Collected by and purchased from O. Fuhrmann. SYNTYPES : two males ; two females. Reg. no. 1928:5:1:84-87 (part). Argelia, nr Viota,Colombia. Collected by and purchased from O. Fuhrmann. Family AGTOECHDAEACTOECIA Dana opihensis Chilton (1901 : 132) SYNTYPES: one male; two females. Reg. no. 1900:11:61-64. Timaru, New Zealand.Presented by C. Chilton. SYNTYPES: one male; one female. Reg. no. 1921:10:18:1072-1073. Timaru, NewZealand. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 219 Family ARMADILLIDAEACANTHONISCUS Kinahan spiniger Kinahan (1859 : 197) HOLOTYPE : female. Reg. no. 1973:478:1 (original registration no. 1845:118). Jamaica,West Indies. Purchased from Mr Gosse. AKERMANIA Collinge spinosa Collinge (1919 : 230) SYNTYPES : four males ; four females ; four other specimens with genitalia missing.Reg. no. 1919:4:26:282-291. Umhlali, Winkle Spruit, South Africa. May 1916. Collectedby C. Akerman. Collinge Collection. SYNTYPES : four females. Reg. no. 1933:1:25:413-417. Umhlali, Winkle Spruit, SouthAfrica. Collected by C. Akerman. Presented by the South African Museum. ANCHICUBARIS CoUinge fongosiensis Collinge (1920 : 484) SYNTYPES : twelve males; twenty females. Reg. no. 1919:4:26:504-518. Mt Fongosi, Zululand, Natal. July 1917. Collected by W. E. Jones. Collinge Collection. SYNTYPES: forty-three males; forty-one females. Reg. no. 1933:1:25:851-870. Mt Fongosi, Zululand, Natal. July 1917. Collected by W. E. Jones. Presented by the SouthAfrican Museum. ARMADILLO Dumeril afflnis (Miers) (i877b : 666) [Cubans affinis] SYNTYPES: one male ; four females. Reg. no. 1879:21:6. Cayenne, Guiana. Presentedby A. Wrzesniowsky.albomarginatus Dollfus (i892C : 3) SYNTYPE : male. Reg. no. 1911:11:8:9755. Route de Ouady-Hafaf at Ain-Djeddy, DeadSea. Collected by T. Barrois. Norman Collection. SYNTYPE: female. Reg. no. 1921:10:18:2434. Karytein, Syria. Collected by T. Barrois.Budde-Lund Collection.albospinosus Dollfus (1900 : 522) SYNTYPE: female. Reg. no. 1904:11:5:8-9 (part). Makaweli, Kauai I., Hawaiian Is.,3000 ft. 1897. Presented by the Joint Committee for investigating the Fauna of the Sand-wich Islands. SYNTYPE: male. Reg. no. 1904:11:5:8-9 (part). Kawaihoa, Oahu I., Hawaiian Is.Collected by Perkins. Presented by the Joint Committee for investigating the Fauna of theSandwich Islands.ausseli Dollfus (1893 : 48) SYNTYPES: three females. Reg. no. 1921:10:18:1095-1096. La Laguna, Tenerife, CanaryIs. 'Agua Garcia, laurel forest (probably Las Mercedes Forest), 800 m.' Collected by H.Aussel. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1908:6:1:44. Tenerife, Canary Is. Purchased from K. W.Verhoeff.bituberculatus Budde-Lund (igi2a : 36) SYNTYPE : female. Reg. no. 1921:10:18:2450. Torbay, South West Australia. 'Sta. 162.'Budde-Lund Collection.brevicornis Budde-Lund (igisa : 69) SYNTYPES : four females. Reg. no. 1921:10:18:2451-2454. New South Wales, Australia.Budde-Lund Collection. 220 R. J. LINCOLN AND J. P. ELLIS cinctus Budde-Lund (1896 : 41) SYNTYPES : two females; one other broken specimen. Reg. no. 1921:10:18:2466-2468.Kasos I., Greece, Mediterranean Sea. Collected by E. von Oertzen. Budde-Lund Collection.collinus Budde-Lund (1895 : 6°4) SYNTYPES : two specimens both in fragments. Reg. no. 1921:10:18:2518-2519. CarimHills, Ascinii Ghecu, Burma. 1300-1400 m. Collected by L. Fea. Budde-Lund Collection.conglobator Budde-Lund (1904 : 124) HOLOTYPE : female. Reg. nos. 1921:10:18:1663 & 1921:10:18:2520 (fragments fromholotype). Pulo, Penang, Malaysia. 600-750 m. February 1879. Collected by L. Feaand Loria. Budde-Lund Collection.egens Budde-Lund (1904 : 124) HOLOTYPE : broken. Reg. no. 1902:12:4:16. Malawi (formerly Nyasaland), CentralAfrica. Collected and presented by H. H. Johnston.erythroleucus Budde-Lund (1904 : 98) HOLOTYPE (?): female. Reg. no. 1921:10:18:2469. Crimea, Ukraine, USSR. Budde-LundCollection.flavus Budde-Lund (igi2a : 37) SYNTYPES: one male; one female. Reg. no. 1921:10:18:2415-2416. Mundaring Weir,West Australia. 'Sta. 101.' 9.8.1905.harsadiensis (Barnard) (1940 : 359) SYNTYPES : one male ; three females ; three other specimens in fragments. Reg. no.1949:2:2:171-178. Shore of Hora Harsadi, lakes of Addas, Ethiopia. 7000 ft. 2-3.12.1926.Collected and presented by J. Omer-Cooper.imtnotus Budde-Lund (1904 : 122) SYNTYPES: one male ; two females. Reg.no. 1921:10:18:2523-2525. Pulo, Penang I.,Malaysia. 600-70001. March 1870. Collected by L. Fea and Loria. Budde-Lund Collection.inconspicuus Miers (1876 : 225) [Transferred to Spherillo danae Heller] HOLOTYPE : female. Reg. no. 1973:477:1 (original reg. no. 1845:4). New Zealand.Presented by Col. Bolton.integer Budde-Lund (i9i2a : 35) SYNTYPES : three males. Reg. no. 1921:10:18:2426-2428. Torbay, Western Australia.'Sta. 162.' 19.8.1905. Budde-Lund Collection.interrnixtus Budde-Lund (1904 : 126) SYNTYPES: two males ; one other specimen in fragments. Reg. no. 1921:10:18:2526-2527.The Caves, Selangore, Malaysia. 1897. Collected by H. N. Ridley. Budde-Lund Collection.SYNTYPES: twenty-nine males; nine females. Reg. no. 1898:4:11:1-10. The Caves,Selangore, Malaysia. 1897. Collected and presented by H. N. Ridley.lifuensis (Stebbing) (1900 : 653) {Cubans lifuensis] SYNTYPES: one male; six females. Reg. no. 1906:4:19:81-88. Lifu L, Loyalty Is.,Pacific Ocean. Presented by A. Willey. SYNTYPES : one male ; two females. Reg. no. 1928:12:1:1553-1555. Lifu L, Loyalty Is.Stebbing Collection.macmahoni Chilton (1901 : 149) SYNTYPES : one male ; one female. Reg. no. 1900:11:1:16-17. Kenepuru, Marlborough,New Zealand. Presented by C. Chilton.mayeti Simon (1885 : 7) SYNTYPES: one male; one female. Reg. no. 1921:10:18:2470-2471. Sfax, Tunisia.Collected by V. Mayer. Budde-Lund Collection.miser Budde-Lund (1904 : 121) HOLOTYPE: male. Reg. no. 1921:10:18:2528. Victoria, Australia. Collected by Haus-child. Budde-Lund Collection.montanus Budde-Lund (1904 : 99) SYNTYPES: one male; five females. Reg. no 1921:10:18:2472-2477. Djebel Tarchuna,Tripoli, Libya. Collected by G. Rohlfs. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 221 nigromarginatus Budde-Lund (1904 : 127) HOLOTYPE : male. Reg. no. 1902:12:4:6. Selangore, Malaysia. 'In caves, 1897.' Col-lected and presented by H. N. Ridley.offlcinalis syriaca Dollfus (18920 : 2) SYNTYPE : female. Reg. no. 1892:12:6:2. Syria. 1886. Collected by L. Bleuse. Pre-sented by D'Arcy Thomson (ex Dollfus collection). SYNTYPES : two females. Reg. no. 1911:11:8:9750-9751. Syria. 1886. Collected byL. Bleuse. Norman Collection. SYNTYPE : female. Reg. no. 1956:10:10:137. Syria. Collected by L. Bleuse. Presentedby University College, Dundee.oxyzomus (Barnard) (1940 : 360) [Cubans oxyzomus] SYNTYPES : two females. Reg. no. 1949:2:2:179-180. Djem Djem Forest, Ethiopia.8000 ft. 20.9.1926. Collected and presented by J. Omer-Cooper.piger Budde-Lund (1896 : 42) HOLOTYPE : female. Reg. no. 1921:10:18:2494. Sitia, Crete. Budde-Lund Collection.pilularis Say (1818 : 432) [Transferred to Armadillidium vulgare (Latr.)] SYNTYPE : female. Reg. no. 1973:510:1. North America. Presented by T. Say.proximatus Budde-Lund (1904 : 122) SYNTYPES: one male ; three females. Reg. no. 1921:10:18:2572-2578. Telang, Borneo.Budde-Lund Collection.pygtnaeus Budde-Lund (i9i2a : 36) SYNTYPES: seven females. Reg. no. 1921:10:18:2495-2501. Guildford, Western Australia.'Sta. 103'. 19.5.1905. Budde-Lund Collection.sharpi Dollfus (1900 : 523) HOLOTYPE: female. Reg. no. 1904:11:5:13. Kauai I., Hawaiian Is. 4000 ft. August1896. Collected by Perkins. Presented by the Joint Committee for Investigating theFauna of the Sandwich Islands.tenuipunctatus Dollfus (18960 : 389) SYNTYPES : two males ; one female. Reg. no. 1896:1:11:2-3. Mustique L, Windward Is.'Beaten from bushes. June.' Collected by H. H. Smith. Presented by the West IndiaCommittee.trifoliutn Dollfus (1890 : i) HOLOTYPE : female. Reg. no. 1927:5:14:1. San Tiago, Cape Verdi Is. August 1873.'Challenger' Collection.viticola Dollfus (18960 : 396) SYNTYPE : male. Reg. no. 1896:1:11:9-11 (part). Chantilly, Grenada L, Windward Is.1400 ft. 23 March. 'Hillside, edge of forest, beaten from vines and brush.' Collected byH. H. Smith. Presented by the West India Committee. SYNTYPES : one male ; one female. Reg. no. 1896:1:11:9-11 (part). Balthazar, GrenadaL, Windward Is. 250 ft. 19 March and 17 August. 'Second-growth woods, beaten fromvines and brush.' Collected by H. H. Smith. Presented by the West India Committee.wilsmorei (Nicholls & Barnes) (1926 : 150) [Cubans wilsmorei] SYNTYPES : four males ; eight females ; five other specimens tightly rolled up. Reg. no.1926:12:1:30-34. Banks of Frankland River, Nornalup, South West Australia. 'Underlogs'. Collected and presented by G. E. Nicholls.zebricolor (Stebbing) (1900:656; {Cubans zebricolor] HOLOTYPE : fragmented. Reg. no. 1906:4:19:93. Lifu I., Loyalty Is. Presented by A.Willey. BETHALUS Budde-Lund barbertoni Barnard (1932 : 312) SYNTYPES : three males ; four females ; two other specimens in fragments. Reg. no.1933:1:25:365-371. Barberton, Transvaal, South Africa. Presented by the South AfricanMuseum. 12 222 R. J. LINCOLN AND J. P. ELLIS SYNTYPES : one male ; three females. Reg. no. 1933:1:25:372-375. Inhambane, Mo9am-bique. Collected by K. H. Barnard. Presented by the South African Museum.barnardi (Collinge) (1920 : 482) [Cubaris barnardi] PARATYPES: forty specimens (males, females and juveniles). Reg. no. 1933:1:25:398-412.Mt Fongosi, Zululand, South Africa. January 1917. Collected by W. E. Jones. Presentedby the South African Museum.carinatus (Budde-Lund) (1908 : 272) [Armadillo carinatus] SYNTYPES: nine males; twenty-three females. Reg. no. 1921:10:18:1189-1200. Tanana-rive, Madagascar. June 1897. Budde-Lund Collection.cassida (Budde-Lund) (1908 : 274) [Armadillo cassida] HOLOTYPE : few fragments only. Reg. no. 1921:10:18:1201. Sahana, Madagascar.September 1904. Budde-Lund Collection.depressus (Dollfus) (i8g6c : 390) [Armadillo depressus] HOLOTYPE : female. Reg. no. 1896:1:11:1. Chateaubelais, St Vincent I., West Indies.August 1889-91. Collected by H. H. Smith. Presented by the West India Committee.etnarginatus Budde-Lund (1910 : 12) SYNTYPES: three males; three females. Reg. no. 1921:10:18:1204-1208. Mkulimusi,nr Tanga, Tanganyika. June 1906. Budde-Lund Collection.ernunitus (Budde-Lund) (1904 : 129) [Armadillo emunitus] SYNTYPES: one male; two females. Reg. no. 1921:10:18:2521-2522. Thailand. Budde-Lund Collection. griseoalbus (Dollfus) (1895 : 347) [Armadillo griseoalbus] [Transferred to Bethalus limbatus(Brandt)] SYNTYPES : two males. Reg. no. 1921:10:18:1218. Matjesfontein, South Africa. Col-lected by E. Simon. Budde-Lund Collection. latifrons (Budde-Lund) (1904 : 132) [Armadillo latifrons'] [Transferred to Bethalus mucidus(Budde-Lund)] SYNTYPES : one male; two females. Reg. no. 1921:10:18:1219-1221. Natal, SouthAfrica. 1840-45. Collected by A. Wahlberg. Budde-Lund Collection.macrodens Barnard (1932 : 311) SYNTYPES: two males ; one female. Reg. no. 1933:1:25:361-364. Groot Vaders Bosch,Langeberg Mts, nr Heidelberg, Cape Province, South Africa. Collected by K. H. Barnard.Presented by the South African Museum.panurus (Budde-Lund) (1904 : 131) [Armadillo panurus] HOLOTYPE: male. Reg. no. 1921:10:18:1132. Natal, South Africa. Collected by A.Wahlberg. Budde-Lund Collection.pretoriensis (Dollfus) (1895 : 348) [Armadillo pretoriensis^ SYNTYPES: one male ; one female (fragments only). Reg. no. 1921:10:18:1227. Pretoria,South Africa. Collected by E. Simon. Budde-Lund Collection.rhodesiae Barnard (1932 : 305) SYNTYPES: one male; two females. Reg. no. 1933:1:25:328-330. Bulawayo, SouthRhodesia. Collected by R. W. E. Tucker. Presented by the South African Museum.stricticauda (Dollfus) (1895 : 348) [Armadillo stricticauda] SYNTYPES: one male; one female. Reg. no. 1921:10:18:1229. Grotte de Makapan,Transvaal, South Africa. Collected by E. Simon. Budde-Lund Collection.tradouwi Barnard (1932 : 310) SYNTYPES: two males; six females. Reg. no. 1933:1:25:353-360. Tradouw Pass,Langeberg Mts, Cape Province, South Africa. Collected by K. H. Barnard. Presented bythe South African Museum. BUDDELUNDIA Michaelson albomaculatus Budde-Lund (igi2a : 33) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:2432-2433. Broome Hill, WestAustralia. 'Sta. 158'. 24.8.1905. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 223 binotatus Budde-Lund (19125,: 24) SYNTYPES : two females. Reg. no. 1921:10:18:1435-2436. Boorabbin, West Australia.'Sta. 95'. 3.7.1905. Budde-Lund Collection.bipartitus Budde-Lund (i9i2a : 24) SYNTYPES: two males; eleven females. Reg. no. 1921:10:18:2437 (fragments); 2438-2449. South West Australia. Budde-Lund Collection.callosus Budde-Lund (i9i2a : 28) SYNTYPES: four males ; three females. Reg. no. 1921:10:18:2455-2461. Northampton,West Australia. 'Sta. 71'. 15.8.1905. Budde-Lund Collection.cinerascens Budde-Lund (191 aa : 26) SYNTYPES: one male ; two females. Reg. no. 1921:10:18:2464-2465. Rottnest I., WestAustralia. 'Sta. 121'. 6-13.10.1905. Budde-Lund Collection.frontosus Budde-Lund (igi2a : 30) HOLOTYPE : male (few small fragments). Reg. no. 1921:10:18:2417. Coolgardie, WestAustralia. 'Sta. 94'. 3.7.1905. Budde-Lund Collection.inaequalis Budde-Lund (igi2a : 27) SYNTYPES: one male ; two females. Reg. no. 1921:10:18:2418-2420. Fremantle, WestAustralia. 20.5.1905. Budde-Lund Collection.labiatus Budde-Lund (i9i2a : 20) HOLOTYPE: male. Reg. no. 1907:5:6:14. Lake Miranda, West Australia. Collected andpresented by G. H. Ince. FRAGMENT OF HOLOTYPE : Reg.no. 1921:10:18:2425. Budde-Lund Collection.laevigatus Budde-Lund (i9i2a : 23) SYNTYPES : two females. Reg. no. 1921:10:18:2430-2431. Day Dawn, West Australia.'Sta. 76'. 9.7.1905. Budde-Lund Collection.lateralis Budde-Lund (igisa : 67) HOLOTYPE : male. Reg. no. 1921:10:18:2429. New South Wales, Australia. Budde-Lund Collection.monticola Budde-Lund (igi2a : 21) HOLOTYPE : female (fragments). Reg. no. 1921:10:18:2478. Mt Robinson, Kalgoorlie,West Australia. 1.7.1905. Budde-Lund Collection.nigripes Budde-Lund (igi2a : 33) SYNTYPES : nine females. Reg. no. 1921:10:18:2479-2487. Bunbury, West Australia.'Sta. 142'. 25.7.1905. Budde-Lund Collection.opacus Budde-Lund (i9i2a : 29) SYNTYPES: three females. Reg. no. 1921:10:18:2491-2493. Gooseberry Hill, WestAustralia. 'Sta. 152'. 31.5.1905. Budde-Lund Collection.quadritracheata Budde-Lund (igi3a : 68) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:2502-2503. Condobolin, NewSouth Wales, Australia. 17.10.1900. Budde-Lund Collection.rugifrons Budde-Lund (igi2a : 28) SYNTYPES : three females. Reg. no. 1921:10:18:2504-2505. South West Australia.Budde-Lund Collection.sulcatus Budde-Lund (igi2a : 22) SYNTYPES : one male; two females. Reg. no. 1921:10:18:2508-2510. Eradu, WestAustralia. 'Sta. 80'. 13.7.1905. Budde-Lund Collection.tomentosus Budde-Lund (i9i2a : 34) SYNTYPE : specimen in fragments. Reg. no. 1921:10:18:2511. Baba Head, Edel Land,West Australia. 7.9.1905. Budde-Lund Collection. DIPLOEXOCHUS Brandt aenigma Barnard (1932 : 372) SYNTYPE : female. Reg. no. 1933:1:25:801-802. Stella Bush, Durban, Natal, SouthAfrica. Collected by K. H. Barnard. Presented by the South African Museum. 224 R- J- LINCOLN AND J. P. ELLIS aerarius Barnard (1937 : SYNTYPES : three females. Reg. no. 1937:11:10:168-169. Steinkopf, Namaqualand,Cape Province. March 1935. Collected by R. F. Lawrence and A. J. Hesse. Presented bythe South African Museum.alberti Barnard (1932 : 352) SYNTYPES: six males; seven females. Reg. no. 1933:1:25:606-616. Zwartberg Pass,Prince Albert Division, Cape Province. 5500 ft. 1929. Collected by K. H. Barnard.Presented by the South African Museum.albescens Budde-Lund (igogb : 56) SYNTYPES : five males ; three females. Reg. no. 1921:10:18:1097-1105. Port Nolloth,Klein-Namaland, South West Africa. April 1905. Collected by L. Schultze. Budde-LundCollection.alticola Barnard (1932 : 348) SYNTYPES : one female ; two other specimens in fragments. Reg. no. 1933:1:25:581-582.Zwartberg Pass, Prince Albert Division, Cape Province, South Africa. Collected by K. H.Barnard. Presented by the South African Museum.bituberculutus Budde-Lund (1910 : n) HOLOTYPE : in fragments. Reg. no. 1921:10:18:1094. Kibonoto, Tanganyika. 1300-1800 m. 'In leaf mould.' November 1905. Collected by Y. Sjostedt. Budde-Lund Collection.canariensis (Dollfus) (1893 : 4§) [Armadillo canariensis'] SYNTYPES: one male; one female. Reg. no. 1911:11:8:9753-9754. Canary Is. Collectedby C. Alluaud. Norman Collection. SYNTYPE : one female and one other specimen, both in fragments. Reg. no. 1921:10:18:1093. Fuerteventura I., Canary Is. Collected by C. Alluaud. Budde-Lund Collection.castor Barnard (1932 : 24) SYNTYPES: nine males ; seventeen females. Reg.no. 1933:1:25:738-754. Kamiesberg,Cape Province, South Africa. 1931. Collected by K. H. Barnard. Presented by the SouthAfrican Museum.celsicauda Barnard (1932 : 366) SYNTYPES : twenty-eight males ; fifty-one females. Reg. no. 1933:1:25:755-774.Vanrhynsdorp, Cape Province, South Africa. 1931. Collected by K. H. Barnard. Presentedby the South African Museum.cingulatus Barnard (1932 : 373) SYNTYPES : one female ; one other specimen in fragments. Reg. no. 1933:1:25:804-805.Stella Bush, Durban, Natal, South Africa. 1912. Collected by K. H. Barnard. Presentedby the South African Museum.color atus Barnard (1932 : 342) SYNTYPES : seven males ; eleven females. Reg. no. 1933:1:25:533-540. Kridouw,between Citrusdal and Clanwilliam, Cape Province, South Africa. 1931. Collected by K. H.Barnard. Presented by the South African Museum.congener (Budde-Lund) (1904 : 108) [Armadillo congener'] SYNTYPES : one male ; one female ; one other specimen in fragments. Reg. no. 1921:10:18:1089-1090. Rio Nabilecche, 'in the interior of Brazil'. July 1897. Collected by G.Boggiani. Budde-Lund Collection.conisaleus Barnard (1932 : 359) SYNTYPES : seven females. Reg. no. 1933:1:25:650-656. Inchanga, Natal, South Africa.1917. Collected by K. H. Barnard. Presented by the South African Museum.crassus (Budde-Lund) (1904 : 105) [Armadillo crassus] SYNTYPE : male. Reg. no. 1921:10:18:1087. Sao Tome", Gulf of Guinea. Collected by R.Greef. Budde-Lund Collection.disjunctus Barnard (1932 : 364) SYNTYPES : one male ; nine females ; one other specimen in fragments. Reg. no.1933:1:25:730-737. Riversdale, Langeberg Range, Cape Province, South Africa, 1926-27.Collected by K. H. Barnard. Presented by the South African Museum. TYPES OF TERRESTRIAL ISOPODS 225 dugesi (Dollfus) (i8g6a : 46) [Armadillo dugesi] SYNTYPE : female. Reg. no. 1921:10:18:1104. Corritos, Silao, Mexico. Collected byDuges. Budde-Lund Collection.ecaudatus Barnard (1932 : 356) SYNTYPES : forty-seven males, females and juveniles. Reg. no. 1933:1:25:626-633.Zwartberg, Caledon, Cape Province, South Africa. 1918. Collected by K. H. Barnard.Presented by the South African Museum.festivus (Budde-Lund) (1904 : 112) [Armadillo festivus] SYNTYPES : two females and one other specimen, all in fragments. Reg. no. 1921:10:18:1106-1107. Port Elizabeth, South Africa. 15.12.1898. Collected by Brauns. Budde-Lund Collection.furcatus Barnard (1932 : 364) SYNTYPES: six males ; twenty-one females. Reg. no. 1933:1:25:720-729. River ZonderEnd Mounts, Cape Province, South Africa. Collected by K. H. Barnard. Presented by theSouth African Museum.galapagoensis (Miers) (i877a : 74) [Cubaris galapagoensis] HOLOTYPE : female. Reg. no. 1876:18:3. Santa Maria I. (formerly Charles I.), GalapagosIs., Pacific Ocean. Collected and presented by W. E. Cookson.gigas (Miers) (i877b : 666) [Cubaris gigas] SYNTYPES : two males ; one female. Reg. no. 1941:6:27:10-13. San Juan, Nicaragua,Central America. Collected by and purchased from Salle".glomus (Budde-Lund) (1898 : 8) [Armadillo glomus] SYNTYPES: three males; seven females. Reg. no. 1921:10:18:1113-1121. Zanzibar,East Africa. Collected by A. Voeltzkow. Budde-Lund Collection. SYNTYPE : male. Reg. no. 1956:10:10:139. Zanzibar I., East Africa. Collected by A.Voeltzkow. Presented by University College, Dundee.gordoniensis Barnard (1932 : 370) SYNTYPES: twenty-four males ; forty-six females. Reg.no. 1933:1:25:781-800. Kaka-mas, Cape Province, South Africa. Collected by K. H. Barnard. Presented by the SouthAfrican Museum.hypselos Barnard (1932 : 357) SYNTYPES: thirty males, females and juveniles. Reg. no. 1933:1:25:634-643. Krantzkop,Natal, South Africa. 1917. Collected by K. H. Barnard. Presented by the South AfricanMuseum.hypsinephes Barnard (1932 : 362) SYNTYPES: three males; nine females. Reg. no. 1933:1:25:675-684. Zwartberg Pass,Prince Albert Division, Cape Province, South Africa. 4000-5000 ft. 1929. Collected byK. H. Barnard. Presented by the South African Museum.kaokoensis Barnard (1932 : 333) SYNTYPES: two males ; one female. Reg. no. 1933:1:25:454-456. Kaoko Otavi, Kaoko-veld, South West Africa. Collected by K. H. Barnard. Presented by the South AfricanMuseum.kogmani Barnard (1931 : 340) SYNTYPES: two males ; sixteen females. Reg. no. 1933:1:25:503-514. Kogman's Kloof,between Ashton and Montagu, Cape Province, South Africa. Collected by K. H. Barnard.Presented by the South African Museum.limenites Barnard (1932 : 361) SYNTYPES : twenty-two males ; eighteen females ; two juveniles. Reg. no. 1933:1:25:659-674. Mossel Bay, Cape Province, South Africa. Collected by W. F. Purcell. Presentedby the South African Museum.longipes (Budde-Lund) (igogb : 55) [Armadillo longipes] SYNTYPE: female. Reg. no. 1921:10:18:1123. Okahandj a, South West Africa. Collectedby L. Schultze. Budde-Lund Collection. 226 R. J. LINCOLN AND J. P. ELLIS makuae Barnard (1932 : 360) SYNTYPE : specimen in fragments. Reg. no. 1933:1:25:657-658. Masiene, PortugueseEast Africa. Collected by R. F. Lawrence. Presented by the South African Museum.meiringi Barnard (1932 : 351) SYNTYPES : two females. Reg. no. 1933:1:25:604-605. Meirings Poort Berg, ZwartbergRange, South Africa. 6900 ft. 1932. Collected by K. H. Barnard. Presented by theSouth African Museum.montagui Barnard (1932 : 571) SYNTYPES: two males; one female. Reg. no. 1933:1:25:571-574. Montagu, Cape Pro-vince, South Africa. 1922. Collected by K. H. Barnard. Presented by the South AfricanMuseum.nanus Budde-Lund (1910 : 12) HOLOTYPE : few fragments only. Reg. no. 1921:10:18:1125. Acacia forest, at Ngare naNyuki, Mt Meru, Tanganyika. Budde-Lund Collection.nebulosus Barnard (1932 : 363) SYNTYPES: five males ; twenty-eight females. Reg. no. 1933:1:25:705-719. SwellendamMts, Langeberg Range, Cape Province, South Africa. 1925. Collected by K. H. Barnard.Presented by the South African Museum. „ obliquidens Barnard (1932 : 330) SYNTYPES : two males; two females; two other specimens in fragments. Reg. no.1933:1:25:430-435. Messina, Transvaal, South Africa. Collected by R. W. E. Tucker.Presented by the South African Museum.orphanus Barnard (1932 : 347) SYNTYPES : five females ; two other specimens in fragments. Reg. no. 1933:1:25:575-580.Weeskind (Orphan) Kop, west edge of the Kamiesberg overlooking Garies, CapeProvince, South Africa. Collected by K. H. Barnard. Presented by the South AfricanMuseum.ovampoensis (Barnard) (1924 : 232) [Cubans ovampoensis] SYNTYPES : two males ; three females ; three other specimens tightly rolled up. Reg. no.1933:1:25:425-429. Namakunde, Ovamboland, South Africa. Collected by K. H. Barnard.Presented by the South African Museum.pachytos Barnard (1932 : 337) SYNTYPES : four males ; twenty-five females ; four other specimens tightly rolled up.Reg. no. 1933:1:25:816-830. Wellington Mts, Cape Province, South Africa. Collected byK. H. Barnard. Presented by the South African Museum.pauperculus Barnard (1932 : 350) SYNTYPES : two males ; five females ; two other specimens in fragments. Reg. no. 1933:1:25:591-600. Fore Bay, nr Mossel Bay, Cape Province, South Africa. Collected by K. H.Barnard. Presented by the South African Museum.pisum (Budde-Lund) (1886 : 32) [Armadillo pisum] SYNTYPES : two males ; two females ; one other specimen in fragments. Reg. no. 1921:10:18:1133-1135. Key West, Florida, North America. Collected by E. Lonnberg. Budde-Lund Collection.pollex Barnard (1936 : 10) SYNTYPE : female. Reg. no. 1936:7:13:50. Le Pouce, Mauritius. 2400 ft. Collectedand presented by R. F. Lawrence.polythele Barnard (1932 : 350) SYNTYPES: one male; one female. Reg. no. 1933:1:25:601-603. Zwartberg Pass,Prince Albert Division, Cape Province, South Africa. 1929. Collected by K. H. Barnard.Presented by the South African Museum.quadritnaculatus Budde-Lund (igogb : 54) SYNTYPES: two males; one female in fragments. Reg. no. 1921:10:18:1138-1140.Keetmanshoop, Gross-Namaland, South West Africa. Collected by L. Schultze. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 227 rhodesiensis Barnard (1932 : 349) SYNTYPES : one male; three females; one specimen in fragments. Reg. no. 1933:1:25:583-590. Bulawayo, South Rhodesia. 'In nests of Pheidole ants.' Collected by W. R. E.Tucker. Presented by the South African Museum.rufescens Budde-Lund (igogb : 56) SYNTYPES: seven males ; six females. Reg. no. 1921:10:18:1141-1152. Klein-Namaland,Kamagas, South Africa. June 1904. Collected by L. Schultze. Budde-Lund Collection.saldanhae Barnard (1932 : 334) SYNTYPES: three males; three females. Reg. no. 1933:1:25:457-461. Saldanha Bay, CapeProvince, South Africa. 1912. Collected by K. H. Barnard. Presented by the SouthAfrican Museum.salisburyensis Barnard (1932 : 327) SYNTYPES : one male ; three females; one other specimen in fragments. Reg. no.1933:1:25:420-424. Salisbury, Rhodesia. Collected by R. W. E. Tucker. Presented bythe South African Museum.silvarum (Dollfus) (i8g6c : 393) [Armadillo silvaruni] SYNTYPES: one male; three females. Reg. no. 1896:1:11:4-7 (part). West Indies.Collected by H. H. Smith. Presented by the West India Committee. SYNTYPE : male. Reg. no. 1896:1:11:4-7 (part). Cumberland Valley, Leeward Is.1000 ft. 2 December. 'Open, damp ground, under a stone.' Collected by H. H. Smith.Presented by the West India Committee. SYNTYPES : two males ; five females. Reg. no. 1896:1:11:4-7 (part). St Vincent, Wind-ward Is. 'Pretty common under rubbish, forest below 2000 ft.' Collected by H. H. Smith.Presented by the West India Committee. SYNTYPE: male. Reg. no. 1896:1:11:4-7 (part). Leeward Is. 'Forest, dry hillside, nearChateaubelair. 1000 ft. October nth. Under stone.' Collected by H. H. Smith.Presented by the West India Committee. SYNTYPE : female. Reg. no. 1921:10:18:1153. St Vincent, Windward Is. Collected byH. H. Smith. Budde-Lund Collection.steenbrasi Barnard (1932 : 335) SYNTYPE : female. Reg. no. 1933:1:25:462-465. Mouth of Steenbras River, South ofGordon's Bay, Cape Province, South Africa. Collected by W. F. Purcell. Presented by theSouth African Museum.tabularis Barnard (1932 : 354) SYNTYPES : fifty males, females and juveniles. Reg. no. 1933:1:25:616-625. Table Mt,Cape Province, South Africa. Collected by K. H. Barnard. Presented by the South AfricanMuseum.tugelae Barnard (1932 : 353) SYNTYPES : two females ; one other specimen in fragments. Reg. no. 1933:1:25:612-615.Krantzkop, nr south bank of Tugela River, Natal, South Africa. 'In ants' nest under stones.'Collected by K. H. Barnard. Presented by the South African Museum.verrucosus (Budde-Lund) (1904 : 104) [Armadillo verrucosus] SYNTYPES : two males; two females. Reg. no. 1921:10:18:1177-1180. Guayaquil,Ecuador. 'Under bark of trees 10.7.1903.' Collected by F. Buchwald. Budde-LundCollection.zigzag (Dollfus) (i8g6c : 397) [Armadillo zigzag] HOLOTYPE : male. Reg. no. 1896:1:11:18. St Vincent, Windward Is. 'Forest, dampground under rubbish. 1000 ft.' Collected by H. H. Smith. Presented by the West IndiaCommittee.zwartbergensis Barnard (1932 : 363) SYNTYPES: fifteen males ; thirty-eight females. Reg. no. 1933:1:25:685-704. Seweweek-spoort Berg, Zwartberg Range, nr Ladysmith, Cape Province, South Africa. 1928. Collectedby K. H. Barnard. Presented by the South African Museum. 228 R. J. LINCOLN AND J. P. ELLIS GLOMERULUS Budde-Lund microps (Budde-Lund) (1904 : 116) [Armadillo microps] SYNTYPES : two males; one female. Reg. no. 1921:10:18:1181-1183. Peak Downs,Queensland, Australia. Budde-Lund Collection. MERULANA Budde-Lund bicarinata (Budde-Lund) (igisa : 65) [Sperillo (Merulana) bicarinata] SYNTYPE : female. Reg. no. 1921:10:18:807. New South Wales, Australia. Collected byMichaelsen. Budde-Lund Collection.canaliculata (Budde-Lund) (1904 : 74) [Spherillo canaliculatus] SYNTYPES : one male ; one female. Reg. no. 1921:10:18:827-828. Chatham I., PacificOcean. Collected by Schauinsland. Budde-Lund Collection.iniqua (Budde-Lund) (1904 : 74) [Spherillo iniquus] SYNTYPES: one male ; two females. Reg. no. 1921:10:18:864-866. Cape York, Queens-land, Australia. Collected by Darnel. Budde-Lund Collection.translucida (Budde-Lund) (1885 : 290) [Armadillo translucidus] HOLOTYPE : female (with two juveniles). Reg. no. 1921:10:18:928. Noumea, NewCaledonia, Pacific Ocean. Collected by Sav6s. Budde-Lund Collection.translucida gracilior Verhoeff (ig26b : 306) HOLOTYPE : female. Reg. no. 1928:7:4:16. Hienghene, New Caledonia. Purchasedfrom K. W. Verhoeff. NESODILLO Verhoeff bocki Verhoeff (i938a : 9) SYNTYPES : two females. Reg. no. 1938:7:7:56-57. Gilbert's I., Pacific Ocean. Collectedon Prof. S. Bock's Pacific Expedition, 1917-18. Purchased from K. W. Verhoeff.sarasini Verhoeff (ig26b : 280) SYNTYPE : male. Reg. no. 1928:7:4:17. New Caledonia. Purchased from K. W.Verhoeff. SYNTYPE : female. Reg. no. 1938:7:7:58. New Caledonia. Purchased from K. W.Verhoeff.scliellenbergi Verhoeff (ig28a : 211) SYNTYPE : female. Reg. no. 1928:7:4:8. Formosa. Purchased from K. W. Verhoeff. ORODILLO Verhoeff sauteri Verhoeff (ig28a : 206) SYNTYPES: one male ; one female. Reg. no. 1928:7:4:1-2. Pilam, Taiwan. Purchasedfrom K. W. Verhoeff. PERICEPHALUS Budde-Lund feae (Budde-Lund) (1895 : 606) [Armadillo feae] SYNTYPES : two males. Reg. no. 1921:10:18:2612. Tagata, Mt Mooleyit, Tenasserim,Burma. 5000-6000 m. March 1887. Collected by L. Fea. Budde-Lund Collection.galeatus (Budde-Lund) (1895 : 605) [Armadillo galeatus] SYNTYPES: one male; one other specimen in fragments. Reg. no. 1921:10:18:2613.Bhamo, Burma. July 1886. Collected by L. Fea. Budde-Lund Collection.marcidus (Budde-Lund) (1904 : 117) [Armadillo marcidus] HOLOTYPE : male. Reg. no. 1921:10:18:2614. Pulo, Penang I., Malaysia, c. 700 m.February 1897. Collected by L. Fea and Loria. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 229 POLYACANTHUS Budde-Lund transvaalensis Barnard (1932 : 321) SYNTYPE : female. Reg. no. 1933:1:25:418-419. Zoutlansberg, Transvaal. Collectedby R. F. Lawrence. Presented by the South African Museum. REDUCTONISCUS Kesselyak fritschei Verhoeff (i93ya : 415) SYNTYPES : three females. Reg. no. 1937:7:6:151-153. Berlin, East Germany. Col-lected by H. Fritsche. Purchased from K. W. Verhoeff. SAIDJAHUS Budde-Lund altimontis Jackson (1936 : 83) SYNTYPES : one male; one female. Reg. no. 1938:5:23:3-4. Pakka, North Borneo.10 ooo ft. 25.3.1929. Presented by R. E. W. Vallentin.creper Budde-Lund (1904 : 49) SYNTYPES: three males; eight females. Reg. no. 1921:10:18:939-949. Telang and Pagat,Borneo. Collected by Grabowski. Budde-Lund Collection. SPHERILLO Dana ambitiosus (Budde-Lund) (1885 : 34) [Armadillo ambitiosus] SYNTYPES: two males; one female. Reg. no. 1921:10:18:801-803. New Zealand.Budde-Lund Collection.bicoloratus (Budde-Lund) (1895 : 602) [Armadillo bicoloratus] SYNTYPES: one male ; four females ; one other specimen in fragments. Reg. no. 1921:10:18:808-813. Mt Carin, Aseinii Cheba, Burma. 1200-1300 m. January 1880. Collected byL. Fea. Budde-Lund Collection.bifrons (Budde-Lund) (1885 : 38) [Armadillo bifrons] SYNTYPES: six males; two females. Reg. no. 1921:10:18:814-821. Rockhampton,Queensland, Australia. Collected by M. Salmin. Budde-Lund Collection. SYNTYPE: female. Reg. no. 1956:10:10:118. Rockhampton, Queensland, Australia.Collected by Salmin. Presented by University College, Dundee.brachycephalus Budde-Lund (1904 : 84) HOLOTYPE (?) : fragments. Reg. no. 1921:10:18:824. Pagat, Borneo, Indonesia. Col-lected by Grabowski. Budde-Lund Collection.caligans Budde-Lund (1904 : 92) SYNTYPE : female. Reg. no. 1921:10:18:826. Bohol I., Philippine Is. Collected by Prof.A. Sempler. Budde-Lund Collection.decor at us Budde-Lund (1904 : 81) SYNTYPE : fragments. Reg. no. 1921:10:18:841. Thailand. Budde-Lund Collection.SYNTYPES : two females. Reg. no. 1921:10:18:842-843. Koh-Chang I., Thailand.Collected by T. Mortensen. Budde-Lund Collection.dispersus Budde-Lund (1904 : 70) SYNTYPES: two males. Reg. no. 1921:10:18:844-845. Botanical Gardens, Hamburg.'Among plants from Bismark Archipelago, Solomon Is.' Collected by L. Reh. Budde-LundCollection.erinaceus (Budde-Lund) (1885 : 36) [Armadillo erinaceus] SYNTYPES : two females. Reg. no. 1921:10:18:847-848. Upolu L, Samoa, Pacific Ocean.Budde-Lund Collection. 230 R. J. LINCOLN AND J. P. ELLIS frontalis (Dollfus) (1900 : 522) [nom. nov. for Armadillo danae Budde-Lund 1904 : 67] SYNTYPES : two males. Reg. no. 1904:11:5:11-12. Makaweli, Kauai I., Hawaiian Is.3000 ft. 1897. Presented by the Joint Committee for Investigating the Fauna of theSandwich Islands.grisescens Budde-Lund (1902 : 380) SYNTYPES: one male; three females. Reg. no. 1921:10:18:849-852. Aring, Kelantan,Malaysia. Collected by the Skeat Expedition. Budde-Lund Collection.grossus (Budde-Lund) (1885 : 19) [Armadillo grossus] SYNTYPES : eight females. Reg. no. 1921:10:18:853-859. Rockhampton, Queensland,Australia. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1956:10:10:119-120. Rockhampton,Queensland, Australia. Presented by University College, Dundee.hebridarum Verhoeff (ig26b : 297) SYNTYPE : female. Reg. no. 1928:7:4:18. New Hebrides, Pacific Ocean. Collected byF. Speiser. Purchased from K. W. Verhoeff.hypotoreus Jackson (1936 : 84) SYNTYPES: two females. Reg. no. 1938:5:23:5-8. Pakka, North Borneo, 10 ooo ft.23.3.1929. Presented by the Raffles Museum.ingens Budde-Lund (1904 : 91) HOLOTYPE : fragments. Reg. no. 1921:10:18:863. Bohol I., Philippines. Collected bySemper. Budde-Lund Collection.lent us Budde-Lund (1904 : 88) SYNTYPES : one male ; two females ; one other specimen in fragments. Reg. no. 1921:10:18:867-870. New Guinea. 1894. Collected by L. Lorja. Budde-Lund Collection.tnaculosus Budde-Lund (1904 : 80) HOLOTYPE : male. Reg. no. 1921:10:18:872. Silhouette I., Seychelle Is., Indian Ocean.Collected by A. Brauer. Budde-Lund Collection.tnarginatus Budde-Lund (1904 : 65) HOLOTYPE : female. Reg. no. 1921:10:18:873. Auckland, New Zealand. Collected bySuter. Budde-Lund Collection.marquesarum Jackson (1933 : 156) [Spherillo (Xestodillo ?) marquesarum] SYNTYPES : one male; three females. Reg. no. 1933:12:20:7-10. Teavanui, Uapou,Marquesan Is. 2900 ft. 27.12.1931. Collected by Le Bronnec. Presented by H. G.Jackson.montivagus (Budde-Lund) (1885 : 35) [Armadillo montivagus'] SYNTYPES : two males. Reg. no. 1921:10:18:874 & 877. Upola Is. Budde-LundCollection. SYNTYPES : one male ; one female. Reg. no. 1956:10:10:138-139. Upola I., Samoan Is.Presented by University College, Dundee.nicobaricus (Budde-Lund) (1885 : 31) [Armadillo nicobaricus] HOLOTYPE : male (fragment only). Reg. no. 1921:10:18:878. 'Pulo Milu', Nicobar I.,Bay of Bengal. Collected by the Galathea Expedition. Budde-Lund Collection.nobilis Budde-Lund (1904 : 90) HOLOTYPE (?) : female. Reg. no. 1921:10:18:879. Oshima, Riu Kiu Is., nr Okinawa Is.Budde-Lund Collection.obliquipes Budde-Lund (1904 : 85) HOLOTYPE (?) : female. Reg. no. 1921:10:18:880. Bohol I., Philippine Is. Collected bySemper. Budde-Lund Collection.parvus (Budde-Lund) (1885 : 25) [Armadillo parvus] SYNTYPES : two males. Reg. no. 1921:10:18:892-893. Silhouette, Seychelle Is., IndianOcean. Collected by Mobia. Budde-Lund Collection.pelt at us Budde-Lund (1904 : 78) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:906-907. Seychelle Is., IndianOcean. Collected by A. Brauer. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 231 perkinsi (Dollfus) (1900 : 522) [Armadillo perkinsi] HOLOTYPE : male. Reg. no. 1904:11:5:10. Haleakala, Kauai I., Hawaiian Is. 5000 ft.April 1894. Collected by Perkins. Presented by the Joint Committee for Investigating theFauna of the Sandwich Islands.purpurascens Budde-Lund (igi^b : 371) HOLOTYPE (?) : female. Reg. no. 1921:10:18:908. Isle of Pines, New Caledonia, PacificOcean. Collected by the 'Sealark' Expedition. Budde-Lund Collection.rufotnarginatus Budde-Lund (1904 : 64) HOLOTYPE : female. Reg. no. 1921:10:18:909. Taranga, New Zealand. Collected by DrThilenius. Budde-Lund Collection.rugulosus (Miers) (1876 : 225) [Cubans rugulosus'] SYNTYPES : male ; female ; two other specimens in fragments (dry). Reg. no. 1845:61.New Zealand. Presented by Dr A. Sinclair.setaceus Budde-Lund (1904 : 89) HOLOTYPE : female. Reg. no. 1921:10:18:924. Auckland, New Zealand. Budde-LundCollection.sellers Budde-Lund (1904 : 55) HOLOTYPE : female. Reg. no. 1921:10:18:925. Rqckhampton, Queensland, Australia.Budde-Lund Collection.spicatus Jackson (1927 : 4) SYNTYPES: three males ; thirteen females. Reg. no. 1928:3:8:11-20. Malololelei, Upola,Samoan Is., Pacific Ocean. 2000 ft. 25.4.1924. Collected and presented by P. A. Buxtonand G. H. E. Hopkins. SYNTYPES: two males ; four females. Reg.no. 1952:4:18:125-128. Malololelei, Upola,Samoan Is. Collected by Buxton and Hopkins. Jackson Collection.tarangensis Budde-Lund (1904 : 67) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:926. Taranga I., New Zealand.Budde-Lund Collection.weberi (Dollfus) (1907 : 364) [Armadillo weberi] SYNTYPE : male. Reg. no. 1921:10:18:1672. Ngalau Cave, nr Pajakomboh, Sumatra.Collected by M. Weber. Budde-Lund Collection. SYNARMADILLO Dollfus clausus Dollfus (i8g2a : 388) SYNTYPE : female. Reg. no. 1921:10:18:1360. Assinie, Ivory Coast, West Africa.(Locality given on label: Habessinia.) Collected by C. Alluaud. Budde-Lund Collection.globus Budde-Lund (1908 : 276) SYNTYPES: two males ; one female. Reg. no. 1921:10:18:1376-1378. Cameroons, WestAfrica. Collected by J. Sjostedt. Budde-Lund Collection. SYNTYPES: four males; eight females. Reg. no. 1921:10:18:1364-1375. Cameroons,West Africa. Collected by S. Sjostedt. Budde-Lund Collection.marmoratus Budde-Lund (1910 : 15) SYNTYPES: ten males ; fourteen females. Reg.no. 1921:10:18:1380-1391. Rainforestat Kibongoto, Kilimanjaro, Tanganyika ; rain forest at Meru, Tanganyika. Collected by Y.Sjostedt. Budde-Lund Collection. SYNTYPES : three females. Reg. no. 1921:10:18:1392-1394. Rain forest at Kibongoto,Kilimanjaro, Tanganyika. Budde-Lund Collection.nigropunctatus (Hilgendorf) (1893 : 154) [Periscyphis nigropunctatus] SYNTYPES: two males; four females. Reg. no. 1921:10:18:1395-1400. Togo, WestAfrica. Collected by Biittner. Budde-Lund Collection.pygmaeus (Budde-Lund) (1898 : 6) [Periscyphis pygmaeus] SYNTYPE : female (?) (fragments). Reg. no. 1921:10:18:1401. Runsoro. Collected byPasch and Stuhlmann. Budde-Lund Collection. 232 R. J. LINCOLN AND J. P. ELLIS simplex Budde-Lund (1910 : 16) SYNTYPES : three females. Reg. no. 1921:10:18:1402-1404. Kibongoto, Kilimanjaro,Tanganyika. 'In leaf mould.' November 1905. Collected by Y. Sjostedt. Budde-LundCollection. VENEZILLO Verhoeff dumorum (Dollfus) (18960 : 391) [Armadillo dumorum] SYNTYPES: two males; three females. Reg. no. 1896:1:11:26-27. Mustique I., WindwardIs., West Indies. 'June. Beating bushes.' 1889-91. Collected by H. H. Smith. Pre-sented by the West India Committee.grenadensis (Budde-Lund) (1893 : IJ5) [Armadillo grenadensis] SYNTYPE : male. Reg. no. 1921:10:18:1122. Grenada, Windward Is., West Indies. May1891. Collected by Meinert. Budde-Lund Collection.pumilus (Budde-Lund) (1893 : JI5) [Armadillo pumilus] SYNTYPE : female (fragments). Reg. no. 1921:10:18:1136. Venezuela. 1891. Collectedby Meinert. Budde-Lund Collection.truncorum (Budde-Lund) (1893 : II6) {Armadillo truncorum] SYNTYPE : fragments. Reg. no. 1921:10:18:1167. Venezuela. Budde-Lund Collection.SYNTYPES: three males ; one female. Reg. no. 1956:10:10:126-129. Caracas, Venezuela.Presented by University College, Dundee.venustus (Budde-Lund) (1893 : 114) {Armadillo venustus] SYNTYPES: one male; six females. Reg. no. 1921:10:18:1169-1174. La Moka, Vene-zuela. August 1891. Collected by Meinert. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1956:10:10:123. La Moka, Venezuela. August 1891.Collected by Meinert. Presented by University College, Dundee. INCERTAE SEDIS KISUMA Budde-Lund papillosa Budde-Lund (i9i2b : 169) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:5139-5140. Semarang, Java,Indonesia. December 1909. Collected by E. Jacobson. Budde-Lund Collection. OURACHAERUS Kinahan caudatus Kinahan (1859 : 198) HOLOTYPE : posterior segments missing (dry). Reg. no. 1973:514:1. Locality and donorunknown. PYRGONISCUS Kinahan cinctutus Kinahan (1859 : 200) HOLOTYPE : female (dry). Reg. no. 1856:85. Eastern Seas. Collected during the voyageof H.M.S. 'Herald'. Donor unknown. REFERENCES ARCANGELI, A. 1913. Isopode terrestre nuovi o poco noti di Italia. Monitore zool. ital. Firenze, 24 : 183-202.AUBERT, A. J. M. & DOLLFUS, A. 1890. Notice sur les Isopodes terrestres de Marseille et de Salon avec descriptions et figures d'especes nouvelles. Bull. Soc. Etud. scient. Paris, 13 : 61-70. TYPES OF TERRESTRIAL ISOPODS 233 BARNARD, K. H. 1924. Contributions to a knowledge of the fauna of South Africa. III.Crustacea Isopoda Terrestria. Ann. S. Afr. Mus. 20 : 231-236. — 1932. Contributions to the Crustacean fauna of South Africa. No. u. TerrestrialIsopoda. Ann. S. Afr. Mus. 30 (2) : 179-388. - 1936. Terrestrial Isopods and Amphipods from Mauritius. Ann. Natal Mus. 8 (i) : 1-17.— 1937. New South African woodlice (Isopoda terrestria). Ann. Natal Mus. 8 (2) : 155-165. 1940. 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Account of the zoological collection made during the visit of H.M.S. 'Peterel' to the Galapagos Islands. V. Crustacea. Proc. zool. Soc. Lond. 73-75.- i877b. On a collection of Crustacea, Decapoda and Isopoda, chiefly from South America, with descriptions of new genera and species. Proc. zool. Soc. Lond. 653-679.NICHOLLS, G. E. & BARNES, H. M. 1926. A description of two new terrestrial Isopods from Western Australia. // R. Soc. W. Aust. 12 (17) : 149-159. OMER-COOPER, J. 1923. The terrestrial Isopoda of Mesopotamia and the surrounding dis-tricts. /. Bombay nat. Hist. Soc. 29 (i) : 391-404. 1926. A revision of the genus Periscyphis Gerst. Proc. zool. Soc. Lond. 349-400. RICHARDSON, H. 1914. Terrestrial Isopods of Colombia. Mem. Soc. neuchdt. Sci. nat. 5 : 29-32.SAY, T. 1818. An account of the Crustacea of the United States. /. Acad. nat. Sci. Philad. 1 : 423-444.SILVESTRI, F. 1897. Descrizione di alcune nuove specie di Isopodi della fauna mediterranea. Annali Mus. civ. Stor. nat. Giacomo Doria, (2a) 18 : 413-416.SIMON, E. 1885. Etudes sur les Crustaces terrestres et fluviatiles recueillis en Tunisie en 1883, 1884 et 1885 par MM. A. Letourneux, M. Sedillot et Valery-Mayet. Exploration Scientifique de la Tunisie. Sciences Naturelles. Crustaces, pp. 5-12. Paris.STEBBING, T. R. R. 1900. On Crustacea from the South Seas. In: WILLEY, A. Zoological Results based upon Material from New Britain, New Guinea, Loyalty Islands and elsewhere collected during the years 1895, 1896 and 1897. Part V, No. 33 : 605-690. Cambridge.STEIN, J. P. E. F. 1859. Einige neue europaische Isopoden-Arten. Berl. ent. Z. 3 : 260- 267.STROUHAL, H. 1927. Zur Kenntnis der Untergattung Armadillidium Verh. (Isop. terr.). Zool. Anz. 74 : 5-34. 1929. Die Landisopoden des Balkans. 3. Siidbalkan. Z. wiss. Zool. 133 : 57-120. TUA, P. 1900. Contribuzione alia conoscenza degli Isopodi terrestri Italiani. Boll. Musei Zool. Anat. comp. R. Univ. Torino, 15 (374) : 1-15- 236 R. J. LINCOLN AND J. P. ELLIS VANDEL, A. 1962. Faunede France. Vol. 66. Isopodes terrestres (DeuxiemePartie), pp. 415-931. Paris. — 1963. Isopodes terrestres recueillis en Amerique du Sud par Claude Delamare Deboutte-ville. Biol. Am. Aust. 2 : 63-100. — 1968. I. Isopodes terrestres. Mission zoologique beige aux lies Galapagos et Ecuador (N.et J. Leleup, 1964-1965), 1 : 37-168. -1970. L'origine et 1'evolution des Trachelipidae (Crustacea; Isopoda ; Oniscoidea). Bull. Soc. zool. Fr. 95 : 321-328.VERHOEFF, K. W. igoia. Uber palaarkitsche Isopoden (3. Aufsatz). Zool. Anz. 24 : 33-41. — igoib. Uber palaarktische Isopoden (4. Aufsatz). Zool. Anz. 24 : 66-72, 73-79. — 19010. Uber palaarktische Isopoden (5. Aufsatz). Zool. Anz. 24 : 135-149. Uber palaarktische Isopoden (7. Aufsatz). Zool. Anz. 24 : 403-408, 417-421.Uber Isopoden. 10. Aufsatz. Zur Kenntnis der Porcellioniden (Kornerasseln). Sber. Ges. naturf. Freunde Berl. no. 8 : 229-281. i907b. Uber palaarktische Isopoden. 9. Aufsatz. Neuer Beitrag zur Kenntnis derGattung Armadillidium. Zool. Anz. 31 : 457-505. igoSa. Uber Isopoden. 15. Isopoden- Aufsatz. Arch, biontol. 2 : 335-387. igoSb. Uber Isopoden. 12. Isopoden- Aufsatz. Neue Oniscoidea aus Mittel- undSiideuropa und zur Klarung einiger bekannter Forrnen. Arch, naturgesch. 74 : 163-198. igoSc. Uber Isopoden. 14. Aufsatz. Armadillidium-Arten, mit besonderer Beriicksich-tigung der in Italien und Sizilien einheimischen. Zool. Anz. 33 : 450-462, 484-492. 1910. Uber Isopoden. 16. Aufsatz, Armadillidium und Porcellio an der Riviera. Jh.Ver. vaterl. Naturk. Wiirtt. 115-143. 1917. Uber mediterrane Oniscoideen, namentlich Porcellioniden. 23. Isopoden- Aufsatz.Jh. Ver. vaterl. Naturk. Wiirtt. 73 : 144-173. 1918. Zur Kenntnis der Ligidien, Porcellioniden und Onisciden. 24. Isopoden- Aufsatz.Arch, naturgesch. 82A (10) : 108-169. i926a. Uber Isopoden der Balkanhalbinsel, gesammelt von Herrn. Dr. I. Buresch. 31.Isopoden- Aufsatz. Izv. bulg. ent. Druzh. 3 : 135-158. i926b. Isopoda terrestria von Neu-Caledonien und den Loyalty-Inseln. 32. Isopoden-Aufsatz. In: SARASIN, F. & Roux, J. Nova Caledonia, A. Zoologie, IV, pp. 243-366.Miinchen. 1927. Uber einige zoogeographisch bedeutsame Isopoden-Arten. 36. Isopoden- Aufsatz.Zool. Anz. 73 : 323-333. ig28a. Isopoden aus Formosa. 39. Isopoden- Aufsatz. Mitt. zool. Mus. Berl. 14 : 200-226. I928b. Uber einige Isopoden der zoologischen Staatsammlung in Miinchen. 38. Isopoden-Aufsatz. Zool. Anz. 76 : 25-36, 113-123. I928c. Uber alpenlandische und italienische Isopoden. 37. Isopoden-Aufsatz. Zool. Jb.(Abt. Syst.) 56 : 93-172. 1929. Uber Isopoden der Balkanhalbinsel, gesammelt von Herrn. Dr. I. Buresch. 33.Isopoden-Aufsatz. Izv. tsarsk. pirodonauch. Inst. Sof. 2 : 129-139. i93oa. Uber einige neue norditalienische Isopoden und einen neuen Typus der Volvation.43. Isopoden-Aufsatz. Zool. Anz. 89 : 162-177. i93ob. Uber Isopoden aus Turkestan. 42. Isopoden-Aufsatz. Zool. Anz. 91 : 101-125. I93QC. Zur Kenntnis osteuropaischer Isopoden. 41. Isopoden-Aufsatz. Zool. Jb.(Abt. Syst.) 59: 1-64. i93ia. Uber Isopoden terrestria aus Italien. 45. Isopoden-Aufsatz. Zool. Jb. (Abt.Syst.) 60 : 489-572. 193 ib. Zur Kenntnis alpenlandischer und mediterraner Isopoda terrestria. 47. Isopoden-Aufsatz. Zool. Jb. (Abt. Syst.) 62 : 15-52. 1933 • Neue Isopoda terrestria aux Mexiko und dem Mediterrangebiet. 50. Isopoden-Aufsatz. Zool. Anz. 103 : 97-119. i936a. Stiidien iiber Isopoda-terrestria. 51. Isopoden-Aufsatz. Mitt. zool. Mus. Berl.21 : 79-163. TYPES OF TERRESTRIAL ISOPODS 237 VERHOEFF, K. W. i936b. t)ber einige Myriapoden und Isopoden aus Dekan, gesammelt von Herrn. S. Jones, Madras. Rec. Indian Mus. 33 : 503-512. I937a. t)ber einige neue und bekannte Isopoda terrestria. 61. Isopoden-Aufsatz. Sber. Ges. naturf. Freunde Berl. pp. 411-430. I93?b. Uber einige Porcellio-Arten aus Marokko gesammelt von Herrn. J. de Le'piney. Zool. Anz. 119 : 302-307.— I938a. t)ber einige polynesische Oniscoideen von Prof. Sixten Bock's Pazifik-Expedition 1917-1918. Arch. Zool. 30A (12) : 1-14. I938b. t)ber Land-Isopoden aus Marokko. II. Teil. Zool. Anz. 124 : 61-67. 1939. Diplopoden, Chilopoden und Oniscoideen, hauptsachlich aus siiditalienischen Hohlen. Zool. Jb. Syst. 72 : 203-224.VOGL, C. VON. 1875. Beitrag zur Kenntnis der Land-Isopoden. Verh. zool.-bot. Ges. Wien. 25 :503-518.WHITE, A. 1847. List of the Specimens of Crustacea in the Collection of the British Museum. 143 pp. London. APPENDIX i The Collection also contains a large number of specimens labelled as types, mostlyfrom the Budde-Lund Collection, for which no reference can be found in publishedliterature. It seems possible that these are specimens with manuscript nameswhich have been incorporated into personal collections but which have never beenpublished. A list of this material comprising 167 items is given below ; and a listof the manuscript names has been deposited in the library of the British Museum(Natural History). Family PORCELLIONIDAE Agabiforius sp. Three specimens. Reg. no. 1921:10:18:4066-4068. Fundo I., Zanzibar. 21.4.1903. Collected by C. W. Pemba (?). Budde-Lund Collection.Hemilepistus sp. Two specimens. Reg. no. 1921:10:18:4113-4114. Asia. Budde-Lund Collection.Leptotrichus sp. One specimen. Reg. no. 1921:10:18:3969. Zanzibar I. 20.5.1888. Collected by Struhl- mann. Budde-Lund Collection.Lucasius sp. Five specimens. Reg. no. 1921:10:18:5177-5181. Philippeville. Budde-Lund Collection.Metoponorthus sp. One specimen. Reg. no. 1921:10:18:5224. Inca, Malorca. Budde-Lund Collection.Metoponorthus sp. One specimen. Reg. no. 1921:10:18:5238. Taurus, Cilicia, Turkey. Budde-Lund Collection.Metoponorthus sp. One specimen. Reg. no. 1921:10:18:5246. Corral, Chile. Budde-Lund Collection.Metoponorthus sp. Two specimens. Reg. no. 1921:10:18:5288-5289. Mt Grappa, Italy. Budde-Lund Collec-tion.Metoponorthus sp. Three specimens. Reg. no. 1921:10:18:5290-5295. Boccadifalco, Italy. 17.2.1896. Col-lected by F. Silvestri. Budde-Lund Collection.Metoponorthus sp. Two specimens. Reg. no. 1921:10:18:5323-5324. East Africa. Collected by Gadat. Budde-Lund Collection. 238 R. J. LINCOLN AND J. P. ELLIS Metoponorthus sp. Six specimens. Reg. no. 1921:10:18:5369-5374. Giilek, Cilicia, Turkey. Budde-Lund Collection.Metoponorthus sp. One specimen. Reg. no. 1921:10:18:5488. Jerusalem, Israel. Budde-Lund Collection.Metoponorthus sp. Six specimens. Reg. no. 1921:10:18:5502-5507. Poizo. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1921:10:18:4241. Monchique, Portugal. 3.5.1888. Collected by Holier. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1921:10:18:3790. Algeria. Collected by E. Simon. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1921:10:18:4259. Asia Minor. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1925:7:452. Siebenburgen. Koch Collection.Porcellio sp. Four specimens. Reg. no. 1921:10:18:4280-4283. Sa. do Gerez Mts, Portugal. June 1899. Collected by Vicira. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1921:10:18:4287. Hercegowina. Budde-Lund Collection.Porcellio sp. One specimen. Reg. no. 1921:10:18:4982. Algeria. 'In monte Djebel-Mahid 2000 m inl. cortice Cedri 1885.' Collected by E. Simon. Budde-Lund Collection.Porcellio sp. Two specimens. Reg. no. 1921:10:18:4295-4296. Mauritius I., Indian Ocean. Collected by Dr Emmery. Budde-Lund Collection.Porcellio sp. Three specimens. Reg. no. 1891:11:15:1-3. Tsu-Shima. Collected by Hoist. Determined/ named by G. Budde-Lund. Purchased from H. Seebohm.Porcellio sp. Ten specimens. Reg. no. 1907:4:30:11-20. Rio de Oro, Morocco. February 1902. Col-lected by W. Riggenbach. Determined/named by G. Budde-Lund. Presented by W. Rothschild. Five specimens. Reg. no. 1921:10:18:4361-4364. Rio de Oro, Sahara. Budde-Lund Col-lection.Porcellio sp. One specimen (dry). Reg. no. 1840:6:26:731. Cape of Good Hope. Purchased from Dr Krauss.Porcellio sp. Three specimens. Reg. no. 1931:4:27:58-60. Palestine. Purchased from K. W. Verhoeff.Porcellio sp. Five specimens. Reg. no. 1921:10:18:4420-4424. Tunis. 27.3.1896. Collected by F. Silvestri. Budde-Lund Collection.Porcellio sp. Two specimens. Reg. no. 1845:112. Norfolk I., Pacific Ocean. Presented by George Newport. Two specimens (dry). Reg. no. 1856:105. Norfolk I., Pacific Ocean. June 1855. Collected during the voygage of H.M.S. 'Herald'. Donor unknown.Porcellio sp. Twelve specimens. Reg. no. 1921:10:18:4441-4452. Coimbra, Portugal. April 1896. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 239 Porcellio sp. Four specimens. Reg. no. 1921:10:18:4462-4465. Naxos I., Greece. Budde-Lund Collec-tion. Porcellio sp. Three specimens. Reg. no. 1921:10:18:3900-3902. Biskra, Algeria. Budde-Lund Collec-tion. Porcellio sp. Two specimens. Reg. no. 1921:10:18:4501-4502. Japan. Collected by Hilgendorf. Budde-Lund Collection. Porcellio sp. Two specimens. Reg. no. 1921:10:18:3904-3905. Jebel Tarhuna Mts, Libya. Budde-Lund Collection. Porcellio sp. One specimen. Reg. no. 1921:10:18:5153. Santorin. Budde-Lund Collection. Threespecimens. Reg. no. 1921:10:18:5154-5156. Favorita. Collected by Silvestri. Budde-Lund Collection. Porcellio sp. Four specimens. Reg. no. 1921:10:18:5157-5160. Palermo, Italy. Budde-Lund Collection. Porcellio sp. One specimen. Reg. no. 1921:10:18:4599. Cansiglio. Budde-Lund Collection. Porcellio sp. One specimen (dry). Reg. no. 1973:480:1. Locality and donor unknown. Porcellio sp. One specimen. Reg. no. 1921:10:18:4887. Algeria. Collected by Beresford. Budde-LundCollection. Porcellio sp. Five specimens (dry). Reg. no. 1845:49. New South Wales, Australia. Purchased fromMr Wood. Porcellio sp. Three specimens (dry). Reg. no. 1973:481:3. Tristan d'Acunha. Presented by CaptainCarmichael. Porcellio sp. Four specimens. Reg. no. 1921:10:18:4948-4951. Liguria, Italy. September 1897. Col-lected by G. Mantero. Budde-Lund Collection. Uramba sp.One specimen. Reg. no. 1921:10:18:4069. No locality. Budde-Lund Collection. Uramba sp. Three specimens. Reg. no. 1921:10:18:4081-4083. Kibwezi, Kenya. Budde-Lund Collec-tion. Uramba sp.One specimen. Reg. no. 1921:10:18:4084. Gansilla, East Africa. Budde-Lund Collection. Family TRAGHELIPIDAE Nagurus sp. Six specimens. Reg. no. 1921:10:18:5741-5746. Japan. Collected by Hilgendorf. Budde-Lund Collection.Nagurus sp. Five specimens. Reg. no. 1921:10:18:1486-1490. Koh-Kahdat. 9.1.1900. Collected by 'Th. Mort.' Budde-Lund Collection.Phalaba sp. One specimen. Reg. no. 1921:10:18:2104. East Africa. Budde-Lund Collection. Two specimens. Reg. no. 1921:10:18:2105-2106. Amani, Tanganyika. Collected by Vosseler. February 1905. Budde-Lund Collection. 24o R. J. LINCOLN AND J. P. ELLIS Phalaba sp. One specimen. Reg. no. 1921:10:18:2107. Ukinga. Budde-Lund Collection.Phalaba sp. Two specimens. Reg. no. 1921:10:18:2108-2109. Amani, Tanganyika. Budde-Lund Collection. Family ARMADILLIDIIDAE Armadillidium sp. Three specimens. Reg. no. 1921:10:18:2683-2685. Hamman Meskhoutin. Budde-Lund Collection. Twelve specimens. Reg. no. 1921:10:18:2686-2689. Constantine, Algeria. 23.3.1901. Collected by M. Kraepelin. Budde-Lund Collection. Three specimens. Reg. no. 1921:10: 18:2698-2700. Algier. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:2701. Stora, Sweden. Collected by E. Simon. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:2702. Ainos, Kefallinia I., Greece. Collected by E. von Oertzen. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:2703-2704. Cyrenaica, Libya. Budde-Lund Collec-tion.Armadillidium sp. One specimen. Reg. no. 1921:10:18:2712. Tunis, Tunisia. Budde-Lund Collection.Armadillidium sp. Six specimens. Reg. no. 1921:10:18:2714-2719. Louk el Arba. Collected by Silvestri. Budde-Lund Collection. One specimen. Reg. no. 1921:10:18:2720. Baboneh. Collected by Silvestri. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:2791-2792. Algiers. Collected by de Gaulle. Budde-Lund Collection. Five specimens. Reg. no. 1921:10:18:2793-2797. Sidi Bel Akaren. Collected by F. Silvestri. Budde-Lund Collection.Armadillidium sp. Twelve specimens. Reg. no. 1921:10:18:2842-2853. San Cataldo. Italy. 1898. Col-lected by Dr Peracca. Budde-Lund Collection.Armadillidium sp. Seven specimens. Reg. no. 1921:10:18:2936-2942. Santa Ninfa, Sicily. 15.3.1896. Collected by F. Silvestri. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:2943. Santa Ninfa, Sicily. Collected by F. Silvestri. Budde-Lund Collection.Armadillidium sp. Five specimens. Reg. no. 1921:10:18:2962-2966. Tunis. 27.3.1896. Budde-Lund Collec-tion.Armadillidium sp. One specimen (dry). Reg. no. 1973:479:1. Sierra Leone. Presented by the Rev. D. S. Morgan.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:2980-2981. Baboneh. Collected by F. Silvestri. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 241 Armadillidium sp. One specimen. Reg. no. 1921:10:18:2982. Capella de S. Antonio (Finalborge). 4.3.1898. Collected by R. Gestro. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:2983-2984. Ulai (?). Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:2985. Amos, Kefallina, Greece. Collected by E. von Oertzen. Budde-Lund Collection.Armadillidium sp. Six specimens. Reg. no. 1921:10:18:2986-2991. Palmi, Italy. 17.5.1893. Scilla, Italy. 24.6.1893. Collected by H. J. Hansen. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:5849. Asia Minor. Collected by Dr Th. Krsyser. (Mus. Berlin.) Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:2993-2994. Corfu I. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3022. Malosin. 5.8.1902. Collected by Mrazek. Budde-Lund Collection. Three specimens. Reg. no. 1921:10:18:3023-3025. Danilov Grad, Yugoslavia. Collected by Mrazek. Budde-Lund Collection. Twelve specimens. Reg. no. 1921:10:18:3026-3037. Celinje, Yugoslavia. Collected by Mrazek. Budde-Lund Collection.Armadillidium sp. Three specimens. Reg. no. 1921:10:18:3082-3084. Zadar (Zara), Yugoslavia. Collected by A. Dollfus. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3085. Eaux Counet, Pyrenees. Collected by Seidtl. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1956:10:10:135-136. Key West, Florida, USA. Presented by University College, Dundee (ex Budde-Lund Collection).Armadillidium sp. One specimen. Reg. no. 1921:10:18:3153. Corfu I. Collected by Gracia (?). Budde-Lund Collection. Five specimens. Reg. no. 1921:10:18:3154-3158. Atri, Italy. Collected by F. Silvestri. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3159. Dalmatia, Yugoslavia. Budde-Lund Collection.Armadillidium sp. Three specimens. Reg. no. 1921:10:18:3160-3162. Meleda (?), Yugoslavia. 13.8.1880. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3163. Corsica. Collected by E. Simon. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:3185-3186. Roumania. Collected by Jaquet. Budde-Lund Collection.Armadillidium sp. c. sixty specimens. Reg. no. 1921:10:18:3187-3210. Njegos, Montenegro, Yugoslavia. Collected by Mrazek. Budde-Lund Collection. Six specimens. Reg. no. 1921:10:18:3211-3216. Podgornica (now Titograd), Yugoslavia. Collected by Mrazek. Budde-Lund Collection. 13 242 R. J. LINCOLN AND J. P. ELLIS Armadillidium sp. Five specimens. Reg. no. 1921:10:18:3217-3221. Yugoslavia. Budde-Lund Collection.Armadillidium sp. Three specimens. Reg. no. 1921:10:18:3223-3225. Zaragoza (Saragossa), Spain. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:3226-3227. Flumentorgne. Budde-Lund Collection.Armadillidium sp. Five specimens. Reg. no. 1921:10:18:3241-3245. Brescia, Italy. Budde-Lund Collection.Armadillidium sp. Three specimens. Reg. no. 1921:10:18:3246-3248. Spezia, Italy. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3264. Yugoslavia. Collected by Uljanin. Budde-Lund Collection.Armadillidium sp. Seven specimens. Reg. no. 1921:10:18:3275-3281. Bocadifalco, Italy. 17.2.1896. Col-lected by F. Silvestri. Budde-Lund Collection. Twenty-two specimens. Reg. no. 1921:10:18:3265-3278 & 3282-3287. Bivona, Sicily. February 1896. Collected by F. Silvestri. Budde-Lund Collection.Armadillidium sp. One specimen. Reg. no. 1921:10:18:3296. Kieff (Kiyev), Ukraine, USSR. Collected by Uljanin. Budde-Lund Collection. Four specimens. Reg. no. 1921:10:18:3297-3301. Sympheropolis. Collected by Uljanin. Budde-Lund Collection.Armadillidium sp. Two specimens. Reg. no. 1921:10:18:3302-3303. Sympheropolis. Collected by Uljanin. Budde-Lund Collection.Armadillidium sp. Six specimens. Reg. no. 1895:1:24:41-46. Cettinje. Named/determined by G. Budde-Lund. Presented by Dr Werner.Armadillidium sp. Four specimens. Reg. no. 1921:10:18:3312-3315. Carthajine. Collected by F. Silvestri. Budde-Lund Collection. Ten specimens. Reg. no. 1921:10:18:3316-3325. Tabarka, Tunisia. 2.8.1896. Collected by F. Silvestri. Budde-Lund Collection. Five specimens. Reg. no. 1921:10:18:3326-3330. Tunis, Tunisia. 27.3.1896. Collected by Silvestri. Budde-Lund Collection. Five specimens. Reg. no. 1921:10:18:3331-3334. Baboneh. Budde-Lund Collection.Eluma sp. Three specimens. Reg. no. 1921:10:18:3760-3761. Oran, Algeria. May 1883. Collected by E. Simon. Budde-Lund Collection.Toradjia sp. One specimen. Reg. no. 1921:10:18:1660. Ceylon. Budde-Lund Collection.Toradjia sp. One specimen. Reg. no. 1921:10:18:1661. East Java. Budde-Lund Collection. Family EUBELIDAE Eubelum sp. One specimen. Reg. no. 1907:5:6:15. Roromo, Ukinga Forest. 7700 ft. Named/ determined by G. Budde-Lund. Presented by R. Cowestay (? or Crawshay). One specimen. Reg. no. 1921:10:18:588. Nawobi (? probably Nairobi). Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 243 Eubelum sp. Four specimens. Reg. no. 1921:10:18:591-594. Gardulla, Ethiopia. Budde-Lund Collec-tion.Eubelum sp. Two specimens. Reg. no. 1921:10:18:595-596. S. Kaffa, Ethiopia. Budde-Lund Collec-tion.Eubelum sp. Three specimens. Reg. no. 1921:10:18:614-616. Ethiopia. Budde-Lund Collection.Eubelum sp. One specimen. Reg. no. 1921:10:18:617. Kaffa, Ethiopia. Budde-Lund Collection.Eubelum sp. One specimen. Reg. no. 1921:10:18:618. San Salvador. Budde-Lund Collection.Eubelum sp. One specimen. Reg. no. 1921:10:18:626. Kaffa, Ethiopia. Budde-Lund Collection.Eubelum sp. Twelve specimens. Reg. no. 1921:10:18:627-638. Gardulla, Ethiopia. 2500-2800 m. 13.1.1901. Collected by O. Neumann. Budde-Lund Collection.Mesarmadillo sp. One specimen. Reg. no. 1921:10:18:729. Niger Delta, W. Africa. Budde-Lund Collection.Mesarmadillo sp. One specimen. Reg. no. 1921:10:18:730. East Africa. Budde-Lund Collection.Mesarmadillo sp. One specimen. Reg. no. 1921:10:18:731. East Africa. Budde-Lund Collection.Mesarmadillo sp. One specimen. Reg. no. 1921:10:18:732. East Africa. Budde-Lund Collection.Microcercus sp. Four specimens. Reg. no. 1921:10:18:1230-1233. Bissao. Collected by H. Ehrhardt (Mus. Hamburg). Budde-Lund Collection.Microcercus sp. Two specimens. Reg. no. 1921:10:18:1268-1269. Langenburg, W. Germany. 27.4.1898. Collected by Fullborn. Budde-Lund Collection.Microcercus sp. Two specimens. Reg. no. 1893:11:9:9-10. Woods near Ngatana. Named/determined by G. Budde-Lund. Collected and presented by J. A. Gregory.Microcercus sp. Four specimens. Reg. no. 1921:10:18:1270-1273. Nakuru, Kenya. Collected by C. Alluaud. Budde-Lund Collection.Microcercus sp. Eight specimens. Reg. no. 1902:12:4:8-15. Cheradguea Mts and its slopes, Nyasaland. July and August 1895. Collected by A. Whyte. Named/determined by G. Budde-Lund. Presented by H. H. Johnston. Eleven specimens. Reg. no. 1921:10:18:1274-1284. Nyasaland. Budde-Lund Collection.Periscyphis sp. One specimen. Reg. no. 1921:10:18:1303. East Africa. Budde-Lund Collection.Periscyphis sp. Five specimens. Reg. no. 1921:10:18:1304-1308. Ethiopia. Budde-Lund Collection.Periscyphops sp. One specimen. Reg. no. 1921:10:18:745. Africa. Budde-Lund Collection.Periscyphops sp. Two specimens. Reg. no. 1921:10:18:746-747. Shoa, Ethiopia. Budde-Lund Collection.Periscyphops sp. One specimen. Reg. no. 1921:10:18:748. No locality. Budde-Lund Collection.Periscyphops sp. One specimen. Reg. no. 1921:10:18:754. East Africa. Budde-Lund Collection. 244 R- J- LINCOLN AND J. P. ELLIS Periscyphops sp. Three specimens. Reg. no. 1921:10:18:757-759. S. Kaffa, Ethiopia. Budde-Lund Collec-tion. Periscyphops sp. Four specimens. Reg. no. 1921:10:18:764-767. Niger delta at Wari. Budde-Lund Collec-tion. Periscyphops sp. One specimen (in fragments). Reg. no. 1921:10:28:768. Misahohe, West Africa. Collectedby Baumann, Budde-Lund Collection. Family SPHAERONISCIDAE Scleropactes sp. Five specimens. Reg. no. 1921:10:18:956-960. Ecuador. Budde-Lund Collection.Scleropactes sp. Two specimens. Reg. no. 1921:10:18:965-967. Ecuador. Budde-Lund Collection.Sphaeroniscus sp. One specimen (in fragments). Reg. no. 1921:10:18:2215. Bogota, Colombia. Collected by Burger. Budde-Lund Collection.Sphaeroniscus sp. Three specimens. Reg. no. 1921:10:18:2216-2218. Costa Rica, South America. Budde-Lund Collection.Sphaeroniscus sp. One specimen (in fragments). Reg. no. 1921:10:18:2219. Bogota, Colombia. Collected by Burger. Budde-Lund Collection.Sphaeroniscus sp. One specimen (in fragments). Reg. no. 1921:10:18:2220. Peru. Budde-Lund Collection.Sphaeroniscus sp. One specimen. Reg. no. 1921:10:18:2221. Paramaribo, Surinam, South America. Budde-Lund Collection. Four specimens. Reg. no. 1921:10:18:2222-2225. Ecuador. Budde-Lund Collection.Sphaeroniscus sp. Six specimens. Reg. no. 1921:10:18:2226-2231. Venezuela. Budde-Lund Collection. Family ARMADILLIDAE Adinda sp. Two specimens. Reg. no. 1921:10:18:1665-1666. Singapore. Collected by H. N. Ridley. Budde-Lund Collection.Adinda sp. One specimen. Reg. no. 1921:10:18:1667. Sicbolangst (?). Collected by Jachan (?). Budde-Lund Collection.Adinda sp. Two specimens. Reg. no. 1921:10:18:1668-1669. Singapore. 1898. Collected by H. N. Ridley. Budde-Lund Collection.Adinda sp. One specimen (in fragments). Reg. no. 1921:10:18:1670. Sumatra. Budde-Lund Collection.Adinda sp. One specimen. Reg. no. 1921:10:16:1671. Pulo Penang, Burma. February 1879. Col-lected by Fea and Loria. Budde-Lund Collection.Armadillo sp. Two specimens. Reg. no. 1921:10:18:2462-2463. Ceylon. Budde-Lund Collection.Armadillo sp. Four specimens. Reg. no. 1921:10:18:2421-2424. South West Australia. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 245 Armadillo sp. One specimen. Reg. no. 1893:12:15:1-8 (part). North Clin Hills, Upper Burma. Named/ determined by G. Budde-Lund. Presented by E. G. Watson.Armadillo sp. One specimen (dry). Reg. no. 1842:22. Philippine Is. Purchased from Mr Cuming.Armadillo sp. One specimen. Reg. no. 1921:10:18:2516. Hackgall, Ceylon. Collected by A. Willey. Budde-Lund Collection. One specimen. Reg. no. 1907:4:30:22. Hackgall, Ceylon. January 1906. Named/deter-mined by G. Budde-Lund.Bethalus sp. Three specimens. Reg. no. 1921:10:18:1185-1187. Fort Dauphin, Madagascar. Budde-Lund Collection.Bethalus sp. One specimen. Reg. no. 1921:10:18:1188. Fort Dauphin, Madagascar. Budde-Lund Collection.Bethalus sp. Two specimens. Reg. no. 1921:10:18:1202-1203. Fort Dauphin, Madagascar. Budde-Lund Collection.Bethalus sp. Nine specimens. Reg. no. 1921:10:18:1209-1217. Fort Dauphin, Madagascar. Budde-Lund Collection.Diploexochus sp. One specimen (in fragments). Reg. no. 1921:10:18:1088. Uvinza, Tanganyika. 16.12.1896. Collected by H. Fremann. Budde-Lund Collection.Diploexochus sp. Thirteen specimens. Reg. no. 1921:10:18:1154-1166. Anjouan I., Madagascar. Budde-Lund Collection.Sperillo sp. Three specimens. Reg. no. 1921:10:18:804-806. Tonkin, Montes Mauson. 2000-3000 ft. Collected by H. Fruhstorfer. Budde-Lund Collection.Sperillo sp. Two specimens. Reg. no. 1921:10:18:822-823. Rio de Janeiro, Brazil. Budde-Lund Collection.Sperillo sp. One specimen. Reg. no. 1921:10:18:825. Tonkin, Montes Mauson. Collected by H. Fruhstorfer. Budde-Lund Collection.Spherillo sp. Fifteen specimens. Reg. no. 1921:10:18:829-840. California, USA. Collected by G. Eisen. Budde-Lund Collection.Spherillo sp. One specimen. Reg. no. 1921:10:18:846. Honolulu, Hawaii. 1874. Collected by d'Alber- tin. Budde-Lund Collection.Spherillo sp. Three specimens. Reg. no. 1921:10:18:1405-1407. Togo. Collected by Dr Biittner. Budde-Lund Collection. INCERTAE SEDISGenus iSpecies a One specimen. Reg. no. 1921:10:18:796. Uvinza, Tanganyika. Budde-Lund Collection.Genus 2Species a One specimen. Reg. no. 1921:10:18:794. Shoa district, Ethiopia. Budde-Lund Collection. 246 R. J. LINCOLN AND J. P. ELLIS Genus 3Species a One specimen. Reg. no. 1921:10:18:5704. Asia Minor. 23.10.1900. Budde-Lund Collec-tion.Species b Two specimens. Reg. no. 1921:10:18:5701-5702. Genargentu. 26.1.1899. Collected by G. Doria. Budde-Lund Collection.Genus 4Species a One specimen. Reg. no. 1921:10:18:5703. Sidi-Bel-Akaren, Algeria. Collected by F. Silvestri. Budde-Lund Collection.Genus 5Species a Two specimens. Reg. no. 1921:10:18:797-798. South Shoa, Ethiopia. Budde-Lund Collection.Genus 6Species a One specimen. Reg. no. 1921:10:18:5706. Paramaibo. Budde-Lund Collection.Genus 7Species a One specimen (in fragments). Reg. no. 1921:10:18:799. Ruanda, East Africa. Budde-Lund Collection.Genus 8Species a Fifteen specimens. Reg. no. 1921:10:18:5707-5721. Brasilia, Brazil. Budde-Lund Collec-tion.Species b One specimen. Reg. no. 1921:10:18:5722. Rio de Janeiro, Brazil. Budde-Lund Collection.Species c Two specimens. Reg. no. 1921:10:18:5723-5724. Rio de Janeiro, Brazil. Budde-Lund Collection.Genus 9Species a Three specimens. Reg. no. 1921:10:18:5726-5728. East Africa. Collected by Langeberg. Budde-Lund Collection.Species b One specimen. Reg. no. 1921:10:18:5725. Nyasa, Malawi. Budde-Lund Collection.Genus 10Species a One specimen (in fragments). Reg. no. 1921:10:18:795. Shoa district, Ethiopia. 10.11.1900. Collected by Erlanger. Budde-Lund Collection.Genus nSpecies a One specimen. Reg. no. 1921:10:18:5747. Futschau. Budde-Lund Collection.Species b One specimen. Reg. no. 1921:10:18:5748. Tsing tua, Hong Kong. Budde-Lund Collection.Species c One specimen. Reg. no. 1921:10:5749. Japan. Collected by Hilgendorf. Budde-Lund Collection.Genus 12Species a One specimen. Reg. no. 1921:10:18:800. Gardulla, Ethiopia. Budde-Lund Collection. INDEX abnormis (Gelsana) 214 abyssinicus (Microcercus) 215 Acaeroplastes 193 Acanthoniscus 219 Actoecia 218 ACTOECIIDAE 218 acutiserra (Porcellio) 197 Adinda 244 aegaeum (Armadillidium) 208 agaeus (Porcellio) 197 aenigma (Diploexochus) 223 aerarius (Diploexochus) 224 affinis (Armadillo) 219 affinis (Cubaris) 219 Agabiformius 193, 237 Agnara 205 Akermania 219 alassiense (Armadillidium) 208, 212 albanicum (Armadillidium) 208 alberti (Diploexochus) 224 albescens (Diploexochus) 224 albicornis (Lucasius) 197 albicornis (Mesarmadillo) 215 albicornis (Porcellio) 197 albifrons (Armadillidium) 208 albinus (Porcellio) 198 albipennis (Tura) 204 albomaculatus (Buddelundia) 222 albomarginatus (Armadillo) 219 albomarginatus (Porcellio) 198 albospinosus (Armadillo) 219 album (Armadillidium) 208 alluaudii (Porcellio) 198 alticola (Diploexochus) 224 altimontis (Saidjahus) 229 ambiguus (Armadillo) ambitiosus (Armadillo) 229 ambitiosus (Spherillo) 229 americanum (Ethelum) 213 americanus (Mesarmadillo) 213 amoenus (Porcellio) 198 Anchicubaris 219 anconanum (Armadillidium) 208 anconanus (Metoponorthus pruinosus) 197 angusta (Tura) 204 angustulus (Porcellio) 198 annulicornis (Cylisticus) 192 apenninorum (Armadillidium) 208 apenninorum (Trachelipus) 207 apenninorum (Tracheoniscus) 207 apfelbecki (Armadillidium) 208 approximatus (Metoponorthus) 196 apulicus (Trachelipus) 207 apulicus (Tracheoniscus) 207 arcuatus (Porcellio) 202 argentarium (Armadillidium) 208 argentarius (Acaeroplastes) 193 argentarius (Metopornorthus) 193 ARMADILLIDAE 219, 244 ARMADILLIDIIDAE 208, 240 Armadillidium 208, 240, 241, 242 Armadillo 219, 244, 245 armatus (Benechinus) 213 asifensis (Metoponorthus) 197 asifensis (Metoponorthus sexfasciatus) 197 assimile (Armadillidium) 209 astriger (Armadillidium) 208 astriger (Armadillo) 208 ater (Porcellio) 198 aternanus (Trachelipus larii) 207 aternanus (Tracheoniscus larii) 207 auritus (Porcellio) 198 ausseli (Armadillo) 219 badium (Armadillidium) 208 badium siculorum (Armadillidium) 208 BALLONISCIDAE 205 Balloniscus 205 balticus (Porcellio) 207 balticus (Trachelipus) 207 balticus burzenlandicus (Porcellio) 207 balticus burzenlandicus (Trachelipus) 207 barbertoni (Bethalus) 221 barnardi (Bethalus) 222 barnardi (Cubaris) 222 barroisi (Porcellio) 198 batesoni (Porcellio) 198 benaci (Metroponorthus) 196 Benechinus 213 bergomaticus (Cylisticus plumbeus) 192 besi (Periscyphis) 216 Bethalus 221, 245 bicarinata (Merulana) 228 bicarinata (Spherillo (Merulana)) 228 bicolor (Exzaes) 212 bicoloratus (Armadillo) 229 bicoloratus (Periscyphis) 216 bicoloratus (Spherillo) 229 bicornis (Mahehia) 195 biellensis (Cylisticus) 192 bifrons (Armadillo) 229 bifrons (Spherillo) 229 binotatus (Buddelundia) 223 bipartitus (Buddelundia) 223 R. J. LINCOLN AND J. P. ELLIS bistriatus (Porcellio) 198bituberculatus (Armadillo) 219bituberculatus (Diploexochus) 224bizonatus (Periscyphops) 217blattarius (Porcellio) 198bocki (Nesodillo) 228bodenheimeri (Hemilepistus) 194borellii (Alloniscus) 205borellii (Plataoniscus) 205brachycephalus (Spherillo) 229brentanus (Trachelipus) 207brentanus (Tracheoniscus) 207brevicornis (Armadillo) 219brevipennis (Porcellio) 198brevis (Ignamba) 215brevis (Phalaba) 206Buddelundia 222buddelundia (Porcellio) 198bulgaricus (Trachelipus) 207bulgaricus (Tracheoniscus) 207burzenlandicus (Porcellio balticus) 207burzenlandicus (Trachelipus balticus) 207 caelatum (Armadillidium) 208 Caeroplastes 193 caligans (Spherillo) 229 callosus (Buddelundia) 223 calmani (Porcellio) 198 canaliculata (Merulana) 228 canaliculatus (Spherillo) 228 canariensis (Armadillo) 224 canariensis (Diploexochus) 224 canariensis (Porcellio) 198 carinatus (Armadillo) 222 carinatus (Bethalus) 222 carinatus (Hemiporcellio) 195 carniolense (Armadillidium) 208 carpathicus (Protracheoniscus saxonicus) 207 carthaginensis (Porcellio) 198 cassida (Armadillo) 222 cassida (Bethalus) 222 castor (Diploexochus) 224 caucasius (Cylisticus) 192 caudatus (Ourachaerus) 232 cavernicola (Periscyphis) 216 cayennensis (Porcellio) 199 celsicauda (Diploexochus) 224 chindeensis (Periscyphis) 216 cilicius (Metoponorthus) 196 cinctus (Armadillo) 220 cinctutus (Pyrgoniscus) 232 cinerascens (Buddelundia) 223 cingendum (Armadillidium) 208 cingendum (Armadillidium maculatum) 208 cingulatus (Diploexochus) 224Circoniscus 218civilis (Periscyphis) 216clausus (Synarmadillo) 231cognatus (Porcellio) 199collicolus (Porcellio (Porcellium)) 206collicolus (Porcellium) 206collinus (Armadillo) 220coloratus (Diploexochus) 224concinnus (Scleropactes) 218congener (Armadillo) 224congener (Diploexochus) 224conglobator (Armadillo) 220conisaleus (Diploexochus) 224contractus (Porcellio) 199corcyreum (Armadillidium) 209coxalis (Metoponorthus) 196crassus (Armadillo) 224crassus (Diploexochus) 224creper (Saidjahus) 229cristatus (Hemilepistus) 194cruentatus (Porcellio) 199CYLISTICIDAE 192Cylisticus 192 dalmatinus frascatensis (Orthometopon) 206 danae (Armadillo) 230 danae (Spherillo) 220 davidi (Armadillidium) 209 debueni (Porcellio) 199 decoratus (Spherillo) 229 dentifrons (Cylisticus) 192 depressus (Armadillo) 222 depressus (Bethalus) 222 Desertoniscus 193 desertorum (Protracheoniscus) 206 dilatatus (Porcellio) dimorphus (Porcellionides) 204 Diploexochus 223, 245 disjunctus (Diploexochus) 224 dispar (Porcellio) 199 dispersus (Spherillo) 229 dollfusi (Eubelum) 214 dugesi (Armadillo) 225 dugesi (Diploexochus) 225 dumorum (Armadillo) 232 dumorum (Venezillo) 232 ecaudatus (Diploexochus) 225egens (Armadillo) 220elbanum (Armadillidium) 209elongatus (Hemilepistus) 194Eluma 212, 242emarginatus (Bethalus) 222 INDEX 2460 emunitus (Armadillo) 222emunitus (Bethalus) 222Ennurensis 193episimus (Hekelus) 213erinaceus (Armadillo) 229erinaceus (Spherillo) 229erythroleucus (Armadillo) 220esterelanum (Armadillidium) 209esterelanus (Cylisticus) 192Ethelum 213EUBELIDAE 213, 242Eubelum 214, 242, 243evansi (Porcellio) 199Exzaes 212 feae (Armadillo) 228 feae (Pericephalus) 228 festivus (Armadillo) 225 festivus (Diploexochus) 225 ficorum (Porcellio) 199 ficulneus (Porcellio) 199 fissifrons (Pagana) 206 flavocinctus (Porcellio) 199 flavoscutatum (Armadillidium) 209 flavovittata (Porcellio) 199 flavus (Armadillo) 220 fongosiensis (Anchicubaris) 219 fontium (Protracheoniscus) 206 formosana (Nagara) 205 formosanus (Nagurus) 205 fragilis (Agnara) 205 frascatensis (Orthometopon dalmatinus) 206 fritschei (Reductoniscus) 229 frontalis (Sphaeroniscus) 218 frontalis (Spherillo) 230 frontexcavatum (Armadillidium) 209 frontirostre (Armadillidium) 209 frontosus (Buddelundia) 223 frontosus (Metoponorthus) 196 furcatum (Armadillidium) 209 furcatus (Diploexochus) 225 fuscomarmoratus (Metoponorthus) 196 galapagoensis (Cubaris) 225 galapagoensis (Diploexochus) 225 galeatus (Armadillo) 228 glaeatus (Pericephalus) 228 gallicus (Porcellio) 199 Gelsana 214 gemmulatus (Porcellio) 199 Gerufa 194 Gerutha 214 gestroi (Armadillidium) 209 gigas (Armadillidium simoni) 211 gigas (Cubaris) 225 gigas (Diploexochus) 225 globus (Synarmadillo) 231 Glomerulus 228 glomus (Armadillo) 225 glomus (Diploexochus) 225 gordoniensis (Diploexochus) 225 gracilior (Merulana translucida) 228 gracilipennis (Cylisticus) 192 graecorum (Armadillidium) 209 grandinatum (Armadillidium) 209 graniger (Porcellio) 199 granulatum (Armadillidium) 210 granulatum peloponnesiaca (Armadillidium) 209 granuliferus (Porcellio) 200gravei (Metoponorthus) 196grenadensis (Armadillo) 232grenadensis (Venezillo) 232griseoalbus (Armadillo) 222griseoalbus (Bethalus) 222grisescens (Spherillo) 230grossus (Armadillo) 230grossus (Spherillo) 230 Haloporcellio 194 Haplarmadillo 215 harsadiensis (Armadillo) 220 hebridarum (Spherillo) 230 Hekelus 213 Hemilepistus 194, 237 Hemiporcellio 195 hermagorensis (Protracheoniscus) 206 herzegovinensis (Porcellio) 206 herzegovinensis (Porcellium) 206 Hiallum 215 hilgendorfi (Eubelum) 215 hilgendorfi (Hiallum) 215 hirsutulus (Protracheoniscus) 206 hirticornis (Gerufa) 194 hirtum (Armadillidium) 209 hirtus (Agabiformius) 193 hirtus (Lucasius) 193 hispida (Metoponorthus) 196 hispida (Porcellio (Porcellionides)) 196 hispidus (Ennurensis) 193 hughscotti (Periscyphis) 216 hybridum (Armadillidium) 213 hybridum (Schizidium) 213 hypotoreus (Spherillo) 230 hypselos (Diploexochus) 225 hypsinephes (Diploexochus) 225 Ignamba 215 R. J. LINCOLN AND J. P. ELLIS ignavum (Eubelum) 214 imbutus (Porcellio) 200 imbutus pellegrinensis (Porcellio) 200 immotus (Armadillo) 220 immsi (Hemiporcellio) 195 immsi (Porcellio) 195 inaequalis (Buddelundia) 223 incanus (Porcellio) 200 Inchanga 195 incicus (Scleropactes) 218 incisa (Nagara) 205 incisus (Nagurus) 205 inconspicuus (Armadillo) 220 ingens (Spherillo) 230 iniqua (Merulana) 228 iniquus (Spherillo) 228 instrenuum (Eubelum) 214 integer (Armadillo) 220 intercalarius (Porcellio) 200 intermixtus (Armadillo) 220 interpolator (Porcellio) 200 japonicus (Porcellio scaber) 202jelskii (Porcellio) 200 kaokoensis (Diploexochus) 225Kisuma 232 kogmani (Diploexochus) 225kunenensis (Periscyphis) 216kunenensis (Periscyphops) 216 labiatus (Buddelundia) 223lacteolus (Metoponorthus) 196laevigatus (Buddelundia) 223laevis (Gerutha) 214laevis (Porcellio) 201lamellatus (Porcellio) 194, 201laminigerum (Armadillidium) 210larii (Trachelipus) 207larii (Tracheoniscus) 207larii aternanus (Trachelipus) 207larii aternanus (Tracheoniscus) 207lateralis (Buddelundia) 223laticauda (Mahehia) 195laticauda (Tura) 204latifrons (Armadillo) 222lagifrons (Bethalus) 222latissimus (Periscyphis) 216latissimus (Porcellio) 200latus (Protracheoniscus) 206lentus (Spherillo) 230lepineyi (Porcellio) 200Leptotrichus 195, 237 leucocephalus (Microcercus) 215leucocephalus (Periscyphis) 215lifuensis (Armadillo) 220lifuensis (Cubaris) 220limbata (Periscyphis) 216limbatus (Bethalus) 222limenetes (Diploexochus) 225littoralis (Metoponorthus) 196longicauda (Porcellio) 200longipennis (Porcellio) 200longipes (Armadillo) 225longipes (Diploexochus) 225lubricum (Eubelum) 214Lucasius 195, 237lugubris orarum (Porcellio) 201lugubris vizzavonensis (Porcellio) 201lusitanus (Porcellio) 202lusitanus (Porcellio scaber) 202lutschniki (Trachelipus) 207lutschniki (Tracheoniscus) 207 macmahoni (Armadillo) 220 macrodens (Bethalus) 222 macrops (Gerufa) 194 maculata (Balloniscus) 205 maculata (Mahehia) 195 maculata (Philoscia) 205 maculatum cingendum (Armadillidium) 208 maculipes (Porcellio) 200 maculosa (Pagana) 206 maculosus (Spherillo) 230 madagascariensis (Agnara) 205 madagascariensis (Metoponorthus) 205 magnincus (Porcellio) 200 Mahehia 195 makuae (Diploexochus) 226 mammillata (Sunniva) 213 marcidus (Armadillo) 228 marcidus (Pericephalus) 228 mereoticum (Armadillidium) 210 marginalis (Uramba) 205 marginatus (Spherillo) 230 marginenotatus (Porcellio) 200 marginepilosa (Gerutha) 215 marinensium (Armadillidium) 210 marmorata (Gerufa) 194 marmoratus (Synarmadillo) 231 marquesarum (Spherillo) 230 marquesarum (Spherillo (Xestodillo?)) 230 mayeti (Armadillo) 220 mehelyi (Protracheoniscus) 206 mehelyi (Protracheoniscus politus) 206 meiringi (Diploexochus) 226 melanurus (Acaeroplastes) 193 INDEX 2460 melanurus (Metoponorthus) 193 meleagris (Metoponorthus) 197 meleagris (Metoponorthus pruinosus) 197 Merulana 228 Mesarmadillo 215, 243 Metoponorthus 196, 237, 238 Mica 197 Microcercus 215, 243 microps (Armadillo) 228 microps (Glomerulus) 228 microps (Ignamba) 215 mildei (Porcellio) 201 minor (Sunniva) 213 minuta (Angara) 193 minuta (Pareluma) 213 minutus (Agabiformius) 193 miser (Armadillo) 220 modestum (Ethelum) 214 modestus (Mesarmadillo) 214 moebiusi (Porcellio) 201 mohamedanicum (Armadillidium) 210 monocellatus (Haplarmadillo) 215 montagui (Diploexochus) 226 montana (Gerufa) 194 montanus (Armadillo) 220 montanus (Porcellio) 203 montanus (Porcellio spinipennis) 203 monticola (Buddelundia) 223 montivagus (Armadillo) 230 montivagus (Spherillo) 230 mucidus (Bethalus) 222 muricatum (Armadillidium) 210 mus (Lyprobius) 205 mus (Uramba) 205 myrmecophilus (Metoponorthus (Myrmecon- iscus) myrmecophilus) 196myrmicidarum (Metoponorthus) 196mystica (Sunniva) 213 Nagurus 205, 239 nana (Nagara) 205 nanus (Diploexochus) 226 nanus (Nagurus) 205 narentanus (Porcellio) 201 nasatum (Armadillidium) 212 nasatum sorrentinum (Armadillidium) 210 nasatus (Cylisticus) 192 natalensis (Inchanga) 195 naupliensis (Porcellio) 201 nebulosus (Diploexochus) 226 Nesodillo 228 nicklesi (Porcellio) 201 nicobaricus (Armadillo) 230 nicobaricus (Spherillo) 230 nigra (Porcellio) 201nigricans (Periscyphis) 217nigripes (Buddelundia) 223nigrobrunneus (Metoponorthus) 196nigromarginatus (Armadillo) 221nigropunctatus (Periscyphis) 231nigropunctatus (Synarmadillo) 231nitidulus (Armadillidium) 210nobilis (Spherillo) 230nodosus (Hemilepistus) 194normani (Lucasius) 201normani (Porcellio) 201 obliquidens (Diploexochus) 226obliquipes (Spherillo) 230obsoletus (Porcellio) 201obtusa (Angara) 193obtusifrons (Porcellio) 201obtusiserra (Porcellio) 201obtusus (Agabiformius) 193occidentalis (Protracheoniscus) 196ocellatus (Porcellio) 201odherni (Armadillidium) 210odherni (Armadillidium peraccai) 210oertzeni (Armadillidium) 213oertzeni (Schizidium) 213officinalis syriaca (Armadillo) 221oliveti (Armadillidium) 210olivieri (Porcellio) 198opacus (Buddelundia) 223opihensis (Actoecia) 218orarum (Porcellio lugubris) 201orarum (Porcellio orarum) 201orarum vizzavonensis (Porcellio) 201ormeanum (Armadillidium) 210Orodillo 228 orphanus (Diploexochus) 226Orthometopon 206Ourachaerus 232ovampoensis (Cubaris) 226ovampoensis (Diploexochus) 226oxyzomus (Armadillo) 221oxyzomus (Cubaris) 221 pachytos (Diploexochus) 226Pagana 206 palaestinus (Hemilepistus) 194pallasi (Armadillidium) 210pallidus (Cylisticus) 192pallidus (Lucasius) 195pallidus (Porcellio) 195panurus (Armadillo) 222panurus (Bethalus) 222papillosa (Kisuma) 232 R. J. LINCOLN AND J. P. ELLIS Paraperiscyphis 216 Pareluma 213 parietinus (Porcellio) 201 parvus (Armadillo) 230 parvus (Porcellio) 201 parvus (Spherillo) 230 pauper (Porcellio) 201 pauperculus (Diploexochus) 226 peleponnesiaca (Armadillidium granulatum) 209 pellegrinense (Armadillidium) 210pellegrinensis (Porcellio imbutus) 200peltatus (Spherillo) 230penicilliger (Haloporcellio) 194peraccae (Armadillidium) 210peraccai odherni (Armadillidium) 210Pericephalus 228Periscyphis 216, 243Periscyphops 217, 243, 244perkinsi (Armadillo) 231perkinsi (Spherillo) 231petraeum (Armadillidium) 210phaeacorum (Porcellio rathkei) 207phaeacorum (Trachelipus) 207Phalaba 206, 239, 240phaleronensis (Orthometopon) 206phaleronensis (Porcellio (Metoponorthus)) 206 philoscoides (Metoponorthus) 196pictus (Rotungus) 218piger (Armadillo) 221pila (Gerutha) 215pilosa (Gerutha) 215pilularis (Armadillo) 221pilum (Eubelum) 215pisum (Armadillo) 226pisum (Diploexochus) 226planarius (Porcellio) 202Plataoniscus 205plumbeus (Cylisticus) 192plumbeus bergomatius (Cylisticus) 192plumbeus umbricus (Cylisticus) 192politus (Leptotrichus) 195politus mehelyi (Protracheoniscus) 206pollex (Diploexochus) 226Polyacanthus 229polythele (Diploexochus) 226pontremolensis (Cylisticus) 192Porcellio 197, 238, 239PORCELLIONIDAE 193, 237Porcellionides 204Porcellium 206 porphyrivagus (Caeroplastes) 193porphyrivagus (Metoponorthus) 193 portofinense (Armadillidium) 210 praeustus (Porcellio) 202 pretoriensis (Armadillo) 222 pretoriensis (Bethalus) 222 Protracheoniscus 206 provincialis (Porcellio) 202 proximatus (Armadillo) 221 pruinosus (Metoponorthus) 199, 200, 204 pruinosus anconanus (Metoponorthus) 197 pruinosus meleagris (Metoponorthus) 197 pseudoratzburgi (Porcellio (Euporcellio)) 207 pseudoratzburgi (Trachelipus) 207 pujetanum (Armadillidium) 211 pujetanus (Porcellio) 202 pulcher (Periscyphis) 217 pumilus (Armadillo) 232 pumilus (Venezillo) 232 purpurascens (Eluma) 212 purpurascens (Spherillo) 231 purpureus (Porcellio) 202 pygmaeus (Armadillo) 221 pygmaeus (Periscyphis) 231 pygmaeus (Synarmadillo) 231 pyrenaeus (Porcellio) 202 Pyrgoniscus 232 quadrimaculatus (Diploexochus) 226quadrimaculatus (Mesarmadillo) 215quadrimaculatus (Periscyphis) 217quadriserriatum (Armadillidium) 211quadritracheata (Buddelundia) 223quietum (Eubelum) 214quinquepustulatum (Armadillidium) 211 ragusae (Porcellio) 202rathkei phaecorum (Porcellio) 207reaumuri (Hemilepistus) 194Reductoniscus 229reflexum (Ethelum) 214reflexus (Mesarmadillo) 214rehobotense (Armadillidium) 211rhodesiae (Bethalus) 222rhodesiensis (Diploexochus) 227ribauti (Sphaerobathytropa) 218riparium (Armadillidium) 211rosai (Armadillidium) 211Rotungus 218 rufescens (Diploexochus) 227ruficauda (Periscyphis) 217rufobrunneus (Agabiformius) 193rufomarginatus (Spherillo) 231rugifrons (Buddelundia) 223rugulosus (Cubaris) 231rugulosus (Spherillo) 231 INDEX 246g sabuleti (Metoponorthus) 197 sabulifer (Porcellio) 202 Saidjahus 229 saldanhae (Diploexochus) 227 salisburyensis (Diploexochus) 227 saltuum (Porcellio) 202 sanctum (Armadillidium) 211 sarajevensis (Porcellio) 202 sarasini (Nesodillo) 228 sarculatus (Porcellio) 202 sauteri (Orodillo) 228 savonense (Armadillidium) 211 saxonicus (Protracheoniscus) 207 saxonicus carpathicus (Protracheoniscus) 207 scaber (Porcellio) 199, 201 scaber japonicus (Porcellio) 202 scaber lusitanus (Porcellio) 202 scaberrimum (Armadillidium) 211 scabrum (Armadillidium) 211 schellenbergi (Nesodillo) 228 schirasi (Hemilepistus) 194 Schizidium 213 Scleropactes 218, 244 serratum (Armadillidium) 211 setaceus (Spherillo) 231 sexfasciatus (Metoponorthus) 197 sexfasciatus asifensis (Metopornorthus) 197 sharpi (Armadillo) 221 siculorum (Armadillidium badium) 208 silvanus (Periscyphops) 217 silvarum (Armadillo) 227 silvarum (Diploexochus) 227 simoni (Armadillidium) 211 simoni gigas (Armadillidium) 211 simplex (Synarmadillo) 232 simulator (Porcellio) 202 sociabilis (Porcellio) 203 sollers (Spherillo) 231 sordidus (Porcellio) 203 sorrentinum (Armadillidium nasatum) 210 spatulata (Porcellio) 203 spatulatus (Porcellio) 200 speyeri (Armadillidium) 212 Sphaerobathytropa 218 SPHAERONISCIDAE 218, 244 Sphaeroniscus 218, 244 Spherillo 229, 245 sphynx (Haloporcellio) 194 spicatus (Spherillo) 231 spinicornis (Porcellio) 203 spiniger (Acanthoniseus) 219 spinipennis (Porcellio) 203 spinipennis (Porcellio spinipennis) 202, 203 spinipennis montanus (Porcellio) 203 spinipennis spinipennis (Porcellio) 203spinipes (Porcellio) 203spinosa (Akermania) 219spinosus (Circoniscus) 218spinosus (Paracubaris) 218spretus (Porcellio) 203squamatus (Periscyphops) 217squamosus (Periscyphops) 217stebbingi (Paraperiscyphis) 216steenbrasi (Diploexochus) 227stipulatum (Eubelum) 214stolikanum (Armadillidium) 212stricticauda (Armadillo) 222stricticauda (Bethalus) 222subdentatum (Armadillidium) 212suberorum (Cylisticus) 192subterraneus (Desertoniscus) 193subtransversus (Periscyphis) 217succinctus (Porcellio) 203sulcatus (Buddelundia) 223Sunniva 213sylvatica (Exzaes) 212Synarmadillo 231syriaca (Armadillo omcinalis) 221 tabularis (Diploexochus) 227tamei (Periscyphis) 217tarangensis (Spherillo) 231tardus (Mica) 197tardus (Porcellio) 197taschkentensis (Protracheoniscus) 207tendanum (Armadillidium) 212tenuipunctatus (Armadillo) 221testacea (Tura) 204tigris (Armadillidium) 212tirolense (Armadillidium) 212tomentosus (Buddelundia) 223Toradjia 242trachealis (Porcellio) 203TRACHELIPIDAE 205, 239Trachelipus 207tradouwi (Bethalus) 222translucida (Merulana) 228translucida gracilior (Merulana) 228translucidus (Armadillo) 228transmutatus (Porcellio) 203transsilvaticus (Cylisticus) 192transvaalensis (Polyacanthus) 229travancoria (Nagara) 205travancorius (Nagurus) 205triangulifera (Uramba) 205triarticulatus (Periscyphis (Periscyphops)) 218triarticulatus (Periscyphops) 218 R. J. LINCOLN AND J. P. ELLIS trifolium (Armadillo) 221trilobatus (Porcellio) 203trivialis (Periscyphis) 217truncorum (Armadillo) 232truncorum (Venezillo) 232tuberculatus (Mesarmadillo) 215tugelae (Diploexochus) 227tunetanum (Armadillidium) 212Tura 204 umbricus (Cylisticus plumbeus) 192undulata (Periscyphis) 217uniformis (Sunniva) 213Uramba 205, 239 vallombrosae (Armadillidium) 212variabilis (Porcellionides) 204venetus (Protracheoniscus) 207Venezillo 232venustus (Armadillo) 232venustus (Venezillo) 232verrucosus (Armadillo) 227verrucosus (Diploexochus) 227versicolor (Armadillidium) 212 vespertilio (Porcellio) 203violaceus (Porcellio) 198, 204virgatus (Metoponorthus) 197viridis (Metoponorthus) 197viticola (Armadillo) 221vittatus (Periscyphis) 217vizzavonensis (Porcellio lugubris) 201vizzavonensis (Porcellio orarum) 201vulgare (Armadillidium) 210, 212, 221 weberi (Armadillo) 231weberi (Spherillo) 231wilsmorei (Armadillo) 221wilsmorei (Cubaris) 221 yemenensis (Porcellio) 204 zachvatkini (Hemilepistus) 194zealandicus (Porcellio) 204zebricolor (Armadillo) 221zebricolor (Cubaris)zigzag (Armadillo) 227zigzag (Diploexochus) 227zwartbergensis (Diploexochus) 227 Dr R. J. LINCOLN JOAN P. ELLIS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.1965. (Out of Print.) £3-75-* 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogyof the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates,77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £970. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : theBiology and Evolution of a Species Flock. Pp. 134 ; I Plate, 77 Text-figures. 1974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU 1 GENERAL *' 27 JAM975 CAMPYLASPIS SPECIES (CRUSTACCUMACEA) FROM THE DEEPATLANTIC N. S. JONES BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 6 LONDON: 1974 GENERAL '' CAMPYLASPIS SPECIES (CRUSTACEA:FROM THE DEEP ATLANTIC BY N. S. JONES ' Department of Marine BiologyUniversity of LiverpoolPort Erin, Isle of Man Pp. 247-300 ; 26 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 6 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 6 of the Zoological series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 23 December, 1974 Price £3.25 CAMPYLASPIS SPECIES (CRUSTACEA : CUMACEA)FROM THE DEEP ATLANTIC By N. S. JONES CONTENTS INTRODUCTION .......... 249 THE GENUS Campylaspis ....... . 249 KEY TO THE SPECIES OF Campylaspis . . . . . . 251 DESCRIPTIONS AND RECORDS ........ 256 ACKNOWLEDGEMENTS ......... 299 REFERENCES ........... 299 SYNOPSIS Thirty-nine species of Campylaspis were collected during a number of cruises, mostly by shipsof the Woods Hole Oceanographic Institution, in depths exceeding 200 m in the Atlantic between1960 and 1969. Twenty-five of these are described as new. A key is included to the 98 speciesknown at the time of writing. INTRODUCTION THIS paper deals with the genus Campylaspis (family Nannastacidae) from a numberof collections made in the deep water of the Atlantic Ocean. Almost all the stationssampled were in depths exceeding 200 m and the great majority of samples wereobtained by epibenthic sled dredges (Sanders & Hessler, 1969). Altogether 39 speciesof Campylaspis were collected, of which 25 are described here as new. A preliminary account of the distribution of the Cumacea from several of thesecollections has been published (Jones & Sanders, 1972) and also descriptions of anumber of new genera and species (Jones, 1973 ; Reyss, 1974). Reyss (1973) hasalso discussed the distribution of Cumacea in the deep waters of the Mediterranean. The material studied was obtained during the following cruises : Gay Head-Bermuda transect (Woods Hole Oceanographic Institution ships),1960-69. 'Chain' cruise (WHOI) off Brazil, April 1973. Dakar- Recife transect ('Atlantis II', WHOI), February 1967. 'Sarsia' cruise (Marine Biological Association) in Bay of Biscay, July 1967. 'Discovery' cruise (National Institute of Oceanography) near the Canary Islands,March 1968. 'Atlantis II' cruise (WHOI) Walvis Bay - Luanda, May 1968. 'Jean Charcot' cruise (Centre Oceanologique de Bretagne) in North Atlantic,August-October 1969. THE GENUS CAMPYLASPIS Including those described in this paper there are now 98 species in the genusCampylaspis known to the author. In spite of this rather unwieldy number there 25o N. S. JONES seem to be no grounds at present for splitting the genus, but it may be divided intofour main groups according to the sculpture or lack of it on the carapace, with twosmaller groups of which one is rather miscellaneous. These are : 1. The rubicunda-group with the following 22 species : alba, nitens, nuda, orientalis, inornata, guttata, pacifica, pulchella, paeneglabra,mauritanica, rubicunda, laevigata, angularis, glabra, bonetti, amblyoda, rufa,kiiensis, thompsoni, similis, roscida, laticarpa. In this group the carapace is smooth, without lateral depressions, and with atmost a pair of low rounded protuberances or with small granulations. 2. The sw/crt/a-group with 20 species : striata, aulacoeis, fusiformis, porcata, canaliculata, granulata, tubulata, pumila,jonesi, latidactyla, sulcata, minor, legendrei, aperta, sticta, echinata, pilosa,redacta, cognata, unisulcata. In these there is a depression on either side of the carapace. If distinct ridgesare present they do not extend to the dorsal hind end of the carapace. Somespines or a few low protuberances may be present but not conical, subcylindricalor rounded tubercles. 3. The costata-group with 24 species : platyuropus, uniplicata, vitrea, pileus, macrophthalma, hartae, sinuosa, undata,johnstoni, ovalis, arcuata, crispa, speciosa, costata, triplicata, umbensis, reticulata,paucispina, caperata, plicata, valleculata, exarata, mansa, bicarinata. These have one or more, usually two or three distinct ridges running hori-zontally or obliquely backwards on either side of the carapace, of which at leastone extends onto the dorsum. Depressions may be present between the ridgesbut they are not defined posteriorly. Tubercles are not present. 4. The verrucosa-group with 24 species : affinis, serratipes, clavata, thetidis, maculata, sagamiensis, intermedia, horrida,submersa, rubromaculata, papillata, paucinodosa, horridoides, glebulosa, frigida,pustulosa, squamifera, torulosa, verrucosa, aspera, nodulosa, globosa, antarctica,multinodosa. These have moderate or large numbers of tubercles on the sides and dorsumof the carapace. 5. The rostrata-group with four species :rostrata, rostellata, aculeata, brevicornis. This is a small group in which the pseudorostrum is prominent and enlarged. 6. The spinosa-group with four species :spinosa, rupta, quadriplicata, bulbosa. These four species have little in common but do not fit into any othergroup. The key which follows is based on that of Hale (1945) with some alterations, but isgreatly enlarged to include more than twice the original number of species. Theshape and markings of the carapace, including the eyelobe and the pseudorostrum,are the most important characters for identification, followed by the relative propor-tions and armature of the second and third maxillipeds, second pereopods and CAMPYLASPIS FROM THE DEEP ATLANTIC 251 uropods. Most characters can be seen under a stereo-microscope without dissectionbut when it is necessary to examine the second maxillipeds it is usually necessary toremove one and to examine it under a magnification of at least x 50. Unfortunatelynot all the important characters have been described or figured for a number ofspecies and identification is not always easy, especially in the rubicunda-group,where carapace markings are of little help, and sometimes where the key dependson relative lengths of appendages or their segments. In case of doubt a more com-plete dissection of appendages may be needed and the original descriptions con-sulted. Undoubtedly many species in the genus remain as yet undescribed. KEY TO THE SPECIES OF CAMPYLASPIS 1 Carapace smooth, without tubercles, spines, ridges or lateral grooves ... 2Carapace with tubercles, spines or ridges, or at least a shallow groove on either side 21 2 Eyelobe small and rudimentary ......... 3 Eyelobe of normal size ........... 5 3 Inner edge of merus of maxilliped 3 not serrated . . . alba Hansen, 1920Inner edge of merus of maxilliped 3 serrated ....... 4 4 Dactyl of pereopod 2 broad and little tapered .... nitens Bonnier, 1896Dactyl of pereopod 2 narrow and tapering ...... nuda sp. nov. 5 Dorsal outline of carapace a little uneven ..... paciftca Sars, 1887Dorsal outline of carapace smoothly rounded ....... 6 6 Eye without lenses ............ 7 Eye with lenses visible ........... 9 7 Maxilliped 3 with ischium, merus, carpus and propodus strongly serrated paeneglabra Stebbing, 1912Maxilliped 3 without conspicuous serrations ....... 8 8 Basis of maxilliped 3 much longer than remaining segments together orientalis Caiman, 1911Basis of maxilliped 3 shorter than the remaining segments together inornata Jones, 1969 9 Uropods at least nearly as long as the last three pleon somites together . . 10Uropods much shorter than the last three pleonites together . . guttata sp. nov. 10 Exopod of uropod a little longer than endopod .... pulchella Sars, 1871Exopod of uropod not longer than endopod . . . . . . . 11 11 Dactyl of pereopod 2 at least as long as carpus and propodus together . . . 12Dactyl of pereopod 2 shorter than carpus and propodus together . . . . 17 12 Dactyl of pereopod 2 longer than the carpus and propodus together . .13Dactyl of pereopod only as long as the carpus and propodus together . .15 13 Distal seta of ischiobasis of maxilliped 2 swollen distally mauritanica Bacescu & Muradian, 1972Distal seta of ischiobasis of maxilliped 2 normally tapered . . . . . 14 14 Peduncle of the uropod twice as long as the endopod . rubicunda (Lilljeborg, 1855)Peduncle of uropod less than twice as long as the endopod . . laevigata sp. nov. 15 Merus of maxilliped 3 narrow, its carpus and propodus not serrated angularis Gamo, 1960Merus of maxilliped 3 broad, its carpus and propodus serrated . . . . 16 16 Basis of pereopod 2 nearly half the total length of the appendage bonetti Bacescu & Muradian, 1972Basis of pereopod 2 little more than a third of the total length . amblyoda Gamo, 1960 17 Dactyl of maxilliped 2 with two or three spines . . . . . . . 18 Dactyl of maxilliped 2 with four spines . . . . . . • • 19 252 N. S. JONES 18 Dactyl of pereopod 2 little tapered. Terminal seta of ischiobasis of maxilliped 2 slender .......... glabra Sars, 1879 Dactyl of pereopod 2 normally tapered. Terminal seta of ischiobasis of maxilliped 2 thickened ........... rufa Hart, 1930 19 Antennal notch deep ........ kiiensis Gamo, 1960 Antennal notch shallow ........... 20 20 Carpus of pereopod i barely longer than propodus . . . thompsoni Hale, 1945Carpus of pereopod i much longer than propodus . . . si mil is Hale, 1945 21 Carapace with lateral furrows or ridges but without spines, tubercles or other large prominences ............ 22 Carapace may or may not have furrows but also at least a pair of tubercles or many spines or large prominences, which may or may not form lateral ridges . . 54 22 Either side of the carapace with a faint groove which is not margined above or below by a distinct ridge or fold . . . ... ... 23 Either side of the carapace with at least one well-formed ridge or fold . . . 31 23 Carapace with three dorso-lateral striations on either side of the mid-line striata Gamo, 1960Carapace without dorso-lateral striations ........ 24 24 Dactyl of pereopod 2 blunt ended and not tapering in distal half . . . . 25Dactyl of pereopod 2 tapering to a narrower distal end ..... 29 25 Carapace with small granules above and below the faint lateral grooves granulata Gamo, 1960 Carapace without granules .......... 26 26 Dactyl of pereopod 2 with robust flattened terminal setae . . porcata sp. nov.Dactyl of pereopod 2 with short terminal setae ....... 27 27 Groove at side of carapace scarcely reaches into posterior half canaliculata Zimmer, 1936Groove at side of carapace extends well into posterior half . . . . . 28 28 Basis of maxilliped 3 much less than half as long as rest of appendage aulacoeis Le Loeuff & Intes, 1972Basis of maxilliped 3 much more than half as long as rest of appendage fusiformis Gamo, 1960 29 Dactyl of pereopod 2 sharply tapered, with short terminal setae . unisulcata Hale, 1945Dactyl of pereopod 2 moderately tapered, with a long end spine .... 30 30 Groove at side of carapace scarcely reaching into posterior half . tubulata Page, 1945Groove at side of carapace broadened posteriorly in female, reaching well into posterior half in male ........ putnila Gamo, 1960 31 Carapace with four irregular, subrectangular, depressed areas on either side, bordered by prominent folds ....... rupta Hale, 1945 Carapace not so sculptured, with carinae on either side subparallel ... 32 32 Carapace with one oblique ridge on either side ....... 33 Carapace with more than one ridge on either side ...... 34 33 Peduncle of uropod very broad and only a little longer than the endopod platyuropus Caiman, 1911Peduncle of uropod slender, three times as long as endopod . . uniplicata Hale, 1945 34 Two oblique carinae arising anteriorly and extending for greater part of length of carapace on either side ........... 35 Three or more oblique carinae extending similarly on carapace .... 49 35 Pseudorostrum long, at least a sixth of the total carapace length .... 36Pseudorostrum shorter, less than a sixth of the total carapace length 37 36 Pseudorostrum turned up at an angle to the line between its hind end and the hind end of the carapace ........ vitrea Caiman, 1906 Pseudorostrum not turned up . . . . . . pileus Foxon, 1932 37 Eyelobe of normal size, with lenses .... . . . . 38 Eyelobe small or absent, without lenses . . . . . . . . 46 CAMPYLASPIS FROM THE DEEP ATLANTIC 253 38 Eyelobe linguiform, narrow and dilated distally . . . macrophthalma Sars, 1879Eyelobe not linguiform, broad at base ........ 39 39 Pseudorostrum turned up at a distinct angle to dorsum of carapace . hart tie Lie, 1969Pseudorostrum little or not turned up ........ 40 40 Peduncle of the uropods not more than one-and-a-half as long as the endopod . 41Peduncle of the uropods more than one-and-a-half as long as the endopod . . 42 41 Uropods shorter than the last three pleonites together . . sinuosa Gamo, 1960Uropods as long as the last three pleonites together . jonesi Bacescu & Muradian, 1972 42 Dactyl of pereopod 2 shorter than the carpus .... uiidata Sars, 1865Dactyl of pereopod 2 longer than the carpus ....... 43 43 Dactyl of pereopod 2 not tapered and ending in a short articulated process latidactyla Hale, 1945Dactyl of pereopod 2 tapered, without an articulated process .... 44 44 Dactyl of pereopod 2 with no terminal setae .... sulcata Sars, 1870Dactyl of pereopod 2 ending in setae. ........ 45 45 Sulcus on side of carapace undivided. ..... minor Hale, 1945 Sulcus divided by a short vertical ridge ..... legendrei Page, 1951 46 Dactyl of pereopod 2 as long as the carpus and propodus together . . . 47Dactyl of pereopod 2 only as long as the carpus . . . aperta Lomakina, 1958 47 Merus and propodus of maxilliped 3 unserrated .... johnstoni Hale, 1937Merus and propodus of maxilliped 3 serrated ....... 48 48 Ridges at side of carapace well separated and extending to hind end ovalis Stebbing, 1912Ridges at side of carapace closer together and not reaching hind end . stictu sp. nov. 49 Carapace much elevated dorsally. Dactyl of pereopod 2 not tapered, without terminal spines or setae ....... arcuata sp. nov. Carapace not much elevated. Dactyl of pereopod 2 tapered, with terminal spines or setae . . . . . . . . . . . .50 50 Uropod distinctly longer than the last two pleonites together, its endopod not very broad ............. 51 Uropod not longer than the last two pleonites together, its endopod broad crispa Lomakina, 1955 51 Lower lateral ridge on carapace branched . . . . . . .52 Lower lateral ridge on carapace unbranched ....... 53 52 Upper horizontal ridge on carapace joined to ridge below by a short vertical ridge near the front ........ speciosa Lomakina, 1955 These two ridges not joined in front by a vertical ridge . . costata Sars, 1865 53 Dactyl of pereopod 2 as long as the carpus and propodus together triplicata Hale, 1945Dactyl of pereopod 2 much longer than the carpus and propodus together utnbensis Gurwitsch, 1939 54 Carapace with many spines .......... 55 Carapace with few or no spines.......... 59 55 Carapace with a depression on either side ........ 56 Carapace without a depression . . . . . . . . . -57 56 Lateral depression deep. Carapace with long hairs dorsally and large reticulations behind the groove ........ reticulata Gamo, 1960 Lateral depression shallow. Carapace without hairs or reticulations echinata Hale, 1945 57 Pseudorostrum long and prominent. Only small spines present on carapace . . 58Pseudorostrum short and strongly upturned. Some stout spines present spinosa Caiman, 1906 58 Spines confined to dorsum of carapace. Dactyl of pereopod 2 shorter than carpus rostellata sp. nov.Spines covering whole of carapace. Dactyl of pereopod 2 longer than carpus aculeata sp. nov. 254 N. S. JONES 59 Carapace with a few low tubercles or with inconspicuous granule-like tubercles . 60Carapace with many conspicuous tubercles ........ 76 60 Carapace with very small granule-like tubercles ....... 61 Carapace with a few low dorsal protuberances ....... 62 61 Ocular lobe narrow, about twice as long as wide. Dactyl of maxilliped 2 with three spines. . . . . • » . . . . laticarpa Hansen, 1920 Ocular lobe wider than long. Dactyl of maxilliped 2 with four spines roscida Hale, 1945 62 Carapace without distinct lateral ridges ........ 63 Carapace with two or more lateral ridges on either side ..... 66 63 Carapace with scattered hairs and a single pair of protuberances dorso-laterally pilosa sp. nov.Carapace without hairs and with several dorsal protuberances .... 64 64 Antenna 2 of male reaching end of uropods ....... 65 Antenna 2 of male not reaching to end of pleon .... brevicornis sp. nov. 65 Dactyl of maxilliped 2 with three spines. Peduncle of uropod smooth affinis Sars, 1870Dactyl of maxilliped 2 with four spines. Peduncle of uropod serrated serratipes Hansen, 1920 66 Carapace with upper lateral ridge recurved above to form a nearly closed loop quadriplicata Lomakina, 1968Lateral ridges on carapace not recurved ........ 67 67 Carapace broad, with two vertical ridges on either side nearly meeting their fellows from the opposite side mid-dorsally ..... bulbosa sp. nov. Carapace without vertical ridges ......... 68 68 Carapace with oblique ridges on either side defining a depressed area which is much broader behind ......... redacta sp. nov. Sulcus if present not much broader behind ....... 69 69 Dorso-lateral protuberances tipped by a short spine and upper oblique ridges with a few small spines ............ 70 Dorso-lateral protuberances and ridges without spines. . . . . . 71 70 Basal segment of maxilliped 2 with peculiarly thickened terminal seta paucispina sp. nov.Terminal seta on basal segment of maxilliped 2 normally tapered. . cognata sp. nov. 71 Dactyl of pereopod 2 much shorter than the carpus . . . . . . 72 Dactyl of pereopod 2 much longer than the carpus ...... 73 72 Basal segment of maxilliped 2 with thickened terminal seta . caperata sp. nov.Basal segment of maxilliped 2 with terminal seta normally tapered . plicata sp. nov. 73 Carapace with three horizontal ridges on either side, the upper ridge meeting its fellow dorsally towards the hind end ..... valleculata sp. nov.Carapace with two ridges on either side, the upper ridge not meeting its fellow on the mid-dorsal line ............ 74 74 Pseudorostrum long and distinctly upturned .... exarata sp. nov.Pseudorostrum short and not upturned . . . . . • - • 75 75 Dactyl of pereopod 2 as long as the carpus and propodus together and with three short stout terminal spines ........ tnansa sp. nov. Dactyl of pereopod 2 longer than the carpus and propodus together and with a single short stout terminal spine and a longer seta . . . bicarinata sp. nov. 76 Carapace with tuberculate ridges or with some of the tubercles situated in rows along the sides ............. 77 Carapace without tuberculate ridges, nor with tubercles situated in rows along the sides .............. 91 77 Dactyl of pereopod 2 much longer than the carpus and propodus together, and with a terminal lobe extending beyond the insertion of the most distal seta . . 78Dactyl of pereopod 2 at most as long as the carpus and propodus together, and with distal setae quite terminal .......... 79 CAMPYLASPIS FROM THE DEEP ATLANTIC 255 78 Pseudorostrum relatively long and prominent. Tubercles on carapace few and large rostrata Caiman, 1905Pseudorostrum shorter and not prominent. Tubercles on carapace small and numerous .......... thetidis Hale, 1945 79 Merus of maxilliped 3 at least one-and-a-half as long as the carpus ... 80Merus of maxilliped 3 very little longer than the carpus . maculata Zimmer, 1907 80 Dactyl of pereopod 2 not longer than the carpus . . . . . . 81 Dactyl of pereopod 2 distinctly longer than the carpus ..... 86 81 Tuberculate ridges at sides and lower edges of carapace joined near front by a small vertical ridge, enclosing a smooth quadrilateral area . clavata Lomakina, 1952 No smooth quadrilateral area at sides of carapace ...... 82 82 Merus of maxilliped 3 very long, three times as long as the carpus sagamiensis Ga.mo,ig6jMerus of maxilliped 3 shorter, less than twice as long as the carpus ... 83 83 Sides of carapace with three folds, the two uppermost bearing large rounded tubercles intermedia Hansen, 1920 Side of carapace with two ridges, formed of conical tubercles .... 84 84 Uropod as long as last three pleonites together .... horrida Sars, 1870Uropod only as long as last two pleonites together ...... 85 85 Carapace with many tubercles. Merus and carpus of maxilliped 3 dentate papillata Lomakina, 1952Carapace with comparatively few tubercles. Merus and carpus of maxilliped 3 not dentate .......... paucinodosa sp. nov. 86 Peduncle of uropod less than one-and-a-half as long as the endopod ... 87Peduncle of uropod more than one-and-a-half as long as the endopod ... 89 87 Merus and carpus of maxilliped 3 without teeth . . . submersa sp. nov.Merus and carpus of maxilliped 3 dentate ........ 88 88 Posterior pereonites and anterior pleonites each with a pair of dorso-lateral tubercles squamifera Fage, 1929Pereon and pleon without dorso-lateral tubercles . . . torulosa sp. nov. 89 Pereopod 2 comparatively stout. Outer edge of merus and carpus of maxilliped 3 without teeth ........ rubromaculata Lie, 1969 Pereopod 2 slender. Outer edge of merus and carpus of maxilliped 3 strongly dentate . . . . . . . . . . ... .90 90 Uropods longer than the last three pleonites together, the peduncle much more than twice as long as the endopod .... horridoides Stephensen, 1915 Uropods only as long as the last two pleonites together, the peduncle much less than twice as long as the endopod ..... glebulosa sp. nov. 91 Merus of maxilliped 3 triangular, expanded distally and as wide as long frigida Hansen, 1908Merus of maxilliped 3 oblong, never as wide as long ...... 92 92 Merus of maxilliped 3 unusually slender, about three times as long as wide pustulosa Hale, 1945Merus of maxilliped 3 not more than twice as long as wide ..... 93 93 Merus of maxilliped 3 as long as carpus and propodus together .... 94Merus of maxilliped 3 much shorter than carpus and propodus together . . 95 94 Tubercles on carapace low and rounded ..... verrucosa Sars, 1865Tubercles on carapace conical and protruding .... aspera Hale, 1945 95 Pleon somites without dorsal teeth, at most with feeble tubercles on the first three. 96Pleon somites with dorsal teeth ......... 97 96 A depressed area on either side of the carapace. Merus of maxilliped 3 triangularly expanded on inner side. Dactyl of pereopod 2 longer than carpus globosa Hansen, 1920No depressed area on sides of carapace. Merus of maxilliped 3 not expanded on inner side. Dactyl of pereopod 2 barely as long as carpus . nodulosa Sars, 1887 256 N. S. JONES 97 Carpus of maxilliped 3 dentate on outer side. Dactyl of pereopod 2 longer than carpus and propodus together ... . . . antarctica Caiman, 1907 Carpus of maxilliped 3 not dentate on outer side. Dactyl of pereopod 2 shorter than carpus and propodus together ...... multinodosa sp. nov. DESCRIPTIONS AND RECORDS Genus CAMPYLASPIS G. O. Sars, 1865 Five pereon somites visible from above. Carapace strongly vaulted in the female,where it often bulges over the anterior pereon somites. Antero-lateral angles ofcarapace little or not at all produced. Eye unpaired and often rudimentary.Mandible with the molar process styliform. The second maxilla reduced to a simpleplate without movable endites. The first maxilliped reduced with only threesegments, the terminal very small. Second maxilliped with the propodus articulatedat nearly a right-angle to the carpus and ending in a broad seta, the dactyl short andending in two or more distal diverging spines. The first pereopod with the ischiumnot specially elongated. Campylaspis alba Hansen, 1920 MATERIAL. 43°40-8'N, 3°35'2'W, 1739 m, 16.7.1967, 2$$; 28°o6'N, i3°28'W,1780 m, 15.3.1968, 2 $$, 2 juv. DISTRIBUTION. Previously recorded only from southwest of the Faroes, 840-918 m, and possibly from west of Ireland (Hansen, 1920), its range is now extendedto the Bay of Biscay and the Canary Islands and its lower limit to 1780 m. Campylaspis nitens Bonnier, 1896 MATERIAL. 43°35-6'N, 3°24-8W, 860 m, 15.7.1967, 2?$; 43°46-7'N, 3°38'W,2379 m> 18.7.1967, 5 $9, 3 $3 ; 43°43'N, 3°47'8'W, 641 m, 19.7.1967, 54 ??, 3 $$,14 juv. ; 27°29-8'N, i5°20-i'W, 2351 m, 18.3.1968, i ? ; 27°i4'9'N, I5°36-3'W,2988 m, 19.3.1968, i <? ; 07°09'S, 34°25-5'W-O7°o8'S, 34°25'W, 770-805 m, 6.4.1963,i 9 ; 7°58'S, 34°22'W, 834-939 m, 18.2.1967, i 3. DISTRIBUTION. Previously recorded from the Bay of Biscay and west of Ireland(Caiman, 1905) but the latter was probably C. alba (see Hansen, 1920), 570-950 m.Its range is now extended to the Canary Islands and the tropical west Atlantic andits lower limit to 2988 m. Campylaspis nuda sp. nov.(fig. i) MATERIAL. 07°58'S, 34°i7'W-07°5o'S, 34°i7'W, 943-1007 m, 20.2.1967, 4$?,3 A? i o8°o3'S, 34°23'W-o8002'S, 34°25'W, 587 m, 21.2.1967, 15 ??, 13 $$. DESCRIPTION. Adult female, length 3-8 mm : carapace (fig. la, b) smooth andvaulted, evenly rounded dorsally, projecting backwards over the first three pereonites,about one-and-a-half as long as high, more than half the total body length without CAMPYLASPIS FROM THE DEEP ATLANTIC 257 FIG. i. Campylaspis nuda. a, $ from side ; b, $ carapace from above ; c, <£ from side ;d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ; h, $ pereopod 2 ;i, $ uropod and end of pleon. appendages. The pseudorostrum is fairly acute, slightly downturned and less thana sixth of the total carapace length. The eyelobe is short and rudimentary. The first antenna (fig. id) has the three segments of the peduncle about equal inlength. The second maxilliped (fig. le) has the dactyl prolonged into three nearly equalspines ; the merus with a tooth. The third maxilliped (fig. if) broad, the ischium,merus, carpus and propodus serrated on the inner edge, the merus and carpus alsoon the outer edge distally ; the basis much curved, a little shorter than the remainingsegments together. The first pereopod (fig. ig) with the basis about as long as the remaining segmentstogether. The second pereopod (fig. ih) with the dactyl distinctly shorter than thecarpus ; its basis is more than half as long as the remainder of the appendage. 258 N. S. JONES Uropods (fig. li) long, about as long as the last five pleon somites together, theendopod less than half as long as the peduncle, which is serrated on its inner edge ;the endopod is also somewhat serrated and has two spines on the inner edge and twounequal end spines. Adult male, length 4-0 mm (fig. ic) : differing from the female in the usual manner,with the carapace less vaulted and less produced backwards over the pereonites.The second antenna with a long flagellum reaching beyond the end of the body.The uropods relatively longer and with more setae and spines. TYPE LOCALITY. o8°03'S, 34°23'W, 587 m. Types deposited at the BritishMuseum (Natural History). Registration numbers : Holotype 1974 : 283 ; Para-types 1974 : 284. REMARKS. C. nuda resembles C. nitens and C. alba in having a smooth carapacewith rudimentary eyelobe but differs from either of these species in the shape of themerus of maxilliped 3, which is serrated internally but broader than in C. nitens.The dactyl of pereopod 2 is narrower than that of C. nitens. All other describedspecies of Campylaspis with a smooth carapace have the eyelobe well developed,although lenses have not always been distinguished, as is the case with C. paeneglabraStebbing. The latter species has a much longer dactyl of pereopod 2 comparedwith the carpus. C. laevigata sp. nov. following has a reasonably well-developedeyelobe with lenses visible and the dactyl of maxilliped 2 ends in four spines insteadof three. C. guttata sp. nov. following also has a well-developed eyelobe with lenses,while the shape and proportions of its appendages differ considerably ; in particularthe uropods are very much shorter. Recorded at present off Recife, Brazil, in depths between 587 and 1007 m. Campylaspis guttata sp. nov. (fig- 2) MATERIAL. 23°o5'S, i2°3i-5'E, 1546-1559 m, 17.5.1968, 2 ?o ; 9°43'5'S, io°57'E-9°29'S, n°34'E, 2644-2754 m, 22.5.1968, i $. DESCRIPTION. Adult female, length 3-7 mm: carapace (fig. 2a, b) smooth,rounded dorsally, less than one-and-a-half as long as high, antero-lateral anglesrounded and not produced. Pseudorostrum blunt and less than a seventh of thetotal carapace length. Eyelobe well developed with lenses faintly visible. Anumber of small irregular reddish pigment spots are present, mainly postero-laterallybut some on the frontal area. Antenna I (fig. 2c) with the first segment of the peduncle stout and elbowed, alittle longer than the second, which is slightly longer than the third. Maxilliped 2 (fig. 2d) with three spines on the dactyl, the middle one short;the carpus and propodus each with a small tooth on the inner edge ; the terminalseta of the ischiobasis is thickened apically. Maxilliped 3 (fig. 2e) with the basislonger than the rest of the appendage, of which none of the segments is serrated ;the merus and carpus are not widened. CAMPYLASPIS FROM THE DEEP ATLANTIC 259 1 mm FIG. 2. Campylaspis guttata $. a, from side ; b, carapace from above ; c, antenna i ;d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropod and end ofpleon. Pereopod i (fig. 2f) with the basis about as long as the rest of the appendage.Pereopod 2 (fig. 2g) with the basis more than half as long as the remaining segmentstogether ; the dactyl is about as long as the carpus. Uropods (fig. 2h) short and broad, much shorter than the last three pleonitestogether, the peduncle less than one-and-a-half as long as the endopod and serratedalong its inner edge ; the endopod with six spines on the inner edge and two endspines, one much the longer. TYPE LOCALITY. 23°05'S, i2°3i'5'E, 1546-1559 m. Types deposited at theBritish Museum (Natural History). Registration numbers : Holotype 1974 : 285;Paratype 1974 : 286. REMARKS. C. guttata is readily distinguished from other described species of thegenus with a smooth carapace by its comparatively short uropods, which in all theothers have the peduncle at least nearly as long as the last two pleonites togetherand in C. guttata is much shorter than these, and proportionately about as short asin C. platyuropus Caiman. The latter has a ridge on either side of the carapace. Three specimens have been collected off. S.W. Africa. Campylaspis rubicunda (Lilljeborg, 1855) MATERIAL. 39°58-4'N, 7O°4O-3'W, 300 m, 28.8.1962, i juv. ; 39°46«5'N,7°°43'3'W, 1470-1330 m, 25.8.1964, i ?, 3 <&?; 4O°i-6'N, 7O°4O-7'W, 196 m,6.7.1965, 3 ??, 5 <J<? ; 39°38'5'N, 7o°36-5'W-39°39'N, 7O°37'i'W, 2178 m, 18.12.1966, 26o N. S. JONES DISTRIBUTION. There have been many records from the N.E. Atlantic betweenNorway and the British Isles, from the Arctic, from the N.W. Atlantic betweenGreenland and New England and from the Okhotsk Sea in the North Pacific, atdepths between 9 and 1977 m. The lower limit is now extended to 2178 m but it isessentially an inhabitant of the shelf and slope. Campylaspis laevigata sp. nov.(fig-3) MATERIAL. 47°40/N> 5°o'W, 119 m, 12.7.1967, i ?, i £; 43°43'N, 3°47-8'W,641 m, 19.7.1967, i ?, i <J ; 46°i7-5'N, 4°45-2'W, 1336 m, 24.7.1967, 3 ?$, 3 &? ;46°i6-3'N, 4°44'W, 1427 m, 25.7.1967, i ?, i <$ ; 22°54'S, I3°32'E, 205 m, 15.5.1968,i ?, 3 juv. ; 22°53'S, i3°3i'E, 220 m, 16.5.1968, 4 ??. DESCRIPTION. Adult female, length 2-8 mm: carapace (fig. 3a, b) smooth,almost evenly rounded dorsally, a little less than twice as long as high, antero-lateralangles slightly produced. Pseudorostrum rather short, fairly acute, about a seventhof the total carapace length. Eyelobe well developed with lenses visible. Antenna i (fig. 3c) with the first segment of the peduncle rather bent, distinctlylonger than the second or third, which are nearly equal. b,e-h FIG. 3. Campylaspis laevigata $. a, from side ; b, carapace and pereon from above ;c, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropodand end of pleon. CAMPYLASPIS FROM THE DEEP ATLANTIC 261 Maxilliped 2 (fig. 3d) with the propodus rather long, ending in a long flattenedseta ; the dactyl with four spines ; merus, carpus and propodus serrated along theirinner edges. Maxilliped 3 (fig. 36) with the basis distinctly shorter than the remain-ing segments together ; the merus serrated on its outer edge and merus, carpus andpropodus on the inner edge ; propodus and dactyl narrow. Pereopod i (fig. 3f) with the basis slightly shorter than the remaining segmentstogether. Pereopod 2 (fig. 3g) with the basis less than half as long as rest ofappendage ; dactyl narrow, longer than the carpus and propodus together. Uropods (fig. 3h) longer than the last four pleonites together, the endopod muchmore than half as long as the peduncle, serrated internally, with three spines on itsinner edge and a long end spine. TYPE LOCALITY. 47°4o'N, 5°o'W, 119 m. Types deposited at the British Museum(Natural History). Registration numbers: Holotype 1974:287; Paratypes1974 : 288. REMARKS. C. laevigata resembles C. bonetti Bacescu & Muradian in having asmooth carapace and well-developed eyelobe with lenses and four spines on thedactyl of maxilliped 2 but the shape of the latter is rather different and the dactyl ofpereopod 2 is proportionately longer. It also resembles in most respects C. paene-glabra Stebbing but the dactyl of maxilliped 2 in that species has only two spinesaccording to Stebbing (1912) and its carapace has a few slight prominences. FromC. mauritanica Bacescu & Muradian it differs in the shape of maxilliped 2 and especi-ally that of the distal seta of the ischiobasis, which is not swollen nor tufted towardsits tip. The endopod of the uropod is proportionately longer than in C. rubicundaSars. The species with a smooth carapace described in Hale (1945) and Gamo (1960)all have the dactyl of pereopod 2 shorter than the carpus and propodus together. Twenty specimens were collected in the Bay of Biscay and also off S.W. Africabetween 119 and 1427 m. Campylaspis glabra G. O. Sars, 1879 MATERIAL. 38°46'N, 70°o6'W, 2886 m, 21.8.1964, i ?. DISTRIBUTION. Many specimens have been recorded from the N.E. Atlanticfrom Norway to Senegal in West Africa and in the Mediterranean. It has also beenrecorded by Fage (1945) from Vietnam but this identification, based on a singleadult male, may be regarded as doubtful, as may also my own record from Senegalbased on two males. The depth range was from 5 to 2827 m. The present recordextends its range to the N.W. Atlantic and the lower depth range slightly to 2886 m.Like C. rubicunda it is an inhabitant of the shelf and slope, with a broadly over-lapping but more southerly distribution. Campylaspis porcata sp. nov.(fig- 4) MATERIAL. 40°37'N, 7o°5o/W-4O°34'N, 70°5i'W, 68 m, 27.11.1967, 82 ??, 20 3$,3 juv. ; 40°io-8'N, 7o°43-6'W-4O0io-6'N, 70°44'W, 123 m, 28.11.1967, i ?. 262 N. S. JONES FIG. 4. Campylaspis porcata. a, ovigerous $ from side ; b, 9 from above ; c, $ from side ;d, $ antenna i ; e, f, $ maxilliped 2 ; g, $ maxilliped 3 ; h, $ pereopod i ; i, j, $ pereopod2 ; k, $ uropod and end of pleon ; 1, $ uropod and end of pleon. DESCRIPTION. Ovigerous female, length 2-3 mm : carapace (fig. 4a, b) twice aslong as high, with on either side a narrow sulcus becoming a little broader towardsits upper end ; the dorsal surface is smooth in outline and little produced backwards.The pseudorostrum is a little less than a sixth of the total carapace length, withbarely a trace of antennal notch below. The eyelobe is well developed but lenses CAMPYLASPIS FROM THE DEEP ATLANTIC 263 are not visible in the preserved specimens although they seem likely to bepresent. Antenna i (fig. 4<i) with the basal segment of the peduncle about one-and-a-halfas long as the distal segment. Maxilliped 2 (fig. 46, f) with the ischiobasis broad proximally, its apical setaslender and not very long ; the carpus and propodus each with a tooth and thedactyl with three rather long fragile spines, of which the middle is a little shorterthan those on either side. Maxilliped 3 (fig. 4g) with the basis a little shorter thanthe rest of the appendage ; ischium with an apical tooth ; merus long and narrowand serrated on either edge. Pereopod i (fig. 4h) with the basis about as long as the following segments com-bined. Pereopod 2 (fig. 4!) with the basis little more than two-thirds as long as therest of the appendage ; dactyl as long as the carpus and propodus together, endingin three spines of which two are rather stout (fig. 4]). Uropod (fig. 4k) fairly short, shorter than the last three pleonites together; endopodmore than half as long as the peduncle, which is strongly serrated along its inner edgeand a little longer than the exopod ; it has four spines with some serrations inter-spersed along the inner edge and two unequal end spines. Adult male, length 3-2 mm (fig. 4c, 1) : differs from the female in the usual manner.The flagellum of antenna 2 reaches to the end of the uropods. TYPE LOCALITY. 40°37'N, 7o°5o'W-4o°34'N, 70°5i'W, 68 m. Types aredeposited in the British Museum (Natural History). Registration numbers : Holo-type 1974 : 289 ; Paratypes 1974 : 290. REMARKS. C. porcata resembles several other species in having a rather narrowsulcus on either side. These include C. canaliculata Zimmer, C. unisulcata Hale, C.pumila Gamo and C. jonesi Bacescu & Muradian but it differs from all these in theshape of the sulcus, which in none of the other species is narrow anteriorly and widerposteriorly. C. porcata differs from other sulcate species in the proportions of someof the appendages and especially in the armature of the dactyl of pereopod 2. It seems possible that Caiman's record of C. affinis (Caiman, 1912), which he de-scribes as differing from Sars' account in having on either side a shallow obliquedepression and which he refers to C. affinis with considerable doubt, should bereferred to C. porcata. They were found off Marthas Vineyard in 66-71 m in thesame area as this species. Many specimens were obtained off Gay Head in 68-123 m. Campylaspis vitrea Caiman, 1906 MATERIAL. 43°43'N, 3°47-8'W, 641 m, 19.7.1967, 7 ??, 2 <$$, 16 juv. DISTRIBUTION. Previously recorded from the Mediterranean south of Sardiniaand near Capri, 950-1100 m and 1227 m, and doubtfully by Caiman (1912) from thecoast of Labrador in 985 m. The present record from the Bay of Biscay extends itsupper depth range to 641 m and shows that it certainly occurs outside as well aswithin the Mediterranean. 264 N. S. JONES Campylaspis macrophthalma G. O. Sars, 1879 MATERIAL. 47°4o'N, 5°o'W, 119 m, 12.7.1967, 5$$. DISTRIBUTION. Recorded from the Mediterranean between Monaco and theStraits of Messina and from lie d'Yeu in the Bay of Biscay, 38-200 m. The presentrecord extends its range a little further to the north. This species is probablyconfined to the shelf. Campylaspis sticta sp. nov.(fig. 5) MATERIAL. 23°o'S, i2°45'E, 1007-1014 m, 16.5.1968, 7$$. DESCRIPTION. Adult female, length 3-3 mm : carapace (fig. 5a, b) more than twiceas long as high, not at all produced over the pereon somites, with a well-definedgroove on either side bounded by distinct ridges ; the dorsal outline is well elevatedand smooth and apart from two small protuberances on the frontal area there areno spines or other prominences. The pseudorostrum is rather upturned, less thana fifth of the total carapace length, with no trace of an antennal notch below. Theeyelobe is fairly large but without lenses. The carapace, pereon and pleon, and mostof the appendages have many small scattered red blotches of pigment. Antenna I (fig. 5c) with the basal segment of the peduncle strongly curved, littlelonger than either of the more distal segments. Maxilliped 2 (fig. 5d) with the ischiobasis long ; carpus with two teeth on its inneredge ; propodus ending in a slender spine and dactyl with four slender spines, of 1mm 1 mm a,b c-h FIG. 5. Campylaspis sticta 9. a, from side ; b, carapace and pereon from above ; c,antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropodand end of pleon. CAMPYLASPIS FROM THE DEEP ATLANTIC 265 which the most basal is longest. Maxilliped 3 (fig. 5e) with the basis curved, aslong as the ischium to propodus combined, widened at the distal end which is pro-duced well beyond the end of the ischium ; the merus and carpus are fairly broadand serrated on both edges and the propodus on the lower edge. Pereopod i (fig. 5f) with the basis almost as long as rest of appendage ; theischium, merus, carpus and propodus are serrated on either edge. Pereopod 2(fig. 5g) with the basis broad, a little more than half as long as the remaining segmentstogether ; the dactyl is tapered and longer than the carpus and propodus together. Uropod (fig. 5h) as long as the last three pleonites together, peduncle fairly broadand about twice as long as the endopod, which is distinctly longer than the exopod ;the peduncle is serrated on both edges but more distinctly on the inner edge, and theendopod has four spines and some serrations on the inner edge and two unequal endspines. TYPE LOCALITY. 23°o'S, i2°45'E, 1007-1014 m. The types are deposited in theBritish Museum (Natural History). Registration numbers : Holotype 1974 : 291 ;Paratypes 1974 : 292. REMARKS. C. sticta is distinguished from most species with a sulcus on eitherside of the carapace by the absence of lenses in the eyelobe. The sulcus differs fromthat of C. porcata in being of much the same width throughout and among otherdifferences the dactyl of pereopod 2 is tapered and without thickened terminalspines. Seven specimens were found off S.W. Africa in 1007-1014 m. Campylaspis arcuata sp. nov.(fig- 6) MATERIAL. 8°48'S, i2°52'E, 527-542 m, 23.5.1968, 2 ??. DESCRIPTION. Adult female, length 3-3 mm : carapace (fig. 6a, b) twice aslong as high, well elevated dorsally, with two long ridges on either side running backparallel with the lower edge of the carapace, joined by a vertical ridge at the hindend but not meeting the ridge from the opposite side dorsally ; a third shorter ridgeis present dorsal to these, running backwards on either side from behind the sideof the frontal area. The pseudorostrum is about a fifth of the total carapace lengthand not inclined upwards but set at a very obtuse angle to the carapace behind theeyelobe ; there is a little trace of an antennal notch below it. The eyelobe is shortand without lenses. Antenna i (fig. 6c) with the segments of the peduncle slender, the first stronglycurved and shorter than the third. Maxilliped 2 (fig. 6d) with the ischiobasis of moderate length, its distal setanormally pinnate ; the propodus has a distal process reaching as far as the end of thespines of the dactyl, of which there are three of about equal length. Maxilliped 3(fig. 6e) with the basis curved, a little shorter than the rest of the appendage, themerus and carpus broadened but not serrated. 266 FIG. 6. Campylaspis arcuata $. a, from side ; b, carapace from above ; c, antenna i ;d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropod and end ofpleon. Pereopod i (fig. 6f) with the basis very little shorter than the remaining segmentstogether ; these are without spines or serrations. Pereopod 2 (fig. 6g) with thebasis less than half as long as rest of appendage ; the dactyl is not tapered and withoutdistal spines ; it is about as long as the carpus and propodus together. Uropod (fig. 6h) a little shorter than the last three pleonites combined ; thepeduncle is serrated on either side and twice as long as the endopod, which is onlyslightly longer than the exopod and has four or five spines and some serrations on itsinner edge and a long end spine. TYPE LOCALITY. 8°48'S, I2°52'E, 527-542 m. Types deposited in the BritishMuseum (Natural History). Registration numbers : Holotype 1974 : 293 ; Para-type 1974 : 294. REMARKS. C. arcuata resembles C. exarata sp. nov. a little in the shape of thecarapace but it is more elevated dorsally and its pseudorostrum is not upturned,while it lacks prominences at the sides of the frontal area. The shape of the dactylof pereopod 2 is quite different in the two species. Two specimens were collected off S.W. Africa in 527-542 m. Campylaspis costata G. O. Sars, 1865 MATERIAL. 39°46-5'N, 7o°43'3'W, 1470-1330 m, 25.8.1964, i <j>. DISTRIBUTION. Many records in the N.E. Atlantic from northern Norway to theBritish Isles, 23-478 m. Although apparently almost confined to the shelf in N.W.Europe this species can apparently occur on the slope off N.E. America down toabout 1470 m. A subspecies, C. c. speciosa, has been described by Lomakina (1955)from the northern Pacific but I am inclined to regard it as a separate species. CAMPYLASPIS FROM THE DEEP ATLANTIC Campylaspis spinosa Caiman, 1906 267 MATERIAL. 28°o6'N, i3°28'W, 1780 m, 15.3.1968, 2 $9, i £ ; c^og'S, 34°25'5'W-o;°o8'S, 34°25'W, 770-805 m, 6.4.1963, i juv. ; 7°58'S, 34°22'W, 834-939 m,18.2.1967, 3??, i& i juv.; 7°58'S, 340i7'W-7°5o'S, 34°i7'W, 943-1007 m,20.2.1967, 9 $$, 2 <?<J, i juv. ; 8°03'S, 34°23'W-8°O2'S, 34°25'W, 587 m, 21.2.1967,56 $$, 9 $$, 8 juv. ; 23°o'S, i2°45'E, 1007-1014 m, 16.5.1968, 2 ??. DISTRIBUTION. Recorded previously from the western Mediterranean (Caiman,1906 ; Reyss, 1972), 950-2924 m. The present records extend its geographicalrange to the Canaries, the tropical West Atlantic and the coast of S.W. Africa andits upper depth limit to 587 m. Campylaspis rostellata sp. nov.(fig- 7) MATERIAL. 27°i3'N, i5°4i'W, 3301 m, 20.3.1968, i $. DESCRIPTION. Adult female, length 4-2 mm : carapace (fig. 7a, b) more thantwice as long as high, smooth at the sides but with fairly numerous small spinesdorso-laterally and on the frontal area. The pseudorostrum is two-ninths of thetotal carapace length, prominent and sharply pointed when viewed from the side ;behind it the side of the carapace is produced into an antero-lateral angle, withserrations extending backwards. The eyelobe is rudimentary. The pereon somites are serrated laterally (fig. 7b), the pleon somites laterally andthe hinder ones also dorsally. a,b c-h FIG. 7. Campylaspis rostellata $. a, from side ; b, carapace and pereon from abovec, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, basis of pereopod i ; g, pereopod 2h, uropod and end of pleon. 268 N. S. JONES Antenna i (fig. 70) with the basal segment of the peduncle elbowed near the base,nearly twice as long as the third segment. Maxilliped 2 (fig. yd) with the ischiobasis long and with a robust but not distallythickened terminal seta ; carpus with two teeth on its inner edge ; propodus with along tooth and dactyl with three equal teeth. Maxilliped 3 (fig. ye) with the basiscurved, longer than rest of appendage ; ischium with a tooth on the lower edge ;merus nearly twice as long as the carpus, both segments with some small serrationson the upper edge. Pereopod i (fig. 71) incomplete in the specimen but the basis curved and serratedon both edges distally. Pereopod 2 (fig. 7g) with the basis about two-thirds as longas remaining segments together ; merus with a terminal tooth on its lower edge ;dactyl narrow and nearly as long as the carpus and propodus together. Uropod (fig. yh) longer than last four pleonites combined ; peduncle with strongserrations along either edge, more than twice as long as the endopod, which has someserrations along the inner edge basally, two spines distally and a stout end spine ;exopod about as long as the endopod, with a long slender terminal spine. TYPE LOCALITY. 27°i3'N, i5°4i'W, 3301 m. Type deposited in the BritishMuseum (Natural History). Registration number : Holotype 1974 : 295. REMARKS. C. rostellata somewhat resembles C. rostrata Caiman and C. aculeatasp. nov. in the shape of the pseudorostrum but is easily distinguished from either bythe markings of the carapace, from C. rostrata by the relative length of the uropodsand from C. aculeata by its much smaller size and lack of spinulation on most of theappendages. A single specimen was obtained off the Canary Islands in 3301 m. Campylaspis aculeata sp. nov.(fig- 8) MATERIAL. o°46'S, 29°28'W-o°46-5'S, 29°24'W, 3459m, 14.2.1967, 3$$, j ^4 juv. DESCRIPTION. Ovigerous female, length 9-6 mm : carapace (fig. 8a) a little morethan twice as long as high, with a mid-dorsal groove reaching from the eyelobe tonear the hind end and then bifurcating obliquely ; the whole surface is covered withnumerous small spines. Pseudorostrum two-ninths of total carapace length,prominent and sharply pointed when viewed laterally ; no antero-lateral angle isvisible but the lower part of the carapace is serrated. The eyelobe is obsolete. The first and second pereonites each have a pair of blunt mid-dorsal spines andthe fourth has a ring of spines. The pleon somites have numerous spines at the sidesand dorso-laterally. Antenna i (fig. 8b) with the first two segments of the peduncle studded with bluntspines or serrations ; the first segment nearly twice as long as the second and morethan twice as long as the third. Maxilliped 2 (fig. 8c) with the ischiobasis fairly long, the terminal pinnate setalong and slender ; there are five spines on the inner edge of the carpus ; the propodus CAMPYLASPIS FROM THE DEEP ATLANTIC 269 b,c FIG. 8. Campylaspis aculeata ovigerous $. a, from side ; b, antenna i ; c, maxilliped 2d, maxilliped 3 ; e, pereopod i ; f, pereopod 2 ; g, uropod and end of pleon. has one terminal spine and the dactyl three of nearly equal length. Maxilliped 3(fig. 8d) with the basis curved, broadened basally, a little longer than the remainingsegments together, with a number of spines on the lower edge ; ischium, merus,carpus and propodus all have one or a number of spines on one or both edges. Pereopod i (fig. 8e) with the basis curved, distinctly longer than the other segmentscombined ; all the segments have a number of spines or serrations, the merus, carpusand propodus on both edges, and the propodus also has a long spine on the lower edge.Pereopod 2 (fig. 8f) with the basis a little shorter than the rest of the appendage,serrated on both edges, as is the carpus along its basal part ; the propodus is muchshorter than the carpus. Uropod (fig. 8g) longer than the last five pleonites together ; the peduncle isnearly two-and-a-half as long as the endopod, which is distinctly longer than theexopod ; the peduncle has many short spines and serrations and the endopod hassix slender spines on its inner edge and a longer end spine. TYPE LOCALITY. o°46'S, 29°28'W-o°46-5'S, 29°24'W, 3459 m. The types aredeposited in the British Museum (Natural History). Registration numbers :Holotype 1974 : 296 ; Paratypes 1974 : 297. 270 N. S. JONES REMARKS. C. aculeata is unusually large for the genus. The shape of thecarapace is rather similar to that of C. rostellata sp. nov. but it is much more spiny.The spines are much smaller than in C. spinosa Caiman and the pseudorostrum is notupturned as in that species. From C. echinata Hale it is distinguished by the shapeof the carapace and of the second maxillipeds and second pereopods, among otherdetails. In general shape and size this species approaches the genus Campylaspides Page,and especially to Campylaspides spinifera Jones (1973). However, although theshape of its second maxillipeds might be thought to represent a transitional stagetowards Campylaspides, having several well-developed carpal spines, the propodusand dactyl retain the form typical of Campylaspis. Eight specimens were found near the mid-Atlantic Ridge between Dakar andRecife in 3459 m. Campylaspis brevicornis sp. nov. (fig. 9)MATERIAL. 23°O2'S, i2°ig'E, 2117-2154 m, 17.5.1968, i <$. DESCRIPTION. Adult male, length 3-8 mm : carapace (fig. ga, b) more thantwice as long as high, with some small rounded prominences dorsally and withnumerous short hairs, the dorsal outline not much elevated and rather irregular. FIG. 9. Campylaspis brevicornis <J. a, from side ; b, carapace and pereon from above ;c, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropodand end of pleon. CAMPYLASPIS FROM THE DEEP ATLANTIC 271 Pseudorostrum fairly large and prominent ; antero-lateral angle not at all pro-duced and some infero-lateral serrations present. Eyelobe rudimentary. The first two pereonites with dorsal projections. The whole body and most of theappendages have numerous small reddish pigment spots. Antenna i (fig. gc) with the segments of the peduncle decreasing successively inlength, the basal segment straight and nearly twice as long as the third. Antenna 2(fig. ga) with the flagellum rather short, not reaching beyond the pereon. Maxilliped 2 (fig. gd) with the ischiobasis broad, its distal seta thickened and witha distal tuft of hairs ; the propodus with a fairly short distal seta and a small toothon its inner edge ; the dactyl with three spines, the middle one small. Maxilliped 3(fig. ge) with its segments fairly narrow, unarmed, the basis as long as the remainingsegments together, with two rather short and thick pinnate setae distally ; the meruslong and narrowed distally, more than twice as long as the carpus. Pereopod i (fig. gf) with the basis broadened proximally, a little shorter than therest of the appendage. Pereopod 2 (fig. gg) with the basis a little more than half aslong as the rest of the appendage ; the dactyl narrowed distally, about as long as thecarpus. Uropod (fig. gh) incomplete in the specimen but longer than the pleon, thepeduncle about twice as long as the endopod ; the peduncle has about eight spineson its inner edge and the endopod at least five. TYPE LOCALITY. 23°O2'S, i2°ig'E, 2117-2154 m. Type deposited in the BritishMuseum (Natural History). Registration number : Holotype ig74 : 2g8. REMARKS. It is difficult to see if the flagellum of the second antenna is completebut it is certainly shorter than usual for the genus and distinguishes the adult malefrom other species with a similarly prominent pseudorostrum. A single male was found off S.W. Africa in 2117-2154 m. Campylaspis pilosa sp. nov.(fig. 10) MATERIAL. 7°o8'S, 34°i7'W-7°5o'S, 34°i7'W, g43-ioo7 m, 20.2.1967, 5 ??, i $ ;8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m, 21.2.1967, 5 ??. DESCRIPTION. Adult female, length 4-0 mm : carapace (fig. loa, b) about twiceas long as high, with a blunt prominence on either side of the frontal area and an ill-defined groove on either side above and below it; a number of moderately long hairsare scattered over the dorsal surface. The pseudorostrum is more than a sixth ofthe total carapace length and the antero-lateral angle is slightly produced on eitherside below it. The eyelobe is rudimentary. Some blunt spines and numerous short hairs are present on the sides of the pleonsomites. Antenna i (fig. loc) with the peduncle segments slender, the first a little curved. Maxilliped 2 (fig. lod) with the ischiobasis broad, its distal seta much thickened atthe distal end ; propodus with a fairly stout spine distally and a prominent tooth on 272 N. S. JONES c-h FIG. 10. Campylaspis pilosa $. a, from side ; b, carapace and pereon from above ;c, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropodand end of pleon. the inner side ; dactyl with three spines, the middle one small. Maxilliped 3 (fig.ice) with the basis a little shorter than the remaining segments together and stronglycurved, the other segments slender and unarmed, the merus nearly twice as long asthe propodus. Pereopod i (fig. lof) fairly slender, with the basis much shorter than the rest of theappendage. Pereopod 2 (fig. log) with the basis about half as long as remainingsegments together ; the dactyl narrow, about as long as the carpus. Uropod (fig. loh) about as long as the last four pleonites together, the endopod alittle less than half as long as the peduncle, which has a few setae along its inneredge ; the endopod has three spines and one terminal spine and is about as long asthe exopod, which has several short spines along its outer edge and a long end spine. TYPE LOCALITY. 8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m. Types deposited inthe British Museum (Natural History). Registration numbers : Holotype 1974 :299 ; Paratypes 1974 : 300. REMARKS. The rather hairy carapace without any other projections except thetwo antero-lateral prominences distinguish this species from others with a lateralgroove. Twelve specimens have been collected off Recife, Brazil, in 587-1007 m. CAMPYLASPIS FROM THE DEEP ATLANTIC Campylaspis bulbosa sp. nov. 273 MATERIAL. 23°O2'S, i2°ig'E, 2117-2154111, 17.5.1968, 5$$. DESCRIPTION. Immature female, length 3-3 mm : carapace (fig. na, b) lessthan twice as long as high, broad and bulbous, with a well-defined ridge on eitherside running from below the pseudorostrum backwards to the dorsal side aboutthree-quarters of the carapace length from the front, where it is joined by a shorttransverse ridge with some blunt teeth to its fellow on the opposite side ; near thelower edge of the carapace is a less prominent ridge ; two further short transverseridges are present on either side of the frontal area, not quite meeting on the mid-line, the more anterior with a low prominence in the middle on either side ; below thelong ridge is a large oval shallow depression and a further small depression is presenton either side dorsally behind the posterior transverse ridge ; there are a fewscattered hairs on the dorsal surface. The pseudorostrum is short, only about aneighth of the total carapace length and there is little trace of an antennal notch.The eyelobe is small but broad, without lenses. Antenna i (fig. nc) with the basal segment of the peduncle curved, more thanone-and-a-half as long as the distal segment. Maxilliped 2 (fig. ud) with the ischiobasis broad, its distal seta rather short butnot thickened ; the propodus has a tooth on its inner edge and a long distal spine 1 mm FIG. ii. Campylaspis bulbosa immature $. a, from side ; b, carapace and pereon fromabove ; c, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ;h, uropod and end of pleon. 274 N. S. JONES and the dactyl has three spines of which the middle is fairly short. Maxilliped 3(fig. ne) with the basis strongly curved, longer than the remaining segmentstogether ; the ischium has two sharp teeth on the lower edge and the merus is fairlynarrow, with a distal tooth on the upper edge. Pereopod I (fig. nf) with the basis curved, as long as the following four segmentstogether. Pereopod 2 (fig. ng) with the basis a little less than a third of the totallength ; the dactyl is broad basally, a little longer than the carpus and propodustogether, and has three long slender terminal spines and a single thicker short bluntspine. Uropod (fig. nh) with the peduncle about as long as the last three pleonites com-bined and serrated along its inner edge ; the endopod is two-thirds as long as thepeduncle and very slightly shorter than the exopod ; it has three slender spines anda few serrations along the inner edge and two unequal end spines. TYPE LOCALITY. 23°02'S, I2°I9'E, 2117-2154 m. Types deposited in the BritishMuseum (Natural History). Registration numbers : Holotype 1974 : 301 ; Para-types 1974 : 302. REMARKS. C. bulbosa is distinguished by the shape and markings of the carapacefrom any other known species in the genus. Five females, none of them fully grown, occurred off S.W. Africa in 2117-2154 m. Campylaspis redacta sp. nov. (fig. 12) MATERIAL. 8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m, 21.2.1967, 15 ?o, 8 &?. DESCRIPTION. Immature male, length 2-2 mm : carapace (fig. I2a, b) less thantwice as long as high, with a large irregular sulcus on either side not bounded byraised carinae and with some hollows and irregularities dorsally. The pseudorostrumis slightly downturned, a little less than a fifth of the total carapace length, and thereis no antennal notch. The eyelobe is rudimentary and minute. The fourth and fifth pereonites have a number of dorsal teeth. Antenna i (fig. i2c) has the third segment of the peduncle almost as long as thebasal segment. Maxilliped 2 (fig. I2d) with the ischiobasis long, its distal seta slender ; the pro-podus is long, with a very long spine, and the dactyl has three equal spines. Maxil-liped 3 (fig. I2e) with the basis short and broad, much shorter than the remainingsegments together ; the merus is long, more than twice as long as the propodus,while the carpus is very short. Pereopod i (fig. 121) with the basis distinctly shorter than the rest of the appendage.Pereopod 2 (fig. I2g) with the basis not much more than half as long as the rest ofthe appendage ; the dactyl is fairly slender and longer than the carpus and propodustogether. Uropod (fig. I2b) slightly longer than the last three pleonites together ; the exopodis nearly as long as the endopod, which is more than half as long as the peduncle ;the peduncle is serrated along its inner edge and has a number of short setae whilethe endopod has five spines on the inner edge and a long end spine. CAMPYLASPIS FROM THE DEEP ATLANTIC 275 FIG. 12. Campylaspis redacta subadult (J. a, from side ; b, from above ; c, antenna i ;d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2. TYPE LOCALITY. 8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m. Types deposited inthe British Museum (Natural History). Registration numbers : Holotype 1974 :303 ; Paratypes 1974 : 304. REMARKS. The length of the largest specimen was 3-0 mm but none was mature.In having a fairly large sulcus on either side of the carapace C. redacta somewhatresembles C. cognata sp. nov. but there are many differences in the appendages. InC. sulcata Sars the sulcus is of a different shape and the latter species has a well-developed eyelobe with lenses. Twenty-three specimens were collected off Recife, Brazil, in 587 m. Campylaspis paucispina sp. nov.(fig- 13) MATERIAL. 39°26'N, 7O°33/W, 2496 m, 21.8.1964, 2 ?? ; 39°46-5'N, 7O°43'3'W,1470-1330 m, 25.8.1964,17 $$, 2 $$ ; 39°48-7'N, 7o°40'8'W, 1102 m, 6.7.1965, 7 Q$ ;39°43-6'N, 70°37-4'W, 2022 m, 4.5.1966, 8??. DESCRIPTION. Adult female, length 4-2 mm : carapace (fig. i3a) twice as longas high, integument rather scabrous, not much elevated, with two ill-defined obliqueridges on either side with a slight hollow between them, somewhat undulating witha few smallish spines on them at intervals ; there are some more spines or slightprominences on the dorsal surface. The pseudorostrum is less than a fifth of thetotal carapace length and is slightly upturned ; the antero-lateral angle is slightly 276 N. S. JONES FIG. 13. Campylaspis paucispina. a, $ from side ; b, $ carapace and pereon from side ;c, 9 antenna i ; d, e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ; h, $ pereopod 2 ;i, $ uropod and end of pleon ; j, $ uropod and end of pleon. produced and the infero-lateral edge is serrated behind it. The eyelobe is rudimen-tary. Antenna i (fig. 130) with the segments of the peduncle differing little in length, thebasal segment curved and serrated below. Maxilliped 2 (fig. I3d, e) has the apical seta on the ischiobasis thickened, with adistal brush of hairs ; the propodus has a longish distal spine and a shorter spine onits inner edge ; the dactyl has three spines of which the middle is very small.Maxilliped 3 (fig. 131) with the basis curved, a little shorter than the remaining seg-ments together ; the merus is long and the carpus broad, both segments serratedalong the upper edge. CAMPYLASPIS FROM THE DEEP ATLANTIC 277 Pereopod i (fig. I3g) with the basis curved, shorter than the rest of the appendage,with some spines on its outer side ; the ischium is serrated below and the carpusabove. Pereopod 2 (fig. I3h) with the basis about half as long as the remainder ofthe appendage ; the dactyl is as long as the carpus and propodus combined ; thecarpus is serrated on its upper edge. Uropod (fig. 131) nearly as long as the last four pleonites together, the peduncleabout one-and-a-half as long as the endopod, which is a little longer than the exopod ;the peduncle is strongly serrated on either edge and the endopod has a few spines onthe inner edge and a longer end spine. Adult male, length 3-8 mm : differs from the female in the usual manner. Thefirst two pereonites are produced dorsally. The second antennal flagellum is long(fig. i3b, j). TYPE LOCALITY. 39°26'N, 7O°33'W, 2496 m, 21.8.1964. Types deposited in theBritish Museum (Natural History). Registration numbers : Holotype 1974 : 305 ;Paratypes 1974 : 306. REMARKS. In having a carapace with comparatively few protuberances and ratherill-defined lateral ridges this species resembles C. intermedia Hansen but in the latterthe prominences are mostly larger and blunter and the antero-lateral angle is moredeveloped. The shape of the distal seta of the ischiobasis of maxilliped 2 resemblesthat of C. affinis Sars and also of some of the other species described here but none ofthese has a similarly marked carapace. A fair number of specimens were collected south of Gay Head between 1102 and2496 m. Campylaspis cognata sp. nov. (fig. 14) MATERIAL. 8°02;S, 34°03'W-7°56'S, 34°O9'W, 1493 m, 19.2.1967, i $ ; 7°58'S,34°i7'W-7°5o'S, 34°i7'W, 943-1007 m, 20.2.1967, 20$$, 6 &? ; 8°o3'S, 34°23'W-8°02'S, 34°25'W, 587 m, 21.2.1967, 10$?, 12 &?. DESCRIPTION. Ovigerous female, length 2-7 mm : carapace (fig. i4a, b) more thantwice as long as high with a sulcus of moderate breadth on either side defined by lowkeels, the uppermost of which has two blunt spines ; dorsal outline fairly smoothbut a fairly large prominence bearing a spine is present on either side below thefrontal area. The pseudorostrum is less than a fifth of the total carapace length andthere is a slightly excavated antennal notch below it. The eyelobe is fairly welldeveloped, though without lenses. Twelve embryos were contained in the broodpouch. The first three pereonites are broad (fig. I4b) and the first three pleonites havesome dorso-lateral spines. Antenna i (fig. I4d) with the basal segment of the peduncle curved, distinctlylonger than the third segment. Maxilliped 2 (fig. 146) with the ischiobasis broad, its terminal feathered setaslender ; propodus with an end spine of moderate length and a prominent tooth onthe inner edge ; dactyl with three spines, the middle one small. Maxilliped 3 278 N. S. JONES FIG. 14. Campylaspis cognata. a, $ from side ; b, $ carapace and pereon from above ;c, o* from side ; d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ;h, $ pereopod 2 ; i, $ uropod and end of pleon. (fig. I4f) with the basis broadened distally, only a little shorter than the remainingsegments combined ; the merus is about twice as long as the carpus and not verymuch longer than the propodus. Pereopod i (fig. I4g) with the basis very slightly shorter than the rest of the ap-pendage ; the merus has a tooth distally and the carpus several serrations along theupper edge. Pereopod 2 (fig. I4h) with the basis less than half as long as the remain-ing segments together ; the dactyl is fairly broad proximally and has a long robustdistal spine ; it is slightly shorter than the carpus and propodus together. Uropod (fig. 14!) longer than the last three pleonites together, the peduncle ser-rated along both edges ; the endopod is rather more than half as long as the peduncleand distinctly longer than the exopod, with some serrations and three spines alongthe inner edge and two unequal end spines. Adult male, length 3-0 mm (fig. i4c) : differs from the female in the usual charac-ters. The antennal notch is deeper and the pseudorostrum more truncated. Thesecond antennal flagellum reaches to the end of the uropods. TYPE LOCALITY. 8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m. Types deposited inthe British Museum (Natural History). Registration numbers : Holotype 1974 :307 ; Paratypes 1974 : 308. REMARKS. The size of the eyelobe and the shape and proportions of maxillipeds2 and 3 and pereopod 2 among other characters differentiate this species from C.redacta. The presence of a few blunt spines distinguish it from other sulcate species. A fairly large number of specimens were obtained off Recife, Brazil, between 587and 1493 m. CAMPYLASPIS FROM THE DEEP ATLANTIC 279 Campylaspis caperata sp. nov. (fig- 15) MATERIAL. io°3o'N, i7°5i-5'W, 1624-1796 m, 5.2.1967, 5 $o, 6 $$. DESCRIPTION. Adult female, length 3-8 mm : carapace (fig. i5a, b) much lessthan twice as long as high, covered with minute spiny scales and with a few hairs,with two oblique ridges on either side, the lower one less well defined, and two moreshort transverse ridges behind the frontal area ; there are a number of undulationsand prominences, especially dorsally. The pseudorostrum is less than a sixth of thetotal carapace length, truncated, with a shallow antennal notch below. The eyelobeis rudimentary. Antenna i (fig. I5c) with the segments of the peduncle fairly slender, the firstdistinctly less than twice as long as the third. Maxilliped 2 (fig. i5d) with the ischiobasis broad, its terminal seta thickened andtufted apically ; propodus with a long terminal spine and dactyl with three spines ofwhich the middle is very small. Maxilliped 3 (fig. I5e) with the basis curved,broadened distally, much shorter than the remaining segments combined ; the merusis about twice as long as the carpus and both are unarmed. Pereopod i (fig. I5f) with the basis nearly as long as the remainder of the appen-dage, all the segments of which are unserrated. Pereopod 2 (fig. I5g) with the basisabout two-thirds as long as the remaining segments together ; the propodus is very FIG. 15. Campylaspis caperata ?. a, from side ; b, from above ; c, antenna i ; d, maxilli-ped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropod and end of pleon. 16 280 N. S. JONES short, only a third as long as the carpus, while the dactyl is a little less than half aslong as the carpus. Uropod (fig. I5h) a little longer than the last three pleonites together ; the endopodis less than half as long as the peduncle, which is serrated along its inner edge ; theendopod has three slender spines on its inner edge and one longer and a shorter endspine ; the exopod is almost as long as the endopod. TYPE LOCALITY. io°3o'N, i7°5i-5'W, 1624-1796 m. Types deposited in theBritish Museum (Natural History). Registration numbers : Holotype 1974 : 309 ;Paratypes 1974 : 310. REMARKS. The combination of ridges and irregular prominences on the carapaceis unlike that of any other species. C. caperata is also distinguished by the shortnessof the dactyl of pereopod 2 compared with other species. Eleven specimens were collected off Dakar in 1624-1796 m. Campylaspis plicata sp. nov.(fig. 16) MATERIAL. 7°58'S, 340i7'W-7°5o'S, 34°i7'W, 943-1007 m, 20.2.1967, 4 $$, 6 $$. DESCRIPTION. Adult female, length 4-8 mm : carapace (fig. i6a, b) about twiceas long as high, with two oblique ridges on either side, the anterior ridge bifurcatingbehind the side of the frontal area to cross the dorsal surface and join with its fellowfrom the other side ; on the dorsal surface there are a number of low spines, with alarger prominence on either side of the frontal area. The pseudorostrum is less thana fifth of the total carapace length and fairly sharply pointed when viewed laterally,with a shallow antennal notch below, followed by some serrations. The eyelobe isrudimentary. The first two pereonites are raised dorsally. Antenna I (fig. i6d) with the basal segment of the peduncle about twice as long asthe third. Maxilliped 2 (fig. i6e) with the ischiobasis fairly long ; the merus has one toothand the carpus two ; the propodus ends in a long slender spine and the dactyl hasthree fairly long spines of almost equal length. Maxilliped 3 (fig. i6f) with thebasis curved, broadened distally and serrated along the distal lower edge ; the merusis a little broadened distally and serrated, and the carpus is fairly broad ; carpusand propodus have serrations along the lower edge. Pereopod i (fig. i6g) with the basis about two-fifths as long as the rest of theappendage, serrated on either edge distally ; the ischium has a single tooth on thelower edge and the merus, carpus and propodus are serrated on either edge. Pereo-pod 2 (fig. i6h) with the basis three-quarters as long as the rest of the appendage ;the dactyl is narrow, about two-thirds as long as the carpus, and ends in a thick spine. Uropod (fig. i6i) about as long as the last four pleonites together, the peduncleserrated on either edge ; the endopod is much longer than the exopod and morethan half as long as the peduncle, with three spines on its serrated inner edge and alonger end spine. CAMPYLASPIS FROM THE DEEP ATLANTIC 281 FIG. 16. Campylaspis plicata. a, $ from side ; b, $ from above ; c, <$ carapace andpereon from side ; d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ;h, $ pereopod 2 ; i, $ uropod and end of pleon ; j, $ uropod and end of pleon. Adult male, length 5-3 mm (fig. i6c, j) : differs from the female in the usual manner.The second antennal flagellum reaches to the end of the uropods which are propor-tionately longer than in the female and have a number of feathered setae along theinner edge and about eight spines on the inner edge of the endopod. TYPE LOCALITY. 7°58'S, 34°i7'W-7°5o'S, 34°i7'W, 943-1007 m. Types de-posited in the British Museum (Natural History). Registration numbers : Holotype1974 : 311 ; Paratypes 1974 : 312. 282 N. S. JONES REMARKS. In general appearance C. plicata somewhat resembles C. caperata butdiffers in the relative shapes and proportions of the appendages, among which maybe mentioned the terminal seta of the ischiobasis of maxilliped 2, which is not thick-ened as it is in C. caperata, while the latter species lacks serrations on maxilliped 3and pereopod i. Ten specimens were found off Recife, Brazil, in 943-1007 m. Campylaspis valleculata sp. nov.(fig- 17) MATERIAL. 38°46'N, 7o°o6'W, 2886 m, 21.8.1964, 29 ?$, 18 &?, 38 juv. ;38°46-7'N, 70°o8-8'W, 2802 m, 21.8.1964, 5 ??, 2 &J ; 38°i6'N, 7i°47'W, 2864m,24.8.1964, 4 9$, i juv. ; 39°38'3'N, 67°57-8'W, 2862 m, 29.6.1965, i o, T juv. ;39°48-7'N, 70°40-8'W, 1102 m, 6.7.1965, i ?, 2 juv. ; 39°37'W, 66°47'W-39°37-5'N,66°44'W, 3806 m, 24.8.1966, 2 ??. DESCRIPTION. Adult female, length 6-7 mm : carapace (fig. i7a, b) more thantwice as long as high, with three well-defined ridges on either side, the upper minutelycrenulated and running backwards from below the front of the pseudorostrum tomeet its fellow from the other side on the mid-line, where it curves forward at ratherless than a sixth of the total carapace length from the hind end ; the middle ridgeruns backwards nearly parallel to the uppermost from well below and behind theantero-lateral angle, with a slight upward curve near the hind end of the carapaceafter which it dips to join its fellow from the other side to define the hind end ; anundulating ridge joins the middle ridge from the top of the upward curve obliquelyforward to the upper ridge ; the lower ridge starts a little below the middle one andruns backwards parallel with the lower edge of the carapace to end below the upwardcurve of the middle ridge ; there are several blunt prominences on either side of thefrontal area and a number of scattered short hairs dorsally. The pseudorostrum isa little less than a sixth of the total carapace length and fairly acute from the side ;there is only a faint indication of an antennal notch and antero-lateral angle. Theeyelobe is fairly long and narrow, without lenses. Antenna i (fig. I7d) with the segments of the peduncle long and narrow, the basalstrongly curved and nearly twice as long as the third. Maxilliped 2 (fig. 176) with the ischiobasis moderately long, its distal seta fairlyshort ; the propodus has a small tooth on the inner edge and a long end process ; thedactyl has three spines, the middle one very small. Maxilliped 3 (fig. i7f) with thebasis only a little shorter than the rest of the appendage ; all the segments are com-paratively slender. Pereopod i (fig. I7g) with the basis curved, about as long as the remaining seg-ments together. Pereopod 2 (fig. I7h) with the basis more than half as long as restof appendage ; the dactyl is tapered, its terminal spines not thickened, and isdistinctly longer than the carpus and propodus together. Uropod (fig. 17!) about as long as the last four pleonites together, the pedunclefaintly serrated and with a few short setae along the inner edge ; it is nearly three CAMPYLASPIS FROM THE DEEP ATLANTIC 283 FIG. 17. Campylaspis valleculata. a, $ carapace from side ; b, $ carapace and pereon fromabove ; c, $ from side ; d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopodi ; h, $ pereopod 2 ; i, $ uropod and end of pleon ; j, £ uropod and end of pleon. times as long as the endopod, which has three slender spines on the inner edge andtwo unequal end spines ; the exopod is only a little shorter than the endopod. Adult male, length 8-1 mm (fig. 170, j) : differs from female in the usual characters ;the second antennal flagellum reaches to the end of the uropods. TYPE LOCALITY. 38°46'N, 7o°o6'W, 2886 m. Types deposited in the BritishMuseum (Natural History). Registration numbers : Holotype 1974 : 313 ; Para-types 1974 : 314. REMARKS. C. valleculata is large for the genus. The shape of the carapace issomewhat similar to that of C. undata Sars and C. mansa sp. nov. but it is distinguished 284 N. S. JONES from either by the presence of a third lower ridge on either side and by a number ofother characters. Many specimens were obtained on the Gay Head - Bermuda transect between1102 and 3806 m. Campylaspis exarata sp. nov.(fig. 18) MATERIAL. 8°O3'S, 34°23'W-8°02'S, 34°25'W, 587 m, 21.2.1967, 6 $?. DESCRIPTION. Adult female, length 4-7 mm : carapace (fig. i8a, b) about two-and-a-half as long as high, produced backwards to overhang the anterior pereonites,with two well-defined, somewhat undulating ridges running obliquely backwards,more or less parallel with the lower edge of the carapace ; the upper edge does notreach as far as the mid-dorsal line ; a fairly large blunt prominence is present oneither side of the frontal area. The pseudorostrum is more than a fifth of the total FIG. 18. Campylaspis exarata $. a, carapace from side ; b, from above ; c, antenna i ;d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropod and end ofpleon. CAMPYLASPIS FROM THE DEEP ATLANTIC 285 carapace length and is obliquely upturned, with a distinctly excavated antennalnotch below. The eyelobe is small and narrow, without lenses. Antenna I (fig. i8c) with the basal segment of the peduncle straight, about one-and-a-half as long as the third segment. Maxilliped 2 (fig. i8d) with the ischiobasis fairly long, its distal seta slender andnormally pinnate ; propodus with a fairly slender terminal process reaching wellbeyond the dactyl, which has three spines, the middle small. Maxilliped 3 (fig. iSe)with the basis curved, distinctly shorter than the rest of the appendage, and with theupper distal part somewhat produced ; the remaining segments are rather slenderand uriserrated. Pereopod i (fig. i8f) with the basis a little shorter than remaining segmentscombined, somewhat curved, with several teeth on the distal upper edge ; the merusand carpus are serrated on the lower edge and have a number of spines on the outerside. Pereopod 2 (fig. i8g) with the basis narrow, little more than half as long as therest of the appendage ; the dactyl is tapered, with several slender spines distally,and is shorter than the carpus and propodus combined. Uropod (fig. i8h) about as long as the last three pleonites together ; the pedunclesare strongly serrated on either side and about twice as long as the endopod, which isa little longer than the exopod and has four spines on its inner edge and two unequalend spines. TYPE LOCALITY. 8°03'S, 34°23'W-8°02'S, 34°25'W, 587 m. Types deposited inthe British Museum (Natural History). Registration numbers : Holotype 1974 :315 ; Paratypes 1974 : 316. REMARKS. C. exarata, with its lateral ridges on the carapace and upturnedpseudorostrum, considerably resembles C. vitrea Caiman. However, the rear dorsalside of the carapace has a different appearance, the uppermost ridge being in C.vitrea joined by its fellow from the opposite side by a short ridge passing straightacross the top of the carapace. Among other differences the dactyl of pereopod2 in C. vitrea is distinctly longer than the carpus and propodus combined. Six specimens were collected off Recife, Brazil, in 587 m. Campylaspis mansa sp. nov.(fig. 19) MATERIAL. 38°46'N, 7o°o6'W, 2886m, 21.8.1964, 2$$, 3 &J; 32°ro-5'N,64°40'7'W, 2291 m, 6.8.1966, 4??; 32°i94/N, 64°34-9'W-32°i9'N, 64°34-8'W,1135-1153 m, 18.8.1966, 97??, ig&J, 10 juv. ; 32°i5'8'N, 64°3i-6'W-3i°i6-i'Nf64°32-6'W, 2095-2223 m, 19.8.1966, 13 $$, 5 $$, 8 juv. DESCRIPTION. Immature male, length 3-5 mm : carapace (fig. iga, b) a littlemore than twice as long as high, not much elevated, with two rather irregular butwell-defined ridges on either side running backwards more or less parallel with thelower posterior edge ; the lower ridge extends back as far as the hind dorsal edge ofthe carapace ; there are a few blunt prominences dorsally and a larger rotundity oneither side of the frontal area. The pseudorostrum is about a seventh of the total 286 N. S. JONES FIG. 19. Campylaspis mansa subadult $. a, from side ; b, carapace and pereon fromabove ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropodand end of pleon. carapace length, with a well-excavated antennal notch below, bounded by an antero-lateral tooth, with some obtuse serrations behind. The eyelobe is rudimentary,fairly narrow. The pereon and pleon somites have some dorsal teeth and lateral serrations. Theanterior pereonites have dorsal elevations. Antenna I (fig. IQC) with the basal segment of the peduncle straight, about one-and-a-half as long as the third segment. Maxilliped 2 (fig. igd) with the ischiobasis broad, its terminal seta slender andnormally pinnate ; there is a tooth on the inner edge of the carpus and of the pro-podus, which has a long end spine ; the dactyl has three spines, the middle rathersmall. Maxilliped 3 (fig. ige) with the basis broad basally, about as long as the restof the appendage ; the merus is narrow and serrated on the lower edge ; the carpushas serrations on the upper edge and one terminal tooth on the lower edge while thepropodus is serrated along the lower edge. CAMPYLASPIS FROM THE DEEP ATLANTIC 287 Pereopod i (fig. 19!) with the basis a little longer than remaining segments to-gether ; the carpus is serrated along the upper edge. Pereopod 2 (fig. igg) with thebasis more than half as long as the rest of the appendage, rather broad ; the merusis serrated on the upper edge ; the dactyl is slender and tapering, as long as the carpusand propodus together, with three stout spines distally. Uropod (fig. igh) distinctly longer than the last three pleonites together, thepeduncle slender, serrated on both edges and more than twice as long as the endopod ;the exopod is about as long as the endopod. Adult male, length 5-0 mm : differs from the immature male in the usual charac-ters. The flagellum of antenna 2 is as long as the body with the uropods. TYPE LOCALITY. 38°46'N, 7O°o6'W, 2886 m. Types deposited in the BritishMuseum (Natural History). Registration numbers: Holotype 1974:317; Para-types 1974 : 318. REMARKS. C. mansa somewhat resembles C. undata Sars in the presence of twowell-defined ridges on either side of the carapace. C. undata, however, has a largeeyelobe with lenses and the dactyl of pereopod 2 is of quite a different shape. Numerous specimens occurred on the Gay Head - Bermuda transect between1135 and 2886 m. Campylaspis bicarinata sp. nov. (fig. 20) MATERIAL. 36°23'N, 67°58'W, 4680 m, 23.8.1964, 2 <?<£ ; 37°59-2'N, 79°26-2'W;3834m, 5-7-i965> 6?$, 2 <fc? ; o°o3'S, 27°48'W, 3730-37^3 m, 13.2.1967, i$,o°46'S, 29°28'W-o°46-5'S, 29°24'W, 3459m, 14.2.1967, 18 ??, u &?; 7°58'S,34°22'W, 834-939 m, 18.2.1967, 3<J<?; 7°58'S, 340i7'W-7°5o'S, 34°i7'W, 943-1007 m, 20.2.1967, 2 $?, i & i juv. DESCRIPTION. Adult female, length 4-2 mm : carapace (fig. 2oa, b) about twiceas long as high, not greatly elevated dorsally, with two well-defined ridges on eitherside running backwards more or less parallel with the lower edge, the lower ridgedefining the hind dorsal edge of the carapace and the upper ridge ending short of themid-line ; there is a small spine on the mid-dorsal line behind the eyelobe but noother prominences. The pseudorostrum is not upturned, less than a fifth of thetotal carapace length, with an obtuse antero-lateral angle below it and some serra-tions behind. The eyelobe is not very large, without lenses. The anterior pereonites are elevated dorsally. Antenna i (fig. 2od) with the segments of the peduncle fairly slender, the firstmoderately curved, about one-and-a-half as long as the third. Maxilliped 2 (fig. 2oe) with the ischiobasis long, its terminal seta fairly short butnot much thickened ; there is a tooth on the inner edge of the carpus and of thepropodus, of which the terminal process reaches beyond the spines of the dactyl ;there are three of these, the middle one very small. Maxilliped 3 (fig. 20!) with thebasis curved, longer than the rest of the appendage ; the remaining segments arefairly slender, the merus and carpus serrated on the upper edge. 17 288 N. S. JONES FIG. 20. Campylaspis bicarinata. a, $ from side ; b, ? carapace from above ; c, $ fromside ; d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ; h, $pereopod 2 ; i, $ uropod and end of pleon ; j, <J uropod and end of pleon. Pereopod i (fig. 2Og) with the basis curved, serrated on the distal lower edge,about as long as the rest of the appendage ; there is a spine on the upper distal endof the merus. Pereopod 2 (fig. 2oh) with the basis about two-thirds as long as theremaining segments together ; the dactyl is slender, little tapered, with some setaeand one short thick spine distally ; it is very little longer than the carpus and pro-podus together. Uropod (fig. 2oi) about as long as the last four pleonites combined, the peduncleserrated on its outer edge, more than twice as long as the endopod, which is distinctlylonger than the exopod ; the endopod has three thin spines on its inner edge and alonger end spine. Adult male, length 4-7 mm (fig. 2oc, j) : differs in the usual respects from thefemale. The flagellum of antenna 2 is probably as long as the body including uro-pods. A male, length 6-0 mm, from the Gay Head - Bermuda transect differs in CAMPYLASPIS FROM THE DEEP ATLANTIC 289 some minor respects from males from the Dakar - Recife transect, especially in thepresence of some small spines dorsally on the carapace and a low tubercle on eitherside of the frontal area ; it also has a number of blotches of red pigment on the cara-pace and pereon somites, especially concentrated along the lateral ridges. These,and some small differences in the appendages, however, do not seem to warrantseparation of the specimens from the two areas. TYPE LOCALITY. o°46'S, 29°28'W-o046-5'S, 29°24'W, 3459 m. Types depositedin the British Museum (Natural History). Registration numbers: Holotype1974 : 319 ; Paratypes 1974 : 320. REMARKS. C. bicarinata is fairly near to C. undata Sars but the latter species hasa well-developed eyelobe with lenses and there are various differences in the append-ages ; in particular the dactyl of pereopod 2 is much more slender and proportionatelylonger in C. bicarinata and the peculiar spine at its distal end is not present in C.undata but somewhat resembles the distal armature of the dactyl of pereopod 2 inC. plicata and C. bulbosa. A moderate number of specimens was obtained at several stations on the GayHead - Bermuda transect and in the western part of the Dakar - Recife transect,between 834 and 4680 m. Campylaspis rostrata Caiman, 1905 MATERIAL. 43035'6'N, 3°24'8'W, 860 m, 15.7.1967. i$; 43°43'N, 3°47'8'W,641 m, 19.7.1967, 4 $$ ; 46°i7-5'N, 4°45-2'W, 1336 m, 24.7.1967, i <j. DISTRIBUTION. Previously recorded from southwest of the Faroes, southwest ofIreland and the Mediterranean off Monaco, between 220 and 1205 m. The presentrecords from the Bay of Biscay extend its lower depth limit slightly to 1336 m.Caiman (1910) records it from the coast of Sudan but this must be a mistake. Thelatitude and longitude given place the position somewhere south of the CanaryIslands. Campylaspis intermedia Hansen, 1920 MATERIAL. 39°5o-5'N, 70°35'W, 823-5 m> 25.5.1961, i $ ; 39°54'5'N, 7O°35'W,466-7-508-7 m, 23.5.1962, i$; 39°46-5'N, 70°43'3'W, 1470-1330 m, 25.8.1964,36 $$, 12 c?o, 4 juv. ; 39°48-7'N, 70°40-8'W, 1102 m, 6.7.1965, 3 $$, 2 $$, 4 juv. ;39°54-i'N, 70°37'W, 478 m, 6.7.1965, 8 ?$ ; 39°5°-6'N, 7i°o3-6'W, 530 m, 5.5.1966,i ? ; 39°38-5'N, 70°36-5'W-39°39'N, 70°37-i'W, 2178 m, 18.12.1966, 2 $$, 4 juv. DISTRIBUTION. The only previous records are those of Hansen (1920) from theDavis Strait and south of Jan May en in 582 and 679 m. The present records extendits geographical range southwards in the N.W. Atlantic and its depth range tobetween 467 and 2178 m. It seems to be confined more or less to the continentalslope. 290 N. S. JONES Campylaspis submersa sp. nov. (fig. 21) MATERIAL. 8°02'S, 34°03'W-7°56'S, 34009'W, 1493 m, 19.2.1967, 2 ?$. DESCRIPTION. Adult female, length 3-4 mm : carapace (fig. 2ia, b, c) distinctlyless than twice as long as high, nearly oval in outline, with many rather large roundedprominences on the dorsal part ; on either side is a shallow indistinct sulcus. Thepseudorostrum is blunt, less than a sixth of the total carapace length, with a distinctantennal notch below and an acute antero-lateral angle. The eyelobe is large andbroad but without lenses. Antenna i (fig. 2id) with the first segment of the peduncle broad, slightly curved,nearly twice as long as the third segment. Maxilliped 2 (fig. 2ie) with the ischiobasis fairly long, its distal seta normallypinnate ; the distal spine of the propodus reaches a little beyond the spines of thedactyl, of which there are three, the middle small. Maxilliped 3 (fig. 2if) with thebasis curved, fairly slender and distinctly longer than the remaining segments to-gether ; the merus is broadened distally. Pereopod i (fig. 2ig) with the basis moderately curved, slender, a little longer thanthe rest of the appendage ; no spines or serrations are present. Pereopod 2 (fig. 2ih)with the basis fairly broad, about half as long as the rest of the appendage ; thedactyl is tapered to a point, with several slender spines at its end, and is shorterthan the carpus and propodus together. FIG. 21. Campylaspis submersa $. a, from side ; b, carapace from side ; c, carapacefrom above ; d, antenna i ; e, maxilliped 2 ; f, maxilliped 3 ; g, pereopod i ; h,pereopod 2 ; i, uropod and end of pleon. CAMPYLASPIS FROM THE DEEP ATLANTIC 291 Uropod (fig. 2ii) fairly broad, shorter than the last three pleon somites together ;the peduncle is serrated on the inner edge and only about a quarter longer than theendopod, which is a little longer than the exopod ; the endopod has some strongserrations and three slender spines on the inner edge, with one end spine. TYPE LOCALITY. 8°02'S, 34°03'W-7°56'S, 34°09'W, 1493 m. Types deposited inthe British Museum (Natural History). Registration numbers : Holotype 1974 :321 ; Paratype 1974 : 322. REMARKS. None of the described species of Campylaspis has precisely the arrange-ment of large rounded prominences on the carapace shown by this species, althoughthere are a number with fairly large tubercles, such as C. globosa Hansen. Thisarrangement and the short, rather broad uropods, as well as the relative proportionsof some of the appendages, should make recognition easy. Two specimens only were collected off Recife, Brazil, in 1493 m. Campylaspis paucinodosa sp. nov. (fig. 22) MATERIAL. 39°54-5'N, 7o°35'W, 466-7-508-7 m, 23.5.1962, i ?. DESCRIPTION. Subadult female, length 3-9 mm : carapace (fig. 22a, b) distinctlyless than twice as long as high, with a relatively small number of fairly large tuberclesconfined to the dorsal half and with a shallow excavation on either side. The FIG. 22. Campylaspis paucinodosa immature $. a, from side ; b, carapace from above ;c, antenna i ; d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h,uropod and end of pleon. 292 N. S. JONES pseudorostrum is inclined slightly upwards and fairly acutely pointed from the side,truncated from above, with a well-defined antennal notch below and a distinct butrounded antero-lateral angle. The eyelobe is large and prominent and has visiblelenses. The fourth and fifth pleonites are serrated dorsally. Antenna i (fig. 22c) with the first segment of the peduncle broad basally, slightlycurved, about twice as long as the third. Maxilliped 2 (fig. 22d) with the basis fairly broad, its distal seta normally pinnate ;the terminal spine on the propodus is fairly long and slender ; the dactyl has two endspines of equal length. Maxilliped 3 (fig. 22e) with the basis strongly curved, longerthan the rest of the appendage ; the merus is fairly broad but none of the segmentshave spines or serrations. Pereopod i (fig. 221) with the basis a little longer than the remaining segmentstogether, with some serrations on the distal lower edge. Pereopod 2 (fig. 22g) withthe basis about two-thirds as long as the rest of the appendage ; the dactyl is short,scarcely tapering, much shorter than the carpus. Uropod (fig. 22h) short, about as long as the last two pleonites together ; thepeduncle is serrated on either edge, about one-and-a-half as long as the endopod,which is scarcely longer than the exopod ; the endopod has some serrations and threefairly stout spines on its inner edge and two unequal end spines. TYPE LOCALITY. 39°54'5'N, 7O°35'W, 467-509 m. Type deposited in the BritishMuseum (Natural History). Registration number : Holotype 1974 : 323. REMARKS. C. paucinodosa shows some resemblance to C. horrida Sars but it hasa smaller number of tubercles and is without dorso-lateral projections on thepereonites and anterior pleonites ; the uropods are also much shorter and the rela-tive proportion of the peduncle to the endopod much smaller than in C. horrida. Caiman (1912) doubtfully referred a dried and broken specimen from 428 fathomsin nearly the same locality as that for C. paucinodosa to C. horrida. It is likely thatit was C. paucinodosa, and not C. clavata, as suggested by Lomakina (1958). A single specimen was obtained south of Gay Head in 467-509 m. Campylaspis horridoides Stephensen, 1915(fig. 23) MATERIAL. 43°35-6'N, 3°24-8'W, 860 m, 15.7.1967, 3 ??, i <$, i juv. ; 38°27'N,4°o8'E, 2447 m, June 1970, i $. DISTRIBUTION. Previously recorded only from the Mediterranean south ofSardinia in 1227 m this species is now recorded also from the Bay of Biscay and itsdepth range between 860 and 2447 m. REMARKS. Although Stephensen's (1915) figures are adequate there is onecharacter which he omits to mention and which may be useful for identifying thespecies. This is the rather broad suture visible from above along the mid-dorsalline (fig. 23) which differs in appearance from closely related species such as CAMPYLASPIS FROM THE DEEP ATLANTIC 293 FIG. 23. Campylaspis horridoides $. Carapace and pereon from above. C. squamifera Fage. Dr Torben Wolff of Universitetets Zoologiske Museum, K0ben-havn, has kindly allowed me to examine Stephensen's original specimens which Ihave checked for this character and from which I have selected lectotypes. Campylaspis squamifera Fage, 1929 MATERIAL. 43°35'6'N, 3°24-8'W, 860 m, 15.7.1967, 5 ??, i <?, 3 juv.; 43°43'N,3°47-8'W, 641 m, 19.7.1967, 3 $$, 5 &£, 19 juv. DISTRIBUTION. Known previously from a single specimen dredged in the southof the Bay of Biscay from 4380 m, those recorded here are from considerably lesserdepths in the more northerly part of the Bay. Campylaspis glebulosa sp. nov.(fig- 24) MATERIAL. o°O3'S, 27°48'W, 3730-3783 m, 13.2.1967, i <$ ; o°46'S, 29°28'W-o°46-5'S, 29°24'W, 3459m, 14.2.1967, i$; 7°58'S, 340i7'W-7°5o'S, 34°i7'W,943-1007 m, 20.2.1967, i $ ; 8°03'S, 34°23'W-8°O2'S, 34°25'W, 587 m, 21.2.1967, i $. DESCRIPTION. Adult female, length 4-5 mm : carapace (fig. 24a, b) a little morethan one-and-a-half as long as high, well elevated, with fairly numerous tubercles,most of which are serrated, and an ill-defined sulcus on either side ; there is a shorttransverse ridge across the middle of the frontal area. The pseudorostrum is lessthan a sixth of the total carapace length, bluntly pointed from the side, with a slightlyexcavated antennal notch below and a slightly projecting antero-lateral angle. Theeyelobe is fairly large but without lenses. The pereon and anterior pleon somites have serrated low dorsal projections. Antenna i (fig. 24d) with the basal segment of the peduncle curved, nearly threetimes as long as the third. Maxilliped 2 (fig. 246) with the ischiobasis long, its terminal seta fairly short andpinnate distally but not thickened ; terminal propodal spine about as long as thespines of the dactyl, of which there are three, the middle very small. Maxilliped 3(fig. 24f) with thebasiscurved, distinctly longer than the rest of the appendage ; thesegments are not broad, the merus and carpus with some strong spines on the upperedge. 294 N. S. JONES a d-i FIG. 24. Campylaspis glebulosa. a, $ from side ; b, $ carapace from above ; c, immaturecJ from side ; d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ; h,$ pereopod 2 ; i, $ uropod and end of pleon. Pereopod I (fig. 24g) with the basis moderately curved, a little longer than therest of the appendage. Pereopod 2 (fig. 2411) with the basis about two-thirds aslong as the rest of the appendage ; the dactyl is tapered but not pointed, withlong slender terminal spines, and is almost as long as the carpus and propodustogether. Uropod (fig. 241) about as long as the last two pleonites together ; the peduncle isserrated on either edge and more than one-and-a-half as long as the endopod, whichis equal in length to the exopod ; the endopod has some serrations on either edgeand three fairly slender spines on the inner edge, with two unequal end spines. TYPE LOCALITY. o°46'S, 29°28'W-o°46-5'S, 29°24'W, 3459 m. Types depositedin the British Museum (Natural History). Registration numbers: Holotype1974 : 324 ; Paratype 1974 : 325. CAMPYLASPIS FROM THE DEEP ATLANTIC 295 REMARKS. C. glebulosa quite closely resembles C. squamifera, but has fewertubercles, especially on the sides of the carapace, and lacks the dorso-lateral processesand strong serrations present on the pereon and pleon of the latter species. Thereare also many differences in the appendages, especially the comparative lack ofteeth or serrations in C. glebulosa and the much longer basis of pereopod 2 comparedwith the rest of the appendage. From C. globosa Hansen this species may be dis-tinguished, among other characters, by the comparative shortness of the uropods,which in C. globosa are as long as the last three pleonites. A few specimens were obtained at several stations in the western part of theDakar - Recife transect, in 587-3783 m. Campylaspis torulosa sp. nov. (fig. 25) MATERIAL. 36°23'N, 67°58'W, 4680 m, 23.8.1964, 1899, 4^; 36°24'4'N,67°56'W, 4749 m, 4.7.1965, 3 $$, 6 <$$, 2 juv. ; 36°2o'N, 67°56'W, 4694 m, 13.12.1965,i $ ; 37°24'N, 65°54'W-37026'N, 6505o'W, 4825 m, 23.8.1966, i $, 4 33, 3 juv. DESCRIPTION. Adult female, length 5-2 mm : carapace (fig. 25a, b) about twiceas long as high, evenly rounded dorsally, somewhat produced backwards over theanterior pereonites, with many fairly large tubercles or blunt spines set in irregularrows, leaving a fairly small area free from tubercles on either side ; the frontal areais crossed by two poorly defined transverse ridges. The pseudorostrum is shortand obtuse, less than a seventh of the total carapace length, with a trace of an antennalnotch below, defined posteriorly by two blunt teeth. The eyelobe is of moderatesize, without lenses. The pereon and the first five pleon somites have blunt lateral projections andthere are some small tubercles on the dorsum of the fifth pleonite. Antenna i (fig. 25c) with the segments of the peduncle fairly stout, the first curvedand nearly three times as long as the third. Maxilliped 2 (fig. 25d) with the ischiobasis broad and truncated at the base ;the carpus has a small tooth on the distal inner edge ; the propodal process reachesbeyond the spines of the dactyl, of which there are two of approximately equallength. Maxilliped 3 (fig. 256) with the basis much longer than rest of appendage ;the ischium has some teeth on the lower edge and there are a number of teeth on thelower edge of the merus, which also has a sharp tooth on the upper edge distally ;there are several sharp teeth on the upper edge of the carpus ; the ischium and merusare fairly broad. Pereopod i (fig. 25f) with the basis distinctly longer than rest of appendage,serrated distally on both edges. Pereopod 2 (fig. 25g) with the basis broad, less thanthree-quarters as long as the remaining segments together, serrated distally on eitheredge ; the dactyl is little tapered, with a blunt end, about as long as the carpus andpropodus together. Uropod (fig. 25h) broad, a little shorter than the last three pleonites together, thepeduncle serrated on either edge and with some short setae on the inner edge, onlyabout one-and-a-quarter as long as the endopod, which is a little longer than the 296 N. S. JONES C-h FIG. 25. Campylaspis torulosa $. a, from above ; b, carapace from side ; c, antenna i ;d, maxilliped 2 ; e, maxilliped 3 ; f, pereopod i ; g, pereopod 2 ; h, uropod and end ofpleon. exopod ; the endopod has three or four spines on the inner edge and two unequalend spines. TYPE LOCALITY. 37°24'N, 65°54'W-37°26'N 65°5o'W, 4825 m. Types depositedin the British Museum (Natural History). Registration numbers: Holotype1974 : 326 ; Paratypes 1974 : 327. REMARKS. C. torulosa seems to be closely related to C. papillata Lomakina butthe arrangement of the tubercles on the carapace is somewhat different, while in C.papillata the dactyl of pereopod 2 is more tapered and the endopod of the uropod isnarrower and has a greater number of spines on its inner edge. It also resembles C.squamifera but has many more tubercles on the dorsum of the carapace than thatspecies. The species was obtained on several occasions on the deeper parts of the Gay Head -Bermuda transect from the abyssal plain, between 4680 and 4825 m. CAMPYLASPIS FROM THE DEEP ATLANTIC 297 Campylaspis verrucosa G. 0. Sars, 1866 MATERIAL. 43°4O'8'N, 3°35'2'W, 1739 m, 16.7.1967, 3 &? ; 43°43'N, 3°47'8'W,641 m, 19.7.1967, 4 $$, i <?<?. DISTRIBUTION. Recorded from the N.E. Atlantic from the Lofotens to the coastof Portugal and from the Mediterranean, between 100 and noo m. The presentrecords from the Bay of Biscay increase the lower limit of known occurrence to 1739 m. Campylaspis globosa Hansen, 1920 MATERIAL. 39°26'N, 7O°33'W, 2496m, 21.8.1964, 20??; 38°46'N, 70°o6'W,2886m, 21.8.1964, i$; 39°55'2'N, 7O°39'5'W, 498m, 27.4.1966, H ??, 6<?<J;39°56-6'N, 7i°o3-6'W, 530 m, 5.5.1966, i ?. DISTRIBUTION. Previously recorded only from the Davis Strait in 2626 m.As with C. intermedia, which has much the same geographical range, its range isextended to the south and its upper depth limit to 498 m. It appears to extend togreater depths than C. intermedia. Campylaspis multinodosa sp. nov.(fig. 26) MATERIAL. 23°oo'S, i2°45'E, 1007-1014 m, 16.5.1968, 12$$, i $; 23°05'S,i2°3i-5'E, 1546-1559 m, 17.5.1968, i ?, 2 juv. ; 23°02'S, I2°I9'E, 2117-2154 m, 17-5-1968, I ?, 2 &?. DESCRIPTION. Adult female, length 3-9 mm : carapace (fig. 26a, b) twice aslong as high, with many small tubercles dorsally and on the sides, some of which areacute, especially on the pseudorostrum and just below it; on either side is a fairlysmall shallow excavation. The pseudorostrum is more than a sixth of the totalcarapace length, truncate in front, with a well-defined antennal notch and antero-lateral angle below it. The eyelobe is fairly large but without lenses. The last pereonite and the first five pleonites have paired dorso-lateral teeth. Antenna i (fig. 26d) with the segments of the peduncle slender, the basal segmentslightly curved and less than one-and-a-half as long as the third. Maxilliped 2 (fig. 26e) with the ischiobasis fairly long, serrated on its distal outeredge, with its distal seta normally pinnate ; the carpus has a few serrations on theouter edge and the propodus a tooth on the inner edge and a slender end spine ; thedactyl has two spines. Maxilliped 3 (fig. 261) with the basis slender, serrated oneither edge distally, much longer than the remaining segments together ; these arenot widened and the merus, carpus and propodus are serrated on the lower edge ;the dactyl is small. Pereopod i (fig. 26g) with the basis about as long as the rest of the appendage,serrated on its distal lower edge ; there are one or two serrations on the lower edgeof the ischium. Pereopod 2 (fig. 26h) with the basis little more than half as long asthe rest of the appendage, serrated along its distal upper edge ; the merus is serrateddistally on both edges and the carpus proximally on the upper edge ; the dactyl is FIG. 26. Campylaspis multinodosa. a, $ from side ; b, $ from above ; c, $ from side ;d, $ antenna i ; e, $ maxilliped 2 ; f, $ maxilliped 3 ; g, $ pereopod i ; h, $ pereopod 2 ;i, $ uropod and end of pleon ; j, $ uropod and end of pleon. not tapered and has several fairly long pinnate end spines ; it is a little shorter thanthe carpus and propodus together. Uropod (fig. 26i) rather longer than the last two pleon somites together, thepeduncle serrated on either edge and about one-and-three-quarters as long as theendopod, which is a little longer than the exopod ; the endopod has some stoutserrations and three spines on the inner edge, with two unequal end spines. Adult male, length 5-3 mm (fig. 26c, j) : differs from the female in the usualcharacters. The sides of the carapace are smoother and there are no dorso-lateralteeth on the pleon somites. The second antennal flagellum reaches to the end of the CAMPYLASPIS FROM THE DEEP ATLANTIC 299 uropods and these are more slender and have more setae and spines but are littleserrated. TYPE LOCALITY. 23°oo'S, i2°45'E, 1007-1014 m. Types deposited in the BritishMuseum (Natural History). Registration numbers : Holotype 1974 : 328 ; Para-types 1974 : 329. REMARKS. C. multinodosa shows a considerable resemblance to C. antarcticaCaiman. The latter has several teeth on the outer side of the carpus of maxilliped 3and the dactjd of pereopod 2 is longer than the carpus and propodus together, butotherwise it is difficult to separate them and the two are very closely related andperhaps identical. A moderate number of specimens were captured at several stations off S.W. Africain 1007-2154 m. ACKNOWLEDGEMENTS I am much indebted to Dr Howard L. Sanders for the material from WHOIcruises and the collections from the Bay of Biscay and the Canaries and to Dr DanielReyss for the specimens collected by the 'Jean Charcot'. REFERENCES BACESCU, M. & MURADIAN, Z. 1972. Nouvelles especes de Nannastacidae (Crustaces, Cumaces)dans les eaux sahariennes de 1'Atlantique. Rev. Trav. Inst. Peches marit. 36 (3) : 255-269. BONNIER, J. 1896. Resultats scientifiques de la campagne du 'Caudan' dans le Golfe deGascogne : Cumacea. Ann. Univ. Lyon, 24 (1895) : 528-562. CALMAN, W. T. 1905. Cumacea. Scient. Invest. Fish. Brch Ire. 1904, No. i : 52 pp. 1906. The Cumacea of the Puritan Expedition. Mitt. zool. Stn. Neapel, 17 : 411-432. 1907. Cumacea. National Antarctic Exped., Natural History, 2 (6) : 6 pp. 1910. Les cumaces des expeditions du Travailleur et du Talisman. Bull. Mus. Hist. nat. Paris, 1910: 180-182. 1911. On new or rare Crustacea of the Order Cumacea from the collection of the Copen-hagen Museum. Pt 2. The Families Nannastacidae and Diastylidae. Trans, zool. Soc.Land. 18 : 341-398. 1912. The Crustacea of the Order Cumacea in the collection of the United States National Museum. Proc. U.S. nat. Mus. 41 : 603-676. PAGE, L. 1929. Cumaces et Leptostraces provenant des campagnes scientifiques du PrinceAlbert ler de Monaco. Result. Camp, scient. Prince Albert I, 77 : 1-50. 1945. Les cumaces du plancton nocturne des cotes d'Annam. Arch. Zool. exp. gin. 84 :165-224. — 1951. Cumaces. Faune Fr. 54 : 136 pp. GAMO, S. 1960. Six new species of cumacean Crustacea Campylaspis (Nannastacidae) fromSagami Bay. Zool. Mag., Tokyo, 69 : 369-387. 1967. Studies on the Cumacea (Crustacea, Malacostraca) of Japan. Part II. Publs Seto mar. biol. Lab. 15 (4) : 245-274. GURWITSCH, G. 1939. The cumacean fauna of the White Sea. Trudy gos. gidrobiol. Inst. 8 : 75-80 (in Russian).HALE, H. M. 1937. Cumacea and Nebaliacea. Rep. B.A.N.Z. antarct. Res. Exped. 1929-31, 4b (2) : 37-56. 1945. Australian Cumacea. No. 9. The Family Nannastacidae. Rec. S. Aust. Mus. 8 : 145-218. 300 N. S. JONES HART, J. F. L. 1930. Some Cumacea of the Vancouver Island region. Contr. Can. Biol. N.S. 6(3):i-i8.HANSEN, H. J. 1908. Schizopoda and Cumacea. Re's. Voyage Belgica, Zool. : 1-21. 1920. Crustacea Malacostraca 4. Dan. Ingolf-Exped. 3 (6) : 86 pp. JONES, N. S. 1969. The systematics and distribution of Cumacea from depths exceeding 200 metres. Galathea Rep. 10 : 99-180. — 1973. Some new Cumacea from deep water in the Atlantic. Crustaceana, 25 (3) : 297-319.JONES, N. S. & SANDERS, H. L. 1972. Distribution of Cumacea in the deep Atlantic. Deep-Sea Res. 19 : 737-745.LE LCEUFF, P. & INTES, A. 1972. Les cumaces du plateau continental de Cote d'lvoire. Cah. O.R.S.T.O.M., sir. Octanogr. 10 (i) : 19-46.LIE, U. 1969. Cumacea from Puget Sound and off the northwestern coast of Washington, with descriptions of two new species. Crustaceana, 17 (i) : 19-30.LILLJEBORG, W. 1855. Om Hafs-Crustaceer vid Kullaberg i Skaane. Ofv. Ak. Fork. 12 : 117-138.LOMAKINA, N. B. 1952. New species of Cumacea of Far-East Seas. Trudy zool. Inst., Leningr. 12 : 115-170 (in Russian). 1955. Cumacea of Far-East Seas. Trudy zool. Inst., Leningr. 18 : 112-165 (m Russian). 1958. Kumovie Raki (Cumacea) Morei SSSR. Opred. Faune SSSR, 66 : 301 pp. (in Russian). 1968. Cumacea of the Antarctic Region. Issled. Faun. Morei, 6 (14) : 97-140 (in Russian).REYSS, D. 1972. Resultats scientifiques de la campagne du N.O. 'Jean Charcot' en Mediter- ran6e Occidentale, Mai-Juin-Juillet 1970. Crustaceana, suppl. 3 : 362-377.- 1973. Distribution of Cumacea in the deep Mediterranean. Deep-Sea Res. 20 (12) : 1119- 1123. 1974- Contribution a I'e'tude des Cumaces de profondeur de 1'Atlantique Nord : le genre Makrokylindrus Stebbing. Crustaceana, 26 (i) : 5-28.SANDERS, H. L. & HESSLER, R. R. 1969. Ecology of the deep sea benthos. Science, N.Y. 163 :1419-1424.SARS, G. O. 1865. Om den aberrante krebsdyrgruppe Cumacea og den nordliske Arter. Fork. Videns.-Selsk. Christiania, 1864 : 83 pp. 1870. Nye Dybvandscrustaceer fra Lofoten. Fork. Videns.-Selsk. Christiania, 1869 : 147-174. 1871. Beskrivelse af de paa fregatten Josephines expedition fundne Cumaceer. Sv. Vet. Ak. Handl. 9 (13) : 1-57. 1879. Nye bidrag till kendskapen om Middelhavets invertebratfauna. 2. Middelhavets Cumaceer. Arch. Math. Naturv. 3-4 : 1-196. 1887. Report on the Cumacea. Rep. sci. Res. 'Challenger', Zool. 19 : 1-73. STEBBING, T. R. R. 1912. The Sympoda. Ann. S. Afr. Mus. 10 : 129-176. STEPHENSEN, K. 1915. Isopoda, Tanaidacea, Cumacea, Amphipoda (excl. Hyperiidea), Rep. Dan. Oceanogr. Exped. 1908-1910, 2 : 29-34.ZIMMER, C. 1907. Neue Cumaceen von der deutschen und der schwedischen Siidpolarexpedi- tion aus der Familien der Cumiden, Vauntompsoniiden, Nannastaciden und Lampropiden. Zool. Anz. 31 : 367-374. 1936. Californian Crustacea of the Order Cumacea. Proc. U.S. natn Mus. 83 : 423-429. DR N. S. JONES Department of Marine Biology THE UNIVERSITY OF LIVERPOOL PORT ERIN ISLE OF MAN A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.1965. (Out of Print.) £3.75. 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogyof the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates,77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa: theBiology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.1974. £3-75. Hardback edition £6. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU - 3 JAN A NEW FAMILY, GENUS AND SPECIES OF BAT (MAMMALIA : CHIROPTERA) FROM THAILAND J. E. HILL BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 27 No. 7 LONDON: 1974 -3 A NEW FAMILY, GENUS AND SPECIES OF BAT (MAMMALIA : CHIROPTERA) FROM THAILAND BY JOHN EDWARDS HILL Pp 301-336; 8 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 7 LONDON : 1974 V ° i J THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted, in 1949, isissued in five series corresponding to the Departmentsof the Museum, and an Historical series. Parts will appear at irregular intervals as theybecome ready. Volumes will contain about three orfour hundred pages, and will not necessarily becompleted within one calendar year. In 1965 a separate supplementary series of longerpapers was instituted, numbered serially for eachDepartment. This paper is Vol. 27, No. 7 of the Zoologicalseries. The abbreviated titles of periodicals citedfollow those of the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 24 December, 1974 Price £2.10 A NEW FAMILY, GENUS AND SPECIES OF BAT (MAMMALIA : CHIROPTERA) FROM THAILAND By J. E. HILL INTRODUCTION WHILE Curator of Terrestrial Vertebrates at the Centre for Thai National ReferenceCollections, Bangkok, the late Kitti Thonglongya became especially interested in thebats of Thailand. His enthusiasm has led in the past few years to the discovery oftwo new species (Rhinolophus marshalli Thonglongya, 1973, Hipposideros lekaguliThonglongya & Hill, 1974) in Thailand, and has added other species (Rhinolophusparadoxolophus, Myotis annectans, Pipistrellus cadornae) to the Thai fauna (Thong-longya, 1973 ; Hill & Thonglongya, 1972). He was also responsible for the discoveryin the collections of the Bombay Natural History Society of an undescribed genusand species of Indian fruit bat (Latidens salimalii Thonglongya, 1972). In thelatter part of 1973 he obtained specimens in southern Thailand which he thoughtmight represent an unknown genus and species justifying perhaps the establishmentof a further higher category within the Microchiroptera. Specimens sent to theBritish Museum (Natural History), London, fully confirmed his initial opinion, andshowed the new genus to differ sufficiently from the established categories of theMicrochiroptera to warrant the proposal of a further family. Originally, the intention had been to describe this remarkable new bat in a jointpaper. With this in view, a detailed comparative study was begun in London toestablish and evaluate its diagnostic features, many already recognized by KittiThonglongya during his preliminary examination in Thailand. As a result of hissudden and untimely death in February 1974, the Applied Scientific ResearchCorporation of Thailand, through Dr Prasert Lohavanijaya, the Director of theCentre for Thai National Reference Collections, has entrusted me with the prepara-tion of the account. In these circumstances I deem it an honour and a privilege torecognize the contribution made by my former friend and colleague by associatinghis name with the most outstanding and significant discovery that he made in hisstudies of the bats of his country. DIAGNOSES AND DESCRIPTION CRASEONYGTERIDAE fam. nov. DIAGNOSIS. Similar in some respects to the Rhinopomatidae and to the Embal-lonuridae but second finger with one very short bony phalange, usually ankylosedto the metacarpal and with an equally short cartilaginous tip. As in these familiesthe premaxillae not fused to surrounding or adjacent parts. However, they areunited anteriorly to enclose a large anterior palatal vacuity and posteriorly to 3o4 J. E. HILL surround the narial aperture ; palatal branches short, solidly ankylosed throughouttheir length to form a posteriorly projecting, tapered spine extending across about onehalf or less of the anterior palatal vacuity which it thus divides into two posteriorlyconfluent foramina ; narial branches extending upwards to lie posteriorly on thesurface of the maxillae and nasals as a thin, laminar, posteriorly rectangular plate,with no trace of a median suture, the inner margin of this structure forming a narrow,sub-tubular, anteriorly directed flange which borders the lateral and upper parts ofthe narial opening. Humerus with trochiter or greater tuberosity as large as trochinor lesser tuberosity, trochiter extending proximally beyond the head of the humerus ;a shallow supraglenoid recess between the proximal base of the head of the humerusand the base of the trochiter ; anterior face of proximal part of shaft of humerusflattened in its central part, lacking any median deltoid ridge, instead a slightlyelevated area at the base of the trochiter, extending distally along the proximal partof the shaft as a short dorsal flange supporting the trochiter, with a similar but shorterventral flange supporting the trochin ; capitellum slightly displaced from line ofshaft; epitrochlea or medial process a little less than half as wide as distal articularsurface, with small epitrochlear or distal spinous process. Sternum not especiallymodified ; shoulder girdle normal; last cervical vertebra not fused with firstthoracic ; lower three thoracic vertebrae fused, their boundaries faintly visible ;lumbar vertebrae except last two solidly ankylosed, the last two free ; sacral verte-brae fused, their boundaries obliterated or nearly so. Pelvis small, weak, ascendingramus of pubis short, without definite acicular or pubic spine, the ventral ramusdelicate, very thin ; ischium with ascending ramus similarly thin and delicate,these structures easily lost in preparation ; fibula thread-like, extending about half-way along the tibia. CRASEONYCTERIS gen. nov. DIAGNOSIS. Most nearly resembling Rhinopoma, the sole genus of the Rhino-pomatidae, but differing as above and further in normal, slightly crescentic nostrilswhich are not slit-like or valvular although as in Rhinopoma they open directlyanteriorly in the face of a thickened, vertical narial pad or plate ; ridge-like dermalprotrusion surmounting narial plate lower and narrower ; ears not joined anteriorlyby a band of integument; tragus narrowed and rounded at the tip, not truncate,and with an oblate swelling in the centre of its anterior margin. Distal phalange ofthird digit very long, reflexed beneath wing in specimens preserved in alcohol;distal phalange of fourth digit similarly reflexed ; no external tail, the uropatagiumnot shortened. Braincase more inflated and more globular than in Rhinopoma ;postorbital region less markedly constricted; lateral rostral swellings less inflated,not projecting anteriorly to surpass the narial aperture which slopes posteriorly andis not nearly vertical as in Rhinopoma ; zygomata less expanded ; coronoid processlow, below height of articular process. Dentition similar to that of Rhinopoma, thedental formula i \, c T, pm i, m f = 28 as in that genus, but upper incisors relativelymuch larger and third upper molar a little more reduced. TYPE SPECIES. Craseonycteris thonglongyai sp. nov. A NEW BAT FROM THAILAND 305 REMARKS. The name of the new genus is derived from Kpavis or Kpdaews, amixing or blending, with wKrepls, a bat, in allusion to the combination of featurespresented by this notable discovery. Since the genus is proposed to include onlythe type species, one description may serve for both. Craseonycteris thonglongyai sp. nov. HOLOTYPE. Thai National Reference Collections No. 54-3871. Adult ^. Cavenear the Forestry Station, Ban Sai Yoke, Kanchanaburi, Thailand, I4°26' N,98°5i' E. Collected 8 December 1973 by Kitti Thonglongya. Original numberKT 5710. In alcohol, skull extracted. OTHER MATERIAL. Thai National Reference Collections (in alcohol unless stated) :(3$ Nos. 54-3213 (skull, left humerus, left femur extracted, skull not seen), 26 October1973 ; 54-3865 (disarticulated skeleton), 54-3866-3870 (skulls extracted), 54-3872-3873, 54-3874-3875 (skuUs extracted), 54-3876, 54-3884 (skull extracted),54-3886, 54-3887-3888 (skulls extracted), 54-3889, 8 December 1973; $$ Nos.54-3230 (skull extracted, right fibula exposed), 28 October 1973 ; 54-3877, 54-3878-3881 (skulls extracted), 54-3882-3883, 54-3885, 8 December 1973. Collected byKitti Thonglongya at or near the type locality. This account is based on the specimens listed. A total of fifty-two numbers isallocated to specimens of this bat in the field catalogue maintained by Kitti Thong-longya : those not examined in the preparation of this study have remained in theThai National Reference Collections and include skins, skulls, skeletons and speci-mens preserved in alcohol (J. T. Marshall, in litt.). The entire representation camefrom one or other of two caves near the Forestry Station, Ban Sai Yoke, probablyabout 2 km along the Kwae Noi River past its junction with the Huay Mae Nam NoiRiver (J. T. Marshall, in litt.). Specimens collected in October 1973 came apparentlyfrom Tham Wang Phra', those in December from 'Namtok Sai Yoke'. The finaldisposition of the series examined at the British Museum (Natural History) has yetto be decided but it is expected that the specimens will remain in London for anindefinite period, some permanently. DESCRIPTION. A very small bat (length of forearm 22-26 mm), dorsally greyishbrown, ventrally rather greyer, some specimens with a slight reddish tinge dorsally(from alcohol), membranes dark. Muzzle (Fig. i) rather suiform, slightly swollenlaterally; anterior part of chin similarly swollen, the swelling faintly dividedmedianly; anteriorly muzzle with a thickened, fleshy vertical narial pad or platesurmounted by a low dermal ridge extending across the central part of the narialpad ; nostrils wide, slightly crescentic, opening directly anteriorly in the face of thenarial pad, sharply inclined to the horizontal, separated by a relatively wide inter-narial septum which broadens above the nostrils to form the base of the low dermalridge. Shallow grooves extend upwards from the upper corner of each nostril toseparate the dermal ridge from the lateral elements of the narial pad ; shallowdepressions extend downwards from the lateral margins of the internarial septumtowards the upper lip ; central part of lower lip raised by a smooth, naked swelling 306 J. E. HILL FIG. i. Craseonycteris thonglongyai. Head. immediately anterior to and extending across the lower incisors. Narial pad andmedian anterior part of lower lip naked; lateral swellings of muzzle and of chinsparsely covered with short, rather stiff hairs, underside of chin and throat onlylightly furred. Eyes small, largely concealed in fur ; ears (Fig. i) very large and rather mem-branaceous, reaching beyond the tip of the muzzle when laid forward. Ear roundedat the tip, anterior margin originating just above eye, without basal lobe, convex,posterior margin moderately concave just beneath tip, then straight, proximal halfstrongly convex, inserted quadrately just behind angle of mouth, antitragus poorlydeveloped, little more than a thickening of the ear membrane. Outer surface ofconch with a few sparse hairs in its proximal half, inner surface with a diffuse bandof sparse hairs posterior and parallel to the proximal part of its anterior margin, afew sparse hairs in its distal part. Tragus (Fig. i) a little less than one half the length of the ear, basally narrowbeneath a small, more or less triangular posterior basal lobe, then broadly expanded,its greatest width at about one half its length, narrowing abruptly to a rounded tip.Anterior part of tragus with distinct oblate thickening about halfway along itslength, the thickened area extending across one half of the width of the tragus atthis point. Tragus with anterior margin slightly concave proximally, then at theoblate thickening slightly convex beneath a shallow concavity, distal part of marginslightly convex to tip ; posterior margin slightly concave distally, then convex andserrated at the widest part of the tragus, curving abruptly, almost angularly to ashallow rounded emargination just above the posterior basal lobe. Propatagium or antebrachium broad, originating proximally at a point level withthe shoulder joint, extending distally to the distal end of the first metacarpal. A NEW BAT FROM THAILAND 307 Thumb short, with well-developed claw ; membrane between thumb and secondmetacarpal broad proximally, anchored to the end of the first metacarpal, taperingto a distal termination at a point about halfway along the second metacarpal.Second digit free anteriorly in its distal part, with long metacarpal and one veryshort bony phalange, usually fused to the metacarpal, the junction visible underrelatively high magnification, often marked by a slight broadening of the metacarpal,the digit tipped by a tapered cartilage which is approximately equal in length to theshort bony phalange. Membrane lying between second and third digits terminatingdistally about one quarter or a little more along the length of the second phalangeof the third digit from its junction with the first phalange. Third digit with twophalanges, no trace of a third, its metacarpal shorter than the metacarpal of thesecond digit, the second phalange very long, about three times as long as the firstphalange and approximately equal in length to the third metacarpal, stronglyreflexed beneath wing in specimens preserved in alcohol. Metacarpal of fourthdigit a little longer than that of third digit but shorter than second metacarpal, itsfirst phalange very short, in length about one sixth that of the associated metacarpal,second phalange long, about three times the length of the first phalange, slightlyreflexed beneath wing. Metacarpal of fifth digit about as long as fourth metacarpal, 2 mm FIG. 2. Craseonycteris thonglongyai. Holotype. Lateral aspect of skull. 308 J. E. HILL its first phalange short, about one quarter the length of its metacarpal, the secondphalange slightly exceeding the first in length. No external tail, uropatagium moderate, narrow proximally between upper partof legs to knees, wider distally, its posterior margin slightly curved, terminatingmedianly at a line a little beyond halfway from knees to ankles, calcar absent,membranes inserted 3-4 mm above the ankle ; a few short hairs on the ventralsurface of the endopatagium, clustered at and about the distal end of the humerus,fur not extending on to the uropatagium. Foot long, narrow and slender, thephalangeal formula 2-3-3-3-3, the toes subequal in length, with a few long hairson the dorsal surface. A rounded glandular swelling at base of underside of throat,well developed and prominent in males, much less so or absent in females. Penisrelatively massive, short and broad, moderately covered with short hairs, preputial 2 mm FIG. 3. Craseonycteris thonglongyai. Holotype. Dorsal aspect of skull. A NEW BAT FROM THAILAND 3°9 2 mm FIG. 4. Craseonycteris thonglongyai. Holotype. Ventral aspect of skull. opening small, rounded ; vulva transverse, set in a fleshy elevation. One pairpectoral and one pair pubic nipples, the latter closely set and situated just anteriorto the genital eminence. Skull (Figs. 2-5) very small (condylobasal length 9-5-10-1 mm) with slightlyinflated, globose braincase ; no lambdoid crests ; prominent sagittal crest extendingfrom rear of braincase to frontal region immediately above narrowest postorbitalpoint, its crest slightly higher frontally than posteriorly; postorbital region notespecially constricted. Rostrum wide, laterally expanded ; no postorbital processesor evident supraorbital ridges ; combined nasals wider than long ; lateral rostralswellings well developed and prominent, not extending laterally to project beyond thenarial aperture at its sides, swellings separated dorsally by a narrow median rostraldepression, deeper posteriorly than anteriorly ; narial opening sloping posteriorly, 310 J. E. HILL 2 mm FIG. 5. Craseonycteris thonglongyai. Holotype. Dorsal aspect of mandible. flanked by the lateral rostral swellings. Premaxillae not fused to adjoining partsbut completely united ; palatal branches small, totally co-ossified with no trace of asuture, anteriorly a U-shaped emargination between the incisors ; posteriorly thefused palatal branches form a narrow, tapered spike extending across about one halfor a little less of the large anterior palatal foramen ; narial branches exceptionallydeveloped to extend upward to the apex of the narial aperture where they are totallyfused, forming a thin lamina or plate, rectangular posteriorly, lying on the maxillaeand nasals, its inner edge produced anteriorly to form a thin, sub-tubular flangearound the lateral and upper parts of the narial opening ; entire premaxillary struc-ture firmly connected by tough tissue to the rear of the narial pad which apparentlyit serves to support. Anteorbital foramen small; zygoma unexpanded, slender butwith well-developed, rounded jugal process ; palate short, wide, with large anteriorforamen closed anteriorly by the premaxillae, its posterior wall extending along aline lying just behind the anterior faces of the upper premolars (pm4~4), partiallydivided medianly by the spike-like rearward extension of the fused palatal branchesof the premaxillae ; maxillary toothrows slightly curved anteriorly; palate ter-minating posteriorly on a line slightly in advance of the posterior faces of the thirdupper molars (m3-3). Mesopterygoid fossa not especially narrow, its width a littleless than one third of the external width across m3-m3 ; no bony post-palate, thepalation a little behind a line joining the posterior faces of m3~3 ; pterygoid wingsmoderately developed, divergent; no basioccipital pits ; tympanic bullae relativelylarge, flattened on the inner face. Coronoid process low, its tip below the level ofthe articular process, angular process slightly deflected. A NEW BAT FROM THAILAND 311 Soft palate (Fig. 6) with six transverse ridges, the first curved on each side to themedian line, second ridge nearly straight, third more definitely incurving to medianline, ridges four and five slightly less curved, sixth ridge straight; first ridge withshallow median division, ridges two to five with small median notch. Dental formula i \, c \, pm \, m f ; tips of upper incisors (i2-2) slightly convergent,i2 relatively large, separated rather widely from canine, flattened antero-posteriorly,longer than wide, with posteriorly a strong basal cingulum, the cusp acutely triangularin frontal aspect. Upper canine with moderately tall, slender shaft, its base tri-angular with narrow internal cingulum and small but prominent antero-internal 2 mm FIG. 6. Craseonycteris thonglongyai. <$ TNRC 54-3872. Palate ridges. cusp, readily visible from the front. Upper premolar (pm4) large, with wide lingualshelf and prominent anterior cingulum cusp, easily seen from the side, the posteriormargin of the tooth centrally slightly concave. First and second upper molars(m1-2) with hypocone markedly lower than the protocone, the hypoconal basin of m1more or less contiguous with the protoconal basin, the two basins in m2 slightly moresharply demarcated from each other in teeth with little wear, the rear face of eachtooth sharply concave in its central part, isolating the hypocone and metacone tosome extent, and producing a wide space between the medio-posterior part of thetooth and the anterior face of the succeeding tooth. Third upper molar (m3) reduced,the metastyle, metacone and hypocone lost, the mesostyle present but displacedinternally, only the first ridge or paracrista and a reduced second ridge or post-paracrista remaining, the latter terminating at the mesostyle which lies posteriorlyin the centre of the rear margin of the tooth, the pre-metacrista and metacrista lost.Lower incisors (i^g) about equal in size, tricuspid, the central cusp slightly the larger,the teeth long and narrow, lacking any low posterior supporting cusps, not imbricated,i2 separated from the canine by a small diastema. Lower canine with moderatelylong, slender shaft, rising from a narrow cingulum, with no accessory cusps. Firstlower premolar (pm2) large, longer than wide, a little longer but slightly narrowerthan the second lower premolar (pm4), with narrow cingulum and large cusp, 312 J. E. HILL triangular in lateral profile. Second lower premolar (pm4) slightly compressed intoothrow, rather caniniform, its base about as long as wide, with narrow cingulumand tall, slender, rounded spike-like cusp, a little exceeding the height of pm2 andequal in height to the first lower molar (mj), the cusp hollowed posteriorly. Lowermolars (nij-g) with no especial peculiarities, m3 a little reduced, its posterior triangleslightly smaller than the anterior triangle, the hypoconid and entoconid reduced. Humerus (Fig. 7) with trochiter or greater tuberosity about as large as trochin orlesser tuberosity, extending proximally beyond the head of the humerus, trochiterwith small articular surface on its posterior face, separated from head of humerus bya distinct groove, the articular surface of head extending across the groove to theventral face of the trochiter ; trochin well developed but not reaching proximally tohead of humerus which is rounded, not compressed laterally. Proximal face ofhumerus lacking distinct ridges, groove between trochiter and head of humerusdeepened at its anterior end to form a shallow supraglenoid fossa. Anterior face ofshaft of humerus flattened medianly in its proximal part, lacking any median deltoidridge, instead a low but distinct elevation extending dorsad from the base of thetrochiter to edge of flattened proximal part of shaft to form a short, narrow dorsalflange beneath the trochiter ; trochin similarly supported by a thin ventral flangeextending along a short part of the opposite side of the shaft, which itself is slightlycurved, the curve simple. Radial fossa indistinct; capitellum slightly displaced 2 mm FIG. 7. Craseonycteris thonglongyai. <J TNRC 54-3865. Left humerus. a. Dorsal aspect,b. Anterior aspect, c. Proximal aspect, d. Ventral aspect, e. Posterior aspect. A NEW BAT FROM THAILAND 313 2 mm FIG. 8. Craseonycteris thonglongyai. $ TNRC 54-3865. a. Dorsal aspect of left scapula,b. Ventral aspect of left scapula, c. Anterior aspect of left scapula, d. Medial aspect ofproximal end of left femur. from line of shaft, its principal articular surface spherical, not compressed or oblique ;lateral surface narrow, not extending distally as far as the principal part of the capitel-lum ; trochlea similarly narrow, extending distally about as far as the principalarticular surface ; epitrochlea or medial process about half as wide as distal articular 3*4 J. E. HILL surface, with small epitrochlear or distal spinous process, extending distally beyondthe articular surface. Sternum not especially modified ; presternum or manubrium T-shaped, withbroad, dorso-ventrally flattened lateral extensions or processes ; anterior part ofpresternum with moderately developed keel, posterior extension broad, flatteneddorso-ventrally, with low keel; mesosternum with longitudinal median ventralridge ; xiphisternum short, parallel-sided, with rounded tip, surrounded by cartilage.Clavicle slender ; scapula (Fig. 8, a-c) large, elongate ; supraglenoid tuberosityrelatively well developed, with small articular face ; coracoid border deeply notched,with narrow, ventrally directed flange ; axillary border slightly notched behindglenoid fossa with a shallow postglenoid depression ; acromion process slender,recurved antero-ventrally; coracoid process well developed, directed laterad,parallel-sided, its tip simple. Supraspinous fossa about one third the area of theinfraspinous fossa, with no especial features ; infraspinous fossa strongly faceted,the antero-medial facet wide, set at a sharp angle to the intermediate facet which iswide distally, the fossa divided more or less medianly by a deep, angular groove ; TABLE i External measurements (in mm) of Craseonycteris thonglongyai Width of narial padWidth of narial ridgeLength of earWidth of earLength of tragusLength of tibiaLength of foot (with claw) Length of forearmLength of thumbLength of IImLength of II1Length of IId cartilageLength of IIImLength of III1Length of III2Length of IV mLength of IV1Length of IV2Length of VmLength of V1Length of V2 Holotype No. of No. of cJTNRC speci- Range Mean speci- Range Mean 54-3871 mens mens 3'9 J7 3-8-4-3 4'1 8 3-8-4-4 4-0 2-2 17 1-9-2-4 2-2 9 2-0-2-5 2-2 9-6 I7 9-0-10-0 9-5 9 9-0-10-2 9-8 7-9 17 7-3-8-5 7'9 9 7-3-8-3 7-9 4-0 J7 3-8-4-2 4-0 9 3-8-4-1 4-0 12-3 16 11-9-12-9 12-3 9 11-7-12-8 12-3 6-024-4 4'925-0 0-52 0-6320-6 6-9 2O-I 22'7 3'910-3 22-7 5'76-0 17 5-9-6-8 6-3 9 5-8-6-6 6-2 16 24-1-25-2 24-6 9 22-5-25-8 24-9 17 4-6-5-3 5'° 9 4-6-5-3 4-9 17 24-1-25-6 24-7 9 23-9-26-5 25-6 17 0-40-0-55 0-50 9 0-45-0-54 0-51 17 0-40-0-63 0-52 9 0-42-0-60 0-51 16 ig-8-21'6 20-7 9 19-7-21-7 2I-I 17 6-4-7-0 6-7 9 6-5-7-1 6-8 16 18-7-21-5 20-3 9 19-4-21-2 2O-I 17 21-7-23-2 22-4 9 21-7-23-8 23-1 *7 3-5-4-2 3-9 9 3-6-4-3 3-9 X7 9-7-10-7 IO-2 9 9-7-10-8 10-3 17 21-7-23-0 22-4 9 21-8-23-7 23-2 17 5-2-5-7 5'4 9 5-2-5-9 5-6 16 6-1-7-0 6-3 9 5-9-6-6 6-2 The nomenclature used for the digits in this Table, in Table 3 and in Table 4 is an extension of Andersen(1905: 246, footnote), i.e. IIId is the third digit, IIIm its metacarpal, and III1 and III2 its first and secondphalanges. A NEW BAT FROM THAILANDTABLE 2 Cranial measurements (in mm) of Craseonycteris thonglongyai Greatest length of skullCondylocanine lengthCondylobasal lengthPalatal lengthPalatilar lengthWidth across ante-orbital foraminaWidth across lacrimalsWidth across rostral swellingsZygomatic widthPostorbital widthWidth of braincaseHeight of braincaseMastoid width Holotype <J TNRC 54-3871 n-i 9-9 10-0 4-0 3-8 3-84'4 5-8 2-2 5-6 No. ofspeci- m3-m3 4-6 Post-palatal width 1-5Width across pterygoid wings 1-8 Length of tympanic bulla 2-6 Width of tympanic bulla 1-8 Length of cochlea 2-1 Width of cochlea 2-2 Width of basioccipital 0-6 c-m3 3-8Length of complete mandible 6-9 Length of right ramus 7-0 c-m3 4-0 Range 10-6-11-5 9-9-10-1 10-0-10-3 3-9-4-23-7-3-9 3-7-4-04-3-4-6 4-0-4-45-5-6-02-1-2-3 5-3-5-74-0-4-35-7-6-02-9-3-2 4-6-4-91-4-1-5 1-8-2-02-4-2-7 1-5-1-9 1-8-2-22-0-2-3 0-5-0-7 3-7-3-9 6-9-7-17-0-7-34-0-4-2 No. of Mean speci- mens II-2 5 10-0 5 10-1 5 4"1 5 3-8 5 3-9 5 4'4 5 4'1 5 5-8 5 2-2 5 5'5 4 4'1 4 5-8 4 3'° 5 4'7 5 i-5 5 1-9 5 2-6 5 1-7 5 2-0 5 2-2 5 0-6 5 3-8 5 7-0 4 7-1 4 4-1 5 Range Mean 10-3-11-1 10-9 9-6-10-0 9-9 9-5-10-1 9-8 3-8-4-1 3-9 3-6-3-9 3'7 3-7-3-9 1-7-2-02-3-2-61-6-1-8 1-8-2-22-O-2-I0-5-0-6 3-6-3-8 6-7-6-96-9-7-13-9-4-° 3-84'4 5 3-9-4-3 4'I 5 5-5-5-7 5-6 5 2-1-2-2 2-1 4 5-4-5-5 5'5 4 3-9-4-0 4-0 4 5-7-5-9 5-8 5 2-8-2-9 2-8 5 4-5-4-6 4-6 5 1-4-1-5 x'4 1-82-61-7 2-O2-1 0-63'7 6-87-04-0 postero-lateral facet about as wide as intermediate facet, with proximally a low ridgeextending from the base of the glenoid fossa along but just within the axillary borderand distally along the further two thirds of the axillary margin ; ventral surface ofscapula with four facets, underside of postero-lateral facet rather narrow. Last cervical vertebra not fused to first thoracic ; lower three thoracic vertebraefused, their boundaries faintly visible ; lumbar vertebrae except last two solidlyankylosed, the last two free ; sacral vertebrae fused, their boundaries obliteratedor nearly so ; two small caudal vertebrae. Pelvis small, weak, the ilia long and veryslender ; ascending ramus of pubis short, the depth of the pelvis less than one halfof its total length ; no definite acicular or pubic spine ; ventral ramus of pubisthin and delicate ; ischium with similarly thin, delicate ascending ramus, thesestructures easily lost. Femur (Fig. 8, d) slender, delicate, its proximal part not 3i6 J. E. HILL markedly deflected, with strong flange on its anterior face ; trochanters small butevident, the lesser trochanter slightly the larger and extending proximally a littlefurther than the greater trochanter. Fibula thread-like, tapered, extending abouthalfway along the tibia. Measurements of Craseonycteris thonglongyai appear in Tables I and 2. REMARKS. The species is named to commemorate the late Kitti Thonglongya,formerly Curator of Terrestrial Vertebrates at the Centre for Thai National ReferenceCollections, whose last discovery it was. His untimely death was a sad loss to hismany friends and to mammalogy in southeastern Asia, especially in Thailand wherehis enthusiasm and energy will be greatly missed. The suiform muzzle of Craseonycteris thonglongyai has suggested (H. ElliottMcClure, in litt.) the vernacular name 'Hog-nosed bat' for this species : it has alsobeen described (J. T. Marshall, in litt.) as the 'Bumble-bee bat'. REVIEW OF DIAGNOSTIC FEATURES The current taxonomic arrangement of the Chiroptera derives largely from thework of Miller (1907) who based the higher categories largely on the structure of theshoulder girdle, the humerus, sternum, spine and pelvis, and of the skull and teeth,remarking (p. 45) that 'in a group of characteristically volant animals the chieftaxonomic importance must be assigned to the development of the wings'. Morerecently, Walton & Walton (1970) and Vaughan (i97oa) have reviewed the structureof the skeleton among the families of bats and Slaughter (1970) has examined theirdentition. Smith (1972) has reviewed a number of characters valuable in theclassification of bats at the familial level and in examining the New World familiesrelied greatly upon a more detailed examination of the humerus, the femur andtragus than did Miller, together with a lesser emphasis upon such features as acousticemissions, hair structure, karyotypes, the histology of facial glands, ectoparasites,brain size and immunology, not always readily available to the systematist. Thenature of the available material of Craseonycteris at present limits any study of thegenus from the familial standpoint chiefly to the traditional characters employed byMiller, and to these have been added a number of features that seem relevant whenconsidering the familial position of this unusual bat. A synopsis of the diagnosticfeatures in the families of bats, with much other information, is given by Koopman& Cockrum (1967). Muzzle The muzzle of Craseonycteris resembles that of the Rhinopomatidae (which includesbut one genus, Rhinopoma) in several respects. As in that family the muzzle islaterally swollen and the nostrils open directly anteriorly in a thickened narial pad,which is surmounted by a low dermal ridge. The nostrils, however, differ sharply :in the Rhinopomatidae they are slit-like and valvular (usually closed in preservedspecimens) and situated quite closely to the upper margin of the narial pad, inclinedat about 30 degrees to the horizontal. In Craseonycteris the nostrils are wide, the A NEW BAT FROM THAILAND 317 inner margin straight, the outer margin sharply curved and they lie nearer to theupper lip, set more sharply to the horizontal at an angle of about 60 degrees. Thereis no definite internarial septum in the Rhinopomatidae, and the superior dermalridge is basically an extension of the narial pad ; in Craseonycteris the nostrils areseparated by a clearly defined internarial septum which broadens in its upper partto form the base of a narrow superior dermal ridge extending over the central partof the narial pad, above the nostrils, and not completely across the pad as in theRhinopomatidae. Among the Emballonuridae the nearest approach to Craseonyc-teris in the structure of the anterior muzzle is found in Balantiopteryx in which thenostrils are of similar shape and open anteriorly in the face of an anteriorly directed,nearly vertical pad and are separated by a wide septum, but there is no superiordermal ridge. It may be conjectured that the low superior dermal ridge in Craseonycteris andperhaps in the Rhinopomatidae represents a rudimentary form of nose-leaf. A broadlysimilar but more developed structure occurs in the Vespertilionidae (in the subfamilyNyctophilinae), while in the Hipposideridae the broadened upper part of the inter-narial septum forms the base of the central part of a sometimes complex nasalfoliation. Furthermore, among the Phyllostomatidae the spear-like nose-leaf is adirect extension of the anterior muzzle. Tragus The tragus of Craseonycteris is well developed as it is in the Rhinopomatidae, buttapered distally and not truncate as in that family. Among the Emballonuridae thetragus is spatulate or pyriform. In Craseonycteris the tragus is remarkable for thecurious oblate swelling at the mid-point of its anterior part, common to both maleand female specimens. A slight swelling of the anterior part of the tragus is some-times to be found in the Rhinopomatidae. Wing structure This in Craseonycteris provides a number of interesting features deserving furthercomment. The propatagium is broad and unusually large, a feature associated byVaughan (igyob : 210) with hovering flight, in allusion to the glossophagine bats.In this respect Craseonycteris differs sharply either from the Rhinopomatidae or fromthe Emballonuridae, in which the propatagium is narrow. The relation of the secondand third metacarpals and their associated phalanges to the intervening wingmembrane that they support follows a similar pattern in Craseonycteris and in theRhinopomatidae, although the method of support differs. In Craseonycteris and inthe Rhinopomatidae that part of the wing membrane lying between the second andthird digits tapers from the end of the second digit to terminate distally about onequarter or a little more of the distance along the second phalange of the third digitfrom its junction with the first phalange. In the Rhinopomatidae this part of themembrane is supported anteriorly by a second digit with a metacarpal similar inlength to the third metacarpal, together with two relatively long phalanges : the 3i8 J. E. HILL total length of the digit is equal almost to the combined lengths of the third meta-carpal and its first phalange. A similar arrangement prevails in Craseonycterisalthough achieved in a different fashion : the second metacarpal is much longerthan the third metacarpal, the very short phalange and its cartilaginous appendagecontributing little to the total length of the second digit, which equals or slightlyexceeds the combined lengths of the third metacarpal and its first phalange. In theEmballonuridae this part of the leading edge of the wing is relatively shorter, themembrane between the second and third digits terminating distally on the thirddigit at or about the junction of its first and second phalanges and supported solelyby the second metacarpal which is shorter than the third metacarpal, although onlyslightly so. The second digit in the Rhinopomatidae has two well-developed bony phalanges :this digit in the Emballonuridae, Nycteridae, Rhinolophidae, Hipposideridae, Nata-lidae, Furipteridae and Thyropteridae lacks phalanges while the Myzopodidae havea cartilaginous rod. The second digit in the remaining extant families of the Micro-chiroptera has one bony phalange but it is minute in the Mystacinidae and rudi-mentary in the Noctilionidae and Molossidae. In Craseonycteris the second digithas one very short, insignificant bony phalange, usually integrated with the meta-carpal, tipped with a short cartilaginous appendage. As in Craseonycteris, there isno distinct third phalange on the third digit in the Rhinopomatidae, Emballonuridae,Nycteridae, Megadermatidae, Rhinolophidae, Hipposideridae and Noctilionidae.The third phalange is cartilaginous, but ossified at its extreme base, in the Natalidae,Furipteridae, Vespertilionidae and Molossidae : in the remaining extant Micro-chiropteran families the third digit has three bony phalanges. In the graduation of the metacarpals, Craseonycteris differs quite markedly eitherfrom the Rhinopomatidae or from the Emballonuridae. In Craseonycteris thesecond metacarpal is the longest, the third the shortest, the fourth a little longerthan the third and the fifth about as long as the fourth. In the Rhinopomatidaethe second and third metacarpals are similar in length, the fourth considerablyshorter, with the fifth longer and similar in length to the second and third. Amongthe Emballonuridae, the second is shorter than the third, the fourth yet shorterand the fifth the shortest: the tendency to shorten the fifth metacarpal in relationto the third and fourth is more pronounced among the larger genera. The relativewing proportions of many of the families of living Microchiroptera were studied byRevilh'od (1916, tabs. 1-5) who for numerous species expressed the lengths of theindividual components of digits three to five as a percentage of the length of the fore-arm : the relative wing proportions for Craseonycteris are given in Table 3. Thepattern of third metacarpal the shortest of metacarpals three to five, followed inlength by the fourth metacarpal and with the fifth subequal to or longer than thefourth occurs most frequently in the Megadermatidae, the Rhinolophidae, infre-quently in the Hipposideridae and in some Phyllostomatidae. The length of the first phalange of the third digit in Craseonycteris is a little morethan one quarter of the forearm length. In this respect Craseonycteris differssharply from the Rhinopomatidae in which the length of the phalange is one sixthor less of the length of the forearm. The Furipteridae (not studied by Revilliod) A NEW BAT FROM THAILAND 319 also have a very short phalange, similar in relative length to that of the Rhino-pomatidae. Among the Emballonuridae the length of the phalange varies from aboutone quarter to two fifths of the forearm length : in the Diclidurinae (not studied byRevilliod), however, the phalange is very short, like that of the Rhinopomatidae.In the Rhinolophidae, Hipposideridae, Noctilionidae, Mormoopidae, Desmodontidaeand Mystacinidae the relative length of the phalange varies from a little less than onefifth to one third or slightly more of the length of the forearm : in the remainingMicrochiropteran families the phalange is usually relatively rather longer althoughin some Phyllostomatidae its length may barely exceed one quarter of the forearmlength. The second phalange of the third digit in Craseonycteris is exceptionallylong, more than three quarters the length of the forearm and similar in length to thethird metacarpal. In the length of this phalange Craseonycteris differs widely fromthe Rhinopomatidae, in which the phalange is about one third of the forearm length,or from the Emballonuridae, in which the phalange, although relatively longer insome species, only rarely equals one half of the forearm length. The relative lengthof the second phalange of the third digit in Craseonycteris is equalled only by theNoctilionidae, and, in the Vespertilionidae, by the Miniopterinae, although it isequalled or exceeded by the combined lengths of phalanges two and three in someMormoopidae, by some species of Phyllostomatidae, notably in the Carolliinae andStenoderminae, and by one of the genera of Desmodontidae. The Megadermatidae,some Kerivoulinae among the Vespertilionidae and some Molossidae approachCraseonycteris in the relative length of the second phalange of the third digit.Furthermore, in Craseonycteris this unusually long phalange is reflexed beneath thewing in specimens preserved in alcohol: reflexion of the proximal phalange of thethird digit above the wing occurs in the Emballonuridae, Mystacinidae and Molos-sidae, and of the terminal phalange below the wing in the Noctilionidae and, amongthe Vespertilionidae, in the Miniopterinae. The extreme shortness of the first phalange of the fourth digit in Craseonycteris,where it is about one sixth of the length of the forearm, is also an unusual feature.In the Rhinopomatidae the phalange is one fifth or more of the forearm length, andas a rule similarly so in the Emballonuridae, although exceptionally in this familythe phalange may be a little less than one fifth of the length of the forearm. Amongthe remaining Microchiropteran families the Noctilionidae and rarely some Rhino-lophidae resemble Craseonycteris in this respect. The first phalange of the fourthdigit is relatively only a little longer in some others of the Rhinolophidae, in somespecies of the Mormoopidae and Desmodontidae, and in the Furipteridae. Thesecond phalange of the fourth digit is shorter than the first phalange in the Rhino-pomatidae, the Emballonuridae, Nycteridae, Thyropteridae (not studied by Revilliod)and Molossidae : in the Hipposideridae it is generally shorter than the first phalange,as it is with some exceptions, notably the Miniopterinae, in the Vespertilionidae.Otherwise, the second phalange of the fourth digit is longer than the first phalange :in Craseonycteris it is very long, its length exceeding two fifths of the length of theforearm and equalled in this respect only by the Noctilionidae and Miniopterinae.As in these, the terminal phalange is reflexed beneath the wing in specimens pre-served in alcohol. 320 J. E. HILL The first phalange of the fifth digit in Craseonycleris is about one quarter thelength of the forearm : in the Rhinopomatidae this phalange is very short, aboutone sixth of the forearm length and in the Emballonuridae rather longer, generally alittle less than one quarter of the length of the forearm but on occasion as long asnearly one third of the forearm length. In the remaining extant Microchiropteranfamilies the length of the phalange is generally between one fifth and one third or alittle more of the length of the forearm but in certain of the Noctilionidae, someMormoopidae and Desmodontidae and in some Vespertilionidae the relative lengthof the phalange is less than one fifth of the forearm length and the Mystacinidaein particular have a very short phalange, like the Rhinopomatidae. The secondphalange of the fifth digit is longer than the first phalange, as in Craseonycteris, inthe Megadermatidae, Rhinolophidae, some Hipposideridae, some Mormoopidae andPhyllostomatidae, the Desmodontidae, some Natalidae, the Mystacinidae and rarelyin the Vespertilionidae. Otherwise it is equal to or shorter than the first phalange. Revilliod (1916) summed the relative lengths of each digital component to providea relative indication of the total length of each digit. This technique shows Craseo-nycteris (Table 3) to have a wing with broadly the same proportions as the wing ofthe Nycteridae. The Rhinopomatidae, Emballonuridae, Mormoopidae and Mysta-cinidae have digital proportions falling short of those for Craseonycteris : the propor-tions for digits three and four in Craseonycteris agree closely with the Noctilionidaebut this family has a shorter fifth digit. The Rhinolophidae, Hipposideridae (exceptfor digit five in Coelops) and Furipteridae also have relatively shorter digits. Thedigital proportions in the Megadermatidae, the Myzopodidae and the Thyropteridaeapproach those of Craseonycteris, as do some Phyllostomatidae and certain of theDesmodontidae and Natalidae : others of the same families exceed them. For themost part the proportions for digits three and four in Craseonycteris are exceeded bythe Vespertilionidae but for digit five only by the Murininae and Kerivoulinae.Similarly, in the Molossidae the relative proportions for digits three and four exceedthose of Craseonycteris, but digit five in the Molossidae is much shorter. In generalterms, the wing of Craseonycteris is relatively wide and long, with a long tip. In the same paper, Revilliod attempted to demonstrate the degree of adaptationto flight in the families that he examined by means of an index obtained by sub-tracting the length of digit five as a total percentage of the forearm length from thetotal percentage length of digit three. This technique emphasizes the position ofspecies with long narrow wings in relation to those with short, broad wings : asmight be expected, the Molossidae, some Vespertilionidae and some Phyllostomatidaeoccupy the highest positions with the greatest values of the index, but the resultsobtained display wide variation with some Vespertilionidae and some Phyllostoma-tidae in relatively low positions. This is perhaps inevitable when one aspect onlyof the chiropteran wing is examined. The figure obtained for Craseonycteris(Table 3) is low, its wing, although long, being comparatively wide, a combinationobscured in the index employed by Revilliod. More recently, the morphological properties of the chiropteran wing have beenexamined in some detail by Findley, Studier & Wilson (1972). These authors havestudied the relation between certain properties of the bat wing to the mode of flight. A NEW BAT FROM THAILAND 321 In particular, the aspect ratio (length from wrist to tip of third digit plus length offorearm, divided by width at fifth digit) and the tip index (length from wrist to tipof third digit, divided by the length of the forearm) seem especially relevant whenconsidering Craseonycteris. In this genus (Table 3) the aspect ratio proves to besomewhat below the average demonstrated by Findley, Studier & Wilson (1972 : 430,tab. i, 434, fig. i) for all bats. However, the tip index is very high, placing Craseo-nycteris among those bats (glossophagines, murinines, kerivoulines) which hover orare thought to do so, or are migrants or high speed foragers (miniopterines, lasiur-ines, molossids) (loc. cit. p. 430, tab. i, 436, fig. 2, 437). Furthermore, Findley,Studier & Wilson suggest (p. 437) that the combination of a below average aspectratio with a high tip index is especially suited for hovering flight, as might be theelongate wing tip. In the Rhinopomatidae there is a complete contrast : accordingto Findley, Studier & Wilson (p. 430, tab. i, 434, fig. i) Rhinopoma has a below averageaspect ratio, combined with the lowest tip index of any bat studied (p. 430, tab. i,436, fig. 2) and, according to Harrison (1964 : 62) has an unusual, fluttering, undulat-ing flight. Little is known of the mode of flight of the Emballonuridae, but in thisfamily the aspect ratio was found by Findley, Studier & Wilson (p. 430, tab. i, 434,fig. i) to be considerably above the average for all bats, and the tip index (p. 430,tab. i, 436, fig. 2) well below. While variation in the proportions of the wing in the various Microchiropteranfamilies is such that these features evidently cannot be used as a ready and reliable TABLE 3 Wing proportions (forearm = 100) and wing characteristics for Craseonycteris thonglongyai ^^ oo oo Holotype No. of No. of cJTNRC speci- Range Mean speci- Range Mean 54-3871 mens mens 102-5 16 97-2-102-8 IOO-I 9 100-0-106-2 102-5 84-4 16 81-9-86-7 84-3 9 82-5-87-5 84-2 28-3 16 26-1-28-5 27-3 9 26-1-29-3 27-2 82-4 15 74-5-88-0 82-4 9 76-2-86-7 80-7 I95-I 14 183-9-201-4 193-4 9 185-8-203-5 192-6 93-0 16 88-6-93-2 90-8 9 90-3-96-4 92-8 16-0 16 14-7-16-9 15-7 9 14-5-16-9 15-5 42-2 16 39-0-44-2 41-5 9 39-9-43-4 41-4 151-2 15 144-5-152-5 148-4 9 145-8-156-4 149-7 93-0 16 88-6-93-0 90-9 9 91-0-96-9 92-9 23-4 16 20-8-23-4 22-O 9 20-3-23-2 22-4 25-0 15 24-5-28-5 25-7 9 23-0-28-0 24-8 141-4 14 136-1-143-0 138-6 9 137-1-148-0 140-0 53-7 14 47-2-61-5 54-8 9 46-5-58-1 52-6 2-09 14 2-06-2-17 2-II 9 2-05-2-15 2-09 i-95 15 1-84-2-01 i-95 9 1-85-2-03 1-92 Urn III1III2 IIId total IV1IV2IVd total V1V2 Vd total IIId total-Vd totalAspect ratioTip index Aspect ratio= IIId from wrist to tip + forearm/width at Vd. Tip index= IIId from wrist to tip/forearm. (IIId from wrist to tip= Illm + mi + HI2; width at Vd= Vn 322 J. E. HILL guide to classification at the familial level, it is clear that Craseonycteris cannot beallied closely either to the Rhinopomatidae or to the Emballonuridae from the struc-ture of its wing, and that it is adapted for a quite different mode of flight from eitherof these. The relative proportions of digits three to five in Craseonycteris correspondreasonably closely to those of the Nycteridae (Revilliod, 1916 : 164, tab. i) or withcertain of the Phyllostomatidae (loc. cit. p. 170, tab. 3). The broad wing of Craseo-nycteris does not imply a bat adapted for fast or sustained flight, and, indeed, thestructure of the wing suggests a hovering or at least a partially hovering species. Tail The tail is variable in the Microchiroptera. It may be present or absent (evenwithin the family) ; its length may exceed the length of the uropatagium, or it maybe subequal to this membrane in length or sometimes shorter, enclosed or partiallyenclosed within it, on occasion projecting from the posterior margin of the uropata-gium or emerging through its upper surface. The Rhinopomatidae are unique amongthe living families in the presence of a long, slender, mouse-like tail, emerging fromthe edge of a reduced uropatagium. In the Emballonuridae the tail is shorterthan the uropatagium and its free tip emerges through the upper surface about half-way or a little less along the length of the membrane. There is no external tail inCraseonycteris, but the uropatagium is moderate and rather full, an unusual featurein a tail-less species : Vaughan (igyob : 214) remarked that a large uropatagium isfound in some 'flycatcher' bats which forage near the ground or among vegetation,and which also have broad wings. Calcar As in the Rhinopomatidae, the uropatagium of Craseonycteris lacks calcarialsupport. Generally present in the Microchiroptera, the calcar is short or absent insome species of Phyllostomatidae, and rudimentary (on occasion not connected withthe membrane) in the Desmodontidae, but in these the uropatagium is reduced :its absence in a species with a moderate membrane is unusual. Throat glands and pubic nipples Throat glands are found in the Emballonuridae, Phyllostomatidae, Vespertilioni-dae and the Molossidae, often associated with a pouch or sac (Quay, 1970 : 23) :pubic nipples have been reported from the Rhinopomatidae, Megadermatidae,Nycteridae, Rhinolophidae, Hipposideridae, from the Nyctophilinae among theVespertilionidae and possibly the Phyllostomatidae (loc. cit. p. 27). Lateral rostral swellings The rostrum is variously swollen or inflated in a number of Microchiroptera.Lateral rostral swellings, however, are noticeably characteristic of the Rhinopomati-dae and Emballonuridae. In the Rhinopomatidae the swellings extend anteriorly A NEW BAT FROM THAILAND 323 beyond the narial aperture : those of Craseonycteris more nearly resemble the lateralswellings of the Emballomiridae. Premaxillae The structure of the premaxillae in Craseonycteris is unique, although it has clearlyan overall resemblance to the arrangement found in the Rhinopomatidae andEmballonuridae. In neither of these, however, do the premaxillae fuse anteriorly,although as in Craseonycteris they are free from the adjacent parts of the skull,with the palatal branch short or absent, nor do their well-developed narial branchesever reach dorsally to the apex of the narial aperture and fuse together. In someEmballonuridae the premaxillae may lie partially on the surface of the maxillae,especially in Taphozous in which there is a very distinct approach towards thecondition found in Craseonycteris. In Taphozous the ascending part of the narialbranch is greatly developed and widened in its upper part, extending dorsad almostto enclose the narial aperture, the two premaxillaries separated at the apex of theaperture by a distance of about one third of the total width of the aperture. Pos-teriorly, the upper part of each narial branch is deflected into a narrow plate orlamina which lies on the surface of the maxilla. The inner edge of the narial branchis thrust forward anteriorly to form a sub-tubular flange at the side of the narialaperture, extending upwards to the outer upper part of the opening and terminatingat the central division between the two branches. The premaxillae, therefore, forma structure similar in many ways to that prevailing in Craseonycteris, that is, a moreor less tubular anterior projection partially surrounding the narial aperture, risingfrom a basal plate which rests on the maxillae and nasals, except that in Taphozousthe surrounding ring and plate is incomplete, the narial branches failing to meet andfuse at the apex of the narial opening. The structure is rather less developed inTaphozous (Saccolaimus) than in T. (Taphozous) or T. (Liponycteris). In some othergenera of Emballonuridae, for example in Coleura, Saccopteryx or Diclidurus, thenarial branch of each premaxilla is widened to lie on the maxilla. A further stagefrom Taphozous is found in the Noctilionidae where the premaxillae are fused an-teriorly and to the maxillae : the narial branches are unusually long and welldeveloped, forming the sides of strongly tubular nares, separated above by the nasals.A similar arrangement prevails among the Mormoopidae. It is perhaps permissibleto speculate that the sub-tubular structure in Craseonycteris supports the narial padto which it is firmly attached by tough tissue. The free premaxillae in the Rhino-pomatidae, the Emballonuridae and in Craseonycteris may serve to increase themobility of the upper lip and the anterior part of the mouth when seizing food.They may also fulfil a similar function in the Nycteridae, Rhinolophidae and Hippo-sideridae but in these the premaxillae are represented only by their palatal branchesand are cartilaginous in the two latter families. Dentition The dentition of Craseonycteris is very similar to that of Rhinopoma, with only afew relatively minor differences. The most obvious of these are the relatively larger 324 J. E. HILL upper incisors, the presence of a small antero-internal canine cusp, and the slightlyfurther reduced condition of the third upper molar. Slaughter (1970 : 66) statedthat in the Rhinopomatidae (mis-headed Rhinolophidae) the third upper molarlacks the pre-metacrista and that the hypocones of the upper molars are completelyseparate from the protoconal basins. However, in the majority of specimens thethird upper molar has, as Miller (1907 : 82) pointed out, a metacone, mesostyle andthree commissures : the mesostyle is invariably displaced and the third commissureor pre-metacrista is very short, terminating in an ill-defined metacone. Wear rapidlyerodes the third ridge and metacone. The hypoconal basin of the first upper molaris broadly contiguous with the protoconal basin and separated only by a low com-missure soon eroded by wear: in the second upper molar the basins are moredefinitely separated particularly in teeth with little wear. Humerus Apart from the early study by Miller (1907), the humerus of bats is discussed inconsiderable detail by Walton & Walton (1970 : 105), Vaughan (i97oa : 117 et seq.)and Smith (1972 : 16). According to Vaughan (i97oa : 130) modifications of thehumerus for improved wing control are proximally the enlargement of the trochiter,the development of a supraglenoid fossa and of a large medial or deltoid ridge and,distally, the development of an epitrochlear or distal spinous process. Thesefeatures are considered in greater detail, but for New World families only, by Smith(1972 : 16 et seq.). Proximal end of humerus Among the Microchiroptera the Rhinopomatidae, Emballonuridae, Nycteridaeand Noctilionidae are sometimes considered primitive in that the trochiter is notgreatly developed and proximally does not extend significantly beyond the head ofthe humerus, and a supraglenoid pit is lacking in the proximal face of the humerusat the anterior edge of the head. To these, Vaughan (i97oa : 131) adds that themedial ridge is weakly developed, but some of these have a prominent medialdeltoid ridge. At the other extreme, the Vespertilionidae, Mystacinidae andMolossidae have a large trochiter which extends proximally well beyond the head,there is a deep supraglenoid fossa and the humerus has a substantial deltoid ridge.The remaining families form an intermediate group in which these characteristicsare developed to varying degrees. The proximal extension of the trochiter apparentlyprovides a locking mechanism which restricts the action of the humerus (Smith,1972 : 28) rather than a 'secondary articulation' or 'double articulation' as Miller(1907) thought. In Craseonycteris the trochiter is about as large as the trochin and extends proxim-ally beyond the head of the humerus and a definite, deep supraglenoid fossa sur-rounded by the proximal part of a deltoid crest is lacking. Instead, the anteriorpart of the groove between the trochiter and the head of the humerus is deepened toprovide a small fossa into which the supraglenoid tuberosity articulates. There is nodeltoid ridge but the anterior face of the humerus is slightly elevated at the base of A NEW BAT FROM THAILAND 325 the trochiter, the elevation passing into a dorsal flange, and a similar ventral flangesupports the trochin. The deltoid ridge is absent or very weak only in the Pteropo-didae but in some Phyllostomatidae (Micronycteris, Glossophaga) and, among theVespertilionidae in the Kerivoulinae, the deltoid crest is displaced dorsad, althoughonly slightly so. The proximal end of the humerus in Craseonycteris thus differssharply from those families considered primitive, and in the proximal extension ofthe trochiter approaches the Vespertilionidae. The lack of a definite deep supra-glenoid fossa corresponds more closely to the allegedly primitive families, althoughthe trochiter in Craseonycteris is separated from the head by a moderate groove witha shallow fossa, and the trochiter itself has small articular surfaces on its posteriorface and on its ventral face opposite the humeral head. In Craseonycteris the headof the humerus is rounded, as in the Rhinopomatidae and in many of the otherMicrochiropteran families : it is elongate or oval in the Emballonuridae, Rhinolo-phidae, Hipposideridae and Noctilionidae, elliptical in the Megadermatidae andMystacinidae and variable in the Phyllostomatidae. Distal end of humerus Certain features of the distal end of the humerus were employed by Miller (1907)in the familial classification of the Chiroptera. These included the size and positionof the capitellum, the development of the lateral epicondyle or external condyle, ofthe trochlea or medial epicondyle and of the epitrochlea, sometimes called theinternal condyle or medial process, and the presence or absence of an epitrochlear ordistal spinous process. These structures are discussed in considerable detail bySmith (1972 : 16), in relation to the New World families, and by Felten, Helfricht &Storch (1973) to European species. Capitellum, lateral epicondyle and trochlea These articular surfaces may vary in shape, size and proportion, and also in theirdegree of displacement dorsally from the axis of the shaft of the humerus. Variationsin shape, size and proportion seem to offer little of familial significance. As a rulethe central part of the capitellum is spherical or slightly oblong and forms the greaterpart of the articular surface : the lateral epicondyle is narrow and the trochleasimilarly undeveloped, each separated from the principal surface by shallow grooves.The trochlea may become widened as in Diclidurus of the Emballonuridae to equalin width the central part of the capitellum, or the central part of the capitellum maybe narrowed as in Mormoops of the Mormoopidae. In distal extension the trochleamay equal the lateral epicondyle, extend beyond it or beyond either the lateralepicondyle or the principal surface of the capitellum. This type of articulation,with some slight modification, is found in the majority of the families of Micro-chiroptera, but not in the Vespertilionidae or Molossidae. In these the principalpart of the capitellum is angular or narrowed, and is tilted diagonally, a conditionfaintly foreshadowed in the Rhinolophidae and Hipposideridae, in which the other-wise spherical principal surface tends to be narrowed distally, or in the Nycteridae,where it is slightly oblique. The articular surfaces in Craseonycteris correspond 326 J. E. HILL closely to the first of these patterns and indeed closely resemble the pattern found inthe Rhinopomatidae, most Emballonuridae and the Phyllostomatidae. The displacement of the articular surfaces also varies between families. Thecapitellum in the Rhinopomatidae is slightly but distinctly displaced from the lineof the shaft: it is similarly or more dorsally displaced in the majority of Microchirop-teran families excepting the Emballonuridae, the Vespertilionidae and the Molossi-dae, although in some, such as the Noctilionidae and the Mormoopidae, as in theRhinopomatidae, the degree of displacement is slight. In the Rhinolophidae andto a lesser extent the Hipposideridae the capitellum is widely displaced dorsally. InCraseonycteris the articular surfaces are slightly removed from the line of the shaftas they are in the Rhinopomatidae. Epitrochlea The epitrochlea or medial process may display varying degrees of development andventral extension. In the Rhinopomatidae and for the most part the Emballonuri-dae, the epitrochlea is short and broad, the ventral portion sometimes curled : asimilar process occurs in the Rhinolophidae and Hipposideridae. In the Nycteridae,Megadermatidae and Noctilionidae the epitrochlea is more massive and projectsrather further ventrally : the Mormoopidae have a moderately developed epitrochlea,while among the Phyllostomatidae the epitrochlea varies from a relatively poorlydeveloped condition to one displaying a moderate degree of development and ventralextension. The small families Natalidae, Furipteridae, Thyropteridae and Myzopo-didae have a broad but rather short epitrochlea. The Vespertilionidae and Molossi-dae present a sharp contrast in a small, undeveloped epitrochlea which projectsventrally only slightly beyond the lateral line of the shaft. The epitrochlea inCraseonycteris is similar to that of the Rhinopomatidae, most Emballonuridae, or tothose of the less developed processes found among the Phyllostomatidae. Epitrochlear process This is the spinous process of Miller (1907) or the distal spinous process of Smith(1972 : 17) which this author defined as the process extending distally from the tipof the epitrochlea or medial process. A second type of process is encountered insome families, arising not definitely from the tip of the epitrochlea, but from itsdistal margin. A similar, less developed structure is found among certain Phyllo-stomatidae, notably in Lonchorhina. Smith (1972 : 21) advanced evidence to suggestthat in this genus at least the structure is not homologous with the distal spinousprocess arising from the tip of the epitrochlea, but is a departure from the basicallybilobed epitrochlea found in certain members of the family. An epitrochlear process occurs in most of the Microchiropteran families, reachingan optimum in the Molossidae. It is absent from the Rhinopomatidae and absentor poorly developed in the majority of Emballonuridae although sometimes welldeveloped as in Taphozous and Dididurus. It occurs in the Rhinolophidae, Hippo-sideridae and Noctilionidae and is well developed in the Mormoopidae. The Phyllo-stomatidae have no epitrochlear process or at least only a weak development of it : A NEW BAT FROM THAILAND 327 a broad, low process is found in the Natalidae, Furipteridae, Thyropteridae andMyzopodidae. In the Natalidae, Furipteridae and Myzopodidae the dorsal base ofthe process is closely adpressed against the rim of the trochlea. The process is lowbut strong, virtually forming a part of the rim of the trochlea, in the majority of theVespertilionidae, but is particularly well developed in the Miniopterinae, to equal thecondition found in the Molossidae, in which the process is long, projecting wellbeyond the articular surfaces, and closely adpressed to the trochlear rim : theMystacinidae also have a well-developed process. In the Nycteridae and theMegadermatidae the process is of the second type described above : in these Miller(1907 : 100) described it as 'styloid' rather than spinous : a similar process is foundin some Rhinolophidae. It is of some interest to remark that the process is widelyremoved from the trochlear edge in the Emballonuridae, Rhinolophidae, Hippo-sideridae, Nycteridae, Megadermatidae, Noctilionidae and Phyllostomatidae. Avariety of positions to one in which the process is almost in contact with the rim isfound in the Mormoopidae (Smith, 1972 : 19, fig. 4). In Craseonycteris the processoriginates from the tip of the epitrochlea rather than from its distal edge, and isremoved from the trochlear rim. It is similar to the process found in the Rhinolophi-dae or Hipposideridae, or to the less advanced of the Mormoopidae. Vaughan (i970a : 130) remarked that the epitrochlear spine is not evident in theRhinopomatidae, Emballonuridae, Noctilionidae and Nycteridae and is only fullydeveloped in the Vespertilionidae, Mystacinidae and Molossidae, with the otherfamilies occupying an intermediate position. He concluded (p. 131) that theprimitive families lack the epitrochlear process while the advanced families possess it.However, the process is present in some of the allegedly primitive families, althoughin these it may take a rather different form from the process in those apparentlymore advanced. Vaughan (p. 132) further considered the development of the epitro-chlear process a factor evolved in response to the need for lightening the distal partsof the wing so that the wings might be controlled more easily. A similar view isadvanced by Smith (1972 : 23) who suggested that the process in the Mormoopidaeand Molossidae forms part of an automatic flexing mechanism. It is perhaps notwithout significance that a well-developed process in one form or another occurs notinfrequently in bats in which the third digit is elongated, even in families, forexample, such as the Emballonuridae and Vespertilionidae in which the processeither is not normally greatly developed or does not normally extend distally to anygreat extent. Scapula The modifications of the chiropteran scapula are briefly discussed by Walton &Walton (1970 : 100) and by Vaughan (i97oa : 128). The scapula in Craseonycterishas a number of relatively unmodified features : it is rather narrow, the supra-spinous fossa is relatively large and lies in the same plane as the post-spinous part ofthe blade, the anterior flange is weak and the coracoid process is directed laterad.The infraspinous fossa, however, is quite strongly faceted, so that while in mostrespects the scapula resembles that of the majority of Microchiropteran families, in 328 J. E. HILL this feature it tends towards those in which the scapula is more specialized, such asthe Molossidae, or Lasiurus or Miniopterus of the Vespertilionidae. This featureoccurs also in Taphozous of the Emballonuridae, and, in this genus, in addition, thearea of the supraspinous fossa is reduced, with a strong anterior flange. There isalso in Craseonycteris a degree of articulation with the trochiter. Fusion of lumbar vertebrae In this respect, Craseonycteris resembles the Natalidae or the Furipteridae, alsosmall, delicately formed bats. However, vertebral fusion occurs in some genera ofthe Hipposideridae and also in Kerivoula of the Vespertilionidae. As Miller (1907 :182) pointed out, the feature is of no value in determining familial position. Proximal end of femur The proximal end of the femur in Craseonycteris resembles the condition found inthe Emballonuridae, with the lesser trochanter a little larger than the greatertrochanter and extending a little further proximally. It differs from the Rhino-pomatidae in which the lesser trochanter is similar in size to the greater trochanterbut stands marginally a little lower. The head and neck of the shaft of the femurare not deflected from the line of the shaft as in the Rhinolophidae and Hipposideri-dae. There is considerable variability in the structure of the head of the femuramong the Microchiropteran families (Smith, 1972 : 29, 30, fig. 8) but the femur hasa more limited value in familial classification than has the humerus, although thisauthor found the Mormoopidae to lack well-developed trochanters, in contrast tothe other families that he had examined. A summary of the more obvious characters discussed appears in Table 4. SYSTEMATIC POSITION OF THE CRASEONYCTERIDAE The new family Craseonycteridae resembles the Rhinopomatidae in some externalfeatures, some cranial characters and especially in the details of its dentition. Inparticular, the narial pad with rudimentary superior dermal ridge, the structure ofthe leading edge of the wing, the lack of calcarial support to the uropatagium, theswollen rostrum, the free premaxillae with prominent narial branch, the shape of thedental arcade and the architecture of the teeth themselves provide good charactersin which it tends towards this monotypic family. At the same time, its unspecializednostrils, large, independent ears, the tragus, the highly modified wing which con-trasts sharply with the rather simple wing of the Rhinopomatidae, the lack of a tailand the features of the humerus, scapula and pelvis set it widely apart from theRhinopomatidae. Although the new family has some features in common with theEmballonuridae, notably in the structure of the nostrils, the presence of rostralswellings and in the structure of the premaxillae which is clearly foreshadowed insome Emballonuridae, the Craseonycteridae differ widely from this family in many A NEW BAT FROM THAILAND 329 other features. Among these the tragus, the lack of a tail, the absence of post-orbital processes and of basioccipital pits, the dentition and the organization of theshoulder joint seem particularly notable. Furthermore, the Craseonycteridaediffer widely from the Emballonuridae in the overall structure and characteristics ofthe wing. The Craseonycteridae differ so widely from the Nycteridae, Megadermatidae,Rhinolophidae and Hipposideridae that any close relation is most unlikely. Thereexist also wide differences from the Noctilionidae and Mormoopidae, especially inthe form and structure of the rostrum, of the palate and teeth, and of the head ofthe humerus. As with all of these families, the premaxillae provide a major distinc-tion between the Craseonycteridae and any of those remaining, although it alsodiffers from these in numerous other features. However, in the structure of thehumeral head the Craseonycteridae approach those families considered moreadvanced in this respect, beginning to some extent with the Phyllostomatidae andterminating with the Vespertilionidae, Mystacinidae and Molossidae, although thehumerus in the Craseonycteridae is less modified in comparison with the last threeof these. My conclusion is, therefore, that the Craseonycteridae must be classifiedwith the Rhinopomatidae and Emballonuridae. The wing in the new family isclearly more modified than in either of these, perhaps for a mode of flight involvinghovering, by the enormous extension of the third digit with the concomitant develop-ment of the humerus. For this reason the Craseonycteridae should follow theRhinopomatidae and the Emballonuridae : few features of the new family suggestthat it in any way intermediates between these two. It is more correct to say that itcarries further features already apparent in one or other, or both, of these. Current classification of the Chiroptera is usually based on Simpson (1945).This author assembled the families delineated by Miller (1907) into superfamilies.Smith (1972 : 39) remarked that these superfamilies were based on Winge's familiesas expressed in his work (1892) on the mammals of Lagoa Santa, Brasil, and that as aresult the Noctilionidae were grouped with the Rhinopomatidae and Emballonuridaein the superfamily Emballonuroidea, while the mormoopids were carried as a sub-family of the Phyllostomatidae into the Phyllostomatoidea. However, Wingeconsistently (1892 : 24 ; 1923 : 235 ; 1942 : 271) placed Noctilio with the mormoopidgenera in the Phyllostomatidae, although Simpson (1945 :180) stated that his classi-fication retains Miller's families but groups them into superfamilies which are thoseof Winge. Possibly in this instance in grouping the Noctilionidae with the Rhino-pomatidae and Emballonuridae Simpson was influenced by the remarks of Miller(1907 : 97) where relationship between these families is suggested. The Craseonyc-teridae stand closer to the Rhinopomatidae and Emballonuridae than to either theNoctilionidae or Mormoopidae. For this reason the new family should be includedwith the rhinopomatids and emballonurids in any further grouping, preferably as asuperfamily, the Emballonuroidea. Within this assemblage, the Craseonycteridaeare the most advanced, especially in wing structure. Smith (1972 : 39 et seq.)advanced reasons for considering the Noctilionidae, Mormoopidae and Phyllo-stomatidae as three distinct families possibly forming another superfamily, thePhyllostomatoidea although he noted (p. 41) that this grouping seemed premature. 330 J. E. HILL H c I -fi +l + * N | +| +| + + | 912plpodozAj\[ I I I + o rr> I I I + + +I I I + O ro I I I + + + + o N I I 9BptdoOUIJOJ\[ tuo bo i 111 MH ° fe ^ b HI g nJ -£ -^ 8 "°- ^ 3 11 .5 .5 g « i> C CH C j? ^ >^o E C o o c a >i <D v> o D a,-4-> -M a> w ^ rD42l<HM-('-l-<r72 CUOlDl-lH •S-SSec^8 -§&3 b St e e 5 *3 5"S13 O U •— "— •<-< F— —' »— W* ^/-\t^K>K>Wt_.r\ Ui : A NEW BAT FROM THAILAND + + +I+II+I I ++I + I+ I+ + + +II + I + + +II+ I ++II++ I++IIII + I + +I + II I + + +II+ I+ + +I+IIIII + + +II+ I + + +II+ +I + IIIIII + + +II+ I + + +II+ +I++I + III + +I + II I + + +II+ +I + III + II + + +II+ I + + +II+ +I++II + II + + +II+ I + + +II+ +I + IIII + I + + +II+ I + + +II+ +I+IIIIIII + + +II+ I ++I + I+ +11 + 111 +111 + +I + I+ I III + I+ +I + IIIIII + III + I I ++I + I+ +I+ + + + +III + III + I I ++I + I+ +I+ + +IIII I I I I I I I IIII++ +I+ + +IIII + III + I I 11 + 11+ +I+ + +III + + 11 + 1+ + + + +III +I + III + II +11+11 I I I I + I + I I +1 I I I I I I + 11 + 11 I 11 + 11+ +IIIIIIII +-> -t-> en en 2 d d 13 o ^ eu cu en £ cu en en X> X5rt oj fl -a 3A 3 M-c T3 V- IH T3 nJ HH en T3 CU cu rt 1 0 0 T! cu fi — O 3 N IH 3 en•a N a OH bo 1 g •2» -2 t> cu flen «3 Od o ;hes of premaxillae shortties of premaxillae short ;hes of premaxillae fusei:hes of premaxillae fus •, lying on maxillae ai;ending distally beyond . fossa in proximal faceierus rounded 13_o 4J1^ O "cuen en3 3l-i !H cu cu ^H r- | ^ o rt ^3en en CU vtj ° Ocu *HH cu • *H en |~H CD d """ ' -^28o ^ ^* process I shoulder girdle specialiand first thoracic verteb ond thoracic vertebrae f;ebrae fused ^cu & 13 CU ^j — , Absent. + , Variablt + 332 J. E. HILL All are clearly distinct from the Rhinolophoidea, including the Nycteridae, Mega-dermatidae, Rhinolophidae and the Hipposideridae : the members of this super-family have diverged considerably in different ways. Smith (1972 : 40, 41) reviewed the possible phylogeny and evolution of the NewWorld families Noctilionidae, Mormoopidae and Phyllostomatidae, and concludedthat for various reasons the commonly accepted view that the Microchiropteran batsevolved from an emballonuroid-like ancestor might be challenged. The furthercorollary to this hypothesis is that the New World noctilionids, mormoopids andphyllostomatids derive from an Old World emballonuroid migrant or migrants.Instead, Smith postulated a 'palaeochiropteran' ancestor in both the Old and NewWorlds, thus providing a base for an autochthonous lineage for the New World batfamilies with which he was concerned. The family Craseonycteridae presents littlethat can be described as truly intermediate between the Rhinopomatidae and Em-ballonuridae on the one hand and on the other hand the Noctilionidae, Mormoopidaeand Phyllostomatidae. However, some features might be considered in this way :the low dermal narial ridge, for example, might be construed as an incipient nose leaf,or the sub-tubular premaxillae, which are foreshadowed in the Emballonuridae andreach a further stage in the Noctilionidae and Mormoopidae, might be thought todisplay an intermediate condition. Slaughter (1970 : 66) discussed the origins of the rhinopomatid dentition andpointed out that it could conceivably have taken origin from within the Emballonuri-dae. However, the structure of the hypo-protoconal basins in the Rhinopomatidaeindicated to this author that this dentition could derive directly from a superfamilyprototype, or from incipient emballonurid stock, the latter seeming to him moreprobable. The conclusion that the dentition of the Rhinopomatidae derives at leastfrom an ancestral prototype rather than from within the Emballonuridae is rein-forced by the Craseonycteridae in which the rhinopomatid dentition appears, littlechanged, in a bat otherwise widely removed either from the Rhinopomatidae or fromthe Emballonuridae, and, indeed, much more widely removed from the Noctilionidaeor Mormoopidae. The new family emphasizes the difficulties and dangers of in-ferring phylogeny from a variety of modern forms each of which exhibits a varyingcombination of different specializations and modifications in different degrees. Allthat can be said is that the Rhinopomatidae, Emballonuridae and Craseonycteridaehad most probably a common ancestry. Miller (1907 : 81) regarded the Rhinopomatidae as the most primitive of livingmicrochiropteran families, largely on account of the presence of two phalanges inthe second digit, the free premaxillae and the primitive shoulder joint. However,there is much to commend the views of Winge (1923 : 267 ; 1941 : 310) who con-sidered Rhinopoma, the sole representative of the family, to be rather isolated amongliving bats. In many respects, he averred, it is highly specialized, as in its dentition,the form of the nasal cavity and the upper arm, while other features, such as the pre-maxillae and the presence of two phalanges in the second digit, indicated a primitiveorigin. Winge noted that Miller had stated that in Rhinopoma there is no 'secondaryarticulation' between trochiter and scapula but that in fact a narrow articulation ispresent. Indeed, the trochiter may on occasion very slightly exceed the head of the A NEW BAT FROM THAILAND 333 humerus. As Winge pointed out, Miller placed much emphasis on the features thathe regarded as primitive. It has long been customary to associate the Rhinopomati-dae with the Emballonuridae (Dobson, 1878 : 353 et seq. ; Winge, 1892 : 33 ; 1923 :235 ; 1941 : 271 ; or Simpson, 1945 : 55) but there seems little in these features tosuggest that the Rhinopomatidae are necessarily the more 'primitive'. The reten-tion of two phalanges in the second digit of Rhinopoma may well be a corollary of theneed to support a long leading edge of the wing, achieved in the Craseonycteridae bya relatively much longer metacarpal with insignificant phalangeal support: thesimple shoulder joint with little modification of the proximal part of the humerus isperhaps a reflection of a correspondingly unspecialized wing. Furthermore, the val-vular nostrils of Rhinopoma seem highly specialized : its long tail is an extreme of avariable chiropteran character and is possibly a 'primitive' feature although it mayserve as a balancing organ in flight. The Emballonuridae themselves are said by Miller (1907 : 84) to combine thegreatest number of primitive characters with the least degree of specialization.Miller appears to have based this conclusion on the presence of postorbital processes,on the free premaxillae and the structure of the proximal part of the humerus. Yetthe Emballonuridae display considerable modification of the premaxillae, varyingfrom a comparatively simple condition in which the narial branches lie alongsidethe lateral margins of the narial aperture to the relatively advanced condition foundin Taphozous in which a sub-tubular structure tending towards that found in theCraseonycteridae lies more or less on the surface of the maxillae. The deep basi-occipital pits, sometimes divided by septa, seem also to be a specialized feature, and,as in the Rhinopomatidae, the number of teeth is reduced, although the process hasnot gone so far as in that family, only Taphozous among the Emballonuridae havinglost one pair of lower incisors, while all emballonurids retain two upper premolars oneach side. The wing in the Emballonuridae also presents some modification, withlengthening of the third digit in Taphozous and Diclidurus, and, throughout thefamily, with the flexing forward of its proximal phalange. The head of the humerusis slightly elongated and oblate in the Emballonuridae, rather than rounded, andslightly tilted, features possibly indicating a degree of specialization as Smith(1972 : 29) suggested for the Mormoopidae. The Rhinopomatidae, with but a single genus, mostly inhabit arid or semi-desertregions in southern Asia and Africa : by contrast, the Emballonuridae are widelydistributed in the tropical and sub-tropical parts of the Old and New Worlds, withnumerous genera and species. The Craseonycteridae have a wing and shoulderstructure very different from either of these families, showing evidence of modifica-tion for specialized flight, most probably of a hovering nature. Nevertheless, thefamily retains the rhinopomatid organization of the anterior edge of the wing, to aconsiderable extent the architecture of the rhinopomatid skull, and especially therhinopomatid dentition, with the premaxillae modified in such a way as to representthe epitome of a tendency foreshadowed in the Emballonuridae. Free premaxillaeare common to all three families and are found (as the palatal rather than the narialbranches) also in the Nycteridae, Rhinolophidae and Hipposideridae. They may wellprovide a flexibility contributing to a feeding mechanism that requires considerable 334 J- E. HILL mobility of the upper lip and anterior mouth and are possibly of value to batsthat glean insects. The enlarged trochiter of the Craseonycteridae suggests con-ventionally a relatively advanced bat, yet in the absence of an evident deltoid ridgeor a definite deep supraglenoid pit it differs from the other families so considered,although in some of these the ridge may be displaced dorsad, and in some the supra-glenoid pit is not exceptionally developed and is displaced dorsad to lie between thebase of the humeral head and the trochiter, as in the Craseonycteridae. The devel-opment of an epitrochlear process seems a corollary of a need to operate and controla long wing with a long tip. My view is, therefore, that this new and striking familyof bats represents a further branch of the Emballonuroidea, modified for a sophis-ticated style of flight, and thus combining some external, cranial and dental featuressimilar to those of the Rhinopomatidae arid Emballonuridae with some of the wingand skeletal features of those families considered conventionally more advanced.Such a combination is perhaps inevitable when the familial classification of theOrder is based to some extent on modifications which improve the flying mechanismor adapt it to particular kinds of flight, reflecting Miller's dictum quoted above thatin a group of characteristically volant animals the greatest taxonomic importancemust be placed on the development of the wings. ACKNOWLEDGEMENTS The late Kitti Thonglongya first involved me in the examination of this new batand initiated the study of the specimens that he had obtained. The Board of theApplied Scientific Research Corporation of Thailand, through the Director of theCentre for Thai National Reference Collections, Dr Prasert Lohavanijaya, generouslyagreed that the work, begun as a joint enterprise, should be completed at the BritishMuseum (Natural History) and made a series of specimens available to supplementthe initial sample first brought to London by Mr Lim Boo Liat. My thanks are dueto Dr H. Elliott McClure, of the United States Army Medical Component, SouthEast Asia Treaty Organization, who helped in these arrangements after the death ofKitti Thonglongya, and to Dr R. Askins, then of the Association for the Conservationof Wildlife (ACW), Thailand, for his courtesy in informing me of that event. I amindebted to Dr J. T. Marshall, also of the United States Army Medical Component,who was instrumental in sending the major series of specimens to me in London, andfor obtaining information and locational data necessary to the description of thenew bat. SUMMARY A new family of bats, the Craseonycteridae, is proposed in the Microchiropterafor a new genus and species from Thailand, described as Craseonycteris thonglongyai.The new family is allied to the Rhinopomatidae and Emballonuridae and is placedin the Emballonuroidea. Its diagnostic features are reviewed in detail, with adiscussion of its taxonomic position. A NEW BAT FROM THAILAND 335 REFERENCES ANDERSEN, K. 1905. On the bats of the Rhinolophus philippinensis group, with descriptions of five new species. Ann. Mag. nat. Hist. (7), 16 : 243-256, i fig., I tab.DOBSON, G. E. 1878. Catalogue of the Chiroptera in the collection of the British Museum. London.FELTEN, H., HELFRICHT, A. & STORCH, G. 1973. Die Bestimmung der europaischen Fleder- mause nach der distalen Epiphyse des Humerus. Senckenberg. biol. 54 : 291-297, 19 figs.FINDLEY, J. S., STUDIER, E. H. & WILSON, D. E. 1972. Morphologic properties of bat wings. /. Mammal. 53 : 429-444, 5 figs., 4 tabs.HARRISON, D. L. 1964. 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(Eds.), About Bats. A Chiropteran Biology Symposium, 51-83, 5 figs. Dallas. SMITH, J. D. 1972. Systematics of the Chiropteran family Mormoopidae. Misc. Publs Mus. nat. Univ. Kans. No. 56 : 1-132, 40 figs.THONGLONGYA, K. 1972. A new genus and species of fruit bat from South India (Chiroptera : Pteropodidae). /. Bombay nat. Hist. Soc. 69 : 151-158, 2 tabs., 6 pis. 1973- First record of Rhinolophus paradoxolophus (Bourret, 1951) from Thailand, with the description of a new species of the Rhinolophus philippinensis group (Chiroptera, Rhinolophidae). Mammalia 37 : 587-597, 2 figs., 2 appendices. & HILL, J. E. 1974. A new species of Hipposideros (Chiroptera) from Thailand. Mam-malia. 38 : 285-294, 2 figs., 2 tabs.VAUGHAN, T. A. i97oa. 3. The skeletal system. In WIMSATT, W. A. (Ed.), Biology of Bats, I : 97-138, 23 figs. New York & London.— I97ob. 5. Flight patterns and aerodynamics. In WIMSATT, W. A. (Ed.), Biology of Bats, I : 195-216, 4 figs., 3 tabs. New York & London.WALTON, D. W. & WALTON, G. M. 1970. Post-cranial osteology of bats. In SLAUGHTER, B. H. & WALTON, D. W. (Eds.), About Bats. A Chiropteran Biology Symposium, 93-126, 6 figs. Dallas. WINGE, H. 1892. Jordfundne og nulevende Flagermus (Chiroptera) fra Lagoa Santa, MinasGeraes, Brasilien. Med Udsigt over Flagermusenes indbyrdes Slaegtskab. K0benhavn. 336 J. E. HILL WINGE, H. 1923. Pattedyr-Slaegter. I. Monotremata, Marsupialia, Insectivom, Chiroptera, Edentata. Kobenhavn. — 1941. The Interrelationships of the Mammalian Genera (Ed. JENSEN, A. S., SPARCK, R.& VOLS0E, H., translated DEICHMANN, E. & ALLEN, G. M.). /. Monotremata, Marsupialia,Insectivora, Chiroptera, Edentata. K0benhavn. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD A LIST OF SUPPLEMENTSTO THE ZOOLOGICAL SERIES OF THE BULLETIN OFTHE BRITISH MUSEUM (NATURAL HISTORY) 1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum(Natural History) described by William Harper Pease. Pp. 96; 14 Plates.1965. (Out of Print.) £3.75.' 2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier andValenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure andMineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ;29 Plates, 77 Text-figures. 1969. £4.50. 4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80. 5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa: theBiology and Evolution of a Species Flock. Pp. 134; i Plate, 77 Text-figures.1974- £375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU