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.) S.RM 1035^ GtNERAL BULLETIN OF I tsmm LIBRARY THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 28 1975 BRITISH MUSEUM (NATURAL HISTORY)LONDON: 1977 DATES OF PUBLICATION OF THE PARTS No. i. . . . . . .19 May 1975 No. 2 . . . . . .21 May 1975 No. 3 . . . . .29 May 1975 No. 4 . . . . .29 May 1975 No. 5 . . . -17 September 1975 No. 6 . . . .5 November 1975 No. 7 . . . .5 November 1975 No. 8 . . . . .16 December 1975 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU CONTENTS ZOOLOGY VOLUME 28 PAGE No. i. A guide to the species of the genus Euplotes (Hypotrichida, Ciliata).By C. R. CURDS ......... i No. 2. Catalogue of the types of terrestrial isopods (Oniscoidea) in thecollections of the British Museum (Natural History) II. Oniscoidea,excluding Pseudotracheata. By J. P. ELLIS and R. J. LINCOLN . 63 No. 3. The larval development of Carcinus maenas (L.) and C. mediter-raneus Czerniavsky (Crustacea, Brachyura, Portunidae) reared inthe laboratory. By A. L. RICE and R. W. INGLE . . . 101 No. 4. A comparative study of the larval morphology of the British por-tunid crabs Macropipus puber (L.) and M. holsatus (Fabricius),with a discussion of generic and sub-familial larval characters withinthe Portunidae. By A. L. RICE and R. W. INGLE . . . 121 No. 5. Miscellanea Streptaxidae from Aldabra Island, Western Indian Ocean. By A. C. VAN BRUGGEN 157 Quickia aldabraensis, a new species of land snail from Aldabra Atoll,Western Indian Ocean. By C. M. PATTERSON .... 177 Notes on some echinoderms from Marion Island. By F. W. E.ROWE and A. M. CLARK ........ 187 A new species of Tilapia in the Zambian Zaire system. By E.TREWAVAS and D. J. STEWART ....... 191 Two new nematodes parasitic in the kiwi in New Zealand. By E. A. HARRIS .......... 199 Description of Pembatoxon insulare gen. n. sp. n. from PembaIsland. By J. VAN GOETHEM ....... 207 A quagga, Equus quagga, at University College, London and a noteon a supposed quagga in the City Museum, Bristol. By A. W.GENTRY ........... 217 A new angelfish of the genus Centropyge from Ascension Island. By R. LUBBOCK and R. D. SANKEY 227 A new species of Nanochromis from the Ogowe System, Gabon. By E. TREWAVAS 233 The first zoeal stages of Cancer pagurus L., Pinnotheres pisum(Pennant) and Macrophthalmus depressus. By A. L. RICE . . 237 No. 6. The hydroid species of Obelia (Coelenterata, Hydrozoa: Cam-panulariidae), with notes on the medusa stage. By P. F. S.CORNELIUS 249 No. 7. Some new and rare species of calanoid copepods from the north-eastern Atlantic. By H. S. J. ROE ...... 295 No. 8. A revision of the species of Lafoeidae and Haleciidae (Coelenterata:Hydroida) recorded from Britain and nearby seas. By P. F. S.CORNELIUS . . . . . . . . . 373 Index ........... 427 INDEX TO VOLUME 28 The page numbers of the principal references and the new taxonomic names are printed in bold type. abbreviatus, Haplophthalmus . . 71 aberrans, Euplotes . . -9, 51, 52 abietina, Abietinaria . . . 379, 380abietina, Campanularia . . . .381 abietina, Grammaria .381, 382, 383, 384, 385abietina, Lafoea .... 379, 381 abietina, Reticularia .... 382 abietina, Salacia . . . . .382 Abietinaria .... 379, 380, 386 abyssalis, Spinocalanus . 297, 298-300, 304 abyssalis pygmaeus, Spinocalanus . . 298acuta, Charybdis .... 146, 147 adelungi, Obelia ..... 266 adonis, Hyloniscus . . . .71 adriatica, Halophiloscia .... 82 adriatica rupium, Halophiloscia . . 82aediculatus, Euplotes . 6, 8, 16, 17-i8, 26, 28Aetideopsis ..... 305, 306 Aetideus ..... 304-305 Aetiteidae ..... 304-311 affinis, Euplotes . . . 5, 16, 18-19, 24affinis, Philoscia . . . . .87 affinis, Philoscia muscorum ... 87affinis tricciratus, Euplotes . . .18agilis, Xanthocalanus . . . 311, 312alata, Metridia . . . 342, 343, 345alatus, Euplotes . . . . 16, 19, 20 alatus, Euplotes patella . . . -41alauda, Ennea .... 158, 160 alba, Pherusa ..... 73 albicincta, Bilawrencia ... 80 albus, Titanethes ..... 73 aldabrae, Buliminus . . . 171 aldabrae, Gulella gwendolinae 158-164, I 7 I 172, 173aldabrae, Rhachis . . . . .171 aldabraensis, Quickia . . 177-i86, i pi.Allocentrotus . . . . -37 Allocentrus . . . . . .22 Alloniscus ..... 78, 82, 97 alpinus, Androniscus .... 70 Alpioniscus ...... 69 altera, Amallophora . . 320-333, 336alternata, Eucope . . . . .273 alternata, Obelia ..... 267 Amallophora . 330, 332-333, 336, 339, 341 Amallothrix . 318, 320, 325-335, 341, 365, 367amieti, Euplotes . . . -17, 19-21Amphisbetia . . . . -379 Amphiura ..... 188-189 Anasterias . . . . . .190 Anchiphiloscia ..... 79 andersoni, Obelia . . . 260, 264, 265 Androniscus .angulosa, Obelia .angusta, Niambia .angusticauda, Philoscia .angusticauda, Setaphoraangusticeps, Haloptilus .angusticeps, Spinocalanusangustissima, Pseudophilosciaannulata, Charybdisannulata, Eucope .annulatum, Heleciumannulipes, Ucaanomala, Philosciaanomalus, Phalloniscusantarcticus, Euplotesantennarius, Canceranthonyi, Cancer .antiquorum, Equus burchelli . 70. 266 779191 345, 347297, 298, 33, 34. 89146, 147. 280 396 245. 87. 87 . 16, 21-22 239238, 239, 246 224 Aphiloscia . . . . -79, 88, 98 apsheronicus, Euplotes . . . 38-39 apterycis, Ascaris . . . . .199 apterycis, Cyrnea . 201-205, Pis. 1-2 apuanus, Haplophthalmus . . .71Arborcinia . . . . . .184 arboreum, Halecium .... 410 arcuatus, Aetideus . . . 304-305 arcuatus, Snelliaetideus .... 304 argi, Centropyge .... 227, 230 Arhina ...... 78 Armadilloniscus . . . . 79, 98 armata, Deto . . . . .81 armata, Paraphiloscia .... 85 arruensis, Obelia ..... 266 articulata, Eucope .... 266 articulata, Obelia . . . . .266 articulosum, Holecium . . 392, 406, 409Ascaris .... 199 aspinosa, Scolecithricella . 321, 322, 323Assiminea . . . . . .166 Asteroidea . . . . . .190 Atelecylus 240 atlanticus, Euaugaptilus . . 349-351 Atoxon 207, 215 Atractylis 4 12 attenuata, Obelia . . 260 Augaptilidae . . . 346-36 1 aurantonotus, Centropyge . . 227, 230auropecten, Amallothrix. 33O-333, 334~33 6 auropecten, Scolecithricella (Amallothrix) 330, 333 auropecten, Scolecithrix . . .321, 333australis, Obelia . . . 266, 271, 272australis, Styloniscus .... 68 428 INDEX australis, Trichoniscusaustriacus, Trichoniscus .austroafricanus, Styloniscusaustroafricanus, Trichoniscusaustrogeorgiae, Laomedeaaustrogeorgiae, Obelia 68 73. . . .68 68 . 280260, 265, 266, 280 backusi, Scottocalanus . . . -313 baloni, Tilapia .... 191-igS balsii, Japanoniscus . . . .83 balssi, Chaetophiloscia . . . .81 balteatus, Euplotes . . 4, 9, 16, 22-23 balticus, Euplotes . . . . 11, 12 balticus, Euplotes vannus . . .11 bargensis, Tiroloscia squamuligera . . 93 Bathynectes . Bathypontia Bathypontiidae Bathytropa . beanii, Halecium . beanii, Thoa . bensoni, Quickia bensoni, Succinea . Benthana Benthanops . beramporia, Heterakis bernardii, Succinea bicolor, Rhyscotus .... 76 bicuspidata, Gonothyrea . . . 260 bicuspidata, Laomedea . . . 260, 261 bicuspidata, Obelia . 260, 261, 262, 264, 265 bicuspidata picteri, Laomedea . . 260, 264 bicuspidata tenuis, Laomedea . bidentata, Laomedea 127, 142, 143, 144, 145 361-364 361-3&9 . 79,83,98 391-393, 406, 408 391 180, 183 . 180 80 88 2OI. 184 260, 264260 bidentata, Obelia 251, 253, 254, 255, 256, 257, 258, 259, 260-265, 278, 279, 280 biellensis, Philoscia muscorum . . 88 bifurca, Obelia ..... 260 biguttata, Xavia . . . . .145 biguttatus, Portumnus . . . .145 billardii, Halecium . . . 412, 413 billardii exigum, Halecium . . .412Bilawrencia ...... 80 bimaculata, Charybdis . . . .146 biserialis, Obelia ..... 267 bistriata, Laomedea . . . .281 bistriata, Obelia . . . . .281 bisulcatus, Euplotes . . . 16, 23 blandfordi, Dotilla . . . 244, 245 bodkini, Calycuoniscus . . . .81 bogorovi, Scaphocalanus . . . 325 boreale, Helecium . . . . .391 borealis, Obelia ..... 266 bosniensis, Trichoniscus .... 73 bougainvillei, Papuaphiloscia ... 84Brachyura . . 101-120, i PI., 237-247 brasiliensis, Campanularia . . . 265brasiliensis, Obelia .... 267 braziliensis, Obelia. .... 267 brembana, Tendosphaera ... 76brentanus, Androniscus .... 70 breuili, Iberoniscus .... 72 brevicaudatus, Spinocalanus 297, 298, 299, 300brevicornis, Pseudophiloscia ... 89brevicornis, Valdiviella . . 309, 310, 311brevicyatha, Grammaria abietina . 382, 385brevis, Alloniscus ..... 78 briani, Philoscia squamuligera . . 93 briani, Tiroloscia squamuligera . . 93bruggeni, Atoxon .... 207, 215 brunnea, Niambia ..... 77 Buddelundiella ..... 74 Buddelundiellidae .... 65, 74 Bugula . . . . . .411 bulgarica, Bureschia .... 70 Buliminus . . . . . .171 Bulinus ...... 163 Burchella 183 burchelli, Equus . . 218, 220, 221-224 burchelli antiquorum, Equus . . . 224burchelli burchelli, Equus . . .224Bureschia ...... 70 Burmoniscus ..... 80 caciniformis, Hydrodendron . . .414caciniformis, Ophiodes . . . .414caciniformis, Ophidissa . . . .414caciniformis, Ophiodissa . . 415, 417 calcivagus, Androniscus .... 70calcuttensis, Quickia . . 180, 183, 184 Calicella 390 callianassa, Charybdis . . . 146, 147Callinectes . . . . . 146, 147 Calmanesia . . . . . .81 Calycuoniscus . . . . .81 Campalecium .... 390, 391 Campanularia 251, 252, 253, 254, 256, 257, 260, 261, 263, 265, 266, 272, 273, 278, 279, 280, 281, 378, 381, 382, 385, 386, 390 Campanulariidae . . 249-293, 377, 390 canariensis, Scolecithricella . . 323-325 Cancer 237-240 capensis, Niambia ..... 77 capensis, Paranotoniscus ... 68 Caphyrinae . . . . . .127 Capsularia . . . . . -378 Carcinus . . 101-120, i PI., 123, 145 Carcininae . 127, 144, 145, 147, 148, 149, 150caribbeanensis, Centropages . . 344~345carinata, Aetideopsis . . . 305, 306carniolense, Lepidoniscus germanicus . 84carynthiacus, Androniscus ... 70cassivelaunus, Corystes. . . 103, 239 castellana, Campanularia . . . 280castellata, Campanularia . . .280 castellata, Obelia . . . . .280 casuarii, Cyrnea .... 204, 205 cateractae, Buddelundiella ... 74 INDEX 429 Catinella .... Catoptrinae .... caudatus, Platyarthrus . caudatus squamatus, Platyarthrus caulini, Campanularia cavalliensis, Nanochromis cavernarum, Androniscus cavernarum strasseri, Androniscus cavernicola, Trichoniscus cavolinii, Campanularia . cavolinii, Sertularia Cellaria .... cellaria, Chaetophiloscia cellaria, Philoscia . Centropages Centropagiidae Centropus .... Centropyge .... cestus, Styloniscus cestus, Trichoniscus Chaetodon .... Chaetophiloscia charon, Euplotes . Charybdis Chavesia chinensis, Obelia . Chiridiella Cichlidae Ciliatea cinerascens, Ligia . cingulata, Philoscia cingulata, Setaphora circularis, Schoblia cithara, Ploesconia Clark, A. M. Clavigeroniscus . 127. 78 78265, 272 234 7070 73 265, 272, 273. 265409, 4118181 . 344. 344204 227-231, i PI.6868230 . 81, 88, 9817, 18, 22, 24, 49146, 147, 14871 . 266306-307 191-197, 233-235. 1-61749191 . 6929187-igo 66-67 Clytia 253, 254, 260, 264, 265, 279, 280, 281 coeca, Philoscia . . . . .91coeca, Setaphora . . . . .91Coelenterata .... 249-294, 375commensalis, Trichoniscus commissuralis, Obeliacomorensis, Gulellacompar, Alloniscuscompta, Philoscia .comta, Setaphora .concisa, Quickiaconcisa, Succinea .congdoni, Laomedeacongdoni, Obelia .contractus, HiatoniscusCopepodaCoptodonCorallinaCordioniscus .Cornelius, P. F. S. .cornuta, Lafoeacornutus, Alloniscuscorona, ObeliaCorsica, Philoscia . 73 265, 271, 272169 . 789191 180, 182, 183, 184 . 182 267 266, 267 . 83 295-372 194, 196 393. 67 249-293, 373-426 . 385, 386 . 78 260 92 Corsica, Tiroloscia .... 92, 93 corsicus, Nesiotoniscus . . . .72 corsicus, Nesiotoniscus corsicus . . 72corsicus, Trichoniscus .... 72 corsicus corsicus, Nesiotoniscus . . 72coruscans, Campanularia . . .278 Corystes ..... 103, 239 costata, Bathytropa .... 79 costata, Bathytropa meinerti ... 79Costigulella . . . . . .167 costulata, Chavesia . . . .71 costulatus, Platyarthrus ... 78 couchi, Halophiloscia . . . 82, 88 coughtreyi, Obelia .... 267 crassa, Ploesconia . . . . .11 crassicornis, Hyloniscus . . . .71 crassus, Euplotes .... 11-12 crenata, Thalamita . . . 146, 147 crenatum, Halecium .... 396 crenosus, Euplotes . . .16, 24-25 crinitus, Macrophthalmus . . 244, 245cristatus, Euplotes . . n, 12-13, 34 Crustacea 63-101, 101-120, i PL, 121-151, 237-247, 295-372cubensis, Rhyscotoides .... 76 cubensis, Rhyscotus .... 76 Cucumaria ..... 187, 188 cunningtoni, Anchiphiloscia . . . 79cupressina, Sertularia . . . 380, 405Curds, C. R. . . . . l-6i cursorium, Ligidium 75 Cyphonetes ...... 73 Cyphoniscellus . . . . 7 1 Cyrnea . . . 201-205, Pis. 1-2 dahli, Titanethes . . -73 daidaleos, Euplotes . . 4, 38, 39, 40 dalmatica, Philoscia . . -87 dalmatica, Philoscia muscorum . . 87 dalmatica, Stenophiloscia . 92 dalmaticus, Hyloniscus . . 7 1 dalmatinus, Armadilloniscus . . 79 damae, Hora . 83 danicus, Haplophthalmus . 71, 79 debilis, Ischioscia ... -83 debilis, Philoscia ... -83 Decapoda . . 237-247 deliculata, Obelia . 280 demarcata, Philoscia . 9 1 demarcata, Setaphora . 9* dentata, Charybdis six- . . 146, 147 dentata, Scolecithricella . 34 1 denticulata, Campanularia . . 279, 280 denticulata, Scolecithricella . . 327denticulatus, Lepidoniscus pruinosus . 84 dentiens, Gulella ... 173 dentiger, Androniscus . 7 dentiger ligulif er, Androniscus . . 7 dentipes, Ligia 75 430 INDEX depressa, Kogmania .... 69depressifrons, Portunus . . . 146, 147depressus, Macrophthalmus . 237, 242-246 Deto 81-82,98 Detonella ...... 98 Diacara ...... 82 Diaixidae ...... 367 diaphana, Eucope . . . . .273 diaphana, Thaumantias . . . .273 dichotoma, Campanularia . . . 265dichotoma, Laomedea . . . 265, 267dichotoma, Obelia . 251, 253, 254, 255, 256, 257,258, 259, 266-272, 273, 277, 278, 279, 281dichotoma, Sertularia . . 253, 254, 265dichotomum, Halecium . . . .410 Didima ...... 82 difficilis, Scaphocalanus . 815-317, 333 dilatatus, Macrophthalmus . . . 245dilectum, Philoscia .... 88 dimidiatus, Nanochromis . . . 235 diminuta, Philoscia .... 88 Diphasia ...... 410 divaricata, Laomedea . . . 265, 266divaricata, Obelia ..... 266 dogieli, Euplotes . . . .16, 25-26 dolinensis, Haplophthalmus fiumaranus . 71dolomiticus, Oroniscus .... 84 dorsalis, Chaetophiloscia . . . .81 dorsispinosus, Spinocalanus . . . 303Dotilla ..... 244-246 droebachiensis, Strongylocentrotus . . 23 Dromia dubia, Amallophora dubia, Heteramalla dubia, Hetermalla . dubia, Obelia dubia, Scopalatum. dumosa, Campanularia dumosa, Capsularia dumosa, Lafoea . . . dumosa, Sertularia . . Dynamena . echinata, Sertularia echinatus, Scaphocalanus Echinoderms. echinoides, Strongylocentrotus edulis, Ostrea elbana, Tiroloscia . . . elbanus, Parastenoniscus . elbanus, Trichoniscus . . elegans, Alloniscus elegans, Centropages . . elegans, Diacara elegans, Euplotes . . . elegans littoralis, Euplotes elegans, Sagitta Ellis, J. P elongata, Chaetophiloscia . . 103 339-34 I339, 341-342 339266, 272 333, 336, 338, 341 385. 386 377,385-390... 385 415 273, 405 318187-190 23,37 394 93. 76 73. 82 344. 82 8, 47, 48. 48 27963-iooe 81,88 elongata, Philoscia elongatus, Euaugaptilus Ennea .... Equidae equilateralis, Obelia Equus . * . esterelana, Paraphiloscia esterelana, Philoscia esterelana, Tiroloscia Euaugaptilus . . 346 Eucope . 252, 254, 265 Eudendrium . Eulafoeinae . euplocami, Cyrnea Euplotes eurycerca, Cyrnea . Euryligia eurystomus, Euplotes . 5, eurystomus, Euplotes patellaeverta, Obeliaexigua exigua, Tirolosciaexigua, Tiroloscia exiguaexigum, billardi, Haleciumexigum, Halecium billardi 81 361 158, 160, 162 217-226, 4 Pis. 267 217-226, 4 Pis.939393 349,351,353-361 266, 271, 273, 280 398, 410 . 378 204 204 746, 8, 10, 16, 17, 19, 26-29, 41 26, 27 267 93 93 . 412. 412 fagorum, Stylohylea .... 72fagorum, Trichoniscus .... 72falcata, Hydrallmania . . . .380falcifer, Amallothrix . . 325, 327, 341falcifer, Scolecithrix . . . -325farciminoides, Salicornaria . . . 409fasciata, Philoscia . . . . .91fasciata, Setaphora . . . .91 fasciatum, Atoxon . . . 207, 215 fecundu, Euaugaptilus . . . 351-352fenestrata, Thuiaria . . . .381ferox, Gaetanus ..... 306ferrani, Scopalatum . . 336, 338, 342Filellum . . . 378, 379, 380, 381, 385filiforme, Halecium . 403, 405, 406, 409 fistulosa, Cellaria .... 409,411fiumaranus, Haplophthalmus . . .71fiumaranus dolinensis, Haplophthalmus . 71flabellata, Campanularia . . . 266 flabellata, Obelia . . . 266, 270, 271 flava, Philoscia 88 flavescens, Niambia .... 77 flavus, Oritoniscus . . . 72, 73 flavus, Trichoniscus .... 72 flexuosa, Campanularia . . 256, 257, 281flexuosa, Sertularia . . . .272 Flustra .... 387,412,414 foliacea, Flustra . . . . .412 formicarum, Niambia .... 77 formosana, Chaetophiloscia . . .81Formososcia ...... 82 foveolatus, Trichoniscus .... 74 fragilis, Allocentrotus .... 37 INDEX 431 fragilis, Allocentrus . . . .22 fragilis, Alpioniscus .... 69 fragilis, Obelia ..... 266fragilis, Pseudophiloscia .... 89fragilis rharelbazi, Trichoniscus . . 73fransiscanus, Strongylocentrotus . . 23frigidana, Philoscia muscorum . . 88 fruticosa, Campanularia .... 386fruticosa, Lafoea .... 386, 389fucorum, Halophiloscia .... 82fulva, Benthanops ..... 88fusiformis, Encope . . . 271, 273 fusiformis, Obelia . . . . .271 gaboniscus, Nanochromis . . 233-235Gaetanus . . . . . . 306 Gaidius ...... 306 gallinarum, Heterakis . . . .201 gangetica, Ilyoplax .... 245 Gastropoda . . 157-176, 177-186, i PL gaudichaudii, Octypode .... 245 gaussi, Obelia .... 273, 277 gelatinosa, Campanularia 265, 266, 279, 280, 281265265272272 251, 253, 254, 255, 256, 257,258, 259, 271, 272-278geniculata, Sertularia . . . 265, 272geniculata subsessilis, Obelia . . .275geniculatum, Halecium . . 393, 396, 409Gentry, A. W. . . 217-226, 4 Pis. Geologia ...... 75 georgensis, Styloniscus .... 68 georgensis, Trichoniscus .... 68 germanicus carniolense, Lepidoniscus . 84gibbera, Scopalatum . 335, 386-338, 342 gibbosulus. Cancer .... 239 glarearum, Stenophiloscia ... 92globiceps, Rhyscotus .... 76 globosus, Scopimera . . . .245 Gonotha ..... 260, 264 Gonothyrea . . . 253, 260, 265, 266, 281 gelatinosa, Laomedeagenicolata, Sertolaregeniculata, Campanulariageniculata, Laomedeageniculata, Obelia gottscheensis, Cyphoniscellusgracilicauda, Heterakisgracilicornis, Halophilosciagracilipes, Ligiagracilis, Campanulariagracilis, Euplotes .gracilis, Laomedeagracilis, Obeliagracillima, Campanulariagracillima, Lafoea .graecus, Labyrinthasius .Grammaria .granulata, Bathytropa .granulatus, Tylos . 71199-201 82 75 . 281 8, 47, 48-49 . 266 266, 281 . 386, 387 386, 389, 390 . 83 378, 379, 381- 3 8 579, 83. 65 granuliferus, Tylos .... 65 graphophasiani, Cyrnea .... 204grarosensis, Philoscia .... 88gravieri, Clytia . . . . .281 grevyi, Equus . . . . .218 grimni, Obelia ..... 266guinasana, Tilapia . . . .196 griseoflavus, Niambia .... 77griseus, Hiatoniscus .... 83guernei, Philoscia ..... 88Gulella ..... 158-173 gwendolinae, Ennea . . . 158, 162gwendolinae, Gulella 158, 162, 163, 164, 171 gwendolinae, Gulella gwendolinae . .163gwendolinae aldabrae, Gulella 168-164, 171, 172, 173 gwendolinae gwendolinae, Gulella . .163gwendolinae mkusiensis, Gulella . 162, 163gwendolinae porrecta, Gulella . . 162, 163gwendolinae scissidens, Gulella . 162, 163gwendolinae tsadiensis, Gulella 158, 162, 163gymnothalma, Obelia . . . .273 375-376, 390- 4 I5393 . 393-396, 399, 49. 4 12391-4U, 412, 414 391, 399 82, 88 72345, 347-349 Haleciidae halecina, Sertularia halecinum, Halecium, Halecium Haloikema .... Halophiloscia halophilus, Miktoniscus . Haloptilus .... hamuligerus, Androniscus roseus . . 70 Hanoniscus ..... 82, 98 Haplophthalmus . . . . 7 1 - 79, 97 harpa, Euplotes . . . . 17, 29, 30 Harris, E. A. . . . 199-2O5, 2 Pis. hastata, Chaetophiloscia . . .81 Hebella 377, 39<> Hebellidae . . -377 hedwigae, Gulella . . . . .167 helecina, Sertularia . . . 39 X 393 helgolandica, Obelia . . .266 Hemiplax ...... 245 hemisphaerica, Medusa .... 254 hendersonae, Clytia . . .281hercegowinensis, Cyphonetes ... 73heroldi, Illyrionethes .... 7 heroldi, Isabelloscia .... 83 herzegowinense, Ligidium 75 Heterakidae 199-201 Heterakis ... . 199-201 Heteramalla .... 339, 34 * 342 Hetermalla ... -339 Heterorhabdidae . . 346 Heterorhabdus . . . 345, 346 Hiatoniscus ...... 83 hirsuta, Halophiloscia . . .82 hirsuta, Nahia . . 84, 88 hirsuta, Niambia ..... 77 432 INDEX hirsuta, Philoscia .hirtipes, Hemiplaxhispana, BathytropaHolothurioideaholsatus, Macropipusholthuisi, Papuasoniscushoplites, SpinocalanusHora ....horae, Styloniscus .horae, Trichoniscushorridus, Spinocalanus .hottentoti, Styloniscus .hottentoti, Trichoniscus .humilis, Didimahyalina, Gonothyreahyalina, ObeliaHydrallmaniaHydranthea .HydrodendronHydroidaHydrozoaHyloniscus .hyperboreus, Euaugaptilushypnorum, LigidiumHypotrichida Iberoniscus . Idiella . ignota, Valdiviella Illyrionethes Ilyoplax immersa, Reticularia, imperfecta, Valdiviella indentatus, Euplotes Indoniscus Indosuccinea inflatus, Hyloniscus inflexa, Pseudophiloscia Ingle, R. W. inkystans, Euplotes insignis, Grammaria insignis, Valdiviella . 84 245 79 187-188 121-152 . 85 298, 303, 304 . 83 68 68 297. 303. 304686882 . 266 265, 266, 267 380 390, 412-414 4*4 375249-293 71-72, 73, 97 359, 360-361 75 . 1-61 72 . 411 . 310 70 244246 378, 379, 380 . 310 8, 45-46 . 67 . 183 71 insulanus, Trichoniscus noricusinsulare, Pembatoxoninsulincola, Gulellaintermedia, Bathypontiaintermedia, Grammaria .intermedius, Scyphaxinvalidus, Scaphocalanusirregularis, Obelia .Isabella, BilawrenciaIsabelloscia .Ischioscia . . ,isolonche, HeterakisIsopoda 101-120, I PI., 121-151 . 17, 29-30382, 383, 385309, 310 jacquelinae, GulellaJapanoniscus 74 . 207-216, i PI.161, 168-169, 171, 172 363. 382. 384 91 317267 80 . 83. 83 2OI63-1006 1 66 83 japonica, Charybdisjaponicum, Nippoligidiumjaponicus, Charybdisjaponicus, Macrophthalmus karongae, Anchiphiloscia kempi, Burmoniscus kenepurensis, Oniscus kermadecensis, Styloniscus kermadecensis, Trichoniscus Kerona kincaidi, Campanularia . Kogmania Komatia Krantzia 149 75147245 7980846868 4 1280698983 396-399, 401, 411 83. 187 187, 188187-188 377. 379, 381, 385-390 375-37 6 , 377-390 406 272, 273, 278. 320 4*4312, 320, 325 . 167399-402 399 labrosum, HaleciumLabyrinthasiuslaevigata, Cucumarialaevigata, Pentactellalaevigatus, PseudocnusLafoea .LafoeidaeLafoeinalairii, Laomedealamellifer, ScolecithricellaLaminaria laminata, Scolecithricellalangi, Gulellalankesteri, Haleciumlankesterii, Haloikema .Laomedea 251, 252, 253, 254, 260, 261, 264, 265, 266, 267, 272, 273, 278, 280, 281lata, Manibia . . . . -77 lateralis, Paraphiloscia .... 85 lateralis, Pseudophiloscia ... 85latifrons, Euaugaptilus .... 353 latipes, Lophothrix . . . 313-315 latipes, Portumnus . . . .145 latissima, Euryligia .... 74 latreillis, Macrophthalmus . . . 245latum, Ligidium . . . . -75 latus, Euplotes .... 16, 30-3 1 latus, Euplotes patella . . . 30, 31, 41latus, Paranotoniscus .... 68 leachii, Chaetodon .... 230 Lepidoniscus .... 84, 98 Leptotrichus . . . . -77 Leucosiidae ...... 242 Lictorella . . . . . -378 Lictorellinae ...... 378 lighti, Halecium ..... 409 Ligia . . . 74-75 Ligidium ...... 75 Ligiidae 65, 74-75 ligulifer, Androniscus dentiger. . . 70Lincoln, R. J. . . . . 63-iooelinearis, Obelia . . . . .280 INDEX 433 linearis, Rhyscotoideslinearis, RhyscotusLithotis littoralis, Armadilloniscuslittoralis, Euplotes eleganslobophora, Amallothrix .longa, Obelialongicarpus, Mictyrislongicauda, Niambialongicirrhus, Euaugaptiluslongicirrus, Haloptiluslongicornis, Haloptilus .longicornis, Philoscialongicyatha, Clytialongicyatha, Gonothalongicyatha, Gonothyrealongicyatha, Laomedea .longicyatha, Obelialongifurca, Scaphocalanuslongipes, Bathynecteslongipes, Spinocalanuslongiseta, Euaugaptilus .longissima, Laomedealongissima, Obelia .longissima, Sertularialongitheca, Clytia .longitheca, Obelia .Lophothrix .loveni, GonothyreaLubbock, R.lubricata, Philoscialubricata, Setaphoralucifera, Charybdislucifera, Obelialymani, Amphiura . 76. 76. 184 79 . 48 329, 335, 336260, 263 245, 246 77355-357 348 348 88 260, 261, 262, 265 260, 264 260, 265 261 260, 261, 264, 280 316-317 . 142 . 301 35i265 256, 258, 271, 278 265260260 313, 321, 323, 325. 281 227-231, i PI.9292 146, 147 251, 272, 278, 279188 macchiae, Tiroloscia .... 93 macrocephala, Rennelloscia ... 89macrodactyla, Chiridiella . . . 306Macrophthalminae . . . 245, 246 Macropipinae . . . . .127 Macropipus . . . . 118, 121-151 Macrothalmus . . . 237, 242-246maculatus, Pinnotheres . . . 240, 242maenus, Carcinus . 101-I2O, i PL, 123, 145 maenus mediterranea, Carcinus . . 118maenus septentrionalis, Carcinus . . 118magellanica, Grammaria . 382, 383, 384 magister, Cancer . . . . .239 magnicirratus, Euplotes . . .16, 31-32magnus, Spinocalanus . . . 297, 298maior, Campanularia .... 265 Mammalia .... 217-226, 4 Pis. Manibia ...... 77 margarica, Atractylis . . . .412 margarica, Hydranthea . . . 412-414margaricum, Halecium . . . .412 marginata, Komatia . . . .89 marginata, Obelia ..... 280 margine papillosa, Niambia . . -77mariae, Hyloniscus . . . .71 Marina ...... 25^ marina, Deto ..... 82 marina, Obelia .... 253, 279 marina, Philougria .... 82 marionensis, Cucumaria serrata . 187, 188marionis, Nullamphiura . . . .188 marionis, Uca ..... 245 Marioniscus ...... 84 marmoreus, Macropipus . 124, 125, 126, 135-142martensi, Ophioglypha . . . .189 martensi, Ophiurolepis .... 189 mascarenensis, Succinea . . .182 mauritiensis, Styloniscus ... 68 mauritiensis, Trichoniscus ... 68maxillaris, Euaugaptilus. . . 353-355mediterraneus, Carcinus 101-I2O, i PI., 145 mediterranea, Carcinus maenus . . 118medius, Androniscus subterraneus . . 70medius, Spelaeonethes .... 73 Medusa .... 252, 253, 254 medusa, Marina ..... 253 medusiferum, Campalecium . . 391 meeusei, Chaetophiloscia . . .81 meinerti, Bathytropa .... 79 meinerti costata, Bathytropa ... 79melanocephala, Ligia .... 75 mendanai, Paraphiloscia ... 86 meridionale, Atoxon . . . 207, 215methueni, Calmanesia . . . .81 Metridia ..... 342, 344 Metridiidae ...... 342 microps, Manibia ..... 77 micros, Trichorhina .... 78 microtaenia, Gulella .... 167 microtheca, Obelia .... 267 Mictyridae ...... 245 Mictyris ..... 245, 246 Miktoniscus ...... 72 miles, Gaetanus ..... 306 mina, Philoscia ..... 92 mina, Setaphora ..... 92 minax, Uca ...... 245 minor, Bathypontia . . . .361 minor, Campanularia spinulosa . .261minor, Laomedea spinulosa . 260, 261, 264minor, Valdiviella .... 309-3 n minuscula, Pupa ..... 166 minuta, Euplotes . . . 9, n, 13-14 minutissima, Trichorhina ... 78mirabilis, Ophiodes . . . .414 mirabilis, Ophiodissa . . . 414-417Mirigulella . . . . . .167 mixtus, Euaugaptilus . . . 358-360mixtus, Trichoniscoides .... 73 mixtus, Trichoniscus 73 mkusiensis, Gulella gwendolinae . 162, 163modesta, Niambia ..... 77 modestus, Trichoniscoides . 73 434 INDEX modestus, Trichoniscus .... 73moebiusi, Euplotes 5, 8, 24, 42, 43, 47, 49-5OMollusca 157-175; 177-186, i PL; 207-216, i PI. monocellatus, Microniscusmonocellatus, StyloniscusMonosklera .montana, Plymophilosciamontanus, Paranotoniscusmontanus, Trichoniscus .montanus, Trichoniscus vividusmoruliceps, Styloniscus .moruliceps, Trichoniscusmultidentata, Obeliamuricata, Sertulariamuricatum, Haleciummurigatum, Haleciummurrayi, Styloniscusmurrayi, Trichoniscusmuscivagus, Trichoniscusmuscicola, Euplotesmuscorum affinis, Philosciamuscorum biellensis, Philosciamuscorum dalmatica, Philosciamuscorum, Euplotesmuscorum frigidana, Philosciamuscorum, Pogonoligia .muscorum triangulifera, Philosciamussaui, Clavigeroniscus 6868 252, 254, 273, 276. 8968737368 . 68. 260402402-405, 409 403. 69. 6974 - 8, 45, 46. 87 . 8747, 50-51 88 75 88 66-67 mutabilis, Euplotes J, 10, ii, 14, 15, 47 nacreus, Alloniscus .... 78 Nahia 84, 88 Nanochromis .... 233-235 nanum, Halecium ..... 399 narentanus, Hyloniscus .... 72 nasatus, Rhyscotus .... 76 natalensis, Ligia ..... 75 neapolitanus, Euplotes . . -17, 32, 33nematodes .... 199-205, 2 Pis. Nemertesia ...... 267 neozealandicus, Tylos .... 65 Neptunus . . . . . .125 Nesiotoniscus .... 72, 97 nevelli, Succinea . . . . .182 Niambia ...... 77 nigra, Obelia . . . . 278, 279 nigrocaulus, Obelia .... 266 Nippoligidium ..... 75 nitida, Philoscia ..... 88 nitida, Philougria ..... 88 nivatus, Trichoniscus .... 74 niveus, Tylos ..... 65 nodosa, Obelia ..... 266 noduliger, Androniscus subterraneus . 70 noliformis, Clytia . . . . .279 noricus insulanus, Trichoniscus . . 74noricus sassanus, Trichoniscus . . 74 noricus sturanus, Trichoniscus . . 74 nova brittanica, Rennellscia ... 89 novemcarinata, Euplotesnudiceps, Nanochromis .nudulus, TylosNullamphiurianyiroensis, Gullella pretiosa Obelaria . Obeletta . Obelia Obelissa obscura, Scolecithricella obtusa, Pseudochirella . obtusidens, Campanularia obtusidens, Obelia obtusidentata, Campanularia obtusidentata, Obelia ocellata, Formososcia ocellatus, Ovalipes ochotensis, Cyrnea octocarinatus, Euplotes . octocirratus, Euplotes Octypoda Octypode Octypodinae Olibrinus oligarthra, Valdiella Oniscidae Oniscoidea Oniscus opercularis, Tylos . operculata, Amphisbetia Ophiacantha Ophidissa Ophiodes Ophiodissa Ophioglypha Ophiuroidea . Ophiurolepis orientalis, Charybdis orientalis, Indoniscus Oritoniscus . ornatus, Paranotoniscus ornatus, Scyphax . Oroniscus ortonedae, Rhyscotoides ortonedae, Rhyscotus Ostrea .... ostreum, Pinnotheres otakensis, Styloniscus otakensis, Trichoniscus . Ovalipes 127, 142, 143, 144, ovata, Chiridiella . ovata, Setaphora . oxydentata, Obelia Oxyloma pacificus, Macrophthalmuspagurus, Cancerpallida, Chaetophiloscia .pallida, Niambia . 51-53 235. 65 188, 189. 167 . 252, 254252, 254249-293252, 254318-319 309 266, 272 267, 272272267 82 127, 142, 143, 144204 . 38, 39-41. 16, 32-33 245 245242, 244, 245 . 8 4. 310 65, 78-93, 97-99 , . 63-iooe . 84 66 379. 189. 414 414 390, 414-417 . 189 188-189 \ . 189 146, H7 . 67 72, 7368 91 . 84. 76. 76 394240, 242 . 69. 69 145, 147, 148, 149 306-307 92 260, 263 177 245237-240, 246 81 77 INDEX 435 pallidemaculata, Setaphora palmetensis, Niambia papillosa, Trichorhina papillosus, Alloniscus . . Papuaphiloscia Papuasoniscus parabyssalis, Spinocalanus parallelus, Rhyscotoides parallelus, Rhyscotus paralongicirrus, Haloptilus Paranotoniscus . . . Paraphiloscia . . . Parascaphocalanus parasitica, Eucope . . Parastenoniscus parkei, Euplotes . . . paru, Pomacanthus . . parvula, Campanularia . parvula, Lafoea patella, Euplotes . . 4, 7, patella, Kerona patella, Ploesconia patella, Trichoda . patella alatus, Euplotes . patella eurystomus, Euplotes . patella latus, Euplotes . patella planctonicus, Euplotes patella typicus, Euplotes patella variabilis, Euplotes patienci, Philoscia . Patterson, C. M. . paululus, Xanthocalanus pauper, Benthana . pauper, Philoscia . peakei, Gulella 161, 164-i68, pelagica, Campanularia . pelagicus, Portunus Pembatoxon Pentactella . perkinsi, Geologia . perkinsi, Ligia personata, Dromia Phaennidae . Phalloniscus . Pherusa Phialella Phialidium . Philaster Philoscia 79, 80, 81, 83, 84, Philougria phormianus, Styloniscusphormianus, TrichoniscusPhymatoniscus picteti, Laomedea bicuspidatapigmentata Ligia .pigmentatus, Olibrinus .pilosa, Setaphora .pinnata, ZygophylaxPinnotheres . 92 77 . 78. 78. 84 85 . 298 . 76 . 76 347-349 68 . 85-87, 93. 321 265. 76 6, 1 6,34 227, 230 . 386 39026, 30, 38, 41-42 4141 . 41 41 26, 27 . 30. 3L 4i41414188 177-186, i PI. . 364-367 79, 80 80 169, 170, 171, 172 266, 281 146, 147 207-216, i PI. 187, 188 75 75. 103 311, 341, 36987, 98 73 . 280. 281 2387-88, 91, 92, 93, 98,99 82, 88 . 69 . 69 72 260, 264 75 . 8492 378, 402237, 240242 pinnotheres, Pinnotheres . . . 242 Pinnotheridae . . . . .242piriformis, Obelia . . . . .266Pisces .... 191-197, 233-235 pisum, Pinnotheres . . 237, 240-242 placunae, Pinnotheres . . . 240, 242plana, Obelia ..... 266 plana, Thaumantias . . . .271planctonicus, Euplotes patella . . 41 Platyarthrus .... 78, 97 platycephala, Ligia .... 75 platycephala, Pogonoligia 75platytarsis, Octypode .... 245pleonalis, Stenoniscus . . . .76plicata, Obelia ..... 266Ploesconia . . . . n, 29, 41 plumipes, Euplotespolygena, Eucope .polystyla, Eucope .plumosa, Blugula .plumosum, HaleciumPlymophilosciapocillum, Hebella .pocillum, Lafoea .pocillum, Lafoea fruticosaPodophthalminae .poecila, Krantzia .Pogonoligia .poljanskyi, Euplotes 5, 26, 27 273. 271. 411 405, 406, 408, 409. 89 390386, 390 . 386, 388 127 . 83 75 16, 25, 34-35, 5i Polybiinae 127, 143, 144, 145, 147, 148, 149 polycarinatus, Euplotes . . 8, 17, 35, 36 polyspina, Pseudochirella . . . 309Pomacanthidae . . . 227-231, i PI.Pomacanthus .... 227, 230 ponticus, Tylos ..... 66pooensis, Gulella .... 166, 167porcellioides, Alloniscus .... 78porcellioides, Arhina .... 78porrecta, Gulella gwendolinae . . 162, 163 PortumnusPortunidaePortuninae Portunuspoutrini, Gulellapretiosa, Gulella pretiosapretiosa nyiroensis, Gulellapretiosa pretiosa, Gulellaprinceps, Metridia .pristis, Idiella .Procyrnea . . . . .productus, Cancer .profunda, Amallothrix .prolifera, Campanularia .prolifera, Sertulariapropinqua, Paraphilosciapropinquus, Euaugaptilusprovisorius, Trichoniscus pusilluspruinosus denticulatus, LepidoniscusPseudocnus . 145 IOI-I2O, I PI., I2I-I5I127, 142, 144, 146, 147, 148, 149,150 123, 125, 142, 146, 147169167 . I6 7. I6 7 344. 411 204, 205 239320, 321 272 272, 27386358, 360 74 . 84187-188 436 INDEX PseudochirellaPseudophilosciapteronus, SpinocalanusPtychotremapuber, Macropipuspugilator, Ucapugnax, Ucapulchella, PhilosciaPulmonata 157-176, pumila, Dynamena punctatus, Ovalipes Pupa ..... purpuratus, Strongylocentrotus pusilla, Monosklera pusilla, Niambia pusillum, Eudendrium . pusillum, Halecium pusillus, Ilyoplax . pusillus, Trichoniscus pusillus provisorius, Trichoniscus pygmaea, Lafoea . pygmaea, Obelia . pygmaea, Philoscia pygmaeus, Phalloniscus . pygmaeus, Spinocalanus abyssalis pygmaeus, Trichoniscus . pyrenaeus, Oritoniscus . pyrenaica, Philoscia pyrenaica, Tiroloscia pyriformis, Eucope pyriformis, Obelia . 307-30985,89 298, 303 . i66 . 121-152 245 245 88 177-186, i PL,207-216, i PL 415127, 144, 147 . 166 23, 37 273, 276 77 398 396 245 7474 390. 266. 87. 87. 298 74 72 9393 . 266266 quadrata, Octypode .... 245quagga, Equus . . . 217-226, 4 Pis.Quickia .... 177-186, i PL quinquecarinatus, Euplotes . .16, 86-36 racemosa, Obelia ..... 267 rafflesi, Philoscia . . . . .92 rafflesi, Setaphora ..... 92 raikovi, Euplotes . . . 6, 38, 42, 51 ramosa, Grammaria . . . 382, 384ramosum, Schizocladium . 266, 272, 273 rariseta, Euplotes . . . 38, 4243, 49reflexum, Halecium . . . 396, 398refugiorum, Hyloniscus .... 72 rendalli, Coptodon .... 196 rendalli, Tilapia ..... 196 Rennelloscia ..... 89-91 resplendens, Centropyge . 227-231, i PLReticularia . . . 378, 380, 381, 382 Rhachis ...... 171 rharelbazi, Trichoniscus fragilis . . 73rhunicola, Obelia ..... 266 Rhyscotidae ...... 76 Rhyscotoides ..... 76 Rhyscotus ...... 76 Rice, A. L. 101-I2O, i PL, 121-151, 237-247rigidus, Euaugaptilus . . . 351, 357riparius, Hyloniscus .... 73riversdalei, Styloniscus .... 69riversdalei, Trichoniscus ... 69 robusta, Deto ..... 82robusta, Grammaria . . . 381, 382robustipes, Amallothrix . . . 365, 367robustum, Halecium . . 399, 401, 402 Roe, H. S. J 295-372 rosacea, Diphasia . . . . .410roscoffensis, Euplotes .... 53roseus, Androniscus .... 70roseus hamuligerus, Androniscus . . 70rotunda, Euplotes .... $3-54 rotundatus, Atelecyclus .... 240 Rowe, F. W. E 187-I9O rupicola, Anasterias . . . .190rupicola, Lithotis . . . . .184rupium, Halophiloscia adriatica . . 82ruweti, Tilapia .... 194, 195 Sabellaria ...... 408 Sagitta ...... 279 Salacia ..... 381, 382 Salicornaria ...... 409 sancristobali, Paraphiloscia ... 86sanguinolentus, Portunus . . 146, 147Sankey, R. D. . . . 227-231, i PL santaisabellae, Paraphiloscia . . .87sapidus, Callinectes . . . 146, 147 sargassi, Laomedea .... 267 Sarotherodon . . . . .196 sarsi, Bathypontia . . . 861-364 sarsi, Heteramalla .... 342 sassanus, Trichoniscus noricus . . 74 sayi, Portunus .... 146, 147 scaber, Androniscus subterraneus . . 70Scandia ...... 377 Scaphocalanus 312, 317~3i8, 320, 321, 325, 329, 333. 339Schiodtia . . . . . . 97 Schizocladium . 252, 254, 266, 272, 273 schneideri, Halecium . . . 396, 399schobli, Platyarthrus .... 78 Schoblia ...... 69 Schobliidae . . . . . 65, 69 scissidens, Gulella gwendolinae . 162, 163 Scolecithricella 312, 318, 320, 321, 323, 329-341Scolecithricidae . . 311-342, 367, 369 Scolecithrix . . .321, 325, 333, 336, 338 Scopalatum . 333, 335, 336, 338, 341, 342 scoparum, Trichoniscoides ... 73Scopimera .... 244, 245, 246 Scopimerinae ..... 245 scotti, Scolecithrix . . . -338 Scottocolanus . . . . -313 scutum, Helecium ..... 391 Scylla ..... 146, 147, 148 INDEX 437 Scyphax ....Scyphoniscussecurifrons, Scottocalanusseptentrionalis, Carcinus maenusseriepunctata, Philoscia .serpens, Capsulariaserpens, Filellum .serpens, Grammariaserpens, Rictulariaserrata, Cucumariaserrata, Scylla serrata marionensis, Cucumariaserratula, Obelia .serrulata, CampanulariaSertolare ....Sertularia 252, 253, 254, 265, 272,385, 391, seruposa, Corallinasessile, Halecium .Setaphora ....setosa, Sagittasilicus, Haplophthalmussima, Thalamitasimoni, Oniscussinensis, Centropussinghi, Cyrneasix-dentata, Charybdis .Slabberia ....smithae, Amallophora .Snelliaetideussolowetzkiana, Obeliasparrmanii, Tilapia . 191, spatium, Gulellaspatulifrons, MarioniscusSpelaeonethes sphaerocephalus, Rhyscotus .sphaerulina, Obeliaspinicarpus, Portunusspinifera, Bathypontia .SpinocalanidaeSpinocalanusspinosa, Sertulariaspinosus, Cordioniscus .spinosus, Spinocalanus .spinosus, Trichoniscusspinulosa, Campanulariaspinulosa, Laomedeaspinulosa, Obelia .spinulosa minor, LaomedeaSpiruridae ....spurca, Quickia . . 177, 178,spurca, Succineasquamata, Niambiasquamatus, Leptotrichussquamatus, Platyarthrus candatussquamiceps, NanochromisSquamiferidae squamuligera bargensis, Tirolosciasquamuligera briani, Philosciasquamuligera briani, Tiroloscia 9191 313. 118 88 378378-381 379, 385 379. 188 146, 147187, 188. 280. 280265 273. 380-381,393- 394. 45 393405, 406-40979, 91-92, 99 279 71146 . 84204204 146, 147. 252332, 336, 342 304, 305. 266194, 195, 196166, 167. 8473 . 76253, 265, 266146 . 364297-304297-304 253. 67 297, 298. 67260, 263260 260, 265260, 261, 264201-205180, 182, 183. 18077 77 78 23565, 77-78, 97 939393 squamuligera tendana, Tiroloscia stammeri, Trichoniscus . stebbingi, Cordioniscus . stebbingi, Trichoniscus . stellatus, Spinocalanus . Stenoniscidae Stenoniscus .... Stenophiloscia stentor, Grammaria Stewart, D. J. 93 74 . 6 7. 6730365, 76. 7692 382, 383, 384, 385191-197 strasseri, Androniscus cavernarum . . 70strasseri, Illyrionethes .... 70 strelkovi, Euplotes . 5, 7, 38, 43-44, 5 1 Streptaxidae .... 157-175 striata, Obelia . . . . .281 Strongylocentrotus . . . 23, 37 sturanus, Trichoniscus noricus . . 74 Stylohylea ...... 72 Styloniscidae . . .65, 66-69, 96-97 Styloniscus .... 68-69, 96, 97 styricus, Cyphoniscellus . . . .71 suarezia, Philosciasuarezia, Setaphorasubbrevicornis, Scaphocalanussubsessilis, Obelia .... subsessilis, Obelia geniculata . subterranea, Philoscia subterraneus medius, Androniscus . subterraneus noduliger, Androniscus subterraneus scaber, Androniscus Succinea Succineidae . sulcata, Dotilla sulcatus, Macrophthalmus superba, Bathynectes surcularis, Obelia . swellendami, Styloniscus swellendami, Trichoniscus syringa, Calicella . syringa, Sertularia 9279,92 317273, 277 275 88 70 7070 177, 180, 182, 184177-186, i PI.244, 245 245142, 143 . 266. 69. 69 390 254 127, tabulae, Styloniscus tabulae, Trichoniscus taylori, Pinnotheres tegulatus, Euplotes Teleostei tendana, Tiroloscia squamuligera Tendosphaera Tendosphaeridae . tenellum, Helicium . 393, 39^ tenuis, Heterorhabdus tenuis, Laomedea bicuspidata tenuis, Obelia terricola, Euplotes tetracythara, Salacia Thalamita Tharybidae . Thaumantias 2 5 2 thermophila, Bathytropa . 69. 69240, 242 8, 9. 45, 46-47227-231, i PI.93 - 7665, 76 >, 398,409-411. 346260, 26426754, 55. 381146, 147 367271, 2 73 79 254, 438 INDEX Thoa . 391, 393 thomasi, Scottocalanus . . -313 thononensis, Euplotes . . . 54-55 thorn elyi, Obelia . . . . .280 Thuiaria . . . . . .381 Tilapia ..... 191-197 Tiroloscia ..... 92-93 Titanethes ...... 73 Titaniidae . . . . . 65, 69 tomentosa, Amphiura . . . 188-189tomentosa, Trichorhina .... 79 torreyi, Halecium . . . 391 toticostata, Gulella . . . .167 tottoni, Laomedea . . . .281 Trewavas, E. . . 191-197, 2 33~ 2 35 triangularis, Uca ..... 245 triangulifera, Philoscia muscorum . . 88tricciratus, Euplotes affinis . . .18Trichoniscidae ... 65, 69-74, 97Trichoniscoides .... 72, 73 Trichoniscus . . 67, 68, 69, 72, 73-74, 97 Trichorhina . . . . . 78, 79 Trichoda . . . . . .41 trisulcatus, Euplotes . . .16, 86-37trituberculatus, Portunus . . 146, 147truncata, Niambia .... 77 truncatella, Philoscia .... 92 truncatella, Setaphora . . . .92 tsadiensis, Gulella gwendolinae 158, 162, 163tuberculata, Pseudochirella . . 307-309tuberculatus, Hanoniscus ... 82tuberculatus, Paranotoniscus ... 68tuberculatus, Phymatoniscus ... 72tuberculatus, Trichoniscoides ... 72Tubularia ...... 414 tuffraui, Euplotes . . 4, 6, 8, 16, 17, 37-38 Tylidae 65-66, 96 Tylos 65-66, 96 Tympanomerus ..... 244typica, Amallophora . . . 335, 338typicus, Euplotes patella . . .41 typicus, Xanthocalanus . . 332, 336, 338tyrrhena, Halophiloscia .... 82 Urocyclidae ....usitatus, Spinocalanus Valdiviella ....valens, AmallothrixVan Bruggen, A. C.Van Goethemvannus, Euplotes .vannus bolticus, Euplotesvariabilis, Euplotesvariabilis, Euplotes patellaventosus, Styloniscusventosus, Trichoniscus .verhoeffi, Trichoniscus .verrucosa, Tendosphaeraverrucosus, Styloniscus .verrucosus, Trichoniscusverticillata, SertulariaVerticillina ....Vesicularia ....veterum, Pinnotheresvilis, Aphilosciavilis, Philosciavillosa, Benthana .villosa, Philoscia .violaceus, Euplotesvittata, Philoscia .vividus, Trichoniscusvividus montanus, Trichoniscusvivipara, Heterorhabdusvivipara, Ophiacantha .volubis, Sertulariavorax, Scolecithrix waitatensis, Scyphoniscuswarreni, Philoscia .Washington!, Halecium . " .woodruffi, Euplotes 207-216, i PI.298, 303. 304 309-311 3i8. 157-175 207-216, i PL 8, 9, ii, 14-i6 ii 8, 26, 27, 2941 . 69. 69 74. 76. 69. 69 254 254 253. 240 79, 88798080ii . 887273 345, 346 . 189 / 254 332, 336, 342 88 409 56 Xanthocalanus 311-312, 332, 335, 338, 864-367Xavia ....... 145 Uca . undotheca, Obelia .undulatum, Halecium 244, 245 . 266 396, 398, 399, 4" Zebra., Equuszenkewitchi, Euploteszosterae, TrichoniscusZygophylax . 218, 220-223 7, 38, 44-45 74 378, 402 . A GUIDE TO THE SPECIES OF THE GENUS EUPLOTES(HYPOTRICHID A, CILIATEA) ( C. R. CURDS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. i LONDON: 1975 BY COLIN R. CURDS Pp 1-61 ; 58 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 28 No. I LONDON: 1975 533.174- 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. 28, No. i, of the Zoology series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 19 May 1975 Price 3.80 A GUIDE TO THE SPECIES OF THE GENUSEUPLOTES (HYPOTRICHIDA, CILIATEA) By COLIN R. CURDS INTRODUCTION ALTHOUGH species of the genus Euplotes Ehrenberg, 1830 are frequently observed inboth marine and freshwater samples it is often not possible for the protozoologist tomake specific identifications. This difficulty is due to at least three major factors ;firstly, many of the features used for distinguishing species within the genus areknown to vary considerably even within clonal cultures, secondly, there is a largeamount of confusion in the literature concerning the identity of certain commonspecies and thirdly, there is the added difficulty of having to search through aconsiderable body of literature before an identification can be attempted. It ishoped that the present paper will help solve some of these problems by gatheringtogether descriptions and diagrams of all those organisms considered to be distinctEuplotes species. Naturally the serious worker will still wish to refer to the originalmaterial but the volume of literature needed to be examined should be significantlyreduced. The revisions of Tuffrau (1960) and more recently the species lists of Borror(1972) have been of considerable value but one is still left with the problem of search-ing through a large amount of original papers. Finally, to the author's knowledgeno attempt has been made to devise a key to the species of Euplotes since that ofKahl (1932). FEATURES OF TAXONOMIC IMPORTANCE In the past 200 years over 80 species and varieties of the genus Euplotes havebeen described. Until relatively recently the species were separated simply on abasis of body size and shape, on the dorsal and ventral ridges and on the arrangementof the cirri on the ventral surface. Several decades were to pass after the intro-duction of the 'wet' silver impregnation method by Chatton and Lwoff (1930)before Tuffrau (1954) first discovered that the silver-line system or argyrome mightbe of taxonomic importance for species determination in the genus Euplotes. Tuffrau(1954) first suggested that the numbers of dorsolateral rows of cilia, the geometry ofthe argyrome, the number of frontoventral cirri and the form of the macronucleuscould be used to distinguish species and later (Tuffrau, 1960) he presented his revisionof the genus based on these ideas. Since that revision more species have beenredescribed and confirmed on the basis of silver preparations (principally Borror1962, 1963, igGSa) and several new species have been introduced. Borror (1972)published a list of species he found acceptable based on modern criteria togetherwith their probable synonyms. In general the present author's findings confirmthose of Borror (1972) although there are several instances where changes have beensuggested. Carter (1972) added four new species to the list of Borror (1972) andredescribed seven others, furthermore he was able to suggest from his investigations 4 C. R. CURDS that the shape of the adoral zone of membranelles (AZM) and the number of mem-branelles therein were also useful additional features for separating species withinthe genus. It is unfortunate that of the taxonomic characters mentioned above only one -the gross geometry of the dorsal argyrome - remains constant. All other featureshave been reported to vary to a greater or lesser extent and an account of some ofthese variations follows. However, the application of these modern criteria nowmakes it possible to examine and reappraise the species and varieties within thegenus. (a) General morphological and ecological features Although Kahl (1932) used the habitat of Euplotes species as a strict taxonomiccharacter he was ill advised to do so since several species are known to be euryhaline.However, specialized habitats such as the digestive tracts of sea urchins may beuseful for species determination. To date only two species have been recorded inurchins, E. balteatus (Dujardin, 1841) which may be found both as a commensal andfree-living and E. tuffraui Berger, 1965 which seems to be limited to a commensalmode of life. The size of a Euplotes species can be used as a taxonomic featurebut must be applied with extreme caution. It is true that the smallest speciesnever attain the size of the largest species and vice versa but it is also known that thesize of a ciliate may vary with several parameters including its rate of growth (Curds,West and Dorahy, 1974), the concentration of food (Curds and Cockburn, 1971) andkind of food (Giese, 1938 ; Tuffrau, 1964). The overall shape of a Euplotes speciesis perhaps a rather more stable feature but is of limited taxonomic value sincefew species have really characteristic outline shapes. Dorsal and ventral ridges are sometimes of value in species determinationparticularly when comparing modern preparations with early descriptions whereridges were often clearly figured. Borror (igGSa) discussed cortical sculpturing andsuggested that the use of nigrosin-HgCl a -formalin method (Borror, i_968b) mighthelp create a union of old and new species within the genus. Coloured species of Euplotes have been encountered for many years ; bothEhrenberg (1840) and Stein (1859) described green species which were likely to beforms of E. patella (Muller, 1773). Later Kahl (1932) listed three 'formae' of E.patella which contained zoochlorellae. It should be emphasized that one mustdistinguish between organisms feeding on green algae and those bearing zoochlorellae.Since Kahl (1932) only Diller and Kounaris (1966) seem to have mentioned zoo-chlorellae in Euplotes, and they described E. daidaleos Diller and Kounaris, 1966, aspecies closely related to E. patella, as continually containing zoochlorellae and thisperhaps is the only valid record although it would be unwise to dismiss the ob-servations of the early workers too readily. (b) Cirri The numbers and arrangement of the cirri on the ventral surface of Euplotes havebeen used as important taxonomic features for many years. Earlier workers THE GENUS EUPLOTES recognized four groups of cirri - frontals, ventrals, anals or transversals and caudals- but now the frontals and ventrals are more commonly treated as a single group -the frontoventral cirri (Fig. i). Several early workers placed a certain amount offaith on the numbers of caudal cirri as a taxonomic character. Kahl (1932), forexample, differentiated between varieties of E. moebiusi Kahl, 1932 and E. affinis(Dujardin, 1841) based on the numbers of caudal cirri and there are many othersimilar examples. However, modern evidence has clearly shown that variation incaudal cirri numbers is a common feature, and indeed seems to be the rule ratherthan the exception in certain species. Hufnagel and Torch (1967), for example,demonstrated in clones of E. vannus (Muller, 1786) that on division the proter alwaysreceived five and the opisthe four caudal cirri no matter how many caudals themother cell originally had ; thus these two morphological variants were found inequal proportions in the total population. Hufnagel and Torch (1967) further Frontals Fronloventrals Ventrals Anals orTransversals Caudals FIG. i. Ventral cirri of Euplotes. a. Wallengren (1900) system of cirrus numeration andolder method of naming cirri groups, b. Modern method of naming groups of cirri. showed that this variation was due to the normal morphogenetic events involvedin the development of the right caudal cirri from the right dorsal kinetics. Inaddition Hufnagel and Torch (1967) observed transient abnormal caudal cirrusdevelopment in E. plumipes Stokes, 1884 (a synonym of E. eurystomus Kahl, 1932)although no adults were found with an abnormal number of caudals. It is evidentfrom this type of information that the number of caudal cirri is of little, if any, valuein the identification of species of Euplotes. The reader is recommended to consultthe paper by Frankel (1973) which gives a good short modern account of morpho-genesis in hypotrichs. To the author's knowledge the numbers of transverse cirri have yet to be reportedto vary in their number which is remarkably constant throughout the genus. Withthe exception of E. strelkovi Agamaliev, 1967 which has 6 transversals all otherspecies have 5 transverse cirri. The number of frontoventral cirri is of greater usefor species determination than the other groups of cirri since although they varyfrom species to species they remain remarkably constant both in number and in 6 C. R. CURDS arrangement within a particular species. Most Euplotes possess 10 or 9 frontoventralcirri, far fewer have 8 while only E. raikovi Agamaliev, 1966 has been reported tohave 7 frontoventrals. With the exception of E. tuffraui the variation in fronto-ventral numbers seems to depend upon streak V (Wallengren, 1900 system, seeFig. i). Euplotes raikovi was reported (Agamaliev, 1967) to exhibit intraspecificpolymorphism in the number of frontoventrals ; he noted that there were 7 or 8frontoventral cirri and indicated that cirrus Vz (Fig. i) was that which did not developin some specimens. Washburn and Borror (1972) described a strain of E. raikovifrom America in which an eighth cirrus (2) never developed although they didobserve a barren plaque in each case. Curds (1974) recently reported a similar casein his description of E. parkei Curds, 1974 where cirrus V2 was present in some andabsent from other specimens even within the same clone. The frontoventral cirriof E. tuffraui show a wider variation than reported in other species and furthermorethe variation is not due to streak V but to streaks III and IV. Berger (1965)stated that the majority of E. tuffraui isolated from the digestive tracts of sea urchinspossess 8 frontoventral cirri although he found n specimens that had 10 fronto-ventrals which was apparently due to the subdivision of the cirrus bases IIl2,IIl3and IV 2. A further two specimens were found with 9 frontoventral cirri where onlycirrus bases III 3 and IV 2 were subdivided. With the exception of the exampleslisted above the number and positioning of the frontoventral cirri neverthelessremain very valuable taxonomic characters that have been widely used in the pastand will do so in the future. (c) The dorsal argyrome The overall geometrical pattern of the dorsal argyrome is one feature of taxonomicimportance that, to date, has not been reported to vary. Tuffrau (1960) first intro-duced this feature for taxonomic purposes and he described three general types whichhe called 'muscicola', 'eurystomus' and 'vannus' after the species in which he firstfound them. Since that time many more silver-line systems of Euplotes specieshave been described in the literature and it is now possible to divide the dorsalargyrome patterns into five or six types as shown in Fig. 2. The simplest type (Fig. 2a) includes six species whose dorsal interkinetal argyromepattern is composed of longitudinal kinetics with simple transverse connectionsbetween them so that there is a single row of polygons between the dorsal cilia orbristles. This group corresponds to the 'vannus' type of Tuffrau (1960) although itis suggested that the term 'single-vannus' is more descriptive and consistent with theterms to be used later. The first complication in the dorsal argyrome pattern is shown in Fig. 2b, wherethe longitudinal rows of polygons are split centrally so that there are two longi-tudinal rows of polygons, approximately equal in width, between the rows of dorsalcilia. This group corresponds to some of those Tuffrau (1960) called the 'eurystomus'type although it would appear that the organism that this author identified as E.eurystomus (Wrzesniowski, 1870) was in fact E. aediculatus Pierson, 1943. However,since both of these species have dorsal argyromes of the same overall pattern andsince 'eurystomus' type is now a widespread term it is suggested that it is kept but THE GENUS EUPLOTES e FIG. 2. Dorsal argyrome patterns of Euplotes. a. Single- vannus type. b. Double-eurystomus type. c-d. Double-patella types, e. Multiple type. f. Complex type. expanded to 'double-eurystomus' type which will serve to distinguish that group of24 species from another group of 8 species which also have double but unequal rowsof polygons between the dorsal cilia. Perhaps the best known species in the lattergroup is E. patella (Muller, 1773) which Tuffrau (1960) included in his 'eurystomus'group and for this reason it is suggested that the pattern be called the 'double-patella' type. In this type the rows of polygons (Figs. 2c, 2d), between the dorsalcilia, are obviously unequal in width ; in all but two species the wide rows of poly-gons are situated on the left of the rows of dorsal cilia (Fig. 2c), but in E. strelkoviand E. zenkewitchi Burkovsky, 1970 the wide rows are on the right and the narrowrows on the left of the kinetics (Fig. 2d). It could be argued therefore that the'double-patella' type should be further divided into these two sub-types, but thisdoes not seem warranted unless more species whose argyrome patterns conform tothe second sub-type are found. In the present author's opinion the 'muscicola' type of Tuffrau (1960) included aheterogeneous assemblage of dorsal argyrome patterns that should now be dividedinto two more natural groups. It is therefore suggested that the term 'muscicola'type be discontinued and replaced by the terms 'multiple' type and 'complex' type.When the dorsal argyrome patterns of these two types are examined it can be seenthat (Figs. 2e, 2f) the tendency for the rows of polygons to become further 8 C. R. CURDS subdivided is continued so that the next complication is that 3 or 4 regular rows ofpolygons of equal width may be found between the rows of dorsal cilia (Fig. ae)and these conditions are found in E. indentatus Carter, 1972 and E. muscicolaKahl, 1932 respectively. It is suggested that argyrome patterns such as thesebe called 'multiple' types. The final complication in dorsal argyrome patterns is that the polygons becomeso subdivided that an irregular network or mesh is formed between the dorsal ciliaso that distinct rows of polygons cannot be distinguished. It is suggested that thispattern be called the 'complex' type (Fig. 2f). The latter pattern is illustrated wellby the species E. gracilis Kahl, 1932, E. muscorum Dragesco, 1970 and E. elegansKahl, 1932 but not so well by E. moebiusi the fourth member of the group. In E.moebiusi the dorsal argyrome consists of a mixture of well-defined regular polygonsinterspersed with an irregular mesh work. With two exceptions the dorsal argyrome patterns of the species of Euplotes thathave been described to date fit neatly into the types outlined above. One of theseexceptions, E. moebiusi has already been mentioned and can readily be fitted intothe 'complex' group by the presence of an irregular meshwork between the dorsalcilia. The other exception, E. tegulatus Tuffrau, 1960, which was previously placedin the 'eurystomus' type (Tuffrau, 1960) is more difficult since it could almost equallywell be fitted into the 'double-eurystomus' or 'multiple' groups. In the case of E.tegulatus the dorsal argyrome pattern (see Fig. 46) consists of two longitudinal rowsof large polygons between the dorsal cilia as in E. eurystomus but here there is also acentral regular line of narrow elongate polygons. Euplotes tegulatus thereforedisplays what could be interpreted as an intermediate stage between the typical'double-eurystomus' type and the typical 'multiple' type as shown in E. indentatus.In the author's opinion it seems more appropriate to place this species in the 'multiple'type group and reserve the 'double-eurystomus' type group solely for those specieswith two equal rows of polygons between the kinetics. Whereas the overall geometrical pattern of the dorsal argyrome appears to be acompletely stable feature, the number of dorsolateral kinetics has been reported tovary in several species including the following ; E. polycarinatus Carter, 1972 (20-21kinetics), E. variabilis Stokes, 1887 (a synonym of E. eurystomus Kahl, 1932, 8-12kinetics), E. eurystomus (a synonym of E. aediculatus, 8-9 kineties) (see Carter, 1972,for all three species), E. vannus (8-9 kineties, see Heckmann, 1963), E. mutabilisTuffrau, 1960 (11-13 kineties, see Tuffrau, 1960), E. eurystomus (8-9 kineties, seeBonner, 1954) and E. tuffraui (9-10 kineties, see Berger, 1965). However, in themajority of species the number of dorsolateral kineties remains constant. In thecase of E. mutabilis, Tuffrau (1960) concluded that the variation in the number ofkineties was accidental and exceptional, although even accidental variations seemto be widespread in nature. In single clonal cultures Bonner (1954), Carter (1972)and Heckmann (1963) have all reported instances of variation in the number ofdorsolateral kineties. Furthermore, Carter (1972) found in three strains of E.eurystomus (a synonym of E. aediculatus) that the number of kineties was constantwithin the clone but different from clone to clone. In spite of variations such asthese the number of kineties in most cases is a good reliable taxonomic character. THE GENUS EUPLOTES g The numbers of kinetosomes or dorsal cilia plaques are more variable than thoseof kinetics, even so variation within the clone is frequently much less than fromspecies to species. Although several authors have recorded the number of dorsalcilia and by doing so have inferred that they are of taxonomic value no one has yetsuggested this number to be of real taxonomic importance. In the key that followsthe number of dorsal cilia has been used frequently but only when the numbers aresufficiently different to warrant their use. In each case the approximate range in themid-dorsal kinetics (where they are most numerous) are given. (d) The ventral argyrome The gross geometry of the ventral argyrome is of little taxonomic value sincedistinctive patterns are not immediately recognizable. However, some authorshave used the general size of the constituent polygons on a comparative basis andthis may be given as additional information. Silver-line preparations of the ventralsurface display the infraciliary network, the cirri plaques and the adoral zone ofmembranelles (AZM) which is of kinetosomal origin. Whereas in the past the sizeand shape of the AZM have often been used as taxonomic features, the number ofmembranelles therein has only recently been introduced. Carter (1972) found, in nspecies that he studied, that the AZM features were even more stable than thosecharacters suggested by Tuffrau (1960). However, all the features of the AZM areknown to vary and the classic example is that of E. balteatus which exhibits poly-morphism depending upon the nature of the food supply. Tuffrau (1964) found thatboth the size and shape of the AZM of E. balteatus was dependent upon the size ofthe cell and the number of membranelles in the AZM varied from 25 to 30 in smallcells and from 70 to 80 membranelles in the case of giant individuals. Fortunatelyvariation of this magnitude seems to be confined to that species, nevertheless variationin the number of membranelles is commonly observed. For this reason the approxi-mate ranges of membranelle numbers are given and their use has been restricted tocases when there is a considerable difference in the numbers between two species, forexample when distinguishing E. minuta Yocum, 1930 (30-40 membranelles) from E.vannus (60-70 membranelles). (e) Nuclear features The protozoan nucleus has been used as an important diagnostic character intaxonomic schemes for many years and the dimorphic nuclei of ciliates have playedan important part in ciliate taxonomy. The macronucleus of Euplotes is elongateand takes a variety of shapes which differ from species to species. Perhaps the mostcommon form is the simple inverted C-shape while in others this has become modifiedto a 3-shape and in others the arms close to form a hoop- or horseshoe-like structure.In addition, several rather more bizarre forms are known. The micronucleus isusually small and round but it does vary in size and shape and in E. tegulatus andE. aberrans Dragesco, 1960 the micronucleus is particularly large. The use of the shape of the macronucleus as a diagnostic feature in the genusEuplotes has been the subject of discussion for several years. Tuffrau (1960) sup-ported the view that the nuclear features were of great taxonomic importance io C. R. CURDS provided the shape of the macronucleus was determined when in a 'quiescent state',that is to say when neither division nor conjugation is in progress. It is true to saythat the macronucleus of Euplotes takes a variety of forms during reorganizationand division and it is likely that these phenomena have been responsible for the con-fusion that has appeared in the literature. Nevertheless, the consensus of opinionseems to be that if the process of reorganization is understood and provided it ispossible for the taxonomist to recognize the interphase macronucleus from one that isin another divisional state, then the interphase macronucleus is constant and charac-teristic of the species. The interphase macronucleus can be recognized by thepresence of replication bands (Gall, 1959 ; Kluss, 1962 ; Prescott, Kimball andCarrier, 1962) which represent bands of DNA synthesis travelling from the two nuclearextremities towards the centre of the macronucleus. Prescott et al. (1962) showedthat the macronucleus of E. eurystomus is in interphase when the replication bandsare present along the macronuclear arms from tips to half the distance to the centre. KEY TO THE GENUS EUPLOTES It is unfortunate that silver-line preparations have not yet been made anddescribed for all species of Euplotes. The key that follows relies heavily upon featuresdisplayed by the silver-line technique but a series of descriptions of undesignatedspecies follows the key (Section F, p. 50) and this includes all those species whosesilver-line systems have not yet been described. Diagrams of silver preparationsare given for all species where available and these have been obtained directly fromoriginal descriptions. Unless otherwise specified all scales given on diagramsindicate io jam. The characters for the key were selected and used in order ofleast variation. Therefore the first division of species into groups is made on abasis of the overall pattern of the dorsal argyrome and further subdivisions are madeusing the number of fronto ventral cirri and dorsolateral kinetics. Wherever pos-sible, reported variations of a feature within a species has been taken into account;thus it is possible to identify E. mutabilis when it has a 'single-vannus' type argyromeor when undergoing reorganization and therefore appears to have a complex argy-rome. Similarly it should be possible to identify E. eurystomus when it has any ofthe reported numbers of dorsolateral kinetics and so on. KEY TO THE MAJOR GROUPS OF SPECIES 1 a Single-vannus type dorsal argyrome (Fig. 2a) with single row of polygons between kinetics .......... 5 (Section A, p. n) b Some other type of dorsal argyrome ..'...... 2 2 a Double dorsal argyrome (Figs, ab, 2C, 2d) with two rows of polygons between kinetics ............. 3 b Multiple or complex dorsal argyrome (Figs. 2e, 21) . . . . . 4 3 a Double-eurystomus type dorsal argyrome (Fig. 2b) with two rows of equal-sized polygons between kinetics . . . ... .10 (Section B, p. 16) b Double-patella type dorsal argyrome (Figs. 2C, 2d) with alternate rows of wide and narrow polygons between kinetics ...... 36 (Section C, p. 38) THE GENUS EUPLOTES n 4 a Multiple type dorsal argyrome (Fig. 2e) with several (more than two) regular rows of polygons between kineties 43 (Section D, p. 45) b Complex type dorsal argyrome (Fig. 2f) with irregular meshwork of small polygons between kineties 45 (Section E, p. 47) SECTION A. KEY TO SPECIES WITH A SINGLE-VANNUS TYPE DORSAL ARGYROME 5 a 8 or less dorsolateral kineties .......... 6 b 9 or more dorsolateral kineties .......... 7 6 a 6 dorsolateral kineties with 7-10 dorsal cilia in central rows . . E. balticusb 8 dorsolateral kineties with 11-15 dorsal cilia in central rows . . . E. cristatus 7 a 9 dorsolateral kineties ........... 8 b 10 or more dorsolateral kineties ......... 9 8 a About 22 dorsal cilia in central kineties and 60-70 membranelles in AZM . E. vannusb About 13 dorsal cilia in central kineties and 30-40 membranelles in AZM . E, minuta 9 a 10 dorsolateral kineties with 26-30 dorsal cilia in central rows . . E. crassusb ii dorsolateral kineties with 11-15 dorsal cilia in central kineties . E. mutabilis SECTION A. DESCRIPTIONS OF SPECIESEuplotes balticus (Kahl, 1932) Dragesco, 1966 Kahl (1932) first described this species as E. vannus var. balticus while laterTuffrau (1960) considered it to be a synonym of E. crassus. However, the silverpreparations of Dragesco (1966) make it clear that E. balticus is a distinct species. DIAGNOSIS. Euplotes balticus (Fig. 3) is a medium-sized (60-100 /am long) ovoidmarine species. The peristome is long and narrow extending down to about f ofthe body length with the AZM containing about 50 membranelles. There are 10frontoventral, 5 transverse and 4 or 5 caudal cirri. The ventral argyrome consistsof a few very large irregular polygons. The dorsal argyrome is of the single- vannustype with 6 dorsolateral kineties carrying about 10 dorsal cilia in the central rows.The macronucleus is C-shaped with a club-like extension on the posterior arm(Fig. 3b) . The micronucleus is situated in a depression of the left anterior edge of themacronucleus. The contractile vacuole may have several satellite vacuoles surround-ing it. Euplotes crassus (Dujardin, 1841) Kahl, 1932 This species was first described by Dujardin (1841) under the name Ploesconiacrassa Dujardin, 1841. Kahl (1932) noted that E. violaceus Kahl, 1928 was in retro-spect a synonym of E. crassus. The silver-line system was first studied by Chattonand Seguela (1940) and fully described by Tuffrau (1960). DIAGNOSIS. Euplotes crassus (Fig. 4) is a large (100-130 /mi long) elongate ovalmarine species. The dorsal surface is strongly sculptured by 8 longitudinal ridges.The peristome is long and narrow while the AZM consists of about 50 membranellesand extends f down the body length. The dorsal argyrome is essentially simple ofthe single type and there are 10 dorsolateral kineties with the central ones bearingabout 26 dorsal cilia. There are 10 frontoventral, 5 transverse and 5 or 6 caudalcirri. The macronucleus is C-shaped with the posterior arm bearing a foot-like 12 C. R. CURDS FIG. 3. Euplotes balticus. a. Ventral aspect, b. Nuclei, c. Ventral and dorsalargyrome patterns. (After Dragesco, 1966.) FIG. 4. Euplotes crassus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) extension. The compact micronucleus is situated anteriorly near the left edge of themacronucleus. Euplotes cristatus Kahl, 1932 This species was originally briefly described by Kahl (1932) and subsequently byTuffrau (1960) and Carter (1972). THE GENUS EUPLOTES 13 DIAGNOSIS. Eiiplotes cristatus (Fig. 5) is a medium (60 /tin long, 45 /mi wide) ovalmarine species. The buccal cavity is narrow and almost covered completely an-teriorly by the lateral edge of the peristomial lip. The AZM is evenly curved, extends-f of the length of the body and is composed of 35-47 membranelles. The dorsalsurface is convex and there are 6 prominent ridges. The dorsal argyrome is of asimple single type with 8 dorsolateral kinetics bearing 11-15 dorsal cilia in the centralrows. There are 10 frontoventral, 5 transverse and 4 caudal cirri. In the originaldescription Kahl (1932) noted the presence of only 3 caudal cirri whereas Tuffrau(1960) and Carter (1972) both found 4 caudals. The macronucleus is C-shaped witha heel-like extension on the posterior arm. The small, compact micronucleus issituated on the upper left border of the macronucleus. Fig 5. Euplotes cristatus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes rninuta Yocum, 1930 Since the original description of E. minuta by Yocum (1930) this species dis-appeared into obscurity until Borror (1962) rediscovered it and described its silver-line system and general morphology. DIAGNOSIS. Euplotes minuta (Fig. 6) is a small (54 /um long, 28 /mi wide) oval marinespecies. The right margin of the peristome is almost straight and extends f downthe length of the body. The AZM consists of 30-40 membranelles. The dorsalargyrome is of the single type with 9 dorsolateral kinetics bearing 12-13 cilia in thecentral dorsal rows. There are 10 frontoventral, 5 transverse and 4 caudal cirri.The macronucleus is C-shaped with the posterior arm carrying a knob on its leftside and a foot-like extension (Fig. 6). The micronucleus is situated anteriorly onthe left edge of the macronucleus. C. R. CURDS FIG. 6. Euplotes minuta. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Borror, 1962.) Euplotes mutabilis Tuffrau, 1960 This species was first found and described by Tuffrau (1960) in his revisionarymonograph. Although the dorsal argyrome is of the single type, Tuffrau (1960)generally observed it as is shown in Fig. 7 where reorganization of the kinetosomalnetwork is in progress. Tuffrau (1960) rarely found specimens in the quiescent statewhich he believed was due to the rapid growth rate of the population that he studied.This species has therefore been included in both single-vannus and complex sectionsof the key since it would depend on the state of the organism into which group itwould appear to fall. DIAGNOSIS. Euplotes mutabilis (Fig. 7) is a medium (95 /u,m long) oval marinespecies that is widest at its posterior extremity. The peristome is quite large andthe right margin extends unevenly down the body to terminate in a spike-likeprojection. The AZM contains about 60 membranelles which extend down almostf of the body length. The dorsal argyrome is single in the quiescent state but withseveral irregular ramifications when undergoing reorganization. There are n dorso-lateral kinetics which bear 12 or 13 dorsal cilia in the central rows. There are 10frontoventral, 5 transverse and 4 or 5 caudal cirri. The macronucleus is C-shapedwith a pointed foot-like structure at the end of the posterior arm. The micronucleusis situated anteriorly. Euplotes vannus (Muller, 1786) Minkjewicz, 1901 This species has a long history which is given in detail by Tuffrau (1960), and thelist of synonyms in Appendix I is indicative of the problems that have arisen in the THE GENUS EUPLOTES FIG. 7. Euplotes mutabilis. a. Ventral argyrome. b. Nuclei, c. Dorsal argyromein state of reorganization. (After Tuffrau, 1960.) past but with the description of the silver-line system by Tuffrau (1960) most of theidentification difficulties were eradicated. DIAGNOSIS. Euplotes vannus (Fig. 8) is a medium-sized (75-100 /*m long) marinespecies. It has an overall oval configuration but is slightly curved towards the right.The peristome is narrow but large and the AZM which extends f down the length ofthe body contains over 60 membranelles. The dorsal argyrome is of the single type FIG. 8. Euplotes vannus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) 16 C. R. CURDS with 9 dorsolateral kinetics carrying about 22 cilia in the central dorsal rows. Thereare 10 strong frontoventral, 5 transverse and 4 fine but rigid caudal cirri. Themacronucleus is an open C-shape with a twisted foot-like extension to the posteriorarm. The micronucleus is compact and lies close to and sometimes overlappingthe macronucleus. SECTION B. KEY TO SPECIES WITH A DOUBLE-EURYSTOMUS TYPE DORSAL ARGYROME 10 a 6 or 7 dorsolateral kinetics .......... 11 b 8 or more dorsolateral kineties ......... 16 11 a 6 dorsolateral kineties . , ! . . . . . . . E. latus b 7 dorsolateral kineties . . . . . . . . . . .12 12 a Less than 10 dorsal cilia in central kineties ...... E. affinis b More than 1 1 dorsal cilia in central kineties ....... 13 13 a 10 frontoventral cirri ........... 14 b 8 or 9 frontoventral cirri . . . . . . . . . .15 14 a 3 caudal cirri, AZM with about 40 membranelles ..... octocirratusb 4 caudal cirri, AZM with about 20 membranelles .... E. trisulcatus 15 a 8 frontoventral cirri . . . . . . . . . E. poljanskyi b 9 frontoventral cirri . . . . . . . . . . E. dogieli 16 a 8 dorsolateral kineties ........... 17 b 9 or more dorsolateral kineties ......... 25 17 a 10 frontoventral cirri ........... 18 b 9 or less frontoventral cirri .......... 22 18 a Macronucleus C- or 3-shaped .......... 19 b Macronucleus hoop- or horseshoe-shaped ........ 21 19 a Slim species, macronucleus 3-shaped ...... E. antarcticus b Ovoid species, macronucleus C-shaped ........ 20 20 a 11-15 dorsal cilia in central kineties ....... E. alatus b 21-25 dorsal cilia in central kineties ....... E. crenosus 21 a Freshwater, 45-55 membranelles in AZM, extra-large cirri bases, 13-17 dorsal cilia in central kineties ....... E. magnicirratus b Marine or in sea urchins, two forms - a small one with 25-30 and a large one with 70-80 membranelles in AZM, 8-n dorsal cilia in central kineties . E. balteatus 22 a 6-10 dorsal cilia in central kineties ......... 23 b 16-35 dorsal cilia in central kineties ........ 24 23 a 8 frontoventral cirri (sometimes 9), 4 caudals and 10 dorsal cilia in central kineties E. parkeib 9 frontoventral cirri, 3 caudals and 6 dorsal cilia in central kineties . E. bisulcatus 24 a Macronucleus flat-backed C-shape with micronucleus distinctly separate. AZM straight E. aediculatus b Macronucleus 3-shaped with micronucleus within a cleft of it. AZM sigmoidal E. eurystomus 25 a 9 dorsolateral kineties ........... 26 b 10 or more dorsolateral kineties ......... 38 26 a Commensal in echinoids .......... E. tujfraui b Free-living ............. 27 27 a 10 frontoventral cirri, AZM with 25-30 membranelles . . E. quinquecarinatusb 9 frontoventral cirri, sigmoidal AZM with 46-65 membranelles . . E. eurystomus 28 a 9 frontoventral cirri ........... 29 b 10 frontoventral cirri ........... 31 29 a Commensal in echinoids, macronucleus C-shaped . -' . . . . E. tujfrauib Free-living, 3-shaped macronucleus ... r ...... 30 THE GENUS EUPLOTES 30 a 10-12 dorsolateral kineties ...... b 14 dorsolateral kineties ....... 31 a 10 dorsolateral kineties ....... b 1 1 or more dorsolateral kineties ..... 32 a 7-15 dorsal cilia in central kineties, commensal in echinoidsb 21-25 dorsal cilia in central kineties, free-living ii or 12 dorsolateral kineties . 33 a E. eurystomusE. amieti3233 . E. tuffrauiE. inkystans 34 b 13 or more dorsolateral kineties ......... 35 34 a ii dorsolateral kineties with about 18 dorsal cilia in central rows and 4 caudal cirri E. neapolitanusb 12 dorsolateral kineties with up to 40 (rarely 18-21) cilia in central rows and 5-8 caudal cirri ........... E. charon 35 a 13 dorsolateral kineties with 36-42 dorsal cilia in central rows . . E. harpab 20-21 dorsolateral kineties with 20-23 dorsal cilia in central rows . E. polycarinatus SECTION B. DESCRIPTIONS OF SPECIESEuplotes aediculatus Pierson, 1943 Euplotes aediculatus was first described by Pierson (1943). Later both Tuffrau(1960) and Carter (1972) described organisms which were obviously E. aediculatusbut named them E. eurystomus. Pierson, Gierke and Fisher (1968) produced well-documented evidence (Figs. 9, 10) on the differences between these two species. DIAGNOSIS. Euplotes aediculatus (Figs. 9, 10) is a large freshwater hypotrich105-160 /Ltm long. The peristome is medium sized, triangular and has two depressionsin the median border. One depression is located anteriorly whilst the other is moreprominent and is situated midway along the peristomial border. The AZM collaris not as prominent as in E. eurystomus, is straight to curved but never sigmoidal FIG. 9. Euplotes aediculatus. a. Ventral cirri, b. Nuclei, c. Dorsal argyrome.(After Pierson, Gierke & Fisher, 1968.) i8 C. R. CURDS FIG. 10. Euplotes aediculatus. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Tuffrau, 1960.) and contains about 40 membranelles. The macronucleus is C-shaped with an archedor flattened back and the micronucleus is distinctly separate from the macronucleus.The dorsal argyrome is of the double-eurystomus type and there are typically 8dorsolateral kinetics with about 20 cilia in the central kinetics on the dorsal surface.There are 9 frontoventral, 5 transverse and 4 caudal cirri. Euplotes affinis (Dujardin, 1841) Kahl, 1932 Until recently E. affinis and its variety with three caudal cirri, E. affinis formatricirratus Kahl, 1932 had not been described using modern techniques. Curds(1974) redescribed a freshwater species which closely resembles the latter form andgave diagrams of the silver-line system. Tuffrau (1960) considered E. affinis to be asynonym of E. charon (Muller, 1773). DIAGNOSIS. Euplotes affinis (Fig. n) is a small (38 ^m long, 26 /mi wide) ovoidfreshwater hypotrich with 9 frontoventral, 5 transverse and 3-4 caudal cirri. Oneof the caudal cirri is larger than the others and is held stiffly out to the right. Theventral surface is sculptured with 3 prominent ridges and the dorsal surface with 5longitudinal ridges. The AZM has 18-20 membranelles and extends f the length ofthe cell. There is a small undulating membrane. The dorsal argyrome is of thedouble-eurystomus type with 7 dorsolateral kinetics and a maximum of 9 dorsalcilia in the central kinetics. The macronucleus is a definite 3-shape and there is asmall compact anterior micronucleus. THE GENUS EUPLOTES FIG. ii. Euplotes affinis. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Curds, 1974.) Euplotes alatus Kahl, 1932 The original description of this species was given by Kahl (1932) and new dataconcerning the silver-line system were added by Borror (igGSa). DIAGNOSIS. Euplotes alatus (Fig. 12) is a small (40 pm long, 30 /urn wide) ovalmarine species. The dorsal surface has several inconspicuous low ridges but thoseon the ventral surface are far more conspicuous. The AZM extends just halfwaydown the length of the body and is composed of approximately 26 membranelles.The dorsal argyrome is of the double-eurystomus type with 8 dorsolateral kineticsbearing 10-12 dorsal cilia in the central rows. There are 10 frontoventral, 5 trans-verse and 4-5 caudal cirri. The macronucleus is C-shaped which has an indentationin the left anterior edge which contains the small compact micronucleus. Euplotes amieti Dragesco, 1970 This species is one of those described by Dragesco (1970) from the Cameroun inAfrica. It has many similarities with E. eurystomus including the presence of aanterior peristomial pouch, a similar nucleus, a sigmoidal AZM which makes thepresent author doubt whether this is a true species or simply a geographical varietyof E. eurystomus. However, because of the very large size, and more particularlythe presence of 14 dorsolateral kinetics, this organism has been treated here as aseparate species until more information concerning E. eurystomus and E. amieti hasbeen gathered. DIAGNOSIS. Euplotes amieti (Figs. 13, 14) is one of the largest (140-240 ^m long,80-160 jLim wide) species of Euplotes so far recorded. It is found in fresh waters 20 C. R. CURDS FIG. 12. Euplotes alatus. a. Ventral aspect, b. Nuclei, c. Dorsal aspect. (After Borror, I968a.) 30um FIG. 13. Euplotes amieti. a. Dorsal aspect showing kinetics, b. Ventral aspect. (After Dragesco, 1970.) THE GENUS EUPLOTES 21 and has a characteristic shape ; the dorsal surface being highly convex while theventral aspect is concave. The peristome is large, open and triangular in appearance.There is a well-developed peristomial collar and anterior pouch present. The AZMconsists of 52-62 membranelles and winds sigmoidally down towards the cytostome.The dorsal argyrome is of the double-eurystomus type with 14 dorsolateral kineticsbearing 28-30 cilia in the mid-dorsal rows. There are 9 fronto ventral, 5 transverseand 4 caudal cirri. The macronucleus is an irregular 3-shape with a compactmicronucleus situated in the left anterior corner of the macronucleus. FIG. 14. Euplotes amieti. a. Ventral and dorsal argyromes. b. Nuclei. (After Dragesco, 1970.) Euplotes antarcticus Fenchel and Lee, 1972 The description of this species depends solely upon the brief original report byFenchel and Lee (1972) based on material collected in Antarctica. DIAGNOSIS. Euplotes antarcticus (Fig. 15) is a medium-sized (85 /xm long, 30 /urnwide) marine species. The shape is unlike that of other species in being veryelongate and almost rectangular in outline except for the pointed posterior region.The peristome is long and narrow and there is a cleft in the right peristomial margin.The AZM is composed of approximately 30 membranelles and extends down ofthe body to the cytostome. The dorsal surface is clearly sculptured with 6 longi-tudinal ridges. The dorsal argyrome was not drawn very clearly but appears to beof the double-eurystomus type with 8 kinetics carrying about 13 cilia in the mid-dorsal rows. There are 10 frontoventral, 5 transverse and 4 or 5 caudal cirri. Themacronucleus is an elongate 3-shape. 22 C. R. CURDS FIG. 15. Euplotes antarcticus. a. Ventral argyrome. b. Macronucleus. c. Dorsalargyrome. (After Fenchel & Lee, 1972.) Euplotes bait cat us (Dujardin, 1841) Kahl, 1932 This species was first described by Dujardin (1841) and although the descriptionsgiven were brief and incomplete they were just sufficient to enable Kahl (1932) toidentify the species and present better diagrams. The morphology of the silver-linesystems were not available until Tuffrau (1964) published his work on polymorphismin the species. In the present author's opinion the E. balteatus described by Burkov-sky (1970) is E. cliaron. DIAGNOSIS. Euplotes balteatus (Fig. 16) is highly variable (30-150 /mi long) insize and the actual size depends to a great extent upon its food source (Tuffrau,1964). This species has been found living in marine waters but has also beenfrequently recorded in the intestinal tract of certain sea urchins (Allocentrus fragilis, FIG. 16. Euplotes balteatus. a. Ventral argyrome. b. Macronucleus. c. Dorsalargyrome. (After Tuffrau, 1964.) THE GENUS EUPLOTES 23 Strongylocentrotus droebachiensis , S. ehinoides, S. franciscanus and 5. purpuratus ~see Berger, 1965). The polymorphism of this organism is correlated with the AZMsize which enlarges considerably when feeding upon ciliates such as Philaster sp.When feeding on bacteria there are 25-30 small membranelles but when feeding onother ciliates the membranelles are larger in size and there are then 70-80 in number.The dorsal argyrome is of the double-eurystomus type but is less regular than isusually found. Tuffrau (1964) reported that there are 8 dorsolateral kinetics withup to ii cilia in the mid-dorsal rows. There are 10 frontoventral, 5 transverse and4 or 5 caudal cirri. The macronucleus is an open C-shape when feeding upon bacteriabut more horse-shoe shaped when feeding on ciliates. Euplotes bisulcatus Kahl, 1932 This species has been reported from marine sponges by Wenzel (1961) since itsfirst description by Kahl (1932). Since that time Borror (1963) isolated it from algalgrowths in tidal marsh ponds and described its silver-line system. DIAGNOSIS. Euplotes bisulcatus (Fig. 17) is a small (40 /urn long, 30 /urn wide) ovalmarine hypotrich. The dorsal surface has prominent double-edged ridges separatedby shallow grooves parallel to the dorsal ciliary rows 4, 5 and 6. The ventral surfaceis also ridged and the central ridge is very conspicuous. The AZM extends almostdown the length of the cell and is composed of about 17 membranelles. The dorsalargyrome is of the double-eurystomus type with 8 dorsolateral kinetics bearing 5-7cilia in the mid-dorsal rows. There are 9 frontoventrals, 5 transverse and 3 caudalcirri. The macronucleus is C-shaped with an adjacent anterior micronucleus. FIG. 17. Euplotes bisulcatus. a. Ventral aspect, b. Nuclei, c. Dorsal aspect. (After Borror, 1963.) 24 C. R. CURDS Euplotes charon (Miiller, 1786) Ehrenberg, 1830 This species has a long history and was described successively by Miiller (1786),Ehrenberg (1830, 1833, 1838), Dujardin (1841) and Stein (1859). These descriptionsvary considerably and it was not until almost a century later that Kahl (1932) gavea good succinct description of this species. Later Tuffrau (1960) and Borror (1963)were to describe the silver-line system. Tuffrau (1960) was of the opinion that E,affmis and E. moebiusi were synonyms of E. charon but recently Curds (1974) hasdemonstrated that these former two organisms are species in their own right. DIAGNOSIS. Euplotes charon (Fig. 18) is a medium (70-96 /u,m long) oval marinespecies which has a very large open triangular peristomial region. The right margin FIG. 18. Euplotes charon. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) of the peristome winds sinusoidally down past the cytostome to about f of the bodylength. The AZM bears approximately 70 strong membranelles while the AZM asa whole extends down to about f of the body length. The dorsal argyrome is of thedouble-eurystomus type with 12 dorsolateral kinetics carrying 35-40 dorsal cilia inthe mid-dorsal rows according to Tuffrau (1960) but only 18-21 according to Borror(1963). There are 10 frontoventrals, 5 transverse and 5-8 caudal cirri. Themacronucleus is horseshoe shaped and there is a small compact micronucleussituated anteriorly. Euplotes crenosus Tuffrau, 1960 This species was first discovered and described by Tuffrau (1960) and has remainedunmentioned since that time. THE GENUS EUPLOTES 25 DIAGNOSIS. Euplotes crenosus (Fig. 19) is a small (50-70 /Am long) freshwateroval hypotrich. There is a prominent notch at the anterior end of the cell which iscoincident with a longitudinal depression on the ventral surface of the cell. Theperistome is quite small and extends just over halfway down the body length. Thereare 25-30 membranelles in the adoral zone. The dorsal argyrome is of the double-eurystomus type with 8 dorsolateral kinetics bearing up to about 23 cilia in the mid-dorsal rows. There are 10 fronto ventral, 5 transverse and 4 caudal cirri. Themacronucleus is C-shaped with the micronucleus situated half-way down the leftborder of the macronucleus. FIG. 19. Euplotes crenosus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes dogieli Agamaliev, 1967 This species has only been described once and that was from material found insand samples taken from the Caspian Sea. DIAGNOSIS. Euplotes dogieli (Fig. 20) is a small (60 /mi long) marine specieswhose shape closely resembles that of E. poljanskyi Agamaliev, 1966 which is ellip-soid. The peristome is of medium size and it stretches about f of the length of thecell. There is a definite curved indentation almost midway down the left peristomialmargin. The AZM is composed of 35-38 membranelles and is rather narrow. Thedorsal argyrome is of the double-eurystomus type with 7 dorsolateral kinetics carryingabout 13 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and3 caudal cirri. The macronucleus is C-shaped with the anterior curve being angularand acute. The compact micronucleus is situated to the left of this anterior angularbend. 26 C. R. CURDS FIG. 20. Euplotes dogieli. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Agamaliev, 1967.) Euplotes eurystomus (Wrzesniowski, 1870) Kahl, 1932 Difficulties over the identification of E. eurystomus and confusion of it with severalother species is well documented but unfortunately is still perpetuated. Wrzes-niowski (1870) originally described an organism which he named E. patella var.eurystomus Wrzesniowski, 1870 which was later elevated to the species level E.eurystomus (Wrzesniowski, 1870) Kahl, 1932. Pierson (1943) made a comparativestudy of several strains of Euplotes and four distinct species were described by him.One of these was E. eurystomus and one was a hitherto undescribed organism thatwas named E. aediculatus Pierson, 1943. It was not until Tuffrau (1960) introducedsilver impregnation methods to Euplotes taxonomy that E. patella could readily bedistinguished from E. eurystomus and E. aediculatus; unfortunately Tuffrau (1960)also introduced a considerable amount of confusion since he described E. aediculatusbut called it E. eurystomus and described E. eurystomus but called it E. plumipesStokes, 1884. Even though Pierson, Gierke and Fisher (1968) pointed out the errorsof Tuffrau (1960), Carter (1972) disregarded the evidence and followed the scheme ofTuffrau (1960). Thus Carter (1972) also described E. aediculatus under the name ofE. eurystomus and E. eurystomus under the name E. plumipes ; furthermore, thislatter author reintroduced the species E. variabilis Stokes, 1887 which in the opinionof the present author is a variant of E. eurystomus. It is evident from the original diagrams given by Wrzesniowski (1870) (see Fig. 2ib)that the shape of the AZM is definitely sigmoidal and Kahl (1932) placed particularemphasis upon this shape as being a distinctive and immediately visible feature. Acomparison of the original diagrams in Figs. 2ia and 2ib of E. plumipes drawn byStokes (1884) and E. patella var. eurystomus drawn by Wrzesniowski (1870) showsthat both have a sigmoidal AZM and in fact have long been regarded as synonymous THE GENUS EUPLOTES species (Kahl, 1932 ; Borror, 1972). The sigmoidal shape of the AZM is also ob-vious in the silver preparations of E. plumipes and E. variabilis (Fig. 22) drawn byCarter (1972). In the descriptions that follow E. eurystomus (Figs. 23, 24) is regardedas was originally intended by Wrzesniowski (1870) and Kahl (1932), not as by Tuffrau(1960) and Carter (1972). DIAGNOSIS. Euplotes eurystomus (Figs. 22, 23, 24) is a large (100-160 /^m long,40-90 /*m wide) ovoid freshwater hypotrich. The buccal overture is triangular 30un 30um FIG. 21. Species synonymous with Euplotes eurystomus. a. Ventral aspect and macro-nucleus of Euplotes plumipes. (After Stokes, 1884.) b. Ventral aspect and macronucleusof Euplotes patella var. eurystomus. (After Wrzesniowski, 1870.) FIG. 22. Euplotes variabilis, a species synonymous with Euplotes eurystomus.a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Carter, 1972.) 28 C. R. CURDS and there is a single anterior peristomial pouch. The AZM collar is high andprominent. The AZM is markedly sigmoid in shape and contains 50-65 membran-elles. There are 9 frontoventral, 5 transverse and commonly 4 caudal cirri althoughCarter (1972) found 5 caudals in one specimen. The arrangement of cirri is shown inFigs. 22, 23 and 24. The dorsal argyrome is typical of the double-eurystomus typewith 10 dorsolateral kinetics although Carter (1972) reported two strains that had a FIG. 23. Euplotes eurystomus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome.(After Pierson, Gierke & Fisher, 1968.) FIG. 24. Euplotes eurystomus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome.(Called Euplotes aediculatus in Tuffrau, 1960.) THE GENUS EUPLOTES 29 variable number (8-12) of kinetics. The dorsal and ventral silver-line systems areshown in Figs. 22, 23 and 24. There are 17-25 dorsal cilia in the central kinetics.The macronucleus is typically 3-shaped and there is sometimes a definite concavenotch which contains the micronucleus. Carter (1972) believed that the nuclearpattern, the sigmoidal AZM, the possession of an anterior peristomial pouch and awide variation in the numbers of dorsolateral kinetics were sufficient to distinguishtwo strains of Euplotes from E. eurystomus and proposed the restoration of the speciesE. variabilis Stokes, 1884. However, a similar nuclear pattern was observed byPierson (1943) in E. eurystomus', the sigmoidal AZM has already been stated to be acharacteristic feature of E. eurystomus which species also has an anterior peristomialpouch. In the present author's opinion the variability of the numbers of dorso-lateral kinetics is not in itself sufficient to distinguish E. variabilis Stokes from E.eurystomus. Euplotes harpa Stein, 1859 This species was first properly described by Stein (1859) although Dujardin (1841)seems to have been the first to see this organism which he called Ploesconia cithara.Wallengren (1900, 1901) gave the first good diagram of E. harpa and these conformwell with the descriptions of Stein (1859). Although Chatton made silver prepara-tions of this species in 1939 these were not published until Tuffrau (1960) did so withnew specimens collected in 1955. DIAGNOSIS. Euplotes harpa (Fig. 25) is a large (150-160 /zm long) marine specieswith a more or less oval shape, although the left side is always more curved than theother. The peristome is large and open and there is a conspicuous lip on the rightmargin. The AZM is very curved and extends about f down the length of the body.There are approximately 65-70 membranelles in the AZM. The dorsal argyrome isof the double-eurystomus type with 13 dorsolateral kinetics bearing 40-45 dorsalcilia in the central rows. There are 10 fronto ventral, 5 transverse and 4 caudalcirri. The macronucleus is an open C-shape with tendencies towards a 3-shape.The micronucleus is in an anterior position. Euplotes inkystans Chatton in Tuffrau, 1960 Although this species was discovered by Chatton in the 1950*3 it was not describedby him but preparations that he made were described by Tuffrau (1960). DIAGNOSIS. Euplotes inkystans (Fig. 26) is a medium (70-80 /urn long) freshwateroval species. There is a prominent notch on the right of the peristomial collarwhich is narrow but rounded. The peristome is long and extends almost f down thelength of the cell. The AZM is composed of approximately 40 membranelles. Thedorsal argyrome is of the double-eurystomus type and there are 10 dorsolateralkineties with about 25 dorsal cilia in the central rows. There are 10 frontoventral,5 transverse and 4 (or rarely 5) caudal cirri. The macronucleus is a simple openC-shape with the micronucleus lying on the left border. C. R. CURDS 3Oum a FIG. 25. Euplotes harpa. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) FIG. 26. Euplotes inkystans. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes latus (Agamaliev, 1967) Euplotes latus was described by Agamaliev (1967) under the name E. patella formalatus Kahl, 1932. However, it is evident from the silver-line system that this speciescannot be a form of E. patella for several reasons (compare Figs. 27 and 40) ; forexample E. latus has a double-eurystomus type of dorsal argyrome whereas E. patellahas a double-patella type, the numbers of kinetics and dorsal cilia are different and THE GENUS EUPLOTES 31 the shapes of the macronuclei are different. However, it is also clear that the or-ganism described by Agamaliev (1967) does not conform to any others described todate and the present author considers it sufficiently different to warrant elevating itto a species in its own right. DIAGNOSIS. Euplotes latus (Fig. 27) is a medium (70 /am long) marine species thatis broadly rounded posteriorly but narrows anteriorly. The peristome is large andextends just over | down the length of the cell. The AZM is broadly curved and iscomposed of 35-40 membranelles. The dorsal argyrome is of the double-eurystomustype with only 6 dorsolateral kinetics bearing up to 15 dorsal cilia in the centralrows. There are 9 fronto ventral, 5 transverse and 4 caudal cirri. The two centrallypositioned frontoventral cirri (V 2 and VI 2 ) are situated very close together. Themacronucleus is an open angular C-shape with a micronucleus situated close to theleft anterior border. FIG. 27. Euplotes latus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome.(Called Euplotes patella forma latus in Agamaliev, 1967.) Euplotes magnicirratus Carter, 1972 This species is one of the four new species described recently by Carter (1972). DIAGNOSIS. Euplotes magnicirratus (Fig. 28) is a small (54 jum long, 40 p.m wide)oval marine species. The peristomial cavity is rather wide and extends approximately| of the length of the cell. The AZM is composed of about 50 membranelles and isrelatively straight for the majority of its length and then sharply curves in towardsthe cytostome. The dorsal surface is convex and prominently ridged. The dorsalargyrome is of the double-eurystomus type but the polygons are rather more squarethan is usual. There are 8 dorsolateral kinetics each containing 13-17 dorsal ciliaexcept the left lateral one which is short and contains only 5-8 cilia. The transversecirri are particularly large and there are 10 frontoventrals, 5 transverse and 4 caudal C. R. CURDS n FIG. 28. Euplotes magnicirratus. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Carter, 1972.) cirri. The macronucleus is an irregular hoop-shape and is highly characteristic ofthe species, since both ends are pointed. The micronucleus is situated in a smalldepression in the upper left border of the macronucleus. Euplotes neapolitanus Wichterman, 1964 This species was first described briefly by Wichterman (19623., b) and later (1964)more precisely from material collected from 50-60 metres in the proximity of theBay of Naples. DIAGNOSIS. Euplotes neapolitanus (Fig. 29) is a large (130 /urn long, 70 /zm wide)marine species that is ellipsoidal in shape although the anterior is often wider andmore truncated than the posterior. The peristome is conspicuous and extendsapproximately f down the body length. The AZM is composed of about 65 mem-branelles. The dorsal argyrome is of the double-eurystomus type consisting of ndorsolateral kinetics with about 18 cilia in the mid-dorsal rows. There are 10 fronto-ventral, 5 transverse and 4 caudal cirri. The macronucleus is C-shaped but botharms point posteriorly and the small spherical micronucleus lies in the middle of thebody on the left of the macronucleus. Euplotes octocirratus Agamaliev, 1967 Agamaliev (1967) described this species from samples of sand collected from theCaspian Sea. DIAGNOSIS. Euplotes octocirratus (Fig. 30) is a small (55-60 /xm long) marinehypotrich that is more or less oval in outline although the posterior is rather narrower THE GENUS EUPLOTES 33 than the anterior. The peristome is quite large and extends f down the length of thecell. The AZM is regularly curved and is composed of about 30 membranelles. Thedorsal argyrome is of the double-eurystomus type with 7 dorsolateral kineticscarrying up to 14 cilia in the mid-dorsal rows. There are 10 frontoventral, 5 trans-verse and 3 caudal cirri. The macronucleus is an angular C-shape with a compactmicronucleus in an anterior position. 30um FIG. 29. Euplotes neapolitanus. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Wichterman, 1964.) FIG. 30. Euplotes octocirratus. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Agamaliev, 1967.) 34 C. R. CURDS Euplotes parkei Curds, 1974 This species is a recent addition to the genus. DIAGNOSIS. Euplotes parkei (Fig. 31) is a small (40 /am long, 30 p.m wide) eury-haline species that is broadly oval in outline. The dorsal surface has 6 inconspicuouslow longitudinal ridges and the ventral surface has 7 minor ridges. The AZM isabout f of the body length and composed of 18 membranelles. A deep pocket nearthe cytostome contains an undulating membrane. There are usually 8 but occasion-ally 9 frontoventral, 5 transverse and 4 caudal cirri. The dorsal argyrome is of thedouble-eurystomus type with 8 dorsolateral kinetics bearing a maximum of n ciliain the mid-dorsal rows. The ventral argyrome consists of a series of few but largepolygons and resembles that of E. cristatus (see Tuffrau, 1960). The macronucleusis C-shaped and the micronucleus is situated close to its left anterior edge. FIG. 31. Euplotes parkei. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Curds, 1974.) Euplotes poljanskyi Agamaliev, 1966 This species has been described on two occasions by Agamaliev (1966, 1967) fromsamples of sand collected from the Caspian Sea. DIAGNOSIS. Euplotes poljanskyi (Fig. 32) is a small to medium-sized (55-70 /milong) marine ciliate. It has an elongated ellipsoidal shape with a medium-sizedperistome that is strongly concave on the right margin. The narrow AZM is com-posed of 36-40 membranelles, and it extends down to about f the body length.The dorsal argyrome is of the double-eurystomus type with 7 dorsolateral kineticsbearing 10-12 dorsal cilia in the central rows. There are 8 frontoventral, 5 trans-verse and 3 caudal cirri. The macronucleus is C-shaped and both ends are bluntlynarrower than the central regions. The micronucleus lies in an anterior position. THE GENUS EUPLOTES 35 FIG. 32. Euplotes poljanskyi. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Agamaliev, 1966.) Euplotes polycarinatus Carter, 1972 Recognition of this species relies upon the recent description of Carter (1972). DIAGNOSIS. Euplotes polycarinatus (Fig. 33) is a medium (90 /u,m long, 80 /miwide) almost triangular shaped marine hypotrich. The shape of the AZM is one ofits most distinctive features as it is very wide and crescent-like with 60-76 mem-branelles. The dorsal surface is slightly ridged and the dorsal argyrome is of thedouble-eurystomus type with 20 narrowly spaced dorsolateral kinetics bearing up to23 dorsal cilia. A few specimens have been seen with 21 dorsolateral kinetics. Theventral surface is flat and there are 10 frontoventral, 5 transverse and 5-9 caudalcirri. The macronucleus is a highly irregular 3-shape with a deep involution on theupper right border within which the compact micronucleus lies. Euplotes quinquecarinatus Gelei, 1950 The general morphology of this species was first briefly described by Gelei (1950)and the silver-line system was added later by Borror (i968a). DIAGNOSIS. Euplotes quinquecarinatus (Fig. 34) is a small (55 ju,m long, 40 /miwide) marine species whose general outline shape tends to be oval although theremay be conspicuous wing-like extensions to the ridges associated with dorsal ciliumrows i, 3 and 7. The AZM is small and extends halfway down the body length andcontains 25-30 membranelles. The dorsal argyrome is of the double-eurystomustype with 9 dorsolateral kinetics carrying 13-15 cilia in the mid-dorsal rows. Thereare 9 frontoventral, 5 transverse and 4 caudal cirri. The macronucleus is C-shapedand the micronucleus lies in an anterior position. C. R. CURDS FIG. 33. Euplotes polycarinatus . a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Carter, 1972.) FIG. 34. Euplotes quinquecarinatus. a. Ventral aspect, b. Nuclei, c. Dorsalaspect. (After Borror, i968a.) Euplotes trisulcatus Kahl, 1932 Kahl (1932) originally found this species in marine aquariums and gave sufficientlyprecise diagrams and descriptions to enable Tuffrau (1960), Borror (1963) and Carter(1972) to identify this species in other marine samples. The descriptions of thesilver-line system of this species given both by Tuffrau (1960) and Carter (1972) areidentical. THE GENUS EUPLOTES 37 DIAGNOSIS. Euplotes trisulcatus (Fig. 35) is a small (40 pm long, 30 pm wide) formthat has been only recorded in marine habitats. The shape of the body is distinctivein that it is prominently narrower at its posterior end. There is a pronouncedextension of the right side of the body beyond the peristomial collar. The peri-tomial cavity is long and narrow extending about f down the body. The AZM isevenly curved along the outer border of the peristome and contains 25-36 membran-elles. The dorsal surface is deeply ridged showing three prominent furrows. Thedorsal argyrome is of the double-eurystomus type and there are 7 dorsolateralkinetics with a maximum of n dorsal cilia widely separated in the central rows.There are 10 frontoventral, 5 transverse and 4 caudal cirri although Borror (1963)reported the presence of only 3 caudals. The macronucleus is a very open C-shapewith angular rather than rounded ends. The micronucleus is compact and is situatedanteriorly near the upper left border of the macronucleus. FIG. 35. Euplotes trisulcatus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes tuffraui Berger, 1965 Euplotes tuffraui was described by Berger (1965) from samples taken from theposterior digestive tract of three species of sea urchins. DIAGNOSIS. Euplotes tuffraui (Fig. 36) is a large (113 pm long, 73 /urn wide)marine species that is found in the digestive tract of strongylocentrotid echinoids.The three urchins that have to date been reported to contain this species are Allo-centrotus fragilis, Strongylocentrotus echinoides and 5. purpuratus. The shape of thebody is narrowly pyriform with the anterior end often being distinctly pointed.The AZM is just over half the length of the body and is composed of 40-45 mem-branelles. The dorsal surface is sculptured longitudinally with 7 ridges. The dorsalargyrome is of the double-eurystomus type but irregular with 10 (rarely 9) dorso-lateral kinetics with 8-u dorsal cilia per kinety. There are 8-10 frontoventralcirri depending upon the degree of fusion of the infraciliary bases in rows III andIV. There are 5 transverse and 4 caudal cirri. The macronucleus is C-shaped witha compact anterior micronucleus. C. R. CURDS FIG. 36. Euplotes tuffratii. a. Ventral aspect, b. Nuclei, c. Dorsal argyrome. (After Berger, 1965.) SECTION C. KEY TO SPECIES WITH A DOUBLE-PATELLA TYPE DORSAL ARGYROME 36 ab 37 ab 38 ab 39 ab 40 ab 41 ab 42 ab Double dorsal argyrome with large polygons on right and small polygons on left of kinetics . . . . . . . . . . . 37 Double dorsal argyrome with large polygons on left and small polygons on right of kineties ............. 38 9 frontoventral and 5 transverse cirri ...... E. zenkewitchi 8 frontoventral and 6 transverse cirri . . . . . . E. strelkovi 7 or less dorsolateral kineties, 7, 8 or 10 frontoventral cirri .... 39 8 or more dorsolateral kineties, 9 frontoventral cirri ...... 40 10 frontoventral cirri, 6 or less dorsal cilia in central kineties . . E. rariseta 7 or 8 frontoventral cirri, 11-15 dorsal cilia in central kineties . . . E. raikovi 8 dorsolateral kineties . E. octocarinatus 9 dorsolateral kineties . . . . . . . . . .41 With symbiotic green algae and about 60 membranelles in AZM . E. diadaleos Usually without symbionts, 30-35 membranelles in AZM ..... 42 11-15 dorsal cilia in central kineties ...... E. apsheronicus 25-30 dorsal cilia in central kineties ....... E. patella SECTION C. DESCRIPTIONS OF SPECIESEuplotes apsheronicus Agamaliev, 1966 This organism was originally described by Agamaliev (1966) from specimenscollected from sand samples from the Caspian Sea. A year later the same authorredescribed the species (Agamaliev, 1967) with a few amendments to the originaldescription. DIAGNOSIS. Euplotes apsheronicus (Fig. 37) is a small (50-60 /urn long) marinespecies that is ellipsoid in shape. There is a distinct peristomial collar and the THE GENUS EUPLOTES 39 peristome is of medium size being just over half the body length. The AZM iscomposed of 30-35 membranelles. The dorsal argyrome is of the double-patellatype with 9 dorsolateral kinetics (not 7-8 as originally given by Agamaliev, 1966)and these bear about 15 cilia in the mid-dorsal rows. There are 9 frontoventral, 5transverse and 4 caudal cirri. The macronucleus is an angular and very open C-shapewith a micronucleus located anteriorly. FIG. 37. Euplotes apsheronicus . a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Agamaliev, 1966.) Euplotes diadaleos Diller and Kounaris, 1966 This species has been recorded on one occasion only when Diller and Kounaris(1966) isolated it from an artificial pond in Pennsylvania, U.S.A. DIAGNOSIS. Euplotes diadaleos (Fig. 38) is a medium (92 ftm long, 57 pm wide)freshwater species that contains symbiotic zoochlorellae. The body is generallyflattened and oval in outline. The peristome extends slightly beyond the midlineof the body. The AZM consists of about 40-45 membranelles that curve smoothlyto the cytostome. The dorsal argyrome is of the double-patella type with 9 dorso-lateral kinetics containing 15-20 cilia in the mid-dorsal rows. There are 9 fronto-ventral, 5 transverse and 4 caudal cirri. The macronucleus is an angular C-shapewith the anterior micronucleus in a shallow depression of the macronucleus. Euplotes octocarinatus Carter, 1972 Euplotes octocarinatus is a recent addition to the genus and the only descriptionavailable is that by Carter (1972). 4 o C. R. CURDS DIAGNOSIS. Euplotes octocarinatus (Fig. 39) is a medium (80 pm long, 50wide) freshwater ellipsoid species. The peristome is triangular in shape and itextends about halfway down the length of the body. The AZM is narrow as itemerges from the dorsal anterior collar and then widens towards the middle of the body,it contains about 36-42 membranelles. There is a well-defined pouch to the right ofthe peristomial cavity. The dorsal surface is convex and ridged. The dorsal FIG. 38. Euplotes diadaleos. a. Ventral aspect, b. Nuclei, c. Dorsal argyrome.(After Diller & Kounaris, 1966.) FIG. 39. Euplotes octocarinatus. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Carter, 1972.) THE GENUS EUPLOTES 41 argyrome is of the double-patella type with 8 dorsolateral kinetics containing 18-21cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and 4 caudalcirri, although occasionally there may be 5 caudals. The macronucleus is a wide-mouthed C-shape and there is a small adjacent micronucleus. Euplotes patella (Miiller, 1773) Ehrenberg, 1838 Euplotes patella has a long historical record and has several synonyms ; it was firstdescribed by Miiller (1773) under the name Trichoda patella and subsequently byMiiller (1786) as Kerona patella. It was called E. patella by Ehrenberg (1838) butwas then named Ploesconia patella by Dujardin (1841). Several other early authorsalso refer to this species and much later Kahl (1932) found it necessary to subdividethe species into five forms which he named formae typicus, latus, alatus, planctonicusand variabilis (Stokes, 1887). These subdivisions were criticized by Pierson (1943)who observed that the first four of the above-mentioned forms were simply temporaryvarieties that could be found within clonal cultures of E. patella. Pierson (1943)also stated that E. patella forma variabilis Kahl, 1932 was in fact a variety of E.eurystomus. It was not until the work of Tuffrau (1960) that E. patella could readilybe distinguished from E. eurystomus but with silver-line preparations this is now quitea simple task. DIAGNOSIS. Euplotes patella (Fig. 40) is a large (no /urn long, 65 /mi wide) ovalfreshwater species that has a pronounced blunt posterior end. The peristome islarge, wide and almost triangular in appearance extending just over halfway downthe length of the body. The AZM is narrow, evenly curved and contains 44-50membranelles. The dorsal argyrome is of the double-patella type with 9 dorsolateral FIG. 40. Euplotes patella, a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) 42 C. R. CURDS kinetics containing about 26 cilia in the mid-dorsal rows, although it should be notedthat Carter (1972) observed rather fewer cilia (13-19). There are 9 frontoventral, 5strong transverse and 4 caudal cirri. The macronucleus is an open C-shape withslightly pointed extremities. The upper arm points down slightly and bears severalindentations. The micronucleus is situated anteriorly. Euplotes raikovi Agamaliev, 1966 This species was first described by Agamaliev (1966) and slightly amended later(Agamaliev, 1967). DIAGNOSIS. Euplotes raikovi (Fig. 41) is a small (50-60 /xm long) marine ciliatewhose outline body shape is broadly rounded. The peristome is of medium sizeand extends about f down the body length. The AZM is regularly curved andcontains 30-35 membranelles. The dorsal argyrome is of the double-patella typewith 7 dorsolateral kinetics bearing up to n cilia in the mid-dorsal rows. There are7 or 8 frontoventral, 5 transverse and 3 caudal cirri. The macronucleus was origin-ally reported to be a simple C-shape by Agamaliev (1966) but less like a C in hislater description (Agamaliev, 1967) (see Fig. 41). The micronucleus is small andsituated anteriorly. FIG. 41. Euplotes raikovi. a. Ventral argyrome. b. Dorsal argyrome. (AfterAgamaliev, 1966.) c-d. Nuclei (after Agamaliev, 1966 and 1967 respectively). Euplotes rariseta Curds, West and Dorahy, 1974 This species was first described by Borror (1963) but he called it E. moebiusi Kahl,1932. Curds, West and Dorahy (1974) isolated a marine hypotrich which in theiropinion did not conform to any previous descriptions and named their organism E. THE GENUS EUPLOTES 43 rariseta. Curds (1974) also described E. moebiusi Kahl, 1932 which agreed preciselywith the descriptions of Kahl (1932) but differed in several respects from E. rariseta.There seems little doubt that E. moebiusi Borror, 1963 is the species described byCurds et al. (1974) and that the E. moebiusi described by Curds (1974) conforms to thedescriptions of Kahl (1932) and to the part of the silver-line system shown by Klein(1958). DIAGNOSIS. Euplotes rariseta (Fig. 42) is a small (30-45 /mi long, 20-31 jam wide)marine hypotrich with 10 frontoventral, 5 transverse and 3 caudal cirri. The cirrusbelow the AZM is stout. The ventral surface is heavily sculptured with 6 posteriorlyprojecting ridges. The dorsal surface has an argyrome of the double-patella typewith 6 dorsolateral kinetics carrying a maximum of 6 cilia in the mid-dorsal rows.The macronucleus is an irregular S-shape. FIG. 42. Euplotes rariseta. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome.(After Curds, West & Dorahy, 1974.) Euplotes strelkovi Agamaliev, 1967 This species was described by Agamaliev (1967) from samples of sand taken fromthe Caspian Sea. DIAGNOSIS. Euplotes strelkovi (Fig. 43) is a small (50 ftm long, 40 /um wide)marine species that is particularly round in outline shape. The peristome is ofmedium size and the right margin is essentially straight. The AZM extends aboutf down the body and is composed of 33-38 membranelles. The dorsal argyrome isof the double-patella type with 6 dorsolateral kinetics bearing up to 10 cilia in themid-dorsal rows. The small polygons of the dorsal argyrome are on the left of thekinetics. There are 8 frontoventral, a unique 6 transverse and 3 caudal cirri. Themacronucleus is an irregular open C-shape with a compact micronucleus situatedanteriorly. 44 C. R. CURDS FIG. 43. Euplotes strelkovi. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Agamaliev, 1967.) Euplotes zenkewitchi Burkovsky, 1970 This species was first described by Burkovsky (1970) and more recently byAgamaliev (1972). DIAGNOSIS. Euplotes zenkewitchi (Fig. 44) is a medium (80 /mi long, 50 /zm wide)marine elongate hypotrich. The peristome is quite narrow and elongate extendingjust over halfway down the body. The AZM contains 50-55 membranelles. Thedorsal argyrome is of the double-patella type except that the narrow polygons lie FIG. 44. Euplotes zenkewitchi. a. Ventral argyrome. b. Nuclei, c. Dorsalargyrome. (After Burkovsky, 1970.) THE GENUS EUPLOTES 45 on the left of the kinetics. There are 10 dorsolateral kinetics with up to 18 dorsalcilia in the central rows. There are 9 frontoventral, 5 transverse and 3 or 4 caudalcirri. The macronucleus is C-shaped with a posteriorly pointing tail. The micro-nucleus is in an anterior position. SECTION D. KEY TO SPECIES WITH A MULTIPLE TYPE DORSAL ARGYROME 43 a Polygons of dorsal argyrome approximately equal in shape and size, with 9 or 10 dorsolateral kinetics ........... 44 b Polygons of dorsal argyrome of two types, squat polygons bordering the dorsal ciliaand very long polygons between squat ones, with 6 dorsolateral kinetics E. tegulatus 44 a 10 frontoventral cirri, 9 dorsolateral kinetics with 11-20 dorsal cilia in central rows E. indentatusb 9 frontoventral cirri, 10 dorsolateral kinetics with over 30 dorsal cilia in central rows E. muscicola SECTION D. DESCRIPTIONS OF SPECIESEuplotes indentatus Carter, 1972 This species is a recent addition to the species of the genus and its descriptionrelies upon that of Carter (1972). DIAGNOSIS. Euplotes indentatus (Fig. 45) is a small (60 /u,m long, 45 /u,m wide)marine species. The body is distinctly oval in outline shape with a prominentanterior notch in the upper border of the dorsal surface. The AZM is composed of42-48 membranelles and extends f of the body length. There are 10 frontoventral,5 transverse and 4 caudal cirri. The dorsal surface is convex and deeply ridged. FIG. 45. Euplotes indentatus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Carter, 1972.) 4 6 C. R. CURDS The dorsal argyrome is of the multiple type with three regular rows of polygonsbetween the kinetics. There are 9 dorsolateral kinetics bearing a maximum of15-21 cilia in the mid-dorsal rows. The macronucleus is a closed C-shape with theanterior micronucleus situated in a slight invagination. Euplotes muscicola Kahl, 1932 Although this species was intially described by Kahl (1932) and its encystment byFaure-Fremiet, Gauchery and Tuffrau (1954) a complete description was not availableuntil Tuffrau (1960) published its silver-line system. DIAGNOSIS. Euplotes muscicola (Fig. 46) is a medium (60-70 /um long) elongateoval freshwater hypotrich. The peristome is long and occupies about f of the anteriorleft side of the ventral surface. The AZM contains about 35 membranelles. Thereare 9 frontoventral, 5 transverse and 4 caudal cirri. The dorsal argyrome is of themultiple type and consists of 4 regular rows of small polygons between the kinetics.There are 10 dorsolateral kinetics bearing up to about 35 cilia in the mid-dorsal rows.The macronucleus is an open C-shape with a compact micronucleus situated abouthalfway down its left side. FIG. 46. Euplotes muscicola. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes tegulatus Tuffrau, 1960 The description of this species relies solely upon that of the original by Tuffrau(1960). DIAGNOSIS. Euplotes tegulatus (Fig. 47) is a large (116 /mi long) marine speciesthat has an elongate ellipsoidal shape. The dorsal surface is prominently sculpturedwith 3 or 4 ridges. The ventral surface has a pronounced longitudinal ridge that THE GENUS EUPLOTES 47 a FIG. 47. Euplotes tegulatus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) ends both posteriorly and anteriorly in spines. The anterior ventral spine is shortbut projects forward beyond the margin of the body. The peristome is long andcurved extending down f of the body length. The AZM consists of about 55 mem-branelles. The dorsal argyrome is of the multiple type but is unique in that thereare two longitudinal fibrils between the dorsolateral fibrils. There are 6 dorsolateralkinetics with up to 15 cilia in the mid-dorsal rows. There are 9 fronto ventral, 5transverse and 4 caudal cirri. The macronucleus is highly angular in appearance ;it is composed of two arms joined anteriorly at an acute angle. The micronucleusis very large for the genus (7-8 ^m diameter) and is situated anteriorly on the leftof the macronucleus. SECTION E. KEY TO SPECIES WITH A COMPLEX TYPE DORSAL ARGYROME 45 a 10 fronto ventral cirri ........... 46 b 9 frontoventral cirri ........... 47 46 a 7 dorsolateral kinetics, complex dorsal argyrome network interspersed with regular rows of polygons ......... E. moebiusi b 1 1 dorsolateral kinetics, complex dorsal argyrome network complete or partially so E. mutabilis (N.B. This species is undergoing reorganization - its description is given in SectionA, p. 14.) 47 a 7 dorsolateral kinetics with 11-15 cma m mid-dorsal rows . . . E. gracilisb 8 dorsolateral kinetics with 20 or more cilia in mid-dorsal rows .... 48 48 a 20-25 dorsal cilia in central kinetics, AZM with 30-35 membranelles . E. muscorumb More than 25 (usually 25-45) cilia in mid-dorsal kinetics, AZM with 40-45 mem-branelles ............ is. elegans C. R. CURDS SECTION E. DESCRIPTIONS OF THE SPECIESEuplotes elegans Kahl, 1932 This species was originally described briefly by Kahl (1932) and then redescribedin detail by both Tuffrau (1960) and Carter (1972). Kahl (1932) and Dragesco(1960) have also described a form of this species, E. elegans forma littoralis, but thisdiffers little from E. elegans and such a differentiation does not appear to be war-ranted. DIAGNOSIS. Euplotes elegans (Fig. 48) is a medium (80 ^m long, 55 /mi wide)euryhaline species that has an oval outline shape. The peristome is large and extendsabout | down the length of the body. The AZM is composed of 40-45 strong mem-branelles. There are 9 frontoventral, 5 transverse and 3 or 4 caudal cirri. Theoriginal description by Kahl (1932) appears to be the only observation of the presenceof 3 caudal cirri. The dorsal argyrome is of the complex type consisting of manyirregular polygons. There are 8 dorsolateral kinetics with a maximum of 30-46dorsal cilia in the central kinetics. The left lateral kinety is short with only 4-8cilia and is easily overlooked. The macronucleus is C-shaped with a small, blunt,knob-like projection on the upper arm and both arms taper to points. The micro-nucleus is small and compact. FIG. 48. Euplotes elegans. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes gracilis Kahl, 1932 First briefly described by Kahl (1932) this species was more fully described byTuffrau (1960). DIAGNOSIS. Euplotes gracilis (Fig. 49) is a small (37-50 /xm long) freshwaterspecies that has an elongate oval shape. The peristome is very deep and extends THE GENUS EUPLOTES 49 about f down the length of the body. The AZM is composed of 30-35 membranelles.There are 9 frontoventral, 5 transverse and 4 caudal cirri. The frontoventral cirriare very long and styliform whereas the caudals are thin. The dorsal argyrome isof a complex nature consisting of an irregular assemblage of polygons between the7 dorsolateral kinetics. There are 10-13 large kinetosomes in the mid-dorsal kine-tics. The macronucleus is C-shaped and the micronucleus is situated approximatelyof the way down its left edge. FIG. 49. Euplotes gracilis. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Tuffrau, 1960.) Euplotes moebiusi Kahl, 1932 Until recently E. moebiusi could only be identified from the brief and incompletedescriptions of Kahl (1932). Borror (1963) described an organism as E. moebiusibut this was later shown by Curds, West and Dorahy (1974) to be a new species, E.rariseta. Photographs of part of the silver-line system were first published by Klein(1958) in order to demonstrate the 'dry' silver method and although these were notsufficiently comprehensive for identification purposes they do conform with thesilver-line systems described by Curds (1974). Tuffrau (1960) considered E. moebiusito be a synonym of E. charon Ehrenberg, 1830. DIAGNOSIS. Euplotes moebiusi (Fig. 50) is a medium (60 jum long, 40 pm wide),ovoid euryhaline hypotrich with 10 frontoventral, 5 transverse and 4 caudal cirri.The ventral surface is heavily sculptured with 7 ridges and the dorsal surface with5 longitudinal ridges. The AZM is composed of 35-40 membranelles which extend| the length of the cell. The dorsal argyrome is unique amongst those speciesdescribed to date ; there are 5 longitudinal rows of narrow polygons interspersedwith an irregular network of larger polygons. The presence of the irregular networkhas been thought to be sufficient to place it within the 'complex' group until data C. R. CURDS concerning other species become available. There are 7 dorsolateral kinetics bearinga maximum of n cilia in the mid-dorsal rows. The macronucleus is 3-shaped andthe micronucleus is situated anteriorly. FIG. 50. Euplotes moebiusi. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Curds, 1974.) Euplotes muscorum Dragesco, 1970 This species was isolated from samples of moss collected in Africa by Dragesco(1970). DIAGNOSIS. Euplotes muscorum (Fig. 51) is a small to medium (50-70 /mi long)elongate freshwater species. The AZM contains approximately 30 membranellesand extends about f down the body. There are 9 frontoventral, 5 transverse and 4long caudal cirri. The dorsal argyrome is of the complex type with many polygonsarranged irregularly between the kinetics. There are 8 dorsolateral kinetics carrying22-28 cilia in the mid-dorsal rows on the peaks of the longitudinal ridges. Themacronucleus is C-shaped with the pointed right arm projecting posteriorly overhalfway down the length of the body. The micronucleus is situated anteriorlyalmost in line with the central longitudinal axis of the body. SECTION F. UNDEFINED SPECIES The silver-line systems of the following seven recognizable species have not yetbeen described. Their descriptions are given in alphabetical order. THE GENUS EUPLOTES FIG. 51. Euplotes muscorum. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Dragesco, 1970.) Euplotes aberrans Dragesco, 1960 This species was briefly and inadequately described by Dragesco (1960) fromsamples of marine sands but for several reasons may be regarded as a distinct speciesuntil further data are gathered. DIAGNOSIS. Euplotes aberrans (Fig. 52) is a medium (70-80 /mi long) marineciliate whose shape is highly elongate. There is a distinct spike-like projection onthe right of the peristomial collar. The dorsal surface bears 4 pronounced ridges andthere are 2 major ones, amongst others, on the ventral surface which travel slightlytrans versally almost the entire length of the body. The peristome is long andnarrow extending about f down the body and is composed of about 50 membranelles.There are 8 frontoventral cirri which is a feature shared by only three other species(E. strelkovi, E. raikovi and E. poljanskyi - all of which have differently shapedmacronuclei) . There are 5 transverse cirri in two groups and 4 caudals. There is avery large subequatorial vacuole which has not yet been seen to contract. Themacronucleus is horseshoe shaped whose ends almost meet one another. The micro-nucleus lies within a definite indentation in the anterior of the macronucleus. Euplotes novemcarinata Wang, 1930 Recognition of this species relies upon the single description by Wang (1930). DIAGNOSIS. Euplotes novemcarinata (Fig. 53) is a small to medium (60-75 /nm long)freshwater ciliate whose deep ridges are distinctive. The overall outline shape ofthe body is oval but there is a wing-like extension on the left side due to the projec-tion of one of the ridges. There are 9 longitudinal ridges, 5 are dorsal, i lateral and C. R. CURDS FIG. 52. Euplotes aberrans. a. Ventral aspect, b. Dorsal aspect, c. Nuclei. (After Dragesco, 1960.) ,0 FIG. 53. Euplotes novemcarinata. a. Ventral aspect, b. Macronucleus. c. Dorsalaspect. (After Wang, 1930.) THE GENUS EUPLOTES 53 3 less prominent ones are restricted to the ventral surface. The anterior end of thebody has a concave notch and the peristome extends down f of the body length.There are 9 frontoventral, 5 transverse and 4 caudal cirri. The macronucleus isC-shaped. Euplotes roscoffensis Dragesco, 1966 Although this species was not described until 1966 it was in fact first studied in1950 long before the importance of silver preparations for Euplotes taxonomy wasrealized. However, because the right-hand border of the peristome is so distinctivethis should be an easily identifiable species. DIAGNOSIS. Euplotes roscoffensis (Fig. 54) is a small to medium (60-70 /nm long)ovoid marine ciliate. The right border of the peristome is straight until it reachesthe level of the undulating membrane just below the middle of the cell, here theperistome border bears a distinctive deep invagination or pocket (Fig. 53) whichappears to be a unique character. The AZM is composed of 40-50 membranelles.There are 10 frontoventral, 5 transverse and 4 caudal cirri. The large contractilevacuole which is situated in the proximity of transverse cirrus VI i often has severalsmall satellite vacuoles surrounding it. The macronucleus is C-shaped with a club-like structure at the posterior end. The micronucleus is small, round and situatedclose to the anterior left edge of the macronucleus. FIG. 54. Euplotes roscoffensis. a. Ventral aspect, b. Nuclei. (After Dragesco, 1966.) Euplotes rotunda Gelei, 1950 The identity of this species relies upon the single brief and inadequate descriptionof Gelei (1950). 54 C. R. CURDS DIAGNOSIS. Euplotes rotunda (Fig. 55) is a small (50 /mi long, 40 /urn wide) roundfreshwater species. The peristome extends down to about the centre of the body andthe AZM carries approximately 25 membranelles. There are 8 dorsolateral kineticswith about 10 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverseand 3 caudal cirri. The macronucleus is a flattened 3-shape. FIG. 55. Euplotes rotunda, a. Ventral aspect, b. Macronucleus. c. Dorsal aspect. (After Gelei, 1950.) Euplotes terricola Penard, 1922 This species relies upon the single description by Penard (1922) who found it insamples of moss. Tuffrau (1960) was unconvinced that this organism is in fact amember of the genus but both Kahl (1932) and Borror (1972) include it in their works. DIAGNOSIS. Euplotes terricola (Fig. 56) is a small to medium (60-65 P l n g)freshwater species that is generally oval in outline but tends to narrow posteriorlyto a blunt point. The peristome is relatively short and extends just to the centre ofthe body and there appear to be somewhere in the order of 20 membranelles in theAZM. The dorsal surface seems to have 6 ridges and there are 9 frontoventral and5 transverse cirri. According to Kahl (1932), there are 10 caudal cirri althoughPenard (1922) originally described these as being very fine marginal cirri. Themacronucleus is a simple hoop-shape but the micronucleus is illustrated as lyinginside the curve of the macronucleus which is an unusual feature for the genus. Itis apparent that there are many doubts about this organism and it requires redescrip-tion before it can be adequately assessed. Euplotes thononensis Dragesco, 1960 Recognition of this organism relies upon the brief and inadequate description byDragesco (1960). THE GENUS EUPLOTES 55 FIG. 56. Euplotes terricola. a. Ventral aspect, b. Nuclei, c. Dorsal aspect. (After Penard, 1922.) DIAGNOSIS. Euplotes thononensis (Fig. 57) is a medium to large (90 /u,m long)marine species whose outline shape is oval except for the pronounced projectingperistomial collar. The dorsal surface carries 12 longitudinal furrows and thereappear to be about n dorsal cilia. The peristome is wide but extends less than half-way down the cell. There are 9 frontoventral cirri, one of which originates on theperistome border beneath the peristomial collar. There are 5 transverse and 4caudal cirri. A group of 3 contractile vacuoles is positioned close to the transversecirri. The macronucleus is an open C-shape with the micronucleus in an anteriorposition. FIG. 57. Euplotes thononensis. a. Ventral aspect, b. Nuclei, c. Dorsal aspect. (After Dragesco, 1960.) 56 C. R. CURDS Euplotes woodruffi Gaw, 1939 This species was first described by Gaw (1939) and more recently by Borror (1963).Unfortunately its silver-line system is still unknown but its unique macronucleusenables it to be easily recognized. DIAGNOSIS. Euplotes woodruffi (Fig. 58) is a large (145 p,m long) euryhaline ovalspecies with a well-defined peristomial collar. The peristome is large, triangular inshape and extends about f of the way down the body. The AZM is composed of60-70 membranelles. The dorsal surface is sculptured with 8-10 grooves and thereare 8 dorsolateral kinetics with about 60 closely set cilia in the central rows. Thereare 9 frontoventral, 5 transverse and 4 caudal cirri. The unique macronucleus isT- or Y-shaped with the micronucleus in an anterior position. FIG. 58. Euplotes woodruffi. a. Ventral aspect, b. Macronucleus. c. Dorsal aspect. (After Borror, 1963.) THE GENUS EUPLOTES 57 APPENDIX I CHECK-LIST AND INDEX OF SPECIES AND SYNONYMS 1. Euplotes aberrans Dragesco, 1960 description on p. 51 2. Euplotes aediculatus Pierson, 1943 17 Euplotes leticiensis Bovee, 1957Euplotes eurystomus Tuffrau, 1960Euplotes eurystomus Carter, 1972 3. Euplotes affinis (Dujardin, 1841) Kahl, 1932 18 Ploesconia affinis Dujardin, 1841Ploesconia subrotundus Dujardin, 1841 Euplotes subrotundus Perty, 1852Euplotes affinis var. tricirratus Kahl, 1932 4. Euplotes alatus Kahl, 1932 19 Euplotes labiatus Ruinen, 1938 5. Euplotes amieti Dragesco, 1970 19 6. Euplotes antarcticus Fenchel and Lee, 1972 21 7. Euplotes apsheronicus Agamaliev, 1966 38 8. Euplotes balteatus (Dujardin, 1841) Kahl, 1932 22 Ploesconia balteata Dujardin, 1841 9. Euplotes balticus (Kahl, 1932) Dragesco, 1966 ii Euplotes vannusvar. balticus Kahl, 1932 10. Euplotes bisulcatus Kahl, 1932 23 11. Euplotes charon (Muller, 1773) Ehrenberg, 1830 24 Trichoda charon Muller, 1773Ploesconia charon (Muller, 1773) Bory, 1826 Euploea charon Ehrenberg, 1830Euplotes appendiculatus Ehrenberg, 1838 Ploesconia charon Dujardin, 1841Ploesconia radiosa Dujardin, 1841Ploesconia longiremus Dujardin, 1841Euplotes balteatus Burkovsky, 1970 12. Euplotes crassus (Dujardin, 1841) Kahl, 1932 ii Ploesconia crassa Dujardin, 1841Euplotes taylori Garnjobst, 1928Euplotes violaceus Kahl, 1928Euplotes salina Yocum, 1930Euplotes crassus var. minor Kahl, 1932 13. Euplotes crenosus Tuffrau, 1960 24 14. Euplotes cristatus Kahl, 1932 12 15. Euplotes diadaleos Diller and Kounaris, 1966 39 Euplotes patella var. alatus Kahl, 1932 16. Euplotes dogieli Agamaliev, 1967 25 17. Euplotes elegans Kahl, 1932 48 17. Euplotes elegans (cont.) Euplotes elegans forma littoralis Kahl,1932 Euplotes elegans forma littoralis Drages-co, 1960 18. Euplotes eurystomus (Wrzesniowski, 1870) Kahl, 1932 26 Himantophorus charon Muller, 1786Euplotes plumipes Stokes, 1884Euplotes variabilis Stokes, 1887Euplotes patella var. eurystomus Wrzes-niowski, 1870 Uronychia paupera Daday, 1907Euplotes patella forma variabilis Kahl, 1932 Euplotes plumipes Tuffrau, 1960Euplotes plumipes Carter, 1972Euplotes variabilis Carter, 1972 19. Euplotes gracilis Kahl, 1932 48 20. Euplotes harpa Stein, 1859 29 Ploesconia cithara Dujardin, 1841 21. Euplotes identatus Carter, 1972 45 22. Euplotes inkystans Chatton in Tuffrau, 1960 29 23. Euplotes latus Agamaliev, 1967 30 Euplotes patella forma latus Agamaliev,1967 24. Euplotes magnicirratus Carter, 1972 31 25. Euplotes minuta Yocum, 1930 13 26. Euplotes moebiusi Kahl, 1932 49 27. Euplotes muscicola Kahl, 1932 46 28. Euplotes muscorum Dragesco, 1970 50 29. Euplotes mutabilis Tuffrau, 1960 14 30. Euplotes neopolitanus Wichterman, 1964 32 31. Euplotes novemcarinata Wang, 1930 51 32. Euplotes octocarinatus Carter, 1972 39 33. Euplotes octocirratus Agamaliev, 1967 32 34. Euplotes parkei Curds, 1974 34 35. Euplotes patella (Muller, 1773) Ehren- berg, 1838 41 Trichoda patella Muller, 1773Kerona patella Muller, 1786Coccudina keronina Bory, 1826Himantopus charon Ehrenberg, 1833Ploesconia patella Dujardin, 1841Euplotes charon var. marina Quenner- stedt, 1867 Euplotes paradoxa Kent, 1880Euplotes carinatus Stokes, 1885Euplotes patella var. alatus Kahl, 1932Euplotes patella var. lemani Dragesco, 1960 C. R. CURDS 36. Euplotes poljanskyi Agamaliev, 1966 34 37. Euplotes polycarinatus Carter, 1972 35 38. Euplotes quinquecarinatus Gelei, 1950 35 39. Euplotes raikovi Agamaliev, 1966 42 40. Euplotes rariseta Curds, West and Dorahy , 1974 42Euplotes moebiusi Borror, 1963 41. Euplotes roscoffensis Dragesco, 1966 53 42. Euplotes rotunda Gelei, 1950 53 43. Euplotes strelkovi Agamaliev, 1967 43 44. Euplotes tegulatus Tuffrau, 1960 46 45. Euplotes terricola Penard, 1922 54 46. Euplotes thononensis Dragesco, 1960 54 47. Euplotes trisulcatus Kahl, 1932 36 48. Euplotes tuffraui Berger, 1965 37 49. Euplotes vannus (Muller, 1786) Mink- jewicz, 1901 14 49. Euplotes vannus (cont.) Kerona vannus Muller, 1786Ploesconia vannus (Muller, 1786) Bory, 1826 Euplotes striatus Ehrenberg, 1838Euplotes longipes Claparede and Lach- mann, 1858 Euplotes extensus Fresenius, 1865Euplotes gabrieli Gourret and Roeser, 1886 Euplotes Worcester i Griffin, 1910Euplotes caudatus Meunier, 1910Euplotes truncatus Meunier, 1910Euplotes marioni Gourret and Roeser, 1886 50. Euplotes woodruffi Gaw, 1939 56 51. Euplotes zenkewitchi Burkovsky, 1970 44 A ddendum Since the preparation of this manuscript it has been brought to the author's notice that asilver-line preparation of E. woodruffi has been published (Magagnini & Nobili, 1964). It isof the double-eurystomus type with 8 dorsolateral kinetics and 25 middorsal cilia. In theexisting key these characters would lead to E. aediculalus and N. eurystomus. All three speciesmay be easily distinguished by their nuclear features. REFERENCES AGAMALIEV, F. G. 1966. 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Leiden, 20 : 243-256.STEIN, F. 1859. Der Organismus der Infusionsthiere nach eingenen Forschungen in systema- tischer Reihenfolge bearbeiteit. I. Leipzig. 2o6pp.STOKES, A. C. 1884. Notices of new fresh-water infusoria, II. Am. mon. microsc. J . 5 : 226- 230. 1885. Some new infusoria from American freshwaters. Ann. nat. Hist. 15 : 437-499. 1887. Some new hypotrichous infusoria from American fresh waters. Ann. nat. Hist. 20 : 104-114.TUFFRAU, M. 1954. Les caracteres specifiques dans le genre Euplotes (Note preliminaire). Bull. Soc. zool. Fr. 79 : 463-465.1960. Revision du genre Euplotes, fondee sur la comparaison des structures superficielles. Hydrobiologia, 15 : 1-77.1964. Le maintien des caracteres specifiques a travers le polymorphisme ^'Euplotes balteatus Dujardin, 1841. Archs Zool. exp. gen. 104 : 143-151. THE GENUS EUPLOTES 61 WALLENGREN, H. 1900. Zur Kenntnis der vergleichenden Morphologic der Hypotrichen. Bih. K. svenska Vetensk. Akad. Handl. 26 : 1-31.- 1901. Zur Kenntniss der Neubildungs- und Resorptionsprocesses bei der Theilung der hypotrichen Infusorien. Zool. Jb. 15 : 1-58.WANG, C. C. 1930. Notes on some new and rare species of hypotrichous infusoria. Contr. biol. Lab. Sci. Soc. China, 6 : 9-18.WASHBURN, E. S. & BORROR, A. C. 1972. Euplotes raikovi Agamaliev, 1966 (Ciliophora, Hypotrichida) from New Hampshire : description and morphogenesis. /. Protozool. 19 : 604-608. WENZEL, F. 1961. Ciliaten aus marinen Schwammen. Pubbl. Staz. zool. Napoli, 32 : 273-277.WICHTERMAN, E. ig62a. Studies on Euplotes. I. Structure and life cycle of a new species of marine Euplotes. Biol. Bull. mar. biol. Lab. Woods Hole, 123 : 516. I9&2b. Ciliate Protozoa from the Bay of Naples. Yb. Am. phil. Soc. 328-332. 1964. Descriptions and life cycle of Euplotes neapolitanus sp. nov. (Protozoa, Ciliophora, Hypotrichida) from the Gulf of Naples. Trans. Am. microsc. Soc. 83 : 362-370.WRZESNIOWSKI, A. 1870. Beobachtungen iiber Infusorien aus der Umgebung von Warsichan. Z. wiss. Zool. 20 : 467-511.YOCUM, H. B. 1930. Two new species of Euplotes from Puget Sound. Publs. Puget Sound mar. biol. Stn. 7 : 241-248. DR. C. R. CURDS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 560 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 884 jNU ,, 0*** CATALOGUE OF THE TYMS^TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) II. ONISCOIDEA, EXCLUDING PSEUDOTR ACHE ATA J. P. ELLIS AND R. J. LINCOLN BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 2 LONDON: 1975 CATALOGUE OF THE TYPES OF TERRE STRIA , ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) II. ONISCOIDEA, EXCLUDING PSEUDOTRACHEATA BY JOAN P. ELLIS AND ROGER J. LINCOLN Pp 63-100 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 2 LONDON: 1975 535. 3-75. 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. 28, 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.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 21 May, 1975 Price 2.55 CATALOGUE OF THE TYPES OF TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) II. ONISCOIDEA, EXCLUDING PSEUDOTRACHEATA By JOAN P. ELLIS AND ROGER J. LINCOLN THE following list incorporates all the families of woodlice represented in the Museumcollection of types with the exception of the pseudotracheate group which are listedin Part I of the type-catalogue (Lincoln & Ellis, 1974). Part II comprises 429separate items referring to 245 species and 76 genera within the following families :Tylidae, Styloniscidae, Titaniidae, Schobliidae, Trichoniscidae, Buddelundiellidae,Ligiidae, Stenoniscidae, Tendosphaeridae, Squamiferidae and Oniscidae. The style and arrangement of this catalogue are similar to Part I, and a list of themanuscript names referred to in the Appendix has been deposited in the library ofthe British Museum (Natural History). In total, this impressive collection of woodlice type material has over nooregistered entries representing 767 species in 153 different genera, and is thus ofconsiderable significance in a group which has only about 300 genera and rathermore than 2000 recognized species (Vandel, 1960). Family TYLIDAE TYLOS Latreille granuliferus Budde-Lund (1885 : 279) [Nom. nov. for Tylos granulatus Miers (1877 : 674)] SYNTYPES : two females, plus one specimen without abdomen (dry). Reg. no. 1847:21.Borneo ; 'among rotten woods, forest of Borneo, Eastern Seas'. Collected by Arthur Adams,H.M.S. 'Samarang'. Presented by Capt. Sir E. Belcher. SYNTYPE : female. Reg. no. 1872:17. Hiogo, Japan. Collected by George Lewis.Presented by F. Smith.neozealandicus Chilton (1901 : 120) SYNTYPE (?) : female. Reg. no. 1900:11:1:55. Wellington, New Zealand. Presented byC. Chilton.niveus Budde-Lund (1885 : 278) SYNTYPES (?) : one male ; one female ; one damaged specimen. (The description states'two examples'.) Reg. no. 1921:10:18:693-696. Key West, Florida, USA. 1878. Budde-Lund Collection.nudulus Budde-Lund (1906 : 76) SYNTYPES : one male ; one female. Reg. no. 1902:12:4:4-5. Christmas I. December1897 ' 'under stones and rotten wood'. Presented by J. Murray. SYNTYPES: two males; one female. Reg. no. 1921:10:18:697-699. Christmas I. Budde-Lund Collection. 66 J. P. ELLIS AND R. J. LINCOLN opercularis Budde-Lund (1885 : 277) SYNTYPE : one specimen in fragments. Reg. no. 1921:10:18:700. Philippine Is. Collectedby Gumming. Budde-Lund Collection.ponticus Budde-Lund (1885 : 274) SYNTYPES : two females. Reg. no. 1921:10:18:701-702. Sevastopol, Ukraine, USSR.Collected by Grebnitzsky. Budde-Lund Collection (ex Uljanin Collection). Family STYLONISCIDAECLAVIGERONISCUS Arcangeli mussaui Vandel (1973 : 20) SYNTYPES : thirty-five specimens : males, females and juveniles. Reg. no. 1970:343:35.Near Kuzi, Kolombangara, Solomon Is. 3-9.9.1965. 250-500 ft ; 'forest litter'. Collectedby P. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : thirty-six specimens : males, females and juveniles. Reg. no. 1970:344:36.Near Kuzi, Kolombangara, Solomon Is. 8.9.1965. 50 ft ; 'valley litter'. Collected byP. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : one male ; three females ; one juvenile. Reg. no. 1970:345:9. N. of Kiai,Kolombangara, Solomon Is. 6.9.1965. 1000 ft ; 'forest litter'. Collected by P. N. Lawrenceand Isiah. Presented by the Royal Society. SYNTYPES : three females. Reg. no. 1970:346:3. S.E. San Jorge, S. of Santa Isabel,Solomon Is. 23.9.1965 ; 'rotten wood litter'. Collected by P. N. Lawrence. Presented bythe Royal Society. SYNTYPES : four females. Reg. no. 1970:347:4. Fulakora Pt, Raja, Santa Isabel, SolomonIs. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES : two males ; nine females ; one juvenile. Reg. no. 1970:348:12. Lilihinia I.,Santa Isabel, Solomon Is. 21.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPES: one male ; eleven females. Reg. no. 1970:349:12. Cockatoo I., Santa Isabel,Solomon Is. 19-20.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPE : male. Reg. no. 1970:350:1. Thousand Ships Bay, opposite Lilihinia I.,Santa Isabel, Solomon Is. 20.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPES : male ; nine females. Reg. no. 1970:351:10. E. central San Jorge I., S. ofSanta Isabel, Solomon Is. 24.9.1965 ; 'forest litter'. Collected by J. Peake and P. N.Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:352:1. S. of San Jorge, Santa Isabel, Solomon Is.22.9.1965 ; 'gulley litter'. Collected by J. Peake and P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : twenty-eight specimens : males, females and juveniles. Reg. no. 1970:353:28.Popamanisiu, Guadalcanal, Solomon Is. 1965. 4400-7000 ft. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : twenty-one specimens : males and females. Reg. no. 1970:354:21. MtGallago, Guadalcanal, Solomon Is. 12.7.1965. 2500-3600 ft. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : two females. Reg. no. 1970:355:10 (part). Monitor Creek, Umasani River,Guadalcanal, Solomon Is. 8.7.1965 ; 'forest litter in hollow'. Collected by P. N. Lawrence.Presented by the Royal Society. TYPES OF TERRESTRIAL ISOPODS 67 SYNTYPES : one male ; seven females. Reg. no. 1970:355:10 (part). Umasani River, nrMt Gallego, c. 6 miles S.W. Tamboko, Guadalcanal, Solomon Is. 4.7.1965 ; 'disturbed forestlitter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : forty-two specimens : males, females and juveniles. Reg. no. 1970:356:42.Mt Austin, nr Honiara, Guadalcanal, Solomon Is. 24.7.1965. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : two females. Reg. no. 1970:357:2. Nuhu, Guadalcanal, Solomon Is. 28-31.10.1965. 1000 ft ; 'forest litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : thirty-six specimens : males, females and juveniles. Reg. no. 1970:358:36.Environs of Wainoni, San Cristobal, Solomon Is. 1965. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES: forty specimens : males, females and juveniles. Reg.no. 1970:359:40. Con-fluence of Warahito and Pagato Rivers, San Cristobal, Solomon Is. 1965. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : two males ; eight females. Reg. no. 1970:360:10. Huni River estuary,c. N.E. Wainoni, San Cristobal, Solomon Is. 9-12.8.1965 ; 'forest litter coral limestone'.Collected by Isiah. Presented by the Royal Society. SYNTYPES: three males; fourteen females. Reg. no. 1970:361:17. Ngaliau Hill, Pawa,Ugi, Solomon Is. 20.7.1965 ; 'secondary forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. CORDIONISCUS Graeve spinosus (Patience) (igo7a : 85) [Trichoniscus spinosus] SYNTYPES : one male ; three females. Reg. no. 1907:5:29:6-9. Springburn Public Park,Glasgow, Scotland ; 'in greenhouse'. Presented and collected by A. Patience. SYNTYPES: four females. Reg. no. 1911:11:8:10984-89. Springburn Public Park,Glasgow, Scotland. 1907 ; 'in greenhouse'. Collected by A. Patience. A. M. NormanCollection (ex Patience Collection).stebbingi (Patience) (i9O7b : 42) [Trichoniscus stebbingi] SYNTYPES: two males; three females. Reg. no. 1907:5:29:1-5. Springburn PublicPark, Glasgow, Scotland. Presented and collected by A. Patience. SYNTYPES: two males ; one female. Reg.no. 1911:11:8:10979-81. Scotland; Hawks-head Asylum, Renfrewshire, and Glasgow ; 'in greenhouses'. A. M. Norman Collection. SYNTYPES : three males. Reg. no. 1921:10:18:250-253. Glasgow, Scotland. Collectedby A. Patience. Budde-Lund Collection (ex Patience Collection). INDONISCUS Vandel orientalis Vandel (1973 : 18) SYNTYPES : one female ; one damaged specimen. Reg. no. 1970:432:6 (part). Popa-manisiu, Guadalcanal I., Solomon Is. 1-4.11.1965. 4400 ft ; 'mossy ridge forest litter'.Collected by P. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : one female ; one juvenile. Reg. no. 1970:432:6 (part). Popamanisiu,Guadalcanal I., Solomon Is. 5.11.1965. 7000 ft ; 'bog masses around trees'. Collected byP. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : three females. Reg. no. 1970:432:6 (part). Popamanisiu, Guadalcanal I.,Solomon Is. 6.11.1965. 7000 ft; 'moss forest'. Collected by P. N. Lawrence and Isiah.Presented by the Royal Society. SYNTYPE : male. Reg. no. 1970:433:1. S.E. Wainoni, San Cristobal, Solomon Is.10.8.1965. 1650 ft ; 'moss forest'. Collected by P. N. Lawrence and Isiah. Presented bythe Royal Society. 68 J. P. ELLIS AND R. J. LINCOLN PARANOTONISCUS Barnard capensis Barnard (1932 : 202) SYNTYPES : three males ; two females. Reg. no. 1933:1:25:934-939. Table Mt, CapeProvince, S. Africa. 2000-3000 ft. Collected and presented by K. H. Barnard.liitus Barnard (1932 : 205) SYNTYPES : three females. Reg. no. 1933:1:25:940-942. River Zonder End Mts, CapeProvince, S. Africa. Collected and presented by K. H. Barnard.montanus Barnard (1932 : 204) SYNTYPES : three males ; seven females. Reg. no. 1933:1:25:943-950. HottentotsHolland Mts, Cape Province, S. Africa. 1916. 4000 ft. Barnard Collection.ornatus Barnard (1932 : 205) SYNTYPES : approx. seven specimens in fragments, mainly ovigerous females. Reg. no.1933:1:25:951-956. Wellington Mts, Cape Province, S. Africa. 2000-3500 ft. Collectedand presented by K. H. Barnard.tuberculatus Barnard (1932 : 204) SYNTYPE : male. Reg. no. 1933:1:25:957. Langeberg Range, Cape Province, S. Africa.1927. 2000 ft. Collected and presented by K. H. Barnard. STYLONISCUS Dana mist rails (Dollfus) (1890 : 6) [Trichoniscus australis] HOLOTYPE : female. Reg. no. 1927:5:14:40. Inaccessible I., Tristan d'Acunha.16.10.1873. 'Challenger' Collection.austroafricanus (Barnard) (1932 : 200) [Trichoniscus austroafricanus'] SYNTYPES : one male ; one female. Reg. no. 1933:1:25:903-904. Table Mt, Cape Province,S. Africa. Collected and presented by K. H. Barnard.cestus (Barnard) (1932 : 201) [Trichoniscus cestus] SYNTYPES : seven females. Reg. no. 1933:1:25:905-910. Riversdale Mts, Cape Province,S. Africa. 1926. Collected and presented by K. H. Barnard.georgensis (Barnard) (1932 : 200) [Trichoniscus georgensis] SYNTYPES : two females. Reg. no. 1933:1:25:911-912. George, Cape Province, S. Africa.1931. Collected and presented by K. H. Barnard.horae (Barnard) (1932 : 200) [Trichoniscus horae] SYNTYPES : four females. Reg. no. 1933:1:25:915-917. Swellendam Mts, Cape Province,S. Africa. 1925. Collected and presented by K. H. Barnard.hottentoti (Barnard) (1932 : 197) [Trichoniscus hottentoti] SYNTYPES : two females. Reg. no. 1933:1:25:913-914. Hottentots Holland Mts, CapeProvince, S. Africa. 1916. Collected and presented by K. H. Barnard.kermadecensis (Chilton) (1911 : 569) [Trichoniscus kermadecensis] SYNTYPES : two females. Reg. no. 1912:5:25:67-68. Sunday I., Kermadec Is. 1908.Collected by W. R. B. Oliver. Presented by C. Chilton.mauritiensis (Barnard) (1936 : 3) [Trichoniscus mauritiensis] SYNTYPES: four males ; eleven females. Reg.no. 1936:7:13:1-5. Curepipe, Mauritius.12.1.1935. Collected and presented by R. F. Lawrence.monocellatus (Dollfus) (1890 : 7) [Microniscus monocellatus] HOLOTYPE : male. Reg. no. 1927:5:14:41. Juan Fernandez beach. 'Challenger' Collec-tion.rnoruliceps (Barnard) (1932 : 199) \Trichoniscus moruliceps~] HOLOTYPE : female (many appendages missing). Reg. no. 1933:1:25:958. JonkershoekMts, Stellenbosch, Cape Province, S. Africa. 1924. Collected and presented by K. H.Barnard. TYPES OF TERRESTRIAL ISOPODS 69 murrayi (Dollfus) (1890 : 5) [Trichoniscus murrayi] HOLOTYPE : male. Reg. no. 1927:5:14:39. Valparaiso, Chile. November 1875. 'Chal-lenger' Collection.otakensis (Chilton) (1901 : 117) [Trichoniscus otakensis] SYNTYPES : eight females. Reg. no. 1900:11:1:27-32. Dunedin, New Zealand. Pre-sented by C. Chilton.phormianus (Chilton) (1901 : 115) [Trichoniscus phormianus] SYNTYPES : nine females. Reg. no. 1900:11:1:1-5. Canterbury, New Zealand. Presentedby C. Chilton. SYNTYPES (?) : two males ; two females. Reg. no. 1921:10:18:414-417. Canterbury,New Zealand. Budde-Lund Collection (ex Dundee University Collection).riversdalei (Barnard) (1932 : 201) [Trichoniscus riversdalei] SYNTYPE : male. Reg. no. 1933:1:25:926. Riversdale Mts, Cape Province, S. Africa.1926. Collected and presented by K. H. Barnard.swellendami (Barnard) (1932 : 201) [Trichoniscus swellendami] SYNTYPES : two females. Reg. no. 1933:1:25:927-928. Swellendam, Cape Province, S.Africa. 1925. K. H. Barnard Collection.tabulae (Barnard) (1932 : 195) [Trichoniscus tabulae] SYNTYPES : four males ; three females. Reg. no. 1933:1:25:30-36. Table Mt, Cape Pro-vince, S. Africa. Collected and presented by K. H. Barnard.ventosus (Barnard) (1932 : 199) [Trichoniscus ventosus] SYNTYPES: three males; two females. Reg. no. 1933:1:25:929-933. Waaihoek Mts,Goudini, Worcester District, Cape Province, S. Africa. 1928. Collected and presented byK. H. Barnard.verrucosus (Budde-Lund) (1906 : 79) [Trichoniscus verrucosus] SYNTYPES : thirty-four specimens : males, females and juveniles. Reg. no. 1921:10:18:422-433. Possession I. 25.12.1901. Budde-Lund Collection. Family TITANIIDAEKOGMANIA Barnard depressa Barnard (1932 : 209) SYNTYPES: one male; one female. Reg. no. 1933:1:25:44-45. Kogmans Kloof, Montagu,Cape Province, S. Africa. 1922. Presented and collected by K. H. Barnard. Family SCHOBLIIDAESCHOBLIA Budde-Lund circular is Budde-Lund (1909 : 66) SYNTYPE : few fragments only. Reg. no. 1921:10:18:451. East Africa. Budde-LundCollection. Family TRIGHONISGIDAEALPIONISCUS Racovitza fragilis Budde-Lund (1909 : 68) SYNTYPES : two males. Reg. no. 1921:10:18:571-572. Sardinia. Budde-Lund Collection. 7 o J. P. ELLIS AND R. J. LINCOLN Subgenus ILLYRIONETHES Verhoeff heroldi Verhoeff (19315 : 22) HOLOTYPE : male. Reg. no. 1931:4:27:73. Hercegovini, Yugoslavia. Verhoeff Collec-tion.strasseri (Verhoeff) (19275 : 270) SYNTYPE : male. Reg. no. 1928:7:4:55. Istrien, Italy. Collected by K. Strasser.Verhoeff Collection. SYNTYPES : three males ; one female. Reg. no. 1930:5:26:76-79. Istrien, Italy. Col-lected by K. Strasser. Verhoeff Collection. SYNTYPES: three males; one female. Reg. no. 1937:7:6:132-135. Istrien, Italy.Collected by K. Strasser. Verhoeff Collection. ANDRONISCUS Verhoeff alpinus Verhoeff (igoSc : 140, 143) SYNTYPE : female. Reg. no. 1908:6:1:30. Lugano, Italy. Verhoeff Collection.brentanus Verhoeff (1932 : 22) SYNTYPES : one male ; two females. Reg. no. 1937:7:6:57-59. Brenta, Italy. Collectedby K. Strasser. Verhoeff Collection.calcivagus Verhoeff (igoSc : 140, 144) SYNTYPES : two males. Reg. no. 1908:6:1:31-32. Lake Como, Italy. Verhoeff Collection.SYNTYPE: female. Reg. no. 1921:6:10:81. Lake Como, Italy. Verhoeff Collection.carynthiacus Verhoeff (igoSc : 136) [Now Androniscus roseus (C. L. Koch)] SYNTYPES : four females. Reg. no. 1908:6:1:26-29. Korinthia, Greece. VerhoeffCollection.cavernarum Verhoeff (igoSc : 136) SYNTYPES : two males. Reg. no. 1908:6:1:24-25. Ukraine, USSR. Verhoeff Collection.cavernarum strasseri Verhoeff (ig28c : 157) SYNTYPES : one male ; one female. Reg. no. 1928:7:4:79-80. Istrien, Italy. VerhoeffCollection.dentiger Verhoeff (igoSc : 139, 143) SYNTYPES : one male ; one female. Reg. no. 1908:6:1:33-34. Italian Riviera. VerhoeffCollection.dentiger ligulifer Verhoeff (igoSc : 139) SYNTYPE : male. Reg. no. 1908:6:1:35. Bologna, Italy. Verhoeff Collection.roseus hamuligerus Verhoeff (ig28c : 158) SYNTYPES : two females. Reg. no. 1928:7:4:81-82. Ukraine, USSR. Verhoeff Collection.subterraneus medius Verhoeff (igsob : 12) SYNTYPES : one male ; one female. Reg. no. 1930:5:26:32-33. Friaul. Verhoeff Collec-tion.subterraneus noduliger Verhoeff (i92gb : 30) SYNTYPES : two males. Reg. no. 1930:5:26:30-31. Friaul. Verhoeff Collection.subterraneus scaber Verhoeff (i93ob : 13) SYNTYPE (?) : male. Reg. no. 1930:5:26:34. Friaul. Collected by K. Strasser. VerhoeffCollection. SYNTYPE : micropreparation. Reg. no. 1931:4:27:98. Villanova, Italy. Verhoeff Collec-tion. BURESCHIA Verhoeff bulgarica Verhoeff (ig26b : 140) SYNTYPE : male. Reg. no. 1928:7:4:78. Bulgaria. Collected by Buresch. VerhoeffCollection. TYPES OF TERRESTRIAL ISOPODS 71 CHAVESIADollius costulata Dollfus (1889 : 2) [Now Haplophthalmus danicus Budde-Lund] SYNTYPES : two females. Reg. no. 1911:11:8:10553. A9ores Is. January 1889. Collec-ted by Lt Chaves. A. M. Norman Collection (ex Dollfus Collection). CYPHONISCELLUS Verhoeff gottscheensis Verhoeff (19273, : 204) SYNTYPES : five males ; one female. Reg. no. 1928:7:4:56-60. Krain. VerhoeffCollection.styricus Verhoeff (19300 : 14) SYNTYPES : two males. Reg. no. 1930:5:26:39-40. Styria. Verhoeff Collection. HAPLOPHTHALMUS Schobl abbreviatus Verhoeff (i928c : 155) SYNTYPES : two males. Reg. no. 1928:7:4:113-114. Ukraine, USSR. Verhoeff Collec-tion.apuanus Verhoeff (19080 : 190, 193) SYNTYPES : two females. Reg. no. 1908:6:1:5. Italian Riviera. Verhoeff Collectiondanicus Budde-Lund (1885 : 250) SYNTYPES: c. 100 specimens, males, females and juveniles. Reg.no. 1921:10:18:311-322.Haunia. Budde-Lund Collection.fiumaranus Verhoeff (19080 : 189, 190) SYNTYPE : female. Reg. no. 1908:6:1:2. Fiume, Italy. Verhoeff Collection.fiumaranus dolinensis Verhoeff (19080 : 189, 192) SYNTYPES : one male, one female. Reg. no. 1908:6:1:3-4. Dolina, Ukraine, USSR.Verhoeff Collection.siculus Dollfus (18960 : 5) SYNTYPES : one male, one female. Reg. no. 1911:11:8:10840-41. Lake Lentini, Sicily,'damp earth near lake'. Collected by A. Dollfus. A. M. Norman Collection (ex DollfusCollection) . SYNTYPES : two females. Reg. no. 1921:10:18:393-395. Lake Lentini, Sicily. Collectedby A. Dollfus. Budde-Lund Collection (ex Dollfus Collection). HYLONISCUS Verhoeff adonis Verhoeff (ig2ja. : 219, 222) SYNTYPES: three males; two juveniles. Reg. no. 1928:7:4:71-75. Krain. VerhoeffCollection.crassicornis Verhoeff (ig26b : 155) SYNTYPE : female. Reg. no. 1928:7:4:77. Bulgaria. Verhoeff Collection.dalmaticus Verhoeff (i93ob : 7) SYNTYPES : one male ; one female. Reg. no. 1930:5:26:37-38. Dalmatia, Yugoslavia.Verhoeff Collection.inflatus Verhoeff (ig27a : 218, 221) SYNTYPE : male. Reg. no. 1908:6:1:42. Siebenbergen. Verhoeff Collection.mariae Verhoeff (igoSa : 376) SYNTYPE: male. Reg. no. 1908:6:1:40. Tatra. Verhoeff Collection. ?2 J. P. ELLIS AND R. J. LINCOLN narentanus Verhoeff (igoSa : 375) [Nom. nov. for Trichoniscus vividus Verhoeff from Herce-govini] SYNTYPES : four females. Reg. no. 1908:6:1:36-39. Hercegovini, Yugoslavia. VerhoeffCollection. SYNTYPES: one male ; one female. Reg. no. 1921:6:10:82-83. Hercegovini, Yugoslavia.Verhoeff Collection.refugiorutn Verhoeff (1933 : 41) SYNTYPES : two males ; three females. Reg. no. 1937:7:6:124-128. Apennine Mts,Europe. Verhoeff Collection. IBERONISCUS Vandel breuili Vandel (1952 : 351) SYNTYPES : nine males ; fifteen females ; four juveniles. Reg. no. 1951:3:30:7-26. OldSt Michael's Cave, Gibraltar. 6.3.1951 ; 'rotten wood; 650 ft from entry'. Collected byT. R. Shaw. Presented by the Cave Research Group of Great Britain. MIKTONISCUS Kesselyak halophilus Blake (1931 : 345) SYNTYPE : male (?) Reg. no. 1931:4:27:69. Massachusetts, USA. Verhoeff Collection. NESIOTONISCUS Racovitza corsicus corsicus (Racovitza) (1907 : 360) [Trichoniscus (Nesiotoniscus) corsicus] SYNTYPES: one male; one female. Reg. no. 1910:1:10:12-13. Grotte de Pietrabello,Corse, France. 9.1.1907. Presented by E. G. Racovitza. ORITONISCUS Racovitza flavus (Budde-Lund) (1906 : 83) [Trichoniscus flavus} [Nom. nov. for Trichoniscus vividusBudde-Lund (1885 : 246) nee. C. L. Koch] SYNTYPES: one male; five females. Reg. no. 1921:10:18:574-579. La Preste, France.Budde-Lund Collection.pyrenaeus Racovitza (1907 : 193) SYNTYPES: one male; two females. Reg. no. 1910:1:10:6-8. Grotte d'Arudy, Arudy,dep. Basses-Pyrenees, France. 6.9.1905. Collected and presented by E. G. Racovitza. PHYMATONISCUS Racovitza tuberculatus (Racovitza) (1907 : 174) \Trichoniscoides tuberculatus] SYNTYPES : three females. Reg. no. 1910:1:10:9-11. Grotte de 1'Herm, Herm, Ariege,France. 30.9.1905. Collected and presented by E. G. Racovitza. STYLOHYLEA Verhoeff Jagorum (Verhoeff) (i93ob : 5) [Trichoniscus (Stylohylea) fagorum] SYNTYPE : male. Reg. no. 1930:5:26:36. Croatia. Verhoeff Collection.SYNTYPE : male. Reg. no. 1970:4:1. Croatia. Larwood Collection (ex Verhoeff Collec-tion) . TYPES OF TERRESTRIAL ISOPODS 73 TITANETHES Schiodte albus (C. L. Koch) (1841 : 24) [Pherusa alba] SYNTYPE : male. Reg. no. 1925:7:22:139. Adelsberg, Krain. Collected by K. Schmidt.Koch Collection.dahli Verhoeff (ig26b : 137) [Nom. nov. for Titanethes albus Verhoeff (1900 : 118)] SYNTYPES : five females. Reg. no. 1928:7:4:83-87. Krain. Verhoeff Collection. Subgenus CYPHONETES Verhoeff hercegowinensis Verhoeff (1900 : 118) SYNTYPES: three males; four juveniles. Reg. no. 1901:9:19:97-103. Hercegovini,Yugoslovia. Verhoeff Collection. SYNTYPE: male. Reg. no. 1921:10:18:470. Hercegovini, Yugoslavia. Budde-LundCollection (ex Verhoeff Collection). TRICHONISCOIDES Sars mixtus (Racovitza) (1908 : 321) [Trichoniscus (Trichoniscoides) mixtus} SYNTYPES: one male; one female. Reg. no. 1910:1:10:1-2. Grotte de Baume-les-Messieurs, Jura, France. Presented by E. G. Racovitza.modestus (Racovitza) (1908 : 306) [Trichoniscus (Trichoniscoides) modestus] SYNTYPES : one male ; two females. Reg. no. 1910:1:10:3-5. Grotte de Rieufourcaud,Ariege, France. Presented by E. G. Racovitza.scoparum Verhoeff (igoSb : 176) [Now Oritoniscus flavus (Budde-Lund)] SYNTYPE : female. Reg. no. 1908:6:1:12. Pyrenees (St Beat). Collected by H. Ribaut.Verhoeff Collection. SYNTYPE : male. Reg. no. 1921:6:10:77. Pyrenees. Verhoeff Collection. TRICHONISCUS Brandt austr incus Verhoeff (igoSa : 376) SYNTYPES : two males. Reg. no. 1921:6:10:79-80. S.E. Alps. Verhoeff Collection.bosniensis Verhoeff (igoib : 75) SYNTYPE : male. Reg. no. 1901:9:10:106-107. Bosna i Hercegovini, Yugoslavia.Verhoeff Collection.cavernicola Budde-Lund (1885 : 2 4 6 ) [Now Spelaeonethes medius (Carl)] SYNTYPES: three males; five females. Reg. no. 1921:10:18:580-587. Pyrenees, 'incaves'. Budde-Lund Collection (ex Simon Collection).commensalis Chilton (1901 : 191) SYNTYPES: five males ; four females. Reg. no. 1952:4:18:1-9. Rai Valley, New Zealand.1902 ; 'in nests of ants'. Collected by J. McMahon. Presented by G. Jackson.elbanus Verhoeff (1931 a : 564) SYNTYPE: female. Reg. no. 1931:4:27:31. Elba I., Italy. Verhoeff Collection.fragilis rharelbazi Racovitza (1908 : 289) SYNTYPES: three males ; one female. Reg. no. 1910:1:10:14-17. Rhar-el-Baz, Algeria.Presented by E. G. Racovitza. montanus Carl (1908 : 143) [Trichoniscus vividus var. montanus] [Now Hyloniscus riparius C. L.Koch] SYNTYPES : one male ; one female. Reg. no. 1921:10:18:438-439. St Gallen, Switzerland.Budde-Lund Collection (ex J. Carl Collection). 74 J. P. ELLIS AND R. J. LINCOLN muscivagus Verhoeff (1917 : 52) SYNTYPES : three females. Reg. no. 1931:4:27:13-16. Salzburg, Austria. VerhoeffCollection.nivatus Verhoeff (1917 : 52) SYNTYPES: two males; one female. Reg. no. 1931:4:27:21-23. Salzburg, Austria.Verhoeff Collection.noricus insulanus Verhoeff (1931 a : 564) SYNTYPES : two females. Reg. no. 1931:4:27:19-20. Elba I., Italy. Verhoeff Collec-tion.noricus sassanus Verhoeff (1931 a : 565) [Now Trichoniscus foveolatus Vandel] SYNTYPE : male. Reg. no. 1931:4:27:18. Lake Maggiore, Italy. Verhoeff Collection.noricus sturanus Verhoeff (1931 a : 565) [Trichoniscus pusillus provisorius Racovitza]SYNTYPE : female. Reg. no. 1931:4:27:17. Piemonte. Verhoeff Collection.SYNTYPES: three females. Reg. no. 1937:7:6:51-53. Piemonte. Verhoeff Collection.pygmaeus Sars (1899 : 162) SYNTYPE: micropreparation. Reg. no. 1911:11:8:564. Christiana, Norway. NormanCollection (ex G. O. Sars Collection). SYNTYPES (?) : three females. Reg. no. 1911:11:8:10964-73. Norway. Norman Collec-tion (ex G. O. Sars Collection).stammeri Verhoeff (1932 : 21) SYNTYPE : female. Reg. no. 1930:5:26:58. Krain. Verhoeff Collection.verhoeffi Dahl (1919 : 209) [Now Trichoniscus pusillus Brandt] SYNTYPES : one male ; four females. Reg. no. 1931:4:27:24-28. Bergamask Alps.Verhoeff Collection.zoster ae Verhoeff (1931 a : 563) SYNTYPES : two females. Reg. no. 1931:4:27:29-30. Elba I., Italy. Verhoeff Collection.SYNTYPE : fragments only. Reg. no. 1970:5:1. Elba I., Italy. Larwood Collection (exVerhoeff Collection). Family BUDDELUNDIELLIDAE BUDDELUNDIELLA Silvestri cater actae Verhoeff (i93ob : 30)SYNTYPE : one specimen.Collection. Reg. no. 1930:5:26:41. Dalmatia, Yugoslavia. Verhoeff Family LIGIIDAEEURYLIGIA Verhoeff latissima Verhoeff (ig26a : 349) SYNTYPES: one male; one female. Reg. no. 1928:7:4:21-22. Canala Berg, New Cale-donia. Verhoeff Collection. LIGIA Fabricius cinerascens Budde-Lund (1885 : 265) SYNTYPES: four males; one female. Reg. no. 1921:10:18:1-5. Japan. Budde-LundCollection. TYPES OF TERRESTRIAL ISOPODS 75 dentipes Budde-Lund (1885 : 268) SYNTYPE : female. Reg. no. 1921:10:18:12. Pulo Milu, Nicobar Is., Bay of Bengal.Budde-Lund Collection.gracilipes Budde-Lund (1885 : 270) SYNTYPES : one male ; two females ; one specimen in fragments. Landana, Angola,S.W. Africa. Budde-Lund Collection (ex Simon Collection).tnelanocephala C. L. Koch (1838 : 18) [Now Ligidium hypnorum (Cuvier)] SYNTYPES: six males ; one female. Reg.no. 1925:7:22:104-110. Sugenheim, Bavaria,W. Germany. Koch Collection.natalensis Collinge (1920 : 474) PARATYPES : two males; eight females. Reg. no. 1919:4:26:371-380. Umhlali, Natal,May 1916, and Winkle Spruit Beach, South Coast, Natal, December 1916. Collected by C.Akerman. Presented by W. E. Collinge.perkinsi (Dollfus) (1900 : 525) [Geoligia perkinsi] SYNTYPES: three females. Reg. no. 1904:11:5:33-37 (part). Olaa, Hawaii. September1896. 2000 ft. Collected by Perkins. Presented by the Joint Committee of the RoyalSociety and British Association for Investigating the Fauna of the Sandwich Is. per Dr DavidSharp. SYNTYPES: one male; one female. Reg. no. 1904:11:5:33-37 (part). Waimea Mts,Kauai I., Hawaiian Is. June 1894. 4000 ft. Collected by Perkins. Presented by theJoint Committee of the Royal Society and British Association for Investigating the Fauna ofthe Sandwich Is. per Dr David Sharp.pigtnentata Jackson (1922 : 699) SYNTYPES: one male (?) ; one female. Reg. no. 1921:10:18:131. Suez. Budde-LundCollection. Subgenus POGONOLIGIA Jackson muscorutn Jackson (1927 : 130) [Now Ligia (Pogonoligia) platycephala (Van Name)] HOLOTYPE : male. Reg. no. 1927:4:4:1. Matacas Waterfall, Trinidad, West Indies. 14.2.1926 ; 'in moss under waterfall'. Collected and presented by C. L. Withycombe. PARATYPES: one male; six females. Reg. no. 1927:4:4:2-6. Matacas Waterfall, Trinidad, West Indies. 14.2.1926 ; 'in moss under waterfall'. Collected and presented by C. L. Withycombe. LIGIDIUM Brandt cursor turn Budde-Lund (1885 : 256) [Now Ligidium hypnorum (Cuvier)] SYNTYPES : four females. Reg. no. 1921:10:18:137-140. Zagreb, Yugoslavia. Budde-Lund Collection (ex Brusina Collection).herzegowinense Verhoeff (1901 a : 41) SYNTYPES : two females. Reg. no. 1901:9:19:39-40. Zenica, Yugoslavia. VerhoeffCollection.latum Jackson (1923 : 834) SYNTYPES: five females. Reg. no. 1921:10:18:168-174. San Francisco, USA. Collectedby G. Eisen. Budde-Lund Collection. Subgenus NIPPOLIGIDIUM Borutsky japonicum Verhoeff (1918 : 119) SYNTYPES : four females. Reg. no. 1938:7:7:69-72. Japan. Verhoeff Collection. 76 J. P. ELLIS AND R. J. LINCOLN Family STENONISCIDAEPARASTENONISCUS Verhoeff elbanus Verhoeff (i93ia : 558) [Now Stenoniscus pleonalis Aubert & Dollfus] SYNTYPES : one male ; two females. Reg. no. 1931:4:27:49-51. Elba I., Italy. VerhoeffCollection. Family TENDOSPHAERIDAE TENDOSPHAERA Verhoeff bretnbana Verhoeff (i93ib : 36) SYNTYPES : two specimens. Reg. no. 1931:4:27:63-64. Alpi Bergamasche, Italy.Verhoeff Collection.verrucosa Verhoeff (i93oa : 166) SYNTYPE : one specimen. Reg. no. 1930:5:26:75. Limone, Italy ('Seealpen'). VerhoeffCollection. Family RHYSCOTIDAERHYSCOTOIDES Arcangeli cubensis (Budde-Lund) (1908 : 300) [Rhyscotus cubensis] HOLOTYPE (?) : fragment only. Reg. no. 1921:10:18:1023. Cuba. Budde-Lund Collec-tion.linearis (Budde-Lund) (1908 : 300) [Rhyscotus linearis] HOLOTYPE : few fragments only. Reg. no. 1921:10:18:1025. Moheli I., Comores Archi-pelago, Indian Ocean. Collected by Voeltzkow. Budde-Lund Collection.ortonedae (Budde-Lund) (1908 : 299) [Rhyscotus ortonedae] SYNTYPES : eight males and two badly damaged specimens. Reg. no. 1921:10:18:1027-1038. Naranjito, Guayas, Ecuador. January 1901. Collected by V. Ortoneda. Budde-Lund Collection.parallelus (Budde-Lund) (1893 : 119) [Rhyscotus parallelus] SYNTYPES : ten males. Reg. no. 1921:10:18:1039-1049. Calvari Hill, Caracas, Venezuela.20.7.1891. Collected by Meinert. Budde-Lund Collection. SYNTYPES : three males. Reg. no. 1956:10:10:164-165. Calvari Hill, Caracas, Venezuela.20.7.1891. Collected by F. Meinert. Presented by University College, Dundee. RHYSCOTUS Budde-Lund bicolor Barnard (1924 : 235) SYNTYPES : thirty-six males ; five females. Reg. no. 1933:1:25:250-256. Ovamboland,S.W. Africa. Barnard Collection.globiceps Budde-Lund (1908 : 301) HOLOTYPE : few fragments only. Reg. no. 1921:10:18:1024. Loango, Congo. 6.7.1892.Collected by H. Brauns. Budde-Lund Collection.nasatus Budde-Lund (1908 : 301) HOLOTYPE : male. Reg. no. 1921:10:18:1026. Realejo, Nicaragua. 18.1.1903. Col-lected by C. F. Baker. Budde-Lund Collection.sphaerocephalus Budde-Lund (1893 : 120) SYNTYPES : two males. Reg. no. 1921:10:18:1050-1051. Caracas, Venezuela. 1891.Collected by Meinert. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 77 Family SQUAMIFERIDAE NIAMBIA Budde-Lund angusta Budde-Lund (1909 : 63) SYNTYPES : three males ; eight females. Reg. no. 1921:10:18:1408-1417. Steinkopf,Cape Province, S. Africa. August 1904. Budde-Lund Collection.brunnea Budde-Lund (1909 : 61) [Now Niambia truncata (Brandt)] SYNTYPES : six males ; twenty females. Reg. no. 1921:10:18:1418-1429. Kamaggas,Cape Province, S. Africa. July 1904. Collected by L. Schultze. Budde-Lund Collection.flavescens Barnard (1924 : 233) SYNTYPES: fifteen males ; twenty-eight females. Reg. no. 1933:1:25:140-149. Ondongua,Ovamboland, S.W. Africa. Collected by K. H. Barnard and R. F. Lawrence. Presented byK. H. Barnard.fortnicarutn Barnard (1932 : 268) SYNTYPES : eight males ; eight females. Reg. no. 1933:1:25:173-180. Matjiesfontein,Cape Province, S. Africa. Collected by W. F. Purcell. Barnard Collection.griseoflavus Barnard (1924 : 234) SYNTYPES : nine males ; five females. Reg. no. 1933:1:25:150-153. Andoni, Ovambo-land, S.W. Africa. 1923. Collected and presented by K. H. Barnard.hirsuta Budde-Lund (1909 : 62) [Now Niambia truncata (Brandt)] HOLOTYPE (?) : male. Reg. no. 1921:10:18:1434. Port Elizabeth, S. Africa. 15.12.1898.Collected by Dr Brauns. Budde-Lund Collection (ex Hamburg Museum).longicauda Barnard (1924 : 235) SYNTYPES : thirteen males ; thirty-four females. Reg. no. 1933:1:25:181-190. Andoni,Ovamboland, S.W. Africa. 1923. Collected and presented by K. H. Barnard.marginepapillosa Budde-Lund (1909 : 64) [Now Niambia capensis (Dollfus)] HOLOTYPE : fragments only. Reg. no. 1921:10:18:1435. Simonstown, Cape Province, S.Africa. Budde-Lund Collection.modest a Budde-Lund (1909 : 62) SYNTYPES: two males ; one female. Reg. no. 1921:10:18:1436-1438. Grootfontein, S.W.Africa. 10.1.1905. Collected by L. Schultze. Budde-Lund Collection.pallid u Budde-Lund (1909 : 61) SYNTYPES: sixteen males ; thirteen females. Reg. no. 1921:10:18:1439-1450. PossessionI. May 1903. Collected by Schultze. Budde-Lund Collection.palmetensis Vandel (1959 : 517) SYNTYPES : one male ; one female. Reg. no. 1973:473:2. Keta, Ghana. April 1958,'diseased crown of coconut palm'. Presented by the Commonwealth Institute of Entomology(coll. no. 16732) (ex Ministry of Food and Agriculture, Kumasi, Ghana).pusilla Budde-Lund (1909 : 63) [Now Niambia capensis (Dollfus)] SYNTYPES: one male ; one female. Reg. no. 1921:10:18:1451-1452. Simonstown, CapeProvince, S. Africa. Budde-Lund Collection.squamata (Budde-Lund) (1885 : 196) [Leptotrichus squamatus] SYNTYPE : male. Reg. no. 1921:10:18:1453. Landana, Congo. Budde-Lund Collection. Subgenus MANIBIA Barnard lot a Barnard (1932 : 270) HOLOTYPE: female. Reg. no. 1933:1:25:191. Sanyati Valley, S. Rhodesia. Collectedby R. H. Stevenson. Barnard Collection.microps Barnard (1932 : 271) SYNTYPES: four females. Reg. no. 1933:1:25:192-194. Maxixe, Mosambique. Collectedby R. F. Lawrence. Barnard Collection. 5*** 78 J. P. ELLIS AND R. J. LINCOLN PLATYARTHRUS Brandt caudatus Aubert & Dollfus (1890 : 10) SYNTYPES : one male; one female. Reg. no. 1911:11:8:10835-6. Marseille, France.Norman Collection (ex Dollfus Collection). SYNTYPE : female. Reg. no. 1921:10:18:1549. Marseille, France. Budde-Lund Collection(ex Aubert Collection).caudatus squamatus Verhoeff (igoSb : 180) [Now Platyarthrus caudatus Aubert & Dollfus] SYNTYPES : two females. Reg. no. 1908:6:1:22-23. Noli, Italian Riviera. VerhoeffCollection.costulatus Verhoeff (igoSb : 179) SYNTYPES: one male ; two females. Reg. no. 1908:6:1:19-21. Italian Riviera. VerhoeffCollection.schobli Budde-Lund (1885 : 200) SYNTYPES : three females. Reg. no. 1911:11:8:10837-39. Bona, Algeria. Collected byMeinert. Norman Collection. SYNTYPES: two females. Reg. no. 1956:10:10:155. Bona, Algeria. Collected by Meinert.Presented by University College, Dundee. TRICHORHINA Budde-Lund micros Budde-Lund (igi^b : 383) HOLOTYPE : fragment only. Reg. no. 1921:10:18:1607. Mauritius, Indian Ocean. Budde-Lund Collection.minutissirna Budde-Lund (igisb : 383) SYNTYPES : one male ; two females. Reg. no. 1913:1:8:123-125. Siren I., CargadosCarajos, Indian Ocean. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner.SYNTYPES: two males; four females. Reg. no. 1921:10:18:1608-1612. Siren I., CargadosCarajos, Indian Ocean. Collected by the 'Sealark' Expedition. Budde-Lund Collection.papillosa (Budde-Lund) (1893 : 123) [Alloniscus papillosus] SYNTYPES : one male ; one female. Reg. no. 1921:10:18:2163-2164. Caracas. Budde-Lund Collection. Family ONISCIDAEALLONISCUS Dana brevis Budde-Lund (1885 : 226) SYNTYPE : male. Reg. no. 1921:10:18:2113. 'Indes'. Collected by J. Ray. Budde-Lund Collection.cornpar Budde-Lund (1893 : I2 4) SYNTYPES : one male ; two females ; one other specimen in fragments. Reg. no. 1921:10:18:2137-2139. Caracas. Collected by Meinert. Budde-Lund Collection.cornutus Budde-Lund (1885 : 228) SYNTYPES: one male; one female. Reg. no. 1921:10:18:2140-2141. California, USA.Budde-Lund Collection.nacreus Collinge (1922 : 108) SYNTYPES : one male; five females. Reg. no. 1922:11:10:1-5. Tamatave, east coast ofMadagascar. Collected by Herscell and Chauvin. Presented by P. A. Methuen.porcellioides (Budde-Lund) (1904 : 45) [Arhina porcellioides] SYNTYPES : two males. Reg. no. 1921:10:18:971-972. No locality. Budde-LundCollection. TYPES OF TERRESTRIAL ISOPODS 79 ANCHIPHILOSCIA Stebbing cunningtoni Stebbing (1908 : 557) SYNTYPES : five males ; fifteen females. Reg. no. 1909:5:1:9-13. Niamkolo Bay, LakeTanganyika, 'under stones'. Collected by W. A. Cunnington. Presented by the TanganyikaExploration Committee. SYNTYPE : one micropreparation. Reg. no. 1909:5:1:14. Niamkolo Bay, Lake Tan-ganyika ; 'under stones'. Collected by W. A. Cunnington. Presented by the TanganyikaExploration Committee.karongae Stebbing (1908 : 556) [Now Setaphora suarezia Budde-Lund] SYNTYPES : two males; four females. Reg. no. 1909:5:1:15-19. Kambwe, nr Karonga,Tanzania. 27.6.1904 ; 'on damp decaying wood close to swamp'. Collected by W. A.Cunnington. Presented by the Tanganyika Exploration Committee. SYNTYPES : six micropreparations. Reg. no. 1909:5:1:20-25. Kambwe, nr Karonga,Tanzania. 27.6.1904 ; 'on damp decaying wood close to swamp'. Presented by the Tan-ganyika Exploration Committee. APHILOSCIA Budde-Lund vilis (Budde-Lund (1885 : 210) [Philoscia vilis] HOLOTYPE : in fragments. Reg. no. 1921:10:18:2090. Cape of Good Hope. Collected byDrege. Budde-Lund Collection. ARMAD1LLONISCUS Uljanin dalmatinus Verhoeff (igoia : 39) [Now Armadilloniscus littoralis Budde-Lund] SYNTYPES: one male; one female. Reg. no. 1970:82:2. Dalmatia, Yugoslavia. LarwoodCollection (ex Verhoeff Collection).littoralis Budde-Lund (1885 : 237) SYNTYPES: one male; one female. Reg. no. 1921:10:18:1057-1058. Venetias. Collectedby Schaufuss. Budde-Lund Collection. BATHYTROPA Budde-Lund granulata Aubert & Dollfus (1890 : 9) SYNTYPE : female. Reg. no. 1911:11:8:10549. Marseille, France. Norman Collection (exDollfus Collection).hispana Dollfus (1893 : 5) [Now Haplophthalmus danicus Budde-Lund] SYNTYPE: female. Reg. no. 1911:11:8:10546. Valencia, Spain. Norman Collection (exDollfus Collection). SYNTYPE : female. Reg. no. 1921:10:18:376. Valencia, Spain. Budde-Lund Collection(ex Dollfus Collection).tneinerti Budde-Lund (1885 : 197) SYNTYPES : two males. Reg. no. 1911:11:8:10550-51. Bona, Algeria. Collected byMeinert. Norman Collection (ex 'Zool. Mus. Haun.'). SYNTYPES : two females; one other specimen in fragments. Reg. no. 1921:10:18:1518-1521. Bona, Algeria. Collected by Meinert. Budde-Lund Collection.meinerti costata Budde-Lund (1885 : 198) [Bathytropa costata] SYNTYPES: two females ; one other damaged specimen. Reg. no. 1921:10:18:1512-1514.Mt Bona, Algeria. Collected by Meinert. Budde-Lund Collection.thermophila Dollfus (i8g6a : 28) [Now Trichorhina tomentosa (Budde-Lund)] SYNTYPES: two females. Reg. no. 1911:11:8:10547-48. Serres du Museum, Paris.Norman Collection (ex Dollfus Collection). 8o J. P. ELLIS AND R. J. LINCOLN BENTHANA Budde-Lund pauper (Jackson) (1926 : 194) [Philoscia (Benthana) pauper} SYNTYPE : one male ; two micropreparations. Reg. no. 1921:10:18:1688. Valparaiso,Chile. Budde-Lund Collection (ex Michaelson Collection, Mus. Hamburg).villosa (Jackson) (1926 : 195) [Philoscia (Benthana) villosa] HOLOTYPE : female ; one micropreparation. Reg. no. 1921:10:18:1674. Matucana, Peru.Budde-Lund Collection. BILAWRENCIA Vandel albicincta Vandel (1973 : 74) SYNTYPES : nine males ; nine females. Reg. no. 1970:414:18. Vulavu, Santa Isabel,Solomon Is. 7.10.1965 ; Village rubbish'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : one male ; one female. Reg. no. 1970:415:1. Fulakora Pt, Raja, SantaIsabel, Solomon Is. 30.9.1965; 'forest litter'. Collected by P. N. Lawrence. Presented bythe Royal Society. SYNTYPES : one male ; nine females. Reg. no. 1970:416:9. Cockatoo I., Santa Isabel,Solomon Is. 19-20.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : four males ; five females. Reg. no. 1970:417:13 (part). Tatamba, SantaIsabel, Solomon Is. 4.10.1965 ; 'mangrove litter'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPE : male. Reg. no. 1970:417:13 (part). Raja, N.E. Tatamba, Santa Isabel,Solomon Is. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : one male ; one female. Reg. 1970:417:13 (part). Tatamba, Santa Isabel,Solomon Is. 6.10.1965 ; 'native gardens'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : two males ; four females. Reg. no. 1970:418:6. Lilihinia I., Santa Isabel,Solomon Is. 21.10.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : one male ; four females. Reg. no. 1970:419:4. E. Central San Jorge, SantaIsabel, Solomon Is. 24.9.1965 ; 'forest litter'. Collected by J. Peake and P. N. Lawrence.Presented by the Royal Society. SYNTYPES : fifty specimens, males, females and juveniles. Reg. no. 1970:421:50. LungaBeach, Guadalcanal, Solomon Is. 31.8.1965; 'lagoon debris'. Collected by P. N. Lawrence.Presented by the Royal Society.Isabella Vandel (1973 : 78) SYNTYPES : one hundred and seven specimens, males, females and juveniles. Reg. no.1970:420:107. Vulavu, Santa Isabel, Solomon Is. 7.10.1965 ; 'village rubbish'. Collectedby P. N. Lawrence. Presented by the Royal Society. BURMONISCUS Collinge kempi Collinge (1916 : 127) PARATYPES : two males ; one female. Reg. no. 1919:4:26:500-502. Maosmai Cave,Cherrapunji, Assam, India. October 1914. c. 4000 ft. Collected by S. W. Kemp. Pre-sented by W. E. Collinge. TYPES OF TERRESTRIAL ISOPODS 81 CALMANESIA Collinge methueni Collinge (1922 : 109) SYNTYPES : one male ; three females. Reg. no. 1922:11:10:6-9. Forest of Folohoy, EastMadagascar. 1911. Collected by Herschell and Chauvin. Presented by P. A. Methuen. SYNTYPES : three females. Reg. no. 1922:11:10:10-12. Analmazotra, Eastern Forest,Madagascar. June 1911 ; 'under rotten logs'. Collected and presented by P. A. Methuen. CALYCUONISCUS Collinge bodkini Collinge (1915 : 509) SYNTYPES : twenty males ; fifteen females. Reg. no. 1919:4:26:520-539. BotanicGardens, Georgetown, Guiana ; 'under bark of trees'. Presented by W. E. Collinge. CHAETOPHILOSCIA Verhoeff balssi Verhoeff (i928c : 170) SYNTYPE : female. Reg. no. 1928:7:4:45. Glasshouse in Munich, Germany. VerhoeffCollection.cellar ia (Dollfus) (1884 : ?) [Philoscia cellana] SYNTYPE : male. Reg. no. 1892:12:6:5. Caves de Beaune, France. Collected by Andr6.Presented by J. D'Arcy Thompson. SYNTYPES: three males ; two females. Reg. no. 1911:11:8:10762-66. Caves de Beaune,France. Collected by E. Andre. Norman Collection (ex Dollfus Collection). SYNTYPES: two males ; two females. Reg. no. 1921:10:18:1739-1742. Caves de Beaune,France. Budde-Lund Collection (ex Paris Museum). SYNTYPES: three males ; two females. Reg. no. 1928:12:1:3667-3670. Caves de Beaune,France. Stebbing Collection (ex Dollfus Collection). SYNTYPES: three males. Reg. no. 1956:10:10:171-173. Caves de Beaune, France.Collected by E. Andre. Presented by University College, Dundee (ex Dollfus Collection).dorsalis Verhoeff (ig28c : 138) SYNTYPE : male. Reg. no. 1928:7:4:43. Italian Riviera. Verhoeff Collection.elongata (Dollfus) (1884 : ?) [Philoscia elongata] SYNTYPES : three females. Reg. no. 1911:11:8:10750-52. St Maxine, France. NormanCollection (ex Dollfus Collection).formosana Verhoeff (ig28a : 221) SYNTYPES : one male ; two females. Reg. no. 1928:7:4:4-6. Formosa (Taiwan). Ver-hoeff Collection.hastata Verhoeff (ig2ga : 133) SYNTYPE : female. Reg. no. 1928:7:4:44. Bulgaria. 12.9.1923. Collected by I.Buresch. Verhoeff Collection.meeusei Holthuis (1946 : 124) SYNTYPES: one male; two females. Reg. no. 1947:4:14:1-3. Victoria Regia House,Royal Botanic Gardens, Kew, England. 15.4.1936 ; 'under stones'. Collected by A. D. J.Meeuse. Presented by the Natural History Museum, Leiden.pallida Verhoeff (ig28c : 141) [Now Chaetophiloscia cellaria (Dollfus)] SYNTYPE : female. Reg. no. 1928:7:4:42. French Riviera. Verhoeff Collection. DETO Guerin armata Budde-Lund (1906 : 84) SYNTYPES: one male ; two females. Reg. no. 1921:10:18:1074-1076. St Paul I., IndianOcean. 26.4.1903. Budde-Lund Collection. 82 J. P. ELLIS AND R. J. LINCOLN marina (Chilton) (1885 : 464) [Philougria marina] SYNTYPE : female. Reg. no. 1921:10:18:1085. Coogee Bay, nr Sydney, New SouthWales. 30.12.1883 ; 'in rock pools'. Collected by C. Chilton. Budde-Lund Collection (exDundee Museum).robusta Budde-Lund (1906 : 87) SYNTYPE : fragments only. Reg. no. 1921:10:18:1086. Auckland I., Pacific Ocean.Budde-Lund Collection. DIACARA Budde-Lund elegans (Dollfus) (1895 : 186) [Alloniscus elegans] SYNTYPE : female. Reg. no. 1921:10:18:1673. Mtgne d'Ambre, Diego-Suarez, Madagas-car. Collected by Ch. Alluaud. Budde-Lund Collection (ex Dollfus Collection). DIDIMA Budde-Lund humilis Budde-Lund (1908 : 292) HOLOTYPE : female. Reg. no. 1921:10:18:1888. Antananarivo, Madagascar. Collectedby F. Sikora. Budde-Lund Collection. FORMOSOSCIA Verhoeff ocellata Verhoeff (ig28a : 219) SYNTYPE : female. Reg. no. 1928:7:4:9. Formosa. Verhoeff Collection. HALOPHILOSCIA Verhoeff adriatica Verhoeff (igoSa : 358) SYNTYPES : two females. Reg. no. 1921:6:10:70-71. Nr Gulf of Fiume (Rijeka), Yugo-slavia. Verhoeff Collection.adriatica ru.piu.rn Verhoeff (i93ia : 548) [Now Halophiloscia couchi (Kinahan)] SYNTYPE : female. Reg. no. 1930:5:26:46. Noli, Italy. Verhoeff Collection.jucorum Verhoeff (i93ob : 36) [Now Halophiloscia couchi (Kinahan)] SYNTYPE : female. Reg. no. 1930:5:26:45. Split, Yugoslavia. 1928. Verhoeff Collec-tion.gracilicornis Verhoeff (1939 : 218) [Now Halophiloscia hirsuta Verhoeff] SYNTYPE : female. Reg. no. 1938:7:7:61. Apulia, Lecce (?), Italy. Verhoeff Collection.hirsuta Verhoeff (ig28c : 132) SYNTYPE : female. Reg. no. 1931:4:27:12. St Maxime, Toscana, Italy. 25th April.Verhoeff Collection.tyrrhena Verhoeff (i928c : 131) SYNTYPES : four females ; one juvenile. Reg. no. 1928:7:4:35-39. Riviera. VerhoeffCollection. SYNTYPES : two males ; three females. Reg. no. 1937:7:6:100-104. Riviera. VerhoeffCollection. HANONISCUS Budde-Lund tuberculatus Budde-Lund (1912 : 42) SYNTYPES: one male; three females. Reg. no. 1921:10:18:5735-5738. W. Australia.Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 83 HIATONISCUS Barnard contractus Barnard (1932 : 285) SYNTYPES : four males ; three females. Reg. no. 1933:1:25:246-249. Swellendam,Langeberg Mts, Cape Province. 1925. Collected and presented by K. H. Barnard.griseus Barnard (1932 : 283) SYNTYPES : twenty-nine males ; twenty-eight females and juveniles. Reg. no. 1933:1:25:234-245. Table Mt, Cape Province, S. Africa. Barnard Collection. HORA Barnard damae Barnard (1932 : 230) SYNTYPES : two males ; one female. Reg. no. 1933:1:25:60-62. Swellendam, LangebergRange, Cape Province. 1925. 3500-4000 ft. Collected and presented by K. H. Barnard. ISABELLOSCIA Vandel heroldi Vandel (1973 : 54) SYNTYPES : four females ; five juveniles. Reg. no. 1970:427:9. Thousand Ships Bay,Lilihinia I., Santa Isabel, Solomon Is. 20.9.1965 ; 'forest litter'. Collected by P. N. Law-rence. Presented by the Royal Society. SYNTYPES: two males ; twelve females ; six juveniles. Reg. no. 1970:428:15. Tatamba,Santa Isabel, Solomon Is. 27.9.1965; 'litter under Casuarina'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : one male ; one female. Reg. no. 1970:429:2. Thousand Ships Bay, oppositeLilihinia I., S.E. Santa Isabel, Solomon Is. 20.9.1965 ; 'forest litter'. Collected by P. N.Lawrence. Presented by the Royal Society. ISCHIOSCIA Verhoeff debilis (Budde-Lund) (1893 : 121) [Philoscia debilis] SYNTYPES : four females. Reg. no. 1921:10:18:2236-2239. La Moka. August 1891.Collected by F. Meinert. Budde-Lund Collection. JAPANONISCUS Verhoeff balsii Verhoeff (i928b : 32) SYNTYPE : female. Reg. no. 1928:7:4:1. Aburatsubo, nr Misaki, Japan. 13.10.1904.Collected by H. Sauter. Verhoeff Collection. KRANTZIA Barnard poecila Barnard (1932 : 281) SYNTYPES : one male ; three females. Reg. no. 1933:3:18:7-11. Krantzkop, Natal, S.Africa. 1917. Presented and collected by K. H. Barnard. LABYRINTHASIUS Verhoeff graecus Verhoeff (igagc : 119) [Now Bathytropa granulata Aubert & Dollfus] SYNTYPE : female. Reg. no. 1930:5:6:26. Crete. 7.1.1929. Verhoeff Collection. 8 4 J. P. ELLIS AND R. J. LINCOLN LEPIDONISCUS Verhoeff germanicus carniolense Verhoeff (19280 : 127) SYNTYPE : female. Reg. no. 1928:7:4:28. Krain. Verhoeff Collection.pruinosus denticulatus Verhoeff (19280 : 126) SYNTYPES : two males ; four females. Reg. no. 1928:7:4:23-27. Italy.Collection. Verhoeff MARIONISCUS Barnard spatulifrons Barnard (1932 : 234) SYNTYPES : ten males ; twelve females. Reg. no. 1933:1:25:67-74.Peninsula, S. Africa. Collected and presented by K. H. Barnard. Hout Bay, Cape NAHIA Budde-Lund hirsuta (Budde-Lund) (1906 : 89) [Philoscia hirsuta] SYNTYPES : mixed with other (non-type) specimens collected by Schultze in Cape Town.Reg. no. 1921:10:18:2022-2033. Simonstown, S. Africa. 19.6.1903. Budde-Lund Collec-tion. OLIBRINUS Budde-Lund pigmentatus Budde-Lund (i9i3b : 390) SYNTYPES : two females. Reg. no. 1913:1:8:154-155. Coin, Peros, Chagos Archipelago,Indian Ocean. Collected by the Percy Sladen Trust Expedition. Presented by J . S. Gardiner. ONISCUS Linnaeus kenepurensis Chilton (1901 : 135) SYNTYPES: four males. Reg. no. 1900:11:1:43-46. Kenepuru, New Zealand. Presentedby C. Chilton.simoni Budde-Lund (1885 : 205) SYNTYPE : female. Reg. no. 1911:11:8:10650. St Jean de Luz, nr Bayonne, France.Collected by E. Simon. Norman Collection (ex Dollfus Collection). SYNTYPE : female. Reg. no. 1921:10:18:5689. Lac Marescot, nr Biarritz, France.Collected by E. Simon. Budde-Lund Collection. ORONISCVS Verhoeff dolomiticus Verhoeff (igoSa : 347) SYNTYPE : female. Reg. no. 1908:6:1:18. Tyrol, Austria. Verhoeff Collection. SYNTYPE : female. Reg. no. 1921:6:10:39. Tyrol, Austria. Verhoeff Collection. SYNTYPE : male (micropreparation) . Reg. no. 1931:4:27:100. Ampezzo, Italy. VerhoeffCollection. SYNTYPES : three females. Reg. no. 1937:7:6:121-123. Dolomite Mts. VerhoeffCollection. PAPUAPHILOSCIA Vandel bougainvillei Vandel (1973 : 57) SYNTYPE : one damaged specimen (abdomen missing). Reg. no. 1970:404:1. Popamani-siu, Guadalcanal, Solomon Is. 1-4.11.1965. 4400 ft ; 'mossy ridge, forest litter'. Collectedby P. N. Lawrence. Presented by the Royal Society. TYPES OF TERRESTRIAL ISOPODS 85 PAPUASONISCUS Vandel holthuisi Vandel (1973 : 25) SYNTYPES : male ; female. Reg. no. 1970:405:2. Raja, nr Fulakora Pt, Santa Isabel,Solomon Is. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPE : male. Reg. no. 1970:406:1. Cockatoo I., Santa Isabel, Solomon Is. 19-20.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : one male ; seven females. Reg. no. 1970:407:7. Lilihinia I., Santa Isabel,Solomon Is. 21.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : two males. Reg. no. 1970:408:16 (part). Tatamba, Santa Isabel, Solomon Is.28-29.9.1965 ; 'coconut and Casuarina litter'. Collected by P. N. Lawrence. Presented bythe Royal Society. SYNTYPE : female. Reg. no. 1970:408:16 (part). Tatamba, Santa Isabel, Solomon Is.27.9.1965 ; 'litter under Casuarina'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPE: female. Reg. no. 1970:408:16 (part). Tatamba, Santa Isabel, Solomon Is.4.10.1965 ; 'mangrove litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES: one male; one damaged female. Reg. no. 1970:408:16 (part). Tatamba,Santa Isabel, Solomon Is. 6.10.1965 ; 'native gardens'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : two males ; six females. Reg. no. 1970:408:16 (part). Raja, N.E. Tatamba,Santa Isabel, Solomon Is. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPES : two males ; one female. Reg. no. 1970:409:3. Opposite Cockatoo I., S.E.Santa Isabel, Solomon Is. 19-20.9.1965 ; 'shore litter'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : one male ; one female. Reg. no. 1970:410:2. Umasami River, 5 mis S.W.Tamboko, Guadalcanal, Solomon Is. 7-10.7.1965 ; 'Areca palm and litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:411:1. Mt Austern, nr Honiara, Guadalcanal, SolomonIs. 24.8.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPE : male. Reg. no. 1970:412:1. nr Mt Gallego, 6 mis S.W. Tamboko, Guadalcanal,Solomon Is. 4.7.1965 ; 'disturbed forest litter'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPES : two males ; one female. Reg. no. 1970:413:3. 7 mis S. of Wainoni, SanCristobal, Solomon Is. 2.7.1965 ; 'litter of palms, vines and ferns'. Collected by P. N.Lawrence. Presented by the Royal Society. PARAPHILOSCIA Stebbing armata Vandel (1973 : 99) SYNTYPES : two males ; eleven females. Reg. no. 1970:386:13. Mt Austen, nr Honiara,Guadalcanal, Solomon Is. 24.8.1965 ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society.lateralis (Budde-Lund) (i9i3b : 372) [Pseudophiloscia lateralis] SYNTYPES : four females. Reg. no. 1913:1:8:35-40. Silhouette, Seychelle Is. Collectedby the 'Sealark' Expedition. Presented by J. Stanley Gardiner. SYNTYPES: two males ; two females. Reg. no. 1913:1:8:41-44. Mt Sebert, Seychelle Is.2.12.1905. 1 800 ft. Collected by the 'Sealark' Expedition. Presented by J. StanleyGardiner. 86 J. P. ELLIS AND R. J. LINCOLN SYNTYPES : four females. Reg. no. 1913:1:8:45-47. Mt Alphonse, Cascade, Mahe I.Seychelle Is. 4.12.1905. 1800 ft. Collected by the 'Sealark' Expedition. Presented by J-Stanley Gardiner. SYNTYPES : six males ; ten females. Reg. no. 1921:10:18:981-992. Palm, Mahe I.,Seychelle Is. 2-8.5.1901. Collected by A. Brauer. Budde-Lund Collection.tnendanai Vandel (1973 : 93) SYNTYPES: sixty specimens : males, females and juveniles. Reg. no. 1970:399:60. Nuhu,Guadalcanal, Solomon Is. 28-31.10.1965. 1000 ft ; 'forest litter'. Collected by P. N.Lawrence. Presented by the Royal Society. SYNTYPES : seven females. Reg. no. 1970:400:20 (part). 5 miles S.W. Tamboko, RiverUmasami, Guadalcanal, Solomon Is. 7.7.1965. 1000 ft ; 'ridge, forest litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : three males ; two females. Reg. no. 1970:400:20 (part). S.W. end ofMonitor Creek, River Umasami, Guadalcanal, Solomon Is. 5.7.1965 ; 'forest litter in ahollow'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES: three males ; three females. Reg. no. 1970:400:20 (part), nr Monitor Creek,Umasami River, Guadalcanal, Solomon Is. 5.7.1965 ; 'forest litter'. Collected by P. N.Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:400:20 (part). Monitor Creek, River Umasami, Guadal-canal, Solomon Is. 5.7.1965 ; 'litter on mat of forest roots'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : two females. Reg. no. 1970:401:4. Mt Gallego, Guadalcanal, Solomon Is.12.7.1965 ; 'ridge forest litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPE : female. Reg. no. 1970:401:4 (part). Mt Gallego, Guadalcanal, Solomon Is.12.7.1965. 3000 ft ; 'moss forest'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES : two males ; three females. Reg. no. 1970:402:5. nr Mt Gallego, 6 mis S.W.Tamboko, Guadalcanal, Solomon Is. 4.7.1965 ; 'disturbed forest'. Collected by P. N. Law-rence. Presented by the Royal Society. SYNTYPE: male. Reg. no. 1970:403:1. Tambeluse, Guadalcanal, Solomon Is. 1.11.1965.c. 1500 ft ; 'dry ridge litter'. Collected by P. N. Lawrence. Presented by the Royal Society.propinqua Vandel (1973 : 90) SYNTYPES : thirty-six specimens : males, females and juveniles. Reg. no. 1970:390:36.nr Kuzi, Kolombangara, Solomon Is. 3-9.9.1965 ; 'forest litter' and 'coral limestone'.Collected by P. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : thirty-five specimens : males, females and juveniles. Reg. no. 1974:59:35.nr Kuzi, Kolombangara, Solomon Is. 3-9.9.1965 ; 'forest litter'. Collected by P. N. Law-rence. Presented by the Royal Society.sancristobali Vandel (1973 : 100) SYNTYPES: two males ; nine females ; four damaged adults. Reg. no. 1970:387:22 (part).Confluence of Warahito and Pagato Rivers, San Cristobal, Solomon Is. 23 and 29.7.1965 ;'rotten wood and forest litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES : one male ; three females. Reg. no. 1970:387:22 (part). 4 miles up Warahitofrom River Pagato, San Cristobal, Solomon Is. 4.8.1965 ; 'forest litter'. Collected by Leoneand Isiah. Presented by the Royal Society. SYNTYPES : one male ; three females. Reg. no. 1970:387:22 (part). Confluence of Wara-hito and Pagato Rivers, San Cristobal, Solomon Is. 1.8.1965; 'stream litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : two females. Reg. no. 1970:388:6 (part), c. 6-75 miles S. Wainoni, SanCristobal, Solomon Is. 24.7.1965; 'forest litter, ridge bottom'. Collected by P. N. Lawrence.Presented by the Royal Society. TYPES OF TERRESTRIAL ISOPODS 87 SYNTYPE : male. Reg. no. 1970:388:6 (part). S.E. Wainoni, San Cristobal, Solomon Is.10.8.1965. 1650 ft ; 'moss forest'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES : one male; two females. Reg. no. 1970:388:6 (part), nr Wainoni, SanCristobal, Solomon Is. 8.8.1965. 1000 ft ; 'ridge litter'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : three males ; four females. Reg. no. 1970:389:8. Huni River estuary, N.E.Wainoni, San Cristobal, Solomon Is. 9-12.8.1965 ; 'forest litter on coral limestone'. Col-lected by P. N. Lawrence. Presented by the Royal Society.santaisabellae Vandel (1973 : 92) SYNTYPES : three males ; nine females. Reg. no. 1970:391:10. Thousand Ships Bay,opposite Lilihinia I., Santa Isabel, Solomon Is. 20.9.1965 ; 'forest litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : seventy-two specimens : males, females and juveniles. Reg. no. 1970:392:72.N.E. Tatamba, Raja, Santa Isabel, Solomon Is. 30.9.1965 ; 'forest litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : six males ; thirteen females. Reg. no. 1970:393:8. Fulakora Pt, Raja,Solomon Is. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : one male ; three females. Reg. no. 1970:394:4. Cockatoo I., Santa Isabel,Solomon Is. 19-20.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : four males ; eight females. Reg. no. 1970:395:12. Lilihinia I., Santa Isabel,Solomon Is. 21.9.1965 ; 'shore litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPE : female. Reg. no. 1970:396:1. S.E. San Jorge, S. of Santa Isabel, Solomon Is.22.9.1965 ; 'gulley litter'. Collected by J. Peake and P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : three females. Reg. no. 1970:397:3. E. central San Jorge I., Santa Isabel,Solomon Is. 24.9.1965 ; 'forest litter'. Collected by J. Peake and P. N. Lawrence. Pre-sented by the Royal Society. PHALLONISCUS Budde-Lund anomalus (Dollfus) (1890 : 4) [Philoscia anomala] SYNTYPES: three males; two females. Reg. no. 1927:5:14:29-38 (part). Valparaiso,Chile. November 1875. 'Challenger' Collection. SYNTYPES : six males ; four females. Reg. no. 1927:5:14:29-38 (part). Juan FernandezI., Pacific Ocean, 'beach'. 'Challenger' Collection.pygmaeus (Budde-Lund) (1885 : 212) [Philoscia pygmaea] SYNTYPES: two females (in fragments). Reg. no. 1921:10:18:1961-1962. Corsica.Collected by E. Simon. Budde-Lund Collection. PHILOSCIA Latreille afflnis Verhoeff (igoSa : 352) [Philoscia muscorum affinis] SYNTYPES: seven females. Reg. no. 1921:6:10:61-65. Southern Italy. VerhoeffCollection. SYNTYPES : seven females. Reg. no. 1928:7:4:29-33. Italian Riviera. Verhoeff Collec-tion. SYNTYPE : female. Reg. no. 1931:4:27:72. Toscana region, Italy. Verhoeff Collection.daltnatica Verhoeff (igoic : 146) [Philoscia muscorum dalmatica] SYNTYPES : two males. Reg. no. 1901:9:19:55-56. Dalmatia, Yugoslavia. VerhoeffCollection. 88 J. P. ELLIS AND R. J. LINCOLN dilectum Collinge (1917 : 579) [Now Aphiloscia vilis (Budde-Lund)] PARATYPES : seven males ; nine females. Reg. no. 1919:4:26:459-468. Pentrich, nrPietermaritzburg, Natal, S. Africa. 14.9.1915. Collected by C. Akerman. Collinge Collec-tion.ditninuta Budde-Lund (1893 : I2 ) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:1785-1786. La Moka, August1891. Collected by F. Meinert. Budde-Lund Collection.flava Budde-Lund (igisa : 70) HOLOTYPE : female. Reg. no. 1921:10:18:1878. Victoria, Australia. Collected byHauschild. Budde-Lund Collection.gravosensis Verhoeff (igoic : 145) SYNTYPE : female. Reg. no. 1901:9:19:54. Dalmatia, Yugoslavia. Verhoeff Collection.guernei Dollfus (1887 : 195) SYNTYPE : female. Reg. no. 1911:11:8:10772. Caldeira volcano, Faial I., Azores.16.7.1887. Norman Collection.longicornis Budde-Lund (1885 : 221) [Now Halophiloscia couchi (Kinahan)] SYNTYPES: four males; one female. Reg. no. 1921:10:18:2232-2235. Algiers. Col-lected by E. Simon. Budde-Lund Collection.muscorutn biellensis Verhoeff (1935 : 108) SYNTYPES : one male ; two females. Reg. no. 1937:7:6:43-45. Piemonte region, Italy.Verhoeff Collection.muscorum frigidana Verhoeff (i928c : 136) SYNTYPE : female. Reg. no. 1928:7:4:34. Italian Riviera. Verhoeff Collection.muscorum triangulifera Verhoeff (1918 : 157) SYNTYPES : two females. Reg. no. 1921:6:10:68-69. Italian Riviera. Verhoeff Collec-tion.nitida (Miers) (1877 : 670) [Philougria nitida] SYNTYPES : six females. Reg. no. 1879:21. Peru or Guiana. Presented by E. J. Miers(ex Wrzesniowsky Collection).patienci Bagnall (1908 : 429) SYNTYPES : five females. Reg. no. 1911:6:6:98-102. Botanical Gardens, Kew, Surrey,'from West Indian plants'. Presented by R. S. Bagnall.pulchella Budde-Lund (1885 : 214) [Now Chaetophiloscia elongata (Dollfus)] SYNTYPES : three males. Reg. no. 1921:10:18:2017-2019. No locality. Budde-LundCollection.seriepunctata Budde-Lund (1893 : I22 ) HOLOTYPE : female. Reg. no. 1921:10:18:1964. Caracas, Venezuela. 14.7.1891. Col-lected by Fr. Meinert. Budde-Lund Collection.subterranea Budde-Lund (1912 : 40) HOLOTYPE: few fragments only. Reg. no. 1921:10:18:1965. Yallingup, W. Australia.5.10.1905. Budde-Lund Collection.vittata Say (1818 : 429) SYNTYPE : female (dry). Reg. no. 1973:511:1. United States. Presented by T. Say.warreni Collinge (1917 : 578) [Now Nahia hirsuta (Budde-Lund)] PARATYPES : three males ; nine females. Reg. no. 1919:4:26:469-478. Umbilo Bush, nrDurban, Natal, S. Africa. 16.9.1915. Collected by E. Warren. Collinge Collection. Subgenus BENTHANOPS Barnard fulva Barnard (1932 : 247) SYNTYPES : eleven males ; thirty-nine females. Reg. no. 1933:1:25:118-129. Table Mt,Cape Peninsula, S. Africa. Barnard Collection. TYPES OF TERRESTRIAL ISOPODS 89 Subgenus KOMA TIA Barnard marginata Barnard (1932 : 240) SYNTYPE : male. Reg. no. 1933:1:25:86. Wanetsi River, Portuguese East Africa.Collected by H. W. Bell-Marley. Barnard Collection. PLYMOPHILOSCIA Warhberg montana Verhoeff (ig26a : 335) SYNTYPE : female. Reg. no. 1928:7:4:20. New Caledonia. Verhoeff Collection. PSEUDOPHILOSCIA Budde-Lund angustissima Budde-Lund (i9i3a : 373) SYNTYPES : one male ; one female. Reg. no. 1913:1:8:48-49. Mt Alphonse, Mahe I.,Seychelle Is. 3.12.1905. Collected by the 'Sealark' Expedition. Presented by J. StanleyGardiner. SYNTYPES : one male ; three females. Reg. no. 1921:10:18:973-976. Palm, Seychelle Is.2.5.1901. Collected by A. Brauer. Budde-Lund Collection.brevicornis Budde-Lund (igisa : 374) SYNTYPE : female. Reg. no. 1921:10:18:977. New Zealand. Budde-Lund Collection.fragilis Budde-Lund (1904 : 43) HOLOTYPE : few fragments only. Reg. no. 1921:10:18:978. Howick, New Zealand.Budde-Lund Collection.inflexa Budde-Lund (1904 : 43) SYNTYPES: one male; one female. Reg. no. 1921:10:18:979-980. Corral, Chile. October1894. Collected by Plate. Budde-Lund Collection. RENNELLOSCIA Vandel tnacrocephala Vandel (1973 : 48) SYNTYPES: five females; two juveniles. Reg. no. 1970:385:8. N. of Kuzi, Kolombangara,Solomon Is. 6.10.1965. 500 ft ; 'forest litter'. Collected by P. N. Lawrence. Presentedby the Royal Society.novabritannica Vandel (1973 : 32) SYNTYPES : two females. Reg. no. 1970:362:2. Thousand Ships Bay, opposite Lilihinia I.,Santa Isabel, Solomon Is. 20.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPES: two males; fourteen females. Reg. no. 1970:363:16. Mt Austern, nrHoniara, Guadalcanal, Solomon Is. 24.7.1965 ; 'forest litter'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : two males ; four females ; one damaged adult. Reg. no. 1970:364:7. Nuhu,Guadalcanal, Solomon Is. 28.10.1965. 1000 ft ; 'forest litter'. Collected by P. N. Law-rence. Presented by the Royal Society. SYNTYPES : one female ; one juvenile. Reg. no. 1970:365:13 (part). Popamanisiu,Guadalcanal, Solomon Is. 1-4.11.1965. 4400 ft ; 'mossy ridge, forest litter'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : three males ; four females ; three juveniles ; one damaged adult. Reg. no.1970:365:13 (part). Popamanisiu, Guadalcanal, Solomon Is. 6-8.11.1965. 7000 ft ; 'forestlitter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:366:24 (part). Umasami River, nr Monitor Creek,Guadalcanal, Solomon Is. 5.7.1965 ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. go J. P. ELLIS AND R. J. LINCOLN SYNTYPES : six males ; fourteen females ; four juveniles ; one damaged adult. Reg. no.1970:366:24 (part). Umasami River, 5 miles S.W. Tamboko, Guadalcanal, Solomon Is.7-10.7.1965 ; ' Areca palm and litter'. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPES : two males ; one female. Reg. no. 1970:367:3. nr Mt Gallego, 6 miles S.W.Tamboko, Solomon Is. 4.7.1965 ; 'disturbed forest'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPE : female. Reg. no. 1970:368:1. Lunga Beach, Guadalcanal, Solomon Is.31.8.1965 ; 'dry grass litter'. Collected by P. N. Lawrence. Presented by the RoyalSociety. SYNTYPES : two females ; two juveniles. Reg. no. 1970:369:5 (part). Popamanisiu,Guadalcanal, Solomon Is. 8-9.11.1965. 5700 ft ; 'forest litter'. Collected by P. N. Law-rence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:369:5 (part). Popamanisiu, Guadalcanal, Solomon Is.5.11.1965. 7000 ft ; 'bog masses around trees'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPES : two females. Reg. no. 1970:370:3 (part). Hidden Valley, nr Mt Gallego,Guadalcanal, Solomon Is. 3.7.1965. Collected by P. N. Lawrence. Presented by theRoyal Society. SYNTYPE : male. Reg. no. 1970:370:3 (part). Summit of Mt Gallego, Guadalcanal,Solomon Is. 12.7.1965. 3600 ft ; 'fern litter and forest mosses'. Collected by P. N.Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:371:1. Lunga Beach, Guadalcanal, Solomon Is.31.8.1965 ; 'tidal debris'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:372:1. Nuhu, Guadalcanal, Solomon Is. 28-31.10.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : one male ; two juveniles. Reg. no. 1970:373:3. Umasami River, c. 6 milesS.W. Tamboko, Guadalcanal, Solomon Is. 1.7.1965 ; 'forest litter, sandy soil'. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : twenty-five specimens : males, females and juveniles. Reg. no. 1970:374:25.6-10 miles S. of Wainoni, San Cristobal, Solomon Is. July and August 1965. Collected byP. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : five males ; six females ; six juveniles. Reg. no. 1970:375:18. Huni RiverEstuary, N.E. Wainoni, San Cristobal, Solomon Is. 9-12.8.1965 ; 'forest litter on corallimestone'. Collected by Isiah. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:376:1. Warahito-Pagato confluence, San Cristobal..Solomon Is. 1.8.1965. 330 ft ; 'arboreal litter'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPE : female. Reg. no. 1970:377:29 (part). 4 miles up Warahito River, from PagatoRiver, San Cristobal, Solomon Is. 4.8.1965 ; 'forest litter'. Collected by Leone and Isiah.Presented by the Royal Society. SYNTYPES : two males. Reg. no. 1970:377:29 (part). Warahito-Pagato confluence, SanCristobal, Solomon Is. 1.8.1965 ; 'stream litter'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPES : twenty-six specimens : males, females and juveniles. Reg. no. 1970:377:29(part). Warahito-Pagato confluence, San Cristobal, Solomon Is. 23-29.7.1965 ; 'rottenwood and forest litter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : thirty specimens : males, females and juveniles. Reg. no. 1970:378:30.Wainoni and environs, San Cristobal, Solomon Is. July and August, 1965. Collected byP. N. Lawrence. Presented by the Royal Society. SYNTYPES : twenty specimens : males, females and juveniles. Reg. no. 1970:379:22. nrKuzi, Kolombangara, Solomon Is. 8.9.1965. 50 ft ; 'valley litter'. Collected by P. N.Lawrence. Presented by the Royal Society. TYPES OF TERRESTRIAL ISOPODS 91 SYNTYPES : three males ; seven females. Reg. no. 1970:380:18. nr Kuzi, Kolombangara,Solomon Is. 6.9.1965. 250 and 500 ft ; 'forest litter'. Collected by P. N. Lawrence.Presented by the Royal Society. SYNTYPES : thirty-four specimens : males, females and juveniles. Reg. no. 1970:381:34.nr Kuzi, Kolombangara, Solomon Is. 3-9.9.1965 ; 'forest litter'. Collected by P. N.Lawrence. Presented by the Royal Society. SYNTYPES : three males ; nine females. Reg. no. 1970:383:9. Ngaliau Hill, Pawa, Ugi,Solomon Is. 20.7.1965 ; 'secondary forest litter'. Collected by P. N. Lawrence. Presentedby the Royal Society. SYNTYPES : three males ; twelve females. Reg. no. 1970:383:15. N. of Kiai, Kolomban-gara, Solomon Is. 6.9.1965. 1000 ft ; 'forest litter'. Collected by P. N. Lawrence. Pre-sented by the Royal Society. SYNTYPES : one male ; six females. Reg. no. 1970:60:7. N. of Kuzi, Kolombangara,Solomon Is. 4.9.1965. 1500 ft ; 'litter mossy wood'. Collected by P. N. Lawrence.Presented by the Royal Society. SCYPHAX Dana intermedius Miers (1876 : 227) [Now Scyphax ornatus (Dana)] HOLOTYPE : female. Reg. no. 1973:476:1. New Zealand. Donor unknown. SCYPHONISCUS Chilton waitatensis Chilton (1901 : 128) SYNTYPE : male. Reg. no. 1900:11:1:51-52. Blueskin Bay, Otago, New Zealand.Presented by C. Chilton. SYNTYPE : male. Reg. no. 1921:10:18:1067. Blueskin Bay, Otago, New Zealand.Budde-Lund Collection (ex Dundee Museum Collection). SETAPHORA Budde-Lund angusticauda (Budde-Lund) (1885 : 216) [Philoscia angusticaudd] SYNTYPES: one male; three females. Reg. no. 1921:10:18:2247-2250. Borneo, Indonesia.Budde-Lund Collection.cingulata (Barnard) (1932 : 244) [Philoscia (Setaphora) cingulata] SYNTYPES : one male ; nine females. Reg. no. 1933:1:25:87-92. Port Shepstone, Natal,S. Africa. 1912. Collected and presented by K. H. Barnard.coeca (Budde-Lund) (1895 : 611) [Philoscia caeca] SYNTYPE : fragments only. Reg. no. 1921:10:18:2251. Moulmein, Burma. January1887. Collected by L. Fea. Budde-Lund Collection.cotnta (Budde-Lund) (1895 : 611) [Philoscia comta] SYNTYPE: male. Reg. no. 1921:10:18:2252. Mt Carin, Asciuii Ghecu, 1300-1400 m.Collected by L. Fea. Budde-Lund Collection.demarcata (Barnard) (1932 : 244) [Philoscia (Setaphora) demarcata] SYNTYPES: four males; three females. Reg. no. 1933:1:25:93-97. Pietermaritzburg,Natal, S. Africa. 1917. Collected and presented by K. H. Barnard.fasciata (Jackson) (1933 : 151) [Philoscia (Setaphora ?) fasciata~\ SYNTYPES : one male ; three females. Reg. no. 1933:12:20:3-6. Penau Ridge, Uahuka,Marquesas Is. 5.3.1931. 2000 ft ; 'in moss'. Presented by H. G. Jackson. SYNTYPES: two males; two females. Reg. no. 1952:4:18:109-111. Hanamiai Valley,Tahuata, Marquesas Is. Presented by H. G. Jackson. 92 J. P. ELLIS AND R. J. LINCOLN lubricata (Budde-Lund) (1895 : 610) [Philoscia lubricata] SYNTYPES : two females. Reg. no. 1921:10:18:2264-2265. Burma. Budde-Lund Collec-tion.mina (Budde-Lund) (1885 : 219) [Philoscia mina] SYNTYPES : two males ; one female. Reg. no. 1921:10:18:1899-1901. Cape, S. Africa.Collected by Drege. Budde-Lund Collection.ovata Budde-Lund (19130 : 386) SYNTYPES: seven males; eight females. Reg. no. 1921:10:18:2291-2303. Seychelles,Indian Ocean. May 1901. Collected by A. Brauer. Budde-Lund Collection.pallidemaculata Budde-Lund (igi^b : 387) SYNTYPES : two males ; two females. Reg. no. 1913:1:8:132-136. Mt Alphonse, Cascade,Mah6 I., Seychelles. 4.12.1905. 1800 ft. Collected by the 'Sealark' Expedition. Pre-sented by J. S. Gardiner. SYNTYPES : one male ; four females. Reg. no. 1913:1:8:137-140. Mahe" I., Seychelles.3.12.1905. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. SYNTYPES: ten males ; nine females. Reg. no. 1921:10:18:2304-2315. Palm, Seychelles,Indian Ocean. April 1901. Collected by A. Brauer. Budde-Lund Collection.pilosa Budde-Lund (igisb : 388) SYNTYPES: five males; one female. Reg. no. 1913:1:8:144-149. Salomon Is., ChagosArchipelago, Indian Ocean. Collected by the 'Sealark' Expedition. Presented by J. S.Gardiner. SYNTYPES: four males; six females. Reg. no. 1921:10:18:2328-2337. Salomon Is.,Chagos Archipelago, Indian Ocean. Collected by the 'Sealark' Expedition. Budde-LundCollection.rafflesi (Jackson) (1936 : 77) [Philoscia (Setaphora) rafflesi] SYNTYPES : two males. Reg. no. 1938:5:23:9-10. nr River Yum, Plus Valley, Perak,Malaysia. 1933. Presented by the Raffles Museum.suarezia (Dollfus) (1895 : 186) [Philoscia suarezia] SYNTYPES : specimens mixed with others from Nossi-Be collected by Voeltzkow. Reg. no.1921:10:18:2351-2358. Diego-Suarez, Madagascar. Budde-Lund Collection (ex DollfusCollection) .truncatella (Budde-Lund) (1902 : 379) [Philoscia truncatella] SYNTYPE : few fragments only. Reg. no. 1921:10:18:2383. Malay Peninsula. Collectedby the 'Skeat' Expedition. Budde-Lund Collection. STENOPHILOSCIA Verhoeff dalmatica Verhoeff (i93ob : 38) SYNTYPE : female. Reg. no. 1920:5:26:44. Dalmatia, Yugoslavia. Verhoeff Collection.SYNTYPE : micropreparation. Reg. no. 1931:4:27:106. Split, Dalmatia, Yugoslavia.Verhoeff Collection.glarearum Verhoeff (igoSa : 359) SYNTYPES : two males ; one female. Reg. no. 1921:6:10:72-74. Sicily. VerhoeffCollection. TIROLOSCIA Verhoeff Corsica (Dollfus) (1888 : 10) [Philoscia Corsica] SYNTYPES: three females. Reg. no. 1911:11:8:10767-10769. Gravone River, Vizzavona,Corsica. July 1881. Collected by E. Che vreux. Norman Collection (ex Dollfus Collection). SYNTYPES : one male ; two females. Reg. no. 1921:10:18:1750-1752. Vizzavona,Corsica. Budde-Lund Collection (ex Dollfus Collection). TYPES OF TERRESTRIAL ISOPODS 93 elbana Verhoeff (i93ia : 542) SYNTYPE : female. Reg. no. 1931:4:27:9. Elba I. Verhoeff Collection.SYNTYPE : female (micropreparation). Reg. no. 1931:4:27:81. Populonia, Italy. Ver-hoeff Collection.esterelana (Verhoeff) (1918 : 155) [Philoscia (Paraphiloscia) esterelana] SYNTYPE : female. Reg. no. 1921:6:10:67. Esterel, France. Verhoeff Collection.macchiae Verhoeff (1931 a : 543) [Now Tiroloscia Corsica (Dollfus)] SYNTYPE : female. Reg. no. 1931:4:27:10. Elba I. Verhoeff Collection.pyrenaica (Dollfus) (1897 : n) [Philoscia pyrenaica] SYNTYPE: female. Reg. no. 1911:11:8:10770. Ahusquy, nr Mauleon, France. Collectedby H. Broelemann. Norman Collection (ex Dollfus Collection).squamuligera bargensis Verhoeff (1935 : 106) SYNTYPES : four females. Reg. no. 1937:7:6:105-108. Apennine Mts, Italy. VerhoeffCollection.squamuligera briani (Verhoeff) (193 la : 545) [Philoscia (Tiroloscia) squamuligera briani] SYNTYPE : female. Reg. no. 1931:4:27:8. Piemonte region, Italy. Verhoeff Collection.squamuligera tendana Verhoeff (i93ia : 545) [Now Tiroloscia exigua exigua (Budde-Lund)] SYNTYPE : female. Reg. no. 1931:4:27:7. Limone ('Seealpen'), Italy. Verhoeff Collec-tion. REFERENCES 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.BAGNALL, R. S. 1908. On Philoscia patiencei, sp. n. a new terrestrial isopod. Ann. Mag. nat. Hist. (8) 1 : 428-431.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. n. Terrestrial Isopoda. Ann. S. Afr. Mus. 30 (2) : 179-388. 1936. Terrestrial isopods and amphipods from Mauritius. Ann. Natal Mus. 8 (i) : 1-17. BLAKE, C. H. 1931. New land isopods from New England. Occ. Pap. Boston Soc. nat. Hist. 5: 34 J -348- BUDDE-LUND, G. 1885. Crustacea Isopoda Terrestria, per familias et genera et species descripta.3i9pp. Hauniae. - 1893. Landisopoder fra Venezuela, insamlede af Dr. Fr. Meinert. Ent. Meddr. 4 : m-129. 1895. Viaggio di Leonardo Fea in Birmania e regioni vicine. 64. Isopodi terrestri. Ann. Mus. Genova, 34 : 602-612.1902. A list of the terrestrial isopods. In : Lanchester, W. F. On the Crustacea collected during the 'Skeat' Expedition to the Malay Peninsula. Pt II. Proc. zool. Soc. Lond., pp. 379-3 81 -1904. A Revision of 'Crustacea Isopoda Terrestria' with additions and illustrations. 2. Spherilloninae. 3. Armadillo, pp. 33-144. Kj0benhaven. - 1906. Die Landisopoden der Deutschen Siidpolar-Expedition 1901-1903 mit Diagnosenverwandter Arten. Deutsche Siidpolar-Expedition 1901-1903, 9, Zoologie i : 69-92.Berlin. - 1908. Isopoda von Madagascar und Ostafrika. Mit Diagnosen verwandter Arten.Wiss. Ergebn. Reise Ostafr. 1903-1905 von A. Voeltzkow. 2. Systematische Arbeiten, pp.265-310. Stuttgart. - 1909. Land-Isopoden. In : Schultze, L. Zoologische und anthropologische Ergebnisseeiner Forschungriese im Westlichen und Zentralen Siidafrika. II. Systematik und Tiero-geographie. Denkschr. med.-naturw. Ges. Jena, 15 : 53 -70. 94 J- P- ELLIS AND R. J. LINCOLN BuoDE-LuND, G. 1912. Oniscoidea, nachgelassenes Fragment. Die Fuana Sudwest Aust-raliens herausgegeben von Prof. Dr. W. Michaelsen und Dr. R. Hartmeyer, 4 (2) : 17-44.Jena. iQisa. t)ber einige Oniscoideen von Australien, nachgelassenes Fragment. Mitt, naturh.Mus. Hamb. 30 : 65-72. igisb. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under theleadership of Mr. J. Stanley Gardiner. IV. No. XXII. Terrestrial Isopoda, particularlyconsidered in relation to the distribution of southern Indo-Pacific species. Trans. Linn.Soc. Lond., Zool. (2) 15 : 367-394. CARL, J. 1908. Monographic der schweizerischen Isopoden. Neue Denkschr. schweiz. naturf. Ges. 42 (2) : 111-242.CHILTON, C. 1885. On a marine species of Philougria. Proc. Linn. Soc. N.S.W. 9 (3) : 463-466. 1901. The terrestrial Isopoda of New Zealand. Trans. Linn. Soc. Lond., Zool. (2) 8 : 99-152- 1911. The Crustacea of the Kermadec Islands. Trans. N.Z. Inst. 43 : 544-573.COLLINGE, W. E. 1915. Description of a new genus and species of terrestrial isopod from British Guiana. /. Linn. Soc. (Zool.), 32 : 509-511. 1916. Contributions to a knowledge of the terrestrial Isopoda of India. Pt II. Rec.Indian Mus. 12 (3) : 115-128. 1917. Contributions to a knowledge of the terrestrial Isopoda of Natal. Pt I. Ann.Natal Mus. 3 (3) : 567-585. 1920. Contributions to a knowledge of the terrestrial Isopoda of Natal. Pt III. Ann.Natal Mus. 4 : 471-490. 1922. On two new terrestrial isopods from Madagascar. /. Linn. Soc. (Zool.), 35 : 107- ii3-DAHL, F. 1919. Reihenfange und die Oekologie der deutschen Landisopoden. Zool. Anz. 50 : 193-203, 209-218.DOLLFUS. A. 1884. Les especes fran9aises du genre Philoscia Latreille (Crustaces Isopodes du groupe des Cloportides) . Bull. Soc. Etud. scient. Paris, 7 : 1-4. 1887. In : De Guerne, J. Notes sur la faune des A9ores. Naturaliste, (2) 1 : 195. 1888. Description d'une espece nouvelle du genre Philoscia. Bull. Soc. Etud. scient.Paris, 11 : 10-11. 1889. Isopodes terrestres recueillis aux A9ores en 1887, 1888 et 1889 par MM. Dr Th.Barrois et le Lieutenant Chaves. Revue biol. N. Fr. 1 (8) : 1-3. 1890. Isopodes terrestres de 'Challenger'. Bull. Soc. Etud. scient. Paris, 12 : 1-8. - 1893. Catalogue raisonne des isopodes terrestres de 1'Espagne (icr Supplement). An.Soc. esp. Hist. nat. 22 : 47-51. 1895. Mission scientifique de M. Ch. Alluaud dans le territoire de Diego-Suarez (Madagas-car-Nord), Avril-Aout 1853 - isopodes terrestres recueillis a Diego-Suarez, a Tamatave et ala Reunion. Mem. Soc. zool. Fr. 8 : 180-187. i896a. Crustaces isopodes receuillis dans les serres du Museum. Bull. Mus. natn. Hist. nat. Paris, 2 : 27-28. i896b. Crustaces isopodes de la Sicile. Notes Faunistiques, Paris, pp. 1-16. 1897. Tableau inconographique des Philoscia d'Europe. Crustaces isopodes terrestres.Feuille jeun. Nat. 27 : 1-13. 1900. Crustacea Isopoda. In : Fauna Hawaiiensis, or the Zoology of the Sandwich(Hawaiian) Islands, 2 (5) : 521-526. Cambridge. HOLTHUIS, L. B. 1946. On a small collection of isopod Crustacea from the greenhouses of theRoyal Botanic Gardens, Kew. Ann. Mag. nat. Hist, (n) 13 : 122-137. JACKSON, H. G. 1922. A revision of the isopod genus Ligia (Fabricius) . Proc. zool. Soc. Lond.,pp. 683-703. 1923. A revision of the isopod genus Ligidium (Brandt). Proc. zool. Soc. Lond., pp.823-839. TYPES OF TERRESTRIAL ISOPODS 95 JACKSON, H. G. 1926. Woodlice from Spain and Portugal, with an account of Benthana, asub-genus of Philoscia - Crustacea. Proc. zool. Soc. Lond., pt. i : 183-201. - 1927. A new subgenus of Ligia, with further observations on the genus. Ann. Mag. nat.Hist. (9) 19: 129-136. 1933. Marquesan terrestrial Isopoda. Pacific Entomological Survey Publication 7, article 10. Bull. Bernice P. Bishop Mus. No. 114 : 145-162. - 1936. Terrestrial isopods from Malaysia. Bull. Raffles Mus. No. 12 : 77-87.KOCH, C. L. 1838. Deutschlands Crustaceen, Myriapoden und Arachniden. Ein Beitrag zur deutschen Fauna. Heft 22. Regensberg. - 1841. ibid., Heft 34. LINCOLN, R. J. & ELLIS, J. P. 1974- Catalogue of the types of terrestrial Isopods (Oniscoidea)in the collections of the British Museum (Natural History). I. Super- family Pseudo-tracheata. Bull. Br. Mus. nat. Hist. (Zool.) 27 (5) : 189-246.MIERS, E. J. 1876. Descriptions of some new species of Crustacea, chiefly from New Zealand. Ann. Mag. nat. Hist. (4) 17 : 218-229.1877. On a collection of Crustacea, Decapoda and Isopoda, chiefly from South America, with descriptions of new genera and species. Proc. zool. Soc. Lond., pp. 653-679.PATIENCE, A. i9O7a. On a new British terrestrial isopod. Ann. Scot. nat. Hist. pp. 85-88.On a new British terrestrial isopod. /. Linn. Soc. (Zool.), 30 : 42-44. RACOVITZA, E. G. 1907. Biospeologica. IV. Isopodes terrestres (premiere serie). ArchsZool. exp.-gen. (4) 7 : 145-225. - 1908. Biospeologica. IX. Isopodes terrestres (seconde serie). Archs Zool. exp.-gen.(4) 9 : 239-415. SARS, G. O. 1899. An Account of the Crustacea of Norway. II. Isopoda, pp. 1-270. Bergen.SAY, T. 1818. An account of the Crustacea of the United States. /. Acad. nat. Sci. Philad. 1 : 423-444.STEBBING, T. R. R. 1908. Zoological results of the third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904-1905. Report on the Isopoda Terrestria. Proc. zool. Soc. Lond. pp. 554-560.VANDEL, A. 1952. Biospeologica. LXXIII. Isopodes terrestres (troisieme serie). Archs Zool. exp.-gen. 88 : 231-262.1959. Description d'une nouvelle espece de 1'Afrique Occidentale appartenant au genre Niambia Budde-Lund. Bull. Mus. Hist. nat. Paris (2) 31 : 516-519. - 1960. Faune de France. Vol.64. Isopodes terrestres (Premiere Partie), pp. 1-416. Paris. - 1973- Les isopodes terrestres (Oniscoidea) de la Melan^sie. Zool. Verh. No. 125 : pp. 160.VERHOEFF, K. W. 1900. Uber palaarktische Isopoden. (2. Isopoden-Aufsatz.) Zool. Anz. 23 : 117-130. - igoia. Uber palaarktische Isopoden. (3. Isopoden-Aufsatz.) Zool. Anz. 24 : 33-41. - igoib. Uber palaarktische Isopoden. (4. Aufsatz.) Zool. Anz. 24 : 66-72, 73-79- - igoic. Uber palaarktische Isopoden. (5. Aufsatz.) Zool. Anz. 24 : 135-149. - igoSa. Uber Isopoden. (15. Isopoden-Aufsatz.) Arch, biontol. 2 : 335-387. - igoSb. Uber Isopoden. (12. Isopoden-Aufsatz.) Neue Oniscoidea aus Mittel- und Siideuropa und zur Klarung einiger bekannter Formen. Arch, naturgesch. 74 : 163-198.igoSc. Uber Isopoden. Androniscus n. g. (13. Aufsatz.) Zool. Anz. 33 : 129-148. - 1917. Zur Kenntnis der Gattungen Trichoniscus und Mesoniscus . (19. Isopoden-Aufsatz . )Zool. Anz. 49 : 40-57. - 1918. Zur Kenntnis der Ligidien, Porcellioiden und Oniscoideen. (24. Isopoden-Aufsatz.)Arch, naturgesch. 82A (10) : 108-169. - ig26a. 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. Munchen. - i926b. Uber Isopoden der Balkanhalbinsel, gessamelt von Herrn. Dr. I. Buresch. (31.Isopoden-Aufsatz.) Izv. bulg. ent. Druzh. 3 : 135-158. - i927a. Uber einige sudosteuropaische Trichonisciden. (34. Isopoden-Aufsatz.) Zool.Anz. 70 : 200-223. 96 J. P. ELLIS AND R. J. LINCOLN VERHOEFF, K. W. 19275. Illyrionethes n. g. eine cavernicole Trichonisciden-Gattung. (35.Isopoden-Aufsatz.) Zool. Am. 72 : 268-274. ig28a. Isopoda aus Formosa. (39. Isopoden-Aufsatz.) Mitt. zool. Mus. Berl. 14 : 200-226. 19280. t)ber einige Isopoden der zoologischen Staatsammlung in Miinchen. (38.Isopoden-Aufsatz.) Zool. Anz. 76 : 25-36, 113-123. 19280. l)ber alpenlandische und italienische Isopoden. (37. Isopoden-Aufsatz.) Zool. Jb. (Systematik) 56 : 93-172.I929a. Dber Isopoden der Balkanhalbinsel, gesammelt von Herrn. Dr. I. Buresch. (33. Isopoden-Aufsatz.) Izv. tsarsk. pirodonauch. Inst. Sof. 2 : 129-139.I929b. Arthropoden aus siidostalpinen Hohlen. Mitt. Hohlen- u. Karstforsch. Pt i : 14- 35- 19290. Eine neue Diplopoden- und eine neue Isopoden-Gattung aus dem LabyrinthKretas. Mitt. Hohlen- u. Karstforsch. Pt 4 : 113-123. 19303. t)ber einige neue norditalienische Isopoden und einen neuen Typus der Volvation.(43. Isopoden-Aufsatz.) Zool. Anz. 89 : 162-177. i93ob. Zur Kenntnis osteuropaischer Isopoden. (41. Isopoden-Aufsatz.) Zool. Jb.(Systematik), 59 : 1-64. i93ia. t)ber Isopoden terrestria aus Italien. (45. Isopoden-Aufsatz.) Zool. Jb. (System-atik) 60 : 489-572. i93ib. Zur Kenntnis alpenlandischer und mediterraner Isopoda terrestria. (47. Isopoden-Aufsatz.) Zool. Jb. (Systematik), 62 : 15-52. 1932. Cavernicole Oniscoideen. (44. Isopoden-Aufsatz.) Mitt. Hohlen- u. Karstforsch. Pt i : 12-24. 1933- Zur Systematik, Geographic und Okologie der Isopoda terrestria Italiens und iiber einige Balkan-Isopoden. (49. Isopoden-Aufsatz.) Zool. Jb. (Systematik), 65 : 1-64. 1935- Studien iiber Isopoda-terrestria. (51. Isopoden-Aufsatz.) Mitt. zool. Mus. Berlin 21 : 79-163. 1939. Diplopoden, Chilopoden und Oniscoideen, hauptsachlich aus suditalienischen Hohlen. Zool. Jb. (Systematik), 72 : 203-224. 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 some 68 items is given below, and a listof the manuscript names has been deposited in the library of the British Museum(Natural History). Family TYLIDAE Tylos sp. Two specimens. Reg. no. 1905:3:13:5-6. Arauco, Chile. Collected by C. S. Reed. Pur-chased from Rosenberg.Fragment. Reg. no. 1921:10:18:646. Arauco, Chile. Budde-Lund Collection. Family STYLONISCIDAE Styloniscus sp. Two specimens. Reg. no. 1921:10:18:408-409. Corral, Chile. October 1894. Collected byDr Plate. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 97 Styloniscus sp. Two specimens. Reg. no. 1921:10:18:420-421. Auckland, New Zealand. Budde-LundCollection. Family TRICHONISCIDAE Haplophthalmus sp. Two specimens. Reg. no. 1921:10:18:301-302. Lake Como, Italy. Budde-Lund Collection.Haplophthalmus sp. Two specimens. Reg. no. 1921:10:18:303-304. Anzio. 4.4.1897. Collected by F. Silvestri. Budde-Lund Collection. Two specimens. Reg. no. 1921:10:18:305-306. Ain Draham. Collected by F. Silvestri. Budde-Lund Collection.Haplophthalmus sp. Four specimens. Reg. no. 1921:10:18:307-310. Klagenfurt, Austria. Collected by M. Latzel. Budde-Lund Collection.Hyloniscus sp. One specimen. Reg. no. 1928:7:4:76. Krain. Verhoeff Collection.N esiotoniscus sp. Four specimens. Reg. no. 1921:10:18:434-437. Sardinia. Collected by Major Forsyth. Budde-Lund Collection.Schiodtia sp. Two specimens. Reg. no. 1921:10:18:445-446. Liguria, Italy. March 1896. Collected by R. Gestro. Budde-Lund Collection.Trichoniscus sp. One specimen. Reg. no. 1921:10:18:193-196. Kirschhorn Pt, Heikelberg, W. Germany. 21.7.1901. Collected and presented by G. Budde-Lund.Trichoniscus sp. One specimen. Reg. no. 1928:7:4:70. Bulgar. Verhoeff Collection.Trichoniscus sp. Three specimens. Reg. no. 1921:10:18:254-256. Gr. Bossea. August 1888. Collected by A. Vacca. Budde-Lund Collection. Family SQTJAMIFERIDAE Platyarthrus sp. Twelve specimens. Reg. no. 1921:10:18:1537-1548. Tunis. Collected by F. Silvestri. Budde-Lund Collection.Platyarthrus sp. Twelve specimens. Reg. no. 1921:10:18:1525-1536. Rome, Italy. Collected by F. Silvestri. Budde-Lund Collection.Platyarthrus sp. One specimen. Reg. no. 1921:10:18:1587. Tunis. Collected by F. Silvestri. Budde-Lund Collection.Platyarthrus sp. Two specimens. Reg. no. 1921:10:18:1603-1604. Mt Capraro (?). Budde-Lund Collection. One specimen. Reg. no. 1921:10:18:1605. Boccadifalco. 17.2.1896. Collected by F. Silvestri. Budde-Lund Collection. Family ONISGIDAE Alloniscus sp. 'Details' (i.e. parts dissected) from one specimen. Reg. no. 1921:10:18:2112. No locality.Budde-Lund Collection. 98 J. P. ELLIS AND R. J. LINCOLN Aphiloscia sp. Four specimens. Reg. no. 1921:10:18:2069-2072. Madagascar. Budde-Lund Collection.Aphiloscia sp. Five specimens. Reg. no. 1921:10:18:2073-2077. Madagascar. Budde-Lund Collection.Armadilloniscus sp. Five specimens. Reg. no. 1921:10:18:1052-1056. Ban Lem Ngop, Thailand. Collected by Th. Mortensen. Budde-Lund Collection.Armadilloniscus sp. One specimen. Reg. no. 1921:10:18:1059. Brasilia, Brazil. Budde-Lund Collection.Bathytropa sp. Three specimens. Reg. no. 1921:10:18:1515-1517. Botanic Gardens, Hamburg. Budde-Lund Collection.Chaetophiloscia sp. Two specimens. Reg. no. 1921:10:18:2020-2021. Corral, Chile. Collected by R. Paessler. Budde-Lund Collection. , Chaetophiloscia sp. Two specimens. Reg. no. 1930:5:26:42-43. Dalmatia, Yugoslavia. Verhoeff Collection.Deto sp. Seven specimens. Reg. no. 1921:10:18:1078-1084. Rottnest I., W. Australia. Budde-Lund Collection.Detonella sp. One specimen. Reg. no. 1938:7:7:46. Sachalin I., Japan. Verhoeff Collection.Hanoniscus sp. Two specimens. Reg. no. 1921:10:18:5729-5730. W. Australia. Budde-Lund Collection.Hanoniscus sp. Four specimens. Reg. no. 1921:10:18:5731-5734. W. Australia. Budde-Lund Collection.Lepidoniscus sp. One specimen. Reg. no. 1930:5:26:56. Piemonte. Verhoeff Collection.Phalloniscus sp. One specimen. Reg. no. 1921:10:18:2037. Stephens Isle, New Zealand. Collected by Schauinsland. Budde-Lund Collection.Phalloniscus sp. Two specimens. Reg. no. 1921:10:18:2038-2039. Marlborough, New Zealand. Budde-Lund Collection (ex Dundee Museum Collection).Phalloniscus sp. Two specimens. Reg. no. 1921:10:18:2040-2041. New Zealand. Collected by C. Chilton. Budde-Lund Collection.Phalloniscus sp. One specimen. Reg. no. 1921:10:18:2052. Auckland, New Zealand. Collected by H. Siiter (Museum Hamburg). Budde-Lund Collection.Phalloniscus sp. Five specimens. Reg. no. 1921:10:18:2053-2057. Auckland, New Zealand. Budde-Lund Collection (ex Museum Hamburg Collection).Philoscia sp. Eight specimens. Reg. no. 1921:10:18:1730-1737. Botanical Gardens, Hamburg. Budde-Lund Collection.Philoscia sp. One specimen. Reg. no. 1921:10:18:1738. No locality. Budde-Lund Collection.Philoscia sp. Two specimens. Reg. no. 1921:10:18:1743-1744. Baboner. Collected by F. Silvestri. Budde-Lund Collection.Philoscia sp. Three specimens. Reg. no. 1921:10:18:1875-1877. Summit of Mt Roraima. 8600 ft. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 99 Philoscia sp. One specimen. Reg. no. 1888:8. Iguarasso, Brazil. Budde-Lund Collection.Philoscia sp. Three specimens. Reg. no. 1921:10:18:1879-1881. Tonkin, Mauson Mts. 2000-3000 ft. Collected by H. Frunstorfer. Budde-Lund Collection.Philoscia sp. Six specimens. Reg. no. 1921:10:18:1882-1887. Cameroons. Collected by Y. Sjostedt. Budde-Lund Collection.Philoscia sp. One specimen. Reg. no. 1921:10:18:1889. Turkestan. Collected by Uljanin. Budde-Lund Collection.Philoscia sp. Two specimens. Reg. no. 1907:4:30:23-24. Trinidad, West Indies. 18.10.1906. Named by G. Budde-Lund. Presented by the Earl of Crawford.Philoscia sp. Four specimens. Reg. no. 1921:10:18:1890-1893. Japan. Collected by Hilgendorf. Budde-Lund Collection.Philoscia sp. Seven specimens. Reg. no. 1921:10:18:1950-1956. Naranjito Province, Guayas. 9.3.1901. Collected by V. Ortoneda. Budde-Lund Collection.Philoscia sp. One specimen. Reg. no. 1907:4:30:21. Hackgall, Sri Lanka. Named by G. Budde-Lund. Presented by Dr Willey.Philoscia sp. One specimen. Reg. no. 1905:3:31:3. Arauco, Chile. Presented by C. S. Reid. One speci-men. Reg. no. 1921:10:18:1966. Arauco, Chile. Budde-Lund Collection.Philoscia sp. Eight specimens. Reg. no. 1921:10:18:1967-1974. Bileundi. August 1891. Collected by Sjostedt. Budde-Lund Collection.Setaphora sp. Eleven specimens. Reg. no. 1921:10:18:2253-2263. Buitenzorg. Collected by K. Kraepelin. Budde-Lund Collection.Setaphora sp. Twelve specimens. Reg. no. 1921:10:18:2266-2277. Formosa. Collected by Bartels. Budde-Lund Collection.Setaphora sp. One specimen. Reg. no. 1921:10:18:2278. Galnit. Collected by Bartels. Budde-Lund Collection.Setaphora sp. Eighteen specimens. Reg. no. 1921:10:18:2279-2290. Formosa. Budde-Lund Collection.Setaphora sp. Twelve specimens. Reg. no. 1921:10:18:2316-2327. Botanical Gardens, Kew, Surrey. Collected by Bagnall. Budde-Lund Collection.Setaphora sp. Twelve specimens. Reg. no. 1921:10:18:2339-2350. Formosa. Budde-Lund Collection.Setaphora sp. One specimen. Reg. no. 1921:10:18:2338. Balavia. Budde-Lund Collection.Setaphora sp. Five specimens. Reg. no. 1921:10:18:2378-2382. Buitenzorg. Collected by K. Kraepelin. Budde-Lund Collection.Setaphora sp. Seven specimens. Reg. no. 1921:10:18:2384-2390. Botanical Gardens, Kew, Surrey. Collected by Bagnall. Budde-Lund Collection. ioo J. P. ELLIS AND R. J. LINCOLN INCERTAE SEDIS Genus iSpecies a One specimen. Reg. no. 1921:10:18:5691. Paraguay. Budde- Lund Collection.Species b Two specimens. Reg. no. 1921:10:18:2165-2166. La Moka. February 1891. Budde-Lund Collection.Species c One specimen. Reg. no. 1921:10:18:5690. Paraguay. Budde-Lund Collection.Species d Six specimens. Reg. no. 1921:10:18:5692-5697. Paraguay. Budde-Lund Collection.Species e Two specimens. Reg. no. 1921:10:18:5698-5699. Corral, Chile. 5.7.1893. Budde-Lund Collection.Species / One specimen. Reg. no. 1921:10:18:5700. Venezuela. Budde-Lund Collection.Genus 2Species a One specimen. Reg. no. 1931:4:27:71. Georgia. Verhoeff Collection.Genus 3Species a One specimen. Reg. no. 1921:10:18:793. No locality. Budde-Lund Collection.Genus 4Species a Two specimens. Reg. no. 1921:10:18:5753-5754. Ralum. 27.2.1897. Budde-Lund Collec-tion. INDEX abbreviatus (Haplophthalmus) 71adonis (Hyloniscus) 71adriatica (Halophiloscia) 82adriatica rupium (Halophiloscia) 82affinis (Philoscia) 87affinis (Philoscia muscorum) 87alba (Pherusa) 73albicincta (Bilawrencia) 80albus (Titanethes) 73Alloniscus 78, 97alpinus (Androniscus) 70Alpioniscus 69Anchiphiloscia 79Androniscus 70angusta (Niambia) 77angusticauda (Philoscia) 91angusticauda (Setaphora) 91angustissima (Pseudophiloscia) 89anomala (Philoscia) 87anomalus (Phalloniscus) 87Aphiloscia 79, 98apuanus (Haplophthalmus) 71Armadilloniscus 79, 98armata (Deto) 81armata (Paraphiloscia) 85australis (Styloniscus) 68australis (Trichoniscus) 68austriacus (Trichoniscus) 73austroafricanus (Styloniscus) 68austroafricanus (Trichoniscus) 68 balsii (Japanoniscus) 83 balssi (Chaetophiloscia) 81 bargensis (Tiroloscia squamuligera) 93 Bathytropa 79, 98 Benthana 80 Benthanops 88 bicolor (Rhyscotus) 76 biellensis (Philoscia muscorum) 88 Bilawrencia 80 bodkini (Calycuoniscus) 81 bosniensis (Trichoniscus) 73 bougainvillei (Papuaphiloscia) 84 brembana (Tendosphaera) 76 brentanus (Androniscus) 70 breuili (Iberoniscus) 72 brevicornis (Pseudophiloscia) 89 brevis (Alloniscus) 78 briani (Philoscia (Tiroloscia) squamuligera) 93 briani (Tiroloscia squamuligera) 93brunnea (Niambia) 77 Buddelundiella 74BUDDELUNDIELLIDAE 74bulgarica (Bureschia) 70Bureschia 70Burmoniscus So calcivagus (Androniscus) 70 Calmanesia 81 Calycuoniscus 81 capensis (Niambia) 77 capensis (Paranotoniscus) 68 carniolense (Lepidoniscus germanicus) 84 carynthiacus (Androniscus) 70 cateractae (Buddelundiella) 74 caudatus (Platyarthrus) 78 caudatus squamatus (Platyarthrus) 78 cavernarum (Androniscus) 70 cavernarum strasseri (Androniscus) 70 cavernicola (Trichoniscus) 73 cellaria (Chaetophiloscia) 81 cellaria (Philoscia) 81 cestus (Styloniscus) 68 cestus (Trichoniscus) 68 Chaetophiloscia 81, 98 Chavesia 71 cinerascens (Ligia) 74 cingulata (Philoscia) 91 cingulata (Setaphora) 91 circularis (Schoblia) 69 Clavigeroniscus 66 coeca (Philoscia) 91 coeca (Setaphora) 91 commensalis (Trichoniscus) 73 compar (Alloniscus) 78 comta (Philoscia) 91 comta (Setaphora) 91 contractus (Hiatoniscus) 83 Cordioniscus 67 cornutus (Alloniscus) 78 Corsica (Philoscia) 92 Corsica (Tiroloscia) 92, 93 corsicus corsicus (Nesiotoniscus) 72 corsicus (Trichoniscus (Nesiotoniscus)) 72 costata (Bathytropa) 79 costata (Bathytropa meinerti) 79 costulata (Chavesia) 71 costulatus (Platyarthrus) 78 couchi (Halophiloscia) 82, 88 crassicornis (Hyloniscus) 71 cubensis (Rhyscotoides) 76 cubensis (Rhyscotus) 76 cunningtoni (Anchiphiloscia) 79 loob J. P. ELLIS AND R. J. LINCOLN cursorium (Ligidium) 75Cyphonetes 73Cyphoniscellus 71 dahli (Titanethes) 73 dalmatica (Philoscia) 87 dalmatica (Philoscia muscorum) 87 dalmatica (Stenophiloscia) 92 dalmaticus (Hyloniscus) 71 dalmatinus (Armadilloniscus) 79 damae (Hora) 83 danicus (Haplophthalmus) 71 debilis (Ischioscia) 83 debilis (Philoscia) 83 demarcata (Philoscia) 91 demarcata (Philoscia (Setaphora)) 91 denticulatus (Lepidoniscus pruinosus) 84 clentiger (Androniscus) 70 dentiger ligulifer (Androniscus) 70 dentipes (Ligia) 75 depressa (Kogmania) 69 Deto 8 1, 98 Detonella 98 Diacara 82 Didima 82 dilectum (Philoscia) 88 diminuta (Philoscia) 88 dolomiticus (Oroniscus) 84 dorsalis (Chaetophiloscia) 81 elbana (Tiroloscia) 93 elbanus (Parastenoniscus) 76 elbanus (Trichoniscus) 73 elegans (Alloniscus) 82 elegans (Diacara) 82 elongata (Chaetophiloscia) 81, 88 elongata (Philoscia) 81 esterelana (Philoscia (Paraphiloscia)) 93 esterelana (Tiroloscia) 93 Euryligia 74 exigua exigua (Tiroloscia) 93 fagorum (Stylohylea) 72 fagorum (Trichoniscus (Stylohylea)) 72 fasciata (Philoscia (Setaphora?)) 91 fasciata (Setaphora) 91 fiumaranus dolinensis (Haplophthalmus) 71 fiumaranus (Haplophthalmus) 71 flava (Philoscia) 88 flavescens (Niambia) 77 flavus (Oritoniscus) 72, 73 flavus (Trichoniscus) 72 formicarum (Niambia) 77 formosana (Chaetophiloscia) 81Formososcia 82foveolatus (Trichoniscus) 74fragilis (Alpioniscus) 69fragilis (Pseudophiloscia) 89fragilis rharelbazi (Trichoniscus) 73frigidana (Philoscia muscorum) 88fucorum (Halophiloscia) 82fulva (Philoscia (Benthanops)) 88 georgensis (Styloniscus) 68georgensis (Trichoniscus) 68germanicus carniolense (Lepidoniscus) 84glarearum (Stenophiloscia) 92globiceps (Rhyscotus) 76gottscheensis (Cyphoniscellus) 71gracilicornis (Halophiloscia) 82gracilipes (Ligia) 75graecus (Labyrinthasius) 83granulata (Bathytropa) 79, 83granulatus (Tylos) 65granuliferus (Tylos) 65gravosensis (Philoscia) 88griseoflavus (Niambia) 77griseus (Hiatoniscus) 83guernei (Philoscia) 88 Halophiloscia 82 halophilus (Miktoniscus) 72 Haplophthalmus 71, 97 hamuligerus (Androniscus roseus) 70 Hanoniscus 82, 98 hastata (Chaetophiloscia) 81 hercegowinensis (Titanethes (Cyphonetes)) 73 heroldi (Alpioniscus (Illyrionethes)) 70heroldi (Isabelloscia) 83herzegowinense (Ligidium) 75Hiatoniscus 83hirsuta (Halophiloscia) 82hirsuta (Nahia) 84, 88hirsuta (Niambia) 77hirsuta (Philoscia) 84hispana (Bathytropa) 79holthuisi (Papuasoniscus) 85Hora 83 horae (Styloniscus) 68horae (Trichoniscus) 68hottentoti (Styloniscus) 68hottentoti (Trichoniscus) 68humilis (Didima) 82Hyloniscus 71, 97hypnorum (Ligidium) 75 INDEX Iberoniscus 72 Illyrionethes 70 Indoniscus 67 inflatus (Hyloniscus) 71 inflexa (Pseudophiloscia) 89 insulanus (Trichoniscus noricus) 74 intermedius (Scyphax) 91 isabellae (Bilawrencia) 80 Isabelloscia 83 Ischioscia 83 Japanoniscus 83 japonicum (Ligidium (Nippoligidium)) 75 karongae (Anchiphiloscia) 79kempi (Burmoniscus) 80kenepurensis (Oniscus) 84kermadecensis (Styloniscus) 68kermadecensis (Trichoniscus) 68Kogmania 69Komatia 89Krantzia 83 Labyrinthasius 83 lata (Niambia (Manibia)) 77 lateralis (Paraphiloscia) 85 lateralis (Pseudophiloscia) 85 latissima (Euryligia) 74 latum (Ligidium) 75 latus (Paranotoniscus) 68 Lepidoniscus 84, 98 Ligia 74 Ligidium 75 LIGIIDAE 74 ligulifer (Androniscus dentiger) 70 linearis (Rhyscotoides) 76 linearis (Rhyscotus) 76 littoralis (Armadilloniscus) 79 longicauda (Niambia) 77 longicornis (Philoscia) 88 lubricata (Philoscia) 92 lubricata (Setaphora) 92 macchiae (Tiroloscia) 93 macrocephala (Rennelloscia) 89 Manibia 77 marginata (Philoscia (Komatia)) 89 marginepapillosa (Niambia) 77 mariae (Hyloniscus) 71 marina (Deto) 82 marina (Philougria) 82 Marioniscus 84 mauritiensis (Styloniscus) 68 mauritiensis (Trichoniscus) 68 medius (Androniscus subterraneus) 70 medius (Spelaeonethes) 73 meeusei (Chaetophiloscia) 81 meinerti (Bathytropa) 79 meinerti costata (Bathytropa) 79 melanocephala (Ligia) 75 mendanai (Paraphiloscia) 86 methueni (Calmanesia) 81 microps (Niambia (Manibia)) 77 micros (Trichorhina) 78 Miktoniscus 72 mina (Philoscia (Setaphora)) 92 mina (Setaphora) 92 minutissima (Trichorhina) 78 mixtus (Trichoniscoides) 73 mixtus (Trichoniscus (Trichoniscoides)) 73 modesta (Niambia) 77 modestus (Trichoniscoides) 73 modestus (Trichoniscus (Trichoniscoides)) 73 monocellatus (Microniscus) 68 monocellatus (Styloniscus) 68 montana (Plymophiloscia) 89 montanus (Paranotoniscus) 68 montanus (Trichoniscus) 73 montanus (Trichoniscus vividus var.) 73 moruliceps (Styloniscus) 68 moruliceps (Trichoniscus) 68 murrayi (Styloniscus) 69 murrayi (Trichoniscus) 69 muscivagus (Trichoniscus) 74 muscorum affinis (Philoscia) 87 muscorum biellensis (Philoscia) 88 muscorum dalmatica (Philoscia) 87 muscorum frigidana (Philoscia) 88 muscorum (Ligia (Pogonoligia)) 75 muscorum triangulifera (Philoscia) 88 mussaui (Clavigeroniscus) 66 nacreus (Alloniscus) 78 Nahia 84 narentanus (Hyloniscus) 72 nasatus (Rhyscotus) 76 natalensis (Ligia) 75 neozealandicus (Tylos) 65 Nesiotoniscus 72, 97 Niambia 77 Nippoligidium 75 nitida (Philoscia) 88 nitida (Philougria ) 88 nivatus (Trichoniscus) 74 niveus (Tylos) 65 noduliger (Androniscus subterraneus) 70 noricus insulanus (Trichoniscus) 74 noricus sassanus (Trichoniscus) 74 jood J. P. ELLIS AND R. J. LINCOLN noricus sturanus (Trichoniscus) 74novabritannica (Rennelloscia) 89nudulus (Tylos) 65 ocellata (Formososcia) 82Olibrinus 84ONISCIDAE 78, 97Oniscus 84 opercularis (Tylos) 66orientalis (Indoniscus) 67Oritoniscus 72ornatus (Paranotoniscus) 68ornatus (Scyphax) 91Oroniscus 84 ortonedae (Rhyscotoides) 76ortonedae (Rhyscotus) 76otakensis (Styloniscus) 69otakensis (Trichoniscus) 69ovata (Setaphora) 92 pallida (Chaetophiloscia) 81 pallida (Niambia) 77 pallidemaculata (Setaphora) 92 palmetensis (Niambia) 77 papillosa (Niambia) 78 papillosus (Alloniscus) 78 Papuaphiloscia 84 Papuasoniscus 85 parallelus (Rhyscotoides) 76 parallelus (Rhyscotus) 76 Paranotoniscus 68 Paraphiloscia 85 Parastenoniscus 76 patienci (Philoscia) 88 pauper (Benthana) 80 pauper (Philoscia (Benthana)) 80 perkinsi (Geoligia) 75 perkinsi (Ligia) 75 Phalloniscus 87, 98 Philoscia 87, 98 phormianus (Styloniscus) 69 phormianus (Trichoniscus) 69 Phymatoniscus 72 pigmentata (Ligia) 75 pigmentatus (Olibrinus) 84 pilosa (Setaphora) 92 Platyarthrus 78, 97 platycephala (Ligia (Pogonoligia)) 75 pleonalis (Stenoniscus) 76 Plymophiloscia 89 poecilla (Krantzia) 83 Pogonoligia 75 ponticus (Tylos) 66 porcellioides (Alloniscus) 78 porcellioides (Arhina) 78 propinqua (Paraphiloscia) 86 provisorius (Trichoniscus pusillus) 74 pruinosus denticulatus (Lepidoniscus) ! Pseudophiloscia 89 pulchella (Philoscia) 88 pusilla (Niambia) 77 pusillus provisorius (Trichoniscus) 74 pusillus (Trichoniscus) 74 pygmaea (Philoscia) 87 pygmaeus (Phalloniscus) 74, 87 pygmaeus (Trichoniscus) 74 pyrenaeus (Oritoniscus) 72 pyrenaica (Philoscia) 93 pyrenaica (Tiroloscia) 93 rafflesi (Philoscia (Setaphora)) 92 rafnesi (Setaphora) 92 refugiorum (Hyloniscus) 72 Rennelloscia 89 rharelbazi fragilis (Trichoniscus) 73 RHYSCOTIDAE 76 Rhyscotoides 76 Rhyscotus 76 riparius (Hyloniscus) 73 riversdalei (Styloniscus) 69 riversdalei (Trichoniscus) 69 roseus (Androniscus) 70 roseus hamuligeris (Androniscus) 70 robusta (Deto) 82 rupium (Halophiloscia adriatica) 82 sancristobali (Paraphiloscia) 86 santaisabellae (Paraphiloscia) 87 sassanus (Trichoniscus noricus) 74 scaber (Androniscus subterraneus) 70 Schiodtia 97 schobli (Platyarthrus) 78 Schoblia 69 SCHOBLIIDAE 69 scoparum (Trichoniscoides) 73 Scyphax 91 Scyphoniscus 91 seriepunctata (Philoscia) 88 Setaphora 91, 99 siculus (Haplophthalmus) 71 simoni (Oniscus) 84 spatulifrons (Marioniscus) 84 sphaerocephalus (Rhyscotus) 76 spinosus (Cordioniscus) 67 spinosus (Trichoniscus) 67 squamata (Niambia) 77 squamatus (Leptotrichus) 77 squamatus (Platyarthrus caudatus) 78 INDEX SQUAMIFERIDAE 77, 97squamuligera bargensis (Tiroloscia) 93squamuligera briani (Philoscia (Tiroloscia)) 93 squamuligera briani (Tiroloscia) 93squamuligera tendana (Tiroloscia) 93stammeri (Trichoniscus) 74stebbingi (Cordioniscus) 67stebbingi (Trichoniscus) 67STENONISCIDAE 76Stenophiloscia 92 strasseri (Alpioniscus (Illyrionethes)) 70strasseri (Androniscus cavernarum) 70sturanus (Trichoniscus noricus) 74Stylohylea 72STYLONISCIDAE 66, 97Styloniscus 68, 97styricus (Cyphoniscellus) 71suarezia (Philoscia) 92suarezia (Setaphora) 79, 92subterranea (Philoscia) 88subterraneus medius (Androniscus) 70subterraneus noduliger (Androniscus) 70subterraneus scaber (Androniscus) 70swellendami (Styloniscus) 69swellendami (Trichoniscus) 69 tabulae (Styloniscus) 69tabulae (Trichoniscus) 69tendana (Tiroloscia squamuligera) 93Tendosphaera 76TENDOSPHAERIDAE 76thermophila (Bathytropa) 79Tiroloscia 92Titanethes 73TITANIIDAE 69 tomentosa (Trichorhina) 79triangulifera (Philoscia muscorum) 88TRICHONISCIDAE 69, 97Trichoniscoides 73Trichoniscus 73, 97Trichorhina 78truncata (Niambia) 77truncatella (Philoscia) 92truncatella (Setaphora) 92tuberculatus (Hanoniscus) 82tuberculatus (Paranotoniscus) 68tuberculatus (Phymatoniscus) 72tuberculatus (Trichoniscoides) 72TYLIDAE 65, 96Tylos 65, 96tyrrhena (Halophiloscia) 82 ventosus (Styloniscus) 69 ventosus (Trichoniscus) 69 verhoeffi (Trichoniscus) 74 verrucosa (Tendosphaera) 76 verrucosus (Styloniscus) 69 verrucosus (Trichoniscus) 69 vilis (Aphiloscia) 79, 88 vilis (Philoscia) 79 villosa (Benthana) 80 villosa (Philoscia (Benthana)) 80 vittata (Philoscia) 88 vividus (Trichoniscus) 72 vividus var. montanus (Trichoniscus) 73 waitatensis (Scyphoniscus) 91warreni (Philoscia) 88 zosterae (Trichoniscus) 74 JOAN P. ELLIS ROGER J. LINCOLN, Ph.D. 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- 3-75- Printed in Great Britain by. John Wright and Sow Ltd. at The Stonebridge Press, Bristol BS4 5NU THE LARVAL DEVELOPMENT OF CARCINUS MAENAS (L.) AND^^iwC. MEDITERRANEUS CZERNIAVSKY (CRUSTACEA, BRACHYURA, PORTUNIDAE) REARED IN THE LABORATORY A. L. RICE AND R. W. INGLE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 3 LONDON: 1975 THE LARVAL DEVELOPMENT OF GARCIA MAENAS (L.) AND C. MEDITERRANEU& **CZERNIAVSKY (CRUSTACEA, BRACHYIPORTUNIDAE) REARED IN THE LABORATORY BY A. L. RICE - -_ Institute of Oceonographic SciencesAND R. W. INGLE Pp. 101-119 ; i Plate ; 8 Text-figures ; 2 Tables BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 3 LONDON: 1975 -342 .- 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. 28, No. 3, 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.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 29 May, 1975 Price 1.35 THE LARVAL DEVELOPMENT OF CARCINUS MAENAS (L.) AND C. MEDITERRANEUS CZERNIAVSKY (CRUSTACEA, BRACHYURA, PORTUNIDAE) REARED IN THE LABORATORY By A. L. RICE AND R. W. INGLE SYNOPSIS The larval stages of the shore crab Carcinus reared from females collected in British watersare compared with those reared from a female collected in Tunisia and slight, but consistent,differences are noted between them. The Atlantic and Mediterranean populations of Carcinushave recently been separated as distinct species on the basis of adult characters, the nameC. maenas (L.) being applied to the Atlantic form and C. mediterraneus Czerniavsky to theMediterranean form. Although the larval differences support the possibility that the twopopulations are genetically distinct, if, as presently seems to be the case, they are found to betotally allopatric with no chance of interbreeding in areas of overlap, it is suggested that theyshould be accorded only subspecific status, a course which would reflect both their close relation-ship and geographical separation. INTRODUCTION IN THE first half of this century the larval stages of the crabs of north-westernEurope were better known than those of any other region, mainly as a result ofLebour's (1928) classic work on the Plymouth Brachyura. Lebour's descriptions,however, were often inadequate and more recent studies on crabs from other areas,and particularly those based on reared material, have provided more detailedaccounts of the larval development than those available for British species. In 1969 therefore, a research programme was started at the British Museum(Natural History) with the intention of rearing as many British crabs as possible andproviding detailed descriptions of all their developmental stages. As a result of thisprogramme descriptions of the sponge crab Dromia personata (L.), and of the maskedcrab Corystes cassivelaunus (Pennant), have already been published (Rice, Ingle &Allen, 1970 ; Ingle & Rice, 1971) and although crabs of any of the fourteen familiesrepresented in British waters will be reared and described as and when ovigerousfemales become available, efforts are now being concentrated on the swimming crabfamily Portunidae and the spider crab family Majidae and the programme has beenexpanded to include the rearing of Mediterranean species belonging to these families. Apart from the spider crabs (Majidae) the portunids are better represented aroundBritish coasts than any other family of crabs, fourteen species and five genera havingbeen recorded. This paper deals with the commonest of these species, Carcinusmaenas (L.) and with the closely related Mediterranean form, C. mediterraneusCzerniavsky. The common shore crab or green crab, C. maenas, is found in the Atlantic on alltypes of shore and sublittoraly to depths of 200 m, from northern Norway to 6* io 4 A. L. RICE & R. W. INGLE Mauritiana in the east and from Nova Scotia to Brazil in the west (Christiansen,1969). It is therefore not surprising that it was one of the first crabs to have itsdevelopment investigated, the first zoea having been hatched and described byCouch in 1840. Since that time there have been many accounts of the various larval stages ofCarcinus, those by Williamson (1900, 1903) being by far the most detailed andcomplete (see Lebour, 1928, for earlier references). Williamson hatched the firstzoea which moulted to the second stage, but he obtained the later zoeae and themegalopa from the plankton and it was not until 1967 that the species was reared inthe laboratory with any degree of success (Williams, 1967). Williams, however,reared Carcinus mainly for behavioural studies and did not describe the larval stageswhich she obtained, so that there is still no published account of the species' develop-ment based on reared material. In 1971 we succeeded in obtaining all four zoeae, the megalopa and the young crabstages of Carcinus maenas from laboratory-reared material. At that time Carcinuswas being used simply as a test animal to assess the efficiency of the rearing method,the shore crab having been chosen for this purpose because of the relative ease withwhich ovigerous females could be obtained. Compared with many other Britishcrabs the larvae of C. maenas were well known and a new account did not seem to bewarranted, even though the description could now be based on a complete series ofreared stages. However, Williamson's papers were published in a journal which isnot now readily accessible to many workers and when one of us (R. W. I.) reared thelarval stages of C. mediterraneus from a female collected in Tunisian waters, it seemedworth while to publish a comparative account of the development of these twoclosely related species. MATERIAL AND METHODS Larvae were reared from two female Carcinus maenas, one collected at Plymouth,Devon, in April 1971 (B.M. reg. no. 1974:331) and the other at Brighton, Sussex, inMay 1973 (B.M. reg. no. 1974:332), and from one female C. mediterraneus collectedat the northern Punic Port, Salammbo, Tunis in February 1974 (B.M. reg. no. 1974:330). The ovigerous crabs were maintained in sea water treated with o-oi N EDTA andin each case hatching occurred over a 48 h period. All larvae were reared at 15 C(but see below) in sea water treated with EDTA (o-oi N), benzylpenicillin (50 ooounits/litre) and reduced glutathione (o-ooi M), the last additive acting as a feedingstimulant. Every other day the culture water was changed and at the same timethe larvae were fed on freshly hatched Ariemia nauplii. The Plymouth C. maenas larvae were reared in compartmented plastic trays,while the Brighton C. maenas and the C. mediterraneus were reared in 'mass culture'polythene jars using a technique similar to that described by Cook (1969) for rearingpenaeid larvae. In these cases 300-400 larvae were placed into each container (seeFig. i) and the large Artemia moults and dead larvae were removed when the water LARVAL DEVELOPMENT OF CARCINUS 105 FIG. I . Mass-culture vessel made from a 500 ml polythene bottle and used to rear Carcinus.i, lid ; 2, nylon mesh secured by a collar ; 3, screw-on jar cap ; 4, air supply and draintube. level was reduced to the 'drain' line at each water change. A gentle air streaminduced sufficient water movement in the jars to keep the larvae circulating. Survivals of both the tray-reared and mass-cultured C. maenas larvae were veryhigh, up to 68 per cent of the stage i zoeae reaching the first crab stage. During theC. mediterraneus culture period, however, an air-conditioner breakdown resulted inseveral days of oscillating water temperatures reaching 25 C. Survivals were there-fore poor, only two animals reaching the first young crab stage and neither survivingto the next moult. The larvae and moults were preserved in 70 per cent ethanol, and cleared anddissected in lactic acid. Drawings and measurements were made with the aid of acamera lucida. The measurements taken were (a) the distance between the tips ofthe dorsal and rostral spines (T.T.), (b) the rostral spine length (R.S.) from the tipof the spine to the lower margin of the eye, (c) the dorsal spine length (D.S.) and(d) the carapace length (C.L.) from between the eyes to the posterio-lateral carapacemargin. RESULTS Adults Until relatively recently the genus Carcinus was considered to contain only thesingle species, C. maenas, which was recorded from the Atlantic, the Mediterraneanand, probably as an introduced form, from the Indo-West Pacific region. io6 A. L. RICE R. W. INGLE Demeusy & Veillet (1953) pointed out differences between the Atlantic andMediterranean populations and Holthuis & Gottlieb (1958) resurrected the name C.mediterraneus Czerniavsky for the Mediterranean form. Four of the features mentioned by Zariquiey Alvarez (1968) as distinguishing C.mediterraneus adults from those of C. maenas, that is the less sharp anterio-lateralcarapace teeth, the denser setation of the anterio-lateral carapace margins, thesharper carpal tooth on the cheliped and the more pronounced anterio-external angleof the merus of the third maxilliped, are not apparent in the females from which thelarvae reported in this paper were obtained. There are, nevertheless, good mor-phological distinctions between the females and these are listed in Table I andillustrated in Plate i. TABLE i Differences between the female C. maenas and C. mediterraneus from which thelarvae were reared (see also Plate i) C. maenas C. mediterraneus 1. Carapace relatively broad (Brighton spec.C.L. 29-5 mm, C.W. 39-0 mm, C.W./C.L.1-32 ; Plymouth spec. C.L. 40-0 mm, C.W.51-5 mm, C.W./C.L. 1-29) (Plate lA) 2. Carapace dorsal surface relatively roughto touch 3. 5th (posterior) pair of anterio-lateral teethdirected forwards 4. Front does not protrude and is not setose(Plate lA) 5. Carapace regions not strongly elevatedand, when viewed from behind, carapacerelatively flat (Plate iB) 6. Outer margin of cheliped carpus not setose(Plate iC) Carapace relatively narrow (C.L. 27-55 mm,C.W. 34-5 mm, C.W./C.L. 1-25) (Plate iD) Carapace dorsal surface smooth 5th pair of anterio-lateral teeth directed more or less outwards Front protrudes and is setose (Plate iD) Carapace regions elevated, and carapacevaulted (Plate lE) Outer margin of cheliped carpus setose(Plate iF) Larval stages The larvae of the two forms are very similar, particularly in the zoeal stages wherethe only morphological distinctions noted were the relative lengths of the dorsaland rostral carapace spines. With the exception of the dimensions, therefore, thefollowing descriptions of the zoeae apply both to C. maenas and C. mediterraneus. FIRST ZOEADimensions Carcinus maenas: T.T. 1-36-1-44 mm, mean (10 specimens) 1-38 mm ; C.L. 0-47-0-53 mm, mean 0-50 mm ; D.S. 0-53-0-57 mm, mean 0-55 mm ; R.S. 0-46-0-51 mm, mean 0-49 mm ; ratio D.S. /R.S. 1-04-1-22, mean 1-12.Carcinus mediterraneus : T.T. 1-36-1-43 mm, mean (10 specimens) 1-38 mm ; C.L. 0-53-0-57 mm, mean 0-56 mm ; D.S. 0-58-0-63 mm, mean 0-60 mm ; R.S. 0-40-0-43 mm, mean 0-41 mm ; ratio D.S.jR.S. 1-34-1-57, mean 1-47. LARVAL DEVELOPMENT OF CARCINUS 107 FIG. 2. Carcinus zoeal stages I and II : (a) C. maenas stage I ; (b) detail of posterio-lateralangles of abdominal somites ; (c) C. mediterraneus stage I ; (d) C. maenas stage II ;(e) C. mediterraneus stage II. Bar scale represents i-o mm. io8 A. L. RICE & R. W. INGLE Carapace (Fig. za, c) : Well-developed backwardly curved dorsal spine and straight or slightly curved rostral spine, but no laterals. A low anterior papilla between the eyes and a pair of small setae lateral to and slightly behind the dorsal spine. Eyes : Partly fused to carapace. Antennule (Fig. 36) : Unsegmented, with two terminal aesthetascs and two setae. Antenna (Fig. y) : Spinous process about half as long as rostral spine in C. mediter- raneus and somewhat less in C. maenas, with two rows of spinules on the distal two-thirds. Exopod about half as long as spinous process, with one long and one short terminal spine each with minute spinules at their bases. Endopod represented by a small bud. Mandible : Without palp. Maxillule (Fig. 3^) : Endopod two-segmented, with 6 and i setae respectively. Basal endite with 4 setose spines and i seta, coxal endite with a total of 5 spines and setae. Maxilla (Fig. 30) : Endopod, basal endite and coxal endite each bilobed, with 5 + 3, 4 + 4 and 3 + 3 setae respectively. Scaphognathite with 4 marginal setae and a long plumose posterior projection. First maxilliped (Fig. 3/) : Basis with 8 or 9 medial setae. Five-segmented endopod with 2, 2, i, 2 and 4+1 setae respectively. Exopod with 4 natatory setae. Second maxilliped (Fig. 3g) : Basis with 4 medial setae. Three-segmented endopod normally with i, i and 5 setae, though the small lateral seta on the terminal segment may be absent. Exopod with 4 natatory setae. Third maxilliped and pereiopods : Unarmed, unsegmented buds. Abdomen (Fig. 3) : Five somites and telson. Somite 2 with forwardly directed dorso-lateral knobs. Somites 2-5 each with rounded posterio-lateral margins with small teeth, and with a pair of small setae near the posterior margin. Telson with 3 pairs of setose processes on the posterior margin, and each fork with one large and one small dorsal spine and a very slender lateral spine. SECOND ZOEADimensions Carcinus maenas : T.T. 1-58-1-85 mm, mean (8 specimens) 1-75 mm ; C.L. 0-60-0-70 mm, mean 0-67 mm ; D.S. 0-55-0-70 mm, mean 0-67 mm ; R.S. 0-52-0-65 mm, mean 0-58 mm ; ratio D.S. /R.S. 1-06-1-26, mean 1-15.Carcinus mediterraneus : T.T. 1-44-1-56 mm, mean (7 specimens) 1-46 mm ; C.L.0-60-0-70 mm, mean 0-62 mm ; D.S. 0-58-0-63 mm, mean 0-59 mm ; R.S.0-36-0-48 mm, mean 0-43 mm ; ratio D.S./R.S. 1-30-1-69, mean 1-38. Carapace (Fig. 2d, e) : A pair of small setae added between the dorsal spine and the anterior papilla. Posterio-lateral margins with 4-6 setae. Otherwise as in the first stage. Eyes : Now stalked. Antennule (Fig. 3^) : Unsegmented, with 4-6 terminal aesthetascs and i or 2 setae. Antenna (Fig. 3;) : Endopod bud slightly larger than in first stage, otherwise unchanged. LARVAL DEVELOPMENT OF CARCINUS 109 FIG. 3. Carcinus zoeal stages I and II : (a) abdomen, stage I ; (b) antennule, stage I ;(c) antenna, stage I ; (d) maxillule, stage I ; (e) maxilla, stage I ; (/) first maxilliped,stage I ; (g) second maxilliped, stage I ; (h) antennule, stage II ; (_;') antenna, stage II ;(k) maxillule, stage II ; (/) maxilla, stage II. Appendages b, c, e, k and / are drawn fromC. maenas specimens and the remainder from C. mediterraneus. Bar scale represents0-2 mm for a, /and g, and o-i mm for the rest. no A. L. RICE & R. W. INGLE Mandibles : Unchanged. Maxillule (Fig. 3^) : Basal endite with 6 or 7 spines and setae, coxal endite with 5 or 6. Exopod seta now present, endopod unchanged. Maxilla (Fig. 3/) : Scaphognathite now with 9-11 marginal setae, the posterior projection no longer being apparent. Endopod and basal endites unchanged, but coxal endite may now carry an extra seta. First and second maxilliped (Fig. 2d, e) : Exopods with 6 natatory setae ; otherwise unchanged. Third maxilliped and pereiopods : Still unsegmented and unarmed buds. Abdomen : Unchanged except that the two smaller spines on each telson fork are either reduced or absent. THIRD ZOEADimensions Carcinus maenas : T.T. 2-13-2-15 mm, mean (3 specimens) 2-14 mm ; C.L. 0-79-0-85 mm, mean 0-82 mm ; D.S. 0-80-0-85 mm mean 0-82 mm ; R.S. 0-70-0-80 mm, mean 0-73 mm ; ratio D.S./R.S. 1-06-1-21, mean 1-14.Carcinus mediterraneus : T.T. 1-93 mm ; C.L. 0-82 mm ; D.S. 0-77 mm ; R.S. 0-51 mm ; ratio D.S.fR.S. 1-52. Carapace (Fig. 40, b) : Posterio-lateral margins with 8-12 setae, otherwise un-changed.Antennule (Fig. 4^) : Three or four terminal and one sub-terminal aesthetascs, plus 1 or 2 terminal setae. Antenna (Fig. 40) : Exopod about two-thirds length of spinous process ; endopod slightly shorter. Maxillule (Fig. 4/) : Basal and coxal endites with 9 and 6 spines respectively ; i or 2 exopod setae ; endopod unchanged. Maxilla (Fig. 4g) : Endopod, basal endite and coxal endite with 4-5 + 3, 3-5 + 4 and 3-4 + 3 setae respectively. Scaphognathite with 19 or 20 marginal setae. First maxilliped (Fig. 40, b) : Basis and endopod unchanged except that terminal segment may carry 4 or 5 + 1 seta ; exopod with 8 natatory setae. Second maxilliped (Fig. 4, b) : Exopod with 8 natatory setae ; otherwise unchanged. Third maxilliped : Now bilobed, but still unsegmented and unarmed. Pereiopods : Unarmed, unsegmented buds, first pair cheliform. Abdomen (Fig. 40, b) : Somite i with single median dorsal seta, somites 2-5 with well-developed pleopod buds, somite 6 separated from the telson and carrying small uropod buds. Telson forks each usually with one large and one small dorsal spine, though the latter may be absent. FOURTH ZOEADimensions Carcinus maenas : T.T. 2-20-2-50 mm, mean (5 specimens) 2-37 mm ; C.L. 1-02-i-io, mean 1-06 mm ; D.S. 0-81-0-91, mean 0-86 mm ; R.S. 0-74-0-83 mm,mean 0-77 mm ; ratio D.S.IR.S. 1-02-1-20, mean i-n. LARVAL DEVELOPMENT OF CARCINUS in FIG. 4. Carcinus zoea stage III : (a) C. maenas ; (b) C. mediterraneus ; (c) telson, mediter-raneus ; (d) antennule, maenas ; (e] antenna, maenas ; (/) maxillule, mediterraneus ;(g) maxilla, mediterraneus. Bar scales represent 0-5 mm for a and b and 0-25 mm for*-/ 112 A. L. RICE & R. W. INGLE Carcinus mediterraneus : T.T. 2-10-2-35 mm, mean 2-23 mm ; C.L. 0-90-0-99 mm,mean 0-95 mm ; D.S. 0-90-0-95 mm, mean 0-93 mm ; R.S. 0-67-0-72 mm,mean 0-70 mm ; ratio D.S./R.S. 1-32-1-34, mean 1-33. Carapace (Fig. 50, b) : Posterio-lateral margin now with 12-16 setae, otherwise unchanged. Antennule (Fig. 5^) : Terminal seta and a total of 6-7 aesthetascs in three groups. Exopod bud present. Antenna (Fig. 50) : Endopod now equal or almost equal to spinous process. Maxittule : Basal endite may have an additional spine, otherwise unchanged. Maxilla (Fig. 5/) : Scaphognathite with 22-24 marginal setae. Setation of endopod and endites falls within the range in the third stage. First and second maxillipeds : Exopods with 10 natatory setae, otherwise unchanged. Third maxilliped and pereiopods : Large and with the beginnings of segmentation, but still unarmed. Abdomen (Fig. 5#, b and c) : Pleopods now as long as succeeding abdominal somites. Telson forks with one large dorsal spine and usually with a minute second spine. MEGALOPA The megalopa stages of the two species are more easily distinguished than thezoeae. However, the differences noted involve only the pleopods, uropods andtelson and with the exception of these features the following description, like thoseof the zoeal stages, applies both to C. maenas and C. mediterraneus. Dimensions Carcinus maenas : C.L. 1-26-1-40 mm ; C.W. 0-96-1-18 mm. Carcinus mediterraneus : C.L. c 1-21 mm ; C.W. c 1-02 mm. Antennule (Fig. 70) : Dorsal flagellum of 4 segments, the distal 3 segments each with3 or 4 aesthetascs. Terminal segment with 2 setae, penultimate segment with alateral seta and with or without a medial seta. Ventral flagellum unsegmentedwith 4 terminal setae, and i or 2 subterminal ones. Antenna (Fig. jb) : Three-segmented peduncle carrying 3, o and i setae respectively.Flagellum of 7 segments, though the septum between segments 2 and 3 is indistinct.Flagellar segments 3, 5 and 7 each carry 4 setae.Mandible (Fig. je] : Two-segmented palp with about 6 terminal setae.Maxittule (Fig. jc] : Endopod with i or 2 terminal setae ; basal endite with a rowof 5-7 marginal spines and a total of about n setae, coxal endite with 7 or 8 setae.Maxilla (Fig. yd) : Scaphognathite with 37-44 marginal setae and with 5 or 6 setaeon the surfaces of the blade. Endopod unarmed or with a single short seta. Lobesof the basal and coxal endites carrying 7, 6-7, 2 and 3-4 setae respectively.First maxilliped (Fig. 7/) : Exopod two-segmented, with 2 setae on proximalsegment and 3-5 on the distal segment. Unsegmented endopod with 4 or 5 marginalsetae. Basal segment with 14-16 marginal and sub-marginal setae. Coxal enditewith 5 or 6 setae. Well-developed triangular epipod. \ FIG. 5. Carcinus zoea stage IV : (a) C. maenas ; (b) C. mediterraneus ; (c) telson, maenas ;(d) antennule, maenas ; (e) antenna, maenas ; (/) maxilla, mediterraneus. Bar scalesrepresent 0-5 mm for a and b., 0-25 mm for c, d and e, and o-i mm for/. Second maxilliped (Fig. 7g) : Exopod two-segmented, with 4 or 5 terminal setae.Endopod of 5 segments, the proximal unarmed. Epipod bilobed.Third maxilliped (Fig. jh) : Exopod two-segmented, with 4 terminal setae. Endopodof 5 segments, armed with numerous spines and setae and with the ischium expandedand carrying 4 or 5 teeth on the medial margin. Elongated epipod with 2 gill buds. A. L. RICE & R. W. INGLE FIG. 6. Carcinus megalopa : (a) dorsal view, C. maenas ; (b and c) anterior-lateral views ofcarapace in C. maenas (b) and C. mediterraneus (c) ; (d and e) ventral views of telson anduropods in C, maenas (d) and C. mediterraneus (e) ; (g and h) dactyl of fifth pereiopod inC. maenas (g) and C. mediterraneus (h) ; (j) dactyl of second pereiopod, C. maenas ; (/)cheliped, C. maenas. Bar scales represent 0-55 mm for a-c, and 0-25 mm for d-j. LARVAL DEVELOPMENT OF CARCINUS FIG. 7. Carcinus maenas megalopa ; (a) antennule ; (6) antenna ; (c) maxillule ; (d)maxilla ; (e) mandible ; (/) first maxilliped ; (g) second maxilliped ; (h) third maxilliped.Bar scales represent o-i mm for (c) and 0-25 mm for the remainder. Pereiopods (Fig. 6a and f-j) : Chelipeds with prominent ischio-basal hook. Legs2-5 without coxal spines. Dactyl of leg 5 narrow (length/width ratio about 7 : i),with 3 long, sub-terminal sensory setae. Abdomen (Fig. 6a, d and e) : Somites 2-4 with slightly variable pattern of dorsalsetae, but usually with 5 pairs on the posterio-dorsal margin and I pair more an-teriorly. Somite 5 with an extra pair. Telson with a pair of setae on both thedorsal and ventral surfaces, somewhat variable in shape, but with consistentdifferences between the two forms ; in C. maenas the telson narrows posteriorly and n6 A. L. RICE & R. W. INGLEd FIG. 8. Carcinus first crab stage : (a-d) Carcinus maenas, (e-h) C. mediterraneus, carapace(dorsal, lateral and frontal views) and abdomen. Bar scale represents i-o mm. the posterior margin is usually straight or convex, and rarely concave ; in C.mediterraneus it is much more square, the lateral margins being more or less parallelor even diverging slightly, the posterio-lateral angles are more abrupt and theposterior margin is always concave, often markedly so. Pleopods and uropods (Fig. 6d, e) : In both C. maenas and C. mediterraneus the pleo-pods are well developed with 3, rarely 4, coupling hooks on each endopod. But thesetation of the exopods of these appendages and of the uropods was consistentlydifferent in the examples of the two forms examined and afforded the clearest meansof separation. In general C. mediterraneus had fewer setae on the pleopods than didC. maenas, though since there was a good deal of overlap the setation of no singleappendage would separate the two forms (see Table 2). When, however, the total LARVAL DEVELOPMENT OF CARCINUS 117 number of setae on the pleopods of the 5th abdominal somite and on the uropodswas combined, a clear distinction was apparent, C. maenas megalopae always havinga total of 32 setae on these appendages, while C. mediterraneus larvae never had morethan 30. TABLE 2 Setation of the pleopods and uropods in the megalopa larvae of C. maenas andC. mediterraneus, based on counts of five specimens in each case C. maenas C. mediterraneus RaiPleopod, somite 2 somite 3 somite 4 somite 5Uropod FIRST CRAB STAGE (Fig. 8) Carcinus maenas : Carapace length 1-49-1-52 mm ; carapace width (across tipsof 5th anterio-lateral carapace spines) 1-36-1-47 mm. Carcinus mediterraneus : Carapace length 1-49-1-57 mm ; carapace width 1-38-1-50 mm. Only two specimens of the first crab stage of C. mediterraneus were obtained sothat little reliance can be placed on their comparison with the corresponding stagein C. maenas. However, in the material examined the two forms were extremelysimilar, the only differences being a slightly more prominent rostrum and dorsalcarapace tubercles and a squarer telson in C. mediterraneus than in C. maenas. Theother characters distinguishing the adults were not apparent in the first crab stagealthough mediterraneus has a slightly lighter build, the carapace being relativelya little narrower, than maenas. DISCUSSION Williamson's (1903) account of the development of Carcinus maenas includes agreat deal of detail and, in this respect, has rarely been equalled by any subsequentdescription of decapod larvae. His larvae were generally a little larger than thosereared from the Plymouth and Brighton females reported here, but otherwise thetwo accounts agree almost without exception. This agreement confirms, if suchconfirmation was needed, firstly the accuracy of Williamson's observations andsecondly that his specimens of the third and fourth zoeal stage and of the megalopa,all of which he obtained from the plankton, were correctly identified. Lebour's(1928) account is much less complete, but where comparisons are possible her descrip-tions also agree with the reared British larvae. Both of these authors mention theloss in the late zoeae of two of the three telson fork spines which are present in theearly stages, whereas a second spine was almost always present in our stage fourlarvae. However, this second spine is extremely small and difficult to see, so that itwas probably simply missed by Williamson and Lebour. The earlier accounts also agree with the present description of the British Carcinuslarvae in those features which distinguish them from the Mediterranean material, Ii8 A. L. RICE & R. W. INGLE and particularly in the setation of the abdominal appendages of the megalopa.Williamson, for instance, gives the setation of the exopods of the pleopods as n,11-13, 12 or 13, an d ii respectively, while both he and Lebour report 5 setae on theuropods as opposed to the 4 setae usually found on our Mediterranean larvae. Theaccount of these Mediterranean larvae is based on material obtained from only onebrood so that the distinctions noted might simply reflect individual variation.But the agreement between all the available descriptions of British larvae arguesagainst this and suggests that definite genetic differences exist between the Atlanticand Mediterranean Carcinus populations. Whether these differences are sufficient to warrant the two populations beingaccorded full specific status is, however, debatable, for they appear to be allopatric,C. mediterraneus never having been reported from outside the Mediterranean, whileC. maenas is not known from within it. In these circumstances the ultimatecriterion of the absence of interbreeding between overlapping populations cannotbe applied and any opinion about their taxonomic status must be somewhat sub-jective. Comparing the degree of difference between the two allopatric forms with thatbetween undoubtedly distinct species in the same group does not provide veryconclusive evidence. For although portunid crabs are often quite difficult toseparate as juveniles, there are usually more distinct differences between the matureforms than those noted between Carcinus maenas and C. mediterraneus, and in thelarval stages the situation is similar. In terms of the number of species of whichlarvae have been described, the best known portunid genus is Macropipus, andalthough the known larval stages of this genus are all very similar, detailed examina-tion has generally revealed better distinctions between the species than thosebetween the two types of Carcinus larvae described here (Rice & Ingle, 1975). It seems, then, that there is no very good larval evidence to support the separationof maenas and mediterraneus as distinct species, despite the existence of consistentdifferences between the adults. A study of both adults and larvae from the vicinityof the Straits of Gibraltar, both in the Mediterranean and in the Atlantic, wouldbe of considerable interest, though if no area was found in which the two formsco-exist, or intergrade, the problem would still be unresolved. Since the adults areso readily distinguishable, however, it seems worth while at present to maintaintheir specific status, but if they are ultimately shown to be totally allopatric themost sensible course would probably be to consider them to be subspecies, as wasapparently the intention of Czerniavsky (1884) when he divided C. maenas into thetwo varieties mediterranea and septentrionalis (see Holthuis & Gottlieb, 1958). Foras Mayr, Linsley & Usinger (1953) point out, the use of trinominals has the advantagein such situations of conveying the important information that the two forms areclosely related and geographically separated. ACKNOWLEDGEMENTS We thank Dr R. B. Manning who sponsored a visit by one of us (R. W. I.) toTunisia under the Smithsonian Institution Foreign Currency Program, partly for LARVAL DEVELOPMENT OF CARCINUS 119 the purpose of obtaining ovigerous material for comparative larval studies. Wealso thank members of the West Wickham Branch of the British Subaqua Clubfor assistance in collecting ovigerous crabs of C. maenas from Brighton, Sussex. REFERENCES CHRISTIANSEN, M. E. 1969. Marine Invertebrates of Scandinavia, No. 2. Crustacea Decapoda Brachyura. University of Oslo. I43pp.COOK, H. L. 1969. A method of rearing penaeid shrimp larvae for experimental studies. FAQ Fish Rep. 57 (3) : 709-715.COUCH, R. Q. 1840. On the metamorphosis of the decapod crustaceans, nth Rep. R. Cornwall polytech. Soc. pp. 28-43.CZERNIAVSKY, V. 1884. Crustacea Decapoda Pontica littoralia. Materialia and Zoographiam Ponticam comparatam. Trudy Obshch. Ispyt. Prir. imp. Khar'kov 13 : 1-268.DEMEUSY, N. & VEILLET, A. 1953. Sur 1'existence de deux populations de Carcinus maenas Pennant et sur les caracteres morphologique qui les distinguent. C.r. hebd. Seanc. Acad. Sci., Paris, 236 : 1088-1090.HOLTHUIS, L. B. & GOTTLIEB, E. 1958. An annotated list of the decapod Crustacea of the Mediterranean coast of Israel, with an appendix listing the Decapoda of the eastern Mediterranean. Bull. Res. Coun. Israel, 7B : 1-126.INGLE, R. W. & RICE, A. L. 1971. The larval development of the masked crab, Corystes cassivelannus (Pennant) (Brachyura, Corystidae), reared in the laboratory. Crustaceana, 20: 271-284.LEBOUR, M. V. 1928. The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Loud. 1928 : 473-560.MAYR, E., LINSLEY, E. G. & USINGER, R. L. 1953. Methods and Principles of Systematic Zoology. McGraw-Hill. 328pp.RICE, A. L. & INGLE, R. W. 1975. A comparative study of the larval morphology of the British Portunid crabs Macropipuspuber (L.) and M. holsatus (Fabricius) with a discussion of generic and sub-familial larval characters within the Portunidae. Bull. Br. Mus. nat. Hist. (Zool). 28 4: 121-151.RICE, A. L., INGLE, R. W. & ALLEN, E. 1970. The larval development of the sponge crab, Dromia per sonata (L.) (Crustacea, Decapoda, Dromiidea), reared in the laboratory. Vie Milieu, 21 : 223-240.WILLIAMS, B. G. 1967. Laboratory rearing of the larval stages of Carcinus maenas (L.) (Crustacea : Decapoda). /. nat. Hist. 2 : 121-126.WILLIAMSON, H. C. 1900. Contributions to the life history of the edible crab (Cancer pagurus) . Rep. Fishery Bd. Scotl. 18 : (3), 77-143.1903. On the larval and early young stages and rate of growth of the shore crab (Carcinus maenas, Leach). Rep. Fishery Bd. Scotl. 19 : (3), 136-179.ZARIQUIEY ALVAREZ, R. 1968. Crustaceos decapodos ibericos. Investigacion pesq. 32 : i-xi + 5iopp. A. L. RICE, Ph.D. INSTITUTE OF OCEANOGRAPHIC SCIENCES WORM LEY GODALMING SURREY R. W. INGLE, Ph.D. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 560 PLATE I Dorsal and posterior views of the carapace and of the outer face of the right cheliped of thefemale C. maenas from Brighton (A, B and C) and of the female C. mediterraneus from Salammbo(D, E and F). Bull. Br. Mus. nat. Hist. (Zool.) 28, 3 PLATE i B 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- 375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU / 1/1/1 7 A COMPARATIVE STUDY OF THE*LARVAL MORPHOLOGY OF BRITISH PORTUNID CRABSMACROPIPUS PUBER (L.) AND M HOLSATUS (FABRICIUS), WITH A DISCUSSION OF GENERIC AND SUB-FAMILIAL LARVAL CHARACTERS WITHIN THE PORTUNID AE A. L. RICE AND R. W. INGLE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 4 LONDON : 1975 A COMPARATIVE STUDY OF THE LARVA/ f 2* J* 44MORPHOLOGY OF THE BRITISH PORTUISCRABS MACROPIPUS PUBER (L.) ANDM. HOLSATUS (FABRICIUS), WITH ADISCUSSION OF GENERIC AND SUB-FAMILIALLARVAL CHARACTERS WITHIN THEPORTUNIDAE BY A. L. RICE v<^ Institute of Oceanographic Sciences AND R. W. INGLE Pp. 121-151; 9 Text-figures ; 9 Tables BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 4 LONDON: 1975 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. 28, No. 4, 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.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 29 May, 1975 Price i-go A COMPARATIVE STUDY OF THE LARVAL MORPHOLOGY OF THE BRITISH PORTUNID CRABS MACROPIPUS PUBER (L.) AND M. HOLSATUS (FABRICIUS), WITH A DISCUSSION OF GENERIC AND SUB-FAMILIAL LARVAL CHARACTERS WITHIN THE PORTUNID AE By A. L. RICE AND R. W. INGLE SYNOPSIS The zoeal and megalopa stages of two portunid crabs, Macropipus puber and M. holsatus, aredescribed from material reared in the laboratory. Comparison of these larvae with one anotherand with those of M. holsatus, the only other species of the genus described in detail, indicatesthat while the larvae of the Macropipus species are all very similar there are clear distinctionsbetween them. The larval information available for the family as a whole generally supports the currentlyaccepted sub-divisions within the Portunidae based on adult characters, but further detailedlarval descriptions, particularly of those sub-families of which the larval stages are at presentunknown, would probably help in the interpretation of phylogenetic relationships both withinthe family and between the portunids and the remainder of the Brachyura. INTRODUCTION THIS paper is the second in what is hoped will be a series describing the larvaldevelopment of the British portunid crabs. The first paper (Rice and Ingle, 1975)described larvae reared from British specimens of the shore crab, Carcinus maenas,and compared them with those of the closely related Mediterranean form. Thissecond paper deals with two of the eight species of the genus Macropipus whichhave been recorded from British waters. Although Lebour's (1928, 1944) classic work on the Plymouth Brachyura estab-lished distinctions between the larvae of several genera and species of British crabs,she was not able to separate satisfactorily the larval and young stages of the Macro-pipus species with which she dealt (under the name Portunus] except on the basis ofchromatophore pattern. Since that time the larvae of only one species of Macropipushave been reared and described in detail (Goldstein, 1971), but with a further twospecies described here it is now possible to discuss the nature of the specific larvalcharacters within the genus. MATERIALS AND METHODS The ovigerous female Macropipus puber was obtained from Lulworth Cove,Dorset, in September 1972 and the M. holsatus from Port Erin, Isle of Man, in 124 A - L - RICE & R. W. INGLE February 1973. In both cases the crabs were maintained in sea-water at a tempera-ture of 15 C and all the larvae were reared at this temperature. The eggs of M. puber hatched on 25 September 1972 and those of M. holsatus on27 February 1973 and both broods were reared partly in mass culture and partlyindividually in compartmented plastic boxes (see Rice and Ingle, 1975). All of thelarvae were fed on a mixture of Prorocentrum and freshly hatched Artemia nauplii. The survival rate through the larval stage was quite good in both species, but wassomewhat better in holsatus than in puber ; of 90 individually reared M. holsatus 25(about 22 per cent) reached at least the first crab stage, whereas of 180 individuallyreared M. puber only 17 (about 9 per cent) reached this stage. Although there was considerable individual variation in the number of days spentin each larval stage the total length of the larval life in both species was very similar,M. holsatus moulting into the first crab stage 43-53 (mean 48) days after hatchingwhile M. puber reached this moult in 46-56 (mean 50) days. The reared larvae and the females from which they were obtained are depositedin the British Museum (Natural History) under registration numbers 1974 : 333(M. holsatus) and 1974 : 334 (M. puber}. MORPHOLOGY OF THE LARVAL STAGES AND SPECIFIC DISTINCTIONSWITHIN THE GENUS MACROPIPUS The larvae of Macropipus puber and M. holsatus are illustrated in Figs. 2-9. Theillustrations are supplemented in Tables 1-6 where these two species are comparedwith Goldstein's (1971) description of the larvae of M. marmoreus. As was to be expected from previous work, the larvae of all three species are verysimilar, but although no single character separates them at all stages there are cleardistinctions between them. Thus the zoeae of puber differ from the other species at all stages in having straightrather than curved dorsal and rostral spines, and in the early stages by being con-siderably larger (Fig. lA-C). This size difference becomes less marked in the laterstages but at this time, that is in the fourth and fifth stages, an obvious distinctionis the loss in puber of one of the spines on the telson forks. The larvae of puber arealso consistently relatively narrower than those of holsatus and, at least in stage I,of marmoreus too (Fig. iD). The absence of the relevant width data in Goldstein(1971) precludes the inclusion of marmoreus in this comparison beyond the firststage, specimens of which are deposited in the British Museum (Natural History).* As in the adults (see Christiansen, 1969), the larvae of holsatus and marmoreus aremore difficult to separate on morphological grounds, both of them having curvedcarapace spines and being similar in size. However, there are a number of fairlyeasily seen differences. First, the posterio-lateral processes on the abdominalsomites are much better developed in marmoreus than in either holsatus or puber,particularly in the late zoeal stages. Secondly, with the possible exception of stageIII, marmoreus resembles puber and differs from holsatus in having the spinous process * Stage V larvae of M. marmoreus subsequently received from Dr Goldstein are intermediate betweenpuber and holsatus in this respect. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 125 of the antenna much shorter relative to the rostral spine. Finally, the lateral knobson the third abdominal somite are lost in stage II in marmoreus, that is one stageearlier than in either holsatus or puber. In addition to these characters there are less obvious differences, particularlybetween the zoeae of marmoreus on the one hand and puber and holsatus on the other,in the number of aesthetascs on the antennules and the setation of the antennae,maxillae and maxillipeds (see Tables 1-5). The megalopae of the three species are also very similar. Although the compara-tive size situation is now the reverse of that in the early zoeal stages, the megalopaeof puber tending to be smaller than those of either holsatus or marmoreus, the sizeranges overlap a good deal. Similarly, while the megalopae of holsatus seem to havea rather lighter build than those of the other two species, this distinction is not veryclear and in any case is obscured by the distortion resulting from preservation.There are, however, good distinctions between the species based on the antennae andtelson. Thus, while puber has an antennal flagellum of seven segments and atelson with three pairs of dorsal setae, holsatus has eight segments in the flagellumand only two pairs of dorsal telson setae, and marmoreus has six segments in theantennal flagellum and four pairs of dorsal telson setae. Finally, the slender dactylof the fifth pereiopod in puber, with its sub-terminal sensory setae, is clearly dis-tinguishable from the broader dactyl and terminal setae of holsatus and marmoreus. Although the features dealt with here clearly enable the larvae of Macropipuspuber, M. holsatus and M. marmoreus to be differentiated, it is not possible toanticipate which of these characters, if any, will be of use in distinguishing betweenthe other species of the genus. However, the fact that such consistent differencesexist, particularly between the larval stages of holsatus and marmoreus which areso similar as adults, indicates that when the larvae of these other species are describedin sufficient detail similarly clear distinctions will be found between them. GENERIC AND SUB-FAMILIAL LARVAL DISTINCTIONS WITHINTHE FAMILY PORTUNIDAE Previous attempts to define generic and sub-familial distinctions amongst portunidlarvae have been clouded by the unfortunate nomenclatural confusion surroundingthis family of crabs (see Opin. Decl. Int. Commn zool. Nom. 1956, vol. 12, Opinion394, pp. 317-336). This confusion was primarily due to the use of the namePortunus for two distinct genera, one mainly confined to European waters, and theother widely distributed in the tropics and sub-tropics. The International Com-mission's decision was to retain the name Portunus Weber, 1795, for the widelydistributed genus, which had also been called Neptunus in the Mediterranean andIndo-West-Pacific, and to adopt the name Macropipus Prestandrea, 1833, for theEuropean genus. These changes have had repercussions in the nomenclature of thesub-familial divisions within the Portunidae, but the situation was clarified byStephenson and Campbell (1960) whose interpretation and nomenclature, based onadult characters, are followed here except that the sub-family name PolybiinaeOrtmann, 1893, is used in place of their Macropipinae. 126 A. L. RICE & R. W. INGLE 0-5 1-8 r MM 0-6 3-0 MM 2-0 3-0 2-6 2-2 ZOEAL STAGE III IV FIG. i. A comparison of the dimensions of the zoeal stages of Macropipus puber (triangles),M. holsatus (circles) and M. marmoreus (squares). (A) Carapace length, (B) distancefrom the tip of the dorsal spine to the tip of the rostrum, (C) distance between the tipsof the lateral carapace spines, and (D) ratio B/C. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 127 FIG. 2. First zoea of M. puber (a, b and c) and of M. holsatus (d, e and/).The bar scale represents i-o mm. Stephenson and Campbell recognized six sub-families (Carcininae, Polybiinae(= Macropipinae), Portuninae, Catoptrinae, Caphyrinae and Podophthalminae), butlarvae of only the first three of these are known. Apart from Macropipus, larvae ofonly two polybiinid genera have been described in detail ; Ovalipes ocellatus (Herbst)was reared through the zoeal stages and the megalopa by Costlow and Bookhout(1966), the megalopa of 0. punctatus was described by Muraoka (1969), while Roberts(1969) reared Bathynectes superba (Costa) to the terminal zoea. (Continued on page 142) 128 A. L. RICE & R. W. INGLE FIG. 3. Appendages of the zoeae of M. puber ; (a) antennule, stage I, (b) antennule,stage II, (c-g) antenna, stages I-V, (h) detail of spinous process, stage I, (_;') endopodand basal endite of maxillule, stage I, (k) endopod and scaphognathite of maxilla,stage II, (I and m) first and second maxillipeds, stage I, (n and o) first and secondmaxillipeds, stage V. The bar scale represents o-i mm for h, j and k, and 0-5 mm forthe remainder. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 129 FIG. 4. Second zoea of M. puber (a, b and c) and of M. holsatus (d, e and/).The bar scale represents i-o mm. 130 A. L. RICE & R. W. INGLE FIG. 5. Third zoea of M. puber (a, b and c) and of M. holsatus (d, e and/).The bar scale represents i-omm. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES FIG. 6. Fourth zoea of M. puber (a, b and c) and of M. holsatus (d, e and/).The bar scale represents i-o mm. 132 A. L. RICE & R. W. INGLE FIG. 7. Fifth zoea of M. puber (a, b and c) and of M. holsatus (d, e and/).The bar scale represents i-o mm LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 133 FIG. 8. Megalopa of M. puber (a-d) and of M. holsatus (e-g). The bar scalesrepresent 0-5 mm for (d) and (g) and i-o mm for the remainder. 134 A. L. RICE & R. W. INGLE FIG. 9. Appendages of the megalopa stage in M. puber and M. holsatus : (a) antenna,M. puber ; (b) antenna, M . holsatus ; (c) antennule, M. puber ; (d, e and /) first, secondand third maxillipeds, M. puber ; (g) cheliped, M. holsatus ; (h, j) second and fourthpereiopods, M. puber ; (k) fifth pereiopod, M. puber ; (/ and m) fifth pereiopod, M.holsatus. The bar scale represents 0-5 mm for a-f and m, and i-o mm for g-l. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 135 TABLE i A comparison of the first zoeal stage in Macropipus puber, M. holsatus and M . marmoreus* M. puber Carapace length : 0-7 mm Tip of dorsal to tipof rostral spines (a) : 1-9-2-2 mmWidth including lateral spines (b) 0-67-0-78 mmRatio a/b : 2-68-2-80Length of dorsal spine : 0-9 mm Carapace : Dorsal spine stout,straight or slightly curved at tip,much longer than carapace.Rostral spine straight or slightlycurved anteriorly, longer thancarapace. Lateral spines welldeveloped. Carapace dorsal surfaceminutely rugose. Eyes sessile(Fig. 2a, b) M. holsatusSIZE 0-4-0-6 mm 1-1-1-3 mm0-61-0-74 mmi -800-4-0-5 mm M. marmoreus* 0-4 mm i-i mm (1-18-1-24 rnmf) - (0-58-0-67 mmf) - (1-84-2-0 mmf)0-5 mm Antennule : Unsegmented, with 2-3 terminal aesthetascs and 1-3 setae (Fig. 3) Antenna : Spinous process less than length of rostral spine. Exopod almost % length of spinous process, with 2 unequal terminal setae (Fig. y) Mandible : without palp Maxillule : Endopod with 5 or 6 setae on distal segment and i on proximal segment. Basal endite with 5 setae, coxal endite with 6 setae (Fig. 3 Maxilla : Endopod bilobed with 6-8 setae, basal endite with 6-8 setae, coxal endite with 7 setae. Scaphognathite with 5 setae, posterior one largest (Fig. 3^) First maxilliped : Basipodite with 10 As puber setae (2, 2, 3, 3), 5-segmented endopod with 2, 2, i, 2 and 5 setae, exopod with 4 natatory setae (Fig. 3/) CEPHALOTHORAX Dorsal spine slender,strongly curvedposteriorly and equal to oronly slightly longer thancarapace. Rostral spinestraight or curvedbackwards, equal to orshorter than carapace.Lateral spines welldeveloped. Carapacesmooth. Eyes sessile(Fig. -zd, e)As puber Similar to holsatus As puber Spinous process aboutas long as rostral spine.Otherwise as puber As puberAs puber As puber Exopod with 3 terminalsetae. Otherwise aspuber As puberAs puber Scaphognathite with 6setae ; otherwise as puber Basipodite with only 6setae ; otherwise as puber 136 A. L. RICE & R. W. INGLE M. puber M. holsatus M. marmoreus* Second maxilliped : Basipodite with As puber As puber 4 setae, 3-segmented endopod with i, i and 4 or 5 setae, exopod with 4 natatory setae (Fig. 3m) Pereiopods : Limb buds just visible Not visible Not visible beneath carapace ABDOMEN (Fig. 2c, /) Five somites plus telson. Somites As puber As puber 2 and 3 with lateral processes, those of somite 3 minute Somites 2 -5 each with pair of As puber As puber, but posterio- minute dorsal setae and with lateral expansions on posterio-lateral margins with somite 3 more prominent rounded expansions bearing small and acute spinules Telson with 2 lateral and i dorsal As puber As puber spines on each fork ; posterior margin with 3 pairs of setae * The data on Macropipus marmoreus in Tables 1-6 are taken from Goldstein, 1971.j- Figures from material deposited in the British Museum (Natural History). TABLE 2 A comparison of the second zoeal stages in Macropipus puber, M. holsatus and M. marmoreusM. puber M. holsatus M. marmoreus SIZECarapace length : 0-85-0-90 mm 0-60-0-70 mm 0-60 mm Tip of dorsal to tipof rostral spines (a) : 2-2-2-5 mm 1-40-1-60 mm 1-40 mm Width including lateral spines (b) : 0-77-0-84 mm 0-71-0-76 Ratio a/b : 2-85-3-01 1-87-2-15 Length of dorsal spine : ci-omm 0-50-0-60 mm 0-50 mm CEPHALOTHORAX Carapace : As in stage I but eyes Similar to stage I but As holsatus now stalked (Fig. 40, b) dorsal spine less than carapace length and eyes stalked (Fig. <\d, e) Antennule : 4-6 aesthetascs and As puber As puber 1-3 setae Antenna : Endopod present as a As puber, but spinous As puber, but exopod with bud. Spinous process < length of process almost as long 3 terminal setae rostral spine. Exopod with 2 as rostral spine terminal setae (Fig. 3^) LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 137 M. puber Mandible : As in stage IMaxillule : Endopod as in stage I.Endites with additional setae.Exopod seta presentMaxilla : Endopod as in stage I.Endites with additional setae.Scaphognathite with 1 1 setae,posterior one no longersignificantly differentFirst maxilliped : Exopod with 6natatory setae ; otherwise as instage I Second maxilliped : Endopodterminal segment with 4 setae.Exopod with 6 natatory setaePereiopods : Limb buds prominent M. holsatus As puberAs puber As puber As puberAs puberLimb buds just visible ABDOMEN (Fig. 40, f)Lateral processes on somite 3 present As puber Posterio-lateral processes of abdominal somites acute but not prominent One lateral spine on telson fork very small or absent Posterior telson margin with 4 pairs of setae As puber All 3 spines on telsonforks well developedAs puber M. marmoreus As puberAs puber As puber Basipodite with 8 setae ;otherwise as puber Endopod terminal segmentwith 5 setae. Otherwiseas puberAs holsatus Absent Posterio-lateral processes of somites 2-4 more prominent As holsatus As puber TABLE 3 A comparison of the third zoeal stages in Macropipus puber, M. holsatus and M. marmoreus M. puber Carapace length : 0-96- i'O mmTip of dorsal to tipof rostral spine (a) : 3-0-3-3 mmWidth including lateral spines (6) i -00-1-04 mmRatio a/b : 2-95-3-17 Length of dorsal spine :1-40 mm Carapace : As in stage II(Fig. 5 a, b) M. holsatusSIZE M. marmoreus 0-80-0-85 mm '7 mm 1-90-2-20 mm 1-70 mm 0-90-0-95 mm 2-10-2-34 c 0-85 mm 0-70 mm CEPHALOTHORAX As in stage II but As holsatus dorsal spine now equalto or slightly greaterthan carapace length(Fig. $d, e) 138 A. L. RICE & R. W. INGLE M. puber Antennule : As in stage IIAntenna : Endopod bud almost Jlength of exopod. Spinousprocess c \ length of rostral spine.Exopod with 2 terminalaesthetascs (Fig. 30)Maxillule : Additional setae onendites. Otherwise as in stage IIMaxilla : Scaphognathite with13-14 setae First maxilliped : Exopod with 8natatory setae. Otherwise as inprevious stages Second maxilliped : Exopod with 8natatory setae. Endopodterminal segment with 4 setae M. holsatus SIZE As in stage IIAs puber but spinousprocess > length ofrostral spine As puberAs puberAs puber Endopod with 3terminal setae.Otherwise as puber M. marmoreus As in stage IIEndopod bud almostlength of exopod.Exopod with 4 terminalsetae. Otherwise asholsatusAs puber As puberAs puber Endopod with 5terminal setae.Otherwise as puber Sixth somite separated from telson Lateral processes of somite 3 now absent Posterio-lateral processes of somites 3-5 less than J length of succeeding somites Telson forks usually with only 2 spines ; if third spine present it is minute Posterior telson margin with 4 pairs of setae ABDOMEN (Fig. y,f) As puberAs puber More prominent, thoseon somite 3 aboutlength of somite 4Telson forks each with 3 spines 4 or 5 pairs of setae As puberAs puber Similar to holsatus As holsatus 5 pairs of setae TABLE 4 A comparison of the fourth zoeal stages in Macropipus puber, M. holsatus and M. marmoreus M. puber Carapace length : i -04- 1 -24 mmTip of dorsal to tipof rostral spine (a) : 3-40-3-60 mmWidth including lateral spines (b) c i -20 mmRatio a/b : 2-86-2-96Length of dorsal spine : 1-30-1-40 mm M. holsatusSIZE M. marmoreus 1-10-1-20 mm 2-60-2-90 mm1-20-1-25 mm2-10-2-291-10-1-20 mm 0-9 mm 2-0 mm 0-90 mm LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 139 M. puber Carapace : As in stage III(Fig. 6a, b) Antennule : 3 or 4 aesthetascs andi or 2 setae. Ventral flagellummay be present as small budAntenna : Spinous process aboutlength of rostral spine. Exopod flength of spinous process, with 2terminal setae. Endopod Jexopod (Fig. 3/) Maxillule : Additional setae on endites. Otherwise as in previous stage Maxilla : Scaphognathite with about 25 marginal setae First maxilliped : Endopod segments with 2, 2, i, 2 and 6 setae. Exopod with 10 natatory seta Second maxilliped : 10 natatory setae. Otherwise as in stage III Pereiopods : First leg now clearlychelate M. holsatus CE PH ALOTH O RAX As in stage III(Fig. 6d, e) As puber (Fig. o) Spinous process >length of rostral spine.Exopod < J spinousprocess, with 2 terminalsetae. Endopod fexopodAs puber As puberAs puber 10 natatory setae.Endopod with 4 terminalsetae, i.e. as in puberAs puber Posterio-lateral processes of somites3-5 longer than in stage III, thoseof somite 3 about J length of somite 4 Telson forks each with 2 spines Posterior margin with 10 or n setae Pleopods and uropods present asbuds ABDOMEN (Fig. 6c, f) More prominent than inpuber, those of somite3 about somite 4 3 spines Posterior margin with 9 or 10 setae As puber M. marmoreus As in stage III 6 aesthetascs in two groups, plus a terminal seta Spinous process -f rostral spine. Exopod < J spinous process, with 3 terminal setae. Endopod < J exopod As puber As puberAs puber 10 natatory setae.Endopod with 5 terminalsetaeAs puber More prominent than inholsatus, those of somite3 > somite 4 3 spines Posterior margin with 10 setae As puber TABLE 5 A comparison of the fifth zoeal stages in Macropipus puber, M. holsatus and M. marmoreus M. puber Carapace length : i -50- 1 -60 mmTip of dorsal to tipof rostral spine (a) : 3-8-4-1 mmWidth including lateral spines (b) 1-60-1-70 mmRatio a/b : 2-4-2-5 M. holsatusSIZE M. marmoreus 1-30-1-48 mm 3-00-3-50 mm1-50-1-60 mm 1-50 mm 3-70 mm 140 M. puber Length of dorsal spine :i -60- 1 -70 mm A. L. RICE & R. W. INGLE M. holsatusSIZE 1-3 mm M. marmoreus Carapace : As in previous stages (Fig. 7 a, b) Antennule : 8-10 aesthetascs in 2 or 3 groups, plus a terminal seta. Ventral flagellum well developed (Fig. 3b) Antenna : Spinous process about length of rostral spine. Exopod about length of spinous process ; 2 terminal setae. Endopod > exopod (Fig. $g) Mandible : Palp now present Maxillule : Similar to stage IV Maxilla : Schaphognathite with 30-36 marginal setae First maxilliped : As in stage IV but with 12 natatory setae (Fig. 3>z) Second maxilliped : Endopod terminal segment with 5 setae. Exopod with 12 natatory setae (Fig. 30) Third maxilliped : Well developed, biramous but unarmed Pereiopods : Limbs clearly segmented CEPHALOTHORAX As in previous stages(Fig. rf, e)As puber Spinous process > flength of rostral spine.Otherwise as puber Posterio-lateral processes ofsomites 3 and 4 about J length ofsucceeding somitesTelson forks with 2 spinesPosterior margin with 10 or, moreusually, 12 setaePleopods as long as succeedingsomites As puberAs puberAs puber As puber Endopod terminalsegment with 4 or 5setae. Exopod with12 natatory setaeAs puber As puber ABDOMEN (Fig. 7 c, f)As puber 3 spines10 setae As puber 1-40 mm As in previous stages 13 aesthetascs plusterminal seta. Ventralflagellum well developed Spinous process < f rostralspine. Exopod < spinousprocess ; 4 terminal setae.Endopod > exopod As puber As puber Scaphognathite with about 40 setae As puber As puber As puberAs puber ? Posterio-lateral process ofsomite 3 > J length ofsomite 43 spines10 setae As puber TABLE 6 A comparison of the megalopa stage in Macropipus puber, M. holsatus and M. marmoreusM. puber M. holsatus M. marmoreus SIZE Carapace lenght (a) : 1-66-2-09 mm 1-86-2-16 mm c 2-0 mm Carapace width (b) : 1-57-1-76 mm 1-65 mm Ratio a/6 : 1-12-1-26 c 1-40 ci-22 LARVAL DEVELOPMENT OF MACRIPOPUS SPECIES 141 M. puber Carapace : Relatively broad (seelength/width ratio above).Rostrum directed ventrally and notprominent in dorsal view. Dorsalsurface with two medial tuberclesand two pairs of lateral ones.These tubercles, and particularlythe posterior medial one, varyconsiderably in prominence. Frontabout maximum carapace width.Posterior margin almost straight andmeeting lateral margins in abruptangles (Fig. 8a, b)Antennule : Three-segmentedpeduncle. Dorsal flagellum of 4segments with 2 terminal setae and 3 groups of aesthetascs (Fig. gc)Antenna : Peduncle of 3 segmentsand flagellum of 7 segments, thefifth bearing 2 long setae (Fig. go)Mandible : Well-developed 2-segmented palp with 8-10 setaeMaxillule : Endopod of 2 segmentscarrying 2 and 4 setae respectively.Endites with many marginal spinesand setae Maxilla : Endopod unarmed or witha single seta. Schaphognathite withabout 50 marginal setaeFirst maxilliped : Exopod with 3-5setae on the proximal segment and 4 or 5 on the distal one. Endopodunsegmented and reduced. Well-developed epipod of fairly constantwidth throughout its length (Fig. gd)Second maxilliped : Exopod2-segmented, proximal with i smallseta, distal with 4 or 5 terminalsetae. Endopod 4-segmented,proximal naked. Epipod small(Fig. ge) Third maxilliped. Exopod with 6terminal setae. Endopod of 5segments, all armed. Epipod long(Fig. 9/) Pereiopods. Cheliped with prominentcurved spine on ischio-basis.Pereiopods 2-4 with straight spineson coxae (Fig. gg, h, j) M. holsatusCEPHALOTHORAX Relatively narrow.Rostrum generallydirected moreanteriorly than in puberand therefore moreprominent in dorsalview. Posterior marginarched and not meetingthe lateral margins inabrupt angles.Otherwise as puber(Fig. 8e,f) As puber Flagellum of 8segments (Fig. gb) As puberAs puber As puberAs puber As puber As puber Pereiopods 1-4 as in puber M. marmoreus Relatively broad.Rostrum prominent indorsal view. Posterio-lateral tubercles absent.Front about maximumcarapace width As puber Flagellum of 6 segments,the long setae being onthe thirdAs puber As puber As puber Epipod expanded basally.Otherwise as puber As puber As puber Pereiopods 1-4 as in puber I 4 2 M. puber A. L. RICE & R. W. INGLE M. holsatus D actyl of pereiopod 5 slender, lengthmore than 5 times maximum width.Sensory setae clearly sub-terminal.Coxal spine minute or absent(Fig. gk) CEPHALOTHORAX Dactyl of pereiopod 5broad, length about 4times maximum width.Sensory setae virtuallyterminal (Fig. gl, m) M. marmoreus As holsatus Somites 2-5 with posterio-lateralexpansions roundedExopods of pleopods on somites2-5 with 17-19, 16-19, 17-18 and13-15 setae respectively Endopods of pleopods with 3, orrarely 4, coupling hooksExopods of uropods with 8-10 setaeTelson usually narrows posteriorly,but may be almost square.Posterior margin usually straightor slightly concave, but may bemarkedly so Telson dorsal surface with 2 pairsof posterio-medial setae arrangedin a rectangle elongatedlongitudinally. Additional pair ofanterio-lateral setae (Fig. 3d) ABDOMEN (Fig. 8c)As puber Exopods of pleopodswith 17-20, 19-20,18-19 and 14-16 setaerespectivelyAs puber 9 or 10 setaeTelson normally square,but may narrowposteriorly. Shapetherefore not consistentlydistinguishable from puberTelson dorsal surfacewith 2 pairs of posterio-medial setae arranged ina rectangle elongatedtransversely. Noanterio-lateral setae(Fig. 8g) As holsatus As puber 8 setae Telson dorsal surfacewith 4 pairs of setae Costlow and Bookhout mistakenly considered the species dealt with by Lebour(1928) under the name Portunus to belong to the sub-family Portuninae, so that withthe exception of an inadequate account of the first zoeal stage of Bathynectes longipesRisso (Lebour, 1931) their description of the development of Ovalipes ocellatusseemed to be the first of a true polybiinid. Their discussion of possible sub-familiallarval characters within the Portunidae was therefore largely invalid, but theynevertheless recognized the possession of sternal cornua in the megalopa stage as aportuninid character (see below). Roberts was similarly confused by the genericnomenclature when he compared his reared Bathynectes larvae with those of otherspecies in the family and consequently did not recognize some larval differenceswhich seem to be generic or even sub-familial. On the other hand, Goldstein (1971) did not mix polybiinid and non-polybiinidspecies in her comparison with the larvae of Macropipus marmoreus, and was ableto point out several differences between M. marmoreus, Ovalipes ocellatus andBathynectes superba. The additional information presented here on the larvae ofM. puber and M. holsatus now warrants a more detailed comparison in order to 143 establish the sort of generic distinctions which are likely to obtain between larvalPolybiinae. The three genera Macropipus, Ovalipes and Bathynectes are thereforecompared in Tables 7 and 8, revealing differences which at the moment appear to begeneric. TABLE 7A comparison of the zoeal stages in Macropipus, Bathynectes Macropipus* Bathynectes^ ABDOMEN Dorso-lateral processes,somite 3 Dorso-lateral processes,somite 4 Dorso-lateral processes,somite 5Posterio-lateralprocesses, somite 4 TELSON Telson fork armature, stage I Telson fork armature, stages II -V Anterior lateral telson spine Posterior margin, stage II Posterior margin, stage III < somite 5 3 spines (i dorsal)2-3 (i dorsal) < J fork length 4 + 4 setae 5 + 5 setae FIRST MAXILLIPED Basipodite medial setae 6-10 Endopod terminal setae, 5 stage III Natatory setae, stages 6, 8, 10, 12 II-V > somite 5 fromstage II 3 spines (i dorsal)3 (i dorsal)> J fork length 3 + 3 4 + 4 10 6 6, 8, 10, 12 and Ovalipes Ovalipes^ All stages Present stages I -III Present all stages < somite 5 2 (both lateral) 2 (both lateral) < J fork length 3 + 3 4 + 4 4-?6 7, 8, 10, 14 SECOND MAXILLIPED Natatory setae,stages II-V 6, 8, 10, 12 6, 8, 10, 12 7, 10, 12, 15 * Based on Macropipus puber and M. holsatus described in this paper, and on M . marmoreus describedby Goldstein, 1971. f Based on Bathynectes superba described by Roberts, 1969. J Based on Ovalipes ocellatus described by Costlow and Bookhout, 1966. 144 A. L. RICE & R. W. INGLE TABLE 8 Lateral knobs on carapace Rostral spine Coxal spines on legs 2-4 Maxillule endopod, basal segment Maxilla endopod Maxilliped i, endopod Dactyl, pereiopod 5 Abdominal somite 5, posterio-lateral margins Pleopods, endopods Pleopods, exopods Uropods, proximal segment Uropods, distal segment A comparison of the megalopa stage in Macropipus and Ovalipes Ovalipes punctatus]Absent Macropipus spp. Ovalipes ocellatus* Absent Present Directed downwardsj Directed forwardsPresent Absent 2 setae 3 setae Unarmed or witha single setaUnsegmented,expanded at tipwith 3-6 setaeNot markedlyflattened ;L: W> 4 : iRounded 3-4 hooks 13-20 marginal setae o-i seta * Data from Costlow and Bookhout, 1966. f Data from Muraoka, 1969. $ Except depurator according to Labour (1928). 7 setae Segmented, notexpanded, about17 setae Markedly flattenedL:W< 3 - 5 :i 8-10 marginal setae ? Directed forwards Absent 2 setae 6 setae Unsegmented, withabout 17 setae As O. ocellatus Acute 7 hooks 31-40 marginal setae 3 setae About 22 marginalsetae In the zoeal phase Ovalipes can be distinguished from the other two genera bythe possession of dorso-lateral processes on the fifth abdominal somite in all stagesand on the fourth somite up to stage III, and in lacking a dorsal spine on the telsonforks. Macropipus differs from both Ovalipes and Bathynectes in losing the dorso-lateral process on the third abdominal somite after the second stage, while in all thezoeal stages Bathynectes has much longer posterio-lateral processes on the fourthsomite and also a much more prominent anterior spine on the telson fork than eitherMacropipus or Ovalipes. Unfortunately Roberts was unable to obtain the megalopa of Bathynectes so thatin this stage Macropipus can be compared only with Ovalipes, and even here muchof the information which would be useful in such a comparison is not available (seeTable 8). Nevertheless, the megalopae of Ovalipes seem to differ from all describedMacropipus megalopae in having more setose appendages, particularly the abdominalones, and in lacking the prominent spines on the coxal segments of legs 2, 3 and 4. Possible sub-familial characters of the Polybiinae can be discussed only in relationto the Carcininae and the Portuninae, for which at least some larval information hasbeen obtained. Within the Carcininae, although Couch hatched and described the LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 145 first zoea of the common and widely distributed shore crab, Carcinus maenas (L.),as early as 1844 and there were many subsequent accounts of some or all of thelarval stages of this species, they have only recently been described in detail fromreared material and compared with the closely related C. mediterraneus Czerniavsky(see Rice and Ingle, 1975). The only other carcininid genus of which larvae areknown is Portumnus, Lebour (i944a) having hatched and reared the first two stagesof P. latipes and attributed to the same species a plankton-caught terminal zoeawhich moulted to the megalopa and then to the first crab stage in the laboratory.Another plankton-caught megalopa which moulted to a first crab had been ascribedto Xavia biguttata (Risso) (as Portumnus biguttatus] in an earlier paper (Lebour,1928) ; these two megalopae were very distinct. There are many morphological details, particularly in the setation of the append-ages, in which the larvae of Carcinus and Portumnus differ from those of the poly-biinid species discussed above. Some of these features may be good sub-familialcharacters, but in addition there are a number of much more obvious differenceswhich appear to enable larval carcininids to be distinguished readily from poly-biinids. Firstly, the two Carcinus species, and probably also Portumnus, passthrough only four zoeal stages, whereas all the known polybiinids have five zoeae.The consequent differences in developmental rates result in distinct combinationsof characters in the two sub-families ; pleopod buds, for instance, occur on zoeaewith 8 natatory setae on the exopods of the maxillipeds in the Carcininae, that is inthe third stage, whereas the pleopods do not appear in the Polybiinae until thefourth zoeae in which the maxillipeds carry 10 setae. This character would not, ofcourse, enable stage I and II carcininid and polybiinid larvae to be separated,but the zoeae of Carcinus and Portumnus differ from those of Macropipus in theseearly stages, and from those of Bathynectes and Ovalipes at all stages, in having nodorso-lateral processes on the third abdominal somite. Carcininid zoeae also differfrom the polybiinids, certainly from the third stage, in having only three pairs ofsetae on the posterior margin of the telson, whereas all the known larval Macropipus,Bathynectes and Ovalipes add at least one extra pair of setae and often more. Butthe most obvious carcininid zoeal character, which distinguishes them from all otherknown portunids, both polybiinids and portuninids (see below), is the total absenceof lateral carapace spines. In the megalopa stage Carcinus and Portumnus differ from Macropipus in havingno coxal spines on pereiopods 2-4, and from Ovalipes in having rounded posterio-lateral margins to the fifth abdominal somite. Both polybiinid genera differ fromCarcinus in having more setae on the pleopods and uropods, the difference beingmore marked in Ovalipes than in Macropipus. In terms of the number of species of which larvae have been described, the Por-tuninae is the best known sub-family of swimming crabs. However, although thereare published accounts of one or more stages of five portuninid genera, many of theseaccounts are inadequate, several are based on material collected from the planktonand therefore of doubtful identity and, of the reared species, the complete develop-ment is known for only four (see Table 9) . Nevertheless, the available data do allowsome general comments about portuninid larvae to be made. 146 A. L. RICE R. W. INGLE TABLE 9 Descriptions of larval Portuninae referred to in the accompanying discussion* SPECIESCallinectes sapidus Scylla serrataScylla serrataCharybdis acuta Charybdis japonica Charybdis luciferaCharybdis annulataCharybdis callianassaCharybdis orientalisCharybdis 6-dentataCharybdis bimaculataPortunus trituberculatusPortunus pelagicusPortunus pelagicus Portunus pelagicusPortunus pelagicus Portunus sayiPortunus depressifronsThalamita crenata Thalamita simaPortunus sanguinoleutusPortunus pelagicusCharybdis orientalisThalamita crenata Zoeae I -IV, megalopa Zoeae I, II and III ? Zoea I ?Zoea I ?Zoea I MegalopaZoea IZoea IZoea IZoea I Hatched and reared Hatched and plankton Plankton Plankton Hatched PlanktonPlanktonPlanktonPlanktonPlankton AUTHOR Costlow and Bookhout,1959 Ong, Kah Sin, 1964Naidu, 1955Kurata and Omi, 1969 Yatsuzuka, 1952 Hashmi, 1970Hashmi, 1970Hashmi, 1970Hashmi, 1970Aikawa, 1937Aikawa, 1937Aikawa, 1937Aikawa, 1937Delsman and De Man, 1925Yatsuzuka, 1962 Prasad and Tampi, 1953 Lebour, 19440Lebour, i944bPrasad and Tampi, 1953 Muraoka, 1969Chhapgar, 1956Chhapgar, 1956Chhapgar, 1956Chhapgar, 1956 * The list is by no means complete, a number of publications having been omitted because the identi-fications are particularly unreliable, the descriptions are inadequate or they have been superseded bysubsequent work. A more serious omission, however, is the excellent account of the larval develop-ment of Portunus spinicarpus by Bookhout & Costlow (1974), which was not seen until after thismanuscript had been submitted. Fortunately, P. spinicarpus is a typical portuninid and its omissiondoes not affect the discussion of sub-familial larval characters in this paper. Firstly, there is considerably more variation in the rate of development in theportuninids than in either the polybiinids or carcininids, the number of zoeal stagesranging from seven in Callinectes sapidus through six in Charybdis acuta and C.japonicus, five in Scylla serrata, to four in Portunus pelagicus (or even three, accordingto Prasad and Tampi, 1953, although the sizes given for their zoeae indicate thatthey may have missed at least one stage). This heterogeneity extends also to somemorphological features, such as the telson fork armature, for while there is a tendency toa reduction of the number of spines on the telson forks in the later stages of portuninids,just as in the other two sub-families, there is considerable variation even in the first LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 147 zoeal stage. Thus while some species (e.g. Scylla serrata, Portunus depressifrons andP. sayi) retain all three spines on each fork, one of the lateral spines is reduced to afine hair in Charybdis callianassa, C. lucifer and C. annulata, and totally disappearsin Portunus trituberculatus, P. pelagicus, P. sanguinolentus, Callinectes sapidus,Charybdis japonicus, C. 6-dentata, C. orientalis and Thalamita crenata. Finally,Chhapgar (1956) illustrates only a single spine on the telson forks of his larvae ofCharybdis orientalis, Thalamita crenata, Portunus sanguinolentus and P. pelagicus. This variability allows the zoeae of individual portuninid species to be distinguishedfairly easily from all known polybiinids, but there does not seem to be a singlefeature which will separate the two sub-families at all zoeal stages. For thosecharacters which are common to all portuninid zoeae, such as the possession of well-developed dorsal, rostral and lateral carapace spines, dorso-lateral processes onabdominal somites 2 and 3, posterio-lateral processes on somites 3-5 in the laterstages, telson forks usually with at least two spines of which one is dorsal, and theaddition of at least one extra pair of setae on the posterior margin of the telson fromstage II, are almost without exception shared also with some or all of the polybiinids.In fact, the only feature we have noticed which seems to be unique to portuninidzoeae is the absence of any setae on the middle segment of the endopod of the firstmaxilliped in the first zoeal stage.* [Hashmi (1970) figures a seta in this position inthe first zoea of Charybdis callianassa, but we have been unable to confirm itsexistence in material deposited by him in the collections of the British Museum(Natural History).] Even this fine distinction disappears in the later stages ofScylla serrata and Charybdis acuta, and as early as the second zoea in Callinectessapidus, and would hardly seem sufficient to support the sub-familial separation ofthe Portuninae and Polybiinae based on adult characters. But in the megalopastage there are much clearer distinctions between the Portuninae, on the one hand,and all other described swimming crabs, on the other. All portuninid megalopae,for instance, have prominent posterio-lateral spines on the fifth abdominal somitewhich are found only in Ovalipes punctatus amongst the Carcininae and Polybiinae.Similarly, there is a tendency in the Portuninae to increase the number of setae onthe uropods beyond that found in the other two families, again with the exception ofOvalipes, but since this armature ranges in Portunus pelagicus from 10 or n (accord-ing to Aikawa, 1937, and Yatsuzuka, 1962) to 20 (according to Prasad and Tampi,1953) the reliability of this character is very doubtful. By far the most strikingportuninid megalopal character, however, is the possession of a pair of prominentposterio-ventrally directed spines, the so-called sternal cornua, arising from thesternum between the last pair of legs. This feature at once distinguishes the por-tuninids, not only from the carcininids and polybiinids, including Ovalipes, but alsofrom all other brachyuran megalopae so far described. These differences between the larvae of the Carcininae, the Polybiinae and thePortuninae are summarized in the following diagnoses. * One other possibly unique portuninid zoeal character is the position of the pair of fine setae on thedorsal surface of the carapace. In the Carcininae and Polybiinae these setae are anterior to the dorsalspine and close together, whereas in the Portuninae they are between the dorsal and lateral carapacespines. However, they may not always be present, for Costlow and Bookhout, 1959, specificallylooked for them in Callinectes and failed to detect them. 148 A. L. RICE & R. W. INGLE Carcininae ZOEAE Four zoeal stages ; carapace without lateral spines ; dorso-lateral projections onabdominal somite 2 only ; posterio-lateral processes of abdominal somites 3 and 4less than half length of succeeding somites in all stages ; telson fork armaturereduced to a single spine in late stages ; telson posterior margin with 3 + 3 setae inall stages ; middle segment of endopod of first maxilliped armed with a single setain stage I. MEGALOPA Rostrum directed forwards or downwards ; no coxal spines on pereiopods ; nosternal cornua ; pleopods with 9-12 (rarely 13) marginal setae ; uropods with 4-10marginal setae ; posterio-lateral spines on abdominal somite 5 absent. PolybiinaeZOEAE Five zoeal stages ; well-developed lateral carapace spines ; dorso-lateral projec-tions on abdominal somites 2 and 3, at least in the early stages ; posterio-lateralprocesses of abdominal somites 3 and 4 usually less than half length of succeedingsomites in later stages ; telson forks with at least 2 spines in all stages ; telsonposterior margin with at least 4 + 4 setae in late stages ; middle segment of endopodof first maxilliped armed in stage I. MEGALOPA Rostrum directed forwards or downwards ; coxal spines on pereiopods 2-4present (Macropipus) or absent (Ovalipes) ; no sternal cornua ; pleopods with 14-20(rarely 13) marginal setae (Macropipus) or 31-40 (Ovalipes) ; uropods with 8-10marginal setae (Macropipus) or c 22 (Ovalipes) ; posterio-lateral spines on abdominalsomite 5 absent (Macropipus) or present (Ovalipes) . PortuninaeZOEAE Four to seven zoeal stages ; well-developed lateral carapace spines ; dorso-lateralprojections on abdominal somites 2 and 3 in all stages ; posterio-lateral processes ofabdominal somites 3 and 4 more than half length of succeeding segments in latestages ; telson forks with at least 2 spines in all stages (except according to Chhap-gar) ; telson posterior margin with at least 5 + 5 setae in late stages ; middle segmentof endopod of first maxilliped unarmed in stage I. MEGALOPA Rostrum directed forwards ; coxal spines absent from pereiopods 3 and 4 but maybe present on pereiopod 2 (Charybdis and Scylla) ; sternal cornua always present ; LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 149 pleopods with 17-26 marginal setae ; uropods with 11-14 marginal setae ; posterio-lateral spines of abdominal somite 5 always prominent and usually over-reachingsomite 6. THE BEARING OF LARVAL CHARACTERS ON PORTUNID CLASSIFICATION The above distinctions between the larval stages of the Carcininae, Polybiinae andPortuninae generally support the current divisions within the family which arebased largely on the degree to which the adults show adaptations to the swimminghabit. These adaptations include a tendency to lighten the integument, to flattenthe carapace and extend it laterally to improve the dynamics of sideways swimming,to change the orientation of the leg articulations and to flatten the limbs and fringethem with setae to produce effective paddles (Hartnoll, 1971). There is a good dealof variation in the extent of these adaptations within the sub-families, but in generalthe Carcininae and Portuninae represent the extreme conditions, the Carcininaeshowing the least and the Portuninae the most modification from the unspecializedbrachyuran form. The Polybiinae are somewhat intermediate between the othertwo sub-families and even within the genus Macropipus there is a considerablerange in swimming adaptations, and particularly in the degree of flattening of thelegs (see Palmer, 1927). This situation is reflected in the larvae, for while the Carcininae and Portuninaeexhibit major differences in both the zoea and megalopa stages, the Polybiinae havezoeae which resemble the Portuninae while the megalopae are more similar to thoseof the Carcininae. This does not, of course, necessarily mean that the Polybiinaestand phylogenetically between the Carcininae and the Portuninae, and the presencein the zoeae of Ovalipes of characters such as the dorso-lateral processes on abdominalsomites 4 and 5, which are apparently unique amongst the portunids, indicates thatthis genus, at least, is well away from any such route. It has been generally accepted that the swimming adaptations of adult portunidsare secondary acquisitions and that the relative absence of such adaptations in theCarcininae is therefore a primitive condition. There have, however, been suggestionsthat the morphological series into which extant swimming crabs can be arrangedmight be read in the opposite direction, so that the loss of swimming adaptationsbecomes an advanced feature (Palmer, 1927; Lebour, 1928). The informationavailable from the larval stages does not provide any clear evidence for one or otherof these views. For the main character distinguishing carcininid larvae from allother portunids, that is the absence of lateral carapace spines in the zoeal stages,occurs sporadically in a number of other brachyuran families and does not seem tobe of any particular phylogenetic importance. On the other hand, the presence ofsternal horns on all portuninid megalopae, a feature which is unique not only amongstthe Portunidae but also amongst the Brachyura generally, indicates that this sub-family probably represents the end of a portunid evolutionary line rather than anintermediate stage. It is possible, of course, that the Polybiinae are phylogenetically more 'primitive'than either the Carcininae or the Portuninae, in the sense that they are closer to the 150 A. L. RICE & R. W. INGLE ancestral stock or stocks of both sub-families. This would require two parallelevolutionary tendencies within the Portunidae, one involving a loss of swimmingadaptations and a return to the relatively unspecialized brachyuran condition of theCarcininae, and the other leading to the increased specialization of the Portuninae.Morphological details of the larvae of more swimming crab species, and particularlythose of the sub-families of which the larval stages are totally unknown at present,might help to clarify these relationships within the Portunidae and between theswimming crabs and other brachyuran families. But the pelagic larvae are generallymuch more similar than the adults since they show none of the specializations forsuch habits as swimming, burrowing or commensalism which characterize thebenthic adult phase. A study of larval systematics will therefore probably requirea numerical approach, making use of a much greater variety of features than hasusually been used in the past. Such an investigation is underway at the momentusing the available published information for all brachyuran larvae, and it is hopedthat this will not only supplement the systematics based on adult characters butwill also help in the identification of unknown plankton-caught larvae. ACKNOWLEDGEMENTS We wish to thank Trevor Davies and members of the Guildford Branch of theBritish Subaqua Club for collecting the female M. puber and Dr D. I. Williamson,Port Erin Marine Biological Station, for obtaining for us the material of M. holsatus. REFERENCES AIKAWA, H. 1937. Further notes on brachyuran larva. Rec. Oceanogr. Wks Japan, 9 : 87-162.CHHAPGAR, B. F. 1956. On the breeding habits of larval stages of some crabs of Bombay. Rec. Indian Mus. 54 : 33-52.BOOKHOUT, C. G. & COSTLOW, J. D. 1974. Larval development of Portunus spinicarpus reared in the laboratory. Bull. mar. Sci. 24 : 20-51.CHRISTIANSEN, M. E. 1969. Marine Invertebrates of Scandinavia, No. 2. Crustacea Decapoda Brachyura. University of Oslo, I43pp.COSTLOW, J. D. & BOOKHOUT, C. G. 1959. The larval development of Callinectes sapidus Rathbun reared in the laboratory. Biol. Bull. mar. biol. Lab. Woods Hole, 116 : 373-396.1966. The larval development of Ovalipes ocellatus (Herbst) under laboratory condi-tions. /. Elisha Mitchell scient. Soc. 82 : 160-171.COUCH, R. Q. 1844. On the metamorphosis of the decapod Crustacea. Rep. R. Cornwall poly tech. Soc. 11 : 28.DELSMAN, H. C. & DeMAN, J. G. 1925. On the 'Radjungans' of the Bay of Batavia. Treubia, 6 : 308-323.GOLDSTEIN, B. 1971. Developpement larvaire de Macropipus marmoreus (Leach) en labora- toire (Crustacea, Decapoda, Portunidae). Bull. Mus. natn. Hist. nat. Paris, 42 : 919-943.HARTNOLL, R. G. 1971. The occurrence, methods and significance of swimming in the Brachyura. Anim. Behav. 19 : 34-50.HASHMI, S. S. 1970. The brachyuran larvae of W. Pakistan hatched in the laboratory. Part II. Portunidae : Charybdis (Decapoda : Crustacea). Pakist. J. scient. Res. 12 : 272- 278.KURATA, H. & OMI, H. 1969. The larval stages of a swimming crab, Charybdis acuta. Bull. Tokai reg. Fish. Res. Lab. 57 : 129-136. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 151 LEBOUR, M. V. 1928. The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Lond- 1928 : 473-560. 1931- Further notes on larval Brachyura. Proc. zool. Soc. Lond. 1931 : 93-96. i944a. The larval stages of Portumnus (Crustacea, Brachyura) with notes on some other genera. /. mar. biol. Ass. U.K. 26 : 7-15. I944b. Larval crabs from Bermuda. Zoologica, N.Y. 29 : 113-128. MURAOKA, K. 1969. On the post-larval stage of two species of the swimming crab. Bull. Kanagawa Pref. Mus. 1 : 1-7.NAIDU, K. G. RAJA BAI. 1955. The early development of Scylla serrata (Forsk.) De Haan and Neptunus sanguinolentus (Herbst). Indian J. Fish. 2 : 67-76.ONG, KAH SIN. 1964. The early development stages of Scylla serrata Forskal (Crustacea, Portunidae), reared in the laboratory. Proc. Indo-Pacif. Fish. Court. 11 : 135-146.PALMER, R. 1927. A revision of the genus Portunus (A. Milne-Edwards, Bell, etc.). /. mar. biol. Ass. U.K. 14 : 877-908.PRASAD, R. R. & TAMPI, P. R. S. 1953. A contribution to the biology of the blue swimming crab, Neptunus pelagicus (Linnaeus), with a note on the zoeae of Thalamita crenata Latreille. /. Bombay nat. Hist. Soc. 51 : 674-689.RICE, A. L. & INGLE, R. W. 1975. The larval development of Carcinus maenas (L.) and C. mediterraneus Czerniavsky, (Crustacea, Brachyura, Portunidae) reared in the laboratory. Bull. BY. Mus. nat. Hist. (Zool.) 28 (4) : 101-119.ROBERTS, M. H. 1969. Larval development of Bathynectes superba (Costa) reared in the laboratory. Biol. Bull. mar. biol. Lab. Woods Hole, 137 : 338-351.STEPHENSON, W. & CAMPBELL, B. 1960. The Australian portunids (Crustacea : Portunidae). IV. Remaining genera. Aust. J. mar. Freshwat. Res. 11 : 73-122.THOMPSON, J . V. 1835. On the double metamorphosis of the decapodous Crustacea exemplified in Carcinus maenas. Phil. Trans. R. Soc. 1835 : 359-362.YATSUZUKA, Ko. 1952. The metamorphosis and growth of the larva of Charybdis japonica A. Milne Edward. Bull. Jap. Soc. Scient. Fish. 17 : 353-358.1962. Studies on the artificial rearing of the larval Brachyura especially of the larval blue-crab, Neptunus pelagicus Linnaeus. Rep. Usa mar. biol. Stat. Kochi Univ. 9 : 1-88. A. L. RICE Ph.D. INSTITUTE OF OCEANOGRAPHIC SCIENCES WORMLEY GODALMING SURREY R. W. INGLE Ph.D. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 560 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 jNU MISCELLANEA BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 5 LONDON: 1975 MISCELLANEA Pp 153-247 ; 9 Plates ; 45 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 28 No. 5 LONDON: 1975 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. These papers form Vol. 28, No. 5 of the Zoologyseries. The abbreviated titles of periodicals citedfollow those of the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.j ISSN 0007 1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 17 September, 1975 Price 6.20 CONTENTS Page Streptaxidae from Aldabra Island, Western Indian Ocean. By A. C. VANBRUGGEN 157 Quickia aldabraensis, a new species of land snail from Aldabra Atoll, Western Indian Ocean. By C. M. PATTERSON ...... 177 Notes on some echinoderms from Marion Island. By F. W. E. ROWE &A. M. CLARK ........... 187 A new species of Tilapia in the Zambian Zaire system. By E. TREWAVAS & D. J. STEWART. .......... igi Two new nematodes parasitic in the kiwi in New Zealand. By E. A. HARRIS 199 Description of Pembatoxon insular e gen. n., sp. n. from Pemba Island. By J. VAN GOETHEM .......... 207 A quagga, Equus quagga, at University College, London and a note on a supposed quagga in the City Museum, Bristol. By A. W. GENTRY . 217 A new angelfish of the genus Centropyge from Ascension Island. By R. LUBBOCK & R. D. SANKEY ........ 227 A new species of Nanochromis from the Ogowe System, Gabon. By E.TREWAVAS 233 The first zoeal stages of Cancer pagurus L., Pinnotheres pisum (Pennant) and M acrophthalmus depressus. By A. L. RICE ..... 237 STREPTAXIDAE (MOLLUSCA, GASTROPODA: PULMONATA) FROM ALDABRA ISLAND, WESTERN INDIAN OCEAN By A. C. VAN BRUGGEN INTRODUCTION THE pulmonate gastropod family Streptaxidae has an almost circumtropicaldistribution. Species are particularly numerous and diverse on the African con-tinent ; in addition the family is well represented on Madagascar, the Comoros,Seychelles and Mascarene Islands in the Western Indian Ocean. Aldabra atoll(924' S 462o' E) is situated in the southwestern Indian Ocean (fig. 5) ; the nearestland of any magnitude is the island of Madagascar at a distance of about 420 kmto the southeast and the African mainland at about 640 km to the west. Apartfrom very small atolls the nearest high islands are the Comoros at a distance ofabout 400 km to the southwest. The Seychelles are further away than both theAfrican continent and Madagascar, viz. about 1200 km to the northeast. Maxwell Smith (1909) was the first to record the presence of streptaxids on theisland of Aldabra. Messrs J. F. Peake and J. D. Taylor have been participating invarious phases of the Royal Society Expedition to Aldabra. In the course of theirwork on the atoll and its satellite island Assumption they have collected extensiveseries of streptaxid shells, the study of which they have entrusted to the presentauthor. Aldabra and Assumption appear to harbour four species of Streptaxidae,three of which are extinct. All are described below followed by a discussion on theirrelationships and possible derivation. The following abbreviations have been used : BMNH British Museum (Natural History), London ; NM Natal Museum, Pietermaritzburg ; PSTE/JCFF Percy Sladen Trust Expedition, leg. J. C. F. Fryer ; RMNH Rijksmuseum van Natuurlijke Historic, Leiden ; RSE Royal Society Expedition ; lid ratio length/major diameter of shells. The l/d has been calculated from micrometer readings, so that these figures maynot always agree with those calculated from the accompanying measurements in mm. Acknowledgements are due to Mr J. F. Peake and Dr J. D. Taylor, and the staffof the Mollusca Section of the British Museum (Natural History) for assistance invarious respects. I am also indebted to The Royal Society, who have fostered andencouraged research on Aldabra Island. Thanks are due to Dr A. Zilch of theSenckenberg-Museum, Frankfurt am Main, for hospitality for comparative studiesat his institute, and to Mr R. N. Kilburn of the Natal Museum for lending the Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 158 A. C. VAN BRUGGEN material described by Connolly in 1925. The illustrations are due to the profes-sional skill of H. Heijn, staff artist of the Department of Systematic Zoology ofLeiden University. Localities have been pinpointed by means of their coordinates on the Royal Societygrid as shown on the map by Stoddart in Westoll & Stoddart (1971) (between pp. 632and 633). The fossils have been registered in the Mollusca Section, Department of Palaeont-ology, and recent specimens bear registration numbers of the Mollusca Section,Department of Zoology, both in the British Museum (Natural History). Gulella gwendolinae (Preston, 1910) Ennea gwendolinae Preston, 1910, Ann. Mag. nat. Hist. (8) 6 : 527, pi. 7, fig. 3 ('Shimbi Hills, British East Africa').Gulella gwendolinae : Verdcourt, 1962, Annls Mus. r. Afr. centr. Sir. 8 Sci. zool. 106 : 8 ('K [= Kenya], Shimba Hills and nearby coastal forests'). Gulella gwendolinae aldabrae n. subsp.Figs, i, 6 Gulella gwendolinae : Connolly, 1925, /. Conch., Lond. 17 : 265 ('Aldabra I.') ; Germain, 1934, 67me Congr. Soc. sav. : 131 (footnote, 'Aldabra').Ennea gwendolinae : Barnacle, 1962, /. Seych. Soc. 2 : 54 ('Aldabra'). Ennea alauda Sykes (nomen nudum) : Barnacle, 1962, /. Seych. Soc. 2 : 54 ('He Picard, Aldabra').Ennea sp. : Smith, 1909, Nautilus 23 : 69 ('Grande Terre. He Picard. lies Vertes'). DIAGNOSIS. A subspecies of Gulella gwendolinae in size and shape in betweenthe typical form and G. g. tsadiensis Blume, but with the aperture smaller than inthe other subspecies ; dentition as in typical form, but with little variation. DESCRIPTION OF SHELL. Shell (fig. i) small, (sub) cylindrical, with open umbilicus,smooth, creamy white. Spire produced, sides subparallel, apex flattened, obtuselyconical. Whorls six to seven, slightly convex, almost completely smooth, onlywith faint traces of costulation or striation below the sutures and around the um-bilicus, initial whorls rarely with indication of spiral sculpture ; sutures shallow,simple, somewhat impressed, (sub)crenellate. Aperture quadrate, rounded at base,peristome incrassate and reflected, white and glossy, with six-fold dentition : areasonably well-developed oblique angular lamella touching the tip of the labrum ;two subequal mid-labral denticles on a slightly raised common base, labral complexcorresponding to shallow but extensive external pit ; a mid-basal denticle, alsocorresponding to a little and shallow outside depression, which is sometimes hardlynoticeable ; bipartite columellar process consisting of two prominent subequaldenticles on a raised common base - the bifurcation of the columellar process mayvary in depth. MEASUREMENTS OF SHELL: 3-6-5-1 x 1-4-1-9 mm, lid 2-23-2-84 (mean 2-53,average of 56 : 2-56), length last whorl 1-8-2-4 mm . aperture length x width1-1-1-6 x 1-0-1-4 rnm, 6-7 whorls. Table i details the measurements of 56 adultshells ; the holotype is no. 34. ALDABRA STREPTAXIDAE TABLE i 159 Measurements of shells of Gulella gwendolinae aldabrae n. subsp. from Aldabra and Assumption length aperture number of no. length x maj. diam. l/d last whorl length x width whorls locality Anse CedresAldabra, SykesAldabra, SykesHe Michel Aldabra, ThomassetAnse CedresAldabra, SykesAssumption, SykesAnse CedresAldabra, ThomassetHe MichelAldabra, SykesHe MichelAldabra (a), SykesAldabra, SykesAnse CedresAldabra, ThomassetAssumption, SykesPlatin Aldabra (a), SykesAssumption, SykesAnse CedresHe MichelHe MichelHe MichelAldabra, SykesAnse CedresAldabra, ThomassetAssumption, SykesAssumption, SykesAnse CedresHe MichelAnse CedresAnse CedresAldabra, ThomassetAldabra, SykesAldabra (a), SykesHe Michel Aldabra, ThomassetAldabra, ThomassetAnse CedresAldabra (a), SykesAssumption, SykesAldabra, SykesAldabra (a), SykesAldabra (a), SykesAssumption, Sykes i6o A. C. VAN BRUGGEN TABLE i (contd) length aperture number of no. length x maj. diam. l/d last whorl length x width whorls locality Assumption He Picard, Sykes Assumption Aldabra (a), Sykes Aldabra (a), Sykes Aldabra (a), Sykes Aldabra (a), Sykes Platin Aldabra (a), Sykes For explanation of the localities see the text. The shells have been enumerated according to sizetaken from micrometer readings, which are much more accurate than their translation into mm (e.g.62 points = 3-9 mm, but also 63 points = 3-g mm, etc.). No. 34 is the holotype (fig. i, BMNH); no. 49 isthe 'type' of Ennea alauda Sykes nom. nud. Specimens nos. 5, 17, 18, and 48 show (very) faint traces ofapical spiral sculpture. MATERIAL EXAMINED : RECENT : Holotype, ALDABRA ISLAND : South Island (Grande Terre), AnseCedres, c. 400 m inland (Grid ref. 359112) under stones in open grassy areaamongst mature bush, ii.ix.ig67, (Peake, RSE) BMNH. No. 197424. Paratypesinclude all material listed below, similar locality, habitat and collecting data asabove, BMNH No. 197425, RMNH Nos. 54937-54939 ; West Island (He Picard),PSTE/JCFF (E. R. Sykes Colin) ['Type' of Ennea alauda Sykes nom. nud.],BMNH No. 197426 ; West Island, near village (Grid. ref. 957102), in litter aroundCasuarina trees, 7.ix.i967 (Peake, RSE), BMNH No. 197427 ; He Michel (CoconutIsland, Grid ref. 325083), under stones, I4.ix.i967 (Peake, RSE) BMNH No.197428 ; South Island, Platin, near pool (Grid ref. 361101), under stones, n.ix.i967(Peake, RSE), BMNH No. 197429 ; ibid, Platin, near runway trace (Grid ref.363095), in litter amongst bush area, 9.ix.i967 (Peake, RSE), BMNH No. 197430.Poorly localized material : ALDABRA, PSTE/JCFF, BMNH and RMNH,(E. R. Sykes Colin) ; ALDABRA, NM (vide Connolly, 1925 : 265) (H. P. Thoma-sett, H. C. Burnup Colin) ; West Island and Takumaka (= Takamaka). N.B.two different localities, shown in Table i as 'Aldabra (a)', PSTE/JCFF, BMNH(E. R. Sykes Colin). ASSUMPTION ISLAND : PSTE/JCFF, BMNH and RMNH(E. R. Sykes Colin) ; Central area, i6.ix.i967 (Peake, RSE), BMNH No. 197433. FOSSIL : ALDABRA ISLAND : Middle Island (He Malabar), Stn 31 (Grid ref. 301123),cavity fill deposit cut into Aldabra Limestone associated with Tropidophora sp.,age less than 125 ooo years BP and possibly 27 ooo years BP (Taylor et al, RSE),BMNH No. 9921200 ; South Island, Stn 390 (Grid ref. 186024), Solution cavityfill cut into Takamaka Limestone buff 'soil' with abundant rootlets and associatedfauna of Tropidophora and Rachis, age probably last glacial, that is post 125 oooyears BP (Taylor et al, RSE), BMNH No. GG2I20I. Note : Juvenile shells and fragments are expressly excluded from the type series. Material recorded, but not examined : ALDABRA ISLAND : He Verte (Smith, 1909 : 69). ALDABRA STREPTAXIDAE 161 1 mm FIGS 1-4. Aldabra Island Streptaxidae. i, Gulella gwendolinae aldabrae n. subsp.,holotype shell, actual length 4-1 mm ; 2, G. peakei n. sp., holotype shell, actual length2-0 mm ; 3, G. peakei n. sp., juvenile shell from below, to show the angular lamella inthe form of a ridge extending beyond the aperture (scale applies to fig. 3 only) ; 4, G.insulincola n. sp., holotype shell, actual length 4-4 mm, broken in the process of drawing,but repaired. All figured specimens in British Museum (Natural History). 162 A. C. VAN BRUGGEN Distribution on Aldabra is shown in fig. 6. There is little variation in the apertural dentition, except for the columellarcomplex, which may vary in the depth of bifurcation. There is no trace of aperturaldentition in juvenile shells. The above numerical data are for all material examined. The population onAssumption Island is presumably genetically separated from those on AldabraIsland proper. Assumption is situated roughly 30 km to the southeast of Aldabra.However, the number of shells available for Assumption amounts to only nine, sothat no conclusions may be drawn. Measurements are in mm and the followingdata are shown from left to right : length x major diameter, l/d, length last whorl,length x width of aperture, number of specimens measured. Aldabra : 3*6-5-1 x 1-4-1-9, 2-23-2-84, 1-8-2-4, i'i-i'6 x 1-0-1-4, 47 Assumption : 3-7-4-7 x 1-5-1-7, 2-40-2-84, 1-9-2-4, 1-2-1-5 x 1-1-1-4, 9Mean l\d, 2-53, average Ijd 2-54 (Aldabra) ; do., 2-62, 2-66 (Assumption). In bothcases the shells have six to seven whorls. The range of measurements is on thewhole smaller on Assumption, but this may reflect limitations imposed by the smallsample. The only local population on Aldabra which may enjoy a certain degree of spatialand consequently genetical isolation is that on He Michel or Coconut Island. Cor-responding figures for this island also show a smaller range of measurements : lie Michel : 3-6-1-4 x 1-6-1-7, 2-28-2-52, 1-9-2-2, 1-2-1-3 x 1-1-1-2, 8In this case mean and average lid have values of 2-40 and 2-42 respectively and theshells have only 6-6| whorls. However, drawing of conclusions based on eightspecimens again seems unwarranted. lie Michel is only about i km from thenearest land, South Island (Grande Terre), so that there is really no question ofeffective isolation. Smith (1909) recorded this species as 'Ennea sp . . . may prove to be new'. Ayear later Preston (1910) described Ennea gwendolinae from Kenya, with whichConnolly (1925) identified his Aldabra material. Gulella gwendolinae is known fromthe coastal area of Kenya (including the Shimba Hills), the Usambaras and Dar-es-Salaam in Tanzania, west of Lake Rudolf in Kenya and southeast of Lake Chad.This includes the following aberrant forms : scissidens Connolly, 1922 (Dar-es-Salaam) ; porrecta Pfeiffer, 1952 (Mombasa) ; mkusiensis Verdcourt, 1953 (W.Usambaras) ; tsadiensis Blume, 1959 (SE Lake Chad) ; var. nov. Verdcourt, 1962(Turkana). Notwithstanding the authority and scientific acumen of Connolly, it seems atfirst somewhat far-fetched to identify an Aldabra snail with a species of the Africanmainland. The gap between the coast of East Africa and Aldabra atoll consists ofroughly 640 km of ocean, a seemingly unsurmountable barrier for a land snail. Onthe other hand, the character of the fauna of many islands in the western IndianOcean is unmistakably African. The following are two quotations from Peake inWestoll & Stoddart (1971 : 581-610) : The affinities of the vertebrate fauna on allislands in the western Indian Ocean are predominantly orientated towards Africa'(p. 586) ; 'The fauna exhibits a wide range of affinities depending on the taxa andtaxonomic levels considered ; those with Africa are dominant, . . .' (p. 606). Cogan, ALDABRA STREPTAX1DAE 163 Hutson & Shaffer in Westoll & Stoddart (1971 : 315-325) write : To summarize itmay be said that the insect fauna of Aldabra is predominantly African in origin, . . .'(p. 324). Wright in Westoll & Stoddart (1971 : 299-313) concludes that the fresh-water snails of the genus Bulinus on Aldabra are also of African origin, althoughsome may have reached the atoll via Madagascar. All this stresses African originand derivation, but so far no non-marine molluscan species on Aldabra has actuallybeen identified with (East) African species. This has, however, occurred in Ento-mostraca (McKenzie in Westoll & Stoddart, 1971), Diptera, and Lepidoptera(Cogan, Hutson & Shaffer in Westoll & Stoddart, 1971). The Comoros are much closer to the African continent. Although no modernsummary is available, it appears from a scrutiny of a series of papers by Morelet(1860-1885) an d material in various collections that African species indeed do occurhere. TABLE 2 Comparison of measurements of the shells of various populations of Gulella gwendolinae aperture number of material length x maj. diam. Ifd length x width whorls n aldabrae 3-6-5-1 x 1-4-1-9 mm 2-23-2-84 1-1-1-6 x 1-0-1-4 mm 6-7 5^ gwendolinae 4-8-5-4 x 1-7-1-8 mm 2-82-3-09 i -6-1-8 x 1-2-1-4 mm 7 7 porrecta 5-5 x 1-9 mm 2-87 1-7 x 1-4 mm 7 i tsadiensis 3-6-4-6 x 1-6-1-9 mm 2-00-2-50 1-7-1-9 x 1-3-1-5 mm 5^-6 29 n = number of specimens examined. Data shown under gwendolinae are from the holotype in the Ter-vuren museum, four paratypes in the British Museum (Natural History), and one paratype each in theFrankfurt and Leiden museums. Data shown under porrecta are from the holotype in the Frankfurtmuseum and those for tsadiensis have been extracted from Blume (1959). In Table 2 the various forms of G. gwendolinae are compared ; the subspecies orvarieties scissidens, mkusiensis and 'var. nov.' have not been taken into account,because these are rather aberrant. The var. porrecta may be a synonym of the typicalform (Verdcourt, 1962 : 8). A warning may be sounded as to the number of speci-mens considered here. This table shows that the Aldabra form is (a) smaller than the typical form, but larger than tsadiensis, although there is awide overlap ; (b) has a smaller major diameter than both the typical form and tsadiensis,although there is an almost complete overlap ; (c) is not as slender as the typical form, but more so than tsadiensis, albeit with asmall overlap ; (d) has a smaller aperture than both the typical form and tsadiensis, with smalloverlaps ; (e) has somewhat fewer whorls than the typical form, but somewhat more thantsadiensis. This shows on the whole that at least three recognizable units may be distinguished,viz. the Aldabra-Assumption populations, the typical form, and the subspeciestsadiensis. Gulella gwendolinae is a variable species with an apparently wide distri-bution. Verdcourt (1962 : 7, footnote) rightly advises : 'Until considerable material 164 A. C. VAN BRUGGEN is available it would be as well not to bestow more names.' However, in view of theisolated location of the Aldabra-Assumption populations, it is proposed here toseparate these as the subspecies aldabrae. Unfortunately anatomical studies arenot as yet possible because of the lack of preserved soft parts. Gulella gwendolinae aldabrae is the only recent streptaxid on Aldabra atoll. It isreasonable to expect it to have been transported from the African mainland to theisland. Aldabra has always been a remote place and in view of the noticeabledifferences found and fossil occurrence it is unlikely that it has arrived in the wakeof Man. Also, there are no records of the species from either the Comoros orMadagascar (cf. Fischer-Piette & Bedoucha, 1964^). Dispersal by tropical stormsis also unlikely because of the direction of these in the area ; the animal is, however,sufficiently light to be carried by high winds. Dispersal by birds is even less likely ;the snail will certainly hardly have been picked up by the birds which now populateAldabra. Rafting may merit consideration, but there are no suitable surfacecurrents to effect a dispersal from Africa to Aldabra. Obviously G. gwendolinae hasgood dispersal ability, a character it shares with many small terrestrial snails (cf.,e.g., Carlquist, 1965 : 292-293). For the time being no reasonably plausible ex-planation for the presence of G. gwendolinae on Aldabra is available. Gulella peakei n. sp. Figs. 2, 3, 7 DIAGNOSIS. A minute species of Gulella with spaced lamellae and smoothinterstices on the whorls, open umbilicus, and dentition consisting of angular lamella,two labral processes and columellar lamella. DESCRIPTION OF SHELL. Shell (fig. 2) small, cylindrical-ovoid, greatest widthabout the middle, with open umbilicus, costulate, creamy white. Spire produced,sides slightly to markedly convex, subparallel, apex somewhat flattened, obtuselyconical. Whorls six to six-and-a-half, convex and sculptured with comparativelyprominent, regular, straight and perpendicular, widely distant, costulae, intersticesmuch wider than riblets, smooth, under high magnification very finely granulate.In front view the holotype shell shows only about eleven costulae on the part ofthe whorl above the aperture ; the last whorl has a total of about fifteen riblets.Initial two whorls smooth, very fine granulate under high magnification ; penultimateand last whorls comparatively small. Sutures shallow, simple to subcrenellate,somewhat impressed. Aperture somewhat oblique, invertedly triangular withsmoothly rounded base, about as high as wide, peristome fairly thick, expanded andsomewhat reflected, dentition more or less four-fold. To the right of the middle ofparies a fairly large, obliquely perpendicular, angular lamella, which is nothing but aV-shaped pleat in the peristome, connected with labrum ; about half-way down thelabrum a superficial swelling may be interpreted as a labral process ; somewhatbelow this, but much deeper inside the aperture, and at a slight distance from thetip of the angular lamella, a blunt inner labral process is seen, slightly above whichthere is a much smaller and less prominent process or mere swelling (not shown in ALDABRA STREPTAXIDAE fig. 2 because hidden behind superficial labral swelling) ; columellar lamella large,blunt and prominent. The main inner labral process corresponds to a shallowdepression on the outside of the aperture ; the columellar lamella corresponds to ashallow furrow on the left of the outside bottom of the aperture, thus adjoining theumbilicus. TABLE 3 Measurements of shells of Gulella peakei n. sp. length aperture number of additional no. length x maj. diam. l/d last whorl length x width whorls data apex damagedapex damaged aperture ratheroblique holotype, fig. 2 aperture damagedaperture obscuredby matrix matter Nos. 1-13 are from Stn 34F, no. 14 from Stn 26F, Aldabra. MEASUREMENTS OF SHELL : 1-7-2-1 x 1-3-1-5 mm, Ijd 1-30-1-57 (mean 1-43,average of 14 : 1-44), length last whorl 0-8-0-9 mm > aperture length x width 0-6-0-7 x 0-6-0-7 mm > 6-61 whorls. Table 3 details the measurements of 14 adultshells ; the holotype is no. n. MATERIAL EXAMINED : FOSSIL : Holotype, ALDABRA ISLAND : Middle Island (He Malabar), Stn 34F (Gridref. 293109), age inferred as 27 ooo BP (Taylor et al, RSE), BMNH No. GG2I2O2.Paratypes, similar locality and information as above, 12 shells in good condition(2-13 of Table 3) BMNH No. GG2I2O3 and RMNH 54940-3, 14 further paratypesin poor condition or still covered by the matrix BMNH No. GG2I204, numerousjuvenile shells and fragments ; South Island (Grande Terre), Stn 26F (Grid ref.337055), age uncertain but possibly same as 34F (Taylor et al, RSE), BMNH No.GG2I2O5, i paratype (14 of Table 3) and 3 juvenile shells. Other material : South Island, Dune d'Messe, Stn 3gA (Grid ref. 186029), ageinferred as 27 ooo BP (Taylor et al, RSE), BMNH No. GG2I206, some shellfragments. Note : Juvenile shells and fragments are expressly excluded from the type series. 166 A. C. VAN BRUGGEN Distribution (fig. 7). Quaternary of Middle Island (He Malabar) and South Island(Grande Terre), Aldabra Island. Dr Taylor has kindly furnished the followingdetails on the localities where Guletta peakei has been obtained. Stn 26F : 'Browncavity-fill deposit in Takamaka Limestone. Many small gastropods. Age un-certain, possibly pre-125 y rs -' Stn 34F : 'Cavity-fill deposit cut into theTakamaka Limestone, buff 'soil' containing abundant large ribbed Tropidophora(only site for this species). Abundant Assiminea, Gulella within cavities of theTropidophora and in the matrix. Almost certain last glacial, post 125 ooo yrs BP.'Stn 3gA : 'Solution cavity-fill cut into Takamaka Limestone. Buff 'soil' withabundant rootlets and associated fauna of Tropidophora and Rachis. Age probablylast glacial, post 125 ooo yrs BP.' The species has been named after Mr J. F. Peake, Deputy Keeper of Zoology andHead of the Mollusca Section of the British Museum (Natural History), as a tokenof friendship and admiration for his island research. The angular lamella is present in all juvenile shells in the form of a long andconspicuous, simple, ridge, usually stretching somewhat beyond the aperture (fig. 3).Damaged shells show that this ridge is being resorbed on the one end in the courseof growth while being added to at the actual aperture. No other dental processesare present in the juvenile shells. Juvenile dentition is rare among species of thegenus Gulella and much more common among representatives of allied genera, suchas Ptychotrema (vide, e.g., van Bruggen, 1971, fig. 2, p. 249). Among the about 125species of Guletta in Southern Africa there are about three species in which thisphenomenon has been described (Burnup, 1925 ; Connolly, 1939), although juvenileshells are as yet unknown for a number of species. At least three of the specieswhich will be considered below when trying to assess the relationships of the newspecies also have juvenile shells with apertural dentition, viz. G. jacquelinae Adam,G. pooensis Ortiz de Zarate & Ortiz de Zarate and G. spatium (Preston) (see Adam,1965 ; Ortiz de Zarate & Ortiz de Zarate, 1956 ; Verdcourt, 1970, the latter asinterpretation of Blume, 1965). Guletta peakei has no allies on Europa Island (so far no streptaxids have beenreported from this island : Fischer-Piette & Bedoucha, ig64a ; Legendre, 1966 ;Fischer-Piette & Vukadinovic, 1971), Madagascar (Fischer-Piette & Bedoucha,1964^, the Mascarene Islands (Germain, 1921 ; Connolly, 1925), the Seychelles(Sykes, 1909 ; Connolly, 1925 ; Barnacle, 1962), the Comoros (Morelet, 1860, 1877,1879, 1881, 1882, 1883, 1885 ; Von Martens, 1876), or Aldabra Island (Von Martens& Wiegmann, 1898 ; Smith, 1909 ; Connolly, 1925 ; Barnacle, 1962). A few of thespecies enumerated by Morelet for the Comoros, such as Pupa minuscula Morelet(1877 : 340, pi. 12, fig. 5), which may be a streptaxid, superficially resemble Gulettapeakei, but never show the peculiar spaced lamellae on the whorls. As regardsPupa minuscula Morelet writes 'obsolete costulata', and 'Le test est orne de cotesfines, espacees sur le dernier tour'. Gulella peakei certainly does not have fine ribs,spaced on the body whorl, but rather shows widely spaced lamellae all over the shellexcept for the apex. The new species obviously belongs to Verdcourt's 'Key 4' (Verdcourt, 1962 : 8).This key features East African species with spaced lamellae on the whorls. This is ALDABRA STREPTAXIDAE 167 probably not a natural group, although some of the species may be allied to eachother. They belong to Costiguletta Pilsbry, 1919, Mirigulella Pilsbry & Cockerell,1933, and Aenigmigulella Pilsbry & Cockerell, 1933 ; these are all considered sub-genera of Gulella L. Pfeiffer, 1856, by Zilch in his manual (Zilch, 1959-60 ; see alsoZilch, 1961). The subgenera Mirigulella (monotypic) and Aenigmigulella (with twospecies, cf. Adam, 1965 : 40) are very probably products of a long and separatedevelopment ; the subgenus Costiguletta with perhaps about ten species altogetheris rather an assemblage of diverse elements. A preliminary assessment of the speciesof Costiguletta shows that this subgenus may consist of three groups, viz. (a) G. langi Pilsbry and G. toticostata Pilsbry, both from the Congo (Zaire), and G.pooensis Ortiz de Zarate & Ortiz de Zarate from Fernando Poo, with lamellae andspiral sculpture on the whorls (Costiguletta s.s.), (b) the Kenya taxa G. adjacens (Preston), G. spatium (Preston), G. p. pretiosa(Preston) and G. p. nyiroensis (Preston), with costulae in between the lamellae on thewhorls (see also Adam, 1965 : 46, who states that all three may also have costulateearly whorls), and (c) the West African species G. hedwigae Degner, with smooth interstices betweenthe lamellae on the whorls. G. microtaenia Pilsbry & Cockerell most probably does not belong to Costigulettas. lat. because of the 'delicate riblets' (Pilsbry & Cockerell, 1933 : 372). For thetime being the present author refrains from naming the subdivisions of Costigulettas. lat., particularly because of the differences in dentition of the shells and the ab-sence of anatomical data. However, zoogeographically the above three groups atfirst sight seem to be fairly natural : group (a) inhabits the West and CentralAfrican equatorial forest, group (b) the East African forests and group (c) thewestern parts of the West African equatorial forest. The subgenera Aenigmigulellaand Mirigulella, both from East African forests on elevated country, may haveancestors in common with group (b). Only Mirigulella has a reduced dentition ;the forest element is usually of a more primitive nature than species or groups outsidethe forest. Therefore one may be tempted to consider all above taxa (all forestdwellers in the uplands, or lower down where the forest comes down to sea level,such as on Fernando Poo) to be more primitive than Mirigulella. Compared with G. peakei all the above species are either too large, or have a dif-ferent dentition or costulation, but usually one finds a combination of all three factors.The writer has been able to study type material or other specimens of most of thesespecies in the museums in London, Frankfurt am Main and Leiden. G. peakei is notto be confused with any of the species under discussion. By virtue of the absenceof sculpture on the interstices between the lamellae on the whorls the new speciesbelongs to group (c) of Costiguletta s. lat. Zoogeographically this is a somewhatsurprising conclusion, because one hardly expects a species from Liberia and onefrom Aldabra to have common ancestors. Of course, the minute terrestrial snailsof continental Africa are still very incompletely known and G. hedwigae or alliedspecies yet to be discovered may well occur further east. Apart from other considera-tions it seems fairly certain that the ancestor of G. peakei is of continental Africanorigin. However, the possibility that a likeness to G. hedwigae may have been caused 168 A. C. VAN BRUGGEN by convergent evolution cannot be ruled out. Only anatomical data are likely tohelp solve this question ; the fact that G. peakei is extinct will, however, frustratesuch a project. G. peakei is obviously extinct on Aldabra Island. Species resembling it are re-stricted to forest habitats in Africa ; there is no more suitable forest habitat availableon Aldabra, indeed the overall impression is that of a 'semi-arid island' (Stoddart inWestoll & Stoddart, 1971 : 8). Aldabra is an elevated atoll consisting of an elevatedreef situated on the summit of a mountain rising from the sea floor. There is evidencethat in the past Aldabra has been much more elevated (Stoddart et al in Westoll &Stoddart, 1971 : 31-66, and personal communication of Dr J. D. Taylor), whichmust have resulted in a moister climate and consequently also the presence of atype of forest suitable for species such as G. peakei. Some of the islands in theWestern Indian Ocean still have a fair amount of forest, but so far no apparent alliesof G. peakei have been obtained here. Gulella insulincola n. sp. Figs. 4, 8 DIAGNOSIS. A small species of Gulella with smooth whorls, open umbilicus, anddentition consisting of angular lamella, two labral processes, a basal denticle, andcolumellar lamella. DESCRIPTION OF SHELL. Shell (fig. 4) small, subcylindriform, with open umbilicus,smooth, creamy white. Spire produced, sides subparallel, apex flattened, obtuselyconical. Whorls six to six-and-a-half, slightly convex, almost completely smooth,only with traces of costulation or striation behind the labrum, initial whorls smooth,very finely granulate under high magnification ; sutures shallow, simple, somewhatimpressed, occasionally subcrenellate. Aperture subquadrate, rounded at base,peristome incrassate and reflected, white and probably glossy when fresh, with five-fold dentition : a reasonably well-developed slightly oblique angular lamella,touching or connected with the tip of the labrum ; two subequal mid-labral denticleson a slightly raised common base, labral complex corresponding to very shallowexternal pit ; a small mid-basal denticle, which usually is situated slightly to theleft of the middle of the base and which may be so small as to be hardly noticeable,in which case the dentition may be interpreted as being four-fold ; columellar lamellasmall and little prominent, blunt and fairly deep-set. MEASUREMENTS OF SHELL : 3-9-4-4 x 1-9-2-0 mm, l/d 2-07-2-29 (mean 2-18,average of 4 : 2-14), length last whorl 2-1-2-3 mm, aperture length x width, 1-3-1-6 x 1-2-1-3 mm, 6-6| whorls. Table 4 details the measurements of four adultshells ; the holotype is no. 4. MATERIAL EXAMINED : FOSSIL : Holotype, ALDABRA ISLAND : Middle Island (He Malabar), Stn 34F (Gridref. 293109), age inferred as 27 ooo BP (Taylor et al, RSE), BMNH No. 21216.Paratypes, similar locality and information as above, 4 shells (Taylor et al, RSE) , ALDABRA STREPTAXIDAE 169 BMNH No. 21217. Collected together with Gulella peakei n. sp. ; for details oflocality see data under 'Distribution' for that species. All specimens are in poorcondition, the shells being worn and very fragile. Distribution (fig. 8). So far only known from the Quaternary of He Malabar orMiddle Island. TABLE 4 Measurements of shells of Gulella insulincola n. sp. from Stn 34F, Aldabra length aperture number of additional no. length x maj. diam. Ijd last whorl length x width whorls data 1 3-9 x 1-9 mm 2-07 2-1 mm 1-3 x 1-2 mm 6 2 4-1 x 1-9 mm 2-09 2-2 mm 1-4 x 1-3 mm 6+ apex damaged 3 4-2 x +2-omm 2-09 2-2 mm 1-4 x 1-3 mm 6J body whorl damaged 4 4-4 x 1-9 mm 2-29 2-3 mm 1-6 x 1-3 mm 6 holotype broken but repairedThe holotype shell, no. 4, was broken, but has been repaired. The specific name insulincola is a noun derived from insula (Lat. : island) andincola (Lat. : inhabitant). The new species represents a common pattern in the genus Gulella. Taxa witha smooth shell and a five-fold dentition have been brought together for SouthernAfrica by Connolly (1939 : 20) as group 4 (ii) (10 species) and the East African onesare treated by Verdcourt (1962 : 20) in part of Key 4 (9 species). Many of thesespecies show a dental pattern similar to that of G. insulincola. Among the islanddwellers G. poutrini (Germain, 1918), which is common on Mauritius, and G.comorensis (von Martens, 1876) of the Comoros exhibit the same pattern. Com-parison with species with the combination of characters as described above hasfailed to provide satisfactory identification, so that we may conclude that G. in-sulincola represents indeed a hitherto undescribed species. Its closest allies areperhaps the above species from Mauritius and the Comoros ; these differ from thenew species in being much larger (sometimes twice as large), having more whorlsand being less slender than G. insulincola. Perhaps the group with a smooth shelland a dental pattern consisting of angular lamella, two labral processes, a basaldenticle, and a columellar lamella, is a natural one distributed over much of West,Central, East and Southern Africa, and on the islands in the Western Indian Oceanas well. The possibility that G. poutrini and G. comorensis may be the componentsof a superspecies or represent the subspecies of a widely-dispersed species may bemerely food for thought in this respect. On the other hand, a shell type such as hasbeen discussed here occurs throughout the genus Gulella and may well be the resultof convergent evolution. G. insulincola is obviously extinct on Aldabra, which may also be due to the factthat suitable habitat has disappeared in the course of the process of the islandbecoming progressively drier. 1 7 o A. C. VAN BRUGGEN Gulella spec. There are two specimens (BMNH) which represent a fourth species of the genusGulella. Both are clearly fossils. The one, from 'Stn 34F' (see sub G. peakei sp. n.),is a juvenile shell of 3-2 x 2-0 mm with five whorls, of which the initial ones aresmooth and the others sculptured with somewhat undulating costulae. The other -10 40 50 , AFRICA -0 60 INDIAN OCEAN 10 20 8- 40 FIG. 5. Map showing position of Aldabra Island, i, Aldabra ; 2, Assumption ; 3,Comoros ; 4, Seychelles ; 5, Europa ; 6, Reunion ; 7, Mauritius ; 8, Rodriguez (6-8Mascarenes) . ALDABRA STREPTAXIDAE 171 specimen is a much younger juvenile shell of probably the same species ; it measuresonly 2 -3 mm and the aperture is filled with matrix matter. This shell was obtainednot far from the settlement on West Island (He Picard), grid ref. 063 E-og8 N (fig. 8),'cavity-fill cut into Basin Cabris calcarenites and Takamaka limestone, 'white soil'also containing Rachis and Tropidophora. Age uncertain ; could be the same as3gA and D'. (Notes supplied by Peake and Taylor ; Stns 3gA and 390 have beendescribed above under the other species.) No opinion as regards identity of thepresent species is ventured here : it is most likely that this is also an extinct species. DISCUSSION A thorough survey of the terrestrial molluscs of Aldabra has revealed the presenceof four species of Streptaxidae, viz. Gulella gwendolinae aldabrae n. subsp., the onlyRecent representative of the family on the island, and three extinct taxa, G. peakein. sp., G. insulincola n. sp. and an incompletely known species, G. spec. The family Streptaxidae has excellent dispersal abilities, at least in the WesternIndian Ocean. Streptaxids have reached both the Seychelles and the MascareneIslands at distances of 1400 and 2500 km from the African mainland respectively.The Mascarene Islands are also situated betweeen 700 and 1300 km from Madagascarfrom whence some streptaxid ancestors may have been derived (fig. 5) . Asia (India)is more than 2500 km from the Seychelles and there are few direct relationships (ifany) with the streptaxids of that continent. G. gwendolinae is a very widely dispersed African species with a tendency to becomelocally separated into reasonably recognizable subspecies. Its dispersal abilitiesare sufficiently illustrated by its wide distribution on Aldabra and Assumption(fig. 6) ; moreover, it is also locally abundant - sufficient reason to consider it asuccessful species in an evolutionary sense. Perhaps the adaptability to widely diver-gent climatic conditions and types of vegetation has been the key to its success in Africa,which at the same time has accounted for its continued survival on Aldabra. Fossiloccurrence shows that it has been on the atoll for a long time, very probably havingbeen already a contemporary of the extinct G. peakei, G. insulincola and G. spec.Streptaxids are carnivores known to feed on soft invertebrates, mainly other ter-restrial molluscs, and particularly snails of the pulmonate family Subulinidae (vanBruggen, 1967 : 186), which family is not (yet?) known to occur on Aldabra. How-ever, there are Subulinidae on the high islands of the Western Indian Ocean. Thereare a few other land snails on the atoll, of which the enid Buliminus (Rhachis)aldabrae von Martens, 1898, is The most common of all the species from Aldabra.'(Smith, 1909 : 70 ; see also Connolly, 1925 : 264-266). Fossil evidence indicatesthat many terrestrial snails were contemporaries of G. peakei, G. insulincola and G.spec. G. peakei, G. insulincola, and G. spec, are extinct and may well have been in-habitants of types of vegetation which have disappeared on Aldabra. The relativesof at least G. peakei are restricted to forest habitats such as are no longer availableon Aldabra. There are still remnants of forest on high islands, e.g. the Comoros,Seychelles and Mascarene Islands, of which the latter two have species of Gulella 172 A. C. VAN BRUGGEN FIGS 6-8. Maps showing the distribution on Aldabra Island of 6, Gulella gwendolinaealdabrae n. subsp. (the arrow points to Assumption) ; 7, G. peakei n. sp. ; 8, G. insulincolan. sp. (dot) and G. spec, (asterisks). which may be the nearest allies of G. insulincola. This species or its ancestors maytherefore have arrived secondarily from these islands, although Aldabra is closer tothe African continent than both the Seychelles or the Mascarene Islands. There is ALDABRA STREPTAXIDAE 173 also other evidence that Aldabra once harboured a much more varied flora withforest components and that a progressive drying out of conditions has caused theseto disappear together with the forest dwellers among the animals that led a shelteredlife in the then available leaf mould. This has caused some of the predators, thethree Gulella species, and perhaps also their prey, species of the family Subulinidac,to disappear for ever. Finally one has to consider how the streptaxids have reached remote Aldabra.The island has never been connected with the African continent, but Aldabrastreptaxids have strong links with those from that continent. Dispersal throughhuman agency, by tropical storms, by birds, and by rafting have all been ruled outwhen G. gwendolinae aldabrae was discussed above. The same applies, mutatismutandis, for the extinct species. Yet, much of the present flora and fauna ofAldabra atoll or their ancestors have obviously come from Africa. Perhaps dispersalalong the usual paths has taken place in the past when the direction of wind and sur-face currents was more favourable to such a process than today. The present article was finalized early in 1973. Therefore the following comprehensivepaper on the land molluscs of the Comoros has not been taken into account : Fischer-Piette, E.& Vukadinovic, D. 1974. Les mollusques terrestres des lies Comores. Mem. Mus. natn. Hist,nat. Paris (N.S.) (A) 84: 1-76. The checklist includes various African species. The familyStreptaxidae appears to occupy a dominant position with 46 species. Gulella gwendolinaealdabrae n. subsp. should be compared to G. dentiens (Morelet, 1883) as figured by Fischer-Piette& Vukadinovic (fig. 18 on p. 59). REFERENCES ADAM, W. 1965. Mission zoologique de 1'I.R.S.A.C. en Afrique orientale (P. Basilewsky et N. Leleup, 1957)- LXXXV. - Mollusca Streptaxidae. Annls Mus. r. Afr. centr. Ser. 8 Sci. zool. 138 : 1-52.BARNACLE, G. A. S. 1962. The land and freshwater shells of the Seychelles group of islands (including the Amirantes, Coetivy, Farquhar, Cosmoledo and Aldabra). /. Seych. Soc. 2 : 53-57-BLUME, W. 1959. Mollusken aus dem Tschadsee-Gebiet. Opusc. zool. Munch. 29 : 1-9. - 1965. Die Mollusken, die Herr Prof. Franz hauptsachlich wahrend seiner letzten Reise inInnerafrika gesammelt hat. Opusc. zool. Munch. 90 : 1-17. BRUGGEN, A. C. VAN. 1967. An introduction to the pulmonate family Streptaxidae. /.Conch., Land. 26 : 181-188. - 1971. Some Streptaxidae (Mollusca) from West and southern Africa with the descriptionof a new species of Gulella. Zool. Meded., Leiden 45 : 245-260. BURNUP, H. C. 1925. On some South African Gulellae, with descriptions of new species and varieties. Part I. Ann. Natal Mus. 5 : 101-158. CARLQUIST, S. 1965. Island Life, A Natural History of the Islands of the World. New York.CONNOLLY, M. 1922. Notes on African non-marine Mollusca, with descriptions of many new species. Ann. Mag. nat. Hist. (9) 10 : 485-517. 1925. Notes on a collection of non-marine Mollusca from the islands of the Indian Ocean./. Conch., Lond. 17 : 257-266.1939. A monographic survey of South African non-marine Mollusca. Ann. S. Afr. Mus. 33 : i -660. CROSSE, H. 1876. Diagnoses Molluscorum novorum. /. Conchyl., Paris, 24 : 166-167.DEGNER, E. 1934. Westafrikanische Landschnecken. I. Streptaxiden, Helicarioniden, Vaginuliden. Zool. Jb. Syst. 65 : 209-308. 174 A - C. VAN BRUGGEN FiscHER-PiETTE, E. & BEDoucHA, J. 1964%. Mollusques terrestres de 1'ile Europa. Bull.Mus. natn. Hist. nat. Paris (2) 36 : 502-505. & 19645. Mollusques terrestres de Madagascar. Famille Streptaxidae. Bull. Mus.natn. Hist. nat. Paris (2) 36 : 368-376. & VUKADINOVIC, D. 1971. Suite aux mollusques terrestres de 1'ile Europa. Bull. Mus.natn. Hist. nat. Paris (2) 42 : 1277-1281. GERMAIN, L. 1921. Faune malacologique terrestre et fluviatile des lies Mascareignes. Paris. 1934. L'origine et la composition de la faune malacologique terrestre et fluviatile des liesSechelles. 6jme Congr. Soc. sav. : 113-133. KOBELT, W. 1905-06. Die Raublungenschnecken (Agnatha). Zweite Abtheilung : Strept-axidae und Daudebardiidae. Syst. Conchyl. Cab. 1 (126, 2) : 1-211. Nuremberg. LEGENDRE, E. 1966. Liste des invertebres terrestres libres actuellement connus de ile Europa(Hexapodes et Arachnides exceptes). Mem. Mus. natn. Hist. nat. Paris (N.S.) (A) 41 : 211. MARTENS, E. VON. 1876. Conchylien von den Comoren. Jb. dt. malakozool. Ges. 3 : 250-253. & WIEGMANN, F. 1898. Land- und Siisswasser-Mollusken der Seychellen nach denSammlungen von Dr. Aug. Brauer. Mitt. zool. Samml. Mus. Naturk. Berl. 1 (i) : 1-96. MOLLENDORFF, O. von & KOBELT, W. 1903-05. Die Raublungenschnecken (Agnatha).Erste Abtheilung : Rhytididae und Enneidae. Syst. Conchyl. Cab. 1 (126, i) : 1-362. MORELET, A. 1860. Ilesorientalesdel'Afrique. Sdries conchyliologiques comprenant I' Enumera-tion de mollusques terrestres et fluviatiles recueillis pendant le cours de differents voyages, ainsique la description de plusieurs especes nouvelles 2 : 37-127. Paris. 1877. Excursion conchyliologique dans 1'ile d'Anjouan (Johanna). /. Conchyl., Paris,25 : 325-347. 1879. Recolte de M. Bewsher a 1'ile d'Anjouan (Comores). /. Conchyl., Paris, 27 : 308-315. 1 88 1. Malacologie des Comores. Recolte de M. Marie a 1'ile Mayotte. /. Conchyl.,Paris, 29 : 212-241. - 1882. Malacologie des Comores. Recolte de M. Marie, a 1'ile Mayotte. /. Conchyl.,Paris, 30 : 185-200. - 1883. Malacologie des Comores. Recolte de M. E. Marie a 1'ile Mayotte. /. Conchyl.,Paris, 31 : 189-216. 1885. Malacologie des Comores. (46 article.) Recolte de M. Humblot a la GrandeComore. /. Conchyl., Paris, 33 : 288-301. ORTIZ DE ZARATE, LOPEZ A. & ORTIZ DE ZARATE, ROCANDIO A. 1956. Contribuciones al conocimiento de la fauna malacologica terrestre de la isla de Fernando Poo. Boln. R. Soc. esp. Hist. Nat. (Biol.), 53 : 75-140. PFEIFFER, K. L. 1952. Neue Landschnecken aus Ostafrika. Arch. Molluskenk. 81 : 89-102.PILSBRY, H. A. 1919. A review of the land mollusks of the Belgian Congo chiefly based on the collections of the American Museum Congo Expedition, 1909-1915. Bull. Am. Mus. nat. Hist. 40 : 1-370. & COCKERELL, T. D. A. 1933- African Mollusca, chiefly from the Belgian Congo. Proc.zool. Soc. Lond. 1933 : 365-375. PRESTON, H. B. 1910. Additions to the non-marine molluscan fauna of British and GermanEast Africa and Lake Albert Edward. Ann. Mag. nat. Hist. (8) 6 : 526-536. SMITH, M. 1909. The land Mollusca of Aldabra. Nautilus, 23 : 69-70. SYKES, E. R. 1909. The land and freshwater Mollusca of the Seychelles archipelago. Trans.Linn. Soc. Lond. (Zool.), 13 : 57-64. TRYON, G. W. 1885. Testacellidae, Oleacinidae, Streptaxidae, Helicoidea, Vitrinidae,Limacidae, Arionidae. Man. Conch. (2) 1 : 1-364. Philadelphia. VERDCOURT, B. 1953. Notes on some East African Gulellae. Basteria, 17 : 36-42. - 1960. Some further records of Mollusca from N. Kenya, Ethiopia, Somaliland and Arabia,mostly from arid areas. Revue Zool. Bot. afr. 61 : 221-265. 1962. Preliminary keys for the identification of the species of the genus Gulella Pfr.occurring in East Africa excluding the sections Primigulella Pilsbry and Plicigulella Pilsbry(Mollusca - Streptaxidae). Annls Mus. r. Afr. centr. Ser. 8 Sci. zool. 106 : 1-39. ALDABRA STREPTAXIDAE 175 VERDCOURT, B. 1970. A reassessment of species described from East Africa by W. Blume. /. Conch., Lond. 27 : 121-125.WESTOLL, T. S. & STODDART, D. R. (eds.) 1971. A discussion on the results of the Royal Society Expedition to Aldabra, 1967-68. Phil. Trans. Roy. Soc., Lond. B 260 : 1-654.ZILCH, A. 1959-60. Gastropoda Euthyneura. Handb. Paldozool. (6) 2 : i-xii + 1-834. Ber-lin -Nikolassee. - 1961. Die Typen und Typoide des Natur-Museums Senckenberg, 24 : Mollusca, Strept-axidae. Arch. Molluskenk. 90 : 79-120. Dr A. C. VAN BRUGGEN Department of Systematic Zoology of the University C/O RlJKSMUSEUM VAN NATUURLIJKE HlSTORIE RAAMSTEEG 2 LEIDEN HOLLAND QUICKIA ALDABRAENSIS (MOLLUSCA,GASTROPODA: PULMONATA, SUCCINEIDAE), A NEW SPECIES OF LAND SNAIL FROMALDABRA ATOLL, WESTERN INDIAN OCEAN By C. M. PATTERSON INTRODUCTION THE Succineidae, a rather diverse land snail family, currently includes 12 recentgenera (Patterson, 1971, I973a). Succinea, Oxyloma and Catinella have received themost attention by researchers, while representatives of the remaining genera havescarcely been studied. Quickia is distinguished from other succineids by the positionof the right tentacular assembly (situated entirely to the inside of the terminalgenitalia) combined with the lack of a penial sheath and penial appendix. Someunique features of Quickia were first reported by H. E. Quick (1936), but the genuswas not named formally until 14 years later by Odhner (1950). The distribution ofQuickia given by Odhner was '. . . from Liberia to the Cameroons and Gabon, onPrince Island and San Thome to East Africa (Zanzibar and Mauritius, as well as,according to Madge, 1938, Rodriguez, Reunion and the Seychelles).' Subsequently,Quickia received no further attention until the reproductive anatomy and chromo-some number of Q. spurca (Gould) was described (Patterson, 1968). Following that,the existence of Quickia in India was reported in a morphological and cytologicalstudy of two Indian species (Patterson, 1970). In addition to its unique genitalmorphology, Quickia is of cytological interest because all five species studied have25 pairs of chromosomes, the highest number known in the Succineidae. The objectives of this report are (i) to describe a new species of Quickia from theIndian Ocean island (atoll) of Aldabra and to record its chromosome number ; (2)to briefly review the distribution of Quickia and (3) to discuss some aspects of thebiology and systematics of the genus. MATERIALS AND METHODS Several living specimens of Quickia aldabraensis were forwarded to me by C. A.Wright of the British Museum (Natural History) in January 1968. Eleven successivegenerations were produced during the following two years. Methods of laboratoryculture are described in Patterson (1971, 1972). The following locality data andhabitat description were provided by J. F. Peake (personal communication) also ofthe British Museum (Natural History). The specimens were collected by J. D.Taylor while participating in the Royal Society Expedition (1967-68) to AldabraAtoll. The snails were found on the platin area at the eastern end of South Island(Fig. i) where the surface limestone is impervious to rain water. They were observed Bull. By. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 1 78 C. M. PATTERSON r Middle Island ALDABRA ATOLL FIG. i. Collecting site for Qiiickia aldabraensis. variously in areas of short grass, bare rock or close to pools of fresh or somewhatbrackish water. They were also found hidden under rock slabs, in cracks or otherplaces of probable protection from desiccation. The climate of Aldabra Atoll isdry for most of the year with a short wet period which does not necessarily occurregularly. Dissections of relaxed, alcohol-preserved specimens were made under 120 x or250 x magnification using a Wild M5 stereoscopic microscope. All drawings wereprepared with the aid of a camera lucida attachment. For details of specimenpreparation and dissection see Patterson (1971). SPECIES DESCRIPTION AND OBSERVATIONS Quickia aldabraensis, sp. n. Shell Shells from both field-collected and laboratory-reared Quickia aldabraensis varyfrom attenuate to somewhat more ovate (Plate i and Figs. 3, 4). The same kind ofvariation was noted among Q. spurca shells (Patterson, 1968). The attenuate formof adult laboratory-reared Q. aldabraensis has three whorls and measures 8-5-9 mmin height and 4-5-5 mm in width. The teardrop-shaped aperture is 575-6 mm highand 4 mm wide. More ovate shells measure 8-5-9 mm m height, 5-5-6 mm inwidth and have an aperture 6 mm high and 4 mm wide. The largest shells mayreach a height of 10 mm. Large adult shells collected from the field (Fig. 3) werecomparable in size to those reared in the laboratory. Shells cleansed with sodiumhypochlorite have a translucent light amber colour. Mud was adherent to mostshells obtained from the field and appeared to be arranged in three spiral ridges onsome specimens while no definite arrangement was discernible on others. Mudridges were not present on shells of laboratory reared snails. Fine growth linesprovide a minimal amount of shell sculpture. The outer shell surface is dull butthe inside is very glossy. There is a well-defined columellar plait and a weaklydeveloped, untwisted columellar fold. The peripheral margin of the shells is roundedwith moderately impressed sutures. The holotype (Plate i) is deposited in themollusc collection of the British Museum (Natural History) (BMNH No. 1973103).Paraty pes (both field collected and laboratory reared) are deposited in the mollusc QUICKIA ALDABRAENSIS SP. N. 179 collections of both the British Museum (Natural History) (BMNH No. 1973104) andthe University of Michigan, Museum of Zoology. Jaw The jaw is small but relatively strong. Jaw colour varies from a medium brownto a translucent amber, with darker brown markings on the cutting piece and basalaccessory plate (Fig. 5). It is higher than wide when measured at its greatest dimen-sions. The arms of the cutting piece are rounded with tapering anterior extremitieswhich bend toward the central longitudinal body axis. The anterior margin of thecutting piece is deeply convex with a median prominence varying from scarcelynoticeable to conspicuous. The sides of the accessory plate slant slightly inwardand the posterior margin is nearly straight with rounded corners. Radula The radula formula for Quickia aldabraensis is 14-15 : 8-9 : i : 8-9 : 14-15. Thereare usually 8 lateral and 14 marginal teeth. The radula has typical succineid-like features (see Patterson, 1971) and possesses no especially distinctive characters. External body morphology The body of living Quickia aldabraensis, especially the head-foot region, has astrikingly red coloration. However, all the red colour is lost in alcohol preservation.In both living and preserved snails, the internal organs positioned within the head-foot are visible through the body wall. Black pigmentation on the head-foot andanterior mantle border varies from almost none to a maximum amount shown inFig. 6. The kidney, visible through both the body wall and shell, is yellow in colour,bi-lobed on the animal's left side and has the shape shown in outline on Fig. 6. The footof adult, relaxed and preserved specimens is 10- 1 1 mm in length and 4-5-5 mm in width . Reproductive anatomy A ventral view of the preserved reproductive system of Quickia aldabraensis isshown in Fig. 7. The ovotestis is of moderate size and consists of 70-80 acini. Ithas a dull, light beige coloration. The ovotestis duct is very narrow as it departsfrom the ovotestis and has a white colour. It soon becomes distended to serve asthe seminal vesicle along most of its length. The seminal vesicle portion is usuallycoiled once and folded three times along its length. The duct again becomes anarrow channel as it opens into the upper portion of the spermoviduct. There aretwo subequal, narrow receptacula seminis and a well-developed fecundation pouch(Fig. 8). Both have a translucent white colour. The cream-coloured albumengland is of moderate size, has a linguiform shape and follicular appearance. Thetubules of the albumen gland are approximately one-half the size of ovotestis acini.The spermoviduct and oviduct have a creamy colour and are compactly folded alongtheir length. The inner folds of tissue along the uteral portion of the oviduct arevisible through the duct wall. The spermatheca is nearly spherical, of moderatesize and cream coloured. Its duct passes along the proximal ventral wall of theuteral portion of the oviduct, courses anteriorly and joins dorsally with the oviduct i8o C. M. PATTERSON at a level slightly posterior to the end of the penis (Fig. 9). Thus, the vagina isquite short in this species. The walls of the anterior portion of the oviduct andvagina become thicker and somewhat muscular in appearance. The prostate glandis of moderate size and white in colour. It is composed of a rather loose collectionof tubules (Fig. 10) which were not separable into distinct bunches as in Q. spurca.There are many channels opening from the prostate gland into the vas deferens(Fig. 10). The vas deferens passes ventrally and anteriorly from the end of theprostate gland. It then passes dorsally over the oviduct and inserts at the apexof the penis. The penis is a simple sheathless tube approximately i mm in length(Fig. 9). It is situated on the right of the vagina in ventral view (Fig. 7). A cross-section shows the inner folds and the lumen of the penis (Fig. n). The long, thinpenial retractor muscle originates adjacent to the vas deferens, passes under theright tentacle assembly and courses posteriorly to its insertion in the tissue coveringthe anterior portion of the digestive gland. The common reproductive channel(Fig. 7) is usually equal to, or occasionally shorter than, the length of the penis. Ithas a muscular appearance and opens to the exterior via the oval-shaped genitalaperture (Fig. 9). Chromosome number The haploid chromosome number of Quickia aldabraensis is n = 25, the samenumber observed in Q. spurca, Q. bensoni and Q. calcuttensis , as well as in members oftwo other unidentified Quickia from Tamilnadu, India. Fig. 2 shows a cameralucida drawing of a cell in late meiotic diakinesis. There were no chromosomalanomalies observed in any cells and chromosome pairing appeared to be completelynormal. FIG. 2. Meiotic chromosomes of Quickia aldabraensis.DISCUSSION AND TAXONOMY Prior to 1968, the genus was considered to be monotypic, containing only the typespecies, Quickia concisa (Morelet). Subsequently, 'Succinea spurca Gould, 'Succineabensoni Pfeiffer and a recently discovered Indian species (Q. calcuttensis Patterson)have also been found to belong to the genus Quickia. Q. aldabraensis is now thefifth species included in the genus. The Liberian Q. spurca differs in several respects QUICKIA ALDABRAENSIS SP. N. 181 ovotestis commonreproductivechannel cutting piece basal ccessory plate ovotestis ductreceptacula seminis fecundation pouchprostate gland ovotestis duct receptacula 3j-fecundation{( pouch 8 lumen vas deferens penis FIGS 3-11. 3, Shell drawing ; 4, Shell shape variability ; 5, Jaw ; 6, Body pigmentation ;7, Ventral view of the reproductive tract ; 8, Receptacula complex ; 9, Terminal genitalia ;10, Prostate gland ; n, Cross section of the penis. 182 C. M. PATTERSON from Q. concisa (Morelet) (Patterson, 1968, 1971). However, Q. spurca and Q.concisa appear to be more closely related to each other than Q. aldabraensis is toeither. For comparative purposes, Table I gives a short summary description ofvarious distinct morphological and antomical characters of these three species. 1 TABLE i Morphological characters of QuickiaQ. aldabraensis Q. spurca non-granulate granulate white charactershell sculptureshell colour radula formula head -foot colourprostate gland receptacula seminis penis position(ventral view) penial retractormuscle insertion amber to reddishamber Q. concisagranulatewhite 14-15:8-9:1:8-9: 12-14:9-10:1:9-10: ii : 10 : i : 10 : i14-15 12-14 reddish white ; branchingtubules ;relatively large highly subequalright of vaginaposteriorly translucent white white ; looselybranching tubulessmall slight subequal toequal usually left ofvagina posteriorly translucent whiteyellow ; small slightly subequal toequal left of vaginaanteriorly Connolly (1925) mentioned the collection of Succinea mascarenensis Nevill (More-let) Nevill from Grand Terre, Aldabra Island. Apparently this is the only occasionof assigning a name to the Aldabran succineid. Madge (1938) correctly establishedthat 'S.' nevellei Crosse and 'S.' mascarensis Nevill are synonyms of 'S.' concisa. Hefurther stated that 'S.' mascarenensis Nevill (Morelet) is probably also a synonym of'S.' concisa. However, the name mascarenensis used by Morelet (1882) does not havevalidity since his name was an alteration of Nevill's name mascarensis and, accordingto Madge (1938), was based on a mis-identification because the specimens were notcomparable to those of Nevill. Quickia aldabraensis is thus the second valid speciesof the genus known to inhabit Indian Ocean islands. The presence of mud ridges on shells of Quickia concisa has, in the past, been a keycharacter used in its identification. However, the presence of mud ridges on shellsof live field-collected Q. aldabraensis indicates they cannot be used as a valid species 1 Descriptions of Q. concisa are taken from Quick (1936) and Odhner (1950). I have verified the anteriorinsertion of the penial retractor muscle in Q. concisa, but the specimens were otherwise unsuitable for amore complete anatomical study. QUICKIA ALDABRAENSIS SP. N. 183 specific character of Q. concisa. From my examination of a specimen of Q. concisafrom Rodriguez and published results of Quick's (1936) observations of Q. concisafrom various localities, the most reliable species specific character for Q. concisa is theanterior insertion of a short penial retractor muscle. Q. concisa appears to be theonly succineid species, studied to date, with such a condition. Based on presentinformation, the accompanying key indicates the most reliable characters for identi-fication of currently recognized Quickia species. All species of Quickia are distinct, although Q. bensoni from India and Q. spurcafrom Liberia, while being the most widely separated geographically, appear to be themost closely related (morphologically) of the species. There is an indication thatQ. concisa is rather distantly related to Q. spurca and Q. bensoni. Q. calcuttensis andQ. aldabraensis have evolved to be most different from the other species. It isdifficult at this time to ascertain the relationship between the latter two species. Quickia has the highest chromosome number known in the family Succineidae.Data for the Subclass Euthyneura indicate that generally higher chromosomenumbers are associated with snail taxa which are considered morphologically moreadvanced by systematists (see Burch, 1965 ; Patterson, 1969). Odhner (1950)stated that 'Quickia and Indosuccinea evidently both represent a more primitivestage of the male genital development than exists in the other species of the Catinel-linae and indeed the most archaic type of male organ in any of the Succineidaehitherto examined'. One is now forced to consider whether simplification of theterminal reproductive system is instead a morphologically more advanced stage ofevolutionary development which was accompanied by an increase in chromosomenumber derived through aneuploidy over a long period of time. Indosuccinea has amore simplified terminal male genital development but a lower chromosome number(n = 24) than Quickia. Indosuccinea could have experienced an aneuploid reductionof one (or more) bivalent (s) from a higher chromosome number or evolved in its owndirection from a predecessor with a lower chromosome number. The question of theevolutionary relationship of the various succineid genera is unsettled, especially inthe Catinellinae (see Patterson, 1972). KEY TO QUICKIA SPECIES i a Penial apron present. ........ Subgenus Burchella Quickia calcuttensis Patterson ib Penial apron absent ....... Subgenus Quickia s. s. . 2 2a Shell amber and without granulations . ..... Q. aldabraensis sp. n. 2b Shell white and granulate ........... 3 3a Penial retractor muscle inserts anteriorly . . . . . Q. concisa (Morelet) 3b Penial retractor muscle inserts posteriorly ........ 4 4a Tubules of prostrate gland very loosely organized Q. spurca (Gould) 4b Tubules of prostrate gland more compactly organized Q. bensoni (Pfeiffer) DISTRIBUTION AND BIOLOGY Fig. 12 shows the known distribution of Quickia. Q. concisa appears to have thebroadest distribution. However, anatomical validation of the species from manyareas is necessary to substantiate such a vast distribution determined largely from i8 4 C. M. PATTERSON identifications often based only on shell characters. There are no records of theoccurrence of Q. aldabraensis on any island in the Indian Ocean other than AldabraAtoll. It would be interesting to know if additional Quickia species have evolved inother island groups. FIG. 12. Distribution map of Quickia. *6, Angola "7, Zanzibar *i, Sierra Leone 2, Liberia *3, Cameroon *8, Seychelles *4, Gabon 9, Aldabra *5, San Thome *io, La Reunion * Denotes localities for Quickia concisa. *n, Mauritius*I2, Rodriquez 13, Tamilnadu State 14, Tamilnadu State 15, Calcutta Most succineids occupy relatively damp or humid habitats, some near sources ofpermanent fresh water. Two species (Lithotis rupicola Blandford and Succineabernardii Recluz) are actually found in running water of falls (Patterson, 1973^.Some tropical succineids are arboreal. Quickia is interesting because some of itsspecies are adapted to areas devoid of permanent standing or running water. Thesespecies depend only on seasonal rains for moisture. Usually the snails are foundon rocks or rock walls or in crevices where they seek shelter in hot, dry weather.Apparently they are able to aestivate for long periods of time (one year or more) inthe absence of monsoon rains. Q. calcuttensis from Calcutta and two other popula-tions of Quickia located in Tamilnadu (Madras), India do inhabit moist areas adjacentto sources of permanent fresh water. Because of its adaptability, Quickia has beenable to colonize areas which would seem to be unsuitable habitats for succineids.Further zoogeographical studies may reveal a much wider distribution of Quickia.It would be particularly interesting to know if Quickia occurs in Australia or if theAustralian succineids living in dry habitats (i.e. Arborcinia] are related to Quickia. QUICKIA ALDABRAENSIS SP. N. 185 ACKNOWLEDGEMENTS I am indebted to The Royal Society for assistance and cooperation. Thanks aredue to John Taylor and C. A. Wright of the British Museum (Natural History) forcollecting and sending the specimens of Q. aldabraensis to me. I would also liketo express my appreciation to John Peake also of the British Museum (NaturalHistory) and J. B. Burch of the Museum of Zoology, University of Michigan forcritically reading the manuscript and for their many kindnesses during the study. SUMMARY (1) A morphological-anatomical description of Quickia aldabraensis, a new speciesfrom Aldabra Atoll, is presented. (2) The chromosome number of Q. aldabraensis is n = 25, the same number charac-teristic of other Quickia species. (3) Quickia occurs in Africa, on some Indian Ocean islands and in India. (4) Based on morphological studies, Quickia species do not seem to be particularlyclosely related to each other. (5) Members of the genus Quickia occupy a variety of habitats including thosewhich are extremely dry while other species are found near areas of permanentfresh water. REFERENCES BURCH, J. B. 1965. Chromosome numbers and systematics in euthyneuran snails. Proc. first Europ. malacol. Congr., 1962, pp. 215-241.CONNOLLY, M. 1925. Notes on a collection of non-marine Mollusca from the islands of the Indian Ocean. /. Conchol. 17 (9) : 257-269.MADGE, E. H. 1938. Notes on some non-marine Mollusca of Mauritius with descriptions of four new species. Mauritius Inst. Bull. 1 (3) : 15-29.MORELET, A. 1882. Observations critiques sur le memoire de M. E. V. Martens, intitule : Mollusques des Mascareignes et des Sechelles. /. Conchyliol. 30 : 85-106.ODHNER, N. H. 1950. Succineid studies : Genera and species of subfamily Catinellinae nov. Proc. malacol. Soc. London, 28 (5) : 200-210.PATTERSON, C. M. 1968. The reproductive anatomy and chromosome number of Quickia spurca (Gould) (Stylommatophora : Heterurethra : Succineidae). Malacol. Rev., 1: 1-13. 1969. Chromosomes of molluscs. Proc. Symp. Moll., II, Mar. biol. Assoc. India, 1969 :635-686. 1970. Morphological and cytological studies of the succineid genus Quickia from India.Malacol. Rev. 3 (i) : 25-36. - 1971. Taxonomic studies of the land snail family Succineidae. Malacol. Rev. 4 (i): 131-202. 1972. The succineid genus Quickia. [Abstract.] Malacol. Rev. 5 (i) : 17. i973a. Generic and specific characters in the land snail family Succineidae. [Abstract.]Malacol. Rev. 6 (i) : 54-56. I973b. Parallel evolution of shell characters in succineids inhabiting waterfalls. [Abstract.] Bull. Amer. malacol. Union, 38 : 28.QUICK, H. E. 1936. The anatomy of some African Succineae, and of Succinea hungarica Hazay and S. australis Ferussac for comparison. Ann. Natal Mus. 8 (i) : 19-45, pis. 1-4. C. M. PATTERSON Museum of Zoology THE UNIVERSITY OF MICHIGAN ANN ARBOR, MICHIGAN 48104 U.S.A. 10 PLATE i Holotype of Quickia aldabraensis a, Holotype (BMNH No. 1973103) collected from the type locality (1968). b, Para type (BMNH No. 1973104) laboratory reared. Measurement line in mm. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE i NOTES ON SOME ECHINODERMS FROMMARION ISLAND By F. W. E. ROWE & A. M. CLARK THE specimens which form the basis of the present report were collected at MarionIsland in the Southern Ocean (approx. 47 S, 37 E) by Mr A. F. de Villiers duringthe 1972/73 South African Expedition sponsored by the Department of Transport atPretoria and sent to the British Museum for identification. Marion and the adjacent Prince Edward Islands were the subject of anotherSouth African expedition in 1965/66, the holothurians from which were reportedon by Pawson and the other echinoderms by Bernasconi in 1971. Both theseauthors remark on the zoogeographical affinities of the fauna with that of Kerguelento the east and the sub- Antarctic Falkland-Magellan area further away to the west. The single species of holothurian taken is discussed here by F. W. E. Rowe andthe remaining echinoderms by A. M. Clark. HOLOTHURIOIDEAPseudocnus laevigatus (Verrill) Pentactella laevigata Verrill, 1876 : 68. Cucumaria serrata var. marionensis Theel, 1886 : 74-75, pi. 4, fig. 3. Cucumaria laevigata : Ekman, 1927 : 396-403, fig. 15. Pseudocnus laevigatus : Pawson, 1968 : 145, figs 2-11 ; 1971 : 288-289. MATERIAL TVLT 7 and 28, under boulders at the sub-littoral fringe and at4 m ; 4 specimens. LD u, undersides of stones at LWS; 7 specimens. Z 9, undersides of holdfasts of the bull kelp Durvillea antarctica at LWN ; 2specimens. Z 32, in a 'lithothamnion'-filled depression at LWS ; 6 specimens. The size of the specimens ranges from length : breadth 8 : 3 mm to 60 : 5 mm.The majority are strongly contracted. Their colour varies from white to light pink.The ten tentacles are more or less equal. The density of the spicules of the bodywall increases posteriorly. The spicules are more or less cone-shaped, ranginggenerally from 90 to 120 /zm x 50 to 70 /*m, though in one specimen (length :breadth 18 : 3 mm) the largest spicules measure 150 /u,m x 90 /am. The average sizeis no jum x 50 /nm. Unfortunately in many cases the spicules have been erodedby initial storage in formalin. However, the complete spicules compare closelywith those figured by Pawson (1968) from what he considers to be the type specimenof Pentactella laevigata Verrill from Kerguelen, though the present specimens are allmuch smaller. Similarly, direct comparison of the spicules of these Marion Islandspecimens with those from syntypes of Cucumaria serrata var. marionensis Theel in Bull. BY. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 i88 F. W. E. ROWE & A. M. CLARK the British Museum collections confirms that Ekman (1927) was correct to treatTheel's C. serrata with its varieties as conspecific with P. laevigata Verrill. Dissection of several specimens shows that hermaphrodite gonads are present inspecimens over 15 mm length (partly contracted), though the egg follicles areempty in the smaller individuals and I doubt whether even the larger specimens arefully mature. No brood pouches are present and only in the largest syntype ofC. serrata var. marionensis, at c. 40 x 9 mm contracted, could I find one. There are four polian vesicles in these specimens. Verrill (1876) noted three inhis specimen and Pawson (1968) found two in specimens from Macquarie Island.I have found two to four in the syntypes of C. serrata var. marionensis. OPHIUROIDEAAmphiura tomentosa Lyman Amphium tomentosa Lyman, 1879 : 23, pi. n, figs 299-301 ; 1882 : 132-133, pi. 29, figs 10-12.Nullamphiura marionis Bernasconi, 1968 : 56-58, 2 figs ; 1971 : 286, pi. 85, fig. 4, pi. 86,fig. 2. MATERIAL Z 25, underside of a stone in a pool connected to the sea by a tunnel;i very small specimen. Z 39, in detritus under boulders at LWS ; 2 specimens. Z 55 (pt), in holdfasts of the giant kelp Macrocystis pyrifera ; 3 specimens. Comparison of this material with the holotype of Amphiura tomentosa Lyman,collected at Kerguelen by the 'Challenger', shows no significant difference thatcannot be attributed to the larger size (d.d. 6-5 mm) of the type, the largest MarionIsland specimen at d.d. 3-5 mm being slightly larger than Bernasconi's holotype ofNullamphiura marionis. The discrepancies in the published descriptions of thetwo nominal species, such as the contiguity of the adoral shields (said to be touchingin A . tomentosa and separate in N. marionis - appearing abnormally so in the pairfigured by Bernasconi in 1968) are attributable to variation. Indeed Bernasconi(1971) notes that the adorals are 'rarely joined'. The distinctive widely separatedradial shields of larger specimens (d.d. 3 mm or more) agree with Bernasconi's andLyman 's descriptions (though in his figures they appear almost contiguous distally).There may also be naked patches of skin in some of the ventral interradii of thepresent Marion Island specimens as well as in the holotype of A. tomentosa. I thinkthat the lack of imbrication in the disc scales of the latter is due to the somewhatdistended condition of the disc. The shape of the oral shields is rather variablebut they never have such a large proximal angle as in Amphiura lymani (Studer)from the vicinity of South Georgia, which is otherwise rather similar, lacking tentaclescales, as Bernasconi notes. Her supposed difference that A. lymani has smallerdistal oral papillae than proximal (infradental) ones, whereas the reverse is thecase in the Marion Island species, is not supported by the present specimens or the'Discovery' material of A. lymani from South Georgia, of which Mortensen's figure(1936, fig. 14, p. 275) is misleading in showing the distal papillae as much smaller ECHINODERMS FROM MARION ISLAND 189 than their true size which I reckon approximately equals the size of the infradentalpapillae. In the Marion Island specimens the distal papillae are variable not onlyin size but also in shape, being either pointed or rounded at the tip. The dorsalarm plates have their distal edges flattened medially in these Marion Island speci-mens and can better be described as fan-shaped than rhombic, as Bernasconidescribes them. The ventral arm plates are pentagonal. As for the generic position of this species, in 1970 I rejected Nullamphiura Fellon the grounds that the number of tentacle scales unsupported by other charactersis inadequate for a generic distinction from Amphiura, being variable in severalspecies and resulting in artificial grouping of otherwise morphologically diversespecies (Clark, 1970). In fact, the holotype of Amphiura tomentosa does have afew pores showing a rudimentary scale, though these are quite lacking in the smallerMarion Island specimens. It is surprising that A. tomentosa has not been reported again from Kerguelendespite extensive collections by the French and the B.A.N.Z.A.R. Expedition. Ophiurolepis martensi (Studer) Ophioglypha martensi Studer, 1885 : 161, pi. 2, fig. 8.Ophiurolepis martensi : Mortensen, 1936 : 321-323, fig. 39. MATERIAL. Z 55 (pt), in holdfasts of the giant kelp Macrocystis pyrifera ; 14specimens. All these shallow-water specimens as well as 57 others taken by the 'Discovery'Investigations off Marion Island in 88-113 m consistently have the disc platingirregular. In contrast, out of a total of 168 specimens from South Georgia - thetype locality - no less than 109 or 65 % have a more or less regular rosette, as Morten-sen's figure (1936) shows ; also of 91 specimens from McMurdo Sound in the RossSea, 70% have regular discs. Possibly this difference is enough to justify a sub-specific distinction of the material from the vicinity of Marion Island but there is noobvious morphological difference between them and those specimens from otherlocalities which share their irregular disc plating. One of these specimens has a number of ? loricates in the grooves between thedisc plates. Ophiacantha vivipara Ljungman See : Mortensen* 1936 : 246-248, pi. 7, fig. 2. MATERIAL. Z 57, in the holdfasts of Macrocystis (rare) ; 5 adult specimens withseveral emerging and loose young. This species was evidently not taken by the 1965/66 expedition, though collectedby the 'Discovery' Investigations at 90 or more metres off Marion Island. igo F. W. E. ROWE & A. M. CLARK ASTEROIDEAAnasterias rupicola (Verrill)See : Bernasconi, 1971 : 285, pi. 85, figs i, 5. MATERIAL. TVLT 14, among boulders at 3-5 m ; 6 specimens.LD 10, in depressions and crevices at LWS ; 3 adults with 3 young.GT 4, on a vertical face at 3 m ; 2 specimens.Z 3, in a rock pool at LWS ; i specimen. Z 31, in the pool connected to the sea by a tunnel ; 2 specimens.The largest one of these specimens has R 58-65 mm and exceeds any other recordfor the species, although one of Bernasconi's had R 49 mm. REFERENCES BERNASCONI, I. 1968. Equinodermos de las Islas Marion y Principe Eduardo, con descricionde una nueva especie de Ofiuroideo. Physis, B. Aires 28 : 55-58, 2 figs., i pi. 1971. Echinodermata. In : Zinderen Bakker, E. M. van, Winterbottom, J. M. &Dyer, R. A. [Eds.] Marion and Prince Edward Islands. Cape Town. pp. 284-287, pis 85,86. CLARK, A. M. 1970. Notes on the family Amphiuridae. Bull. Br. Mus. nat. Hist. (Zool.) 19 : 1-81, ii figs. EKMAN, S. 1927. Holothurien. Dt. Sudpol Exped. 19 Zool. No. n : 361-419, 18 figs.LYMAN, T. 1879. Ophiuridae and Astrophytidae of the "Challenger" Expedition. 2. Bull. Mus. comp. Zool. Harv. 6 (2) : 17-83, 8 pis. 1882. Ophiuroidea. Rep. scient. Results Voy. "Challenger" (Zool.) 5 : 1-386, 46 pis.MORTENSEN, T. 1936. Echinoidea and Ophiuroidea. "Discovery" Rep. 12 : 199-348, 53 figs, 9 pis.PAWSON, D. L. 1968. Some holothurians from Macquarie Island. Trans. R. Soc. N.Z. (Zool.) 10 : 141-150, 13 figs.- 1971. Holothuroidea. In : Zinderen Bakker, E. M. van, Winterbottom, J. M. & Dyer, R. A. [Eds.] Marion and Prince Edward Islands. Cape Town. pp. 288-289, 2 fig 8 -STUDER, T. 1885. Die Seesterne Siid-Georgiens nach der Ausbeute der deutschen Polar-station in 1882 u. 1883. Jb. hamb. wiss. Anst. 11 : 143-166, 2 pis.THEEL, H. 1886. Report on the Holothurioidea dredged by H.M.S. "Challenger" during the years 1873-1876. Part 2. Rep. scient. Results Voy. "Challenger" (Zool.) 39 11-290, 1 6 pis.VERRILL, A. E. 1876. Echinoderms. In : Kidder, J. H. Contribution to the Natural History of Kerguelen Island. Washington, pp. 68-75. [Also in : Bull. U.S. natn. Mus. 3.] F. W. E. ROWE Department of Marine Invertebrates THE AUSTRALIAN MUSEUM 6-8 COLLEGE STREET SYDNEY, N.S.W. 2000 AUSTRALIA AILSA M. CLARK Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 560 A NEW SPECIES OF TILAPIA (PISCES,CICHLIDAE) IN THE ZAMBIAN ZAIRE SYSTEM By ETHELWYNN TREWAVAS & DONALD J. STEWART SYNOPSIS A new species of the cichlid genus Tilapia A. Smith, 1840, is described from the Luongo River,a tributary of the Luapula in northern Zambia. It is assigned to the subgenus Tilapia. Com-pared to its near relative, T. sparrmanii, the new species is more elongate, with higher modalnumber of vertebrae (28, cf. 27) and a slightly longer caudal peduncle and has relatively shorterpectoral fins, features suggesting adaptation to flowing waters. The new species also appearsto exhibit sexual dichromatism in the dorsal fin, an unusual feature in substrate-spawningTilapia. INTRODUCTION THE new Tilapia described herein was collected by Dr Eugene K. Balon* while hewas stationed at the Central Fisheries Research Institute (UNDP/FAO), Chilanga,Zambia, in 1970. Fish samples were taken in the Luongo River as part of DrBalon's stock and production assessment programme for Lake Mweru and itsdrainage area. Analysis of the Luongo fish collections is continuing ; a completetaxonomic and zoogeographic survey of the fish fauna will be published later. The Luongo River is an eastern head-water stream of the Zaire basin and entersthe Luapula River about 20 km upstream from Johnston Falls. On the Luongoabout 20 km upstream from its mouth Musonde Falls may have historically formeda barrier to upstream movement of some fish species. Recently a dam was built inthe vicinity of Musonde Falls, completely isolating upstream areas. The newTilapia was collected at two localities above Musonde Falls : (a) Nsenga (or Insenga)stream, a shallow, clear forest stream, 2 -6m wide with banks overgrown withbrush, (b) Luongo River mainstream, a sluggish, turbid stream 18-20 m wide,0-5-4 m deep, in open savannah with a deep, grassy ravine. ABBREVIATIONS AND MEASUREMENTS BMNH = British Museum (Natural History)ROM = Royal Ontario MuseumSL = Standard length The upper jaw is measured from the anterior point of the upper lip to the posteriorend of the maxilla, the lower from the anterior point of the lower lip to the posteriorend of the jaw, here the position of the retroarticular. The preorbital bone is measured from the middle of its orbital rim along a linecontinuing the radius of the eye at that point. * Present address : Department of Zoology, University of Guelph, Guelph, Ontario, Canada. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 192 E. TREWAVAS & D. J. STEWART The last ray of the dorsal and anal fins is counted as one if it is separate from thepenultimate at its base, even if it is smaller than the penultimate. A simple rayfused basally to the penultimate is not counted. Tilapia baloni n. sp. HOLOTYPE. ROM 28120 : male, 136 + 30 mm, Nsenga stream, tributary ofUpper Luongo. PARATYPES. ROM 28120 : 5 males, 86 + 23-5 to 114 + 28 mm, 6 females, 83 + 22to 106 + 25-5 mm an( i a juvenile of 50 + c, same field data as holotype. ROM 28071 : 3 males, 90 + 26-5 to 113 + 29 -5 mm from Luongo River aboveMusonde Falls. BMNH 1974.4.23.1 : male, 138 + 36-5 mm from Luongo River above MusondeFalls (formerly ROM 28071). BMNH 1974.4.23.2-4: female, 101 + 27 mm an d 2 males, 100 + 26 and 85-5 +22 mm from Nsenga stream (formerly ROM 28120). DESCRIPTION. Meristic characters from holotype and all paratypes, proportionsfrom holotype and 6 male and 5 female paratypes of 82-5-138 mm SL. Proportions as %SL. Depth of body 36-5-41-7 (over 40-5 in only 2) ; lengthof head 30-4-32-0 ; length of pectoral fin 23-0-29-0 (over 28 in only 2) ; length ofcaudal peduncle 13-3-17-2, 14 or more in all but i specimen (0-96-1-25 times itsdepth, less than i-o in only 2 specimens). Proportions as % length of head. Length of snout 33-9-38-4 ; diameter of eye24-6-29-0 ; depth of preorbital 18-0-21-6 ; interorbital width 33-0-36-6 ; lengthof upper jaw 29-5-34-6, of lower jaw 35-5-38-5. Maxillary not extending to below eye. Teeth in 3-5 rows in upper jaw, 3 or 4in lower, 40-50 in outer row of upper jaw. Outer teeth bicuspid with the maincusp obliquely or rectangularly truncate ; inner tricuspid with subequal cusps. Gill-rakers on first arch (1-2) + ! + (8-10, usually 8 or 9), short, those near thejoint often bluntly bifid. Microbranchiospines present on outer sides of 2nd, 3rdand 4th arches. Lower pharyngeal bone (Fig. 2) with a median length of 24-3-29-4% length ofhead, and width 31-4-36-3%, with short blade, 0-55-0-63 length of toothed area ;teeth slender, the posterior bicuspid, the anterior kukri-shaped. Scales on cheek in 3 horizontal rows ; in lateral line series 29 or 30, betweenorigin of dorsal and lateral line 3^ or (usually) 4 ; around caudal peduncle 16.Circuli granular, mostly in a roman pattern but occasionally a few gothic. Dorsal XIV 10 (1.2), XV 9 (f.i), XIV n (f.i), XV 10 (f.i6) or XV u (f.2) ; lastspine 14-5-17-8% SL, soft rays not greatly prolonged. Anal III 9, third spine I4-I5%SL. Pelvics not quite reaching vent. Caudal truncate with angular or roundedcorners, scaly only at the base and not densely. Vertebrae 27 (f.3), 28 (f.i7) or 29 (f.i). A NEW TILAPIA 193 10mm FIG. i. Tilapia baloni, holotype. Genital papilla of male conical or bluntly bifid, of female a short pigmented tubewith scalloped rim. Loose ovarian eggs with long diameter about i -9 mm in a fishof 100-5 mm. Intestine in a specimen of SL 95 mm a little over 3 times SL, this and the stomachcontaining fine dark debris including parts of vascular plants, sparse fragments offilamentous algae and some desmids. FIG. 2. Lower pharyngeal bone of Tilapia baloni, holotype. Colour (from colour photographs taken by Dr Balon from recently killed fishes).Snout, top of head and dorsum dark blue-green, nearly black in adult male ; flanksmore diffuse green with 9 or 10 dark vertical bars and 2 horizontal bands present orabsent (present in all preserved fish) ; scales of lower half of flanks and caudalpeduncle each with a dark spot either centrally or at base. In the larger malemain part of operculum and flanks around, below and behind pectoral crimson ;fainter indications of same colour in other males. 194 E - TREWAVAS & D. J. STEWART Spinous dorsal dusky with vague paler spots and a submarginal black band ;soft dorsal proximally with spots or a dark reticulum with light interstices, distallywith dark streaks between the rays ; a tilapia-mark on the dorsal fin in all specimens.In males dorsal lappets whitish with a pinkish tinge ; upper edge of soft dorsaland postero-dorsal corner of caudal the same colour. In females lappets dark ordusky, but soft dorsal and corner of caudal sometimes with narrow pale edge. Anal dusky, with or without a pale tip. Pelvics dusky, pectoral transparent.Caudal with horizontal dark streaks between the rays. T. baloni is so far known only from the Luongo River above Musonde Falls. Asearch through the British Museum and University of Michigan collections ofT. sparrmanii from the Luapula basin and the Bangweulu Region revealed nooverlooked specimens of the new species and none was encountered during thesurvey of the Kafue Flats in which one of us (D. J. S.) took part in 1969/70. Wehave both examined catches in the Mweru basin without finding it. Dr Balon informs us that there is topographical and geological evidence that theupper Luongo once drained to the Kalungwishi, an eastern tributary of LakeMweru. Some headwater streams of the Kalungwishi River rise in the swamps ofthe Kawambwe plateau, whose waters also feed the Luongo. The fish fauna of theupper Kalungwishi is not well known. (The 'undescribed species from the Lake Mweru area' related to T. sparrmaniimentioned by Thys (1968 : 369) was T. ruweti and not T. baloni.} AFFINITIES. T. baloni most resembles T. sparrmanii A. Smith, within whosegeneral area of distribution it is found, but T. baloni is a more attenuate fish with amore acute head. The different shape of the body is expressed in the lower ratioof depth to length and the shape of the caudal peduncle as well as the number ofvertebrae (Table i). There is also a higher modal number of scales, but since thisis true not only of the lateral line series but also the number between lateral line andorigin of dorsal fin and rows on the cheek it may not be related to the presence of anadditional metamere, the basic difference being the smaller size of the scales. Thepectoral fin is usually relatively shorter, a feature which with the slender formprobably indicates a stronger swimmer, with less use of the pectorals as paddles andbalancers and more use of the tail for propulsion. At the sizes of our examplessome individuals of T. sparrmanii have an additional row of teeth, but this is notconstant. The long intestine and its contents in the one specimen examined do notsuggest a sharp difference in feeding habits between this and T. sparrmanii. Duerre(1969) found a preponderance of periphyton in the diet of the latter in the upperZambezi, with organic debris an important supplement. The two species differ also in colour. In T. sparrmanii the red colour on theflanks takes the form of red edges to the otherwise green scales around and behindthe pectoral. In the biggest photographed (male) T. baloni it is a red flush involvingthe greater part of the operculum and the abdominal region at and below the levelof the pectoral nearly to the ventral surface. It is more metallic in appearance thanthe red of the flanks characteristic of most species assigned to subgenus Coptodon.In the latter and T. sparrmanii males and females are coloured alike in body and fins. A NEW TILAPIA 195 TABLE i Contrasts between T. baloni (22 specimens, i of 50 mm, 21 of 82-136 mm in SL) and T. sparrmanii (proportions in 16 specimens 82 -126 mm SL, meristic characters also in others) from the Luapula basin and Bangweulu region T. sparrmanii T. baloni Depth of body (% SL) 39-5-50-0 36-5-41-7 (only 2 < 41-5) (only 2 > 40-5) Length of pectoral (% SL) 24-0-32-5 23-0-29-0 23-0-27-9 f. 4 f.2o 28-0-29-0 2 2 29-1-32-5 10 o Length of caudal peduncle (% SL) 12-0-15-3 I 3'3-i7'2 divided by its depth 0-66-0-99 0-96-1-25 (only 2 < i-o) Scales : 1.1. series 26 f . 2 f. o 27 18 o 28 17 o 2 9 5 I5j 30 2 6D-l.l. 3 f.i 9 f. o 3i 24 6 4 5 15 4i i i cheek rows 2 f.37 f. o 3 7 22 Dorsal spines XIII f. 2 f. o XIV 38 3 XV 9 19 Total dorsal rays 23 f. i o 24 ii 3 25 3 17 26 7 2 Modal formula XIV n XV 10 Vertebrae 26 f. 2 f. o 27 22 3 28 8 17 29 o i T. ruweti (Poll & Thys van den Audenaerde) is also found on the Kawambwaplateau in waters including tributaries of the Luongo (specimens in Chilanga FisheriesResearch Station, examined by E. T., others collected by Dr Balon examined byD. J. S.). Its modal dorsal formula is XIV 10, nearly approached by XIV u, andthe modal number of vertebrae, as in T. sparrmanii, is 27. Its caudal fin is rounded.It has not been recorded at sizes greater than 80 mm SL and is more slender thanT. baloni, with an even shorter pectoral fin. Distinctive elements of the colourpattern are the tricolour band edging the dorsal fin and the light blue and purple-redspots on soft dorsal and caudal fins. No sexual dichromatism has been reported. T. sparrmanii is present below Musonde Falls, but has not been caught in theUpper Luongo. T. ruweti is sympatric with T. baloni in the Upper Luongo, justas elsewhere (e.g. in the Mweru lagoons) it is caught together with T. sparrmanii. I 9 6 E. TREWAVAS & D. J. STEWART The fact that to this extent T. baloni occupies the niche elsewhere belonging toT. sparrmanii suggests that it is either a vicariating species or a subspecies of thelatter. We have no evidence so far that the differences are less than specific, andthe probability that there are colour and size differences between the sexes inT, baloni is further support for its specific status. Although T. sparrmanii isreported from rivers, it is not a fish of the main streams, but occupies lagoons andbackwaters rich in vegetation. T. baloni is probably derived from T. sparrmaniior a sparrmanii-like common ancestor in response to life in a river-bed of steepergradient than that favourable to T. sparrmanii, with all that it means in terms oflocomotion and food. Sexual dichromatism in the dorsal fin of T. baloni is a significant difference betweenit and other species of Tilapia (as distinct from Sarolherodon) and it suggests thatthe sexes may differ more in their courtship roles than is usual in Tilapia. We place the new species in subgenus Tilapia, which includes the two specieswith which we have compared it (as well as T. guinasana in the opinion of one of us,E. T.) because of the bicuspid pharyngeal teeth (tricuspid in most species of subgenusCoptodon), low total numbers of dorsal rays and the relatively narrow preorbitalbone. But there are two features in which T. baloni resembles the species includedin Coptodon - the modal number of vertebrae, also 28 in Coptodon, and the presenceof 3 rows of scales on the cheek. The species, T. (Coptodon) rendalli Boulenger,that is found in the same geographical area as T. baloni and T. sparrmanii is the onemost unlike these species, having a modal 29 vertebrae, and is more advanced insome other ways, and this is another reason for relating T. baloni to the species ofT. (Tilapia). ACKNOWLEDGEMENTS We have pleasure in recording our gratitude to the following : Dr Eugene K.Balon, University of Guelph, Ontario, who collected the species, for putting hisnotes and colour slides at our disposal ; Drs W. B. Scott and E. J. Grossman,Royal Ontario Museum, Toronto, for making available material under their cura-torial care ; Dr R. M. Bailey, University of Michigan Museum of Zoology, AnnArbor, Michigan, for providing D. J. S. with facilities for work on the Luongo Rivercollections ; the authorities of the British Museum (Natural History) for the facilitiesenjoyed by E. T. including the making of radiographs (by M. McLellan) for vertebralcounts and use of the drawing by Sharon Chambers ; Mr Graham Bell-Cross forintroducing T. ruweti to E. T. in the Mweru basin in 1965. REFERENCES DUERRE, D. C. 1969. Report to the Government of Zambia on fishery development in the central Barotse floodplain. Second phase. FAO TA 2638.POLL, M. & THYS VAN DEN AUDENAERDE, D. 1965. Deux Cichlidae nouveaux du sud du bassin du Congo. Rev. Zool. Bot. afr. 72 : 322-333, text-figs i & 2. A NEW TILAPIA 197 SMITH, A. 1840. Illustrations of the Zoology of South Africa. Vol. 4: Pisces. Pis 1-31. 4 London.THYS VAN DEN AUDENAERDE, D. 1968. An annotated bibliography of Tilapia (Pisces, Cichlidae). Documn zool. Mus. r. Afr. centr. No. 14. xl+ 4o6pp. Dr ETHELWYNN TREWAVAS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD Dr DONALD J. STEWARTLaboratory of LimnologyUNIVERSITY OF WISCONSINMADISON, WISCONSINU.S.A. TWO NEW NEMATODES PARASITIC IN THEKIWI IN NEW ZEALAND By EILEEN A. HARRIS CONTENTS PageSYNOPSIS ........... 199 INTRODUCTION ........... 199 DESCRIPTION OF SPECIES . . . . . . . . . 199 Heterakis gracilicauda ......... 199 Cyrnea (Cyrnea) apterycis . . . . . . . .201 ACKNOWLEDGEMENTS ......... 205 REFERENCES ........... 205 SYNOPSIS Two new species of nematodes, Heterakis gracilicauda sp. nov. and Cyrnea (Cyrnea) apterycissp. nov., are described from the kiwi (Apteryx sp.) in New Zealand. INTRODUCTION THROUGH the kindness of Mr D. M. Rutherford of the Whangarei Animal HealthLaboratory, Whangarei, South Island, New Zealand, a small collection of nematodesobtained from a kiwi (Apteryx sp.) was received for study. So far as the writer isaware, the only record of nematodes occurring in Apteryx appears to be that ofChatin (1884, 1885), who gave a very brief and inadequate description of a form thathe called Ascaris apterycis. This species does not seem to have been met withagain, and it is therefore not surprising that the present material has proved to bevery interesting, consisting as it does of two new species which are described below. DESCRIPTION OF SPECIES Family HETERAKIDAE Railliet & Henry, 1914Heterakis gracilicauda sp. nov. (Figs i & 2) DESCRIPTION. This description is based upon specimens from the caecum ofApteryx sp. in South Island, New Zealand. The male measures 3-3-4-4 mm in length and 0-17-0-33 mm in maximumthickness and the female 5-6-4 mm and 0-19-0-3 mm, respectively. The headbears the usual three lips without interlabia, and no teeth have been made out(Fig. i). The diameter of the head taken just at the base of the lips is approxi-mately 42-49 /Am. There is a short pharynx measuring 30 /xm in length, and thisleads into an oesophagus terminating inwardly in a distinct bulb, which measures Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 JOO E. A. HARRIS 0-1 mm FIG. I. Heterakis gracilicauda sp. nov. : lateral view of head. 0-1mm 0-1 mm FIG. 2. Heterakis gracilicauda sp. nov. : (a) ventral view of male tail ;(b) tip of spicule. 0-14-0-16 mm in length and 0-11-0-13 mm m width. The total length of thepharynx together with the oesophagus, including the bulb, is 0-52-0-7 mm inthe male and 0-7-0-85 mm in the female. The nerve-ring is situated at approxi-mately 0-17 mm from the anterior end of the body. The excretory pore occurs justposteriorly to the nerve-ring at 0-26 mm from the anterior end. No cervicalpapillae have been observed. The caudal end of the male is long, slender and tapering, ending in a long, thinprocess. It measures 0-49-0-52 mm from the cloaca to the extreme tip of the tail.The caudal alae are narrow and little developed. The sucker is situated at 59-70 /u,m from the cloaca, and has a well-defined, chitinous rim. It measures 48-52 /Ltmin diameter from the outside of the sucker rim. There appear to be ten pairs of TWO NEW NEMATODES IN THE KIWI 201 caudal papillae arranged as shown in Fig. 2a. These are disposed as two pairs ofpedunculate papillae around the sucker, two pairs of sessile papillae lateral to thecloaca and two large pairs of pedunculate papillae lateral to these. The other fourpairs are distributed along the tail, and it is just posterior to the hindmost pair thatthe tail begins to narrow gradually. The spicules are short, alate and constantly subequal, measuring 0-24-0-29 mmfor the left and 0-16-0-18 mm for the right. They have a granulated appearance,are narrow proximally, but expanded slightly in their distal regions. Theleft spicule is constantly longer than the right and each spicule bears a smallprocess at the tip, as shown in Fig. 2b. The vulva is situated anteriorly to themiddle of the body, at approximately 2 mm from the anterior end. This leads intoa muscular vagina which runs posteriorly for a short distance before opening intothe uterus. The eggs measure 58-60 ^m x 30-35 p.m. MATERIAL. British Museum (Nat. Hist.) Reg. no. 1974 : 577-627 (syntypes). DISCUSSION. Inglis, Schmidt & Kuntz (1971) have divided the genus Heterakisinto species-groups, based upon the structure and the relative lengths of the spicules.The present species in having subequal alate spicules falls into their 'Heterakisgallinarum group. Of the species in this group, H. gracilicauda is most similar toH. isolonche Linstow, 1906, but differs from it in the length of the spicules, thoseof the new species being less than half the length of those reported for H. isolonche.The ratio of spicule-length : body-length in H. isolonche is I : 9 for the left spiculeand i : 7 for the right, in H. gracilicauda it is I : 20 for the right spicule and i : 15for the left. It appears that in all the male specimens of H. gracilicauda examinedthe left spicule is longer than the right. This is somewhat unusual, because in othermembers of the 'H. gallinarum' group, the relationship of the left spicule to the rightis reversed. It is interesting to note that in the collections of the British Museum (NaturalHistory) there are specimens of H. gallinarum and of H. isolonche from Australiaand of H. beramporia from New Guinea*. It would seem, therefore, that the 'H.gallinarum' species-group as denned by Inglis, Schmidt & Kuntz is quite widespreadin the Australasian region. Family SPIRURIDAE Oerley, 1885 Cyrnea (Cyrnea) apterycis sp. nov. (Figs 3, 4, 5 & 6 ; Pis i & 2) DESCRIPTION. This description is based upon specimens from the gizzard ofApteryx sp. in South Island, New Zealand. The body is cylindrical and the cuticle is provided with fine, transverse striations.The male measures 3-5-6-8 mm in length and 0-19-0-26 mm in maximum thickness,and the female 4-8-7-4 mm and 0-2 mm, respectively. The head has a diameter of30-40 /mi. The dorsal and ventral lips are well developed and rather deeplybilobed, each bearing a median process. Both the lobes of each lip carry twon 202 E. A. HARRIS papillae, a large one adjacent to a small one. The two lateral lips or 'pseudolips'carry at their bases the three pairs of teeth typical of the subgenus (Cyrnea) (Pisi & 2). A large papilla-like structure is visible on each of the lateral lips andposteriorly to this there is a small amphid. The oesophagus is, as usual, in two parts, a short anterior portion and a longerposterior portion, the total length being 1-1-1-8 mm in the male and 1-6-2-3 mmin the female. The nerve-ring is situated anteriorly to the junction of the twooesophageal regions, at about 0-20-0 -26 mm from the anterior end of the body.No cervical papillae have been observed. The features of the caudal end of the maleare exceedingly difficult to make out, as the tail of each male specimen is, as so oftenhappens in spiruroid nematodes, coiled in a tight, double spiral. Nevertheless, thecaudal alae are wide and well developed, and the tail is covered over a wide area ofits ventral surface with cuticular elevations, which are arranged in longitudinalrows, extending forward to a position just anteriorly to the beginning of the alae.There are eleven pairs of papillae, which are disposed in the manner usually foundin the genus Cyrnea, and shown in Fig. 3. There are four pairs of large, peduncula-ted, pre-anal papillae, situated just laterally and anteriorly to the cloaca, and two 0-1 mm FIG. 3. Cyrnea (Cyrnea) apterycis sp. nov. : ventral view of male tail. pairs posterior to the cloaca, whilst the remaining five pairs are very small and aredistributed at the tip of the tail. They are arranged in two rows, each of fourpapillae, with the fifth pair lying centrally between the two rows. The spiculesare unequal and dissimilar. The left spicule is long and slender, measuring 0-9-1-3 mm in length, and possesses a barbed tip (Fig. 4b). The right spicule is shorterand stouter, measuring only 0-18-0-25 mm in length and ends in a fine point.The proximal end is cup-shaped, as shown in Fig. 43.. In lateral view, thegubernaculum appears triangular in outline and measures 0-46-0-48 mm by 0-36-0-38 mm. TWO NEW NEMATODES IN THE KIWI 203 a 0-05mm 0-3mm FIG. 4. Cyrnea (Cyrnea) apterycis sp. nov. : (a) lateral view of male tail ;(b) tip of left spicule. The tail of the female is conical and in some specimens it bears a small, flattenedmucron at the tip (Fig. 5). The vulva is situated at 2-6-4-6 mm from the anteriorend. From the vulva, the long muscular vagina runs backwardly to enter the 0-3 mm FIG. 5. Cyrnea (Cyrnea) apterycis sp. nov. : ventral view of female tail. 204 E. A. HARRIS uterus, which bifurcates. One of the two uterine branches appears to run anteriorlyfor a short distance before bending posteriorly to run parallel with the other branchof the uterus (Fig. 6). The eggs measure 40-50 p.m by 24-28 | 0-3 mmFIG. 6. Cyrnea (Cyrnea) apterycis sp. nov. : vulval region of female. MATERIAL. British Museum (Nat. Hist.) Reg. no. 1974:628-667 (syntypes). DISCUSSION. Chabaud (1958) has divided the genus Cyrnea into two subgenera,Cyrnea (Cyrnea} and Cyrnea (Procyrnea}. The position of the teeth on the 'pseudo-lips', situated deep in the buccal capsule, places the present specimens in the sub-genus Cyrnea (Cyrnea}. In another work published later in the same year, Chabaud& Rousselot (1958) presented a differential key to the species of Cyrnea (Cyrnea),and this key divided the subgenus into ten groups. In this key, the present speci-mens fall into their group 9, which includes C. (C.) eurycerca Seurat, 1914. This isregarded by these authors as the only valid species known hitherto in this group, forboth C. (C.) graphophasiani Yamaguti, 1935, and C. (C.) euplocami Maplestone,1930, are considered to be synonymous with C. (C.) eurycerca. C. (C.) apterycisdiffers from the latter species in having shorter spicules, and in the more anteriorposition of the vulva. It may also be distinguished by the number and distributionof the caudal papillae on the male tail. In C. (C.} apterycis there are six pairs oflarge, pedunculated papillae plus five pairs of small, sessile papillae, whilst inC. (C.) eurycerca there are only five pairs of large papillae, one pair of adanal and fourpairs of small sessile papillae. The new species also possesses a barb-like tip to theleft spicule. Of the species described since Chabaud & Rousselot presented their key, onlyC. (C.) ochotensis Belogurov & Zueva, 1967, and C. (C.) singhi Ali, 1961, can bereferred to the C. (C.) eurycerca group. C. (C.} ochotensis, described from Falcosubbuteo in the U.S.S.R., differs from the present form in the greater length of itsspicules. C. (C.) singhi, from Centropus sinensis in India, can be distinguished fromC. (C.} apterycis by its much longer right spicule, by the number of cloacal papillaeand by the apparent possession of only two pairs of teeth on the 'pseudolips'. C. (C.} casuarii (Maplestone, 1932) has been placed by Chabaud and Rousselotin a separate group of its own. This species is mentioned here because it appearsto be the only member of the subgenus to have been described hitherto from anAustralian bird- Casuarius biarunculatus. This nematode differs from the form TWO NEW NEMATODES IN THE KIWI 205 described above in its large size, being 4-7 times as long as C. (C.) apterycis, and inthe number and arrangement of the caudal papillae of the male tail. It is interesting to note that according to Mawson (1968), although Cyrnea spp.are fairly common in Australian birds, all have been determined as belonging tothe subgenus Cyrnea (Procyrnea). She appears to have ignored C. (C.) casuarii,presumably because the host was resident in the Calcutta Zoo at the time of itsdeath, but it is more probable that the infestation was a natural one rather thanone acquired in India. ACKNOWLEDGEMENTS I wish to express my thanks to Mr S. Prudhoe for his constant help and encourage-ment during this study, and to Dr D. I. Gibson and the Electron Microscope Unitof the British Museum (Natural History) for the electron-micrographs. Thanks arealso due to Mr D. M. Rutherford of the Whangarei Animal Health Laboratory,New Zealand, for providing the material. REFERENCES ALT, S. M. 1961. On some new nematodes (Habronematinae) from birds in Hyderabad,India and the relationships of the genus Habronema. J. Helminth. 35 : 1-48. BELOGUROV, O. I. & ZUEVA, L. S. 1967. New and rare species of helminths from the birdsof the Far East. Zool. Zh. 46 : 999-1008. CHABAUD, A. G. 1958. Essai de classification des nematodes Habronematinae. AnnlsParasit. hum. comp. 33 : 445-508. & ROUSSELOT, R. 1958. Description d'un nematode Habroneme : Cyrnea (Cyrnea)antennifera n. sp. interessant par ses caracteres cephaliques. Bull. Soc. zool. Fr. 82 :420-429. CHATIN, M. J. 1884. Parasites de I'Apterix. C. r. Seanc. Soc. Biol. Ser. 8, 1 : 770-771. 1885. Helminthes de 1'ile Campbell et de la Nouvelle-Zelande. Bull. Soc. philomath.Paris Ser. 7, 9 : 36-43. INGLIS, W. G., SCHMIDT, G. D. & KUNTZ, R. E. 1971. Nematode parasites of Oceanica. XII. A review of Heterakis species, particularly from birds of Taiwan and Palawan. Rec. S. Aust. Mus. 16 : 1-14.MAPLESTONE, P. A. 1932. Parasitic nematodes obtained from animals dying in the Calcutta Zoological Gardens. Parts 9-11. Rec. Indian Mus. 34 : 229-261.MAWSON, P. M. 1968. Habronematinae (Nematoda : Spiruridae) from Australian birds. Parasitology 58 : 745-767. Mrs E. A. HARRIS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD PLATE iCyrnea (Cyrnea) apterycis - stereoscan micrographs of head (a) Lateral view. (b) Dorsal view. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE i PLATE 2 Cyrnea (Cyrnea) apterycis - stereoscan micrographs of head (a) En-face view. (b) Teeth at base of 'pseudolips'. Bull. Brit. Mus. mat. Hist. (Zool.) 28, 5 PLATE 2 DESCRIPTION OF PEMBATOXON INSULARE GEN. N., SP. N. FROM PEMBA ISLAND(MOLLUSCA PULMONATA, UROCYCLIDAE) By JACKIE VAN GOETHEM INTRODUCTION FROM previous work on slugs of the family Urocyclidae, it appears that the lack of astimulatory organ (diverticulum atrii, stylophorus or sarcobelum) in different speciescannot be the peremptory argument for uniting them in a single genus. Forexample, in a recent revision of the group (Van Goethem, in press) it was clearlydemonstrated that three species formerly united in a single genus, Atoxon Simroth1888, could be divided into two groups on the basis of a wide variety of morphologicalcharacters. These groups have been recognized as separate genera ; the firstincludes Atoxon fasciatum Verdcourt 1965, while the second is represented byAtoxon meridionals Forcart 1967 and Atoxon bruggeni Forcart 1967. The specimens described below certainly represent a new species and though itlacks a stimulatory organ, it cannot be included in any described genus (e.g. Atoxon).A new taxon is, therefore, described here. Abbreviations used in text : h. height ; M median tooth of radula ; Ig. length ; w. width. SYSTEMATICS PEMBATOXON gen. n.* TYPE SPECIES : Pembatoxon insulare sp. n. DIAGNOSIS : Tail long and strong, with visceral cavity extending to its posteriorend. Mantle posteriorly completely fused to the cephalopodium ; covering aninternal shell ; mantle aperture present, forming a small, medio-dorsal slit at theposterior end of the mantle. Shell unguiform, presenting a more or less bilateralsymmetry ; nucleus at the posterior end, medially ; protoconch oval. Jaw stronglyarcuate, with a median projection on the ventral margin. Radula with medianand laterals tricuspid ; marginals lacking the endocone, bicuspid (some mainmarginals unicuspid, the ectocone being reduced or lacking), very slender, muchlonger than the laterals ; mesocone of the marginals extending far beyond theposterior margin of the basal tooth plate. Pulmonary cavity little vascularized ;heart transverse ; aorta divided in two branches immediately after leaving theventricle. Reproductive system : epiphallus with a small bursa calcifera and along tubular caecum ; hermaphrodite gland situated well behind the albumengland, in the middle part of the tail ; stimulatory organ absent ; penis long, with * Pembatoxon results from the contraction of Pemba and Atoxon. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 208 J. VAN GOETHEM a spiral torsion ; penial sheath only near the proximal end separated from thepenial tube ; penial papilla simple, very small ; praeputium very short ; vaginaabsent ; epiphallus 2, very short. Spermatophore helicoid, without an angle ;presenting scaly spines over about its whole length ; top blunt ; filiform part veryshort. Alimentary tract forming 3 loops (intestine with one forwardly directedloop) and presenting a spiral torsion. Retractor muscles : the right upper tenta-cular retractor passes over the proximal end of the penis. Pembatoxon insulare sp. n.TYPE LOCALITY. Pemba Island, E. Africa.GEOGRAPHICAL DISTRIBUTION. Only from the type locality. MATERIAL. Pemba Island, E. Africa. Collected by C. Crossland, 1901. Materialin British Museum (Natural History) : holotype (Reg. No. 1909.5.10.52) and 5paratypes (Reg. No. 1909.5.10.53-57). MEASUREMENTS, (in mm) Holotype Paratype Paratype (1909.5.10.54) (1909-5-10.55) Body length 28-5 28-5 29-5 Body width at middle 5-5 6 6 Tail length 15 15-5 16-5 Mantle length 11-5 11-5 11-5 Mantle width at middle 9-5 10 10-5Distance between respiratory orifice and anterior mantle margin 6-5 6-5 6-5 Length of respiratory orifice 0-5 0-7 0-5Distance between respiratory orifice and posterior mantle margin 4-5 4-3 4-5 Height : maximum 6 6-5 6-5 at proximal end of tail 5 5-5 5-5 at posterior end of tail 2 2-5 2 Sole length 27 28-5 29 Sole width 2-8 3-5 3-5 Mid-area width of sole i-o 1-2 1-2 Length of mantle aperture 0-8 0-7 0-8 Width of mantle aperture 0-4 0-2 0-2Distance between mantle aperture and posterior mantle margin 0-7 0-6 0-8 DESCRIPTION.External characters (PI. i, fig. 1-2 ; Fig. lA-C) Small slug with an adult length of c. 28 mm (in alcohol). Genital opening situatedjust behind and beneath the right lower tentacle, above the right anterior angleof the sole and just above the right mouth lobe. Tail long and well developed ;its posterior end, at caudal gland level, slightly laterally compressed ; not keeled ;tapering backwards and ending in a well developed caudal horn (Fig. lA) ; with finelongitudinal grooves (irregular on the dorsal part) limiting small polygonal tuberclesand running to a fine irregular suprapedal groove. The latter being connected to the PEMBATOXON INSULARE GEN. N., SP. N. 209 FIGS 1-5. Pembatoxon insulare gen. n., sp. n. Fig. i. Holotype ; A : Caudal end of the tail, lateral view ; B : Posterior part of themantle, dorsal view ; C : Anterior part of the body, lateral view. Fig. 2. Shell, holotype ; A : Dorsal view ; B : Lateral view. Fig. 3. Shell, paratype I9O9-5-IO-53; A: Dorsal view ; B: Lateral view. Fig. 4. Jaw, holotype ; A : Frontal view ; B : Ventral view ; C : Semi-frontal view. Fig. 5. Jaw, paratype IQO9-5-IO-53, semi-frontal view. (go, genital opening ; It, lower tentacle ; m, mouth ; ma, mantle aperture ; ml, mouth lobe ;pg, pedal groove ; pgo, pedal gland opening ; pn, pneumostome ; s, shell ; spg, suprapedalgroove ; ut, upper tentacle) pedal groove by fine vertical grooves. Caudal aperture running to the posteriorend of the sole. Mantle finely and very distinctly granulated. Mantle aperturesmall, slit like (Fig. iB). Respiratory orifice, see Fig. iC. Sole rather narrow.Colour (in alcohol) : ground colour sandy ; on the tail with a brown lateral stripeand a very fine, yellowish, medio-dorsal line (holotype) ; the ventral half of the tail,vaguely brown mottled. Posterior half of mantle with a trace of a brown lateralstripe. Sole unicoloured. Paratypes (1905.5.10.53, 56-57) sandy, unicoloured. 2io J. VAN GOETHEM Shell (Figs 2-3) Ovate, rather convex, concentrically striate, very thin, whitish. Nucleus raised,median, posterior, white. Periostracum pale yellow.Holotype, Ig. 3-2 mm ; w. 2*4 mm ; h. 0-6 mm.Paratype (1909.5.10.53), Ig. 3-5 mm ; w. 2-6 mm ; h. 07 mm.Paratype (1909.5.10.55), Ig. 3-6 mm ; w. 2-8 mm ; h. 0-7 mm. Jaw (Figs 4-5) Strongly arcuate, with a rounded median projection on the ventral margin. Holotype, w. 1-6 mm ; Ig. 0-5 mm. Paratype (1909.5.10.53), w. 1-5 mm ; Ig. 0-5 mm. Paratype (1909.5.10.55), w. 17 mm ; Ig. 0-6 mm. Radula (Figs 6-8) Holotype (Fig. 6), formula: (5I + 3 + I4 + M + -) x 124. There is a ratherdistinct transition between laterals and marginals ; only 3 transitional teeth ;median tooth slightly asymmetrical. Median and laterals tricuspid. Marginalswithout endocones, very slender, with a high mesoconal supporting ridge ; mainmarginals (teeth 25-40) very long, with generally a reduced ectocone (posteriormargin of the basal tooth plate not reaching half the tooth length. Only c. 4irregular teeth, with a blunt mesocone and several ectocones ; generally i rudimentaltooth. Size : 3-1x1-3 mm ; c. 43 transversal rows/mm ; length of M : 32-35 /u,m. Paratype (1909.5.10.53) (Fig. 7), formula: (53 + 2 + J.6 + M+ -) x 130. Endo-cone lacking from the 2Oth-2ist tooth onwards. Most marginals between 20th and42nd without an ectocone (unicuspid). Only c. 4 irregular teeth ; generally Irudimental tooth. Size: 3-4 x 1-5 mm; c. 39 transversal rows/mm; length ofM: 34-37/zm. Paratype (1909.5.10.55) (Fig. 8), formula: (60 + 3 + J.6 + M+ -) x 127. Endo-cone lacking from the 22nd-23rd tooth onwards. Most marginals between 27th and43rd without an ectocone (unicuspid). Mesoconal supporting ridge of marginalsillustrated in Fig. 8. Only 4-5 irregular teeth ; generally i rudimental tooth.Size : 3-3 x 1-5 mm ; c. 39 transversal rows/mm ; length of M : 34-37 ^m. Pallial organs (Fig. 9) Pulmonary cavity little vascularized. Kidney quadrangular. Heart transverse.Aorta divided into two branches immediately after leaving the ventricle ; theanterior branch curving round the anterior loop of the intestine. Reproductive system (Figs 10-13) Holotype (Figs loA-C, n) : Hermaphrodite gland (ovotestis] yellowish white,long and slender, with a weak torsion, situated in the middle part of the tail. Herma-phrodite duct (ductus hermaphroditicus] rather long, convoluted proximally. Albu-men gland (glandula albuminalis] strongly developed, its middle part situated on alevel with the posterior mantle margin. Common duct (spermoviductus] short, PEMBATOXON INSULARE GEN. N., SP. N. 211 15 10 2 1 M 71 67 60 50 40 30 20 FIGS 6-9. Pembatoxon insulare gen. n., sp. n.Fig. 6. Radula, holotype.Fig. 7. Radula, paratype i9O9'5-io-53. Fig. 8. Two marginal teeth in almost lateral view, paratype I9O9-5-IO-55.Fig. 9. Pallial organs, ventral view, holotype. (ao, aorta ; au, auricle ; k, kidney ; lu, lung (roof of pulmonary cavity) ; me, columellarretractor muscle ; per, pericardium ; pn, pneumostome ; re, rectum ; rp, penial retractormuscle ; ur i, primary ureter ; ur 2, secondary ureter ; ve, ventricle) voluminous. Vas deferens short, passing under the spermathecal duct. Epiphalluswith a small, ovoid bursa calcifera (Fig. loB) at its distal end, and a tubular caecum(6-2 mm long) near its proximal end. Epiphallus i, 12-5 mm long, wound aroundthe penis and epiphallus 2 over its whole length. The latter very short (1-2 mm).Penis tubular, thick-walled, spirally torsive, clearly thicker than the epiphallus.Penial sheath (tunica penis) individualized at the proximal part of the penis (Fig.n). Prepuce (praeputium) very short, separated from the genital atrium by athick ring- wall. Penial papilla (glans penis) simple, very small (c. 0-25 mm long).Penial retractor muscle (retractor penis) very thin, long, arising from the left side ofthe diaphragm near the posterior margin of the kidney (Fig. 9), slightly adheredto the inner side of the penis windings and ending at about the distal end of thepenial sheath (Fig. loA, indicated by a small arrow ; Fig. n). Oviduct (oviductus) J. VAN GOETHEM 10 A 11 10A,C,12,13 PEMBATOXON INSULARE GEN. N., SP. N. 213 very short. Oviduct gland (glandula oviductus} ovoid, with 4-5 rounded, well-developed irregular folds. Spermatheca (bursa copulatrix) consisting of a long,slender sac (corpus bursae), and a short, wide duct (ductus bursae), internally withnumerous high, narrow folds ; containing 2 spermatophores. In situ, the end of thespermathecal sac situated ventrally, on the same level as the insertion of the colu-mellar retractor muscle. Atrium (atrium genitale) well developed, with numerousretractor muscles : at the issue of the oviduct gland, 3 long and strong ventralretractor bundles inserting at the left side of the foot-sole (on a level with the lung)and at the ventral side of the left body wall ; at the issue of the penis and betweenpenis and spermatheca several retractor muscles inserting at the right body wall.The ventral side of the atrium shows strong muscles (Fig. loC, stipplings). Paratype (1909.5.10.53) (Fig. I3A-B) : Albumen gland quadrangular with aterminal lobe on the left side. Vas deferens wider at its distal end. The firstsection of the epiphallus, 14-5 mm long. Caecum, 9-0 mm long. Penis: distalend of penial sheath (indicated by a small arrow, Fig. I3A) is coincident with theinsertion point of penial retractor muscle. Penial papilla very small, smaller thanin the holotype, very near to the penial opening. Spermatheca containing a sperma-tophore. Paratype (1909.5.10.55) (Fig. 12) : Vas deferens a little wider at its distal end.The first section of the epiphallus, 13-5 mm long. Caecum, 8-0 mm long. Penis :distal end of penial sheath and insertion point of penial retractor muscle indicatedby a small arrow on Fig. 12. Penial papilla not distinct ; however, a small lobecan possibly be interpreted as a penial papilla. Spermatophore (PI. i, figs 3-4 ; Fig. I4A-I) Holotype (Fig. I4A-I) : Helicoid, with 6 windings. Length: c. 16-5 mm.Apex blunt, rounded. Only the apical part of the spermatophore (for nearlyi mm) smooth. Outer side of the whorls with 4-5 irregular rows of forwardlydirected scaly spines, each spine about o-i mm long. Posteriorly the number ofrows decreases. Filiform section of the spermatophore, very short (c. 1-4 mm),with i sharp ridge (Fig. I4A, H) and i row of forwardly directed denticles (Fig.I4A, G, I), each denticle about 80-90 /mi long. Terminal opening present. A FIGS 10-13. Pembatoxon insulare gen. n., sp. n. Genitalia. Fig. 10. Holotype; A: General view ; B: Detail of bursa calcifera ; C: Ventralview of atrium and adjacent organs. Fig. ii. Holotype, longitudinal section of atrium and adjacent organs.Fig. 12. Paratype i9O9'5-io-53. Fig. 13. Paratype i909'5-io-53 : A : General view ; B : Detail proximal part of herma-phrodite duct, partly unrolled. (ag, atrium genitale (genital atrium) ; bca, bursa calcifera ; ca, caecum ; cb, corpus bursae(spermathecal sac) ; db, ductus bursae (spermathecal duct) ; dh, ductus hermaphroditicus(hermaphrodite duct) ; ep 1,2, epiphallus 1,2 ; gl, glans penis (penial papilla) ; gla, glandulaalbuminalis (albumen gland) ; glo, glandula oviductus (oviduct gland) ; od, oviductus (oviduct) ;ot, ovotestis (hermaphrodite gland) ; pe, penis ; pr, praeputium (prepuce) ; rp, retractor penis(penial retractor muscle) ; spo, spermoviductus (common duct) ; tp, tunica penis (penialsheath) ; vd, vas deferens). 214 J. VAN GOETHEM FIG. I4A-I. Pembatoxon insulare gen. n., sp. n. Spermatophore, holotype. A : General view ; B : Detail apical part, lateral view ; C :Detail apical part, dorsal view ; D : Transversal section at D in Fig. 14 A ; E : Detail ofspines at E in Fig. I4A ; F : Detail of spines at F in Fig. I4A ; G : Detail of denticles at Gin Fig. I4A ; H : Transversal section at H in Fig. I4A ; I : Detail terminal end. second spermatophore, with the median area partially digested, measures c. 16 mmin length (the filiform part being c. 1-4 mm). Paratype (1909.5.10.53) (PI. i, figs 3-4) : Length : c. 16 mm (the filiform partmeasuring c. i -5 mm) . Visceral organs Visceral cavity extending to the posterior end of the tail. Alimentary tractforming 3 loops (intestine with one forwardly directed loop), presenting together PEMBATOXON INSULARE GEN. N., SP. N. 215 nearly a complete circumvolution. Posterior tip of the hermaphrodite gland ex-tending behind the first loop of the alimentary tract. Anterior tip of the digestivegland extending to the inner side of the forwardly directed intestinal loop. Rectumpassing above the basal part of the columellar retractor muscle (Fig. 9, ventralview). Retractor muscles The right upper tentacular retractor passes over the proximal end of the penis.Retractor muscles of the genitalia, see above. DISTINGUISHING FEATURES OF THE GENUS AND SPECIES Pembatoxon gen. n. resembles the genus Atoxon Simroth, 1888, in the absence of astimulatory organ, but differs from that genus in : 1. the spermatophore having a very short filiform section ; a blunt, rounded apexand numerous forwardly directed spines over almost the whole length of thefusiform part ; 2. the penis having a short penial sheath and a very small penial papilla ; 3. the atrium having retractor muscles inserting at the left side of the foot-sole andat the ventral side of the left body wall ; 4. the absence of a vagina ; 5. the radula with relatively smaller number of laterals in comparison with thenumber of marginals. Pembatoxon insulare gen. n., sp. n. differs from Atoxon fasciatum Verdcourt, 1965(classified in a new genus, see J. Van Goethem in press) in : 1. the form of the spermatophore (see above) ; 2. the penis with a spiral torsion ; 3. the very short epiphallus 2 ; 4. the absence of a vagina ; 5. the aorta divided in two branches immediately after leaving the ventricle ; 6. the radula. with median and laterals not having a very long mesocone. Pembatoxon insulare gen. n., sp. n. differs from Atoxon meridionale Forcart, 1967,and Atoxon bruggeni Forcart, 1967 (both species classified in a new genus, see J.Van Goethem, 1973) in : 1. the form of the spermatophore (see above) ; 2. the penis with a very small, simple penial papilla ; 3. the absence of a vagina ; 4. the radula with most of the marginals bicuspid ; 5. the right upper tentacular retractor passing over the proximal end of the penis. ACKNOWLEDGEMENTS I am grateful to Mr J. F. Peake for reading the manuscript, to Mrs J. Van Melderenfor tracing over the drawings in ink, to the Trustees of the British Museum (Natural 216 J. VAN GOETHEM History) for the loan of the specimens and to the 'Komitee voor Elektronenmikro-skopie van de vaste stof, Katholieke Universiteit Leuven' for enabling me to makethe photographs of the spermatophore (PL i, figs 3-4). REFERENCES FORCART, L. 1967. Studies on the Veronicellidae, Aperidae and Urocyclidae (Mollusca) of Southern Africa. Ann. Natal Mus. 18 (3) : 505-570, figs 1-46.VAN GOETHEM, J. 1973. Atoxonoides aberrans gen. n., sp.n. du Malawi (Mollusca, Pulmonata, Urocyclidae). Bull. Inst. r. Sci. nat. Belg. 49 (8) : i-n, figs 1-14.VAN GOETHEM, J. in press. Revision systematique des Urocyclinae (Mollusca, Pulmonata, Urocyclidae). Annls Mus. r. Afr. cent. Sc. Zool., 215.VERDCOURT, B. 1965. Report on a further collection of East African slugs (Urocyclidae). Revue zool. hot. afr. 71 (3-4) : 274-296, figs 1-15. Dr JACKIE VAN GOETHEM INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE BRUXELLES PLATE i Pembatoxon insular e gen. n., sp. n. FIG. i. Holotype, lateral view, xa-i.FIG. 2. Holotype, dorsal view, x2-i.FIG. 3. Spermatophore i909'5-io-53 (Cambridge Stereoscan), fragment of third winding, XI25- FIG. 4. Spermatophore i909'5-io-53 (Cambridge Stereoscan), denticles on one row, justbefore the filiform part, x 440. Bull. Br. Mus. not. Hist. (Zool.) 28, 5 PLATE i A QUAGGA, EQUUS QUAGGA (MAMMALIA, EQUIDAE), AT UNIVERSITY COLLEGE, LONDON AND A NOTE ON A SUPPOSED QUAGGA IN THE CITY MUSEUM, BRISTOL By A. W. GENTRY CONTENTS Page INTRODUCTION . . . . . . . . . . 217 COMPARISONS ........... 218 DISCUSSION ........... 222 A SUPPOSED QUAGGA SKULL IN THE CITY MUSEUM, BRISTOL . . . 224 ACKNOWLEDGEMENTS ......... 225 REFERENCES ........... 225 SYNOPSIS A mounted equid skeleton in the Department of Zoology, University College, London, isidentified by its skull characters as Equus quagga, the extinct quagga of South Africa. Anequid skull in the City Museum, Bristol, is not a quagga although accessioned as one. INTRODUCTION IN the summer of 1972 I was asked to identify a mounted equid skeleton, Z$8i, inthe museum of the Department of Zoology, University College, London. Nowritten information existed about its history, but the number 41 had been writtenon the mandible a long time ago, and the late Professor D. M. S. Watson had onceinformed Dr K. A. Kermack that it was a quagga. Z58i lacks canines and is therefore a female. It is adult ; all the permanentcheek teeth are in wear and their occlusal surfaces much flattened. Infundibula,otherwise known as marks or central cavities, are still present in the upper andfirst lower incisors, so that the animal could have been 10-15 years old, were itsageing at all comparable with that of horses. Upper first premolars are present onboth sides. 7581 is not a horse, hemione or ass. Horse is ruled out by the relatively greatwidth of the skull across the zygomatic arch compared with that across the orbits,the rather squared outline of the occipital in rear view, insufficient narrowing of themore dorsal part of the premaxilla, and deep V-shaped internal sulci of the lowermolars. The zygomatic width and the normal instead of diminutive size of theoccipital rule out the hemione. It cannot be an ass because of the great zygomaticwidth, insufficient narrowing of the premaxilla, the lack of any marked backwarddeflection of the top of the occipital in side-view, and the curved rather than the Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 12 2i8 A. W. GENTRY flattened lateral walls between the styles of the upper molars. There is no doubtthat Z^8i is a zebra, of which four species are known : Grevy's zebra, Equus grevyi Oustalet, 1882, of parts of Somalia, eastern andsouthern Ethiopia and northern Kenya. Mountain zebra, E. zebra Linnaeus, 1758, of the coastal region of the Cape Provinceof South Africa and southern South West Africa. Burchell's zebra, E. burchelli Gray, 1824, with a range extending from the southernparts of the Sudan, Ethiopia and Somalia southwards to the Orange Free Stateand northernmost Cape Province. Quagga, E. quagga Gmelin, 1788, an incompletely striped zebra formerly foundin South Africa to the south of the Vaal River, in the Orange Free State andthe southern Cape Province. It was heavily slaughtered in the 18505 andi86os and became extinct in the 18705. Specimens in European zoos surviveduntil 1872 (London), 1875 (Berlin) and 12 August 1883 (Amsterdam). Anumber of specimens are available in museums, mostly as mounted skins,together with about a dozen skulls and five skeletons (Ridgeway, 1909 ; Hilz-heimer, 1912 ; Rzasnicki, 1949 ; Lundholm, 1951 ; Willoughby, 1966 ; Rau,1974)- COMPARISONS The skull of Z$8i was kindly lent to me for further study at the British Museum(Natural History). I established that it could not be a Grevy's zebra because itwas too small, its face was short, and the premaxilla did not narrow in its upper parts.I then made a detailed comparison with up to 55 adult skulls of Burchell's zebra,10 of mountain zebra and the museum's single example of a quagga skull, 1864.7.2.3,this last being from a complete skeleton of a male animal that lived in the LondonZoo from 1858 until 1864 (Sclater, 1901). Its tooth wear is similar to Z$8i. Theresults of the comparison will be discussed character by character. The positionsof all characters except the first are shown on a quagga skull in Fig. I and the skullof Z58i is illustrated in Plates 1-4. 1. The skull is smaller than in Equus zebra. Many E. burchelli approach it moreclosely, while the E. quagga skull is about the same size. It should be rememberedthat the quagga is a zoo specimen, that in the hemione and wild ass zoo specimenshave skulls with linear measurements up to 10 % smaller than their wild counterparts(Groves, 1966), that the same may be expected for zebras, and that Z58i may havebeen a zoo specimen itself. Thus the size of Z58i is appropriate for either burchelli orquagga. The back of M 3 is more posterior relative to the front of the orbit in Z58iand the quagga skull than in zebra, while burchelli is intermediate. Hence E. quaggaand Z58i are rather short faced, but this character appears to be linked with overallskull size in living equids, and both the large Grevy's zebra and large domestichorses have long faces. 2. The interorbital area of the f rentals shows a slight doming in burchelli but isflatter in zebra. The quagga skull and Z58i both resemble zebra. There is a A QUAGGA AT UNIVERSITY COLLEGE 219 D FIG. i. Diagrams of a quagga skull to show the positions of characters 2 to 16, discussed inthe text. A. Lateral view. B. Palatal view. C. Dorsal view of postorbital part of skull. D. Dorsal view of front of mandible. 220 A. W. GENTRY possibility of regional variation within burchelli in that the doming seems less markedin specimens from South West Africa. Furthermore Dr D. A. Hooijer has toldme that the quagga skull in the Leiden Museum shows a frontals' profile like burchelli.Nevertheless, Z$8i would still lie beyond the burchelli range for the character, sothat its affinities are with quagga or zebra. 3. The level of fusion of the two temporal lines to form a median sagittal line ismore anterior in burchelli than in zebra. In 36 out of 50 burchelli it was forward ofthe rearmost level of the zygomatic arch in dorsal view ; in the remainder it waslevel or behind. In 2 zebra it was anterior, and in 8 level or behind. The quaggaskull and Z$8i have the fusion in the more posterior position, thus being unlikemost burchelli but not unlike zebra. 4. The external auditory meatus is slanted upwards as it passes laterally in burchellibut emerges horizontally in zebra in which most individuals also have a larger meatalorifice. Both Z$8i and the quagga skull resemble burchelli. 5. The maxilla-nasal suture dips anteriorly as it approaches the top of thepremaxilla in both zebra and burchelli, but the phenomenon is more pronounced inzebra in which the top of the premaxilla suture is also more rounded so that a flangeof the nasal bone descends a short way behind the premaxilla. Unfortunately thesutures become obliterated in older individuals. They are no longer visible withcomplete certainty in the quagga skull, but in 7581 they resemble the burchellipattern. 6. The alisphenoid area on either side of the basisphenoid antero-medially to thearticulation for the lower jaw is more nearly horizontal in ventral view in zebra thanin burchelli. 7581 and the quagga skull are like burchelli. 7. E. zebra has a longer styloid process between the auditory bulla and the meatusthan does burchelli. Z$8i is like burchelli and the quagga like zebra. 8. In side-view the top of the occipital is drawn out posteriorly in zebra, but notnearly so much in Z$8i or the quagga. Among the sample of burchelli 17 resemblezebra, 17 resemble Z$8i and quagga, while 15 are intermediate and indeterminate.This character can also be seen in rear view as a relatively high occipital in zebraand a relatively low occipital in 7581 and the quagga. 9. The mastoid bone is wider in zebra than in Z$8i or the quagga skull. In burchelliit is narrow in 37 and wide in 18 skulls. The flange of the squamosal between themastoid and the external auditory meatus is also sometimes wide and sometimesslim, but may well be associated with the mastoid width. Of the 37 burchelli withnarrow mastoids only n also have a slender squamosal flange, but among the 18with wide mastoids n is again the number with a slender squamosal. A QUAGGA AT UNIVERSITY COLLEGE 221 10. In zebra, the top edge of the zygomatic arch is fairly straight in lateral viewand remains at more or less the same level. In the quagga and Z58i the top edge ismore strongly curved and ascends posteriorly. Of the burchelli sample 27 skulls arelike zebra and 25 like quagga. 11. On the upper molars the protocone or inner pillar is elongated anteriorly in40 out of 50 burchelli individuals but not in zebra. The quagga skull and Z58iboth resemble the majority of burchelli. 12. Cement is better developed around the exposed sides of the cheek teeth of zebrathan of burchelli. A crude assessment of 'more' or 'less' cement showed that inonly 9 among 48 burchelli skulls was there as much cement as in 7 out of 10 zebra.Both the quagga and Z58i skulls had 'more' cement and can be considered to bemore like zebra than burchelli. 13. A small P 1 is present in 5 out of 10 zebra, these being mostly the youngerindividuals. It is present on at least one side in 30 of 51 burchelli, is often largerthan in zebra, and there are indications that it is less frequent among the olderindividuals ; it also seems to occur less often in burchelli from South West Africa.Erz (1964) believes that the tooth is part of the deciduous dentition. When presentin burchelli it is often accompanied by a concavity in the front part of the medialwall of P 2 . A relatively large deciduous P 1 is present in Z58i, with only a slightconcavity on P 2 , but it is absent on the quagga skull. For this character Z58i thusresembles burchelli and differs from quagga, but the character must be very unim-portant for classification. 14. Infundibula, marks or central cavities, are present in the lower incisors ofzebra although less marked than those appearing in the uppers. This conditionagrees with horses (Grossman, 1953 : 400). In old individuals the infundibuladisappear from upper and lower incisors. However in burchelli infundibula werepresent in the lower incisors of only 6 among the 42 individuals in which theyoccurred in the upper incisors, and in 2 of the 6 they were present in I 1 s alone. TheLondon sample of burchelli again gives a hint of regional variation in that those fromSouth West Africa appear to have lower incisor infundibula more frequently. InZ58i and the quagga skull infundibula were present in the upper incisors and Ij.This intermediate state may best be counted as a difference from both zebra andburchelli. 15. The posterior edge of the mandibular symphysis in the median line is high andmore or less upright in burchelli. In zebra the top edge dips gently as it passesbackwards and so curves round at a lower level to become the posterior edge. Theheight of the posterior edge is thus less than in burchelli. This character is difficultto assess except in sectioned mandibles, but is real enough as can be seen fromLundholm's illustration (1951, fig. 2). Both the quagga skull and Zs8i resembleburchelli. A. W. GENTRY 16. The top surface of the mandibular symphysis is noticeably flat behind theincisors in both the quagga skull and Z$8i. It could be linked with the incisors'occlusal surface appearing to be very little upturned. This character contrasts withmost burchelli and zebra, even when the latter have very well worn teeth. It doesnot appear to be linked with a regression of bone around the incisor roots such ashas taken place in both the quagga skull and DISCUSSION It seems from the 16 characters considered above that Z$8i can be taken quitesafely as E. quagga. The characters can be summarized as in Table i, from which itis seen that Z$8i differs from zebra in 13 of the 16 characters, from burchelli in 5 ofthe 16, but from quagga in only 2 of 15 characters, one of the two being the scarcelysignificant presence of P l s. Accepting Z$8i as quagga, one can suggest that skullsof quagga are like burchelli but differ in flatter frontals, a shorter united sagittal lineon the cranial roof, more cement around the molars, better developed infundibula TABLE i Summary of cranial characters 7581 quagga 1. Size, smaller or larger smaller 2. Frontals, convex or flat flat 3. Level of fusion of temporal lines, anterior or posterior posterior 4. External auditory meatus, slanted or horizontal slanted 5. Nasal flange behind premaxilla, absent or present 6. Plane of alisphenoid, slanted or horizontal 7. Styloid process, short or long 8. Occiput, drawn out posteriorly or more upright 9. Mastoid, narrow or wide 10. Upper edge of zygomatic arch, curved and rising high or straight and low 11. Anterior part of protocone, elongated or short 12. Cement around cheek teeth, much or less 13. Deciduous P 1 , present or absent 14. Infundibula of lower incisors, present or absent 15. Shape of mandibular symphysis pos- teriorly, tall or low 16. Surface behind lower incisors, flat or concave flat V = agreement with Z58i ; x = difference from burchell' Vx zebra x V v \/ = character not visible. A QUAGGA AT UNIVERSITY COLLEGE 223 in the lower incisors and a flatter mandibular symphyseal surface behind the lowerincisors. The poor extent to which the occipital top is drawn out backwardly andthe strong curvature of the zygomatic arch are also characters which appear to bemore distinctive of quagga than of burchelli. These findings are in agreement withthe photographs of quagga skulls in Hilzheimer (1912, pi. 7, fig. 4, pi. 8, pi. 9, figs3-4). The anterior elongation of the protocone is poorly developed in the Berlinskull A26i7 of Hilzheimer, pi. 9, fig. 2, but this skull is not of a quagga (Antonius Three other skull characters have been discussed in the past in relation to quaggabut have not been used in the comparisons in this paper : diastema length, a pre-orbital cheek depression, and an upwardly bent profile of the anterior part of thenasals. Z$8i has a longer diastema than the London quagga skull, as seen in thetable of measurements, but the latter is unusual among quaggas, Hilzheimer(1912 : 100) giving much greater diastema lengths at 87, 97, 99 and 94 mm, as wellas a juvenile at 55 mm. Lydekker (1904 : 428) observed the preorbital depression inthe London quagga skull and took it as a differentiating character of the species,but similar vestigial depressions appear in some males of other zebras. There is nopreorbital depression in 581. The front part of the nasals is bent slightly upwardsin the London quagga skull but not in Z^Si. Such an outline is present in about athird of the burchelli sample, although perhaps less frequent in specimens fromSouth West Africa. Hilzheimer (1912 : 93) attributed the strength of this characterin the London quagga skull to its having come from an animal which had been captivefor a long period. EQUUS BURCHELLI" ZEBRA" QUAGGA FIG. 2. Map of southern Africa to show the approximate original distributions of three zebraspecies. The original distributions of E. quagga and E. zebra are very uncertain, and theiroverlap may have been less than indicated here. Political boundaries are shown for guidance. 224 A. W. GENTRY No known skulls of the extinct southernmost race of Burchell's zebra, E. burchelliburchelli, were available for use in the comparisons and it might be alleged that theirskulls could have resembled quagga skulls more closely than do those of otherburchelli races. As far as skin characters go, it is known that southern populationsof burchelli tended to lose the posterior striping, and the beginnings of this trend canbe seen even in surviving populations of E. burchelli antiquorum, the next southern-most race to burchelli burchelli. However, this resemblance to quagga may be moreapparent than real ; quagga had a brown body colour posteriorly, its posteriorstriping assumed a pattern unlike burchelli as can be seen on mounted skins in Tringand Vienna (Rau, 1974, figs 19, 21), it had no lighter 'shadow' stripes between themore posterior black stripes as seen in southern burchelli, and its dark stripes werewider relative to the white ones than in other zebras (also see Cabrera, 1936 : 91-2).It seems likely that both zebra and quagga were originally zebras of the more aridcountry lying to the south-west of the range of burchelli (Fig. 2), and that there is noneed to suppose that skulls of burchelli burchelli would have approached those ofquagga in their morphology. A SUPPOSED QUAGGA SKULL IN THE CITY MUSEUM, BRISTOL A male equid skull, Aa 3294, in the City Museum, Bristol, was accessioned inthe latter half of 1927 and thought to be of a quagga. A pencilled note in themuseum register reads : '? S. H. Swayne, 28 June 1872'. I was able to see this skull on 24 January 1974, and it did not appear to be aquagga or any other species of zebra. It showed the following differences fromzebras : overall size is too small ; skull width across the orbits exceeds that across the back of the zygomatic arch ; premaxilla is narrow in side-view (even narrower than in Grevy's zebra), andnarrows still more as it rises towards the nasal ; occiput is rather small ; braincase roof is too strongly curved in profile. There are additional differences between the Bristol skull and the quagga skullin the British Museum (Natural History) : the exposed part of the mastoid is too irregular in shape ; vertical part of the rear median wall of the mandibular symphysis is too low ; upper edge of zygomatic arch is insufficiently curved in side view ; too little cement around the side walls and the cheek teeth ; upper surface of the mandibular symphysis behind the incisors is more concave. It is not necessary in this paper to provide an alternative identity for Aa 3294.However, its size and morphology agree well with two skulls in the British Museum(Natural History) of a Dartmoor pony and a Shetland pony. The rather smallcanines agree well with the Shetland pony, Osteology register 1952.4.1.3, and raisethe possibility that Aa 3294 comes from a castrated domestic animal. The irregu-larly shaped mastoid is commoner in horses and ponies than in other equids. A QUAGGA AT UNIVERSITY COLLEGE 225 TABLE 2 Measurements taken on the skulls discussed in this paper (mm) Equus quagga 2581 Aa 3294 1864.7.2.3 Skull length, top of occipital crest to front premaxilla 494 482 431 Skull length, anterior edge foramen magnum to front premaxilla 446 442 398 Top of occipital crest to back of orbit 186 181 - Back of orbit to front premaxilla 349 335 - Skull width across posterior orbital rims 182 179 183 Skull width across zygomatic arch 174 178 177 Median length of nasals 194 171 Nasals breadth above infraorbital foramen 50-2 49 Occipital condyle to back M 3 at occlusal surface 209 208 174Back M 3 at occlusal surface to front pre-maxilla 274 273 254Diastema length, I 3 to P 1/2 (right side) 73-5 92 65Occlusal length, M 1 to M 3 73 69-8 71Occlusal length, P 1/2 to P 4 88-2 87-6 89 ACKNOWLEDGEMENTS I thank Dr Pamela Robinson for allowing me to study the University Collegequagga and Mrs R. M. Down for help in doing so. I thank Mr P. F. Bird for assis-tance in the City Museum, Bristol. Dr G. B. Corbet, Dr J. Jewell, Dr Q. B. Hendey and Madame V. Eisenmann readthe paper and offered helpful suggestions. REFERENCES ANTONIUS, O. 1951- Die Tigerpferde ; die Zebras. Monographien der Wildsdugetiere 11 : 1-124.CABRERA, A. 1936. Subspecific and individual variation in the Burchell zebras. /. Mammal. 17 : 89-112.ERZ, W. 1964. Tooth eruption and replacement in Burchell's zebra, Equus burchelli Gray 1825. Arnoldia (Rhodesia) 1, 22 : 1-8.GROSSMAN, J. D. 1953. Sissoris Anatomy of Domestic Animals, 4th edn. Philadelphia : Saunders. 972 pp.GROVES, C. P. 1966. Skull-changes due to captivity in certain Equidae. Z. Sdugetierk. 31 : 44-46.HILZHEIMER, M. 1912. Die in Deutschland aufbewahrten Reste des Quaggas. Abh. senckenb. naturforsch. Ges. 31 : 83-105, pis 4-9.LUNDHOLM, B. 1951. A skull of the true quagga (Equus quagga) in the collection of the Transvaal Museum. 5. Afr. J. Sci. 47 : 307-312.LYDEKKER, R. 1904. Note on the skull and markings of the quagga. Proc. zool. Soc. Lond. (i) : 426-431.RAU, R. E. 1974. Revised list of the preserved material of the extinct Cape Colony quagga, Equus quagga quagga (Gmelin). Ann. S. Afr. Mus. 65 : 41-87, 25 figs.RIDGEWAY, W. 1909. Contributions to the study of the Equidae ; ii. On hitherto unrecorded specimens of Equus quagga. Proc. zool. Soc. Lond. (2) : 563-586. 13 226 A. W. GENTRY RZASNICKI, A. 1949. Complete list of the specimens of skeletons and skins of Equus quaggaquagga (GM.) preserved in the museums of the whole world in 1939. Annls Mus. zool.polon. 14, 5 : 69-73. SCLATER, P. L. 1901. List of the specimens of the quagga that have lived in the Society'smenagerie. Proc. zool. Soc. Lond. (i) : 165-166. WILLOUGHBY, D. P. 1966. The vanished quagga. Nat. Hist. N.Y. 75, 2 : 60-63. A. W. GENTRY Department of Palaeontology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD PLATE i Dorsal view of skull of Equus quagga, University College Z$8i. The scale inthis and succeeding plates is marked in centimetres. Bull. BY. Mus. nat. Hist. (Zool.) 28, 5 PLATE i PLATE 2 Lateral view of same skull. The old number '41' is written on the mandiblejust above the number '581'. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE 2 PLATE 3Palatal view of same skull. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE 3 PLATE 4 A. Occipital view of same skull. B. Occlusal view of lower dentition of same skull. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE 4 A NEW ANGELFISH OF THE GENUS CENTROPYGE (TELEOSTEI: POMACANTHIDAE) FROM ASCENSION ISLAND By ROGER LUBBOCK & RICHARD D. SANKEY SYNOPSIS A new species of Centropyge is described from Ascension Island ; it is the second pomacanthidto be recorded from Ascension, and the third Centropyge to be recorded from the tropicalAtlantic ; it appears to be derived from tropical western Atlantic forms. INTRODUCTION To DATE only one species of angelfish (family Pomacanthidae) has been recordedfrom Ascension Island in the south Atlantic, namely the widespread Pomacanthusparu (Bloch), known otherwise only from the tropical western Atlantic (see 'Remarks'below) ; specimens of P. paru from Ascension were recorded by Fowler (1919, 1936)and Fraser-Brunner (1933). We examined living juvenile and subadult specimensof P. paru from Ascension and various West Indian localities, and on coloration andsuperficial morphology, they clearly appear to be conspecific. We now report asecond species of angelfish from Ascension, belonging to the genus Centropyge.Two species of Centropyge have previously been recorded from the tropical westernAtlantic, C. argi Woods & Kanazawa, 1951 and C. aurantonotus Burgess, 1974 ; thepresent species is new to science, and is the third Atlantic species of Centropyge.Type specimens are deposited at the British Museum (Natural History) (BMNH)and the United States National Museum (USNM). Centropyge resplendens n. sp. Resplendent angelfish(Fig. i, Plate i) DESCRIPTION. Based on three fishes, 29-0-35 -3 mm SL (standard length),from Ascension Island. Measurements are those of the holotype, followed inparentheses by those of the larger and smaller paratypes respectively where thesediffer from the holotype. Dorsal fin rays XIV 16 (last ray divided to base) ; anal fin rays III 17 (last raydivided to near base in holotype) ; pelvic fin rays I 5 ; pectoral fin rays 16, first rayrather closely applied to second ray, all rays branched except for upper 2 and lower2 rays ; 17 principal caudal fin rays, branched with the exception of the uppermostand lowermost rays, which do not quite reach posterior margin ; three smallsupplementary rays above and below principal caudal fin rays ; pored lateral linescales 34 (35, 34) ; oblique scale rows from upper end of gill opening to caudal base Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 228 R. LUBBOCK & R. D. SANKEY 5 mm FIG. i. Holotype of Centropyge resplendens n. sp. 35-3 mm SL. Ascension Island. Drawing by G. Howes. 44 (43, 45) ; scales above lateral line to origin of dorsal fin 6 (7, 6) ; scales belowlateral line to origin of anal fin 14 (14, 16) ; circumpeduncular scales 16 ; gill-rakerson first arch 5 or 6 + 1 + 15 or 16 = 22 > the first very small ; branchiostegal rays 6 ;upper teeth in outer series 53 (51, 55) ; lower teeth in outer series 46 (54, 47). The following measurements are presented as percentages of the SL. Totallength 123-8 (123-8, 122-9) '> nea d length (measured from front of upper lip to endof opercular membrane) 30-0 (30-7, 31-3) ; snout length 9-9 (9-0, 10-4) ; orbitdiameter 11-9 (12-1, 13-3) ; bony interorbital width 7-1 (7-6, 6-4) ; predorsallength 36-3 (37-2, 40-2) ; depth at first dorsal spine 43-1 (44-1, 47-4) ; body widthbehind gill opening 18-7 (16-9, 16-5) ; least depth of caudal peduncle n-6 (11-4,1 2-0) ; length of caudal peduncle (measured horizontally from rear base of dorsalfin to caudal base) 7-6 (8-6, 7-2) ; length of longest pectoral fin ray 22-1 (24-1, 24-1) ;length of longest pelvic fin ray 26-1 (28-6, 28-9) ; dorsal fin base length 65-7 (65-5,64-3) ; lengths of first, second, third and fourth dorsal spines 4-5 (6-6, 5-6), 8-8(11-4, 11-2), 11-9 (12-4, 16-9) and 11-3 (12-4, 14-5) respectively; anal fin baselength 35-4 (35-9, 36-1) ; lengths of first, second and third anal spines - (9-7, 11-2),- (12-1, 14-1), and - (13-4, 14-5) respectively (anal spines clearly deformed in holo-type, not measured). Body deep and relatively compressed. Mouth small, terminal, the gape horizon-tal ; maxilla reaching slightly posterior to vertical from anterior nostril ; upper lipapproximately as broad (vertically at front) as lower, upper lip height about a thirdof orbit diameter. Teeth slender and elongate, close-set, flexible, tricuspid (the central cusp notablylonger and broader than the lateral ones), in two principal rows in jaws, the teeth ofthe outer row largest ; no teeth on roof of mouth ; tongue short and broadly rounded. A NEW ASCENSION I. ANGELFISH 229 Gill membranes narrowly attached to isthmus ; 5 (5, 7) fleshy papillae in mid-ventral line between chin and isthmus ; anterior and posterior nostrils separatedby a space about a third to a half of length of posterior nostril ; posterior nostriloval, slightly larger than round anterior nostril, separated from eye by a spaceabout equal to its width ; anterior nostril with a short membraneous tube with anelevated posterior flap. A stout spine at corner of preoperculum, its length about three-quarters of orbitdiameter ; 20 (25, 23) small spines of somewhat irregular size along upper marginof preoperculum and one spine on lower edge of preoperculum, about a third (abouta fifth in paratypes) as long as spine at angle ; 2 smallish spines on interoperculum ;a few small spines on suboperculum ; operculum with 2 broad flat spines, one nearupper end of gill opening and the other at about the level of the lower edge of eye ;preorbital with 3 (2, 2) spines along ventral edge. Scales coarsely ctenoid (up to about 20 ctenii on margins), the exposed portionridged ; from fourth dorsal spine onwards, dorsal fin with scaly base, scalationextending over three-quarters if not virtually all of fin on median and posteriorportion ; anal fin with scaly base, scalation extending over three-quarters or moreof fin on median and posterior portion ; scales on basal half of caudal fin, with smallscales extending out on fin rays for about a third more of length of fin ; pectoralfins scaled only basally ; pelvic fins with small scales extending out on fin rays forup to three-quarters of length of spine. Head almost completely scaled. Lateral line steeply arched anteriorly, ending below posterior part of dorsal fin. Origin of dorsal fin slightly posterior to vertical through upper angle of gill opening;membranes of anterior portion of dorsal fin, between first and second, second andthird, and third and fourth spines, deeply incised, while those between posteriorspines and rays only slightly if at all indented. Membranes of anterior portion ofanal fin, between first and second, and second and third spines deeply incised,while those between posterior rays not indented. Caudal fin nearly truncate.Pectoral fins more or less rounded, not reaching to level of origin of anal fin. Originof pelvic fins below base of pectoral fins ; first pelvic soft ray prolonged, reachingspinous part of anal fin base, about a third as long again as pelvic spine. Coloration: In life, head and body bright to dark blue ; snout bright yellow-orange ; chest with orange tinges ; bright yellow-orange to orange stripe fromsnout between eyes along dorsal body contour onto upper part of caudal peduncle.Orbit ringed with bright blue ; iris orange. Dorsal fin bright yellow-orange toorange with dark blue distal margin ; anal fin bright to dark blue, becoming bluishorange distally and posteriorly, with bright blue distal margin ; pelvic fins duskybluish orange with bright blue anterior margin ; pectoral fins hyaline ; caudal finbright yellow-orange to orange with dark blue distal margin. In alcohol, bright to dark blue markings become brown to dark brown ; yellow-orange to orange markings become beige on body, beige to hyaline on fins. REMARKS. As was pointed out by Randall & Caldwell (1973), 'All of the species ofCentropyge are very similar in body form and counts, but they are distinctive incolour pattern'. The colour pattern of Centropyge resplendens (see above and Plate i) 14 230 R. LUBBOCK & R. D. SANKEY is quite distinctive and separates it clearly from all known species. As might beexpected, C entropy ge resplendens is closest to C. argi Woods & Kanazawa and toC. aurantonotus Burgess, both restricted to the tropical western Atlantic. Centropygeargi has a dark blue body, but only the head and chest are yellow to orange (seecolour illustrations in Burgess, 1974, and Bohlke & Chaplin, 1968). Centropygeaurantonotus also has a dark blue body but with orange head, chest, dorsal bodycontour and dorsal fin (see colour illustration in Burgess, 1974). Centropyge auranto-notus differs from C. resplendens in the coloration of the caudal fin and of thedorsal margin of the caudal peduncle, which are dark blue in C. aurantonotus butyellow-orange to orange in C. resplendens ; also, the head of C. aurantonotus isorange, while that of C. resplendens is bright to dark blue, with only the snout anddorsal contour yellow-orange to orange. Briggs (1974) stated that much of Ascension Island's shore fauna was apparentlyderived from the western Atlantic, and that some justification existed for placingAscension in the West Indian Province of the Western Atlantic Region. Only twoPomacanthidae are known from Ascension : Centropyge resplendens, which isclosely related to tropical western Atlantic forms (no Centropyge is known as yetfrom the eastern Atlantic), and Pomacanthus paru, which is otherwise recorded onlyfrom the tropical western Atlantic. (The only record of P. paru from the easternAtlantic is that of Fowler, 1936, who considered that Chaetodon leachii Bowdichfrom Madeira was probably P. paru ; having read the original description of C.leachii, we find it most unlikely that C. leachii is synonymous with P. paru.} Thederivation of Ascension Pomacanthidae appears prima facie to be in accordance withBriggs' statement, although it should be emphasized that the pomacanthid faunaof the eastern Atlantic is still relatively poorly known, making genuine comparisondifficult. ETYMOLOGY. The name resplendens is derived from the Latin for resplendent,and refers to the beautiful coloration of this species. MATERIAL EXAMINED, (a) Holotype, 35-3 mm SL, among coral boulders at 20 m,Ascension Island, coll. G. Earnshaw in December 1974 ; BMNH 1974.12.20.1.(b) Paratype, 29-0 mm SL, coll. with (a) ; BMNH 1974.12.20.2. (c) Paratype,24-9 mm SL, coll. with (a) ; USNM. ACKNOWLEDGEMENTS P. J. P. Whitehead of the British Museum (Natural History) kindly reviewed thetext and offered suggestions. We are also grateful to George Earnshaw, whocollected the specimens, and to Gordon Howes of the British Museum (NaturalHistory), who provided us with drawings. REFERENCES BOHLKE, J. E. & CHAPLIN, C. G. C. 1968. Fishes of the Bahamas and adjacent tropical waters. Philadelphia, xxiii + yyipp. BRIGGS, J. C. 1974. Marine zoogeography. New York. 475pp.BURGESS, W. E. 1974. Centropyge aurantonotus, a new species of pygmy angelfish from the southern Caribbean. Trop. Fish Hobby, 23 (November) : 90-97. A NEW ASCENSION I. ANGELFISH 231 FOWLER, H. W. 1919. The fishes of the United States Eclipse Expedition to West Africa. Proc. U.S. natn. Mus. 56 : 195-292. 1936. The marine fishes of West Africa. Bull. Am. Mus. nat. Hist. 70 : i-vii + 1-1493.FRASER-BRUNNER, A. 1933. A revision of the chaetodont fishes of the subfamily Poma- canthinae. Proc. zool. Soc. Lond. : 543-599.RANDALL J. E. & CALDWELL D. K. 1973. A new butterflyfish of the genus Chaetodon and a new angelfish of the genus Centropyge from Easter Island. Contr. Sci. (237) : i-n.WOODS, L. P. & KANAZAWA, R. H. 1951. New species and new records of fishes from Bermuda. Fieldiana Zool. 31 : 629-644. R. LUBBOCK Department of ZoologyUNIVERSITY OF CAMBRIDGEDOWNING STREETCAMBRIDGE R. D. SANKEY24 HILL CLOSESTANMOREMIDDLESEX 14* PLATE i Holotype of Centropyge resplendens n. sp. 35-3 mm SL. Ascension Island.Aquarium photograph by R. Lubbock. Bull Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE i V* *% *' A NEW SPECIES OF NANOCHROMIS (PISCES, CICHLIDAE) FROM THE OGOWE SYSTEM, GABON By ETHELWYNN TREWAVAS AMONG the fishes collected by the Cambridge Expedition of 1957 to Gabon is asingle specimen of a cichlid of an undescribed species. Unwilling to describe itfrom a single specimen, I had put it aside, but recent renewed interest in the ich-thyology of Gabon persuades me to call attention to it and name it. RecentlyThys van den Audenaerde & Loiselle (1971) have described two species of Nano-chromis from Ghana and Ivory Coast, one of which is so similar to the Gabonesefish as to require close comparison with it. Nanochromis gabonicus sp. nov. HOLOTYPE: $, 47 + 13 mm, from a pool by the road from Mitzic to Medouneu,probably in the drainage basin of R. Okano, tributary of the Ogowe. BMNH1967.10.12.57. CHARACTERS OF GENERIC IMPORTANCE. Pad on roof of pharynx with a groovearound its antero-lateral edge and covered with tubercles resembling, under lowmagnification, the batteries of taste-buds of the pad of Chromidotilapia (seeTrewavas, 1974, pi. 4). Microbranchiospines absent. Outer ceratobranchialgill-rakers with two subequal tubercles. At least in female, first and second softpelvic rays approximately equal, not produced, but coming to a blunt point. Lastscale of upper lateral line separated from dorsal fin by only a half-scale. Scalescycloid, the circuli parallel to the edge of the scale (roman), except a few with agothic arrangement. Although the base of the skull was not examined in this unique specimen, theother characters indicate that it belongs to the group in which the apophysis for thepharyngeals is formed by the parasphenoid alone. DISTINGUISHING SPECIFIC CHARACTERS. A black spot at the tip of each pelvicfin ($). Chest completely covered with imbricating scales. DESCRIPTION. Proportions as % SL : length of head 35, of pectoral fin 20-5,length and depth of caudal peduncle 13-8. Proportions as % length of head : snout 39-4, diameter of eye 27, depth of pre-orbital 21, interorbital width 21, length of lower jaw 36-5. Two series of small scales on the left cheek, two and a short third series on right ;a naked patch below them. Teeth all unicuspid, in two series in upper jaw, 2-3 in lower ; 52 in outer series ofupper jaw. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 234 E. TREWAVAS FIG. i. Nanochromis gabonicus, holotype. Lower pharyngeal bone deeply indented posteriorly, with short blade and ratherfew, pointed, bicuspid teeth. Gill-rakers 5 + 1 + 8 on first arch. Scales 28 in the lateral line series, the upper lateral line piercing 19 scales, thelower 4 on the left side, 8 on the right, the last of which is on the caudal fin ; 3between origin of dorsal and lateral line ; 3 between bases of pectoral and pelvicfins ; immediately in front of this row the scales are smaller, but the chest is com-pletely covered with imbricating scales ; 12 around the caudal peduncle. Vertebrae 27 (12 + 15). Dorsal XV 9 ; last spine 12% SL. Anal III 7 ; third spine 13-8% SL. Pectoral fin rounded. Pelvic with first two soft rays approximately equal, notproduced, just reaching vent when laid back. Caudal rounded at the corners, thestraight edge in the middle involving 3^ upper and 3! lower rays. Colour in general pale, top of head darker. A dark bar from eye to top of oper-culum and a dark band along middle of side to base of caudal, emphasized in five orsix places ; traces of three broad blotches from dorsum, tapering towards lateralband. Dorsal lappets and upper edge of soft dorsal black. Postero-dorsal edge ofcaudal black with a white submarginal band ; a black spot at the tip of each pelvicfin. The ovaries of this small fish are partly discharged, still containing a few eggs ofabout 1-3 mm diameter as well as some smaller oocytes. This specimen (and probably the species to which it belongs) differs from N.cavalliensis Thys & Loiselle in the blunt pelvic fin and the absence of any pro-longations of caudal rays, but it is smaller than the holotype (also a $) of thatspecies. It also has a wider interorbital region, more teeth in the upper jaw (cf. 42in N. cavalliensis) and a slightly shorter pectoral fin. No black marginal and whitesubmarginal bands were described on the caudal of N. cavalliensis and in thatspecies ripe eggs were 2-0 x 1-5 mm. Those in the type of N. gabonicus may ofcourse be unripe, of a batch to follow those recently shed. A NEW SPECIES OF NANOCHROMIS 235 DISCUSSION. The type species of Nanochromis Pellegrin, N. nudiceps (Boulenger),inhabits swiftly running water and has characters associated with this habit, namelyscaleless chest, middle soft rays of pelvic fin longer than the others. In it scales arealso absent from the cheek, top of head and a narrow strip along the back on eachside of the base of the dorsal fin ; both lateral lines are interrupted. Matthes (1964 : 134) has re-examined the types of the other two nominal species,N. dimidiatus Pellegrin, 1900 (from Banghi) and N. squamiceps (Boulenger) fromR. Lindi, Congo system, and finds that both fall within the range of variation ofspecimens from Lake Tumba basin regarded by him as a single species. In the.British Museum four specimens of N. dimidiatus collected at Banghi on two separateoccasions show that the incomplete squamation is maintained, at least in thatpopulation. Thys, in an addendum to his paper of 1968, claims to have evidencefrom living specimens that they are distinct. Thys also refers to an unnamed (new) species from R. Ivindo, tributary of theOgowe, which he relates to N. squamiceps. From his brief characterization of thisI believe it must be a species of which Dr Gery has sent me a few specimens. Imention it here to state that it differs from N. gabonicus in having the pelvic fin alittle more acute in the female (much more in the male), with a black streak alongthe first soft rays but no black spot at its tip, a broad naked area on the chest oneither side of a median row of scales and very small scales on the nape and occiput.As in N. gabonicus, the vertical fins of the female are clear or lightly peppered exceptat the edges ; in the male they bear conspicuous series of spots. REFERENCES BOULENGER, G. A. 1899. Materiaux pour la faune du Congo. Poissons nouveaux. Cinquiemepartie. Ann. Mus. Congo Zool. 1 : 97-128, pis 40-47. (P. nudiceps, p. 122, pi. 44, fig. 6.) 1915. Cat. Afr. Fish. Ill, 526 pp. (Nanochromis p. 375.) MATTHES, H. 1964. Les poissons du Lac Tumba et de la region d'Ikela. Etude systematique,ecologique et zoogeographique. Theses of Amsterdam Museum, April, 1964 : 201 pp.,6 pis. PELLEGRIN, J. 1900. Poissons nouveaux du Congo fran^ais. Bull. Mus. nat. Hist. Paris1900 : 98-101. 1904. Contribution a 1'etude anatomique, biologique et taxonomique des poissons de lafamille des cichlides. M6m. Soc. zool. Fr. 16 : 41-401, pis 4-7. THYS VAN DEN AUDENAERDE, D. F. E. 1968. A preliminary contribution to a systematicrevision of the genus Pelmatochromis Hubrecht (sic) sensu lato (Pisces, Cichlidae). Rev.Zool. Bot. afr. 77 : 349-391. & LOISELLE, P. V. 1971. Description of two new small African cichlids. Rev. Zool. Bot.afr. 83 : 193-206, figs 1-5. TREWAVAS, E. 1974. The freshwater fishes of Rivers Mungo and Meme and Lakes Kotto,Mboandong and Soden, West Cameroon. Bull. Brit. Mus. nat. Hist. (Zool.) 26 : 329-419,pis 1-5. DR. E. TREWAVAS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD THE FIRST ZOEAL STAGES OF CANCERPAGURUS L, PINNOTHERES PISUM (PENNANT)AND MACROPHTHALMUS DEPRESSUS RUPPELL (CRUSTACEA, DECAPODA, BRACHYURA). By A. L. RICE SYNOPSIS The first zoeal stages in Cancer pagurus, Pinnotheres pisum and Macrophthalmus depressus aredescribed and larval characters within their respective families are discussed. INTRODUCTION ALTHOUGH there are published accounts of one or more larval stages of severalhundreds of brachyuran species, less than one hundred of these descriptions includea significant amount of detail. Consequently, while there has been a welcome tend-ency in recent years for larval papers to deal with all the developmental stages ofcrabs reared in the laboratory, there is still a need for detailed re-descriptions, evenof single zoeal stages, where the previous accounts are clearly inadequate. Thepurpose of this paper is to provide such details of the first zoeal stages of three crabsfrom material in the larval collections of the British Museum (Natural History). Cancer pagurus Larvae hatched from a female collected at Lulworth Cove, Dorset, June, 1973. B.M.(N.H.) registration no. 1975 : 66. Dimensions : Tip of dorsal to tip of rostral spines : 2-4-2-6 mm.Tip to tip of lateral carapace spines : i-o-i-i mm.Carapace (Fig. la, b) : Dorsal carapace spine straight or with a very slight backward curve, about twiceas long as the carapace and slightly longer than the straight rostral spine. Rostralspine with minute spinules. Lateral spines about half carapace length. Carapacewith a low anterio-median papilla, a pair of setae at the base of the dorsal spine andposterio-lateral margins without sub-marginal setae.Antennule : Simple, with 2 or 3 aesthetascs and a single seta. Antenna (Fig. ic) : Spinous process slightly more than half length of rostrum, withspinules on the distal two-thirds increasing in size towards the tip. Exopod aboutone-third length of spinous process, with i long terminal seta and 2 shorter ones, ofwhich i is fused. Bull. BY. Mus. nat. Hist. (Zool.) 28, 5 Issued 17 September, 1975 238 A. L. RICE FIG. i . Cancer pagurus, first zoea : a, lateral view ; b, frontal view ; c, antenna ; d,maxillule ; e, maxilla ; f, first maxilliped (exopod omitted) ; g, endopod of secondmaxilliped ; h, telson ; j, detail of telson ; k, detail of telson in the third zoea of C.anthonyi. Bar scale represents 0-5 mm for a, b and h, and 0-25 mm for c-g, j and k. Maxillule (Fig. id) : Endopod of 2 segments, with i and 6 setae respectively.Maxilla (Fig. le) : Endopod bilobed with 3 + 5 (occasionally 4) setae; scapho-gnathite with 4 marginal setae and a long plumose posterior projection. ZOEAE OF BRACHYURA 239 First maxilliped (Fig. if) : Basis with 10 setae (arranged 2, 2, 3, 3) ; endopod seg-ments with 3, 2, i, 2 and 4 + 1 setae ; exopod with 4 natatory setae.Second maxilliped (Fig. ig) : Basis with 4 setae ; endopod of 3 segments with i, iand 6 setae. Abdomen (Fig. la) : Somite 2 with forwardly directed dorso-lateral knobs ; somites2-5 each with a pair of dorso-posterior setae and slight projections on the posterio-lateral margins. Telson (Fig. ih, j) : Forks long, slender and divergent, each with one lateral and onedorsal spine, and minute spinules distally ; inner posterior margin with a deep,rounded median notch and three pairs of processes armed with short setules andspinules, those of the distal one-third of the outer process being particularly stoutand tooth-like. DISCUSSION. The larvae of Cancer pagurus were first hatched by Thompson in 1829 and havesubsequently been described many times (see Lebour, ig28a). However, with theexception of Williamson's (1910) account, most of these descriptions are very in-adequate and the larvae of several Pacific species of the genus are much better knownthan are those of C. pagurus. Thus, Aikawa (1937) published details, includingappendage setation, of the first stage of C. gibbosulus (de Haan), Mir (1961) comparedthe first zoeae of C. magister Dana, C. antennarius Stimpson and C. anthonyi Rathbun,Poole (1966) described the complete development of C. magister, and Trask (1970,1974) has similarly dealt with C. productus Randall and C. anthonyi. From theseaccounts, including the details of C. pagurus given here, it is now possible to definegeneric, and possibly familial, zoeal characters. All of the Cancer zoeae so far known possess well-developed dorsal, rostral andlateral carapace spines, antennae with spinous processes more than half as long asthe rostrum, the two-segmented endopod of the maxillule with 1 + 4 to i + 6 setae,the endopod of the maxilla with at least six setae, usually arranged in two distinctgroups, lateral knobs on only the second abdominal somite and the telson furcaewith one lateral and one dorsal spine. In all the species, with the exception ofC. anthonyi, the proximal segment of the endopod of the first maxilliped is describedas carrying three setae and this, together with the characters listed above, dis-tinguishes these Cancer zoeae from those of all other known brachyurans. Mir alsorecorded three setae on this segment in his first-stage C. anthonyi but Trask (1974)reported only two setae in all five stages of his reared material. This character seemsgenerally to be rather conservative amongst brachyuran zoeae, at least withingenera* and often within whole families, so that it seemed at least possible that Traskwas mistaken. However, I have been able to examine Trask's material and can con-firm that C. anthonyi does differ from the other known species in this respect. Butthis species possesses the strong tooth-like spines on the outer posterior telsonprocess (see Fig. ij, k) which seem to be typical of Cancer zoeae and which have * For instance, a re-examination of the material of Corystes cassivelaunus described by Ingle & Rice(1971) revealed that the first zoea carries three setae on this segment, as in the later stages, and nottwo as figured in that paper. 240 A. L. RICE otherwise been reported only in the closely related Atelecydus rotundatus (Olivi)(Bourdillon-Casanova, 1960). Pinnotheres pisum (Pennant) Larvae hatched in July, 1971, from a female collected at Plymouth, Devon. B.M.(N.H.) registration no. 1975 : 67.Dimensions : Tip of rostral spine to mid-dorsal point of the carapace : 0-60 mm. Tip to tip of lateral carapace spines : 0-55 mm.Carapace (Fig. 2a, b) : Rostral spine about half carapace length, with a slight forward curve. Lateralspines arise close to the posterio-lateral carapace margins, directed downwards andslightly backwards. No dorsal spine, but with a pair of setae close to the mid-dorsalline of the carapace. Antennule (Fig. 2d) : Reduced to small hemispherical buds carrying 2 aesthetascsand a single seta.Antenna : Totally absent. Maxillule (Fig. 2e) : Endopod of two segments carrying o and 5 setae respectively.Maxilla (Fig. 2f) : Coxal and basal endites not clearly bilobed ; endopod with 3setae, of which 2 are more or less terminal ; scaphognathites with 3 marginal setaeand a long plumose posterior process. First maxilliped (Fig. 2g) : Basis with 2, 2, 3, 3 setae ; endopod segments with2, 2, i, 2 and 4 + 1 respectively ; exopod with 4 natatory setae.Second maxilliped (Fig. 2h) : Basis with 4 setae ; endopod of 2 segments with o and4 or 5 setae ; exopod with 4 natatory setae. Abdomen (Fig. 2c) : Somites 2 and 3 with small dorso-lateral knobs, those of thesecond somite joined by a slight ridge over the dorsal surface. Somites 4 and 5widening to the tri-lobed telson ; rounded median lobe not protruding beyond theacute lateral lobes ; three setose processes between the median and lateral lobes oneach side. DISCUSSION. Lebour (i928a, b) described the first stage of P. pisum hatched from the egg andattributed to the same species a second-stage zoea taken in the plankton. Herdescription is very inadequate, but where comparison is possible it agrees with thatgiven here except that Lebour was able to detect the rudimentary antenna and foundthe median telson lobe to overreach the lateral lobes. The available descriptions of the larvae of other species of Pinnotheres indicatethat there is a good deal of morphological variation within the genus. Thus, whilethe zoeae of P. pinnotheres (L.) ( = veterum) and P. maculatus Say possess dorsal,rostral and lateral carapace spines (see Hyman, 1925 ; Labour, I928a, b ; Costlow& Bookhout, 1966), the dorsal spine is absent in P. pisum and P. placunae (Hornelland Southwell) (Lebour, i928a, b ; Hashmi, 1970 ; this paper), the laterals areabsent in P. taylori Rathbun (Hart, 1935), while none of the carapace spines arepresent in P. ostreiim (Say) (Hyman, 1925 ; Sandoz & Hopkins, 1947). Similarly, 2 4 I FIG. 2. Pinnotheres pisum, first zoea : a, lateral view ; b, frontal view ; c, abdomen ;d, antennule ; e, maxillule ; f, maxilla ; g, first maxilliped ; h, second maxilliped.Bar scale represents 0-5 mm for a and b, and 0-25 mm for c-h. 242 A. L. RICE while P. pisum, P. pinnotheres, P. placunae and P. ostreum possess the very charac-teristic trilobed telson which has been found only in the Pinnotheridae, in P. maculatusand P. taylori the telson is the much more typical brachyuran forked type. Thedegree of development of the antennules and antennae is also very variable, theseappendages being greatly reduced or even absent, at least in the early stages, inP. ostreum, P. placunae and P. pisum, moderately developed in P. taylori, and quitenormal in P. maculatus. The number of zoeal stages also varies within the genus,with P. pinnotheres and P. taylori having only two zoeae, P. placunae probablyhaving three, P. ostreum four and P. maculatus five. One result of this variation indevelopment rate is that the pleopods make their appearance at different stages inthe different species, these appendages appearing as buds in the third stage in P.ostreum and the fourth in P. maculatus, but being well developed by the third stagein P. placunae or even as early as the second stage in P. pinnotheres and P. taylori.Finally, as Costlow & Bookhout (1966) point out, P. maculatus seems to have a farmore typically 'brachyuran' development than any of the other described species,since it is apparently the only one in which the sixth abdominal somite is separatedfrom the telson in the late zoeal stages. Clearly, then, there is no difficulty in distinguishing between the known zoeaelarvae of Pinnotheres species. A more difficult problem may be the recognition ofcharacters common to the zoeae of Pinnotheres or of the Pinnotheridae generally,which will distinguish them from other crab larvae even where, as in P. taylori,neither the characteristic trilobed telson nor the posterio-ventrally directed lateralcarapace spines are present. Firstly, in all those species for which the information is available the endopod ofthe maxilla carries only three setae and these are arranged in a single more or lessterminal group, or at least not clearly divided into two distinct groups. Thischaracter at once distinguishes the Pinnotheridae from almost all other brachyuranlarvae, for such a setal armature has been recorded outside this family only in theLeucosiidae and the ocypodid sub-family Ocypodinae (see also below). The three-segmented endopod of the second maxilliped readily distinguishesthese ocypodids from both the Leucosiids and the pinnotherids in which this endopodnever has more than two joints. Finally, while there are a number of differencesbetween the pinnotherids and the leucosiids in the detailed morphology of the ap-pendages, the most obvious distinction is the simple triangular telson with the closelyspaced row of six processes on the relatively straight posterior margin which hasbeen found in every leucosiid zoea so far described ; this contrasts strongly with boththe tri-lobed telson and the more typical fork found in the Pinnotheridae. Macrophthaltnus depressus Riipell Larvae hatched in Bahrain, Arabian Gulf, from a female collected from theforeshore at Jufair in March, 1974. B.M.(N.H.) registration no. 1975 : 68. Dimensions : Tip of dorsal to tip of rostral spines : 066 nvm.Maximum width across carapace : 0-29 mm. ZOEAE OF BRACHYURA 243 Carapace (Fig. 3a, b) : Slender dorsal and rostral spines, each about half the carapace length ; lateralspines absent. A pair of short setae at the base of the dorsal spine. Posterio-ventral edge of carapace with a small tooth and a slightly crenulate margin, but with-out sub-marginal setae. FIG. 3. Macrophthalmus depressus, first zoea : a, lateral view ; b, frontal view ; c,abdomen ; d, detail of telson ; e, antenna ; f , endopod of maxillule ; g, maxilla ; h, firstmaxilliped ; j, second maxilliped. Bar scale represents 0-5 mm for a-c, 0-25 mm fore-j, and o-i mm for d. 244 A - L - Antennule (Fig. 3b) : Simple, with 2 terminal aesthetascs and i seta. Antenna (Fig. 36) : Spinous process slightly shorter than rostrum ; exopod a simple, unarmed spine slightly more than half the spinous process. Maxillule (Fig. 3!) : Endopod of 2 segments with i and 5 setae respectively. Maxilla (Fig. 3g) : Endopod with 2 + 2 setae ; scaphognathite with 4 marginal setae and a long, plumose posterior process. First maxittiped (Fig. 3h) : Basis with 9 or 10 setae ; endopod segments with 2, 2, i, 2 and 4 + 1 setae ; exopod with 4 natatory setae. Second maxilliped (Fig. 3]) : Basis with 4 setae ; endopod of 3 segments with o, i and 5 setae respectively ; exopod with 4 natatory setae. Abdomen (Fig. 3c) : Somites 2 and 3 with dorso-lateral knobs ; somites 2-5 each with short posterio-lateral processes and a pair of short setae close to the dorso- posterior margin. Telson widening only slightly posteriorly, the forks about the same length as the body of the telson. Posterior margin with three pairs of process which are naked distally (see Fig. 3d). Telson forks unarmed except for two rows of minute spinnules basally. DISCUSSION. Hashmi (1969) described the first zoeae of five species of Macrophthalmus, in-cluding M . depressus. In general, Hashmi's account agrees very closely with thatgiven here, except in the details of the setation of the maxillipeds. For instance,Hashmi gives the setal formula of the endopod of the first maxilliped as i, 2, i, 2, 5,whereas with the exception of his own account of M. crinitus Rathbun and the larvaeof Dotilla blanfordi Alcock and D. sulcata (Forskal) (Rajabai, 1959 ; Ramadan, 1940)the proximal segment carries two setae in every ocypodid larva for which this infor-mation is known, and the typical setal formula for the family seems to be 2, 2, i, 2, 5. Similarly, Hashmi records a seta on the basal segment of the endopod of thesecond maxilliped whereas I was unable to find a seta in this position in my materialand it seems to be unusual in the family as a whole (Table i). Using characters of the carapace spines, antennae, telson, maxillae and maxillipedsAikawa (1937) was able to separate zoeae of the ocypodid genera Macrophthalmus,Tympanomerus (= Ilyoplax), Scopimera and Uca into groups corresponding to thethree sub-families based an adult taxonomy. Thirty years later Wear (1968) re-examined the larval situation within the Ocypodidae, information on the larvae offourteen species belonging to seven genera by then being available, but was unableto obtain any support for the adult classification. However, in separating the larvaeinto groups Wear gave greatest significance to the presence or absence of lateralcarapace spines, and somewhat lesser significance to the form of the abdomen andthe degree of development of the antennal exopod. There is, however, a good dealof evidence to suggest that the setation of the mouthparts may reflect taxonomicdivisions between larvae more effectively than these more 'obvious' characters.Certainly, on the basis of the setation of the endopods of the maxillules, maxillaeand of the second maxillipeds the described ocypodid larvae fall into distinctgroups, which correspond rather well with the accepted sub-families (Table i). ZOEAE OF BRACHYURA 245 TABLE i Setation of the maxillae and the second maxilliped, the number of telson fork spines, and thepresence or absence of lateral carapace spines in described ocypodid zoeae MACROPHTHALMINAE Macrophthalmus depressusMacrophthalmus depressusMacrophthalmus depressusMacrophthalmus dilatatusMacrophthalmus japonicusMacrophthalmus sulcatusMacrophthalmus latreillisMacrophthalmus pacificusMacrophthalmus crinitusHemiplax hirtipes OCYPODINAE Ocypode quadrata Ocypode platytarsisOcypode gaudichaudiiUca annulipesUca annulipesUca marionisUca pugilatorUca pugnaxUca minaxUca triangularis SCOPIMERINAE Scopimera globosusDotilla sulcata Dotilla sulcataDotilla blanfordiIlyoplax pusillusIlyoplax gangetica MICTYRIDAE ? Mictyris longicarpus Maxillule Maxilla Maxilliped 2endopod endopod endopod , 5 o, 50,4 o, 4o, 4o, 4o,4 2 (3) 2, 3 (5)2, 3 (5) 2, 2 (4) 2, 3 (5) 2, 3 (5) (4-5) , o, 5 o, i, 6 0, 2, 4 1, 1,4i, i, 5 o, 5 i, 5 2, 2 (4) i, i, 6 Telson Lateralfork carapacespines spines Source This paper Hashmi, 1969Aikawa, 1929Aikawa, 1929Aikawa, 1929 Hashmi, 1969Hashmi, 1969Hashmi, 1969 + Hashmi, 1969+ Wear, 1968 + Diaz & Costlow, 1972 + Rajabai, 1951+ Crane, 1940Feest, 1969 Hashmi, 1968Hashmi, 1968 Hyman, 1920Hyman, 1920Hyman, 1920 Feest, 1969 + Aikawa, 1929+ Gohar & Al- Kholy, 1957+ Ramadan, 1940 Rajabai, 1959+ Aikawa, 1929 Feest, 1969 Cameron, 1965 Thus the zoeae of the genera Macrophthalmus and Hemiplax (sub-family Macroph-thalminae) all have the basal segment of the endopod of the maxillule armed with asingle seta, the endopod of the maxilla armed with a total of four or five setae, andthe middle segment of the endopod of the second maxilliped carrying a single seta.In contrast, the genera Ocypoda and Uca (sub-family Ocypodinae) have both thebasal segment of the endopod of the maxillule and the middle segment of the endopodof the second maxilliped unarmed, while the endopod of the maxilla carries a totalof only three setae. Larvae of the third sub-family, the Scopimerinae, represented 246 A. L. RICE by the genera Scopimera, Ilyoplax and Dotilla, show a combination of these charac-ters which tend to exclude them from both of the other two groups. Finally, thefirst zoea of Mictyris longicarpus Latreille has setal characters similar to those of theMacrophthalminae but, as pointed out by Wear (1968), it possesses other characters,including the absence of both dorsal and lateral carapace spines and the form of thetelson, which may support Balss' (1957) separation of Mictyris into a distinct family.Table I also includes data on the armature of the telson forks and the presence orabsence of lateral caparace spines, showing that these characters are not correlatedwith the sub-family groups but vary even within the same genus. ACKNOWLEDGEMENTS My thanks are due to Dr R. W. Ingle for making material in the collections of theBritish Museum (Natural History) available for study, to divers of the GuildfordBranch of the British Sub-Aqua Club who collected the adult Cancer pagurus, andto Dr R. S. K. Barnes for confirming the identity of the Macrophthalmus depressus.Finally, I am indebted to Dr T. Trask for allowing me to examine his reared materialof Cancer anthonyi REFERENCES AIKAWA, H. 1929. On larval forms of some Brachyura. Rec. oceanogr. wks Jap. 2 : 1-55. 1937- Further notes on brachyuran larvae. Rec. oceanogr. wks Jap. 9 : 87-162. BALSS, H. 1957- Decapoda. In : H. G. Bronn (Ed.), Klassen und Ordnungen des Tierreichs, 5 (i), Buch 7, Lief. 12 : 1505-1672.BOURDILLON-CASANOVA, L. 1960. Le meroplancton du Golfe de Marseille : Les larves de crustacesdecapodes. Rec. Trav. Stat. mar. d'Endoume. 30 (18) : 286pp.CAMERON, A. M. 1965. The first zoea of the soldier crab, Mictyris longicarpus (Grapsoidea : Mictyridae). Proc. Linn. Soc. N.S.W. 9 : 222-224.CRANE, J. 1940. Eastern Pacific expeditions of the New York Zoological Society. XVII. On the postembryonic development of brachyuran crabs of the genus Ocypode. Zoologica, N.Y. 25 : 65-82.COSTLOW, J. D. & BOOKHOUT, C. G. 1966. Larval stages of the crab, Pinnotheres maculatus, under laboratory conditions. Chesapeake Sci. 7 : 157-163.DIAZ, H. & COSTLOW, J. D. 1972. Larval development of Ocypode quadrata (Brachyura : Crustacea) under laboratory conditions. Mar. Biol. 15 : 120-131.FEEST, J. 1969. Morphophysiological studies on the ontogeny and sexual biology of Uca annulipes and Uca triangularis as compared to Ilyoplax gangetica. Form. Fund. 1 : 159- 225.GOHAR, H. A. F. & AL-KHOLY, A. A. 1957. The larvae of some brachyuran Crustacea. Publ. mar. biol. Sta. Al Ghardaga, 9 : 145- 176.HART, J. F. L. 1935- The larval development of British Columbia Brachyura. I. Xanthidae, Pinnotheridae (in part) and Grapsidae. Can. J. Res. 12 : 411-432.HASHMI, S. S. 1968. Study on larvae of (Gelasimus) (Ocypodidae) reared in the laboratory (Decapoda : Crustacea). Pak. J. Sci. Res. 20 : 50-56. 1969. Studies on larval Ocypodidae (Macrophthalmus) hatched in the laboratory (Deca-poda : Crustacea). Pak. J. Sci. Res. 21 : 42-54.1970. The larvae of Elamena (Hymenosomidae) and Pinnotheres (Pinnotheridae) hatched in the laboratory (Decapoda : Crustacea). Pak. J. Sci. Ind. Res. 12 : 279-285. ZOEAE OF BRACHYURA 247 HYMAN, O. W. 1920. The development of Gelasimus after hatching. /. Morph. 33 : 485-525. 1922. Adventures in the life of a fiddler crab. Rep. Smithson. Instn, 1920 : 443-59. 1925. Studies on the larvae of crabs of the family Pinnotheridae. Proc. U.S. natn. Mus. 64: 1-9.INGLE, R. W. & RICE, A. L. 1970. The larval development of the masked crab, Corystes cassivelaunus (Pennant) (Brachyura, Corystidae), reared in the laboratory. Crustaceana, 20 : 271-284.LEBOUR, M. V. i928a. The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Lond. 1928 : 473-56o-ig28b. Studies on the Plymouth Brachyura. II. The larval stages of Ebalia and Pinnotheres. J. mar. biol. Assoc. U.K. 15 : 109-118.MIR, R. D. 1961. The external morphology of the first zoeal stages of the crabs, Cancer magister Dana, Cancer antennarius Stimpson, and Cancer anthonyi Rathbun. Calif. Fish and Game, 47 : 103-111.POOLE, R. I. 1966. A description of laboratory-reared zoeae of Cancer magister Dana, and megalopae taken under natural conditions (Decapoda : Brachyura). Crustaceana, 11 : 83-97.RAJABAI, K. G. 1951. Some stages in the development and bionomics of Ocypode platytarsis. Proc. Indian Acad. Sci. 33 : 32-40.1959- Studies on the larval development of the Brachyura. I. The early and post larval development of Dotilla blanfordi Alcock. Ann. Mag. nat. Hist. ser. 13, 2 : 129-135.RAMADAN, M. M. 1940. On the first zoeal stage of Dotilla sulcata (Forskal). Ann. Mag. nat. Hist. Ser. n, 5 : 253-255.SANDOZ, M. & HOPKINS, S. H. 1947. Early life history of the oyster crab Pinnotheres ostreum (Say). Biol. Bull. 93 : 250-258.TRASK, T. 1970. A description of laboratory-reared larvae of Cancer productus Randall (Decapoda, Brachyura) and a comparison to larvae of Cancer magister Dana. Crustaceana, 8 : 133-146.1974. Laboratory reared larvae of Cancer anthonyi (Decapoda : Brachyura) with a brief description of the internal anatomy of the megalopa. Mar. Biol. 27 : 63-74.WEAR, R. G. 1968. Life history studies on New Zealand Brachyura. 3. Family Ocypodidae. First stage zoea larva of Hemiplax hirtipes (Jacquinot, 1853). N ' .Z. ] . mar. freshw. Res. 2 : 698-707.WILLIAMSON, H. C. 1910. On the larval and later stages of Portunus holsatus, Portunus puber, Portunus depurator, Hyas araneus, Eupagurus bernhardus, Galathea dispersa, Crangon trispinosus, Cancer pagurus. Fish. Scot. Sci. Invest. 1909 : 2opp. Dr A. L. RICE INSTITUTE OF OCEANOGRAPHIC SCIENCES WORMLEY GODALMING SURREY 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.) 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- 3-75- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU THE HYDROID SPECIES OF OBELIA (COELENTERATA, HYDROZOA : CAMPANULARIIDAE), WITH NOTES ON THE MEDUSA STAGE P. F. S. CORNELIUS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 6 LONDON: 1975 THE HYDROID SPECIES OF OBELIA (COELENTERATA, HYDROZOA :CAMPANULARIIDAE), WITH NOTES ON THE MEDUSA STAGE BY PAUL FREDERICK SINEL CORNELIUS Pp. 249-293 ; 5 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 6 LONDON: 1975 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. 28 No. 6 of the Zoology series.The abbreviated titles of periodicals cited follow those ofthe World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.). ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 533,7-4-. 1 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 5 November, 1975 Price 2-50 THE HYDROID SPECIES OF OBELIA (COELENTERATA, HYDROZOA:CAMPANULARIIDAE), WITH NOTES ON THE MEDUSA STAGE By P. F. S. CORNELIUS CONTENTS PageSYNOPSIS. ........... 251 INTRODUCTION ........... 251 GENERIC DIAGNOSIS AND IDENTIFICATION OF THE HYDROID STAGE . . 252MORPHOLOGICAL AND OTHER VARIATIONS ...... 256 Obelia bidentata .......... 260 Obelia dichotoma . . . . . . . . . .265 Obelia geniculata . . . . . . . . . . 272 THE MEDUSA PROBLEM ......... 278 SPECIES TRANSFERRED TO OTHER GENERA ...... 279 ACKNOWLEDGEMENTS . . . . . . . . .281 REFERENCES ........... 282 INDEX ............ 291 SYNOPSIS The genus Obelia Peron & Lesueur, iSioa, is redefined and nominal taxa already describedfrom the hydroid stage are assessed. Three species are considered valid, O. bidentata Clarke,1875, O. dichotoma (Linnaeus, 1758) and O. geniculata (Linnaeus, 1758). Each is diagnosed,redescribed and illustrated with notes on morphological variation, nematocysts, identification,nomenclature, synonymy, type specimens and distribution. Owing to the paucity of informationthe many species described from the medusa stage cannot yet be evaluated or related to thehydroids. The affinities of those species of hydroids no longer assigned to Obelia are discussedbriefly. The genus Laomedea Lamouroux, 1812, is reduced to a synonym of Obelia but the genusCampanularia Lamarck, 1816, is regarded as valid. INTRODUCTION IN many genera of hydromedusae there is specific diversity of the medusa generationwhile the hydroid stage appears uniform. However, in the genus Obelia Peron &Lesueur, iSioa, the reverse is the case and it is the medusae which cannot be dis-tinguished. Thus, as summarized by Russell (1953), medusae liberated from thehydroid species Obelia dichotoma (Linnaeus, 1758) and 0. geniculata (Linnaeus,1758) and reared to maturity appear morphologically identical, both being referableto the medusa 0. lucifera (Forbes, 1848) as known from the plankton. Further 252 P. F. S. CORNELIUS confusion results from the large number of species that has been described fromthe hydroid, some seventy having been proposed between 1830 and 1948. Itseemed timely to evaluate the systematic criteria on which these species were pro-posed, hopefully to provide a basis for evaluating taxa described from the medusa.Previously, the genus had been fully revised only in the publications of Bedot(1901, 1905, 1910, 1912, 1916, 1918, 1925) who recognized thirty-seven hydroid speciesdescribed up to the end of 1910. In the present work, however, these and sub-sequently described species are referred to only three nominal species. Some speciespreviously assigned to Obelia are removed to allied genera. Although an attempt hasbeen made to consider all described hydroid species, only the more important usagesof the name of each have been cited. Full reference lists are already available inthe works of Bedot and in abstracting journals. The taxonomic status of all of the numerous species described from the medusageneration remains problematical. The difficulties outlined by Russell (1953) andKramp (1961) still remain, and further rearing work may be necessary before thehydroid and medusa species can be related. References to species described fromthe medusa before 1910 were provided by Mayer (1910) and Bedot, while thosedescribed between then and 1959 were listed by Kramp (1961). The material examined during the present study was drawn mainly from thecollections of the British Museum (Natural History). The figures were preparedwith the aid of a camera lucida. GENERIC DIAGNOSIS AND IDENTIFICATION OF THE HYDROID STAGE Genus OBELIA Peron & Lesueur, iSioa 1 Sertularia Linnaeus, 1758 : 807 (part). Medusa : Slabber, 1769 : 67, pi. 9, figs 5-8 (part) ; Slabber, 1775 : 40, pi. 9, figs 5-8 (part) ; Modeer, 1791 : 25 (part).Obelia Peron & Lesueur, iSioa 1 : 355 ; Oken, 1815 : 115, pi. 5, fig. 3 ; Deshayes & Edwards, 1840 : 170-171 ; McCrady, 1857 : 197-198, pi. n, figs 5-7 ; Hincks, 1868 : 146 (part) ; Fraser, 1937 : 82 ; Fraser, 1944 : 151 ; Russell, 1953 : 296 ; Naumov, 1960 : 260 (part); Naumov, 1969 : 281 (part) ; [non Obelia : Lamouroux, 1821 : 81 ; Deshayes & Edwards, 1836 : 245-246 ; Michelin, 1847 : 321 ; = Bryozoa (d'Orbigny, 1853 : 751 ; Gregory, 1909 : 47 ; Buge, 1951 : 464)].Laomedea Lamouroux, 1812 : 184 (part) ; Johnston, 1847 : 101 (part) ; Kramp, 1935 : 106 (part).Slabberia Oken, 1815 : 828 (rejected work, Opinion 417, International Commission on Zoological Nomenclature ; Ben them Jutting, 1970 : 60).Campanularia Lamarck, 1816: 112-113 (part).Thaumantias : Forbes, 1848 : 41 (part).Eucope Gegenbauer, 1856 : 241 (part) ; [syn. nov.].Schizocladium Allman, 1871 : 18. Obelaria Haeckel, 1879 : 173 (nom. nov. pro hydroid stage of Obelia).Obeletta Haeckel, 1879 : 173 ; [syn. nov.].Obelissa Haeckel, 1879 : 175 ; [syn. nov.].Monosklera von Lendenfeld, i885b : 910. 1 See footnote on facing page. HYDROID SPECIES OF OBELIA 253 TYPE SPECIES. Obelia sphaerulina Peron & Lesueur, iSioa 1 (nom. nov. proMedusa marina Slabber, 1769) ; by monotypy ; for nomenclatural purposes takenas conspecific with the hydroid 0. dichotoma (Linnaeus, 1758) (van der Hoeven,1862 : 280 ; Russell, 1953 : 297), not 0. geniculata (Linnaeus, 1758) as proposed byNaumov (1960 : 260 ; 1969 : 281). DIAGNOSIS. Colonial Campanulariidae (sensu Russell, 1953) with free medusae.Polyp generation forming upright colonies, branched or unbranched, variablyflexuose ; internodes annulated proximally, supporting hydrothecal pedicel ondistal lateral process. Hydrotheca bell-shaped, hydranth with prominent sphericalhypostome. Gonotheca inverted cone-shaped, usually with raised tubular aperture,occasionally simply truncate. Medusa umbrella flat, eversible, mesogloea thin ;mouth of manubrium 4-sided, lacking tentacles ; marginal tentacles 16 + on release,numerous in adult ; gonads 4, spherical, on radial canals. REMARKS. This restricted diagnosis agrees with that of Fraser (1937, 1944) inexcluding from the genus species with no medusa generation (accommodated in thegenera Campanularia Lamarck, 1816, and Gonothyrea Airman, 1864) and also thosein which the medusa has only 4 tentacles on release and which, when sexually mature,has a hemispherical umbrella (referred to the genus Clytia Lamouroux, 1812).There seems little justification in synonymizing the first two genera plus LaomedeaLamouroux, 1812, with Obelia as has been proposed by Naumov (1960, 1969). Theappearance of the adult medusa of 0. bidentata Clarke, 1875, is as yet unrecorded,but the characters of its young stages and of the hydroid fall within the abovediagnosis. The polyphyletic genus Medusa Linnaeus (1758 : 659) was disbanded by Peron &Lesueur (iSioa) who assigned the originally included species to other genera. Nospecies of Obelia was included in the original scope of Medusa, however, and Peron& Lesueur were justified in forming the genus Obelia to accommodate Medusamarina Slabber (1769). They also provided a new trivial name for the species,calling it Obelia sphaerulina. d'Orbigny (1853 : 684, 751) cited earlier uses of Obelia by Peron in 1803 and 1804,but gave no bibliographic information. Biographies about Peron (Alard, i8na,b ; Audiat, 1855 ; Girard, 1857) do not list any publication by him in 1803, whilePeron's 1804 papers (Peron, i8o4a-f) contain no mention of Obelia. It seems thatd'Orbigny was mistaken in giving these dates and that the name Obelia was notintroduced until 1810, by Peron & Lesueur. The genus Laomedea Lamouroux, 1812 : 184 was proposed to accommodate twospecies, Sertularia dichotoma Linnaeus, 1758, and 5. spinosa Linnaeus, 1758. Thefirst-named had, however, previously been removed from Sertularia under the nameObelia sphaerulina Peron & Lesueur, iSioa, while the second species is currentlyreferred to the bryozoan genus Vesicularia Thompson, 1830 (Prenant & Bobin, 1 Although dated 1809, Peron & Lesueur's paper was not published until January 1810 (Sherborn,1929 : 4455). They later published a second designation of Obelia, dated May 1810, but this work waslargely a reprint of part of the earlier paper (Pe"ron & Lesueur, iSiob). A footnote in the first paperstates that the plates did not appear with it, and although Lesueur (1811) later published some plates,those from the iSioa paper apparently remain unpublished. 254 p - F - s - CORNELIUS 1956 : 276). S. dichotoma Linnaeus, 1758, is here selected as type species of thegenus Laomedea which can thus be considered a junior synonym of Obelia. The nextavailable name in place of Laomedea auct. appears to be Campanularia Lamarck,1816, which originally included the four species Sertularia verticillata Linnaeus,1758, S. volubilis Linnaeus, 1758, S. syringa Linnaeus, 1767, and 5. dichotomaLinnaeus, 1758. Although Nutting (1915 : 28) made Sertularia verticillata genotypeof Campanularia, Naumov (1960 : 249) later nominated S. volubilis Linnaeus, 1758(not sensu Ellis & Solander, 1786), as type-species of Campanularia and madeSertularia verticillata genotype of a new genus, Verticillina Naumov, 1960 : 269.Millard (1966 : 477) commented on this confusion and nominated S. volubilis sensuEllis & Solander, 1786 : 51 as type-species of Clytia Lamouroux, 1812 : 184. Sheshowed the correct identification of this type-species to be Medusa hemisphaerica 1Linnaeus, 1767 : 1098. However, this species had previously been nominatedtype-species of the medusoid genus Thaumantias Eschscholtz, 1829 : 102, by Forbes(1848 : 41) and Thaumantias can be regarded a junior objective synonym of Clytia.The status of these genera and their genotypes will be elucidated further by referenceto the International Commission on Zoological Nomenclature. The availability of the generic name Eucope Gegenbauer (1856) was discussed byRees (1939)- The genus Schizocladium Allman (1871) was regarded as a junior synonym ofObelia by Bedot (1910 : 470). The sole included species is reduced to a synonym ofObelia dichotoma in the present review (p. 272) . The genus Obelaria Haeckel (1879) was proposed as a nom. nov. for the hydroidstage of Obelia, of which it is a junior objective synonym, as recognized by Bedot(1912 : 326). The two subgenera Obeletta Haeckel (1879) and Obelissa Haeckel(1879) were introduced to accommodate taxa defined from the medusa stage.Although thus outside the scope of the present survey they nevertheless at presentseem superfluous and can be regarded as junior synonyms of Obelia. The genus Monosklera von Lendenfeld, i885b : 910, was synonymized underObelia by Bedot (1916 : 152). The sole included species is regarded as a synonymof Obelia geniculata in the present review (p. 273), following Vanhoffen (1910). Three species are recognized from the hydroid stage, 0. bidentata Clarke, 1875,0. dichotoma (Linnaeus, 1758) and O. geniculata (Linnaeus, 1758). They canusually be identified using the characters shown in Table I, but occasional specimensoccur with characters apparently intermediate between 0. dichotoma and 0. genicu-lata. Usually such specimens are examples of 0. dichotoma with a slight thickeningof the internodal perisarc, and can nevertheless be identified from the charactersshown. 1 Although this species name has been attributed to Gronovius (1760 : 38), his usage was not strictlybinominal (Millard, 1966 '.477). HYDROID SPECIES OF OBELIA 255 256 P. F. S. CORNELIUS MORPHOLOGICAL AND OTHER VARIATIONS THE large number of species of Obelia described from the hydroid stage reflects thehigh degree of morphological variation present. Although between-colony variationis readily apparent and many characters are inconstant in expression, neverthelesssuch characters have been freely used in diagnosing new species. That muchvariation is phenotypic has long been suspected (Mayer, 1910 ; Hammett, 1943)but does not seem to have been given due attention in systematic accounts of thegenus. All of the new synonymies proposed here result from the adopting of widerspecific limits to take account of this variation. The characters used in definingthe specific limits are now discussed. COLONY SIZE. Crowell & Wyttenbach (1957) indicated that there are probablyno endogenous factors operating to limit colony-size in Campanularia flexuosa(Hincks, in Alder, 1856), and it would seem likely that this also holds in Obelia. Itfollows that colony-size is probably of little or no systematic value. They suggestedthat colonies of C. flexuosa continue their apical growth until natural breakageoccurs. It is relevant that the long colonies referred to 0. longissima by some authors(pp. 256-266) are found in places of limited wave-action, such as broad estuaries[the late D. N. Huxtable (personal communication) ; personal observation], whileshorter colonies hitherto regarded as more typical of 0. dichotoma occur intertidallyand sublittorally where wave-action is greater. Colonies of intermediate lengths occurand on present evidence it appears that colony-size in 0. dichotoma s. lat. is relatedto wave-action. The status of 0. longissima is discussed further below (p. 271). 0. geniculata colonies at a single locality in Massachusetts were usually shorterthan 25 mm (Hammett, 1943). Ralph (1956), however, demonstrated a correlationbetween length and latitude in populations between 35 S and 41 S around NewZealand. Cold-water specimens were up to 40 mm long, while those from thewarmest places measured only 5 mm. Later Ralph & Thomson (1968) showed thatin Wellington harbour colonies formed in the austral winter were longer than thoseformed in the summer. Nevertheless, it appears that variationin colony-size in0. geniculata is less than in 0. dichotoma. Information on 0. bidentata is apparentlylacking. MONOSIPHONIC VERSUS POLYSiPHONic STEMS. Old stems of 0. bidentata areinvariably polysiphonic, those of 0. dichotoma are usually monosiphonic and those of0. geniculata always so. BRANCHING OF COLONY. Several species of Obelia have been proposed on the basisof the pattern of branching of the colonies. However, this appears not to becorrelated with other morphological features. Similarly the length of the branchesseems unimportant. The influence of habitat on branching is poorly documented,but Ralph & Thomson (1968) reported that low temperatures induced branching in0. geniculata. Occasional dichotomously branched specimens occur in 0. dichotoma,but it is questionable whether such specimens are truly dichotomous. INTERNODES. Inter-colony variation occurs throughout the genus in internodelength, breadth, length : breadth ratio, curvature, amount of asymmetric internalperisarc thickening, angle of flexure and number of annulations. Of these only the HYDROID SPECIES OF OBELIA 257 amount of perisarc thickening seems to have systematic value, being pronounced andasymmetric (although variable) in 0. geniculata and usually slight and symmetricalin the other two species. Occasional specimens of 0. dichotoma occur in which theinternodal perisarc is slightly thickened on one side, but such specimens can beidentified by their hydrothecae. Asymmetric internodal thickening is furtherdiscussed below (pp. 271, 277). DARKENING OF PERISARC. In large colonies of 0. dichotoma darkening of theinternodal perisarc proceeds with age, older parts being dark brown, grading tobrown, horn-coloured or transparent in younger parts of the colony. Colonies of0. geniculata do not become so dark, while the limited material of 0. bidentataavailable suggests that no darkening occurs. Histochemical studies by Knight(1970) showed that in the normal perisarc tanning process of Campanulariaflexuosaseveral of the biochemical precursors of melanin are formed and it is possible thatthis is the dark pigment in 0. dichotoma. HYDROTHECA. Many hydrothecal characters appear to be systematically sound,and they are here regarded as important in defining specific limits within the genus.The hydrothecal rim may be bimucronate or mucronate as in 0. bidentata (Fig. 2),even, sinuous or castellate as in 0. dichotoma (Fig. 4) or invariably even as in 0.geniculata (Fig. 5). The greater variability shown by 0. dichotoma is not understood,but colonies otherwise similar may differ in having one type of rim or another.The factors involved in the rupture of the embryonic operculum and the consequentformation of the hydrothecal rim are only partly known (Knight, 1965) and it wouldseem unwise to attach systematic importance to rim-variation until more is knownof the factors responsible. The shape of the hydrotheca appears to be variable in length : breadth ratio, inthe angle between the sides as seen in median vertical section and in the degree towhich the basal region is curved inwards. Hydrothecae of 0. bidentata are tubularwith a curved basal region while those of 0. geniculata usually have walls curvedthroughout their length (Figs 2, 5). Those of 0. dichotoma are more variable, beingtubular to conical (Fig. 4), with or without a curved basal region. A variableamount of thickening of the basal region of the hydrothecal wall occurs in 0. genicu-lata but apparently not in the other two species. The possession of an oblique hydrothecal diaphragm has been used to delimitspecies otherwise resembling 0. dichotoma, but it appears to be a variable characterand such species are not here regarded as valid (see p. 272). In 0. geniculata,however, the diaphragm is always transverse while in 0. bidentata it is alwaysoblique. It has been noted that oblique diaphragms seen in optical section can,from certain angles, appear transverse (Mammen, 1965). HYDROTHECAL PEDICEL. Although the number of annulations of the hydro-thecal pedicel has been reported as constant at 4-6 in 0. geniculata (Hammett,1943), in 0. dichotoma the number varies between the approximate limits 2-10 andin 0. bidentata between 3-26 (Mammen, 1965). In both there is often a smoothcentral portion. There seems to be no indication at present whether this variationis phenotypic or genotypic. 258 P. F. S. CORNELIUS GONOTHECA. Gonotheca shape is similar in the three species and with theexception of the terminal region is approximately constant. In 0. bidentata andto a lesser extent in 0. dichotoma gonothecae with apparently truncate ends havebeen described, there being only a trace of a raised central aperture (Fig. 2 ; pp.263-264). NEMATOCYSTS. Previous work, and also that reported here, suggests thatnematocysts do not provide useful specific criteria in Obelia. a. 0. dichotoma and 0. geniculata. The lengths of fresh, undischarged microbasicmastigophores from living hydranth tentacle-tips of the two species are shown inTable 2. The sizes varied to the extent that the range of measurements obtained TABLE 2 Measurements of undischarged nematocysts from the tips of hydranth tentacles of Obelia spp., 0-2 [zm SPECIES LOCALITY LENGTH WIDTH REMARKS S.E. O. geniculata Cornwall (1973.9.24.2)Cornwall (1973.9.24.3) O. dichotoma Devon (1973.7.23.1)Devon (1973.9.24.4) O. bidentata Norfolk (1953.11.16.1) 10 5-6 0-0897 specimen resembledO. longissima sensuAlderAll 1-5 Formalin-preserved n= number in sample, ~x= mean, S.E. = standard error. 'Fresh' nematocysts are those taken fromliving hydranths. from one species fell within the range recorded from the other. Application ofStudent's /-test showed nematocyst-length to be similar in the two 0. geniculatapopulations (p <o-ooi), but dissimilar in the two groups of 0. dichotoma (p >o-32).Two types of nematocyst have previously been recorded from 0. geniculata.Ito & Inoue (1962) reported microbasic mastigophores (measuring 4-9-5 -2 /mix1-3-1-5 /mi undischarged) from the hydranths, while Weill (1934^ identifiedbasitrichous isorhizas (length 5 /mi) from the tentacles of both hydranth andmedusa. Only the latter type has been recorded from 0. dichotoma, from themedusa (Westfall, 1966, as 0. longissima ; measurements not stated). Thus thepresent recording of microbasic mastigophores in 0. dichotoma is new. However,the identification of the two kinds of nematocyst depends on the presence or absenceof a butt (Fig. i), a feature which can be ascertained only in discharged capsules(Weill, i934a). Maybe a butt is not always present. It is certainly so small inObelia as to be difficult to observe with the light microscope. Possibly more exten-sive observations on Obelia from various localities will explain the reported occur-rence of both types. HYDROID SPECIES OF OBELIA 259 FIG. i. Obelia geniculata. Microbasic mastigophores from live hydranths. SW. England( I 973-9-24-2). (a) Discharged. Capsule, 7 (im ; butt, 6 (xm ; barbs c. 1-5 (zm ; thread,63 (Jim (not all shown). Note distinct butt at base of thread, (b-d) Undischarged,different scales. Lengths, 6-8 (jim ; breadths, 2 (j.m (see Table 2 for standard error).One side is slightly flatter than the other, and the tip of the capsule inclines to one side. b. 0. bidentata. Measurements of undischarged nematocysts from the tentacle-tips of formalin-preserved hydraiiths are shown in Table 2, living material beingunobtainable. The preserved nematocysts at least were similar in size to thosefrom live hydranths of the other two species. Identification was not attempted asno discharged nematocysts were seen, but the undischarged ones closely resembledthose of the other species and may prove to be of the same kind. BIOLUMINESCENCE. Although Hincks (1868) reported bioluminescence only in0. geniculata, it has now been reported in all three Obelia hydroids (Morin & Cooke,1971). SUBSTRATE. 0. geniculata has been widely recorded on the blades of laminarianand fucoid algae, whereas 0. dichotoma occurs usually on animal and inert substratesand less frequently on algae (Hincks, 1868 ; Hammett, 1943 ; Barrett & Yonge,1958 ; Fey, 1969). Manton (1942) examined the attachment of the hydrorhiza ofObelia sp. to an unspecified substrate, and found that the coenosarc was not involved.Sections cut during the present study along the stolons of 0. geniculata(1973.7.23.2-3) and into the algal substrate showed no tissue connections betweenhydroid and alga. It seems likely, therefore, that the growth-form of O. geniculatais not related to substrate. The occasional occurrence of each species on the sub-strate more usual for the other is additional evidence that substrate does not influencemorphology. The substrate-preference of 0. bidentata is less well known but issaid to be for inert solid substrates and also sand (Vervoort, I946a) to which neitherof the other^two species attaches directly. MEDUSA GENERATION. The medusae of the three hydroid species, althoughvariable, cannot at present be distinguished (see pp. 278-279). 260 P. F. S. CORNELIUS Obelia bidentata Clarke, 1875(Fig. 2) Obelia bicuspidata Clarke, 1875 : 58, pi. 9, fig. i ; Bedot, 1912 : 326 ; Nutting, 1915 : 80, pi. 20,figs 5-6 (= O. bidentata Clarke) ; Bedot, 1916 : 160 ; Bedot, 1918 : 195 ; Bedot, 1925 : 298-299 (= O. bidentata Clarke) ; Fraser, 1944 : 153-154, pi. 27, fig. 125 (= O. bidentata Clarke ;L. spinulosa var. minor Leloup) ; Deevey, 1950 : 343 (= O. oxydentata Stechow ; L. spinulosavar. minor Leloup) ; Leloup, 1952 : 157, fig. 89 (= C. spinulosa Bale) ; Vannucci, 1954 : 108-110, pi. 2, figs 2-7, 9-10 (= O. bidentata Clarke ; C. spinulosa Bale ; O. bifurca Hincks ;Obelia sp. Clarke; O. oxydentata Stechow) ; Millard, 1958 : 174; Mammen, 1965 : 11-13,figs 37-38 ; Millard & Bouillon, 1973 : 56 (= G. longicyatha : Jarvis). Obelia bidentata Clarke, 1875 : 58-59, pi. 9, fig. 2 ; Pictet, 1893 : 25-26, pi. i, figs 20-21 ;Jaderholm, i9O4a : 270 271 (= O. bicuspidata Clarke); Jaderholm, 190413 : vii (= O.bicuspidata Clarke) ; Jaderholm, 19050 : 17 ; Mayer, 1910 : 254 [= O. bicuspidata Clarke ;? = O. austrogeorgiae Jaderholm ; (see below, p. 280)] ; Bedot, 1912 : 326 ; Billard, 1912 :463, fig. 2 (= O. bicuspidata Clarke); Bedot, 1916:160; Bedot, 1918:195; Kramp,1961 : 162 ; Teissier, 1965 : 16 ; Fey, 1969 : 393. ? Obelia longicyatha Allman, 1877 : 10, pi. 7, figs 4-5 ; (see below, p. 264). Campanularia spinulosa Bale, 1888 : 756-757, pi. 12, figs 5-7. Obelia andersoni Hincks, 1889:132-133, pi. 12, figs 2-4; Thornely, 1904:113; Ritchie,1910 : 810 ; [syn. nov.]. Obelia bifurca Hincks, 1889 : 133, pi. 12, fig. i ; Mayer, 1910 : 494 ; [syn. nov.]. Gonothyrea longicyatha Thornely, 1899 : 454-455, pi. 44, figs 4, 4a (non O. longicyatha Allman,1877). Obelia corona Torrey, 1904 : 14, figs 5-6 ; Nutting, 1915 : 79, pi. 20, figs 1-2 ; [syn. nov.]. Obelia sp. Clarke, 1907 : 10-12, pi. 5, figs 5-7. Obelia bifurcata Thornely, 1908 : 81-82, pi. 9, fig. 2 (nom. nov. pro O. bifurca Hincks). Laomedea bidentata : Babid, 1913 : 284-286, fig. I (= O. bicuspidata Clarke). Obelia multidentata Fraser, 1914 : 154, pi. 17, fig. 56 ; Fraser, 1937 : 89-90, pi. 18, fig. 93 ; [syn.nov.]. Obelia oxydentata Stechow, 1914 : 131-132, fig. 7 (nom. nov. pro Obelia sp. Clarke) ; Stechow,1919 : 50 ; Vannucci Mendes, 1946 : 555-556, pi. 2, fig. 22 ; Hirohito, 1969 : 9-10, fig. 8a-b. Gonothyrea bicuspidata : Stechow, I9i9:5o-5i(= O. bidentata Clarke ; G. longicyatha Thornely) ;Vannucci Mendes, 1946 : 556-557, pi. 3, fig. 23 [= O. bidentata Clarke ; ? = O. austrogeorgiae :Nutting, 1915 ; (see below, p. 280)]. Obelia longa Stechow, ig2ia : 221-223, fig. i ; Stechow, 1925 : 436-437, figs i2b, 13 ; [syn.nov.]. Gonotha longicyatha (sens. Thornely) : Jarvis, 1922 : 336. Clytia longitheca Hargitt, 1924 : 484, pi. 3, fig. 9 ; [syn. nov.]. Obelia longitheca Hargitt, 1924 : 484-485, pi. 3, fig. 10 ; [syn. nov.]. Obelia attenuata Hargitt, 1924 : 486, pi. 3, fig. n ; [syn. nov.]. Obelia spinulosa : Billard, 1927 : 333-334, fig- 2. Laomedea bicuspidata var. picteti Leloup, 1932 : 151-153, pi. 17, figs 4, 4d, text-fig. 19. Laomedea spinulosa var. minor Leloup, 1932 : 155-158, pi. 17, figs 6, 6a, text-figs 24-25. Laomedea bicuspidata: Hummelinck, 1936 : 53-57, fig. 8a-v (= O. bidentata Clarke ; C. spinulosaBale ; G. longicyatha Thornely ; Obelia sp. Clarke ; O. oxydentata Stechow ; L. bicuspidatavar. picteti Leloup ; L. spinulosa var. minor Leloup) ; Vervoort, i946a : 298-300, fig. I32a-f( = O. bidentata Clarke ; C. spinulosa Bale ; G. longicyatha Thornely ; O. oxydentata Stechow) ;Vervoort, I94&b : 344-345, fig. loa-b ; Hamond, 1957 : 312-313, figs 20-21 ; Vervoort,J 959 ' 3 I 5 ' Vervoort, I972a : 92-93, fig. 26d (= O. bidentata Clarke; C. spinulosa Bale;L. spinulosa var. minor Leloup ; Obelia sp. Clarke ; O. oxydentata Stechow). ? Clytia longicyatha (sens. Allman) : Fraser, 1944 : 142, pi. 25, fig. 114 ; (see below, p. 264). Clytia longicyatha (sens. Allman) : Rees & White, 1966 : 276. HYDROID SPECIES OF OBELIA 261 Laomedea bicuspidata var. tennis Vervoort, 19465 : 345-346, fig. ice (nom. nov. pro L. spinulosa var. minor Leloup). ? Laomedea longicyatha (sens. Allman) : Vervoort, 19465 : 343-344.Laomedea (Obelia) bicuspidata: Vervoort, 1968 : 19-21, fig. 7 (= O. bidentata Clarke; C. spinulosa Bale ; Obelia sp. Clarke ; C. spinulosa var. minor Leloup).Laomedea (Obelia) longicyatha (sens. Allman): Vervoort, 1968:21-22, fig. 8; Vervoort, I972a : 93.non Clytia longicyatha (sens. Allman) : Pictet, 1893 : 28-29, pi- 2 , figs 22-23 ( Clytia sp., see below, p. 264). TYPE LOCALITY. Greenport, Long Island, New York, U.S.A., on wharf piles(Clarke, 1875). DIAGNOSIS. Obelia hydroid usually with branched, erect hydrocaulus ; poly-siphonic basally, internodes straight, narrow, lacking internal thickening of perisarcand not usually strongly tanned ; rim of hydrotheca with a variable number ofcusps, usually bimucronate. DESCRIPTION. Mature colony comprising several erect, sometimes flexuose,polysiphonic stems up to 350 mm, alternate lateral hydrocauli bearing the hydro-thecae ; basal hydrorhiza on sandy substrates a tangled mass of stolons. Lateral a Fig. 2. Obelia bidentata. (a) Nigeria (1966.10.6.1). Part of hydrocaulus and ripegonotheca with aperture only slightly raised. Scale = 500 (im. (b) Sierra Leone(1966.10.7.5). Bimucronate hydrothecal rim slightly irregular. Scale = 50 [xm. (c)Nigeria (1966.10.8.117). Hydrothecal rim with indentations of similar depth (? atypical).Scale = 50 [xm. 262 P. F. S. CORNELIUS hydrocauli delicate, slightly flexuose ; internodes long, annulated proximally,with lateral process distally to which hydrothecal pedicel is attached. Pedicelringed throughout or with central smooth portion ; hydrotheca 1-3 times long asbroad, bell-shaped, slightly asymmetric, sometimes having folds in the hydrothecalwall between cusps running proximally from the rim ; diaphragm oblique (canappear transverse in optical section) ; rim with 10-20 bimucronate cusps (rarely,simply cusped ; Fig. 2a-c) ; hydranth undescribed. Gonotheca usually an invertedcone with raised aperture ; occasionally truncated with broad aperture, lackingraised structure. Measurements - see Table 3. TABLE 3 Measurements of the hydroid stage of Obelia bidentata in \j.m. NIGERIA SIERRA LEONE SOUTH YEMEN NORFOLK, ENGLAND (1966.10.6.1 ; (1966.10.7.8) (1966.11.15.2) (1953.11.16.1)Fig. aa) HYDROTHECA Length (diaphragm to tips of cusps) 400-470 340-490 320-430 380-430 Breadth at rim 210-270 120-230 180-280 200-220 HYDROTHECAL PEDICELS Length 070-240 070-580 060-410 070-240 INTERNODES Length 450-680 400-570 320-480 480-710 Maximum breadth 080-130 070-080 070-090 080-120 Length/breadth ratio c. 6 5-6 5-6 5-6 GONOTHECA Length 550-600 Maximum breadth 220-260 MATERIAL EXAMINED. Atlantic Ocean - Greenport, Long Island, New York,U.S.A., 5 August 1874, several infertile hydrocauli in spirit, syntypes, Yale PeabodyMuseum of Natural History no. 3119. Thimble Island, Branford, Connecticut,U.S.A., 23 September 1874, three fragments in spirit, syntypes of 0. bicuspidataClarke, Yale Peabody Museum of Natural History no. 7265. Hunstanton, Norfolk, England, 26 September 1953, several large colonies inspirit, coll. R. Hamond, I953.ii.i6.! 1 (Table 2). Shelmess, Isle of Sheppey, Kent,England, strandline, 23 September 1973, two abraded colonies in spirit, coll. P. F. S.Cornelius, 1973.9.24.1. Zeeland Province, Netherlands, 20 August 1946, several colonies in spirit, LeidenRijksmuseum van Natuurlijke Historic no. 3687. Monte Brazil W., Terceira I., Azores, 29 m, 27 July 1959 and August 1959,several colonies in spirit, coll. Imperial College Azores Expedition, 1962.1.15.17, 21(Rees & White, 1966, as Clytia longicyaiha ; see p. 264). 1 Registered numbers of this format refer to British Museum (Natural History) collections, unlessotherwise stated. HYDROID SPECIES OF OBELIA 263 Trinidad, West Indies, August 1966, three microslides of fragments, coll. J. H.Wickstead, 1966.11.11.1. Port Harcourt, Nigeria, n July - 14 November 1957, 26 microslides of fragments,coll. H. G. Stubbings, 1966.10.8.14, 31, 44, 55, 56, 58, 70, 74, 92, 93, 95, 112, 117,119, 120, 124 (Fig. 2c). Lagos, Nigeria, 1957, four microslides of fragments, coll.M. B. Hill, 1958.3.1.1-4. Lagos Harbour, Nigeria, January 1959, i m, one micro-slide of fragments, coll. M. B. Hill, 1966.10.6.1 (Fig. 2a). Bunce Island, Sierra Leone, 19 February 1955, seven microslides of fragments,coll. A. Longhurst, 1966.10.7.5-11 (Fig. 2b). Indo-Pacific Ocean - Sapper Bay, Aden, South Yemen, 17 August 1966, coll. K. W.England, one microslide of fragments, 1966.11.15.2. Amoy, China, December 1925, one colony in spirit, coll. C. Ping, 1926.3.17.36. On Hong Kong to Manilla cable, 2057' N, ii523' E, c. 200 m, 21 February1929, one microslide of fragments, coll. British East India Company, 1929.4.18.20. OTHER MATERIAL RECORDED. North Atlantic - West European waters fromHelgoland (Kramp, 1961), Netherlands, Belgium, northern France (Vervoort, I94&a ;Leloup, 1952 ; Teissier, 1965 ; Fey, 1969); Mediterranean Sea (Vervoort, I946b),Algeria (Picard, 1955), Azores (Rees & White, 1966), Ghana (Buchanan, 1957),tropical W. Africa (Vervoort, 1959). North American coast from Casco Bay,Maine, south to Caribbean and Panama (Fraser, 1946 ; Vervoort, i946b, 1968). South Atlantic - South American coast from Brazil (Fraser, 1946 ; VannucciMendes, 1946) and near Tierra del Fuego (Vervoort, i972a). Pacific - San Francisco Bay (Fraser, 1937, 1946), Hawaii (Vervoort, I946b),Japan (Hirohito, 1969, as 0. oxydentata), New Britain Island (Vervoort, i946b),Port Jackson, New South Wales, Australia (Bale, 1888, as Campanularia spinulosa). Indian Ocean - Several localities off India and Malaya (Vervoort, ig46b ; Mam-men, 1965), Kerguelen Island (Stechow, 1925, as 0. longa), Natal (Millard, 1958),Mozambique and Seychelles (Millard & Bouillon, 1973, 1974). DISTRIBUTION. Continental shelf depths in tropical, sub-tropical and sometemperate seas in both northern and southern hemispheres. Records furthest fromthe equator are Helgoland (54 N), South Georgia and Tierra del Fuego (both 54 S).Reported unrecorded from Argentina (Vervoort, I972a), New Caledonia (Redier,1966), Tasmania (Hodgson, 1950), South Australia (Blackburn, 1942), New Zealandand the Chatham Islands (Ralph, 1957, 1961), south and west coasts of the Republicof South Africa (Millard, 1957). REMARKS. The number of bimucronate cusps on the hydrothecal rim may varybetween 10 and 20 (Nutting, 1915 ; Mammen, 1965), although at any one localitythe range of variation is less. Most specimens have alternating deep and shallownotches around the hydrothecal rim but in some the gaps are of almost equal depthand the bimucronate condition is obscured (Fig. 2b-c). Two kinds of gonothecae have been recorded. The more usual kind is typicalof Obelia, with a tubular aperture (Nutting, 1915 ; Hamond, 1957 ; Hirohito, 1969)while the other is truncated distally and opens directly (Babic, 1913 ; Fraser,1937 ; Mammen, 1965). Possibly a reduced tubular aperture was overlooked by 264 P. F. S. CORNELIUS authors describing a truncate gonotheca. Medusa release was recorded fromnormal gonothecae by Hamond, while a BM(NH) specimen shows medusaedeveloping within an apparently truncate gonotheca (Fig. 2a). The two nominal species 0. bicuspidata and 0. bidentata were first described onthe same page by Clarke (1875). They were synonymized under 0. bidentata byJaderholm (i904a, b, 19053.), whose usage as first reviser was followed by Mayer(1910), Billard (1912) and Babic (1913). Nutting (1915), however, used 0. bicuspidataas the senior synonym, although including Jaderholm's I904a paper in his synonymy.Since then bicuspidata has been used more frequently than bidentata but Jaderholm'susage gives bidentata priority. Two varieties of the species have been described. Laomedea spinulosa var.minor Leloup, 1932, was erected to accommodate forms differing only in the heightof the colony. Vervoort (i972a), however, placed var. minor Leloup together withL. bicuspidata var. tenuis Vervoort, I946b (a new name for Leloup's variety), in thesynonymy of 0. bicuspidata. The second variety, 0. bicuspidata var picteti Leloup,1932, was distinguished on specimens with long gonothecae. As such variation iscommon and has no apparent taxonomic value, the separation is not upheld here. There is some confusion concerning Obelia longicyatha Allman, 1877, and anothernominal species of the same name, 0. longicyatha (Thornely, 1899). The typespecimens of neither species could be located. Allman's species, described frominfertile material, was said to have long pointed cusps on the hydrothecal rimreminiscent of Clytia, to which genus the species has been referred by several authors(references in Fraser, 1944). Nevertheless, the form of the colony as illustrated byAllman equally suggests 0. bidentata, and the affinities of the species remain unclear.Pictet (1893) referred to Allman's species material with developing medusae clearlynot of Obelia type and referred his material to Clytia. Vervoort (i946b) describedunder Allman's species material identical with 0. bidentata except that the hydro-thecae were unusually long (900-950 /urn). The gonothecae were typical for thespecies. Vervoort (1968) later described other, infertile, material with hydrothecaenearer in size (580-620 /mi) to the measurements given here (320-490 /u,m) andillustrated an oblique hydrothecal diaphragm characteristic of 0. bidentata.Vervoort (ig72a) subsequently referred another infertile specimen to Allman'sspecies, distinguishing it from 0. bidentata on the basis of hydrothecal length.However, it seems unclear at present whether or not specimens of such dimensionsfall within the range of variation of 0. bidentata, and the identity of Vervoort 'smaterial is problematical. In contrast the material identified with Allman's speciesby Rees & White (1966) and re-examined here seems identical with 0. bidentata,having characteristic bimucronate hydrothecal rims and falling within the normalsize-range. The other species, 0. longicyatha (Thornely, 1899), was stated in theoriginal description to have such hydrothecal rims and Stechow (1919) and Hummel-inck (1936) justifiably referred it to 0. bidentata. Jarvis (1922) referred to Gonothalongicyatha specimens with bimucronate hydrothecal cusps and truncate gonothecaewhich seem also to have been typical 0. bidentata. Obelia andersoni Hincks, 1889, from the Mergui Archipelago, can be referred to0. bidentata. It was originally distinguished on the basis of a sharp demarcation HYDROID SPECIES OF OBELIA 265 between the cylindrical side of the hydrotheca and its inward-sloping base. Thornely(1904) found similar material in which there was no sharp demarcation, and thespecies appears invalid. The affinities of 0. austrogeorgiae Jaderholm, 1904^ are discussed on page 280. MEDUSAE. It has often been established that 0. bidentata releases a medusa(Hincks, 1889, as 0. andersoni ; Pictet, 1893, as 0. bidentata ; Thornely, 1899, asGonothyrea longicyatha ; Billard, 1927, as 0. spinulosa ; Fraser, 1944, as Clytialongicyatha sensu Allman ; the following, as 0. bicuspidata : Vervoort, I946a,b ; Vannucci, 1954 ; Hamond, 1957 ; Mammen, 1965 ; and Fey, 1969, as 0.bidentata}. However, the medusae were apparently not recorded as resemblingthose of the other Obelia species until Billard (1927) identified them in the gonotheca.Subsequently Hamond (1957) and Mammen (1965) confirmed the resemblance byobserving newly liberated medusae. Adult specimens have not been described andthe best available description, of the young stage, is that by Mammen. The youngstage at least is very similar to that in the other two species. Obelia dichotoma (Linnaeus, 1758)(Figs 3 & 4) Sertularia dichotoma Linnaeus, 1758:812; Linnaeus, 1767:1312 (= S. longissima Pallas) ; Maratti, 1776:34; Rees, 1819 : unpaginated ; Dalyell, i836a:9i-92, 94, fig. i; Dalyell, i836b : 84-85, 87, fig. i. Sertularia longissima Pallas, 1766 : 119-121 (nom. nov. pro 5. dichotoma Linnaeus).Sertolare genicolata Cavolini, 1785 : 205, pi. 8, figs 1-4 (lapsus pro Sertularia geniculata Linnaeus) .Laomedea dichotoma: Lamouroux, 1812:184; Johnston, 1838:150-151, pi. 22, figs 1-2 (= Sertularia longissima Pallas); Johnston, 1847:102-103, 119, pi. 26, figs 1-2 (= S. longissima Pallas) ; Alder, 1857 : 121 (= Campanularia gelatinosa : van Beneden) ; Hincks, 1861 : 258. Sertularia geniculata : Sprengel, 1813 : 95-97, pi. 8, figs 3-4.Campanularia dichotoma : Meyen, 1834 : 193-195, pi. 30, figs 1-4, pi. 31, fig. i ; [non C. dichotoma : Grant, 1826 : 150-156 (= Laomedea flexuosa Hincks, in Alder, 1856)].Campanularia maior Meyen, 1834 : 196-197, pi. 32, figs 1-4 ; Bedot, 1905 : 53 ; [syn. nov.].Campanularia brasiliensis Meyen, 1834:198, pi. 32; Nutting, 1915:77, pi. 18, figs 8-9; [syn. nov.].Campanularia cavolinii Deshayes & Edwards, 1836 : 133 (nom. nov. pro Sertularia geniculata : Cavolini) ; [syn. nov.].Campanularia caulini Chiaje, 1841 : 143 (unjustified emendation of C. cavolinii Deshayes & Edwards). Sertularia cavolinii : Kolliker, 1843 : 81 ; [syn. nov.].Campanularia gelatinosa : van Beneden, 1844 : 33-34, pis 1-2 ; Maitland, 1876 : 13 (= Obelia sphaerulina Peron & Lesueur, iSioa) ; (see p. 279).Laomedea longissima : Alder, 1857 : 121-122 ; Hincks, 1861 : 259.Obelia commissuralis McCrady, 1857 : 197-198, pi. n, figs 5-7 ; Agassiz, 1862 : 315-321, pis 33~34> n g s 10-21 ; Agassiz, 1865 : 91-92, figs 134-135 (= Laomedea dichotoma: Leidy ; L. gelatinosa : Gould, Stimpson) ; Norton, 1896 : 291-296, figs 1-12 ; Nutting, 1915 : 83, pi. 21, figs 1-5 ; Berrill, 1949 : 235-264 ; Vannucci, 1951 : 80-81, pi. 2, figs 8-9 ; Mammen, 1965 : 14-15, fig. 41 (? = O. hyalina Clarke) ; [syn. nov.].Laomedea divaricata McCrady, 1857 : 195-196 ; Agassiz, 1865 : 91.Eucope parasitica Agassiz, 1865 : 87 ; [syn. nov.]. 266 P. F. S. CORNELIUS Eucope pyriformis Agassiz, 1865 : 88-89 ', Mayer, 1910 : 247 (? = Laomedea divaricataM.cCr3.dy) ; [syn. nov.]. Eucope articulata Agassiz, 1865 : 89-90, figs 130-131. Campanularia flabellata Hincks, 1866 : 297 (nom. nov. pro C. gelatinosa : van Beneden).Obelia longissima : Hincks, 1868 : 154-155, pi. 27 (= Campanularia gelatinosa : van Beneden) ; Russell, 1953 : 303, fig. 1850 ; Naumov, 1960 : 263-264, figs 149-151 ; Naumov, 1969 : 284, figs 149-151.Obelia dichotoma : Hincks, 1868 : 156-157, pi. 28, fig. i, la-b ; Mayer, 1910 : 245-246, 248, pi. 30, figs 1-2 ; text-figs 125-127 (= Eucope articulata Agassiz ; O. sphaerulina : Haeckel ; ? O. australis von Lendenfeld ; O. rhunicola Billard) ; Bedot, 1925 : 301 (= O. rhunicola Billard) ; Russell, 1953:303, fig. 1850 ; Millard, 1966:483 (= O. dubia : Vanhoffen ; Campanularia obtusidens Jaderholm).Obelia flabellata : Hincks, 1868 : 157-158, pi. 29 ; Nutting, 1915 : 84-85, pi. 22, figs 3-4 ( = O. plana : Mayer ; see footnote, p. 271).Obelia plicata Hincks, 1868 : 159, pi. 30, figs i, la ; Nutting, 1915 : 78, pi. 19, figs 5-6 ; [syn. nov.]. Schizocladium ramosum Allman, 1871 : 18-21, pi. 2, figs 1-8 ; [syn. nov.].Obelia pygmaea Coughtrey, 1876 : 25, pi. 3, fig. 3 ; Ralph, 1957 : 832 [? = O. longissima (Pallas)] ; [syn. nov.]. Obelia hyalina Clarke, 1879 : 239, 241-242, pi. 4, fig. 21 (non Gonothyrea hyalina Hincks, 1866).Obelia adelungi Hartlaub, 1884 : 164-165, text-fig, i ; [syn. nov.].Obelia helgolandica Hartlaub : 1884 : 165-167, text-fig. 2 ; [syn. nov.].Obelia australis von Lendenfeld, i885a : 604, 630 ; Bale, 1888 : 753-754, pi. 12, figs 1-2 ; Ralph, 1957 : 830, fig. 4a-h ; Mammen, 1965 : n. Obelia angulosa Bale, 1888 : 752-753, pi. 12, fig. 3 ; Mayer, 1910 : 257 ; [syn. nov.].Obelia chinensis Marktanner-Turneretscher, 1890 : 209-210, pi. 3, figs 6-7 ; Mayer, 1910 : 242 (? = O. plana : Haeckel) ; [syn. nov.]. Obelia arruensis Marktanner-Turneretscher, 1890 : 210, pi. 3, fig. 8 ; [syn. nov.].Obelia nigrocaulus Hilgendorf, 1898 : 203-204, pi. 17, figs I, la ; Bale, 1924 : 230.Obelia gracilis Calkins, 1899 : 353-354, pi. 3, figs 13, I3a-c, pi. 6, fig. i3d [non Laomedea gracilis Sars, 1850 = Campanularia pelagica van Breemen, 1905 (Vervoort, I946a : 285)] ; Nutting, 1915 : 78, pi. 19, figs 2-4 ; Hargitt, 1927 : 504-505 ; Ling, 1938 : 183 ; Blanco, 1967 : 130-134, figs 1-16; [syn. nov.].Obelia surcularis Calkins, 1899 : 355, pi. 3, figs 14, I4a-b, pi. 6, fig. I4C ; Nutting, 1915 : 84, pi. 22, figs 1-2 ; [syn. nov.].Obelia fragilis Calkins, 1899 : 355-356, pi. 3, figs 15, i5a-b, pi. 6, fig. I5C ; Nutting, 1915 : 87, pi. 33, fig. 6 ; [syn. nov.].Obelia griffini Calkins, 1899 ' 357, pi- 4, figs 18, i8a-c, pi. 6, fig. i8d ; Nutting, 1915 : 87, pi. 23, figs 4-5 ; Vannucci Mendes, 1946 : 552-553, pi. 2, figs 16-17 ; [syn. nov.].Obelia rhunicola Billard, igoib : 522-523.Obelia borealis Nutting, 1901 : 174, pi. 19, figs 4-6 ; Nutting, 1915 : 85, pi. 22, figs 5-7 ; [syn. nov.].Obelia dubia Nutting, 1901 : 174, pi. 20, fig. i ; Nutting, 1915 : 77, pi. 19, fig. i ; Mayer, 1910 : 248 (? = O. dichotoma : Hincks) ; Vanhoffen, 1910 : 307-308, fig. 27 ; Bedot, 1925 : 302 (= Campanularia obtusidens Jaderholm).Obelia solowetzkiana Schydlowsky, 1902 : 123-125, pi. 3, figs 20-22 (nom. nov. pro O. flabellata : Schlater) ; Jaderholm, 1909 : 63 ; [syn. nov.].Campanularia obtusidens Jaderholm, i905a : 2, pi. i, fig. i. Obelia congdoni Hargitt, 1909 : 375-376 (nom. nov. pro O. hyalina Congdon) ; [syn. nov.].Obelia articulata : Mayer, 1910 : fig. 126. Obelia pyriformis : Mayer, 1910 : 240, 247, fig. 128 (= Laomedea divaricata McCrady).Obelia piriformis Bedot, 1910 : 342 (lapsus pro pyriformis). Obelia undotheca Stechow, ig23a : 4 ; Stechow, ig23b : 115-117, fig. O ; [syn. nov.].Obelia nodosa Bale, 1924 : 230, fig. i ; Ralph, 1957 : 832, fig. 5i-k ; [syn. nov.]. HYDROID SPECIES OF OBELIA 267 Obelia coughtreyi Bale, 1924 : 230-231, fig. 2 ; [syn. nov.]. Obelia obtusidentata Bedot, 1925 : 302 (lapsus pro obtusidens). Obelia everta Hargitt, 1927 : 505, fig. 4 ; [syn. nov.]. Obelia alternata Fraser, 1938 : 35-36, pi. 8, fig. 38 ; [syn. nov.]. Obelia equilateralis Fraser, 1938 : 36-37, pi. 9, fig. 39 ; Fraser, 1944 : 157, pi. 28, fig. 128 ; [syn. nov.]. Obelia microtheca Fraser, 1938 : 37, pi. 9, fig. 40 ; [syn. nov.].Obelia tennis Fraser, 1938 : 38-39, pi. 9, fig. 42 ; [syn. nov.].Obelia racemosa Fraser, 1941 : 82, pi. 15, fig. 7 ; [syn. nov.]. Obelia irregularis Fraser, 1943 : 77 ; Fraser, 1944 : 162, pi. 29, fig. 132 ; [syn. nov.].? Obelia obtusidens : Fraser, 1944 : 163-164, pi. 29, fig. 134 ; (see p. 272).Obelia braziliensis Vannucci Mendes, 1946 : 553-555, pi. 2, figs 20-21 (lapsus pro brasiliensis) .Obelia biserialis Fraser, 1948 : 213, pi. 24, fig. 6 ; [syn. nov.].Laomedea (Obelia} dichotoma : Vervoort, 1959 : 315-316.Laomedea (Obelia) congdoni : Vervoort, 1968 : 23 (= O. hyalina Clarke ; L. sargassi Leloup). TYPE SPECIMEN AND TYPE LOCALITY. Ellis, 1755 I 21-22, pi. 12, fig. A, but not fig. a ; coast of SW. England. 1 Location of specimen unknown. DIAGNOSIS. Obelia hydroid usually with branched, flexuose, monosiphonichydrocauli ; internodes long, straight or slightly curved, without asymmetricperisarc thickening. Perisarc colourless in small specimens ; tanned to a horn-colour, brown or black in large colonies. Hydrothecal pedicels long, ringed, some-times with smooth central portion. Hydrothecae straight-sided, curving basally,often slightly flared at rim, which may be even, castellate or sinuous, frequentlywith minute longitudinal folds in the hydrothecal wall. DESCRIPTION. Main stems of colonies usually monosiphonic, flexuose,I0 ~35 mm > racemose hydrocauli with shorter lateral branches ; occasionalspecimens apparently dichotomous (see p. 256). Main hydrocauli frequentlytanned to a horn-colour, often brown or black, darker basally ; internodes long,nodes usually annulated, process supporting hydrothecal pedicel distal (Fig. 3).Hydrothecal pedicel long, annulated throughout or with smooth central portion ;hydrotheca bell-shaped, occasionally flared distally ; length 1-2 times greatestbreadth ; rim usually even but often sinuous or castellate (Fig. 4), frequently withminute longitudinal folds extending proximally from centre of each indentation ;ornamented rims easily abraded even ; diaphragm usually transverse but sometimes 1 In the absence of Linnaean material, type selection depends on the sole work cited by Linnaeus(1758), i.e. Ellis (1755). The woodcut figure is a good representation of part of an Obelia colony. Itshows four internodes with three attached gonothecae but no hydrothecae. The figure is thus com-patible with Linnaeus' diagnosis, which similarly does not mention hydrothecae ('Sertularia denticulisobsoletis, calycibus [= gonotheca] obovatis axillaribus, pedunculis intortis, caule dichotomo geniculato').Some of Ellis' hydroid material was eventually incorporated in Sir Hans Sloane's herbarium whichformed the basis of the biological collections of the British Museum; but there are no specimens ofO. dichotoma in that herbarium. Other Ellis hydroid material, some of it probably figured and henceeligible for typification, was until recently preserved in the Hunterian Museum of the Royal College ofSurgeons of England (Royal College of Surgeons of England, 1830, 1860; Harmer, 1931). Fragments ofO. dichotoma were evidently included (Royal College of Surgeons of England, 1860 : 137, as Laomedea)although it is not certain that they originated from Ellis. However, the bulk of the Ellis material wasdestroyed during the Second World War, and it is virtually certain that only a single specimen, ofNemertesia Lamouroux, 1812, survived (Dobson, 1971; Miss E. Allen, personal communication). Theexquisite drawings listed by Harmer from which most of the plates of Ellis (1755) and Ellis and Solander(1786) were prepared still survive, however. Linnaeus gave no locality for the species, but Ellis recordedit as common on the coast of SW England. 268 P. F. S. CORNELIUS FIG. 3. Obelia dichotoma. (a) SW. England (1959.9.17.43). Part of hydrocaulus.(b) Ireland (1959.9.17.35). Detached gonotheca. Scale = 1000 [j.m for both diagrams. oblique. Gonothecal pedicel annulated, situated in axil of hydrothecal pedicel ;gonotheca inverted cone-shaped, apex domed with central tubular aperture.Measurements - see Table 4. TABLE 4 Measurements of the hydroid stage of Obelia dichotoma in (j.m HYDROTHECA Length (diaphragm to rimBreadth at rim HYDROTHECAL PEDICELS Length INTERNODES Length Maximum breadthLength/breadth ratio GONOTHECA LengthMaximum breadth 280-790 750-1200080-150 C. IO 200-730 930-1150 070-110 C. IO 750-800(obscure) 210-790 830-930120-190 c. 7 870-1000260-320 120-450 430-500080-110c. 4 500-590210-270 HYDROID SPECIES OF OBELIA 269 MATERIAL EXAMINED. Atlantic Ocean - Oban, Argyll, Scotland, 1877, severalcolonies in spirit, coll. A. M. Norman, 1912.12.21.276. North end of Loch Sween,Argyll, Scotland, I m, on Halidrys siliquosa (L.) Lyngbye, iSig, 1 31 May 1962,numerous colonies in spirit and microslide, coll. W. J. Rees, 1962.6.19.13. CaolScotnish, Loch Sween, Argyll, Scotland, I m, on Halidrys siliquosa, 30 May 1962,numerous colonies in spirit and two microslides, coll. W. J. Rees, 1962.6.19.23.Cuan Sound, Argyll, Scotland, LWST, on Halidrys siliquosa, 2 June 1962, numerouscolonies in spirit and microslide, coll. W. J. Rees, 1962.6.19.11 (Fig. 4f). Millport,Isle of Cumbrae, Bute, Scotland, 17 September 1902, several colonies in spirit andmicroslide, coll. E. T. Browne, 1959.10.17.1-2. Clyde Sea, Scotland, 35m, onstem of Thecocarpus myriophyllum (Linnaeus, 1758), 27 August 1920, several hydro-cauli in spirit and microslide, coll. L. P. W. Renouf, 1920.9.10.1. Isle of Man, FIG. 4. Obelia dichotoma. Several hydrothecae showing variation in shape of rim andangle of diaphragm, (a) England (1973.7.23.1). (b) England (1929.1.1.1). (c)Probably British Isles (1920.3.1.1). (d) Massachusetts (1915.3.6.37) (Table 4). (e)Ireland (1967.6.15.106). (f) Scotland (1962.6.19.11). Scale = 250 j.m. 1 The scientific names of algae follow Parke & Dixon (1968). 270 P. F. S. CORNELIUS British Isles, 10 September 1894, colony on microslide, coll. E. T. Browne,1959.9.17.47. Port Erin, Isle of Man, British Isles, 6 October 1892, colony onmicroslide, coll. E. T. Browne, 1959.9.17.37. Martin's Beach, Marloes Peninsula,Pembrokeshire, Wales, colony on microslide, coll. P. Dick, 1959.9.23.2. Ireland,24 July 1902, colony on microslide, coll. E. T. Browne, 1967.6.15.106 (Fig. 46).Ireland, 2 May 1922, five colonies on microslide, coll. E. T. Browne, 1959.9.17.35(Fig. 3b). Plymouth, England, 14 September 1897, several colonies on piece ofwood, in spirit, coll. E. T. Browne, 1954.8.3.71. Plymouth, England, 14 March1898, colonies on two microslides, coll. E. T. Browne, 1959.9.17.32, 43 (Fig. 3a).Millbay Dock, Plymouth, England, 2 November 1906, several hydrocauli in spirit,coll. E. T. Browne, 1954.8.3.77. Mewstone Ledge, Plymouth, England, colony onmicroslide, coll. R. Davis, 1962.8.8.1. Tinside, Plymouth, England, intertidal,on Fucus vesiculosus L., 22 September 1972, several colonies in spirit and microslide,coll. P. F. S. Cornelius, 1973.7.23.1 (Fig. 4a). River Tamar, Devon, England,0-8 km upstream of Cargreen, sublittoral, 4 October 1972, several colonies in spirit,coll. P. F. S. Cornelius, 1973.9.24.4. Weymouth Bay, Portland, Dorset, England,on test of Ascidiella sp. Roule (Tunicata), 20 m, several hydrocauli in spirit, coll.R. Kirkpatrick, 1897.8.9.12. Hastings, Sussex, England, several colonies in spirit,coll. & det. T. Hincks (as Obelia flabellata) , 1899.5.1.148 (non-type). SouthendPier, Essex, England, hydrocauli in spirit, coll. R. Kirkpatrick, 1897.8.9.12. South-end Pier, Essex, England, 24 July 1927, two microslides, coll. F. J. Lambert,1927.9.7.3. Havengore Creek, Rushey I., near Foulness L, Essex, England, micro-slide with three colonies, coll. F. J. Lambert, 1929.1.1.1 (Fig. 4b). 'ProbablyBritish', two colonies on microslide, 1920.3.1.1 (Fig. 4c). Banyuls, S. France,intertidal, 19 October 1959, several colonies on alga, in spirit and microslide, coll.W. J. Rees, 1959.11.17.4. Froggy Pond, station CPi6, University of Cape Town Ecological Survey, Republicof South Africa, three fragments of colony on microslide, 1962.10.2.3 (mentioned,Millard, 1957 : 198). Casco Bay, Maine, U.S.A., on unidentified plant, several hydrocauli in spirit andmicroslide, pres. Smithsonian Institution, Washington, D.C., 1880.9.27.93. W'oodsHole, Massachusetts, U.S.A., on alga sp., 23 July 1911, several hydrocauli in spirit,coll. C. M. Fraser, 1915.3.6.37 (Fig. 4d). Mediterranean I Red Seas - Suez Canal, 13 December 1924, several hydrocauli onwood in spirit and microslide, coll. Cambridge University Expedition to the SuezCanal, 1928.5.31.47, 49. Pacific Ocean - Port Jackson, New South Wales, Australia, several colonies onalgal stipe, in spirit, coll. R. von Lendenfeld, 1886.6.8.102. No locality - Pseudo-dichotomous colony on microslide, prep. H. J. Waddington,ex A. M. Norman coll., 1919.5.26.12. OTHER MATERIAL RECORDED. North polar region - Entire north coast of Russia(Linko, 1911 ; Naumov, 1969), Greenland, Bering Sea, Northern Canada and Alaska(Calder, 1970). Atlantic - coasts of western Europe (Hincks, 1868), Mediterranean Sea and SuezCanal (Deevey, 1950), Azores (Rees & White, 1966), Ghana (Buchanan, 1957), HYDROID SPECIES OF OBELIA 271 Tropical W. Africa (Vervoort, 1959), coast of North America from arctic to Caribbean(Fraser, 1944 ; Vervoort, 1967 ; Calder, 1970), Republic of South Africa (Millard,1958). Pacific - Coast of North America from arctic regions to southern U.S.A. (Fraser,1937), Japan (Hirohito, 1969), China Sea, Galapagos Islands and coast of Ecuador(Deevey, 1950), Tasmania (Briggs, 1939, as 0. australis), New Zealand (Ralph,1957), Mozambique (Millard & Bouillon, 1974). DISTRIBUTION. Widespread in the northern hemisphere, but less frequentlyrecorded south of the equator. Most northerly record probably 77 N, 138 E,north of New Siberian Islands (Linko, 1911, as 0. longissima) ; most southerlyrecord probably 61 S, 45 E, South Orkneys (Ritchie, 1909, as 0. longissima). Occurs intertidally and common at depths down to 100 m, seldom below 300 m,deepest record probably 510 m (Broch, 1918; Naumov, 1969). A world dis-tribution map was given by Deevey (1950). REMARKS. Specimens having exceptionally thickened internodal perisarcconsequently appear intermediate between this species and 0. geniculata. Constantdifferences between the hydroid stages are given in Table i. Colonies of 0. dichotoma showing variations in the length and nature of branchingof the hydrocaulus and in the shape of the hydrothecal rim have been given specificstatus by some authors (see also p. 256). Alder (1857), Hincks (1868) and somesubsequent authors assigned specimens with long, strongly tanned, dark mainhydrocauli to 0. longissima (Pallas, 1766), retaining only specimens with shorter,less tanned main hydrocauli in 0. dichotoma. However, although specimensresembling 0. longissima sensu Alder are distinctive, Obelia specimens in the B.M.(N.H.) collection form a continuous series between the two taxa. In addition,isolated second-order hydrocauli of 'longissima' specimens cannot be distinguishedfrom hydrocauli of 0. dichotoma, and the two taxa appear inseparable. Thusspecimens hitherto assigned to longissima are probably simply older colonies ofdichotoma s. str. Originally Pallas (1766) introduced longissima as an alternative,perhaps more appropriate, name for Linnaeus' species but Linnaeus (1767) gave hisown name priority. Johnston (1838, 1847) also regarded longissima as the juniorsynonym. Obelia flabellata (Hincks, 1866) represents an intermediate point in thisseries and can be regarded as conspecific with 0. dichotoma. 1 Obelia commissuralis McCrady, 1857, was founded on a medusa of which thehydroid was imperfectly known. Agassiz (1862) provided detailed descriptionsof both stages and subsequent descriptions of the hydroid alone were given byNutting (1915) and Fraser (1944). Berrill (1949), although describing the develop-ment of stolons, hydranths and medusae, did not describe systematic characters ofhis material. The species has been distinguished from 0. dichotoma only once, by 1 Certain species of medusae have been assigned to one or other hydroid species but owing to theconfusion surrounding the validity of the medusa species it is probably best to regard these synonymieswith caution. Thus, Sars (1835) described the medusa Thaumantias plana which Nutting (1915) andStechow (igalb) referred to the hydroid Obelia flabellata (Hincks, 1866) (= O. dichotoma); Gegenbauer(1856) described the medusa Eucope polystyla which Mayer (1910) provisionally referred to 0. longissimahydroid (= O. dichotoma); and Agassiz (1865) described the medusa Eucope fusiformis which Bedot(1910) synonymized under O. fusiformis (= O. dichotoma). 272 P. F. S. CORNELIUS Nutting (1915) in a key to the genus, on the basis of the number of annuli perinternode and the arrangement of the side-branches. These characters are variablein 0. dichotoma and on present evidence 0. commissuralis appears invalid. The original description of 0. australis von Lendenfeld, i885a, is inadequate foridentification, but the subsequent restriction by Bale (1888) is clearly referable to0. dichotoma, as also is the non-type material identified as 0. australis by vonLendenfeld (1886.6.8.102). Recently the species was distinguished by Ralph(1957) solely on the presence of an oblique hydrothecal diaphragm, a character notedby Bale and present in von Lendenfeld's material. However, this is a commonvariation in specimens of 0. dichotoma from British localities (Fig. 4e-f) and itsoccurrence in southern populations cannot be considered grounds for maintaining0. australis distinct. Schizocladium ramosum Allman, 1871, was referred to Obelia sp. by Billard (igoia,I904a) and to 0. geniculata by Bedot (1918, 1925). However, Allman originallylikened the species to 0. dichotoma and his illustration, showing a much-branchedcampanularian hydroid with unthickened internodal perisarc, has greater resem-blance to 0. dichotoma to which the species is here referred. Following Millard (1966 : 483) Campanularia obtusidens Jaderholm, igosa, isassigned to 0. dichotoma. Bedot (1925 : 302) had previously placed it in 0. dubiaNutting, 1901, which Millard also assigned to 0. dichotoma. The original descriptiondid not mention gonothecae, and those described by Fraser (1944) differed from thoseof 0. dichotoma only in being stolonal. It seems probable that Fraser's materialwas simply a growth-form of 0. dichotoma. Campanularia obtusidentata is a name ascribed to Vanhoffen (1910 : 272) byBedot (1925 : 302) in the latter's synonymy of 0. dubia Nutting. However, thename does not occur in Vanhoffen's paper and seems to have been used by Bedotalone. It is clearly a lapsus for obtusidens. MEDUSAE. Those reared from both this species and from 0. geniculata have beenfound to resemble 0. lucifera (Forbes, 1848) (summary in Russell, 1953). Theirrelation with the hydroid is discussed below (p. 278). They were first recorded byBaster (1762) who saw their release on 3 June 1757. The subsequent history of theelucidation of the life-cycle was reviewed by van Beneden (1844) and briefly byHincks(i868). Obelia geniculata (Linnaeus, 1758)(Figs i, 5) Sertularia geniculata Linnaeus, 1758:812; Pallas, 1766:117-119; Linnaeus, 1767:1312 (= S. flexuosa Linnaeus) ; Maratti, 1776 : 34. Laomedea lairii Lamouroux, 1816 : 207 ; Lamouroux, 1821 : 14, pi. 67, fig. 3.Campanularia geniculata : Meyen, 1834 : 195-196, pi. 31, figs 3-5 ; Bedot, 1905 : 51-52 (= C. prolifera Meyen ; C. cavolinii Deshayes & Edwards 1 ; C. caulini Chiaje 1 ).Campanularia prolifera Meyen, 1834 : 198-201, pi. 33, figs 1-5.Sertularia prolifera : Deshayes & Edwards, 1836 : 139.Laomedea geniculata : Johnston, 1838 : 151-152, pi. 21, figs 1-2 ; Johnston, 1847 : 103-104, pi. 25, figs 1-2 ; Gosse, 1853 : 84-90, pi. 4 ; Vervoort, ig^6a : 294-298, figs 129-131 (= L. 1 C. cavolinii and C. caulini are here assigned to O. dichotoma. HYDROID SPECIES OF OBELIA 273 /ami Lamouroux ; Sertularia prolifera : Deshayes & Edwards ; Campanularia cavolinii Deshayes & Edwards 1 ; Schizocladium ramosum Allman 1 ).Eucope diaphana L. Agassiz, 1862 : 322-325, pi. 34, figs 1-9 ; (non A. Agassiz, 1865 : 83-85, figs 115-125 ; indeterminate).Obelia geniculata : Allman, 1864:372; Hincks, 1868: 149-151, pi. 25, fig. i, la (= Eucope diaphana L. Agassiz; E. alternata A. Agassiz) ; Bedot, 1910: 338-340 (= Laomedea lairii Lamouroux ; Schizocladium ramosum Allman 1 ; O. gymnopthalma Spagnolini) ; Mayer, 1910:249-252, figs 132-133 (= E. diaphana L. Agassiz; E. alternata A. Agassiz; ? E. polygena A. Agassiz; E. fusiformis A. Agassiz); Vanhoffen, 1910:304-306, fig. 25 ( = Monosklera pusilla von Lendenfeld) ; Bedot, 1916 : 162-164 (= E. diaphana L. Agassiz) ; Bedot, 1925:304-307 (= E. polygena A. Agassiz); Vannucci Mendes, 1946:551-552, pi. 2, figs 14-15 ; Russell, 1953 : 302, fig. i85a ; Naumov, 1960 : 261-263, figs 147-148 ; Naumov, 1969 : 282-283, figs 147-148.Eucope alternata A. Agassiz, 1865 : 86 (nom. nov. pro E. diaphana L. Agassiz, non Thaumantias diaphana A. Agassiz). Eucope polygena A. Agassiz, 1865 : 86-87, fig- I2 ^ ' Bedot, 1912 : 332.Eucope fusiformis A. Agassiz, 1865 : 90, figs 132-133.Obelia gymnopthalma Spagnolini, 1871 : 186 [nom. nov. pro medusa stage of O. geniculata (Linnaeus)]. Monosklera pusilla von Lendenfeld, 18850 : 911-912, pi. 40, figs 1-3.Obelia geniculata var. / Marktanner-Turneretscher, 1890 : 207-208.Obelia geniculata var. // Marktanner-Turneretscher, 1890 : 208.Obelia geniculata var. /// Marktanner-Turneretscher, 1890 : 208.Campanularia coruscans Schneider, 1897 : 482 ; [syn. nov.].Obelia geniculata f. subsessilis Jaderholm, i9O5a : 2, pi. i, fig. 2.Obelia geniculata f. gaussi Vanhoffen, 1910 : 305, fig. 25d.Obelia geniculata f. subtropica Ralph, 1956 : 285.Obelia geniculata f. intermedia Ralph, 1956 : 285.Obelia geniculata f. subantarctica Ralph, 1956 : 285. TYPE SPECIMEN AND TYPE LOCALITY. Ellis, 1755 I 22, pi. 12, fig. B, but not fig. b 2 ; Dover, Kent, England ; intertidal, on fucoid alga. Present location ofspecimen unknown. 1 C. cavolinii, and S. ramosum are here assigned to 0. dichotoma. 2 Linnaeus (1758) gave no type locality and cited only Ellis' description and figure. The illustrationshows a specimen having typical Obelia gonothecae but lacking hydrothecae. Linnaeus' designation'Sertularia denticulis obsoletis, calycibus [= gonotheca] obovatis subrostratis, caule geniculato flexuososimplici' similarly describes the gonotheca, but omits reference to the hydrotheca. It is likely that thespecimens in the Linnean herbarium of the Linnean Society of London (Savage, 1945) were not beforeLinnaeus when he wrote his designation since they possess both hydrothecae and gonothecae (see foot-note, p. 267). It seems probable, therefore, that Linnaeus based his designation solely on Ellis' figure.The locality from which the figured specimen was obtained was not given precisely, although Ellisstated: 'This coralline was found at Dover; and I have lately received some specimens from Harwich;so that I believe it is not uncommon on our coasts.' The figured specimen is perhaps more likely tohave come from the earlier locality, Dover, as the Harwich material possibly came to Ellis too late foran illustration to be prepared. He recorded it on 'podded Fucus' , which could refer either to Fucusvesiculosus or Ascophyllum nodosum (L.) Le Jol (J. M. Price, personal communication). As with O. dichotoma, it is virtually certain that the figured specimen no longer survives (see foot-note, p. 267). However, it remains possible that some other hydroid specimens in the Linnaean her-barium are Ellis' figured material and hence type, since as late as i January, 1767, Linnaeus had re-ceived hydroid material from no source but Ellis (Smith, 1821 : 196). The type-series of Sertulariaechinata Linnaeus, 1761, to be described in a later paper (Cornelius, 1975: in press], indicates that it atleast had reached Linnaeus by or during 1761. This is evident since the original designation of S. echinataincludes characters of both species represented in the type-series, which was mixed. Thus Harmer(1930 : 84) was apparently wrong in suggesting that Linnaeus received his first batch of Ellis' hydroidmaterial at the end of 1766. Unfortunately the correspondence between Linnaeus and Ellis during1763 and 1764 is missing (Smith, 1821 : 164 (footnote); Ahrling, 1885 : 55, 77; Savage, 1948 : 23-25;T. O'Grady, personal communication) so it is not at present clear which if any specimens were sent toLinnaeus during those two years. 274 P. F. S. CORNELIUS FIG. 5. Obelia geniculata. Parts of three hydrocauli to show differing extents of inter-nodal thickening, (a) Republic of South Africa (1964.8.7.80). (b) SW. England(1966.10.28.7). (c) Republic of South Africa (1936.2.4.13), probably the extreme ofshortening and thickening. Scale = 500 [xm throughout. DIAGNOSIS OF SPECIES. Obelia hydroid usually with unbranched, erect, flexuose,monosiphonic hydrocauli ; internodes short, curved, with prominent internalthickening of perisarc below origin of pedicel ; hydrotheca even-rimmed. DESCRIPTION. Colony comprises attached stolons from which arise verticalmonosiphonic hydrocauli, usually unbranched, up to 40 mm (Fig. 5). Hydrocaulusflexuose ; internodes short, curved, internally thickened below origin of eachpedicel, usually with one to five proximal annulations. Pedicel variable in length,attached to short lateral process near distal end of internode, annulated throughoutor with smooth central portion. Hydrotheca broad, even-rimmed, bell-shaped,length about equal to width at rim. Gonotheca situated on short annulated pedicelin axil of hydrotheca ; conical, wider distally, apex domed with narrow tubularaperture. Measurements - see Table 5. MATERIAL EXAMINED. Atlantic Ocean -No locality (? Baltic), probably after1758, coll. Linnaeus, two herbarium sheets, Linnean Society of London cat. no.1298.19-20 (Savage, 1945). x Balta Sound, Shetland, on Laminaria sp., 1867, 1 Sheet 1298.19 bears a specimen of a fucoid alga with an attached colony of O. geniculata comprisingabout 20 erect hydrocauli. All the hydrothecae are missing, and only two gonothecae remain. Sheet1298.20 bears four specimens, one at the top of the sheet, two side by side in the centre and one at thebottom. The top specimen is a piece of Laminaria sp. Lamour. to which a colony of O. geniculata isattached. It has many hydrothecae but no gonothecae. Of the two centre specimens that on the leftis a piece of alga with no hydroids attached, and that on the right is another piece of alga, probablyFucus sp. L. or Laminaria sp. with colonies of O. geniculata attached. A few hydrothecae are present,but no gonothecae. The specimen at the bottom of the sheet is not Obelia, and is not readily identifiedas it is decayed. It appears from an attached label to have been added later. These specimens areprobably not type material (see footnote, p. 273). HYDROID SPECIES OF OBELIA 275 TABLE 5 Measurements of the hydroid stage of Obelia geniculata in \t,m. SW ENGLAND REPUBLIC OF SOUTH AFRICA W SCOTLAND (1973.9.26.1) (1936.2.4.13 ; Fig. 50)* (1962.6.19.5)! HYDROTHECA Length (diaphragm to rim) 240-290 220-270 210-300 Breadth at rim 270-390 250-310 230-320Maximum thickness of hydrothecal perisarc 20 50 20 HYDROTHECAL PEDICELS Length 070-170 070-140 240-370 INTERNODES Length 600-700 470-590 600-820 Maximum breadth 160-250 290-440 210-270 Length/breadth ratio c. 3 c. i - 1 -5 c. 3Maximum diameter of asymmetric thickening 050-100 270 100 GONOTHECA Length 700-800 820-940 1030-1070 Maximum breadth 240-300 290-340 270-320 GONOTHECAL PEDICELS Length 030-080 020-030 090-100 * Specimen resembles 0. geniculata 'var. subsessilis' Jaderholm.f Specimen has long internodes in distal parts. several colonies in spirit and microslide, coll. A. M. Norman, 1912.12.21.262.Clachan Bridge, Seil, Argyll, Scotland, i June 1962, two hydrocladia on microslide,coll. W. J. Rees, 1962.6.19.5. Creagan Narrows, Argyll, Scotland, MLWST, onFucus vesiculosus, 10 September 1970, several hydrocladia, spirit, coll. P. F. S.Cornelius, 1971.5.11.18. Port Erin, Isle of Man, British Isles, 26 September 1892,two hydrocauli on microslide, coll. E. T. Browne, 1959.9.17.29. Port St Mary,Isle of Man, British Isles, on Laminaria sp., 30 m, several hydrocladia in spirit,coll. J. Lomas, 1886.1.9.2. St Ives, Cornwall, England, hydrocaulus on microslide,coll. R. E. W. Vallentin, 1935.8.12.24. Looe, Cornwall, England, LWM, on Cysto-seira sp. Agardh, 21 September 1972, several hydrocauli in spirit, coll. P. F. S.Cornelius, 1973.9.24.2 (Fig. i). Looe, Cornwall, England, LWM, on Laminariasaccharina (L.) Lamour., 21 September 1972, several hydrocauli, coll. P. F. S.Cornelius, 1973.9.24.3. Drake's Island, Plymouth, Devon, England, June 1965,three hydrocauli on microslide, coll. R. C. Vernon, 1969.12.1.20. Torpoint pontoon,Plymouth, Devon, England, 13 August 1963, four hydrocauli on microslide, coll.R. C. Vernon, 1973.9.26.1. Plymouth, Devon, England, 29 September 1947,coll. E. White, two hydrocauli on microslide, 1947.10.8.1. Plymouth, Devon,England, 29 August 1947, coll. E. White, hydrocaulus on microslide, 1947.10.8.3.R. Yealm, Devon, England, MLWST, on Fucus serratus L., 4 July 1973, several 276 P. F. S. CORNELIUS colonies in spirit and six microslides, coll. P. F. S. Cornelius, 1973.7.23.2. R.Yealm, Devon, England, MLWST, on Laminaria sp., 4 July 1973, several coloniesin spirit and six microslides, coll. P. F. S. Cornelius, 1973.7.23.3. Start Bay, Devon,England, on Laminaria sp., 10 m, spirit material and microslide, coll. R. Kirk-patrick, 1893.8.7.7. Eddystone grounds, western English Channel, 2 September1898, several hydrocauli on microslide, coll. E. T. Browne, 1959.9.17.30. GreatBritain Rock, St Mary's, Scilly Isles, on Laminaria sp., 10 m, 22 July 1964, severalcolonies in spirit and two microslides, coll. Queen Mary College expedition,1966.10.28.2. Great Britain Rock, St Mary's, Scilly Isles, 15 m, 23 July 1964,three hydrocauli on microslide, coll. Queen Mary College expedition, 1966.10.28.7(Fig. 5b). South of Fugloy, Espegrend, Norway, 40 m, on bryozoan, 7 August 1962, spiritmaterial and microslide, coll. W. J. Rees, 1962.11.7.47. Nez de Joburg, NW. of Cherbourg Peninsula, France, on Laminaria sp., 3 May1965, several colonies in spirit, coll. A. M. Clark, det. W. J. Rees, 1965.5.14.2. Newport, Rhode Island, U.S.A., on wood, 25 m, several colonies in spirit andmicroslide, pres. Smithsonian Institution, 1890.8.23.14. Potts Point, SouthHarspowell, Maine, U.S.A., on Fucus sp., seven hydrocauli on microslide, coll. C. M.Fraser, 1915.3.6.38. Vineyard Sound, Massachusetts, U.S.A., on Laminaria sp.spirit material and microslide, pres. Smithsonian Institution, 1880.9.27.94. Gough Island, Tristan da Cunha, 22 April 1904, hydrocaulus on microslide, coll.Scottish National Antarctic Expedition, det. J. Ritchie, 1964.8.7.79. Oudekraal, Republic of South Africa, two microslides, University of Cape TownEcological Survey, 1936.2.4.13 (Fig. 5c). Entrance to Saldanha Bay, Cape Colony,Republic of South Africa, 50 m, 21 May 1904, two hydrocauli on microslide, coll.Scottish National Antarctic Expedition, det. J. Ritchie, 1964.8.7.80 (Fig. 5a). Pacific Ocean - New Zealand, 20 hydrocauli on algal thallus in spirit, coll. R. vonLendenfeld, 1886.6.8.107, (Monosklera pusilla', det. von Lendenfeld, not type).Off Port Ross, Auckland Islands, New Zealand, on Laminaria sp., 28 March 1904,several colonies in spirit, coll. 'Discovery' Antarctic Expedition, 1907.8.20.44. OTHER MATERIAL RECORDED. North Atlantic - White Sea (Linko, 1911), JanMayen Island, Iceland, coast of Norway at least to 68 N (Broch, 1918) ; BarentsSea, European coast and Mediterranean Sea (Naumov, 1969) ; fjords of WestGreenland (Calder, 1970) ; whole Atlantic coast of North America (Fraser, 1944) ;Coats Island, Hudson Bay (63 N) (Calder, 1970) ; parts of Caribbean (Vervoort,1968). South Atlantic - Zaire (Leloup, 1939), Ghana (Buchanan, 1957), Luderitz Bay(26 S) and other localities in Republic of South Africa (Broch, 1914 ; Millard,1957, 1966) ; Kerguelen Island (Vanhoffen, 1910) ; Falkland Islands (M. W. Robins,personal communication) ; South Georgia (Deevey, 1950) and Brazil (VannucciMendes, 1946). North Pacific - Japan (Yamada, 1958) ; British Columbia and whole of U.S.A.coast except Alaska (Fraser, 1937). South Pacific - Chile (Jaderholm, igo^b), Indonesia and Galapagos Islands(Deevey, 1950), West Australia, South Australian Bight, Victoria, New South HYDROID SPECIES OF OBELIA 277 Wales and Tasmania (Hodgson, 1950), Macquarie Island (Briggs, 1939), New Zealand,adjacent seas between 35 S and 49 S and the Chatham Isles (Ralph, 1956, 1961). Indian Ocean -Two localities in southern India (Mammen, 1965), Mozambique(Millard & Bouillon, 1974). DISTRIBUTION. Almost cosmopolitan in continental shelf seas. Deevey (1950)stated that 0. geniculata is the most widely distributed of all hydroids, being absentonly from the northern Indian Ocean and the tropical West Atlantic (althoughsubsequently recorded from the Caribbean). Apart from South Georgia andMacquarie Island there appear to be no records also from the Southern Ocean, andthe species appears unrecorded from much of the Pacific, including northern partsof the Australian mainland, Brisbane and the Great Barrier Reef (Ralph, 1956). In the northern hemisphere the species extends almost as far north as 0. dichotoma,being known from the White Sea. Although it was reported absent from Greenlandby Broch (1918) and was not found by Kramp (1932) in some west Greenlandcollections, Calder (1970) nevertheless recorded the species from both east and westGreenland, but did not cite material. REMARKS. Variation in the hydroid stage of this species occurs mainly in theangle of flexure between internodes, the asymmetric thickening of the internodalperisarc, the length to breadth ratio of the internodes and the shape of the hydro-theca. Apart from the form of the hydrotheca, these characters can be placed in aseries ranging from specimens with shallow angles between internodes, little inter-nodal thickening and proportionately long internodes to more flexuous specimenswith greater internodal thickening and shorter internodes (Fig. 4a-c). The occurrence of these variations has prompted authors to recognize severalvarieties and formae. Vanhoffen (1910), however, was of the opinion that thefollowing of these taxa fall within the limits of normal variation : Varieties /, //and III of Marktanner-Turneretscher, 1890 ; forma gaussi Vanhoffen, 1910 (nom.nov. pro var. / of Marktanner-Turneretscher) and forma subsessilis Jaderholm,I905b. Further varieties (subtropica, intermedia and subantarctica) were describedby Ralph (1956) based on variations in length of hydrocaulus, number of annulationsof the internodes, amount of branching and linear dimensions of all structures.These characters are known to be variable (p. 256) and it is perhaps best at presentnot to regard them as systematically valid. The species appears to be variable within the above limits over the whole ofits range. Thus, specimens showing the extreme of shortening and internodalthickening have been reported from Chile, Kerguelen Island and the Republic ofSouth Africa (Jaderholm, igosb ; Vanhoffen, 1910 ; Fig. 5c), while specimens withboth minimal and maximal thickening are known from the English Channel(1959.9.17.30). Almost the full range of variation is represented in British specimensin the BM(NH) collection, and the naming of distinct varieties seems unnecessary. Hammett & Hammett (1945) followed the seasonal morphological changes incolonies of 0. geniculata during several summers in Massachusetts but variationsin features of accepted systematic importance were not reported. More recentlydetailed accounts of variation in New Zealand populations by Ralph (1956) andRalph & Thomson (1968) demonstrated that over a wide geographical range and 278 P. F. S. CORNELIUS from season to season at a single locality certain morphological characters variedwith temperature. Low temperatures induced longer colonies with longer inter-nodes, and colonies from warmer localities showed a reduction in branching. Laomedea lairii Lamouroux, 1816, was placed in the present species by Bedot(1901) and also by Billard (1909) who examined the type specimen. The specimenwas subsequently destroyed in the Second World War (Redier, 1967). Campanularia coruscans Schneider, 1897, was originally stated to release a medusaand was subsequently assigned to Obelia by Stechow (i92ib, 1923^. The widedistal shelf in each internode supporting the hydrotheca, as described by Schneider,suggests that C. coruscans is referable to 0. geniculata. MEDUSAE. Those reared from both this species and 0. dichotoma have been foundto resemble 0. lucifera (Forbes, 1848) (summary in Russell, 1953). Their relationwith the hydroid is discussed below. The earliest record of medusa release in 0.geniculata seems to be that of F. W. L. Thomas (in Johnston, 1847 : 467), whocommented on the similarity between the medusa of this species and that of 0.dichotoma. Medusae of 0. geniculata were apparently first illustrated by Gosse (1853). THE MEDUSA PROBLEM Many nominal species of Obelia are based solely on the medusa stage, most havingbeen described in the past 100 years (references in Mayer, 1910 ; Bedot, 1901-25 ;Kramp, 1961). However, they are very similar and it is likely that many areconspecific (Kramp, 1961). As noted by Russell (1953), it is at present impossibleto relate them to the hydroid species, and it may remain so until further rearingwork has been done and the characters of the medusae reassessed. Although all three species recognized from the hydroid stage occur in Britishseas, only two nominal species of medusae are known. The five taxa recorded areas follows (modified from Russell, 1953) : HYDROIDS 0. bidentata Mature medusa not described 0. dichotoma Mature medusa resembles 0. lucifera 0. geniculata Mature medusa resembles 0. lucifera MEDUSAE 0. lucifera Reared to maturity from both 0. dichotoma and 0. geniculata, and known from the plankton0. nigra Browne, 1900 Known only from the plankton It has been suggested by elimination (Browne, in Kramp, 1927 ; Russell, 1953)that 0. longissima might prove to be the hydroid of 0. nigra. However, thiscannot be the whole answer if, as is shown above, 0. longissima is conspecific with0. dichotoma from which 0. lucifera has been reared. The occurrence of the hydroid0. bidentata in the North Sea and English Channel, and the fact that the appearanceof its medusa when adult is unrecorded, add further complications. HYDROID SPECIES OF OBELIA 279 The possibility that 0. nigra alone is released from 0. bidentata is unlikely as thetwo species have different distributions in western Europe. Thus, the hydroidoccurs no further north than the coasts of France, Belgium and Holland and thesouthern North Sea (Vervoort, ig^6a. ; Leloup, 1952 ; Hamond, 1957 ; Kramp,1961 ; Teissier, 1965), while the medusa is known from as far north as westernScotland, Iceland and Bergen, Norway (Browne, 1905 ; Thiel, 1932 ; Kramp,1939 ; Rees, 1953). Sir Frederick Russell (personal communication) is of the opinion that in the seasaround the British Isles 0. nigra is associated with mixed oceanic and coastal watertypified by the presence of Sagitta elegans (Chaetognatha), while 0. lucifera occursin coastal water of which S. setosa is characteristic (Meek, 1928). 0. nigra is saidto be larger than 0. lucifera and to have darker pigmentation around the tentaclebases (Russell, 1953). It has long been suspected that the mixed water is moreproductive than British coastal water (Russell, 1939), and as suggested by Browne(in Kramp, 1927) the larger size characteristic of 0. nigra might simply reflectbetter feeding opportunities. Thus it is possible that medusae from all threehydroid species develop 0. lucifera characters in coastal water and those of 0. nigrain more productive areas. It is noteworthy that although Browne recorded bothspecies of medusae off Plymouth between 1897 and 1899, published records suggestthat he never obtained them together (Marine Biological Association, 1957). Thisperhaps indicates that the furthest extent of mixed water into the English Channelthen varied in position around Plymouth. Although mixed oceanic and coastalwater frequently reached Plymouth in the 1920*3, it did not do so for some decadesafter the early i93o's (Russell et al., 1971). During that time 0. lucifera was theprevalent species (Russell, personal communication), further suggesting a correlationwith water mass and, possibly, feeding opportunity. Although it might be inferred from the available evidence that the two medusaspecies are conspecific, the possibility that they are genetically distinct neverthelessremains open, while another possibility is that the hydroid of 0. nigra is as yetundescribed. SPECIES TRANSFERRED TO OTHER GENERA Although Campanularia gelatinosa (Pallas, 1766) was reported to release a medusaby van Beneden (1844), his illustrations identify his material as Obelia dichotoma.Hincks (1868) was aware of this misidentification, but independently described amedusa stage in C. gelatinosa and placed the species in Obelia. Later Maitland(1876 : 13) identified the medusa Obelia marina (Slabber, 1769) with Campanulariagelatinosa auct. (? sensu Hincks). A medusa has not been reported since in thespecies, and recent opinion (Vervoort, ig46a ; Naumov, 1969 ; J. Clare, personalcommunication) is that one is not released. It is possible that Hincks mistook thelarge ova characteristic of C. gelatinosa for developing medusae. Campanularia denticulata Clarke (1876 : 9, pi. i, fig. 4) has usually been referred toObelia (references in Bedot, 1912-25) although Pictet (1893) assigned it to Clytianoliformis McCrady, 1857. Mammen (1965) did not agree with Pictet's synonymy 280 P. F. S. CORNELIUS and Clarke's nominal species still stands as Campanularia denticulata. However,the acutely cusped hydrothecal rim originally described suggests that the speciesshould nevertheless be referred to Clytia. Obelia marginata Allman (1877 : 9-10, pi. 6, figs 1-2) is now accepted as a speciesof Cnidoscypus Splettstosser, 1929 (Vervoort, 1968). The two nominal species Obelia longicyatha Allman, 1877, and 0. longicyathaThornely, 1899, are discussed above under 0. bidentata (p. 264). Eucope annulata von Lendenfeld (i885a : 602-603, pi. 28, figs 53-57) was placedin Obelia by Bedot (1925 : 298), but von Lendenf eld's illustration shows an approxi-mately hemispherical medusa quite unlike Obelia and Kramp (1961) referred thespecies to Phialella Browne, 1902. Campanularia serrulata Bale (1888 : 757, pi. 12, fig. 4) was referred to Obelia byThornely (1899) and Mayer (1910, mis-spelt as 0. serratula) but later transferred toClytia by Bedot (1918, 1925). Although the triangular cusps on the hydrothecalrim shown by both Bale and Mayer are typical of Clytia, the gonotheca described byThornely was unlike that of Clytia in having a rounded, truncate top. However, asthe gonotheca may have been immature, it is still possible that the three authorswere describing the same species. Nutting (1927), however, provided the new nameObelia thornelyi for Thornely's material. Nevertheless, the nature of the hydro-thecal rim described by both Bale and Mayer suggests that their material at leastshould be referred to Clytia. Campanularia castellata Clarke (1894 : 71-72) (= Obelia castellata Clarke, 1894 : 73,pi. i, fig. 3-8, pi. 2, fig. 9) is referable to Campanularia gelatinosa (Pallas, 1766) as ithad a castellated hydrothecal rim. Bedot (1918 : 196 ; 1925 : 300) mis-spelt thename as castellana. Campanularia kincaidi Nutting (1899 : 743-744, pi. 62, figs 2a-c) was referred toLaomedea by Leloup (1940 : 21) and to Obelia by Rees & White (1966 : 277). Itscreeping habit and singly cusped hydrothecal rim are reminiscent more of Clytia,however ; and in the absence of information on the life-history the species is probablybest excluded from Obelia. Obelia linearis Thornely (1899 : 453, pi. 44, fig. 6), although retained in Obeliaby Mayer (1910 : 257), was transferred to Clytia sp. by Mammen (1965 : 21). Thepresence of a deeply cleft hydrothecal rim and a reflexed gonothecal aperturesupport Mammen's opinion. Obelia delicatula Thornely (1899 : 453, pi. 44, fig. 7) was assigned to Clytia byStechow (i923b) and Blackburn (1942). Although the medusa is unknown, itsstolonal gonotheca and sharply-cusped hydrothecal rim are features typical ofClytia and the species is for the present probably best assigned to that genus. Obelia austrogeorgiae Jaderholm (i904b : 7 ; igo^b : 17, pi. 17, figs 1-2) has notbeen shown to produce a medusa and is here provisionally referred to the genusCampanularia. The species has also been recorded by Nutting (1915) and Vervoort(i972b, as Laomedea (Obelia} austrogeorgiae). As reported by Jaderholm, gonothecaeare absent from the schizoholotype material (1960.8.29.34) which was examinedduring the present work, and have not yet been recorded for the species. At present,therefore, its generic affinity is obscure, but the bimucronate hydrothecal rim HYDROID SPECIES OF OBELIA 281 originally described suggests that Mayer (1910) may have been correct in assigningthe species to 0. bidentata. Obelia striata Clarke (1907 : 9-10, pis 6-7) is a distinctive hydroid known sincethe first description to release a medusa with four tentacles. It was, therefore,referred to Clytia sp. by Rees & Thursfield (1965) and to Clytia gravieri (Billard,I904b) by Millard & Bouillon (1973), while Vervoort (1966) assigned it to Laomedea(Phialidium) . As adult medusae are unrecorded it is difficult to assign the speciesto a particular genus with confidence, but the four tentacles of the young medusasuggest that it is not an Obelia species. Vervoort (1968 : 19) considered thatLaomedea tottoni Leloup, 1935, was very similar to 0. striata Clarke. Laomedea (Obelia} bistriata Leloup (1931 : 4-6, figs 8-n) was referred to Clytiahendersonae Torrey, 1904, by Mammen (1965) and to C. gravieri (Billard, 1904^ byVervoort (1967), Schmidt (1972) and Millard & Bouillon (1973). The spines associ-ated with the cusps on the hydrothecal rim were illustrated by Leloup (1931, 1932)and Vervoort, and resemble closely those of 0. bidentata. Although such spines areotherwise unreported from Clytia, Vervoort's observation that the developingmedusa has a 'strongly convex umbrella', and Millard & Bouillon's that it has fourtentacles, are evidence that the species should be assigned to the genus Clytia. As stated in the generic diagnosis, several of the species admitted to the genusObelia by Naumov (1960, 1969) are here referred to the genera Campanularia andGonothyrea. The species involved are Campanularia gelatinosa (Pallas, 1766), C. gracilis 1 Sars, 1850 (non 0. gracilis Calkins, 1899, = 0. dichotoma), C. flexuosa(Hincks, in Alder, 1856) and G. loveni (Allman, 1859), none of which releases amedusa. ACKNOWLEDGEMENTS I am grateful to R. W. Sims and Dr W. Vervoort for detailed criticism of themanuscript, and to Dr Elaine Robson and Sir Frederick Russell, F.R.S., for com-menting on parts of it. The last-named kindly allowed me to quote unpublishedobservations on the distribution of Obelia medusae in the British Isles, and Dr M. W.Robins provided an unpublished locality record of 0. geniculata, in the FalklandIslands. I am grateful also to the following for helpful conversation and cor-respondence : Miss P. L. Cook, J. Clare, Dr C. Edwards, Dr R. Hamond, the late D. N. Huxtable, J. M. Price and Dr M. W. Robins. D. W. Cooper kindly prepared serial sections of Obelia geniculata, and A. Varley,Librarian at the Marine Biological Association's Plymouth Laboratory, helped withliterature. I was fortunate in being permitted to work at that Laboratory and tomake use of its research vessels, and similarly fortunate in being able to work for ashort time at the Laboratory of the Scottish Marine Biological Association, Oban,and to collect from its vessel. T. O'Grady of the Linnean Society of London kindlyallowed me to study Linnaeus' hydroid material and correspondence with JohnEllis, and Miss Elizabeth Allen of the Hunterian Museum of the Royal College ofSurgeons of England provided information about the Ellis material held there. 1 Vervoort (19463 : 285) had previously suggested that the correct name for Sars' species is C. pelagicavan Breemen, 1905. 2 8a P. F. S. CORNELIUS Specimens were lent by Dr W. D. Hartman of the Peabody Museum of NaturalHistory, Yale, and Dr W. 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Report on the Hydroida collected by Professor Herdman, at Ceylon, in 1902. Rep.Govt. Ceylon Pearl Oyster Fish. Gulf Manaar, Suppl. Rep. 8 : 107-126. 1908. Reports on the marine biology of the Sudanese Red Sea. X. Hydroida collectedby Mr C. Crossland from October 1904 to May 1905. /. Linn. Soc. 31 : 80-85. TORREY, H. B. 1904. The hydroids of the San Diego region. Univ. Calif. Publs Zool. 2 : 1-43.VANHOFFEN, E. 1910. Die Hydroiden der Deutschen Siidpolar-Expedition 1901-1903. Dt. Sudpol.-Exped. XI, Zoologie, 3 : 269-340.VANNUCCI MENDES, M. 1946. Hydroida Thecaphora do Brasil. Archos Zool. Est. S. Paulo, 4:535-597-VANNUCCI, M. 1951. Hydrozoa e Scyphozoa existentes no Institute Paulista de Oceanografia. Bolm Inst. Oceanogr. S. Paulo (i) 2 : 69-104.1954. Hydrozoa e Scyphozoa existentes no Institute Oceanografico, II. Bolm Inst. Bolm Oceanogr. S. Paulo, 5 : 95-149.VERVOORT, W. I94&a. Hydrozoa (C i). A. Hydropolypen. Fauna Ned. 14 : 1-336. i946b. Exotic hydroids in the collections of the Rijksmuseum van Natuurlijke Historicand the Zoological Museum at Amsterdam. Zool. Meded. Leiden, 26 : 287-351. - 1959. The Hydroida of the tropical west coast of Africa. Atlantide Rep. 5 : 211-325. 1966. Bathyal and abyssal hydroids. Galathea Rep. 8 : 97-174. - 1967. The Hydroida and Chondrophora of the Israel South Red Sea Expedition, 1962.Bull. Sea Fish. Res. Stn Israel, 43 : 18-54. 1968. Report on a collection of Hydroida from the Caribbean region, including an annota-ted checklist of Caribbean hydroids. Zool. Verh. Leiden, 92 : 1-124. I972a. Hydroids from the Theta, Vema and Yelcho cruises of the Lamont-DohertyGeological Observatory. Zool. Verh. Leiden, 120 : 1-247. I972b. Hydroids from submarine cliffs near Arthur Harbour, Palmer Archipelago,Antarctica. Zool. Meded. Leiden, 47 : 337-357. WEILL, R. I934a. Contribution a 1'etude des cnidaires et de leurs nematocystes. I.Recherches sur les nematocystes. Trav. Stn zool. Wimereux, 10 : 1-347. i934b. Contribution a 1'etude des cnidaires et de leurs nematocystes. II. Valeurtaxonomique du cnidome. Trav. Stn zool. Wimereux, 11 : 347-701. WESTFALL, J. A. 1966. The differentiation of nematocysts and associated structures in the Cnidaria. Z. Zellforsch. mikrosk. Anat. 75 : 381-403.YAMADA, M. 1958. Hydroids from the Japanese Inland Sea, mostly from Matsuyama and its vicinity. /. Fac. Sci. Hokkaido Univ. (6) Zool. 14 : 51-63. HYDROID SPECIES OF OBELIA 291 INDEX Valid names are printed in Roman type and names regarded in this paper as synonyms areprinted in italic type. Main page references also are in italics. adelungi, Obelia 266alternata, Eucope 273alternata, Obelia 267andersoni, Obelia 260, 264, 265angulosa, Obelia 266annulata, Eucope 280arruensis, Obelia 266articulata, Eucope 266articulata, Obelia 266attenuata, Obelia 260australis, Obelia 266, 271, 272austrogeorgiae (Campanularia, Laomedea &Obelia) 260, 280-281 bicuspidata, Gonothyrea 260 bicuspidata, Laomedea 260 261, 264 bicuspidata, Obelia 260-262, 264 bidentata, Laomedea 260 bidentata, Obelia 253, 254, 255, 256-259 (variation), 260-265 (main section), 27^- 279 bifurca, Obelia 260bifurcata, Obelia 260biserialis, Obelia 267bistriata, Laomedea (Obelia) 281borealis, Obelia 266brasiliensis, Campanularia 265braziliensis, Obelia 267 Campanularia 254 (synonymy), 256 Campanularia 252, 255, 281 castellana, Campanularia 280 castellata, (Campanularia & Obelia) 280 caulini, Campanularia 265, 272 cavolinii, Campanularia 265, 272 cavolinii, Sertularia 265 chinensis, Obelia 266 Clytia 264 Clytia 253-254, 260, 279-281 Cnidoscyphus 280 commissuralis, Obelia 265, 27 j -272 congdoni, Laomedea 267 congdoni, Obelia 266 corona, Obelia 260 coruscans, Campanularia 278 coruscans, Obelia 273, 27$ coughtreyi, Obelia 267 delicatula (Clytia & Obelia} 280denticulata, Campanularia 279-2^0 diaphana (Eucope & Thaumantias) 273dichotoma (Campanularia & Laomedea) 265dichotoma, Obelia 251, 253, 254, 255, 256- 259 (variation), 265-272 (main section), 275-279 dichotoma, Sertularia 253-254, 265divaricata, Laomedea 265, 266dubia, Obelia 266, 272 echinata, Sertularia 273equilateralis, Obelia 267Eucope 252, 254everta, Obelia 267 flabellata, Campanularia 266 flabellata, Obelia 266, 270, 271 (text and footnote) flexuosa, Campanularia 256, 257, 281flexuosa, Laomedea 265flexuosa, Sertularia 272fragilis, Obelia 266fusiformis, Eucope 271 (footnote), 273 gaussi, forma of Obelia geniculata 273, 277 gelatinosa, Campanularia 265, 266, 279, 281 gelatinosa, Obelia 281 genicolata, Sertolare 265 geniculata (Campanularia & Laomedea} 272 geniculata, Obelia 251, 253, 254, 255, 256- 259 (variation), 271, 272-27$geniculata, Sertularia 265, 272Gonotha 260Gonothyrea 253Gonothyrea 281gracilis, Campanularia 281gracilis, Laomedea 266gracilis, Obelia 266, 281gravieri, Clytia 281griffini, Obelia 266gymnopthalma, Obelia 273 helgolandica, Obelia 266 hemisphaerica (Medusa & Thaumantias) 254 hendersonae, Clytia 281 hyalina, Gonothyrea 266 hyalina, Obelia 265, 266 intermedia, forma of Obelia geniculata 273, 277irregularis, Obelia 267 2Q2 P. F. S. CORNELIUS kincaidi (Campanularia, Clytia, Laomedea &Obelia) 280 lairii, Laomedea 272, 273, 278 Laomedea 252, 253-254 linearis (Clytia & Obelia) 280 longa, Obelia 260 longicyatha, Clytia 261, 265 longicyatha, Gonotha 260 longicyatha, Gonothyrea 260, 265 longicyatha, Laomedea 261 longicyatha, Obelia 260, 264, 280 longissima, Laomedea 265 longissima, Obelia 256, 258, 266, 271, 278 longissima, Sertularia 265 longitheca (Clytia & Obelia} 260 loveni, Gonothyrea 281 lucifera, Obelia 251, 272, 278-279 maior, Campanularia 265 marginata (Cnidoscyphus & Obelia) 280 marina, Medusa 253, 254 marina, Obelia 253, 279 Medusa 252, 255 microtheca, Obelia 267 minor, Laomedea spinulosa 260, 261, 264 Monosklera 252, 254, 273 multidentata, Obelia 260 Nemertesia 267 (footnote)nigra, Obelia 278-279nigrocaulus, Obelia 266nodosa, Obelia 266noliformis, Clytia 279 Obelaria 252, 254Obeletta 252, 254 Obelia 257-259 (introduction, generic syn-onymy, variation and identification)Obelia sp. 260 Obelia spp. See under specific nameObelissa 252, 254 obtusidens, Campanularia 266, 267, 272obtusidens, Obelia 267obtusidentata, Obelia 267, 272oxydentata, Obelia 260 parasitica, Eucope 265pelagica, Campanularia 266, 281Phialella 280 Phialidium 281 picteti, var. of Laomedea bicuspidata 260, 264 piriformis, Obelia 266 plana, Obelia 266 plana, Thaumantias 271 (footnote) plicata, Obelia 266 polygena, Eucope 273 polystyla, Eucope 271 (footnote) prolifera (Campanularia & Sertularia) 272 pusilla, Monosklera 273, 276 pygmaea, Obelia 266 pyriformis (Eucope & Obelia) 266 racemosa, Obelia 267 ramosum, Schizocladium 266, 272, 273 rhunicola, Obelia 266 sargassi, Laomedea 267 Schizocladium 252, 254, 272 serratula, Obelia 280 serrulata, Campanularia 280 Sertolare 265 Sertularia 265 Sertularia 252 Slabberia 252 solowetzkiana, Obelia 266 sphaerulina, Obelia 253, 265, 266 spinosa, Sertularia 253 spinulosa, Campanularia 260 spinulosa, Obelia 260, 265 striata (Clytia, Laomedea & Obelia) 281 subantarctica, var. of Obelia geniculata 273, 277 subsessilis, var. of Obelia geniculata 273, 277subtropica, var. of Obelia geniculata 273, 277surcularis, Obelia 266syringa, Sertularia 254 tenuis, Obelia 267 tenuis, var. of Laomedea bicuspidata 261, 264 Thaumantias 252, 254 thornelyi (Clytia & Obelia) 280 tottoni, Laomedea 281 undotheca, Obelia 266 verticillata, Sertularia 254Verticillina 254Vesicularia 253volubilis, Sertularia 254 HYDROID SPECIES OF OBELIA 293 DR P. F. S. CORNELIUS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 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.) 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 ; 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. Hardback edition 6. Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 fNU SOME NEW AND RARE SPECIES OF CALANOID COPEPODS FROM THE NORTHEASTERN ATLANTIC H. S. J. ROE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 7 LONDON: 1975 SOME NEW AND RARE SPECIES OF CALANOID COPEPODS FROM THE NORTHEASTERN ATLANTIC BY HOWARD STANLEY JAMES ROE Institute of Oceanographic Sciences Pp 295-372 ; 33 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 7 LONDON: 1975 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in five series corresponding to the ScientificDepartments of 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. 28 No. 7 of the Zoology series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) ISSN 0007-1498 Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 5 November, 1975 Price 4-75 SOME NEW AND RARE SPECIES OF CALANOID COPEPODS FROM THE NORTHEASTERN ATLANTIC By H. S. J. ROE SYNOPSIS One new genus, nine new species, and nineteen undescribed males of known species of calanoidcopepods are described. A further eighteen rare or poorly known species are described orcommented upon. Five existing species are transferred to the new genus ; new synonymiesare proposed for six species, and two species are recorded as incertae sedis. All the specimenswere taken from vertical series of closing net hauls made in the northeastern Atlantic. INTRODUCTION FOR some years the Institute of Oceanographic Sciences has been investigating thevertical distributions and migrations of zooplankton and nekton in the northeasternAtlantic. These investigations have been based largely upon a series of verticalhauls made with horizontally towed opening and closing net systems. The calanoidcopepods have now been analysed from two of these series. The results of the first,the RRS 'Discovery' SOND cruise in 1965, have been given previously (Roe, 19720,b, c, d). The SOND series was taken with a modified Indian Ocean Standard Net(the Nii3H), fitted with a catch dividing bucket, in a position off Fuerteventurain the Canary Islands. Details of the sampling methods used in this series are givenby Foxton (1969), Angel (1969) and Roe (19720). The second series was made in 1969 in a position centred on i8N 25W near theCape Verde Islands. A day and a night series of hauls was made with an acousticallycontrolled net system. This comprises two rectangular midwater trawls, one of8 m 2 mouth area and mesh of 4-5 mm (RMT 8), the other of i m 2 mouth area andmesh 0-32 mm (RMT i), combined within the same frame. This net system,designated the RMTi + 8, is described by Baker, Clarke & Harris (1973). Theseries was taken at 1250-0 m depth and was fished in the manner described byBaker et al. Both these series contained a number of very rare calanoid copepods, severalundescribed sexes of known species and some new species. These specimens are alldescribed below. The geographical positions given for each species are approximate ;more accurate positions are listed at the Institute of Oceanographic Sciences. TAXONOMY Family SPINOCALANIDAE The RMT i collections contain females of the following species of Spinocalanus :S. abyssalis Giesbrecht, 1888 ; 5. magnus Wolfenden, 1904 ; S. spinosus Farran,1908 ; S. horridus Wolfenden, 1911 ; S. angusliceps Sars, 1920 ; 5. brevicaudatus 298 H. S. J. ROE Brodsky, 1950 ; S. parabyssalis Park, 1970 ; 5. hoplites Park, 1970 ; 5. usttatusPark, 1970 ; 5. pteronus Park, 1970. These females all agree with previous de-scriptions and all except 5. angusticeps fit Park's (1970) partial key to the genus.Together with these females were the males of five species. Three of these aremore or less tentatively identified as 5. abyssalis, S. brevicaudatus and 5. sp. Grice& Hulsemann, 1967, but the remaining two could not be attributed to any knownspecies. Since the males of many species of Spinocalanus are unknown and as thepresent specimens are all incomplete they are not considered here to be new species. Spinocalanus ? abyssalis Giesbrecht, 1888 MATERIAL EXAMINED : 56 males from 13 RMT i hauls made at 1020-300 mdepth in a position i8N 25W, between 12 and 16 November 1969. 44 specimensdeposited at the British Museum (Natural History), reg. no. 1974 : 462-482. DESCRIPTION. Male (Fig. la-j) : Body length 1-06-1-22 mm with a mean ofi -14 mm (35 specimens). The cephalothorax is stout and 1-7 times as long as the5-segmented abdomen. In lateral view the cephalothorax is arched dorsally.The head and ist thoracic segment are completely fused and thoracic segments 4 and5 are partially so. The 3rd thoracic segment has protruding corners in dorsal view.There is no rostrum. The anal segment is very short and telescoped into the 4thabdominal segment. The ist antenna has 19 free segments and reaches to abouthalfway along the 2nd abdominal segment ; the basal eight segments have largeflattened sensory filaments ; the ist segment has a small group of spines. Theremaining mouthparts and legs are as shown. A satisfactory preparation of theist maxilla was not obtained but it appears to be similar to that of S ? brevicaudatus(p. 300). The 2nd maxilla and maxilliped are weakly chitinized and have smallfeeble setae. The swimming legs of all specimens are damaged. The 5th pair oflegs reaches back to the hind edge of the 2nd abdominal segment ; the right endopodand the terminal spine of the left exopodite are broken off in all the dissected speci-mens. REMARKS. Brodsky (1950) and Park (1970) discussed the synonomy of this species.The present males do not fit the description of any known Spinocalanus male.They were taken at depths where the most abundant spinocalanids were females ofS. abyssalis (2022 specimens), S. spinosus (336), S. magnus (in), 5. brevicaudatus(73) and 5. angusticeps (50). S. magnus, the male of which is described by Tanaka(1956), and 5. angusticeps are larger species, and the presumed male of S. brevicauda-tus is described below. According to Brodsky (1950) the male of S. abyssalis isunknown, although Vervoort (1946) mentioned a male of 5. abyssalis var. pygmaeusmeasuring 1-14 mm. Presumably the males listed by Vervoort (1957) and thatdescribed by Bradford (19710;) are too large (1-7-2-2 mm) to be 5. abyssalis. Themales of all species in which the female has a spiny cephalothorax are also unknown.The present specimens seem to be the right size to be the males of S. abyssalis - thefemales of this species here average 1-04 mm in length (1055 measured specimens),whereas those of S. spinosus are larger (1-79 mm from 156 specimens). They are NEW AND RARE CALANOID COPEPODS 299 FIG. i. a-j, Spinocalanus ? abyssalis $. a, base of ist antenna; b, 2nd antenna;c, mandible ; d, 2nd maxilla ; e, maxilliped ; f, ist leg ; g, 2nd leg ; h, 3rd leg ; i, 4thteg I j 5th leg ; k-u, Spinocalanus ? brevicaudatus <J. k, body, dorsal ; 1, 2nd antenna ;m, mandibular palp ; n, ist maxilla ; o, 2nd maxilla ; p, maxilliped ; q, ist leg ; r, 2ndleg ; s, 3rd leg ; t, 4th leg ; u, 5th leg. Bar scale o-i mm unless indicated. 300 H. S. J. ROE recorded here as 5. abyssalis although in the absence of complete animals thisidentification remains tentative. Spinocalanus ? brevicaudatus Brodsky, 1950 MATERIAL EXAMINED : 21 males from 10 RMT i hauls made at 1250-300 m depthin a position i8N 25W, between 12 and 17 November 1969. 17 specimens de-posited at the British Museum (Natural History), reg. no. 1974 : 483-493. DESCRIPTION. Male (Fig. ik-u) : Body length 1-29-1-44 mm with a mean ofi -38 mm (16 specimens). The cephalothorax is elongated and just over twice aslong as the 5-segmented abdomen. The head and ist thoracic segment are com-pletely fused and thoracic segments 4 and 5 are partially so. In lateral view thecephalothorax is not strongly arched. The ist antenna has 22 segments and reachesto the hind edge of the 2nd abdominal segment ; it is similar in structure to that ofS ? abyssalis (p. 298) but has no spines on the ist segment. The remaining mouth-parts and the legs are as shown. The ist and 2nd maxillae and the maxilliped areweakly chitinized, and the three inner lobes of the ist maxilla bear very rudimentaryshort setae. The outer edge spine of the ist exopodite segment of the ist leg isshort. The exopodites of legs 3 and 4 are damaged in all specimens. The 5th pairof legs reaches back to the hind edge of the 2nd abdominal segment. REMARKS. Brodsky (1950) discussed the synonomy of this species. The presentmales seem to agree with previous descriptions of the male of 5. brevicaudatus,especially in their rather elongate body and structure of the 5th pair of legs. Thereare, however, some differences : With's (1915) specimen had more setae on all lobesof the ist maxilla, though these are mostly rudimentary and their numbers mayvary, and Sars' (1903) specimen had only two rows of spines on the 2nd endopoditesegment of the 4th leg whereas the present males have three. These specimens can be distinguished from the previous species by the slightlylarger size, the shape and proportions of the body, the details of some of themouthparts and spinulation of the swimming legs, and by the structure of the 5thlegs. They are tentatively identified here as S. brevicaudatus although since themales of so many Spinocalanus spp. are unknown specific determination of thesespecimens is doubtful. Spinocalanus sp. Grice & Hulsemann, 1967 MATERIAL EXAMINED : n males from 5 RMT i hauls made at 1250-800 m depthin a position i8N 25W, between 12 and 18 November 1969. 10 specimens de-posited at the British Museum (Natural History), reg. no. 1974 : 494-504. DESCRIPTION. Male (Fig. 2) : Body length 1-22-1-29 mm with a mean of 1-25 mm(7 specimens). The cephalothorax is 3 times as long as the 5-segmented abdomen.The head and ist thoracic segment are fused but thoracic segments 4 and 5 areseparate. The ist antenna has 19 segments and reaches to the hind edge of the istabdominal segment ; the ist segment has no group of spines. The remainingmouthparts are as shown ; the maxillae and maxilliped are weakly chitinized. NEW AND RARE CALANOID COPEPODS0-5 301 FIG. 2. Spinocalanus sp. <J. a, cephalothorax, dorsal ; b, abdomen and 5th legs,lateral ; c, 2nd antenna ; d, mandibular palp ; e, ist maxilla ; f, 2nd maxilla ; g,maxilliped ; h, ist leg ; i, 2nd leg ; j, 3rd leg ; k, 4th leg ; 1, 5th leg. Bar scale o-i mmunless indicated. Legs 2-4 are badly damaged in all the specimens. The 5th pair of legs extendsback to the hind edge of the 3rd abdominal segment : the left exopodite is verylong and the endopodite is reduced to a spine ; the right leg is uniramous and doesnot reach the end of the 2nd basipodite of the left leg. REMARKS. The present specimens seem to be conspecific with the male describedas Spinocalanus sp. by Grice & Hulsemann (1967). They described only the 5thpair of legs of their specimen, which differs slightly from that of the present malesin the relative length and segmentation of the right leg. S. longipes Tanaka, 1956is similar to the present specimens but has a much longer left 5th leg. Grice &Hulsemann did not describe the length of this leg. Spinocalanus sp. A MATERIAL EXAMINED : 14 males from 7 RMT i hauls made at 1220-610 m depth in a position i8N 25W, between 13 and 18 November 1969.British Museum (Natural History), reg. no. 1974 : 505-515. Deposited at the 302 H. S. J. ROE FIG. 3. Spinocalanus sp. A cj. a, body, dorsal ; b, base of ist antenna ; c, 2nd antenna ;d, mandibular palp ; e, ist maxilla ; f, 2nd maxilla ; g, maxilliped ; h, ist leg ; i, andleg ; j, 3rd leg ; k, 4th leg ; 1, 5th leg. Bar scale o-i mm unless indicated. DESCRIPTION. Male (Fig. 3) : Body length 1-75-2-20 mm with a mean of1-91 mm (n specimens). The cephalothorax is 1-6 times as long as the 5-segmentedabdomen. The head and ist thoracic segment are fused but thoracic segments4 and 5 are separate. The cephalothorax is slightly arched in lateral view. Theanal segment is very short and telescoped into the 4th abdominal segment. Theist antenna has 19 segments and reaches to the hind edge of the 2nd abdominalsegment ; the ist segment has a stellate group of spines on the dorsal surface andthe first eight segments have large flattened sensory filaments. The remainingmouthparts and the legs are as shown. The ist and 2nd maxillae and the maxillipedare weakly chitinized ; the terminal segment of the maxilliped has a group of spines.The 3rd exopodite segment of the ist leg has a patch of fairly large surface spines ; NEW AND RARE CALANOID COPEPODS 303 the exopodites of the remaining swimming legs are missing in all the specimens.The 5th pair of legs reaches back to the hind edge of the 2nd abdominal segment. REMARKS. These specimens differ from any known Spinocalanus male. 5.stellatus and 5. dorsispinosus both have stellate bundles of spines on the ist antenna,but the males of both species are larger than the present specimens and are struc-turally quite distinct. Females of 5. horridus, S. angusticeps, S. hoplites, S. usitatusand S. pteronus, for all of which the males are unknown, were found in the samehauls as these specimens. Since they may be the undescribed male of a knownspecies they are not considered here to be a new species. Spinocalanus sp. B 2 males taken in an RMT i haul made at 1220-1000 m Deposited at the British MATERIAL EXAMINED depth in a position i8N 25W on 18 November 1969.Museum (Natural History), reg. no. 1974 : 516 and 517. DESCRIPTION. Male (Fig. 4) : Body length 2-05 mm (both specimens). Thecephalothorax is slightly distorted in both specimens and is about 2-6 times as longas the 5-segmented abdomen. The head and ist thoracic segment are fused but a FIG. 4. Spinocalanus sp. B cT. a, body, dorsal ; b, and antenna ; c, ist maxilla ; d, 2ndmaxilla ; e, maxilliped ; f, ist leg ; g, 2nd leg ; h, 3rd leg ; i, 4th leg ; j, 5th leg. Barscale o-i mm unless indicated. 304 H. S. J. ROE thoracic segments 4 and 5 are separate. The forehead has a distinct dorsal pro-trusion. The ist antennae are broken on both specimens ; the ist segment has nospines. The mandible is similar to that of the preceding species, and the remainingmouthparts are as shown. The ist leg has a group of spines on the surface of the2nd exopodite segment ; legs 2-4 are badly damaged in both specimens. The 5thpair of legs reaches back to the hind edge of the 3rd abdominal segment. REMARKS. These relatively large specimens do not agree with the description ofany known Spinocalanus male. They were taken in a haul containing females ofS. horridus, S. hoplites, S, usitatus, S. angusticeps and S. abyssalis. Since they maybe the undescribed male of a known species they are not considered here to be anew species. Family AETIDEIDAEAetideus arcuatus (Vervoort, 1949) Snelliaetideus arcuatus Vervoort, 1949 : 3-7, fig- i ; Park, 1970 : 475 ; Bradford, 19716 : 32. MATERIAL EXAMINED : 14 females and i male taken in 4 RMT I hauls made at790-610 m depth in a position i8N 25 W, between 12 and 14 November 1969.6 females and the male deposited at the British Museum (Natural History), reg. no.1974 : 518-524. DESCRIPTION. Female (Fig. 5a-d) : Body length 1-52-1-67 mm with a meanof i -60 mm (12 specimens). The present females agree generally with Vervoort's(1949) account but show the differences noted by Bradford (19716). Hence theendopod of the ist maxilla has 12 setae instead of 9, the endopodite of the 2nd leghas only one segment - but with a more or less obvious line of fusion, and there is agroup of small spines at the base of the seta on the ist basipodite of the 4th leg.The 2nd maxilla was obscure in Vervoort's specimen ; it is shown here and hassix fine setae on the endopodite - not five as stated by Vervoort. Male (Fig. 5e-m) : Body length 1-22 mm. The head and ist thoracic segmentare fused, as are thoracic segments 4 and 5. There is no rostrum and the hindmargin of the cephalothorax is rounded. The abdomen has four segments, and thefurcal rami are about 3 times longer than wide. The ist antenna has 20 segmentsand reaches to the hind edge of the 2nd abdominal segment. The 2nd antenna andmandibular palp are as in the female ; the mandible blade is only a simple lamella.The ist maxilla is reduced as shown (Fig. 5]) ; the 2nd maxilla is very reduced, andthe maxilliped is similar to that of the female but lacks setae on the ist basipodite.The swimming legs are similar to those of the female. The endopodite of the 2ndleg has one segment but has an obvious line of fusion. There are no spines on theist basipodite of the 4th leg. The 5th leg is uniramous and present on the left sideonly. It has five segments, the last of which has patches of spinules and terminatesin a group of modified setae. REMARKS. Except for the absence of spines on the ist basipodite of the 4th legthe present male differs from the females only in the normal sexual characters. It NEW AND RARE CALANOID COPEPODS 305 FIG. 5. a-d, Aetideus arcuatus ?. a, body, dorsal ; b, forehead and rostrum, lateral ;c, and maxilla ; d, 4th leg. e m, Aetideus arcuatus <$. e, body, dorsal ; f, abdomenand 5th leg, lateral ; g, ist antenna ; h, 2nd antenna ; i, mandibular palp ; j, ist maxilla ;k, maxilliped ; 1, 5th leg ; m, terminal segment, 5th leg. n-p, Aetideopsis carinata ?.n, body, dorsal ; o, rostrum ; p, hind margin of cephalothorax, lateral. Bar scaleo-i mm unless indicated. is immediately distinguished from all the other males in the genus by having nopoints on the hind margin of the cephalothorax. On the basis of the femalesBradford ( 19716) concluded that the only difference between Snelliaetideus Vervoort,1949 and Aetideus Brady, 1883 was the absence of postero-lateral thoracic points.This discovery of the male confirms Bradford's opinion and Snelliaetideus should,I believe, be amalgamated with Aetideus. 306 H. S. J. ROE Aetideopsis carinata Bradford, 1969 Aetideopsis carinata ? ; Roe, igjza : 294, 302, 308. MATERIAL EXAMINED : 14 females taken in 10 Nii3H hauls made at 940-450 mdepth off Fuerteventura (28N i4W), between 15 and 27 November 1965. 2 femalestaken in 2 RMT i hauls made at 500-410 m depth in a position i8N 25W on16 November 1969. REMARKS. The present females (Fig. 5n-p) agree completely with Bradford's(1969) account except for slight differences in the cephalothorax. The notchbetween the rostral points is more rounded than described by Bradford and theshoulders on the head are more pointed - in some specimens they are almost hook-like. Dr Bradford kindly sent me some specimens for comparison and there is nodoubt that they are conspecific. The present specimens measure 2-66-2-96 mmwith a mean of 2-79 mm (16 specimens). Gaetanus ferox With, 1915* Gaetanus sp. Roe, 197205 : 286, 302, 308. MATERIAL EXAMINED : 2 males taken in 2 Nii3H hauls fished at 510-450 mand 300-240 m depth respectively, off Fuerteventura (28N I4W) on 15 and 16November 1965. REMARKS. These specimens conform with the descriptions of With (1915) andGrice and Hulsemann (1967). The right exopodite of the 5th leg has two segmentsthereby agreeing with With's account rather than Grice & Hulsemann's. Thepresent specimens measure 3-12 and 3-27 mm and are slightly smaller than previousrecords. Chiridiella ovata Deevey, 1974 ? Chiridiella macrodactyla ; Scott, 1909 : 79, pi. 36. MATERIAL EXAMINED : i female taken in an RMT i haul made at 1200-1000 mdepth in a position i8N 25W on 18 November 1969. REMARKS. I was preparing to describe this specimen as a new species untilDr G. B. Deevey kindly sent me a copy of her then unpublished manuscript onChiridiella (Deevey 1974). The present female (Fig. 6) agrees almost completelywith her description of C. ovata. The ist maxilla, however, is variable. On theleft side (Fig. 6f) the 3rd inner lobe is absent - contrasting with Deevey's specimens,and the exopodite has three setae ; on the right (Fig. 6g) the 3rd inner lobe ispresent and has one seta, but the exopodite has only two. The ist maxilla ofScott's (1909) specimen differs from both this and Deevey's account. The 2nd lobeof the 2nd maxilla is very small in the present specimen, and the seta on the ist * Park has recently concluded that G. ferox is the male of Gaetanus miles, Giesbrecht 1888. (Park,T. S. 1975. Calanoid copepods of the genera Gaetanus and Gaidius from the Gulf of Mexico. Bull. mar.Sci. 25 : 9-34.) NEW AND RARE CALANOID COPEPODS 37 FIG. 6. Chiridiella ovata ?. a, body, dorsal ; b, body, lateral ; c, 2nd antenna ; d,mandibular palp ; e, mandible blade ; f, ist maxilla, left ; g, ist maxilla, right ; h, 2ndmaxilla ; i, maxilliped ; j, ist leg ; k, 2nd leg ; 1, 4th leg. Bar scale o-i mm unlessindicated. basipodite of the maxilliped is present on one side only. The endopodites of the3rd and 4th legs have only one segment but the lines of fusion are clearly visible. Pseudochirella tuberculata Tanaka, 1957 MATERIAL EXAMINED : 4 females and I male taken in 4 RMT i hauls made at1250-700 m depth in a position i8N 25W, between 12 and 17 November 1969.The male is deposited at the British Museum (Natural History), reg. no. 1974 : 525. 3 o8 H. S. J. ROE FIG. 7. Pseudochirella tuberculata <J. a, body, dorsal ; b, and antenna ; c, mandible ;d, ist maxilla ; e, 2nd maxilla ; f, maxilliped ; g, ist leg ; h, 2nd leg ; i, 4th leg ; j, 5thleg ; k, 5th leg, left exopodite ; 1, 5th leg, right 3rd exopodite segment. Bar scale o-i mmunless indicated. DESCRIPTION. Male (Fig. 7) : Body length 576 mm. The cephalothorax isrobust and oval ; it is 3-1 times as long as the 5-segmented abdomen. The headand ist thoracic segment are fused. Thoracic segments 4 and 5 are separate ; the5th segment is rounded laterally and has no spines. The rostrum is a single spikedirected ventrally. The ist antenna has 21 segments and reaches to the hind edgeof the 3rd abdominal segment. The remaining mouthparts are as shown. The NEW AND RARE CALANOID COPEPODS 309 exopodite of the and antenna is 1-4 times as long as the endopodite. The ist and2nd maxillae and the mandible blade are reduced. The exopodite of the ist leghas three segments and the endopodite has a comb of spines. The endopodite ofthe 2nd leg has only one segment but with an evident line of fusion ; the 3rd and4th legs have 3-segmented endopodites. The 5th pair of legs is as shown : on theleft side (Fig. 7k) the 2nd exopodite segment has two strong points, and the 3rdsegment is kidney-shaped with bunches of hairs ; the 3rd segment of the rightexopodite (Fig. 7!) has two lamellae - one on a central swelling and one terminatingthe segment. REMARKS. This specimen does not fit the description of any known male Pseudo-chirella. It is nearest to Tanaka & Omori's (1969) account of P. polyspina Brodsky,1950, but it is larger and differs in the detail of the mouthparts and 5th pair oflegs - especially the right exopodite. It was found in association with females ofP. tuberculata and P. obtusa (Sars, 1905). It does not agree with the descriptionof the male P. obtusa given by Vervoort (1949). Apart from sexual characters, itdiffers from Tanaka's (1957) account of the female P. tuberculata only in the ist leg,where his specimen had only two exopodite segments and had no comb of spines onthe endopodite. The present females, however, show some variation in the degreeof fusion of these exopodite segments and they do have a comb of spines. Specificattribution of Pseudochirella males is difficult but because the structures common toboth sexes are so similar I believe that this specimen is the hitherto unknown male ofP. tuberculata. This is the first Atlantic record of this species. Valdiviella minor Wolfenden, 1911 ? Valdiviella brevicornis ; Scott, 1909 : 78, pi. 22 ; Brodsky, 1950 : 223, fig. 137 (male only). MATERIAL EXAMINED : 2 females and i male from 2 RMT i hauls made at1250-1000 m depth in a position i8N 25W on 17 and 18 November 1969. Themale is deposited at the British Museum (Natural History), reg. no. 1974 : 526. DESCRIPTION. Male (Fig. 8) : Body length 4-96 mm. The cephalothorax is3 times as long as the 5-segmented abdomen. The head and ist thoracic segmentare completely fused ; thoracic segments 4 and 5 are partially so. The head hasprominent shoulders and the rostrum has two short spikes. The hind margin ofthe cephalothorax is rounded laterally and has no spines. The ist antennae areboth broken off. The remaining mouthparts are identical to those described bySewell (1929) for the male V. insignis. The legs are as shown. The ist leg hasthree exopodite segments but the 2nd, 3rd and 4th legs have only two. The 2ndleg has a single endopodite segment and legs 3 and 4 have two endopodite segments.The exopodites and endopodites of legs 2-4 all show signs of fusion between seg-ments. The 5th pair of legs is typical of the genus. The basipodite segments aremuch longer on the left side than on the right, but conversely the left endopod ismuch shorter than the right. The ist and 2nd exopodite segments of the left legeach have a small spine. The ist and 2nd exopodite segments of the right leg arepartially fused and are drawn out into a long curved point. H. S. J. ROE FIG. 8. Valdiviella minor <$. a, body, dorsal ; b, forehead and rostrum, lateral ; c, 2ndantenna ; d, mandibular palp ; e, ist maxilla ; f, 2nd maxilla ; g, maxilliped ; h, istleg ; i, 2nd leg, j, 3rd leg ; k, 4th leg ; 1, 5th leg. Bar scale o-i mm unless indicated. REMARKS. This male was found with two undoubted females of V. minor. Boththe females have the very characteristic chitinous ridge on the dorsal side of thegenital segment and are identical to the descriptions of Wolfenden (1911) and Sewell(1929). They both measure 5-44 mm and are thus slightly larger than previousrecords. The present male is distinguished both in structure and size from males ofV. insignis, V. oligarthra and V ' . ignota. Of the three smaller species, V. imperfectais very poorly known, but females of V. brevicornis and V. minor can be identified NEW AND RARE CALANOID COPEPODS 311 easily. Scott (1909) described a male which he doubtfully attributed to V. brevi-cornis. Sewell (1929) also described a male as V. brevicornis and noted slightdifferences between his specimen and Scott's. Vervoort (1957) stated that Sewell'sspecimen (s) were 'undoubted males of V. brevicornis'. The present male is verysimilar to that described by Scott. As far as can be seen from his description itdiffers only in having the 4th and 5th thoracic segments and the ist and 2nd exo-podite segments of the right 5th leg partially fused instead of completely so. Sewell'sspecimen had a short spine on the posterior thoracic margin and a 3-segmentedexopodite in the 2nd leg. I believe that Scott's male and the present specimen areconspecific. If the differences between them and Sewell's specimen of V. brevicornisare of specific value they may well be the hitherto undescribed male of V. minor,especially in view of the present association with females of this species. Family PHAENNIDAEXanthocalanus agilis Giesbrecht, 1892 MATERIAL EXAMINED : i male taken in an RMT i haul made at 200-110 m depthin a position i8N 25W on 16 November 1969. Deposited at the British Museum(Natural History), reg. no. 1974 : 527. REMARKS. The present male (Fig. ga-d) measures 2-58 mm and agrees withGiesbrecht's (1892) description except for the 5th pair of legs. Giesbrecht describedonly the elongate left leg but it is not clear from his account whether the absence ofthe right leg was accidental or not. Giesbrecht & Schmeil (1898) said that the rightleg is absent. Other than restatements of Giesbrecht's original account I know ofno other description of the male X. agilis. The general appearance of the presentmale is so similar to that of the female X. agilis that there can be no doubt that theyare conspecific. The right leg of the 5th pair has five segments and is very short,reaching only to the end of the ist segment of the left leg. I believe that Giesbrecht'sdescription is incomplete and that the male X. agilis has a pair of 5th legs in commonwith other males of this genus. Family SCOLECITHRICIDAE Several authors have commented on the difficulties in classifying species withinthis large and complex family. Most recently, Bradford (1973) partially reviewedboth the Phaennidae and the Scolecithricidae, redefining both families and some oftheir genera. Within the Scolecithricidae she found apparently consistent differ-ences in the setation of the ist maxilla of various genera, although the number ofspecies which she examined was rather small. She was, however, unable to placedefinitely a large number of species into her redefined genera, either because of theirstructure or, presumably, because their descriptions are inadequate and many oftheir males unknown. It is therefore impossible, at the moment, to verify hernarrower generic definitions for many species in this family. Re-examination ofmore species will clarify the acceptable limits for each genus. 17 312 H. S. J. ROE FIG. 9. a-d, Xanthocalanus agilis <?. a, body, dorsal ; b, last thoracic segment andabdomen, lateral ; c, 5th leg ; d, 5th leg, terminal segment left leg. e-g, Scottocalanusthomasi ?. e, body, dorsal ; f , forehead and rostrum, lateral ; g, genital segment,half lateral, h-r, Scaphocalanus sp. $. h, body, dorsal ; i, and antenna ; j, mandibularpalp ; k, ist maxilla ; 1, 2nd maxilla ; m, maxilliped ; n, ist leg ; o, 2nd leg ; p, 4th leg,basipodite segments ; q, 5th leg ; r, 5th leg, 3rd exopodite segment left leg. s, Scolecithri-cella laminata ?, 2nd maxilla endopod. t-v, Scolecithricella laminata <$. t, body, dorsal ;u, 5th leg ; v, 5th leg, 3rd exopodite segment left leg. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 313 The present collections contain several rare and new scolecithricid species.Some of these agree well with Bradford's generic definitions but some do not. A newgenus is described for some of the latter species but others are maintained withinexisting genera pending further study of the variation within these. Scottocalanus thomasi Scott, 1909 ? Scottocalanus backusi Grice, 1969 : 451, figs 16-34. MATERIAL EXAMINED : 3 females and i male taken in 3 RMT i hauls made at500-210 m depth in a position i8N 25W, between 12 and 16 November 1969. REMARKS. The present females (Fig. ge-g) agree with Grice's (1969) descriptionof 5. backusi which is, in turn, very similar to 5. thomasi. The females of thesetwo species apparently differ only in the structure of their genital segment whereS. backusi has a pair of spine-like protrusions. Sewell (1929), however, describedthe genital segment of 5. thomasi as having 'a finger-like backwardly directedprojection near the posterior margin a little to the left of the middle line'. Scott(1909) made no mention of this. (Sewell's description of three endopodite segmentson the ist maxilla of S. thomasi may be atypical since Scott described this as being'nearly similar' to that of S. securifrons which has only two endopodite segments.)I have examined a female specimen of 5. thomasi in the collections of the BritishMuseum (Natural History), which was taken at 'Investigator' Station 670 andpresumably identified by Sewell. It has two spine-like projections on its genitalsegment exactly as shown here (Fig. 96, g). These projections are not alwaysconspicuous, especially if they are bent back alongside the genital segment, and theycan be easily overlooked, as they apparently were on this specimen. For thisreason I think it probable that 5. backusi and 5. thomasi are conspecific. Thepresent female specimens measure 5-92-6-08 mm and the male 5-92 mm. Lophothrix latipes (T. Scott, 1894) MATERIAL EXAMINED : 5 females and 12 males taken in 7 Nii3H hauls madeat 800-90 m depth off Fuerteventura (28N I4W), between 13 and 28 November1965. 13 females and 28 males taken in 9 RMT i hauls made at 700-49 m depthin a position i8N 25W, between 13 and 16 November 1969. 7 males are depositedat the British Museum (Natural History), reg. no. 1974 : 528-535. DESCRIPTION. Male (Fig. 10) : Body length 2-96-3-19 mm with a mean of3-05 mm (26 specimens). The cephalothorax is 3-4 times as long as the 5-segmentedabdomen. The head and ist thoracic segment are fused ; thoracic segments 4 and 5are separate. The head has a low crest and the forehead is angular in lateral view.The rostrum has two thick filaments. The head has prominent shoulders. Thehind margin of the ist thoracic segment is undulating and the ventro-lateral marginof the 5th thoracic segment is abruptly cut off. The ist antenna has 20 segmentsand reaches to the end of the cephalothorax. The remaining mouthparts are asshown. The mandible blade, ist and 2nd maxillae and the maxilliped are reduced. H. S. J. ROE FIG. 10. Lophothrix latipes (?. a, body, dorsal ; b, forehead and rostrum, lateral ; c,thoracic segments 4 and 5, lateral ; d, 2nd antenna ; e, mandibular palp ; f, ist maxilla ;g, 2nd maxilla ; h, maxilliped ; i, ist leg ; j, 2nd leg ; k, 3rd leg ; 1, 4th leg ; m, 5th leg ;n, 5th leg, last exopodite segment right leg ; o, 5th leg, terminal exopodite segmentsleft leg ; p, 5th leg, last endopodite segment left leg. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 315 The endopod of the 2nd maxilla has six sensory setae, none of which are brush-like.The first four pairs of legs are identical to those of the female. The 5th pair of legsare asymmetric and reach back to the hind edge of the 3rd abdominal segment.In the right leg, the exopodite has four segments, the last of which is flattened androughly triangular in shape ; the endopodite is short and has two segments. Theleft leg has longer basipodite segments, the exopodite has three segments and theendopodite has two ; the last exopodite segment has two wide setae and numerousspines ; the endopodite terminates in a ribbon-like seta. REMARKS. Wilson (1950) described a male as L. latipes which differs in manyrespects from the present specimens, particularly in its mouthparts, which weredescribed as 'like those of the female', and in its 5th pair of legs. There is someconfusion between Wilson's description and his figures. His figure of the lateralview of the complete animal shows a distinct crest on the forehead, but he describedthis as 'evenly rounded with no trace of a crest'. The same figure shows a prominentspine on the posterior margin of the last thoracic segment ; this is described as'smoothly rounded' in the text. The present males are so similar to the femalesin all structures common to both sexes that I have no doubt that they are con-specific. The mouthparts are typically reduced but the cephalothorax, particularlythe characteristic low crest and last thoracic segment, and the swimming legs, areidentical to those of the female. Scaphocalanus difficilis sp. nov. MATERIAL EXAMINED : 64 females from 8 RMT i hauls made at 700-210 m depthin a position i8N 25W, between 12 and 17 November 1969. The holotype wascaught between 600 and 515 m on 14 November 1969. Deposited at the BritishMuseum (Natural History), holotype reg. no. 1974 : 536, paratypes reg. no.1974 : 537-557- DESCRIPTION. Female (Fig. n) : Body length 1-29-1-44 mm with a mean of1-38 mm (44 specimens). The holotype measures 1-37 mm. The cephalothoraxis elongate and is 3-5 times as long as the 4-segmented abdomen. The head andist thoracic segment are fused as are thoracic segments 4 and 5. The rostrum hastwo tapering filaments. The hind margin of the cephalothorax is flattened inlateral view. The abdominal segments and furcal rami have the following pro-portional lengths : Seg. 1 + 23 4 5 F. rami 30 : 19 : 19 : 13 : 19 = 100 The furcal rami are twice as long as wide. The ist antenna is broken in allspecimens ; the near-basal segments are flattened and typical of the genus. Inthe 2nd antenna the endopodite is 1-4 times longer than the exopodite. In theist maxilla the ist inner lobe has only two surface setae and the 3rd inner lobe hasonly three setae. The endopod of the 2nd maxilla has five brush-like setae, ofwhich two are enlarged, and three flattened worm-like setae. The ist pair of legshas no external spine on either the ist or 2nd exopodite segments. The outer spine H. S. J. ROE n FIG. ii. Scaphocalanus difficilis ?. a, body, dorsal; b, abdomen and last thoracicsegment, lateral ; c, rostrum, lateral ; d, rostrum, ventral ; e, 2nd antenna ; f, istmaxilla ; g, 2nd maxilla ; h, maxilliped ; i, ist leg ; j, 2nd leg ; k, 3rd leg ; 1, 3rd leg,ist basipodite segment (paratype) ; m, 4th leg ; n, 5th leg. Bar scale o-i mm unlessindicated. on the ist exopodite segment of the 2nd leg is short. The basipodite segments ofthe 3rd and 4th legs have varying numbers of small surface spines. The 5th leghas a single segment on each side carrying two spines ; the inner spine is 1-8 timeslonger than the terminal one. REMARKS. The small size of the present species distinguishes it from all otherScaphocalanus species which have a well-developed 5th pair of legs. S. longifurca(Giesbrecht, 1888) is only slightly larger ($ 1-48-1-90 mm -limits from Tanaka,1961, and Grice & Hulsemann, 1965). The present species differs from Giesbrecht's(1892) description of 5. longifurca in the following characters : (a) the endopod of the2nd antenna longer than the exopod, (b) a relatively shorter abdomen, (c) differentproportional lengths of the abdominal segments, (d) no outer spine on the 2nd NEW AND RARE CALANOID COPEPODS 317 exopodite segment of the ist leg, (e) a more or less straight outer spine on the istexopodite segment of the 2nd leg, and (f) different proportional lengths of the spineson the 5th leg. Giesbrecht (1892), Vervoort (1951) and Tanaka (1961) all describedspines on the basal segments of legs 2-4 in 5. longifurca. Vervoort said that theseform a 'dense covering' on the 3rd leg and according to Tanaka these same jointsare 'coarsely covered'. Without illustrations it is difficult to decide on the degree ofspinulation in these descriptions. The present specimens have a varying number ofsmall spines on these basal segments but they are relatively few in number.Tanaka's (1961) specimens of S. longifurca have the endopod of the 2nd antennalonger than the exopod, and the cephalothorax is 3-5 times as long as the abdomen.They differ from the present females, however, in the proportional lengths of theabdominal segments and in the shape of the hind margin of the cephalothorax. 5. siibbrevicornis (Wolfenden, 1911) is very similar, if not identical, with 5. longi-furca (Farran, 1929 ; Vervoort, 1951 ; Tanaka, 1961). In some of the slight differ-ences between these two species - the proportions of the rami of the 2nd antenna,the straight outer spine of the ist exopodite segment of the 2nd leg of 5. subbrevi-cornis, and the degree of spinulation of the basal joints of legs 2-4-5. subbrevicornisis structurally more similar to the present species. Moreover, the relative length ofthe abdomen and the proportional lengths of the abdominal segments are alsosimilar to the present females (Tanaka, 1961). 5. subbrevicornis is, however,larger, females measuring 1-60-2-10 mm (extremes from Vervoort, 1957, and Grice& Hulsemann, 1965) and there are differences in the ist and possibly also the 5thpair of legs. Whether or not these slight differences merit separation into three species isquestionable. It may be necessary to amalgamate two or all of them in futurebut for the moment at least they can remain separate. The ist maxilla of 5.difficilis differs from Bradford's (1973) definition of the genus in having only twosurface setae on the ist inner lobe and three setae on the 3rd inner lobe. This lastcharacter is shared with other scaphocalanids, however, e.g. 5. similis and S. invalidus(Hure & Scotto di Carlo, 1968), and the general structure of the present species is sosimilar to Scaphocalanus that there can be no doubt that it belongs to this genus. Scaphocalanus sp. MATERIAL EXAMINED : 3 males taken in 2 RMT i hauls, one at 900-800 m depthand the other at 500-410 m, in a position i8N 25W on 14 and 16 November 1969.Deposited at the British Museum (Natural History), reg. no. 1974 : 558-560. DESCRIPTION. Male (Fig. gh-r) : Body length 1-98-2-13 mm with a mean of2-05 mm (3 specimens). The cephalothorax is 1-9 times as long as the 5-segmentedabdomen. The head and ist thoracic segment are completely fused, thoracicsegments 4 and 5 are partially so. The rostrum has two thickened filaments. Theabdominal segments and furcal rami have the following proportional lengths : Seg. i 2 3 4 5 F. rami 13 : 26 : 21 : 29 : 3 : 8 = 100 318 H. S. J. ROE The ist antenna has 20 segments and reaches back to the hind edge of the 2ndabdominal segment ; the first eight segments have long sensory setae. The re-maining mouthparts are as shown. The swimming legs are damaged in all specimens.The outer spine on the ist exopodite segment of the 2nd leg is moderately long andstraight. The basipodite segments of legs 2-4 are covered with small surface spineswhich are particularly dense on legs 3 and 4. The 5th pair of legs reaches back tothe end of the abdomen. On the left side the exopodite is much shorter thanthe endopodite ; the last exopodite segment has some coarse setae and a lamella,and the endopodite has no marked swelling along its length. REMARKS. These specimens cannot be identified with any of the known Scapho-calanus males. The abdomen, with an unswollen 2nd segment, is similar to thatdescribed for 5. echinatus by Tanaka (1961) but the present males are larger andhave a different 5th pair of legs. As the males of several Scaphocalanus species areunknown and as these specimens are all damaged they cannot be described as newspecies. Scolecithricella obscura sp. nov. MATERIAL EXAMINED : 10 females taken in 3 RMT i hauls made at 1250-800 mdepth and i female taken in an RMT i haul at 500-410 m. The holotype wastaken at 1250-1000 m on 16 November 1969. All hauls were made in a positioni8N 25W, between 14 and 18 November 1969. Deposited at the British Museum(Natural History), holotype reg. no. 1974 : 561, paratypes reg. no. 1974 : 562-570. DESCRIPTION. Female (Fig. 12) : Body length 1-82 -2-13 mm with a mean of2-04 mm (10 specimens). The holotype measures 1-90 mm. The cephalothoraxis an elongated oval, and is 5-9 times as long as the 4-segmented abdomen. Thehead and ist thoracic segment and thoracic segments 4 and 5 are fused. Therostrum has two thick filaments. The hind margin of the cephalothorax is indentedlaterally. The genital segment is as long as the combined length of the two sub-sequent segments. The furcal rami are about as long as wide. The ist antenna isbroken in all specimens. The remaining mouthparts are as shown. The endopodof the 2nd maxilla has five brush-like setae, two of which are much larger than theothers, and three flattened worm-like setae. In the maxilliped the ist basipoditeis about two-thirds the length of the second, and carries one brush-like and twoworm-like setae. The legs are as shown. There is no external spine on the istexopodite segment of the ist leg. All the outer edge spines on the exopoditesegments of the 2nd leg are long. The 5th pair of legs is uniramous and symmetrical.On each side there is a single segment, incompletely divided into three, having asmall external spine - which is absent on one side of one of the paratypes, a longerterminal spine and a long stout internal spine garnished with teeth. REMARKS. This species agrees generally with Scolecithricella as defined byBradford (1973). It can be distinguished from all others in the genus by its 5th pairof legs, which are more like those found in Amallothrix and are very similar to thoseof ? Amallothrix valens (Farran, 1926). The great development of two of the NEW AND RARE CALANOID COPEPODS 319 FIG. 12. Scolecithricella obscura ?. a, body, dorsal ; b, rostrum, ventral ; c, last thoracicsegment and abdomen, lateral ; d, and antenna ; e, mandibular palp ; f, ist maxilla ;g, 2nd maxilla ; h, 2nd maxilla endopod (paratype) ; i, ist leg ; j, 2nd leg ; k, 3rd leg,exopodite ; 1, 3rd leg, endopodite segments 2-3 ; m, 4th leg; n, 5th leg. Bar scaleo-i mm unless indicated. 320 H. S. J. ROE brush-like setae of the 2nd maxilla approaches the condition seen in Amallothrixalter a (p. 331). Few authors have figured these setae in detail in Scolecithricellabut in some species two of the brush-like setae are shorter and thicker than theothers, though not, apparently, to the extent seen here (for example, see Rose,1942). Scolecithricella laminata (Farran, 1926) Scolecithrix laminata Farran, 1926 : 265, pi. 8 ; Bradford, 1973 : 143, 145. Scolecithricella laminata ; Grice & Hulsemann, 1965 : 239, figs i4g-m, I5a-j ; 1967 : 16 ; Park, 1960 : 476 ; Wheeler, 1970 : 8 ; Roe, 1972^ : 304, 310 ; 1972^ : 540.S. lamellifer Tanaka, 1962 : 78, fig. 146.? Amallothrix profunda Brodsky, 1950 : 263, fig. 172 ; Bradford, 1973 : 144. MATERIAL EXAMINED : 3 females and i male from an Nii3H haul made at960-800 m depth off Fuerteventura (28N I4W) on 25 November 1965. 15females and 3 males from 5 RMT i hauls made at 1250-700 m depth in a positioni8N 25W, between 12 and 18 November 1969. 3 females and the males aredeposited at the British Museum (Natural History), reg. no. 1974 : 571-577- DESCRIPTION. Female (Fig. 95) : Body length 1-82-2-28 mm with a mean of1-94 mm (14 specimens). Redescription of these females is unnecessary as theygenerally agree with Farran's (1926) account but have the modifications describedby Grice & Hulsemann (1965) and Tanaka (1962). The structure of the ist and 2ndmaxillae, however, seems in need of clarification. The ist maxilla has the followingnumber of setae : 5 large and 2 small on the ist outer lobe, 8 on the exopod, 5 on thefused 2nd and 3rd endopodite segment, 2 on the ist endopodite segment, 6 on the2nd basipodite, 4 on the 3rd inner lobe, 2 on the 2nd inner lobe and 13 on the istinner lobe - of which 4 are on the surface as opposed to the edge. The 2nd maxilla(Fig. 95) has 8 sensory setae on the endopodite of which only 2 are short and brush-like and 6 are long and worm-like. Male (Fig. 9t-v) : Body length 2-20-2-43 mm with a mean of 2-35 mm (3 speci-mens). The cephalothorax is 2-3 times as long as the 5-segmented abdomen.The head and ist thoracic segment and thoracic segments 4 and 5 are fused. Therostrum has two stout filaments. The hind margin of the cephalothorax is smoothlyrounded. The ist antenna has 19 segments and reaches to the end of the furcalrami. The remaining mouthparts are generally slightly smaller and more weaklychitinized than in the female ; their setation patterns are identical to those of thefemale except in the following respects. The basipodite of the mandibular palphas no setae ; in the ist maxilla the ist outer lobe has six large and two small setae,the exopodite has nine and the 2nd basipodite has five setae. The endopodite of the2nd maxilla is as in the female with six long filaments and two shorter brush-likesetae. In all the specimens the 2nd and 3rd exopodite segments of legs 2-4 aremissing but the remainder of the swimming legs are identical to those of the female.The 5th pair of legs are asymmetrical and resemble those of Scaphocalanus. In theright leg the exopodite has three segments, the last of which is roughly triangularin shape. In the left leg the exopodite is much shorter than the 2-segmented NEW AND RARE CALANOID COPEPODS 321 endopodite ; the 3rd exopodite segment has bunches of fine hairs and two largesetae. REMARKS. Apart from the normal sexual characters these males are so similar tothe females that there is no doubt that they are the same species. Brodsky's (1950)description of A. profunda is incomplete but the structure of the 5th pair of legs,especially the short exopodite of the left leg, is very similar to that of the presentspecimens. In her review of the Scolecithricidae, Bradford (1973) could not place this speciesinto a genus. She suggested that it may belong to Amallothrix or to an unnamedgroup typified by Scolecithrix auropecten Giesbrecht, 1892. The combination ofsix worm-like and two brush-like setae on the endopod of the 2nd maxilla wouldexclude this species from the Scolecithricidae as defined by Bradford. However,this definition of the 2nd maxilla would also exclude Parascaphocalanus Brodsky,1955 although Bradford admits this genus to the family. In other respects thepresent species is a typical scolecithricid. The combination of the 2nd maxilla,a Lophothrix-type ist maxilla with four surface setae on the ist inner lobe, well-defined rostral filaments, a spine on the ist exopodite segment of the ist leg, and,in the male, relatively unreduced mouthparts and a Scaphocalanus-type 5th pair oflegs, excludes this species from those genera defined by Bradford and from thosenot described by her. As mentioned earlier (p. 311), she had difficulty in definitelyassigning a large number of species to genera, and rather than create a new mono-typic genus for the present species it seems better to place it into Scolecithricella asdefined by Sars (1902), until examination of more species clarifies the limits of anynarrower generic definitions. Scolecithricella aspinosa sp. nov. MATERIAL EXAMINED : 9 females taken in 2 RMT i hauls - one fished at1250-1000 m depth on 17 November 1969 and the other (containing the holotype)at i22O-iooom on 18 November 1969. Both hauls were made at i8N 25W.Deposited at the British Museum (Natural History), holotype reg. no. 1974 : 578,paratypes reg. no. 1974 : 579 -586. DESCRIPTION. Female (Fig. 13) : Body length 3-19-3-65 mm with a mean of3-36 mm (8 specimens). The holotype measures 3-19 mm. The cephalothorax isoval and robust, and is 3-6 times as long as the 4-segmented abdomen. The headand ist thoracic segment are completely fused, thoracic segments 4 and 5 arepartially so. The forehead is rounded in lateral view ; the rostrum has two fila-ments with thick sausage-shaped bases. The hind edge of the cephalothoraxis slightly indented. The genital segment is longer than the combined length of thethree subsequent segments. The furcal rami are about as long as wide. Theist antenna has 23 segments and reaches to the end of the furcal rami. Theremaining mouthparts are as shown. The exopod of the ist maxilla has a groupof spines. The endopod of the 2nd maxilla has five brush-like setae and threeworm-like setae. In the holotype the worm-like setae are very wrinkled but this may 322 H. S. J. ROE a FIG. 13. Scolecithricella aspinosa $. a, body, dorsal ; b, forehead and rostrum, lateral ;c, last thoracic segment and abdomen, lateral ; d, 2nd antenna ; e, mandibular palp ;f, ist maxilla ; g, 2nd maxilla, exopod ; h, 2nd maxilla, endopod ; i, maxilliped ; j, istleg; k, 2nd leg; 1, 2nd leg, exopodite segments 2-3 (paratype) ; m, 3rd leg; n, 4thleg ; o, 4th leg endopod anterior surface ; p, 5th leg. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 323 be an artifact of preservation as in one of the paratypes these setae are simpleflattened filaments. The basal segment of the maxilliped has one brush-like andtwo worm-like setae. The legs are as shown. The ist leg has no outer spine oneither the ist or 2nd exopodite segment. In the 2nd leg the external spine of the istexopodite segment is short, and the 2nd endopodite segment has a raised lobecarrying four spines. The endopods of legs 2-4 all have long spines on their pos-terior surfaces ; their anterior surfaces generally have small spines but those on the4th leg are fairly large. The 5th leg is symmetrical and uniramous. On each sidethere are two segments with the second having a distinct line of fusion and carryinga short terminal spine and a very long internal spine. REMARKS. Scolecithricella aspinosa and the following species are distinct inhaving no external spines on the ist and 2nd exopodite segments of the ist leg.Discussion of the affinities of both these species follows the description of Scole-cithricella canariensis on p. 325. Scolecithricella canariensis sp. nov.Scolecithricella sp. 3 Roe, 19720 : 304, 310 ; 19726 : 540. MATERIAL EXAMINED : 2 females from 2 Nii3H hauls made at 960-825 m depthoff Fuerteventura (28N I4W) on 25 and 27 November 1965. 9 females from6 RMT i hauls made at 1220-610 m depth in a position i8N 25W, between12 and 18 November 1969. The holotype was taken at 790-700 m on 12 November1969. Deposited at the British Museum (Natural History), holotype reg. no.1974 : 587, paratypes reg. no. 1974 : 588-597. DESCRIPTION. Female (Fig. 14) : Body length 3-12-3-42 mm with a meanof 3-30 mm (n specimens). The holotype measures 3-19 mm. The cephalothoraxis elongate and 5-1 times as long as the 4-segmented abdomen. The head and istthoracic segment are fused, as are thoracic segments 4 and 5. The forehead issomewhat angular in dorsal view. The rostrum has two fairly long filaments. Thehind margin of the cephalothorax is indented laterally. The genital segment isa little longer than the subsequent segment ; the furcal rami are slightly longer thanwide. The ist antenna has 23 segments, the last of which extends past the furcalrami. The remaining mouthparts are as shown. The endopod of the 2nd maxillahas three ribbon-like setae and four brush-like setae. The basal segment of themaxilliped has two worm-like and one brush-like seta. The legs are as shown.The ist leg has no external spine on either the ist or 2nd exopodite segment. The2nd leg has a short outer edge spine on the ist exopodite segment and a distinctraised lobe bearing six spines on the 2nd endopodite segment. The 5th pair of legsis symmetrical and uniramous. On each side there is a single segment, which bearstraces of fusion, attached to a common basal segment. Each terminal segment hasa short end spine and a longer robust internal spine armed with short teeth. REMARKS. Scolecithricella canariensis and S. aspinosa are closely related. Bothhave (a) no trace of an external spine on the ist and 2nd exopodite segments of theist leg, (b) a Lophothrix-type ist maxilla, (c) uniformly small heads to the brush-like 324 H. S. J. ROE FIG. 14. Scolecithricella canariensis ?. a, body, dorsal ; b, forehead and rostrum, lateral ;c, last thoracic segment and abdomen, lateral ; d, 2nd antenna ; e, mandibular palp ;f, ist maxilla ; g, 2nd maxilla, exopod ; h, 2nd maxilla, endopod ; i, maxilliped ; j, istleg ; k, 2nd leg ; 1, 3rd leg ; m, 4th leg ; n, 5th leg. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 325 setae of the 2nd maxilla (S. canariensis apparently only has four of these setae),(d) a similar-shaped maxilliped, and (e) a short external spine on the ist exopoditesegment of the 2nd leg and, in the same limb, a raised lobe on the 2nd endopoditesegment. The two species are clearly distinct, however, differing from each otherin body shape, the detailed setation of the ist maxilla and the structure of the legs.They both show similarities with Scaphocalanus bogorovi Brodsky, 1955. Theexopodite of the ist leg is identical (see Grice & Hulsemann, 1965), and the maxil-liped and the endopod of the 2nd maxilla are also very similar. There are differences,however, in the structure of the ist maxilla and in the legs. 5. bogorovi seems atbest a very peculiar Scaphocalanus and should probably be removed from this genus.As with 5. laminata (p. 321) it is impossible to place either of the two presentspecies into any of the scolecithricid genera defined by Bradford (1973), or into anyof the remaining genera in this family. Both have some affinity to Lophothrixbut also show a general resemblance to Amallothrix. The peculiar structure oftheir ist leg, however, is constant in all the present specimens and it may be necessaryto establish a new genus for these species and also perhaps for S. bogorovi. Pendingthe discovery of the males and determination of the acceptable limits within eachgenus both species are temporarily placed within Scolecithricella Sars, 1902. Amallothrix falcifer (Farran, 1926) Scolecithrix falcifer Farran, 1926 : 262, pi. 8 ; Bradford, 1973 : 143.Scolecithricella sp. 2 Roe, 1972^ : 296, 304, 310 ; 1972^ : 540. MATERIAL EXAMINED : n females and 2 males taken in 10 Nii3H hauls made at960-510 m depth off Fuerteventura (28N I4W), between 24 and 28 November1965. 10 females and 7 males taken in 7 RMT i hauls made at 1250-700 m depthin a position i8N 25W, between 12 and 17 November 1969. One female and5 males are deposited at the British Museum (Natural History), reg. no.1974 : 598-603. DESCRIPTION. Female (Fig. I5a-m) : Body length 1-82-2-20 mm with a meanof 2-00 mm (20 specimens). The cephalothorax is oval and robust ; it is 3-8 timesas long as the 4-segmented abdomen. The head and ist thoracic segment are fused,as are thoracic segments 4 and 5. The forehead is rounded, the two rostral filamentsare thickened over most of their length. The hind margin of the cephalothorax isslightly indented. The genital segment is almost as long as the combined length ofthe three subsequent segments. The furcal rami are only slightly longer thanwide. The ist antenna has 23 segments and reaches back to the anal segment.The maxilliped has a sensory seta in the middle of the ist basipodite. The re-maining mouthparts are as shown. The 2nd maxilla has five brush-like setae on theendopodite and at least two ribbon-like setae. The legs are as shown. The internaledge of the 2nd basipodite of the 2nd leg has a characteristic group of spines. Thesurfaces of legs 2 and 3 are heavily ornamented with spines. Male (Fig. I5n-r) : Body length 1-98-2-81 mm with a mean of 2-50 mm (9specimens). The cephalothorax is 3 times longer than the 5-segmented abdomen. 326 H. S. J. ROE FIG. 15. a-m, Amallothrix falcifer ?. a, body, dorsal ; b, forehead and rostrum, lateral ;c, last thoracic segment and abdomen, lateral ; d, 2nd antenna ; e, mandibular palp ;f, ist maxilla ; g, 2nd maxilla ; h, 2nd maxilla, brush-like seta head ; i, ist leg ; j, 2ndleg ; k, 3rd leg ; 1, 4th leg ; m, 5th leg. n-r, Amallothrix falcifer <$. n, body, dorsal ;o, 2nd maxilla, endopod ; p, 5th leg ; q, 5th leg, last exopodite segment right leg ; r, 5thleg, terminal segments left leg. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 327 The head and ist thoracic segment and thoracic segments 4 and 5 are fused. Theanal segment is very short. The ist antenna has 19 segments and reaches back tothe hind edge of the 3rd abdominal segment. The setation of the mouthparts isidentical to that of the female except in the following respects. In the 2nd antennathe exopodite has three terminal setae and the ist endopodite segment has a bunchof hairs. The ist maxilla is slightly variable ; in one specimen the ist outer lobehas eight setae, in another the 3rd inner lobe has three, and in both the 2nd basi-podite has five and the ist endopodite has three setae. In the 2nd maxilla lobes2-5 each has an enlarged seta typical of other Amattothrix males and the endo-podite has five brush-like setae and three ribbon-like setae. Two of the brush-likesetae are short and one of these is conspicuously larger than the others. Theswimming legs are identical to those of the female. The 5th pair of legs reachesback to the end of the abdomen. On the right leg the endopod is a short singlesegment and the last exopodite segment is leaf-shaped. On the left leg the endopodends in a flagellum and the 3rd exopodite segment has a row of setae and a lamella. REMARKS. A . falcifer is difficult to recognize from Farran's (1926) account andapart from Wilson's (1950) figure of the female 5th legs it has never been redescribed.I have compared the present material with the type specimen held in the BritishMuseum (Natural History). The ist maxilla of the type is not mounted and theendopodite of the 2nd maxilla is rather obscure - it does, however, have at leastfour brush-like setae and three ribbon-like setae. In all other respects the presentfemales are identical to the type. The 2nd leg of Farran's specimen has the charac-teristic group of spines on the 2nd basipodite although this is neither described norfigured by him. Apart from sexual characters and the slightly variable setation ofsome of the mouthparts the males are identical to the females. They have thedistinct group of spines on the 2nd basipodite of the 2nd leg. There are, apparently,two size groups in the males. Two specimens each measured 1-98 mm but theremaining seven were between 2-58 and 2-81 mm in length. There are no structuraldifferences between the large and small specimens and both are considered to beconspecific. Scolecithricella denticulata Tanaka, 1962 has a similar group of spineson the basipodite of the 2nd leg but its 5th pair of legs differs from the present males.Bradford (1973) could not definitely place A. falcifer into Amallothrix following herrevision of the Scolecithricidae. The present specimens are in complete agreementwith her definition of this genus. Amallothrix sp. MATERIAL EXAMINED : 6 males from 5 RMT i hauls made at 1220-610 m depthin a position i8N 25W, between 12 and 18 November 1969. Deposited at theBritish Museum (Natural History), reg. no. 1974 : 604-609. DESCRIPTION. Male (Fig. 16) : Body length 2-05-2-81 mm with a mean of2-43 mm (5 specimens). The cephalothorax is 2-5 times as long as the 5-segmentedabdomen. The head and ist thoracic segment and thoracic segments 4 and 5 arefused. The rostrum has two stout filaments. The ist antenna has 19 segments andreaches to the hind edge of the 2nd abdominal segment. The remaining mouthparts 18 328 H. S. J. ROE 0-01 FIG. 1 6. Amallothrix sp. g. a, body, dorsal ; b, forehead and rostrum, lateral ; c, 2ndantenna ; d, mandibular palp ; e, ist maxilla ; f, 2nd maxilla, endopod ; g, ist leg ;h, 2nd leg ; i, 3rd leg ; j, 4th leg ; k, 5th leg ; 1, 5th leg, last exopodite segment left leg ;m, 5th leg, terminal segments (different specimen). Bar scale o-i mm unless indicated. are as shown. There is some slight variation in setation of the ist maxilla whereone specimen has nine setae on the exopodite. The swimming legs are incompletein all specimens. The 3rd exopodite segment of the ist leg has surface spines ;the ist exopodite segment of the 2nd leg has a long curved outer spine. The basi-podites of legs 2-4 are covered with surface spines. The 5th pair of legs reachesback about half the length of the abdomen. On the left side the exopodite isslightly longer than the endopodite and the last exopodite segment has a lamellaand groups of coarse setae. REMARKS. I cannot definitely identify these specimens with any of the knownmales of Amallothrix. They show a general similarity to the male of Amallothrix NEW AND RARE CALANOID COPEPODS 329 propinqua, described by Tanaka (1962), but differ in the spinulation of the swimminglegs and in the structure of the 5th pair of legs. They agree with Bradford's (1973)redefinition of Amallothrix, but as they may be the undescribed male of a knownspecies they are not considered here to be a new species. Amallothrix lobophora (Park, 1970) Scolecithricella lobophora Park, 1970 : 511, figs 188-201. MATERIAL EXAMINED : n females and 4 males taken in 8 RMT i hauls made at785-300 m depth in a position i8N 25W, between 13 and 17 November 1969.2 females and the males are deposited at the British Museum (Natural History),reg. no. 1974 : 610-615. DESCRIPTION. Male (Fig. 17): Body length 1-44-1-67 mm with a mean of1-56 mm (4 specimens). The cephalothorax is 2-8 times as long as the 5-segmentedabdomen. The head and ist thoracic segment and thoracic segments 4 and 5 arefused. The forehead is flattened anteriorly in dorsal view ; the rostrum has twolarge swollen filaments. The hind margin of the cephalothorax is bluntly triangularin lateral view. The anal segment is very short and is telescoped into the 4thabdominal segment. The ist antenna has 19 free segments and reaches to the hindend of the abdomen ; the proximal segments have large sensory filaments. Themandible blade is similar to that of the female but smaller. The 2nd maxilla hasfive exopodite lobes but two have been omitted in Fig. I7h for clarity ; the endo-podite has five brush-like setae, two of which have enlarged heads, and three worm-like setae. The other mouthparts and the legs are as shown. The external spineon the 2nd exopodite segment of the ist leg varies slightly in length ; in anotherspecimen it is longer than shown here. The 5th pair of legs is biramous and reachesback to the end of the abdomen. It is similar to that found in Scaphocalanus withthe left endopodite longer than the exopodite. The left exopodite terminates in alamella, which is broken in the illustrated specimen, and the last endopodite segmentis knobbled. In the right leg the last exopodite segment is roughly triangular inshape. REMARKS. These males were taken in the same hauls as several female A.lobophora. The present females agree with Park's (1970) description except that theist inner lobe of the ist maxilla has three surface setae and not two. Some of themale mouthparts are reduced but the characteristic rostrum and the swimming legsare identical to those of the female. Bradford (1973) transferred this species from Scolecithricella Sars, 1902 to Amallo-thrix Sars, 1925. The present specimens differ from her redefinition of Amallothrixin having a Scaphocalanus-type. ist maxilla with three surface setae on the ist innerlobe. The 5th legs of both sexes also seem to be atypical of this genus and are verysimilar to those of Amallothrix auropecten (Giesbrecht, 1892). A discussion of thepossible affinities of these two species is given after the account of A. auropectenon p. 334. 330 H. S. J. ROE FIG. 17. Amallothrix lobophora <?. a, body, dorsal ; b, rostrum, ventral ; c, rostrum,lateral ; d, last thoracic segment and abdomen, lateral ; e, 2nd antenna ; f, mandibularpalp ; g, ist maxilla ; h, 2nd maxilla, lobes 4 and 5 omitted ; i, maxilliped ; j, ist leg ;k, 2nd leg ; 1, 3rd leg ; m, 4th leg ; n, 5th leg ; o, 5th leg, last exopodite segment left leg.Bar scale o-i mm unless indicated. Amallothrix alter a (Farran, 1929) Amallophora altera Farran, 1929 : 252-4, fig. 19 ; Tanaka, 1960 : 102.Scolecithricella sp. i (part) Roe, igjia : 304, 310 ; igj2c : 540.? S. (Amallothrix) altera ; Vervoort, 1965 : 6973, figs 14-16.? S. (Amallothrix) auropecten ; Vervoort, 1951 : 101-3, n g- 54- not Amallophora altera ; Vervoort, 1957 : 94~95> fig s 77~79 ', Bradford, 1973 : 134, 136, 141,144-5, figs 2-3. NEW AND RARE CALANOID COPEPODS 331 MATERIAL EXAMINED : i female from an Nii3H haul fished at 700-550 m depthoff Fuerteventura (28N I4W) on 24 November 1963. 3 females from 2 RMT ihauls fished at 600-410 m depth in a position i8N 25W on 15 and 16 November1969. Deposited at the British Museum (Natural History), reg. no. 1974 : 616-619. DESCRIPTION. Female (Fig. 18) : Body length 1-98-2-20 mm with a mean of2-09 mm (4 specimens). The cephalothorax is 4-1 times as long as the 4-segmentedabdomen. The head and ist thoracic segment are fused ; thoracic segments 4 and 5are separate. The rostrum is large and has two filaments which are thickened FIG. 18. Amallothrix altera ?. a, body, dorsal ; b, rostrum, half lateral ; c, rostrum,ventral ; d, last thoracic segment and abdomen, lateral ; e, and antenna ; f, mandibularpalp ; g, ist maxilla ; h, and maxilla ; i, maxilliped ; j, ist leg ; k, and leg ; 1, 3rd leg ;m, 4th leg ; n, 5th leg. Bar scale o-i mm unless indicated. 33 2 H. S. J. ROE basally. The last thoracic segment protrudes laterally. The genital segment isslightly longer than the combined length of the three subsequent segments. Thefurcal rami are about i times as long as wide. The ist antenna has 24 segments(the last is very small), and reaches to the hind edge of the genital segment. Theremaining mouthparts are as shown. The endopodite of the 2nd maxilla has fivebrush-like setae, of which two are enlarged, and three flattened worm-like setae.The maxilliped has a large brush-like seta on the ist basipodite. The legs are asshown. The endopodite of the ist leg has no lobe and no surface spines. The 5thleg has three segments on each side. REMARKS. The present females have been compared with the type specimens ofAmallophora alter a held in the collections of the British Museum (Natural History).They are smaller than Farran's types but are structurally identical except that theholotype has two setae on the basipodite of the mandibular palp and no apparentsegmentation line between endopodite segments 4 and 5 of the maxilliped. Farran(1929) could not ascertain the structure of the sensory setae of the 2nd maxilla.However, both holotype and paratype are identical to the females described hereand have two large brush-like setae and three smaller ones. There is no doubt thatthe present females are conspecific with Amallophora alter a Farran. The identity of this species is confused and it has been mixed up with Amallothrixauropecten (Giesbrecht, 1892). For comparison the latter species is described below(p. 333). Bradford (1973) described as Amallophora altera a species having one verylarge and four small brush-like setae on its 2nd maxilla. This arrangement isconspicuously different from both the present material and Farran's types.Bradford's specimen is incompletely described but it also differs from Amallothrixaltera in the structure of its rostrum, its ist maxilla and in its ist and 5th legs. It isclearly a different species. Bradford used her specimen to typify a group containingAmallophora smithae Grice, 1962, Scolecithrix vorax Esterley, 1911 and Xanthocalanustypicus Farran, 1908. This group of species is considered here to comprise a newgenus which is described later on p. 335. The true Amallothrix altera does not belongin this new genus. Vervoort's (1957) description is also incomplete but his specimens differ from thepresent species by (a) a short outer edge spine on the ist exopodite segment of the2nd leg, (b) an incompletely segmented 5th leg, (c) differences in the setation ofthe ist maxilla and (d) apparent differences in the endopodite of the 2nd maxilla,although the description is not very clear on this. These specimens may be A.auropecten but their identification cannot be certain from his description. Similarlythe definite identity of his 1951 specimen cannot be determined. It has a 3-seg-mented 5th leg but Vervoort (1957) stated that it was conspecific with his latermaterial. In 1965 Vervoort described a further specimen which may be Amallothrixaltera, but the rostrum appears to be smaller and the endopod of the 2nd maxilla isdifferent, although this may be due to the difficulty in seeing the small brush-likeheads. A . altera is generally similar to A . auropecten (Fig. 19) but differs in the followingrespects, (a) the rostrum of A . altera is larger and of a different shape, (b) the setationof the ist maxilla is different, especially of the exopodite, (c) the endopodite of the NEW AND RARE CALANOID COPEPODS 333 2nd maxilla has two of the five brush-like setae enlarged in A. alter a but not inA. auropecten, (d) the ist basipodite of the maxilliped has a large brush-like seta inA. altera but not in A. auropecten, (e) the endopod of the ist leg has a comb ofsurface spines in A. auropecten but not in A. altera, and (f) the 5th leg has threesegments in A. altera but is incompletely segmented in A. auropecten. This lastdifference may be variable but all the examined specimens of A. altera have acompletely segmented 5th pair of legs whereas those of A. auropecten are normally2-segmented or incompletely divided into three (Fig. 19 and see Giesbrecht, 1892 ;Rose, 1942 ; Wilson, 1942, 1950 ; Park, 1968 ; Bradford, 1973). One of thepresent specimens of A. auropecten has an abnormal 5th pair of legs similar to thatfigured by Park (1968, pi. 9, fig. 3). In 1965 Vervoort transferred Farran's Amallophora altera to Amallothrix. Themain apparent difference between the species and Bradford's redefinition of Amallo-thrix is its Scaphocalanus-type ist maxilla with three surface setae on the ist innerlobe. This character is shared with A. lobophora (p. 329) and A. auropecten (p. 334)and discussion of the affinities of all three species follows on p. 334. The enlargementof two of the brush-like setae in the present species is not, by itself, a generic characterand it is shown by several other species and genera within the Scolecithricidae(see, for example, A. falcifer, p. 325, Scolecithricella obscura, p. 318, Scaphocalanusdifficilis, p. 315, and Scopalatum dubia, p. 338). Amallothrix auropecten (Giesbrecht, 1892) Scolecithrix auropecten Giesbrecht, 1892 : 266, pi. 13, figs 8, 18, 22, 27, pi. 37, figs 3, 10. Scolecithricella auropecten ; Rose, 1933 : 158-9, fig. 175. 5. sp. i (part) Roe, igj2a : 296, 304, 310 ; I972C : 540. ? Amallophora altera ; Vervoort, 1957 : 94-5, figs 77-79. ? not Scolecithricella (Amallothrix) auropecten ; Vervoort, 1951 : 101-3, fig- 54- MATERIAL EXAMINED : 2 females and 21 males taken in n Nii3H hauls madeat 950-475 m depth off Fuerteventura (28N I4W), between 24 and 28 November1965. 6 females and 6 males taken in 7 RMT I hauls made at 785-300 m depth ina position i8N 25W, between 13 and 16 November 1969. DESCRIPTION (Fig. 19) : Body length of the females is 1-9-2-05 mm with a meanof 1-96 mm (7 specimens) ; of the males it is 2-20-2-51 mm with a mean of 2-28 mm(21 specimens). The present females agree with the descriptions of Giesbrecht(1892) and Rose (1942). They are figured here for comparison with A. altera (p. 330)where their structure is commented upon. The endopod of the 2nd maxilla hasfive brush-like setae, all with small heads, but only four could be seen in the specimenillustrated. The males agree with Rose's account and redescription is unnecessary.The male described by Wilson (1950) as Scolecithricella auropecten is not of thisspecies. REMARKS. Rose (1942) transferred Scolecithrix auropecten to Amallothrix.Bradford (1973) omitted it from her redefined Amallothrix and considered that itshould typify an unnamed group, together with Scolecithrix laminata Farran, 1926 334 H. S. J. ROE n FIG. 19. Amallothrix auropecten ?. a, body, dorsal ; b, rostrum, lateral ; c, rostrum,ventral ; d, last thoracic segment and abdomen, lateral ; e, 2nd antenna ; f, mandibularpalp ; g, ist maxilla ; h, and maxilla ; i, maxilliped ; j, ist leg ; k, 2nd leg ; 1, 3rd leg ;m, 4th leg ; n, 5th leg. Bar scale o-i mm unless indicated. and Scolecithricella modica Tanaka, 1962. Scolecithrix (= Scolecithricella} laminatadiffers in many respects from the present species (see p. 320) and is not, in myopinion, closely related. A. altera and A. lobophora, however, do seem to be relatedto the present species. All three differ from Bradford's Amallothrix in having aScaphocalanus-type ist maxilla (p. 329). The female 5th legs of all three speciesand the male 5th legs of A . lobophora and A . auropecten are also atypical of this genus.There are, however, differences between these three species. A. altera and A.auropecten are distinct (p. 332), and A. lobophora differs from both by lacking an NEW AND RARE CALANOID COPEPODS 335 outer edge spine on the ist exopodite segment of the ist leg and by having spineson the ist basipodite of the 3rd leg. The rostrum of A . lobophora is similar to that ofA . alter a but the structure of its 5th legs (both sexes) , the spiny lobe on the endopodof the ist leg, and the uniformly small brush-like setae on the female 2nd maxilla areall more like A. auropecten. Whether or not these three species are sufficiently alikeand sufficiently distinct from others to stand as a separate genus within the Scole-cithricidae is not yet clear. Discovery of the male of A . alter a and determination ofthe acceptable generic limits, especially regarding the ist maxilla, will clarify theirposition. Meanwhile all three species can conveniently remain within Amallothrix. SCOPALATUM gen. nov. DIAGNOSIS. Head and ist thoracic segment fused ; thoracic segments 4 and 5fused or separate. Rostrum with two filaments, ist antenna with 23 segments inthe female, 20 in the male ; there are large sensory filaments on the proximalsegments of the male ist antenna. 2nd antenna exopod slightly longer than theendopod. ist maxilla with two posterior surface setae on the ist inner lobe, twosetae on the 2nd inner lobe and four on the 3rd inner lobe ; in the female the endopodis ornamented with spinules. 2nd maxilla endopod with five brush-like setae andthree flattened worm-like setae ; in the female one of the brush-like setae is greatlyenlarged ; in the male two are enlarged. MaxiUiped with one brush-like seta on theist basipodite. The male mouthparts are weakly chitinized and slightly reduced. ist leg has a 3-segmented exopod and a single endopodite segment : all threeexopodite segments have an outer spine ; the endopod has a lobe carrying smallspines. 2nd leg has a 3-segmented exopod and a 2-segmented endopod ; the outeredge spines on all exopodite segments are large ; the 2nd and 3rd exopodite segmentsand the 2nd endopodite segment carry prominent surface spines. 3rd leg has a3-segmented exopodite and endopodite : the ist basipodite segment has a group oflong spines ; the 2nd and 3rd exopodite and endopodite segments have surfacespines. 4th leg has a 3-segmented exopodite and endopodite : the ist basipoditemay have a group of long spines ; the 2nd and 3rd endopodite segments have veryfew small surface spines ; the exopodite segments have no surface spines. Female5th leg is uniramous ; on each side there are two segments, which may be fused,attached to a common basal segment ; there are 1-3 spines. Male $th leg similarto that of Scaphocalanus : left exopod is shorter than the endopod ; right exopod ismuch longer than the endopod. Scopalatum gibbera sp. nov. is nominated as the type species of the genus. The name Scopalatum is derived from the Latin scopae - a broom and latum -carried ; it refers to the sensory setae of the 2nd maxilla. DISCUSSION. T. Scott (1894) described a new subgenus, Amallophora, for a malecopepod, A. typica, having a peculiar enlarged sensory seta on its 2nd maxilla.Giesbrecht (1892) had previously described a similar structure in the male ofXanthocalanus and considered that Amallophora was a junior synonym of Xantho-calanus (Giesbrecht & Schmeil, 1898). Farran (1908) named a female which hebelieved to be conspecific with Scott's male as X. typicus. Recently, Bradford 336 H. S. J. ROE (1973) separated these two specimens, reiterating Giesbrecht's opinion that Scott'smale is a xanthocalanid but transferring Farran's female to the Scolecithricidae.She placed this, as X. typicus Farran, into a group together with Amallophorasmithae Grice, 1962, Scolecithrix vorax Esterley, 1911 and Amallophora altera Farran,1929. Her inclusion of A. altera in this group however is erroneous, see p. 332 ;Bradford's specimen of A. altera is not conspecific with Farran's, which belongs toan entirely different group. Nevertheless, I believe she was correct in collectingtogether her specimen of 'A . altera' and the other three species into a distinct groupwithin the Scolecithricidae. Bradford briefly characterized this group but did not name it as a separate genus.The discovery of a female of a new species which clearly belongs in this group (seebelow) and the redescription of what, in my opinion, is a male of this group (p. 338)lead me to believe that this collection of species should be given generic statuswithin the Scolecithricidae. This new genus is denned above (p. 335). The genusScopalatum at present contains 5. farrani sp. nov., 5. vorax (Esterley, 1911), 5.smithae (Grice, 1962), 5. dubia (T. Scott, 1894) and 5. gibber a sp. nov. It alsoincludes the species 'Amallophora altera' wrongly identified by Bradford (1973)which should be further described and named. Grice's (1962) account of S. smithaeis complete and also conforms with the new genus. The descriptions of 5. vorax(Esterley, 1911) and 'A. altera' of Bradford (1973) are incomplete, however, par-ticularly regarding the swimming legs. The present generic definition may have tobe modified when these latter two species are fully described. Details of S. gibberasp. nov., 5. farrani sp. nov. and 5. dubia (T. Scott, 1894) are given below. Scopalatum agrees in all respects with Bradford's (1973) redefinition of theScolecithricidae. The combination of characters given on p. 335, especially thoseof the ist and 2nd maxillae and the legs, distinguishes this new genus from allothers in the family. It is related to A . altera, A . auropecten and A . lobophora, seep. 334. These three species may have to be removed from Amallothrix but theydiffer from the present genus in the structure of their ist maxilla and, in the females,of the 2nd maxilla. The 2nd maxilla of the male A. auropecten is also distinct, andboth this species and A. altera are further distinguished by having no basipoditespines on either their 3rd or 4th legs. A. lobophora has a similar 2nd maxilla in themale to Scopalatum and both sexes have a group of spines on the ist basipodite ofthe 3rd leg. They do not, however, have an external spine on all exopodite segmentsof the ist leg. Scopalatum gibbera sp. nov. Amallophora sp. Roe, 19720 : 295, 309. MATERIAL EXAMINED : i female in an Nii3H haul fished at 310-240 m depth offFuerteventura (28N I4W) on 13 November 1965. 2 females in 2 RMT i haulsfished in a position i8N 25W ; one (the holotype) caught at 300-210 m depth on12 November 1969 and the other at 400-305 m depth on 17 November 1969.Deposited at the British Museum (Natural History), holotype reg. no. 1974 : 620,paratypes reg. no. 1974 : 621-622. NEW AND RARE CALANOID COPEPODS 337 m FIG. 20. Scopalatum gibbera ?. a, body, dorsal ; b, body, lateral ; c, rostrum, ventral ;d, ist antenna ; e, 2nd antenna ; f, mandible ; g, ist maxilla ; h, 2nd maxilla, exopod ;i, 2nd maxilla, endopod ; j, maxilliped ; k, ist leg ; 1, 2nd leg ; m, 2nd leg, exopoditesegments 2-3 (paratype) ; n, 3rd leg ; o, 4th leg ; p, 5th leg. Bar scale o-i mm unlessindicated. 33 8 H. S. J. ROE DESCRIPTION. Female (Fig. 20) : Body length 2-58-2-81 mm with a mean of2-73 mm (3 specimens). The holotype measures 2-81 mm. The cephalothorax isrobust and 3-9 times as long as the 5-segmented abdomen. The head and istthoracic segment are fused, as are thoracic segments 4 and 5. The mid-dorsalregion of the head has a very characteristic protuberance which is particularlyconspicuous when seen from the side. The rostrum has two filaments with thickenedbases. The hind margin of the cephalothorax is deeply indented laterally. Thegenital segment has a slight ventral protrusion and is longer than the combinedlength of the three following segments. The furcal rami are about as long as wide.The ist antenna has 23 segments and does not quite reach the end of the cephalo-thorax. The remaining mouthparts and the legs are as shown. The endopod of theist maxilla is covered with small spines. The endopodite of the 2nd maxilla hasfive brush-like and three flattened worm-like setae, one of the brush-like setae isgreatly enlarged. The ist basipodite of the maxilliped has one brush-like seta.The ist basipodite of the 3rd leg has two conspicuous groups of long spines. Theterminal spines on legs 2-4 have laminae. The 5th pair of legs is uniramous withtwo fused segments on each side attached to a basal segment ; the line of fusion isclearly visible. The terminal segment on each side has one long and one shorterspine. There is an additional small spine on one side only, but this is clearly variablesince in one of the paratypes it is about halfway down the outside edge of the fused2nd and 3rd segments. REMARKS. 5. gibbera has all the female characters defined for the genus on p.335. A discussion of its affinities has been given earlier. Scopalatum farrani sp. nov. Xanthocalanus typicus ; Farran, 1908 : 47-48, pi. 4, figs 15-17 ; Bradford, 1973 : 145.Amallophora typica ; Sars, 1924 : pi. 38, figs 8-18 ; 1925 : 140-142 (part) ; Rose, 1933 : 134- 135 (part), fig. 129 ; Wilson, 1942 : 170-171, fig. i ; Wilson, 1950 : 159 (part), pi. 20, fig. 275 ; Vervoort, 1957 : 94-95 (part) ; Tanaka, 1960 : 102 (part) ; Vervoort, 1965 : 27-28 (part) ; Binet & Dessier, 1971 : 430, 453 ; Bradford, 1973 : 138-139 (part).? Amallophora typica ; Grice & Hulsemann 1965 -.221, 223 ; Harding, 1972 : 58.not Xanthocalanus typicus ; Giesbrecht, 1897 : 254 ; Giesbrecht & Schmeil, 1898 : 50 ; Wolfen- den, 1908 : 35.not Amallophora typica T. Scott, 1894 : 54, pi. 3, figs 39-46, pi. 6, figs 1-4 ; A. Scott, 1909 : 85, pi. 36, figs 1-8. The female described by Farran (1908) as X. typicus was misidentified and belongsto a new species which I have named Scopalatum farrani. The female specimens ofS. farrani in the collections of the Institute of Oceanographic Sciences agree com-pletely with the generic definition of Scopalatum given earlier. Scopalatum dubia (T. Scott, 1894) Amallophora dubia T. Scott, 1894 : 55, pi. 4, figs 10-18. Scolecithrix scotti Giesbrecht, 1897 : 254 ; Giesbrecht & Schmeil, 1898 : 46 ; Wolfenden, 1911 :250. NEW AND RARE CALANOID COPEPODS 339 Heteramalla dubia ; Sars, 1907 : 17 (part) ; A. Scott, 1909 : 86-87 (part) ; Sars, 1925 : 142-144(part) ; Rose, 1933 : 135 (part) ; Alzamora, 1940 : 6 ; Wilson, 1942 : 189 (part) ; Vervoort,1965 : 28-30 (part). Heleremalla dubia ; Rose, 1929 : 26. Hetermalla dubia ; Roe, 19720 : 303, 309. ? Heteramalla dubia ; Wilson, 1950 : 239 ; Grice & Hulsemann, 1967 : 16. not Heteramalla dubia ; A. Scott, 1909 : pi. 33, figs 1-9 ; Sars, 1924 : pi. 39 ; Rose, 1933 : fig.130 ; Grice & Hulsemann, 1965 : 221, 223, 235. MATERIAL EXAMINED : i male in an Nii3H haul made at 660-510 m depth offFuerteventura (28N I4W) on 26 November 1965. 8 males taken in 5 RMT ihauls made at 900-305 m depth in a position i8N 25W between 12 and 17 Novem-ber 1969. 7 males deposited at the British Museum (Natural History), reg. no.1974 : 623-629. DESCRIPTION. Male (Fig. 21) : Body length 2-36-2-74 mm with a mean of2-48 mm (9 specimens). The cephalothorax is 2-7 times as long as the 5-segmentedabdomen. The head and ist thoracic segment are fused ; thoracic segments 4 and5 are separate. The rostrum has two filaments attached to a common base. Thehind margin of the cephalothorax has a slight lateral indentation. The ist antennareaches to the hind edge of the 2nd abdominal segment ; both ist antennae have20 segments and the proximal 12 segments on each side carry large flattened sensoryfilaments. The mouthparts and legs are as shown. The endopod of the 2ndmaxilla has five brush-like setae and three flattened worm-like setae ; the heads ofthree of the brush-like setae are small whereas two are greatly enlarged - one of themespecially so. All the 'brush' heads consist of fine threads ending in roundedswellings. The ist basipodite of the 3rd leg has two groups of long narrow spines.The 5th pair of legs is similar to that of Scaphocalanus and reaches back to the endof the furcal rami. On the right leg the exopodite terminates in a claw-like segment.On the left leg the exopodite is shorter than the endopodite ; the 3rd exopoditesegment is covered with hairs and has a terminal lamella. REMARKS. I have compared the present males with the type specimens of Amallo-phora dubia T. Scott, 1894 held in the collections of the British Museum (NaturalHistory). The type material consists of several slides on which are mounted themixed appendages and abdomens of two specimens. There is only one example ofmost appendages - including the 2nd maxilla, and there is no specimen of the istmaxilla. Neither of the two cephalosomes is in the Museum's collection. Thematerial is very faded and it is impossible to ascertain the detailed structure of someof the appendages. An attempt to remount some of the specimens failed becausethe mountant would not dissolve. The only observable difference between thepresent males and Scott's type is in the 3rd leg where the type does not, apparently,have spines on the ist basipodite. This limb is figured by Scott but wronglydescribed as the 4th leg (Scott, 1894 : PL 4, fig. 16). According to Scott the rightist antenna has 18 joints and is indistinctly geniculate and the left ist antenna has23 joints. There are three ist antennae in the type collection ; one has 19 segmentsbut the others are very faded and have, I believe, 20 and 21 segments respectively.None of these antennae are in any way geniculate and their segmentation agrees 340 H. S. J. ROE u FIG. 21. Scopalatum dubia $. a, body, dorsal ; b, rostrum and forehead, lateral ; c,rostrum, ventral ; d, last thoracic segment, lateral ; e, ist antenna ; f, and antenna ;g, mandibular palp ; h, mandible blade ; i, ist maxilla ; j, ist maxilla (differentspecimen) ; k, 2nd maxilla ; 1, 2nd maxilla, head of small brush-like seta ; m, maxilliped,basal segment (different specimen) ; n, ist leg ; o, 2nd leg ; p, 3rd leg ; q, 4th leg ; r,5th leg ; s, 5th leg, last exopodite segment right leg ; t, 5th leg, last exopodite segmentleft leg; u, 5th leg, last endopodite segment left leg (different specimen). Bar scaleo-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 341 better with the present male's antennae, which have 20 segments on each side, thanwith Scott's description. The type material of the 2nd maxilla is in very poorcondition and it is impossible to see any structure except the general presence of twolarge brush-like setae. Despite the possible difference in the 3rd leg I believe thatthe present males are conspecific with Scott's Amallophora dubia. Sars (1907) erected a genus, Heteramalla, for a female which he believed wasconspecific with Scott's male. In 1925, however, he expressed doubts as to thecorrectness of his assumption, and in 1965 Vervoort also mentioned the possibilitythat Sars' females and Scott's male were not the same species. Further malespecimens have been recorded by Wolfenden (1911) and by Rose (1929) but theywere not described. Bradford (1973) transferred Heteramalla from the Phaennidae to the Scolecithridae.The present males are clearly scolecithricids according to her redefinition of thefamily. I have examined female specimens of H. dubia Sars, 1907 taken at 1020-800 m depth in a position i8N 25W. These specimens agree with Sars' (1924,1925) description except that they have no 5th pair of legs (thereby conformingwith Grice & Hulsemann's (1965) specimen), and there is no seta on the 2nd innerlobe of the ist maxilla. The endopod of the 3rd leg has two segments only. Thesefemales also agree with Bradford's definition of the Scolecithricidae. I believe,however, that Sars was mistaken in identifying his female with Scott's male. Apartfrom the normal sexual differences they differ in the following respects : (a) therostrum of $ H. dubia is strong and plate-like, in the male it has two filaments,(b) the ist maxilla of the males has two setae on the surface of the ist inner lobe,two on the 2nd inner lobe and four on the 3rd inner lobe ; the corresponding numberof setae in the female are i, o and 3, (c) the ist leg of $ H. dubia has a 2-segmentedexopod with no external spine on the ist exopodite segment ; the endopod has nolobe and has a group of large spines ; in the male the ist leg has three exopoditesegments, all with outer spines, and the endopod has a lobe with a comb of smallspines only, (d) $ H. dubia has no surface spines on the exopodites of legs 2-4-these legs are heavily spinulated in the male, and (e) the 3rd leg of the $ H. dubiahas a 2-segmented endopod and has no spines on the ist basipodite segment ; the3rd leg of the male has three endopodite segments and two groups of spines on theist basipodite. I agree with Bradford (1973) that the genus Heteramalla Sars, 1907 belongs tothe Scolecithricidae. It is at present a monospecific genus known only from thefemale (see p. 342). Scott's male is here considered to belong to the new genusScopalatum described on p. 335. The most striking difference between S. dubia and the females of Scopalatum is inthe development of the sensory setae of the 2nd maxilla. The females have onebrush-like seta much enlarged but the male has two. Enlargement of one or two ofthese setae is, however, common to the males of many species and genera of theScolecithricidae (see, for example, Amallothrix falcifer, p. 325, and Scolecithricelladentata (Rose, 1942 : 55). I believe that the increased size of one seta in thepresent males is a sexual character. Apart from the normal sexual differences inthe ist antenna and more weakly chitinized and slightly reduced mouthparts, the 342 H. S. J. ROE appendages of these males are very similar to those of the female 5. gibber a sp. nov.,5. farrani sp. nov. and 5. smithae Grice, 1962. They are so similar to 5. gibberathat these two species may be conspecific. They differ in the length of the outeredge spines of the exopodite of the ist leg, and the males do not have the curiousdorsal swelling of the female. In addition to these differences 5. farrani has groupsof spines on the ist basipodite of the 4th leg. 5. smithae and S. vorax are muchsmaller species. Heteramalla sarsi sp. nov. Heteramalla dubia ; Sars, 1907 : 17 (part) ; A. Scott, 1909 : 86-87 (P ar t)> pi. 33, figs 1-9 ;Sars, 1924 : pi. 39 ; 1925 : 142-144 (part) ; Rose, 1933 : 135 (part) fig. 130 ; Wilson, 1942 :189 (part) ; Grice & Hulsemann, 1965 : 221, 223, 235 ; Vervoort, 1965 : 28-30 (part). ? Heteramalla dubia ; Wilson, 1950 : 239 ; Grice & Hulsemann, 1967 : 16. not Heteramalla dubia : Alzamora, 1940 : 6. The female described by Sars (1907) as H. dubia was misidentified and belongs to anew species which I have named H. sarsi. The validity of this species is consideredearlier together with the discussion of Scopalatum dubia (T. Scott). Family METRIDIIDAEMetridia alata sp. nov. MATERIAL EXAMINED : 6 females and 6 males taken in 6 RMT i hauls made at1220-700 m depth in a position i8N 25W, between 12 and 18 November 1969.The holotype (female) and allotype (male) were caught on 18 November 1969 ati22O-iooom. Deposited at the British Museum (Natural History), holotypereg. no. 1974 : 630, paratypes reg. no. 1974 : 631-641. DESCRIPTION. Female (Fig. 22) : Body length 8-64-10-24 mm with a mean of9-09 mm (5 specimens). The holotype measures 9-12 mm. The cephalothorax is1-2 times as long as the 3-segmented abdomen. It is very robust and in lateral viewthe anterior part of the head protrudes sharply from the hind part. The right lateralmargin of the head is produced into a very conspicuous wing ; this wing protrudesboth laterally and ventrally and is present on the right side only. The head and istthoracic segment are separate ; thoracic segments 4 and 5 are fused. The rostrumhas two strong spikes pointing ventrally. The hind edge of the cephalothorax issmooth and rather abruptly rounded. The genital segment is swollen both laterallyand ventrally and is longer than the combined length of the two subsequent segments.The furcal rami are 0-36 times the length of the abdomen and are 6 times longer thanwide. The ist antenna is slightly longer than the body : it has 25 segments, seg-ments 7 and 8 are partially fused and the last segment is small ; the ist segment hasthree hooks and the 2nd, 4th and 5th segments each have a single hook. Theremaining mouthparts and legs are as shown. The endopodite of the mandibularpalp has a group of large spines on the ist segment and many small spines on the 2nd.The mandible blade has numerous small spines in addition to the large teeth. The NEW AND RARE CALANOID COPEPODS 343 FIG. 22. Metridia alata ?. a, body, dorsal ; b, body, lateral ; c, ist antenna, basalsegments ; d, 2nd antenna ; e, mandibular palp ; f, mandible blade ; g, ist maxilla ;h, 2nd maxilla ; i, maxilliped ; j, ist leg ; k, 2nd leg ; 1, 3rd leg ; m, 4th leg ; n, 5th leg.Bar scale o-i mm unless indicated. 19 344 H - s - J- ROE ist endopodite segment of the 2nd leg has three strong claws. The 5th pair of legsis symmetrical ; on each side the basipodite has a bunch of long hairs, the ist seg-ment has a plumose outer edge seta, the 2nd a short spine and the 3rd terminateswith three plumose setae. Male (Fig. 23a-b) : Body length 8-00-8-64 mm wrt h a mean of 8-51 mm (6specimens) . The allotype measures 8-64 mm. The appearance of the cephalothoraxis as in the female ; the head has a conspicuous wing on the right side only. Theabdomen has five segments. The left ist antenna is geniculate ; both ist antennaeare slightly longer than the body. The remaining mouthparts and swimming legsare as in the female. The 5th pair of legs is asymmetrical : on the left side, the basalsegment is longer than the right and has a bundle of hairs, the 4th segment ter-minates with two small spines ; on the right side there are five segments, the 2ndbearing a long curved process and the 5th ending in three small spines. REMARKS. The appendages of the present species are very similar to those ofMetridia princeps Giesbrecht, 1892, but it can be distinguished easily from this andfrom all other species in the genus by the peculiar wing-like development of the rightside of the head. Family CENTROPAGIIDAECentropages caribbeanensis Park, 1970 MATERIAL EXAMINED : 43 females and 21 males from 6 RMT i hauls made at194-0 m depth in a position i8N 25W, between 15 and 18 November 1969. 5males are deposited at the British Museum (Natural History), reg. no. 1974 :642-646. DESCRIPTION. Male (Fig. 23c-e) : Body length 1-82-2-13 mm with a mean of1-95 mm (15 specimens). The general appearance of the cephalothorax is as for thefemale ; it is 2-2 times as long as the 5-segmented abdomen. The furcal rami areabout 3 times longer than wide. The ist antenna exceeds the body length by threeor four segments. The right ist antenna is geniculate and has 22 segments, segments17, 18 and the proximal part of 19 have a row of fine teeth, the hinged joint isbetween segments 18 and 19. The remaining mouthparts and swimming legs areas in the female. The 5th pair of legs is asymmetrical. On the right side theterminal exopodite segment forms a large chela ; on the left side the last exopoditesegment has two large spine-like processes. REMARKS. The present males were found in the same hauls as females whichagree in every respect with Park's (1970) description. The male is similar to thatof C. elegans Giesbrecht, 1895 described by Park (1968). It can be distinguishedfrom this latter species by the 5th pair of legs. In C. caribbeanensis the spines onthe terminal segment of the left exopodite are longer than in C. elegans and are notpitted distally. The shorter arm of the chelate right exopodite is pointed in C.caribbeanensis and rounded in C. elegans. The present specimens are only the secondrecord of this species and the first from the eastern north Atlantic. NEW AND RARE CALANOID COPEPODS 345 FIG. 23. a-b, Metridia alata <$. a, body, dorsal ; b, 5th leg. c-e, Centropages carib-beanensis 3- c, body, dorsal ; d, last thoracic segment and abdomen, lateral ; e, 5thlegs, f, Heterorhabdus vipera <$ 5th legs, g, Heterorhabdus tenuis <$ 5th legs, h-k,Haloptilus angusticeps <$. h, body, dorsal ; i, forehead and rostrum, lateral ; j, lastthoracic segment and abdomen, lateral ; k, 5th legs. Bar scale o-i mm unless indicated. 346 H. S. J. ROE Family HETERORHABDIDAEHeterorhabdus vipera (Giesbrecht, 1892) MATERIAL EXAMINED : 14 females and 4 males from 12 Nii3H hauls fished at590-90 m depth off Fuerteventura (28N i4W), between n and 26 November1965. 22 females and 3 males from 12 RMT i hauls made at 1250-110 m depth ina position i8N 25W between 12 and 17 November 1969. REMARKS. The males of H. vipera and H. tenuis Tanaka, 1964 have been confused.Park (1970) described the male of H. tenuis as H. vipera (Roe, 1972^), and since thepresent material includes both species it seems advisable to illustrate the differencesin their fifth pair of legs. H. vipera (Fig. 23f) has a strongly asymmetric fifth pair of legs. The endopoditesconsist of three segments on both sides and have been omitted for clarity. The 3rdexopodite segment of the right leg is abruptly terminated and about as long as thecombined length of the two previous segments. The present specimens differslightly from that of Giesbrecht (1892) by having a spine on the 2nd basipoditesegment of both legs, and by having an outer edge spine on the 2nd exopodite of theright leg. The similarity is such that I have no doubt that they are conspecific. Heterorhabdus tenuis Tanaka, 1964 Heterorhabdus vipera ; Park, 1970 : 523-525, figs 250-252 (male only). MATERIAL EXAMINED : 7 males from 5 Nii3H hauls made at 660-460 m depth offFuerteventura (28N 14 W), between u and 26 November 1965. 14 males from6 RMT i hauls made at 600-210 m depth in a position i8N 25W, between 12 and16 November 1969. REMARKS. The present males agree with the descriptions of Tanaka (1964) andPark (1970 - as H. vipera <$}. The differences in the fifth pair of legs between H.tenuis and H. vipera can be seen clearly by comparing Figs 23g and 23f. In H.tenuis the 2nd basipodite of the left leg has a haired inner margin and the 3rdexopodite segment of the right leg is a totally different shape to that of H. vipera.Both endopodites have three segments and are omitted for clarity. Family AUGAPTILIDAE Several of the genera in this family, particularly Euaugaptilus, have recentlybeen reviewed by Matthews (1972). Because of the high degree of intraspecificvariation within this genus he used a system of numerical taxonomy, based upon35 structural characters, to identify the various species. Tanaka & Omori (1974)also used a similar list of 21 characters to identify the Euaugaptilus spp. off Izu,Japan. These latter authors also recognized the great variation within species inthis genus but retained a normal diagnostic key for their identification. Several species of Euaugaptilus are described here, and all of them show markedintraspecific variations. This variability emphasizes the difficulties in recognizingor describing species based upon single specimens. Some Euaugaptilus species are NEW AND RARE CALANOID COPEPODS 347 so similar to one another (see, for example, Matthews, 1972 : 66), that the discoveryand description of more specimens is needed before firm conclusions can be reachedregarding their validity. Haloptilus angusticeps Sars, 1907 MATERIAL EXAMINED : 2 males and 7 females from 5 Nii3H hauls made at625-0 m depth off Fuerteventura (28N I4W), between 17 and 26 November 1965.12 females from 5 RMT I hauls made at 700-305 m depth in a position i8N 25W,between 14 and 17 November 1969. The males are deposited at the British Museum(Natural History), reg. no. 1974 : 647-648. DESCRIPTION. Male (Fig. 23h-k) : Body length 3-04-3-12 mm. The cephalo-thorax is long and slender and slightly more than 4 times as long as the 5-segmentedabdomen. The head and ist thoracic segment are separate ; thoracic segments 4and 5 are fused. The rostral base is rounded and bears two fine fairly long filaments.The hind margin of the cephalothorax is smoothly rounded. The ist antennaereach to the end of the furcal rami ; the left ist antenna is geniculate and has 20segments, the hinge joint is in segment 18. The remaining mouthparts and swim-ming legs are similar to those of the female. The 5th pair of legs is slightly asym-metrical. On each side, the ist basipodite segment has a pointed projection whichis larger on the left, and the 2nd basipodite has a patch of hairs. Both endopodshave three segments and are omitted here for clarity. The 2nd and 3rd exopoditesegments are almost fused and only a faint line of separation remains. On the leftside the 3rd exopodite segment has three small spines (the outer one is missing on theillustrated specimen) ; on the right the 3rd exopodite has one long curved spine andtwo of medium length. REMARKS. Except for sexual differences these males agree so closely with thefemale H. angusticeps that there is no doubt that they are conspecific. Haloptilus paralongicirrus Park, 1970 MATERIAL EXAMINED : 8 males and 160 females from 7 RMT i hauls made at4~55 m depth in a position i8N 25W, between 12 and 16 November 1969. 5males are deposited at the British Museum (Natural History), reg. no. 1974 : 649-653. DESCRIPTION. Male (Fig. 24) : Body length 1-67-2-20 mm with a mean ofi -87 mm (6 specimens). The shape of the cephalothorax is similar to that of thefemale. The head and ist thoracic segment are fused, as are thoracic segments 4and 5. In dorsal view the forehead is shaped like a rounded triangle, and in somespecimens it is more rounded than shown here. The rostral filaments are fairlylong and have swollen bases. The hind margin of the cephalothorax is smoothlyrounded. The abdomen has five segments, the furcal rami are about twice as longas wide. The ist antennae are longer than the body. The left ist antenna isgeniculate and has 21 segments ; the proximal 12-13 segments have long sensorysetae, segments 16, 17 and 18 have an undulating lamella, and segment 18 contains H. S. J. ROE FIG. 24. Haloptilus paralongicirrus <J. a, thorax and abdomen, dorsal ; b, head, dorsal(different specimen) ; c, rostrum, lateral ; d, rostrum, ventral ; e, ist antenna, segments15-20 ; f, 2nd antenna ; g, mandibular palp ; h, mandible blade ; i, 2nd maxilla ;j, maxilliped ; k, ist leg ; 1, 2nd leg ; m, 5th legs (different specimen). Bar scale o-i mmunless indicated. the hinge joint. (The last segment is missing in the figure.) The remaining mouth-parts are very feebly chitinized and are similar in structure to those of the female.A satisfactory preparation of the ist maxilla could not be made. The 2nd, 3rd and4th pairs of legs are similar. The 5th pair of legs is slightly asymmetrical ; thepointed projection on the ist basipodite is larger on the left side, and the 2ndexopodite segment of the right leg bears a small haired lobe. Both endopoditeshave three segments and are omitted here for clarity. REMARKS. Haloptilus paralongicirrus was described from the female (Park, 1970)and is closely related to Haloptilus longicornis (Claus, 1863) and H. longicirrusBrodsky, 1950. The females of the present species differ from those of H . longicirrusonly in the shape of the forehead and their smaller size. Park suggested that thesevery slight differences may only be variations caused by differences in their verticaldistributions - H. longicirrus living deeper than H . paralongicirrus. The presentcollections contain females of all three species and the males of two (that of H .longicirrus is unknown). H. longicornis was the shallowest of the three and H.longicirrus the deepest. The mean body lengths of H. longicornis were 2-14 mm(females) and 1-18 mm (males), of H. paralongicirrus they were 2-59 mm (females)and 1-87 mm (males), and of H. longicirrus 3-08 mm (females). All three species NEW AND RARE CALANOID COPEPODS 349 were non-migrants. These results agree with those of Park (1970). The presentmales were found in association with female H. paralongicirrus and since their struc-ture, body size and depth distribution accord with those of the female I believethat they are conspecific. Euaugaptilus atlanticus sp. nov. MATERIAL EXAMINED : 7 females and 2 males from 2 RMT i hauls made in aposition i8N 25W ; one, containing the female holotype, at 400-305 m depth on17 November 1969 and the other, with the male allotype, at 400-300 m depth on13 November 1969. Deposited at the British Museum (Natural History), holotypereg. no. 1974 : 654, paratypes reg. no. 1974 : 655-662. DESCRIPTION. Female (Fig. 25a-g) : Body length 2-81-2-96 mm with a mean of2-92 mm (7 specimens). The holotype measures 2-89 mm. The cephalothorax islong and slender and about 3-5 times as long as the 3-segmented abdomen. Thehead and ist thoracic segment are separate but thoracic segments 4 and 5 are fused.In dorsal view the head protrudes slightly and is rounded. The rostrum has twofine short filaments. The hind margin of the cephalothorax is smoothly rounded.The genital segment is elongated and over twice as long as the combined length ofthe remaining two segments ; it is slightly swollen laterally and more distinctlyso ventrally. The furcal rami are about twice as long as wide ; the inner furcalsetae are almost as long as the cephalothorax. The first antenna has 25 segments, the last five of which extend past the end ofthe body. The remaining mouthparts are as shown. The mandible blade has thesame structure on both the right and left side. The swimming legs are identicalin both sexes. The base of each external spine on the exopodite of the ist leg hasa conspicuous comb of curved spines. The terminal exopodite spine on the 2nd,3rd and 4th leg is long and bears numerous small spinules. The 5th pair of legs issymmetrical. The 2nd basipodite has a very long seta which is longer than theremainder of the leg. The 2nd exopodite segment has a massive internal spine,carrying many coarse teeth, but no external spine. The endopodite has threesegments. Male (Fig. 25h-m) : The allotype is slightly crushed and could not be measured ;the other male is 2-58 mm long. The general appearance is as for the female. Theabdomen has five segments. The left ist antenna is geniculate but broken in bothspecimens, the right ist antenna reaches to the end of the furcal rami. The mouth-parts are identical to those of the female except that the exopodite of the 2ndantenna has three terminal setae not two, and the mandible blade on the right sidehas a minute extra tooth. The swimming legs are as shown, the 4th leg is similarto the 3rd. The 5th pair of legs is slightly asymmetrical. The ist exopodite seg-ment of the left leg has no outer spine and the 3rd exopodite segment of this leg hasa short claw and a long curved spine. On the right leg the second exopodite segmenthas a haired lobe on its inner edge, and the terminal segment has three spines. Onboth legs the inner corner of the 2nd basipodite bears a curious structure shaped likea segmented hemisphere. The endopodites of both legs have three segments. 350 H. S. J. ROE m FIG. 25. a-g, Euaugaptilus atlanticus ?. a, body, dorsal ; b, forehead and rostrum,lateral ; c, 2nd antenna ; d, mandible right side ; e, 2nd maxilla ; f, maxilliped ; g, 5thleg. h-m, Euaugaptilus atlanticus $. h, ist maxilla ; i, ist leg ; j, ist leg, outer edgeof exopodite segments i and 2 ; k, 2nd leg ; 1, 3rd leg ; m, 5th legs. Bar scale o-i mmunless indicated. NEW AND RARE CALANOID COPEPODS 351 REMARKS. The female of E. atlanticus resembles that of E. longiseta Grice &Hulsemann, 1965. I have examined the type specimen of this latter species at theBritish Museum (Natural History) and it differs from E. atlanticus in the structureof the mandible blade, the ist leg and the 2nd exopodite segment of the 5th leg.The male of E. longiseta is smaller than the present species and has a different 5thpair of legs. The copepodite E. sp. i Tanaka & Omori, 1974, is also similar to thepresent species but it is larger and differs in the structure of the ist maxilla andmandible blade. Euaugaptilus fecundus Tanaka & Omori, 1974 Euaugaptilus sp. 3 Grice & Hulsemann, 1967 : 37, figs 231-240 ; Matthews, 1972 : 45. MATERIAL EXAMINED: i female from an RMT i haul made at 1250-1000 mdepth in a position i8N 25W on 17 November 1969. Deposited at the BritishMuseum (Natural History), reg. no. 1974 : 663. DESCRIPTION. Female (Fig. 26a-h) : Body length 6-40 mm. I was preparingto describe and name this specimen until I saw the recent account of E. fecundus.The present female differs from Tanaka & Omori's specimen only in the followingminor details, (a) the endopod of the 2nd antenna is about 4 times longer than theexopod, instead of 2-6 times, (b) there are 24 setae on the endopod of the 2ndmaxilla instead of 19, and (c) the 3rd endopodite segment of the 2nd leg has sevensetae instead of six. The setation of the distal segments of the maxilliped isslightly obscure in the present female but in all other respects it is identical toE. fecundus. REMARKS. I believe that this species is the female of the male described asEuaugaptilus sp. 3 by Grice & Hulsemann (1967). Except for the ist antenna and2nd maxilla the mouthparts of the present female are identical to those of the male.The ist lobe of the 2nd maxilla bears a single seta in both the present female andthat of Tanaka & Omori ; this is not shown by Grice & Hulsemann but it is easilydamaged when the appendage is dissected off the body. The endopod of the 2ndmaxilla in the present specimen has more setae than either the male or Tanaka &Omori's female. The proportions of the rami in the 2nd antenna are similar in thepresent female and the male. The endopodite of the ist leg has three segments inthe male but only two in both females ; the remaining swimming legs of the maleare not described in detail by Grice & Hulsemann. Except for normal sexual differences the only real difference between the femalesand the male is in the segmentation of the ist leg. Intraspecific variation occursin Euaugaptilus, however, both in the setation of the mouthparts and in the seg-mentation of the ist leg (Matthews, 1972). Grice & Hulsemann (1967) and Matthews(1972) were unable to refer the male to a known species but pointed out similaritieswith E. rigidus (Sars, 1907). The true male of E. rigidus is described below (p.357), and the agreement between the present female and the male E. sp. 3 is so goodthat I have no doubt that they are conspecific. 352 H. S. J. ROE FIG. 26. a-h, Euaugaptilus fecundus ?. a, body, dorsal ; b, forehead and rostrum,lateral ; c, mandible ; d, ist maxilla ; e, maxilliped ; f, ist leg ; g, 2nd leg ; h, 5th leg.i-q, Euaugaptilus latifrons ?. i, body, dorsal ; j, forehead and rostrum, lateral ; k, lastthoracic segment and abdomen, lateral ; 1, 2nd antenna ; m, mandibular palp ; n,mandible blade ; o, ist maxilla ; p, 2nd maxilla ; q, ist leg. Bar scale o-i mm unlessindicated. NEW AND RARE CALANOID COPEPODS 353 Euaugaptilus latifrons (Sars, 1907) MATERIAL EXAMINED : 2 females from 2 Nii3H hauls made at 800-475 m depthoff Fuerteventura (28N I4W) on 26 November 1965. 3 females from 2 RMT ihauls made at 700-505 m depth in a position i8N 25W, between 14 and 15 Novem-ber 1969. DISCUSSION. The present females (Fig. 261- q) agree generally with the descrip-tions of Sars (1924, 1925), Sewell (1932) and, particularly, Tanaka & Omori (1974).There is tremendous variation in the number of setae on the 2nd maxilla in thisspecies. Tanaka & Omori discussed this in relation to their specimen and those ofSars, Sewell and Vervoort (1965). The present females have the following numbersof setae on the various lobes of the 2nd maxilla : Station no. of Lobes of 2nd maxilla specimen 123456 Endopod 7089 no. 16 i o i 2 5 6 16 (fig. 26p) no. 1 6 (2nd specimen) 10122 14 no. I7\ . 101235 J 2 (right side) ' >same specimen j. * . , . ' no. 17 J 10125 I 7 (leftside) 5825 no. 4 i o i 2 3 17 no. i 1012248 In the 2nd maxilla of some of these females lobe 6 and the endopod cannot bedistinguished, and in one specimen there is some variation between the left and rightside. The present specimens measure 4-96-6-72 mm with a mean of 5-63 mm. Euaugaptilus maxillaris Sars, 1920 MATERIAL EXAMINED : 3 females from 3 Nii3H hauls made at 960-550 m depthoff Fuerteventura (28N I4W), between 24 and 28 November 1965. 3 females and2 males from 3 RMT i hauls made at 785-610 m depth in a position i8N 25W,between 13 and 14 November 1969. 5 females and the males are deposited at theBritish Museum (Natural History), reg. no. 1974 : 664-670. DESCRIPTION. Female (Fig. 27a-g) : Body length 3-27-5-76 mm with a meanof 5-11 mm (6 specimens). The cephalothorax is long and narrow and just over3-5 times as long as the 3-segmented abdomen. The head and ist thoracic segmentare separate. The forehead and hind margin of the cephalothorax are rounded.The genital segment is 1-8 times as long as the combined length of the two subsequentsegments. The ist antenna has 25 segments and exceeds the body length by4-5 segments. Except for the ist maxilla the remaining mouthparts are as figuredby Sars (1924). The distal setae of the 2nd maxilla and maxilliped have very small'buttons'. The ist maxilla (Fig. 276, f) differs from the descriptions of Sars (1924,1925) and Tanaka & Omori (1974) in the development of the endopodite. BothSars' and Tanaka & Omori's specimens had a single endopodite segment with fivesetae. One of the females illustrated here has two endopodite segments bearing 354 FIG. 27. a-g, Euaugaptilus maxillaris ?. a, body, dorsal ; b, rostrum, ventral ; c,mandibular palp ; d, mandible blade ; e, ist maxilla ; f, ist maxilla, small specimen ;g, 2nd maxilla, h-i, Euaugaptilis maxillaris $. h, body, dorsal ; i, 5th legs. Barscale o-i mm unless indicated. three and four setae (Fig. 276). Both the segmentation and setation of this endo-podite are subject to variation however, since two other specimens have only asingle segment with six and seven setae respectively. The endopodite of the istmaxilla of the smallest female (3-27 mm, Fig. 27f) agrees with Sars' account but the2nd and 3rd endites have two and one setae respectively - the converse of thearrangement in both previous accounts and in the other specimens here. Theswimming legs are as described by Sars. Male (Fig. 27!!, i) : Body length 5-12 mm (both specimens). The cephalothoraxis 3-5 times as long as the 5-segmented abdomen. The head and ist thoracic seg-ment are separate ; thoracic segments 4 and 5 are fused. In dorsal view the fore-head protrudes more than in the female ; the rostrum has two fairly long filaments.The furcal rami are twice as long as wide. The left ist antenna is presumably NEW AND RARE CALANOID COPEPODS 355 geniculate but is broken in both specimens ; the right ist antenna has 25 segmentsand exceeds the body length by five segments. The remaining mouthparts areidentical to those of the present females. Both specimens have only a single endo-podite segment in the ist maxilla, carrying seven and eight setae respectively ;otherwise this appendage is as shown in Fig. 276. The swimming legs are the same as in the female. The 5th pair of legs is slightlyasymmetrical. The external exopodite spines on the left leg are missing in theillustrated specimen but present in the other. The 2nd basipodite segment of bothlegs has a small patch of spines. The endopodites are 3-segmented on both sidesand are omitted here for clarity. REMARKS. The female of E. maxillaris has only been described and figured bySars (1924, 1925) and by Tanaka & Omori (1974). The male has hitherto beenunknown. The present female specimens agree with these descriptions and I haveno doubt that they are conspecific. The variation in structure of the ist maxillaaccords with the intraspecific variations noted in Euaugaptilm by Matthews (1972)and by Tanaka & Omori (1974). Similar differences are also present in several ofthe species described here. Euaugaptilus ? longicirrhus (Sars, 1905) MATERIAL EXAMINED : i female taken in an NiiaH fished at 800-680 m depthoff Fuerteventura (28N I4W) on 26 November 1965. 3 females and I male from2 RMT i hauls made at 500-410 m depth in a position i8N 25W, between 22 and24 November 1969. 3 females and the male are deposited at the British Museum(Natural History), reg. no. 1974 : 671-674. DESCRIPTION. Female (Fig. 28a-c) : Body length 4-18-4-56 mm with a meanof 4-31 mm (4 specimens). The general appearance of the body is as shown by Sars(1924). The rostral filaments are very long. Except for the 2nd maxilla themouthparts of the present females are the same as those of the male, and fit Sewell's(1947) description better than Sars'. The mandibular palp is very reduced with theexopod having four and the endopod two setae ; the blade has two more teeth thaneither Sars' or Sewell's specimens. The ist maxilla is reduced and agrees withSewell's description. The 2nd maxilla has 3 (2 long and i spike), i, 2, 3, 2 and 3setae on lobes 1-6 respectively, whereas Sars' specimen had , i, 2, 3, 3, 3 setaeand Sewell's 2, ,2,3,2 and 3. The legs all have 3-segmented exopods and endo-pods - the ist pair thereby agreeing with Sewell's account rather than Sars'. Male (Fig. 28d-h) : Body length 3-42 mm. The cephalothorax is similar inshape to the female but has slightly more conspicuous shoulders. It is 3-7 times aslong as the 5-segmented abdomen. Thoracic segments 4 and 5 are fused and thehind margin of the cephalothorax is rounded. The rostral filaments are very long.The left ist antenna reaches to the end of the abdomen ; it is geniculate and has 21segments. The right ist antenna is broken. The mouthparts are as shown and areidentical to the female except that the ist lobe of the 2nd maxilla has only one seta.The swimming legs are the same as in the female. The 5th pair of legs are slightly 356 H. S. J. ROE n FIG. 28. a-c, Euaugaptilus longicirrhus $. a, body, dorsal ; b, rostrum, ventral ; c, istleg. d-h, Euaugaptilus longicirrhus <J. d, body, dorsal ; e, mandible ; f, ist maxilla ;g, 2nd maxilla ; h, 5th legs, i-o, Euaugaptilus rigidus <$. i, head, dorsal ; j, foreheadand rostrum, lateral ; k, last thoracic segment and abdomen, lateral ; 1, mandible ; m,ist maxilla ; n, 2nd maxilla ; o, 5th legs. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 357 asymmetrical. The 2nd exopodite segment of the right leg has a small haired lobe ;the 2nd basipodite segment of both legs has a patch of raised spots on its internaledge. The endopods of both legs are 3-segmented. REMARKS. E. longicirrhus was described by Sars (1924, 1925) from an adultfemale. The only other description is by Sewell (1947) who attributed a stage Vfemale copepodite to this species. Matthews (1972) agreed with Sewell and kindlyidentified the present female taken off Fuerteventura (Roe, 1972^). The appendagesof Sewell's specimen differ from those of Sars' in several respects - the exopodite ofthe ist pair of legs has three segments not two, and the mandibular palp and istand 2nd maxillae differ in details of their setation. The present specimens showalmost complete agreement with Sewell's account and if his copepodite is E. longi-cirrhus then the specimens described here are also this species. Euaugaptilus rigidus (Sars, 1907) MATERIAL EXAMINED : 2 females from 2 NiisH hauls made at 590-450 m depthoff Fuerteventura (28N I4W), between 15 and 26 November 1965. i female andi male from 2 RMT i hauls made at 785-300 m depth in a position i8N 25 W,between 13 and 14 November 1969. The male is deposited at the British Museum(Natural History), reg. no. 1974 : 675. DESCRIPTION. Male (Fig. 28i-o) : Body length c. 4-3 mm but as the cephalotho-rax was fractured accurate measurements are impossible. The general appearanceof the body, especially the shape of the head and hind margin of the cephalo-thorax, are as for the female. The head and ist thoracic segment are separate butthoracic segments 4 and 5 are fused. The rostrum has two filaments of mediumlength. The abdomen has five segments ; the furcal rami are about 1-5 times aslong as wide. The ist antenna are broken off on both sides. Except for the 2ndmaxilla the other mouthparts are identical to the female described by Sars (1924,1925). On the present specimen the first six lobes of the 2nd maxilla bear 2, i, i,3, 2 and i setae respectively ; whereas Sars' female had i, i, i, 2, 2 and i setae onthese lobes. Slight variation in the setation of this appendage occurs, however,since the female described by Tanaka (1964) had 2, i, i, 3, 2 and 2 setae and thepresent females have 2, i, i, 3, 2, i (two specimens) and 2, , i, 3, 2, i setae respect-ively. The first four pairs of legs are identical to the female. The 5th pair of legsis slightly asymmetrical. The 2nd exopodal segment of the right leg has a pointedlobe on its inner margin ; the outer spine of this segment is missing and apparentlybroken off. The 2nd basipodite segment of both legs has a small corrugated patch.The endopodite on both sides has three segments and is omitted for clarity. REMARKS. The agreement between this specimen and the female is so good thatI have no doubt that they are conspecific. The slight variation in setation of the2nd maxilla is not surprising since Matthews (1972) found a considerable degree ofintraspecific variation in this genus. Vervoort (1965) reported a damaged male ofthis species, measuring 4-30 mm, but it has hitherto been undescribed. The maleE. sp. 3 of Grice & Hulsemann (1967) has been discussed earlier (p. 351). 358 H. S. J. ROE Euaugaptilus mixtus (Sars, 1907) Euaugaptilus propinquus Sars, 1920 : 17 ; 1924 : pi. 102 ; 1925 : 297 ; Rose, 1933 : 221, 230,fig. 283 ; Vives, 1970 : 554-555 ; 1972 : 221-222 ; Matthews, 1972 : 39 ; Tanaka & Omori,1974 : J 93 J 97> J 99 2OO > 248-250, figs 3h, 26. MATERIAL EXAMINED : 2 females in 2 Nii3H hauls made at 960-750 m depthoff Fuerteventura (28N I4W), between 25 and 28 November 1965. 5 femalestaken in 5 RMT I hauls made at 900-410 m depth in a position i8N 25W, between12 and 16 November 1969. 6 females are deposited at the British Museum (NaturalHistory), reg. no. 1974 : 676-681. DESCRIPTION. Female (Fig. 2911-0) : Body length 3-50-3-80 mm with a meanof 3-65 mm (7 specimens). The general appearance of the body is as shown by Sars(1924), Hulsemann (1967) and Tanaka & Omori (1974). The cephalothorax is2-6 times as long as the 3-segmented abdomen. The head and ist thoracic segmentare separate ; thoracic segments 4 and 5 are fused. The forehead is rounded andthe rostrum has two filaments of medium length. The hind margin of the cephalo-thorax is rounded in dorsal view but slightly angular when seen laterally. Thegenital segment is about 1-5 times as long as the combined length of the two subse-quent segments. The furcal rami are twice as long as wide. The ist antenna has25 segments and exceeds the body length by 2-3 segments. The remaining mouth-parts are as shown but there is slight variation in the setation of both the 2nd antennaand ist maxilla, and in the number of mandibular teeth. The 2nd antenna wasexamined in three specimens, two of which had nine large setae on the endopoditeand the 3rd (Fig. 29]) had eight. The ist outer lobe of the ist maxilla has six largeand one small setae in three specimens, six large in one, five ? large in one, and fivelarge and one small in another (Fig. 29m). The mandibular teeth are in groups of2, 2 and 3 in four specimens (Fig. 29!), but in two of these four examples the secondtooth of the middle group is minute and could easily be overlooked ; a further speci-men has 2, 2 and 2 teeth, and another 2, i and 3. The distal setae of the 2nd maxillaand maxilliped have well-developed 'buttons'. The legs agree with previousdescriptions. REMARKS. E. mixtus was described briefly by Sars in 1907 from Trincesse Alice'station 1794. It does not, however, appear in the list of copepods taken at thisstation in Sars (1925). The only other record of this species is by Lysholm &Nordgaard (1945). This specimen was probably identified by Sars and was re-described by Hulsemann (1967). E. propinquus was described briefly by Sars in1920 and more fully in 1924 and 1925. It was recorded by Vives (1970, 1972) and afurther description has recently been given by Tanaka & Omori (1974). Sars listedhis specimens as coming from Trincesse Alice' stations 1781, 2738 and 3021. I haveborrowed a female specimen of E, propinquus from the Oceanographic Museum,Monaco, which was caught at station 1768 in 1904. This specimen is not mentionedby Sars and it is not on his list of species taken at this station. Hulsemann (1967) pointed out the similarity between E. mixtus and E. propinquusand said that they could be distinguished by (a) a slightly different shaped forehead,(b) the mandibular tooth arrangement is 2, i and 3 in E. mixtus and 2, 2 and 2 in NEW AND RARE CALANOID COPEPODS 359 n FIG. 29. a-g, Euaugaptilus hyperboreus <J. a, 2nd antenna ; b, mandibular palp ; c,mandible blade ; d, ist maxilla ; e, 2nd maxilla ; f, maxilliped ; g, 5th legs, h-o,Euaugaptilus mixtus ?. h, body, dorsal ; i, forehead and rostrum, lateral ; j, andantenna (different specimen) ; k, mandibular palp ; 1, mandible blade ; m, ist maxilla ;n, 2nd maxilla ; o, maxilliped (different specimen). Bar scale o-i mm unless indicated. 20 360 H. S. J. ROE E. propinquus, and (c) the ist maxilla of E. mixtus has one seta on the 2nd basipoditeand six large setae on the exopodite, whereas these segments in E. propinquus havetwo setae and five large and one small seta respectively. The female described by Tanaka & Omori (1974) is identical to the present'Discovery' specimens. These latter females are intermediate between E. mixtusand E. propinquus (Roe, 1972^). In all of them the setation of the ist maxillaagrees with that of E. mixtus but the arrangement of the mandibular teeth varies.One specimen has the pattern described by Hulsemann for E. mixtus, one thatdescribed by Sars for E. propinquus, and four have a varying and intermediatearrangement. Without dissection it is difficult to ascertain the setation of the istmaxilla of the 'Princesse Alice' specimen. However, it definitely has five large andone small seta on the exopod but I could see only one seta on the 2nd basipodite.The mandibular tooth arrangement of this specimen differs from any describedabove and is I, I and 4. The cephalothorax of this specimen is distorted and theshape of the forehead cannot be accurately determined. The differences between these two species are very slight. The presence of anadditional seta on the ist maxilla and one of a different size seem well within therange of intraspecific variation seen in several Euaugaptilus spp. (for example, seeP- 355)- The shape of the forehead may well be variable and the mandibular toothpattern certainly is. Furthermore, it is clear that the detailed structure of Sars'material was not uniform. I believe that there is no essential difference betweenE. mixtus and E. propinquus and that they are synonymous. Euaugaptilus hyperboreus Brodsky, 1950 Euaugaptilus hyperboreus ? ; Roe, 19720 : 299, 311 ; 19720" : 1035.E. elongatus ; Roe, 19720 : 299, 306, 311. MATERIAL EXAMINED : 2 females and i male taken in 3 Nii3H hauls made at950-600 m depth off Fuerteventura (28N I4W), between 26 and 28 November1965. 2 females taken in 2RMT I hauls made at 1220-800 m depth and I femalein an RMT 8 fished at 1020-910 m in a position i8N 25W, between 14 and 18November 1969. DISCUSSION. Tanaka & Omori (1974) have recently clarified the differencesbetween this species and Euaugaptilus elongatus (Sars, 1905). In the course of thepresent work I borrowed a male and a female specimen of E. elongatus from theOceanographic Museum, Monaco, both of which were taken at 'Princesse Alice'station 1781 on 21 August 1904. These specimens and the present 'Discovery'material confirm the differences in the setation of the ist and 2nd maxillae observedby Tanaka & Omori. These differences are shown in Table I. Slight variation occurs in the setation of the ist maxilla endopodite of E. hyper-boreus, but none of the present specimens, nor those examined by Tanaka & Omori,have three setae on this lobe. Some differences are also present in the setation ofthe 2nd basipodite of the ist maxilla, where the 'Discovery' male (Fig. 2ga-g) has NEW AND RARE CALANOID COPEPODS 361 TABLE i The number of endopodite setae on the ist and 2nd maxillae ofEuaugaptilus hyperboreus and E. elongatus Specimen Species ist maxilla 2nd maxilla 'Princesse Alice' Station 1781 $ E. elongatus 3 8 'Princesse Alice' Station 1781 <? E. elongatus 3 8 (right) 9 (left) 'Discovery' station 5825 no. 7 $ E. hyperboreus 2 15 'Discovery' station 5827 no. 3 $ E. hyperboreus 2 15 'Discovery' station 5827 no. 4 J E. hyperboreus i 15 'Discovery' station 7089 no. 12 $ E. hyperboreus i 15 'Discovery' station 7089 no. 14 $ E. hyperboreus i 14 'Discovery' station 7089 no. 34 $ E. hyperboreus 2 14 three setae on the left side (Fig. 2gd) but four on the right. Similar variation occurson the endopod of the 2nd maxilla of the male E. elongatus from Monaco (Table i).The specimens of E. hyperboreus have a spinous ridge at the base of the externalexopodite spines of the ist leg as described by Tanaka & Omori. Although closelyrelated, these two species show consistent differences, especially in the number ofendopodite setae on the 2nd maxilla. They should, I believe, be maintained asseparate species. Family BATHYPONTIIDAEBathypontia sarsi Grice & Hulsemann, 1965 Bathypontia minor Sars, 1907 : 27 ; 1924 : pi. 127 ; 1925 : 360 ; Lysholm & Nordgaard, 1945 :42 ; Wilson, 1950 : 171, fig. 303 ; Owre & Foyo, 19640, : 343 ; 19646 : 367 ; 1967 : 100,figs 720-722, 724-726 ; 1972 : 494 ; Tanaka, 1965 : 379. MATERIAL EXAMINED : 2 females and 3 males from 5 RMT i hauls made at1250-700 m depth in a position i8N 25W, between 14 and 18 November 1969.Deposited at the British Museum (Natural History), reg. no. 1974 : 682-686. DESCRIPTION. Female (Fig. 30a-i) : One specimen is 3-04 mm in length and theother is too badly damaged to measure. The cephalothorax is oval and 2-9 timesas long as the 4-segmented abdomen. The head and ist thoracic segment areseparate, as are thoracic segments 4 and 5. The rostrum is large, rounded andcurved downwards ; it has no filaments or points. The last thoracic segment issymmetrical with ventrally directed points. The 2nd and 3rd abdominal segmentshave small spines on their hind margins. The furcal rami are slightly longer thanwide. The ist antenna has 23 segments and reaches to the hind edge of the genitalsegment. The remaining mouthparts are as shown. The mandible blade has alarge tooth (Fig. 3od). The left 2nd maxilla has only five large hooked setae (Fig.3of), but the right one has seven. Swimming legs 1-4 are similar to those of themale. The external exopodite spines on the 2nd pair of legs are symmetrical. The5th pair of legs is uniramous with three segments on each side ; the terminal segment 362 H. S. J. ROE FIG. 30. a-i, Bathypontia sarsi ?. a, body, dorsal ; b, last thoracic segment, lateral ;c, 2nd antenna ; d, mandible ; e, ist maxilla ; f, 2nd maxilla, left ; g, maxilliped ;h, 5th leg ; i, 5th leg, other specimen, j-s, Bathypontia sarsi <$. ], body, dorsal ;k, last thoracic segment, lateral ; 1, mandible blade ; m, ist leg ; n, 2nd leg, left ; o, 2ndleg, outer edge of right exopodite ; p, 3rd leg ; q, 4th leg ; r, 5th leg, right ; s, 5th leg,left. Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 363 has one long and one short spine. The arrangement of these spines is slightlyasymmetric in one specimen (Fig. 30!), but in the other both sides are as shown inFig. 3oh. Male (Fig. 30] -s) : Body length 2-74-3-04 mm with a mean of 2-91 mm (3specimens). The general appearance of the body is the same as the female. Theabdomen has five segments. The ist antenna reaches to the end of the cephalo-thorax. The right ist antenna is geniculate and has 20 segments. The remainingmouthparts are identical to those of the female but the right 2nd maxilla has sevenlarge setae and the left only six. The swimming legs are as shown. The externalspine on the 2nd exopodite of the 2nd pair of legs is slightly larger on the left side.The 5th pair of legs is uniramous and completely asymmetrical. The right leg(Fig*. 3or) is longer than the left and has four segments ; the last segment is roughlytriangular in shape, terminates in two unequal spines, has a central pore and has asmall lamella in the distal corner. The left leg (Fig. 305), has five segments of whichsegments 2-5 have bundles of hairs of varying length ; the 5th segment terminatesin one long and one short spine. REMARKS. B. sarsi is a very rare species which has never previously been fullydescribed or figured. Sars (1924, 1925) gave the most complete account, and toverify the identification of the present specimens I borrowed a female and a maleB. sarsi from the collections of the Oceanographic Museum, Monaco. The femalewas taken at 'Princesse Alice' station 1871 in 1904, but this specimen is not men-tioned by Sars, nor is station 1871 included in his station list. The male has nostation number or date but it was probably taken on a cruise to the Azores in 1904-5(G. Testa, pers. comm.). These omissions add to the existing confusion surroundingSars' station listings for this species (Owre & Foyo, 19646). The present 'Discovery' specimens differ from Sars' description in the shape ofthe last thoracic segment and possibly in the structure of the mandible blade. The5th thoracic segment of these specimens has a fairly blunt ventrally directed point,whereas Sars figures this as being very sharp and directed straight back. In the'Princess Alice' male, however, this point is identical to that of the 'Discovery'specimens (Fig. 30k), and in the 'Princess Alice' female it is also blunt but does gostraight backwards. Sars described the mandible blade as having an externaltooth which is only a little larger than the two following teeth. This tooth on thepresent 'Discovery' specimens seems to be larger than described by Sars but it isdifficult to be certain. Owre & Foyo's (1967) photograph of the mandible bladeshows no external tooth. It is impossible to see the mandible blade on either ofthe 'Princess Alice' specimens without dissection. The remaining mouthparts andthe swimming legs of B. sarsi have never been described. In these 'Discovery'specimens they are very similar to those of B. similis Tanaka, 1965. The female5th pair of legs agrees with previous accounts. Their asymmetry in one specimenapproaches the condition seen in B. intermedia Deevey, 1973. B. sarsi and B.intermedia differ in several other respects, however, and presumably the slightasymmetry seen here is merely a variation similar to that shown in the setation ofthe 2nd maxilla. The 'Discovery' male's 5th pairs of legs are identical to that of the'Princess Alice' male. All differ from Sars' figure in the shape of the last segment of 36 4 H. S. J. ROE FIG. 31. Bathypontia spinifera <J. a, body, dorsal ; b, last thoracic segment, lateral ;c, 2nd legs ; d, 5th legs. Bar scale o-i mm unless indicated. the right leg, but this is probably due to a difference in the angle at which Sars'figure was drawn. Bathypontia spinifera A. Scott, 1909 Bathypontia sarsi ; Wheeler, 1970 : 12, figs 77-90. MATERIAL EXAMINED : i male taken in an RMT i fished at 700-610 m depth ina position i8N 25W on 14 November 1969. DESCRIPTION. Fig. 31 shows the very characteristic 2nd pair of legs and the 5thpair of legs of the male B. spinifera. REMARKS. Deevey (1973) pointed out the overall similarity between B. spiniferaand B. similis Tanaka, 1965. Dr Tanaka kindly sent me some specimens of B.similis, the male of which can easily be distinguished from B. spinifera as it doesnot have the very long spine on the right 2nd leg. The present specimen measured2-66 mm. Incertae sedis' Xanthocalanus' paululus Park, 1970 ? Amallothrix robustipes Grice & Hulsemann, 1965 : 239, fig. I3f-k ; 1967 : 26 ; Bradford,1973 : 147- MATERIAL EXAMINED : 23 females and 2 males taken in 4 RMT i hauls made at1250-800 m depth in a position i8N 25W, between 12 and 18 November 1969. NEW AND RARE CALANOID COPEPODS 365 9 females and the males are deposited in the British Museum (Natural History),reg. no. 1974 : 687-697. DESCRIPTION. Female (Fig. 32a-n) : Body length 1-14-1-29 mm with a meanof i-22 mm (19 specimens). The cephalothorax is oval and 4-8 times as long as the4-segmented abdomen. The head and ist thoracic segment are almost completelyfused but the 4th and 5th thoracic segments are separate. The head is character-istically helmet-shaped ; there are no apparent rostral filaments. The 5th thoracicsegment protrudes laterally and is indented. The genital segment is longer than thecombined length of the three subsequent segments. The furcal rami are abouttwice as long as wide. The ist antenna reaches to the hind edge of the genitalsegment. In the 2nd maxilla, lobes 4 and 5 each have a large curved seta armed withteeth, and the endopodite has six setae of which at least two are plumose but none,as far as can be seen, are brush-like. The remaining mouthparts and the legs areas shown. Male (Fig. 32 o-r) : Body length 1-29-1-37 mm. The cephalothorax is 3-2 timesas long as the 5-segmented abdomen. The head and ist thoracic segment areseparate, as are thoracic segments 4 and 5. In lateral view the head is the sameshape as in the female ; there are no rostral filaments. The ist antenna has 22segments and exceeds the body length by 2-3 segments. Some of the mouthpartsare reduced as follows. The mandible blade is smaller than that of the female.The ist maxilla has fewer setae on almost all lobes - 5 on the ist outer lobe, 2 onthe exopod, and i, I and 3 on the 3rd, 2nd and ist inner lobes respectively. Allthe setae except those on the outer lobe are very small and feeble. The 2nd maxillahas small, weak setae on all five lobes with a single larger seta on the 5th lobe ; theendopod has six sensory seta, all of which are, apparently, worm-like. The istbasipodite of the maxilliped has fewer setae than in the female. Legs 1-3 are iden-tical to those of the female ; the 4th pair of legs is broken in both specimens. The5th pair of legs is asymmetrical and reaches back to the middle of the abdomen.The right leg has a 2-segmented exopodite and one short endopodite segment. Inthe left leg the endopodite is also only a single segment but it is almost as long asthe 3-segmented exopodite ; the 2nd exopodal segment has a lamella which partiallyoverlaps the 3rd segment ; this last segment bears clusters of hairs of varying length. REMARKS. Despite the reduction in their mouthparts the present males are sosimilar to the females in the curious shape of the head and the structure of theswimming legs that I believe that they are conspecific. The present females differfrom Park's (1970) description in only a few details. Park's specimens had onemore seta on the ist maxilla, two more on the endopod of the 2nd maxilla, longerist antennae and two fine rostral filaments. I could see no rostral filaments on thepresent specimens but this, together with the differences in setation, may be dueto the difficulties in accurately observing these. Overall agreement is so good,however, that there is little doubt that they are the same species. X. paululus is very similar to Amallothrix robustipes Grice & Hulsemann, 1965.I have examined the type of this latter species in the British Museum (NaturalHistory). The shape of the cephalothorax is the same as that of the present femalesbut unfortunately the slide (s ?) of the appendages is not in the Museum's collection. 366 H. S. J. ROE FIG. 32. a-n, ' Xanthocalanus' paululus $. a, body, dorsal ; b, head, lateral ; c, lastthoracic segment and abdomen, lateral ; d, ist antenna ; e, 2nd antenna ; f, mandible ;g, ist maxilla ; h, 2nd maxilla ; i, maxilliped ; j, ist leg ; k, 2nd leg ; 1, 3rd leg ; m, 4thleg ; n, 5th leg. o-r, 'Xanthocalanus' paululus $. o, body, dorsal ; p, last thoracicsegment and abdomen, lateral ; q, 5th leg ; r, 5th leg, exopodite segments 2 + 3 left leg.Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 367 Of the mouthparts of A. robustipes only the ist antenna and the ist maxilla haveever been described (Grice & Hulsemann, 1965, 1967). The ist maxilla is verysimilar to that of the present species, and the swimming legs are apparently identicalexcept for the absence of small spines on the endopod of the 2nd leg. The 5th legdiffers in having the terminal spine longer than the internal spine. Without adescription of the remaining mouthparts, however, especially the 2nd maxilla, it isimpossible to ascertain whether or not A . robustipes and X. paululus are synonymous. At the moment it is not possible to place this species into any of the four familieswhich have sensory setae on the endopod of the 2nd maxilla. Bradford (1973) wasunable to assign either A . robustipes or X. paululus to either the Phaennidae or theScolecithricidae. This discovery of the male X. paululus confirms her opinion.This species differs from both these families (as defined by Bradford) in the followingrespects : (a) the endopod of the 2nd maxilla apparently lacks any brush-like setae,(b) there are no large spines on the posterior surfaces of the swimming legs, and (c)the male 5th pair of legs are biramous and have symmetrical basipodites. Thepresence of enlarged setae on lobes 4 and 5 of the 2nd maxilla further distinguishesit from the Scolecithricidae. It is distinct from the Diaixidae and Tharybidae inhaving reduced mouthparts in the male. It is further distinguished from theDiaixidae by the female possessing a 5th pair of legs and by the relatively simplenature of the male 5th legs. This species is apparently closest to the Tharybidae but pending a revision ofthis family in relation to the three others mentioned here it seems advisable to recordit as incertae sedis. To avoid any future additional synonomy I have retained thegeneric name Xanthocalanus for this species although it is certainly not of this genus,nor, if A. robustipes is synonymous, of Amallothrix either. Species 1 MATERIAL EXAMINED : 2 females taken in 2 RMT i hauls made in a positioni8N 25W ; one at 1010-900 m depth on 13 November 1969 and the other at900-800 m on 14 November 1969. Deposited at the British Museum (NaturalHistory), reg. no. 1974 : 698-699. DESCRIPTION. Female (Fig. 33) : Body length 1-29 mm and 1-44 mm. Thecephalothorax is oval and about 5 times longer than the 4-segmented abdomen.The head and ist thoracic segment are separate, as are thoracic segments 4 and 5.The rostrum is a curved spike containing a central groove - presumably it is formedof two more or less fused filaments. The last thoracic segment is slightly producedlaterally. The ist antennae are broken in both specimens. The other mouthpartsare as shown. The illustrated ist maxilla (Fig. 33f ) has a damaged ist outer lobe ;the other ist maxilla of the same specimen has seven large and two small setae onthis lobe. In the 2nd maxilla all the lobes are at the distal end of the appendage.Lobe 4 has three large curved setae, furnished with many small spines, lobe 5 hastwo such setae and two small ones. The endopodite has eight long thin filamentswhich, as far as can be seen, all taper to a fine point ; none of them is brush-like.The ist leg has a long curved spine on the outer edge of each exopodite segment. H. S. J. ROE a FIG. 33. Species i ?. a, body, dorsal ; b, forehead and rostrum, lateral ; c, last thoracicsegment and abdomen, lateral ; d, 2nd antenna ; e, mandible ; f, ist maxilla ; g, 2ndmaxilla; h, maxilliped ; i, ist leg; j, 2nd leg; k, 3rd leg; 1, 4th leg; m, 5th legs.Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 369 Legs 2 to 4 are incomplete but the endopodites of legs 2 and 3 are covered with fairlylarge spines. The 5th pair of legs is small and uniramous. Each side has threesegments, the third having a long internal spine and two shorter stout spines ; onone side the surface of the 3rd segment has two small spines. REMARKS. As with the preceding species I am unable to identify these femaleswith any of the four families having sensory setae on the 2nd maxilla endopod.These setae are all simple and worm-like in the present specimens, thereby excludingthem from either the Phaennidae or Scolecithricidae according to Bradford (1973).The presence of a 5th pair of legs distinguishes them from the Diaixidae. Theymay be members of the Tharybidae but as their legs are incomplete it is impossibleto be certain. It seems best to record them as incertae sedis until complete specimensare found. ACKNOWLEDGEMENTS I wish to thank Dr J. M. Bradford of the New Zealand Oceanographic Institute,Dr A. L. Rice, then of the British Museum (Natural History), Dr O. Tanaka, c/oOcean Research Institute, Japan, and Dr G. Testa of the Oceanographic Museum,Monaco, for the loan of specimens used in this work ; also Dr G. B. Deevey of theFlorida State Museum, who very kindly sent me a copy of her manuscript on Chiridi-ella ; and Mrs C. Darter who prepared the drawings for reproduction. REFERENCES ALZAMORA, M. M. 1940. Los copepodos pelagicos del mar de Baleares. Notas Resum. Inst. esp. Oceanogr. Ser. 2, no. 99 : 1-15.ANGEL, M. V. 1969. Planktonic ostracods from the Canary Island region ; their depth distributions, diurnal migrations, and community organisation. /. mar. biol. Ass. U.K. 49:515-553.BAKER, A. DE C., CLARKE, M. R. & HARRIS, M. J. 1973. The N.I.O. combination net (RMT 1+8) and further developments of rectangular midwater trawls. /. mar. biol. Ass. U.K. 53: 167-184.BINET, D. & DESSIER, A. 1971. 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Bathypontia (Copepoda : Calanoida) : Six species, one new, from the Sargasso Sea. Proc. biol. Soc. Wash. 86 : 357-372.I974- Chiridiella Sars (Copepoda : Calanoida) : descriptions of species, six new, from the Sargasso Sea. Bull. mar. Sci. 24 : 439-472. 3?o H. S. J. ROE FARRAN, G. P. 1908. Second report on the Copepoda of the Irish Atlantic Slope. Scient.Invest. Fish. Brch. Ire. 1906, II : 3-104. - 1926. Biscayan plankton collected during a cruise of H.M.S. 'Research' 1900. Part XIV.The Copepoda. /. Linn. Soc. (Zool.), 36 : 219-310. - 1929. Copepoda. Nat. Hist. Rep. Br. antarct. Terra Nova Exped. (Zool.), 8 : 203-306.FOXTON, P. 1969. SOND cruise 1965. Biological sampling methods and procedures. J.mar. biol. Ass. U.K. 49 : 603-620. GIESBRECHT, W. 1892. Systematik und Faunistik des pelagischen Copepoden des Golfesvon Neapel and der angrenzenden Meeresabschnitte. Fauna Flora Golfo Napoli, 19 : 1-831. 1897. Notizen zur Systematik der Copepoden. Zool. Anz. 20 : 253-255.& SCHMEIL, O. 1898. Copepoda. i. Gymnoplea. Das Tierreich, 6 : 1-169. GRICE, G. D. 1962. Calanoid copepods from equatorial waters of the Pacific Ocean. FisheryBull. U.S., 61 : 171-246. - 1969- Calanoid copepods from the Caribbean Sea and Gulf of Mexico, i. New speciesand new records from midwater trawl samples. Butt. mar. Sci. 19 : 446-455. & HULSEMANN, K. 1965. Abundance, vertical distribution and taxonomy of calanoidcopepods at selected stations in the North-east Atlantic. /. Zool. 146 : 213-262. - 1967. Bathypelagic calanoid copepods of the western Indian Ocean. Proc. U.S. natn. Mus. 122 : 1-67.HARDING, G. C. H. 1972. Ecological observations on north Atlantic deep-sea copepods. Ph.D. Thesis, Dalhousie Univ., Halifax, aoipp.HULSEMANN, K. 1967. Re-description of Euaugaptilus mixtus (Sars) (Copepoda, Calanoida). Crustaceana, 12 : 163-166.HURE, J. & SCOTTO Di CARLO, B. 1968. Two new species of Scaphocalanus (Copepoda, Calanoida) from the Mediterranean Sea. Pubbl. Staz. zool. Napoli, 36 : 152-166.LYSHOLM, B. O. & NORDGAARD, K. F. 1945. Copepoda from the 'Michael Sars' North Atlantic Deep-sea Expedition, 1910. Rep. scient. Results Michael Sars N. Atlant. deep Sea Exped. 5 : i -60.MATTHEWS, J. B. L. 1972. The genus Euaugaptilus (Crustacea, Copepoda). New descriptions and a review of the genus in relation to Augaptilus, Haloptilus and Pseudaugaptilus. Bull. Br. Mus. nat. Hist. (Zool.), 24 : 3-71.OWRE, H. B. & FOYO, M. 19640. Plankton of the Florida Current. Part IX. Additions to the list of Copepoda, with descriptions of two rare species. Bull. mar. Sci. Gulf Caribb. 14:342-358. 19646. Report on a collection of Copepoda from the Caribbean Sea. Bull. mar. Sci. Gulf Caribb. 14 : 359-372. 1967. Copepods of the Florida Current. Fauna Caribaea, No. i, Crustacea, Pt i : Copepoda : 1-137. 1972. Studies on Caribbean zooplankton. Description of the program and results of the first cruise. Bull. mar. Sci. 22 : 483-521.PARK, T. S. 1968. Calanoid copepods from the central North Pacific Ocean. Fishery Bull. U.S. 66 : 527-572.1970. Calanoid copepods from the Caribbean Sea and Gulf of Mexico. 2. New species and new records from plankton samples. Bull. mar. Sci. 20 : 472-546.ROE, H. S. J. 19720. The vertical distributions and diurnal migrations of calanoid copepods collected on the SOND cruise, 1965. I. The total population and general discussion. /. mar. biol. Ass. U.K. 52 : 277-314. 19726. The vertical distributions and diurnal migrations of calanoid copepods collectedon the SOND cruise, 1965. II. Systematic account. Families Calanidae up to and includingthe Aetideidae. /. mar. biol. Ass. U.K. 52 : 315-343. I972C. The vertical distributions and diurnal migrations of calanoid copepods collectedon the SOND cruise, 1965. III. Systematic account. Families Euchaetidae up to andincluding the Metridiidae. /. mar. biol. Ass. U.K. 52 : 525-552. NEW AND RARE CALANOID COPEPODS 371 ROE, H. S. J. igjzd. The vertical distribution and diurnal migrations of calanoid copepodscollected on the SOND cruise, 1965. IV. Systematic account of families Lucicutiidae toCandaciidae. The relative abundance of the numerically most important genera. /. mar.biol. Ass. U.K. 52 : 1021-1044. ROSE, M. 1929. Copepodes pelagiques particulierement de surface provenant des campagnesscientifiques du Prince Albert ler de Monaco. Result. Camp, scient. Prince Albert i, 78 :1-123. 1933. Copepodes pelagiques. Faune Fr. 26 : 1-374. 1942. Les Scolecithricidae (Copepodes pelagiques) de la Baie d'Alger. Annls Inst.oceanogr. Monaco, 21 : 113-170. SARS, G. O. 1902, 1903. Copepoda Calanoida. Crustacea of Norway, 4 : 1-171. 1907. Notes supplementaires sur les Calanoidds de la Princesse-Alice (Corrections etadditions). Bull. Inst. ocdanogr. Monaco, 101 : 1-27. 1920. Calanoides recueillis pendant les campagnes de S.A.S. le Prince Albert de Monaco.(Nouveau Supplement.) Bull. Inst. oceanogr. Monaco, 377 : 1-20. 1924, 1925. Copepodes particulierement bathypelagiques provenant des campagnesscientifiques du Prince Albert ler de Monaco. Result. Camp, scient. Prince Albert i, 69,pis 1-127 (1924) : 1-408 (1925). SCOTT, A. 1909. The Copepoda of the Siboga Expedition, Part i, Free-swimming, littoral and semi-parasitic copepoda. Siboga Exped. 29a : 1-323.SCOTT, T. 1894. Report on Entomostraca from the Gulf of Guinea, collected by John Rattray, BSc. Trans. Linn. Soc. Lond. Series 2, Zool, 6 (i) : 1-161.SEWELL, R. B. S. 1929. The Copepoda of Indian Seas. Calanoida. Mem. Indian Mus. 10 : 1-221. 1932. The Copepoda of Indian Seas. Calanoida. Mem. Indian Mus. 10 (contd) : 223-407. 1947. The free-swimming planktonic copepoda. Systematic account. Scient. Rep.John Murray Exped. 8 : 1-303. TANAKA, O. 1956. The pelagic copepods of the Izu region, middle Japan. Systematicaccount. II. Families Paracalanidae and Pseudocalanidae. Publs Seto mar. biol. Lab.5 : 367-406. 1957. The pelagic copepods of the Izu region, middle Japan. Systematic account. IV.Family Aetideidae (Part 2). Publs Seto mar. biol. Lab. 6 : 169-207. 1960. The pelagic copepods of the Izu region, middle Japan. Systematic account. VI.Families Phaennidae and Tharybidae. Publs Seto mar. biol. Lab. 8 : 85-135. 1961. The pelagic copepods of the Izu region, middle Japan. Systematic account. VII.Family Scolecithricidae (Part i). Publs Seto mar. biol. Lab. 9 : 139-190. -1962. The pelagic copepods of the Izu region, middle Japan. Systematic account. VIII.Family Scolecithricidae (Part 2). Publs Seto mar. biol. Lab. 10 : 35-90. 1964. The pelagic copepods of the Izu region, middle Japan. Systematic account. XI.Family Augaptilidae. Publs Seto mar. biol. Lab. 12 : 39-91. -- 1965. The pelagic copepods of the Izu region, middle Japan. Systematic account.XIII. Parapontellidae, Acartiidae and Tortanidae. Publs Seto mar. biol. Lab. 12 : 379-408. & OMORI, M. 1969. Additional report on calanoid copepods from the Izu region. Part 2.Euchirella and Pseudochirella. Publs Seto mar. biol. Lab. 17 : 155-169. 1974. Additional report on calanoid copepods from the Izu region. Part 5. Euaugaptilus. Publs Seto mar. biol. Lab. 21 : 193-267.VERVOORT, W. 1946. The bathypelagic Copepoda Calanoida of the Snellius Expedition, i.Families Calanidae, Eucalanidae, Paracalanidae and Pseudocalanidae. Biological resultsof the Snellius Expedition, XV. Temminckia, 8 : i 181. 1949. Some new and rare Copepoda Calanoida from East Indian seas. Zool. Verh.Leiden, 5 : 3-53. 372 H. S. J. ROE VERVOORT, W. 1951. Plankton copepods from the Atlantic sector of the Antarctic. Verh. K. ned. Akad. Wet. Sect. 2, 47 : 1-156.1957. Copepods from Antarctic and sub- Antarctic plankton samples. Rep. B.A.N.Z. antarctic Res. Exped. Ser. B, 3 : 1-160.1965. Pelagic Copepoda. II. Copepoda Calanoida of the families Phaennidae up to and including Acartiidae, containing the description of a new species of Aetideidae. Atlantide Rep. 8 : 9-216.VIVES, F. 1970. Distribucion y migraci6n vertical de los copepodos planct6nicos (calanoida) de SO. de Portugal. Investigation pesq. 34 : 529-564.1972. Los copepodos del SW. de Portugal en junio y julio de 1967. Investigation pesq. 36 : 201-240.WHEELER, E. H. JR 1970. Atlantic deep-sea calanoid Copepoda. Smithson. Contr. Zool. 55 : 1-31.WILSON, C. B. 1942. The copepods of the plankton gathered during the last cruise of the 'Carnegie'. Scient. Results Cruise VII Carnegie, Biology- i : 1-217.1950. Copepods gathered by the United States fisheries steamer 'Albatross' from 1887 to 1909, chiefly in the Pacific Ocean. Bull. U.S. natn. Mus. 100, 14 : 141-441.WITH, C. 1915. Copepoda. i . Calanoida Amphascandria. Dan. Ingolf Exped. 3 (4) : 1-260.WOLFENDEN, R. N. 1 908. Crustacea VIII. -Copepoda. In: National Antarctic Expedition 1901-1904. Natural History IV Zoology : 1-46.1911. Die marinen Copepoden der Deutschen Siidpolar-Expedition 1901-1903. II. Die pelagischen Copepoden der Westwinddrift und des siidlichen Eismeers. Dt. SiidpoL- Exped. 12 (Zool. 4) : 181-380. H. S. J. ROE INSTITUTE OF OCEANOGRAPHIC SCIENCES WORMLEY GODALMING SURREY 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.) 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 BSj jNU > , 01 I A REVISION OF THE SPECIES OF LAFOEIDAE AND HALECIIDAE (COELENTERATA: HYDROIDA) RECORDED FROM BRITAIN AND NEARBY SEAS P. F. S. CORNELIUS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 8 LONDON: 1975 A REVISION OF THE SPECIES OF LAFOEIDAE AND HALECIIDAE (COELENTERATA: HYDROIDA) RECORDED FROM BRITAIN AND NEARBY SEAS BY PAUL FREDERICK SINEL CORNELIUS Pp. 373-426 ; 14 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)ZOOLOGY Vol. 28 No. 8 LONDON: 1975 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, isissued in Jive series corresponding to the ScientificDepartments of 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. 28, No. 8, of the Zoology series.The abbreviated titles of periodicals cited follow thoseof the World List of Scientific Periodicals. World List abbreviation :Bull. Br. Mus. nat. Hist. (Zool.) ISSN 0007-1498Trustees of the British Museum (Natural History), 1975 TRUSTEES OFTHE BRITISH MUSEUM (NATURAL HISTORY) Issued 1 6 December, 1975 Price 3.60 A REVISION OF THE SPECIES OF LAFOEIDAE AND HALECIIDAE (COELENTERATA: HYDROIDA) RECORDED FROM BRITAIN AND NEARBY SEAS By P. F. S. CORNELIUS CONTENTS SYNOPSIS .INTRODUCTION . Page375375 Family LAFOEIDAE 377 Sub-family EULAFOEINAE 377 Genus Filellum .......... 378 F Helium serpens . . . . . . . -378 Genus Grammaria . . . . . . . . .381 Grammaria abietina ........ 382 Genus Lafoea .......... 385 Lafoea dumosa ......... 385 Family HALECIIDAE 390 Genus Halecium . . . . . . . . . .391 Halecium beanii . . . . . . . .391 Halecium halecinum . . . . . . . -393 Halecium labrosum ........ 396 Halecium lankesteri ........ 399 Halecium muricatum ........ 402 Halecium sessile ........ 406 Halecium tenellum ........ 409 Genus Hydranthea . . . . . . . . . 412 Hydranthea margarica . . . . . . . 412 Genus Ophiodissa . . . . . . . . . 414 Ophiodissa mirabilis . . . . . . . . 414 ACKNOWLEDGEMENTS ......... 417 REFERENCES ........... 418 INDEX ............ 423 SYNOPSIS Nominal species of hydroids belonging to the families Lafoeidae and Haleciidae recordedfrom Britain and neighbouring seas are revised. Three species of Lafoeidae and nine speciesof Haleciidae are accepted. INTRODUCTION IDENTIFICATION guides covering the Hydroida faunas of the majority of countriesbordering the southern North Sea have appeared during the last fifty years (Broch,1927, 1928 ; Kramp, 1935 ; Vervoort, 1946 ; Leloup, 1952). In contrast the most 376 P. F. S. CORNELIUS FIG. i. Map of part of western Europe showing the faunal area (cross-hatching),delimited in most places by the 100 fm (183 m) depth contour (continuous thin line). recent work describing all the hydroids of British coastal waters is that of Pennington(1885) in which the hydroid chapters were based largely on the monographs ofHincks (1868) and Allman (1871) . Perhaps owing to the excellence of its illustrationsHincks' somewhat outdated work still often serves as an identification guide to theBritish thecate hydroids. The large amount of European work produced duringthe present century makes a systematic revision of the British hydroids now timely,and two of the thecate/leptomedusan families are revised here. All nominal taxa of hydroids of the families Lafoeidae and Haleciidae recordedfrom the area defined below are evaluated. No new species are proposed. Indeed,it seems necessary to interpret the specific limits of some species more widely thanhitherto, and some features previously regarded as good specific characters areshown to be unreliable. BRITISH LAFOEIDAE AND HALECIIDAE 377 A redefinition of the family Lafoeidae was given recently by Naumov (1960, 1969).He did not, however, treat the genus Hebella Allman (1888 : 39), which has unclearaffinities. It has variously been included in the families Campanulariidae (byAllman, 1888) and Lafoeidae (by Ralph, 1958 ; Vervoort, 1968, 1972), and removedto a separate family, the Hebellidae (by Nutting, in Fraser, 1912 ; Nutting, 1927 ;Fraser, 1937, 1944, 1946). The Hebellidae was established to accommodate generawith separate gonothecae, a hydrothecal diaphragm and a conical hypostome to thehydranth (said to be spherical in Campanulariidae). The two originally includedgenera, Hebella and Scandia Fraser (1912 : 371), were separated as having respec-tively free medusae and fixed sporosacs. It seems logical at present to retain thefamily Hebellidae to accommodate these two genera, which are implicitly excludedfrom the Lafoeidae in Naumov's diagnosis also. The British records of Hebella arediscussed below under Lafoea dumosa. None appears valid. The faunal area outlined in Fig. i corresponds approximately with the continentalshelf. Unavoidably it is arbitrary, some species being limital and none restrictedto it. It was found that to include the whole of the English Channel, southernNorth Sea and all Danish waters would add only a few species to the list of all thecatehydroids known from Britain, whereas to include the coast of Norway would addsub-arctic species. Oslofjord and the whole of the Swedish west coast are included.To the south the latitude of the Isle d'Ouessant (Ushant) forms the boundary, sothat the whole of the English Channel is included ; while to the west, north andnorth-east the 183 m (600 ft) depth contour provides the arbitrary limit. Taking thethecate hydroid fauna as a whole there appears to be only a few species confined tothe deeper parts of the area thus defined, and the British species form, therefore,a recognizable continental shelf assemblage. The material used was drawn mainly from the collections of the British Museum(Natural History), and carries registered numbers of the format 1894.3.5.11. Thenumbers reflect the approximate dates, in reverse, on which the specimens wereregistered and not the dates of collection or deposition in the Museum. Specimensloaned from other museums are so indicated. Scientific names of British algae mentioned follow the checklist of Parke & Dixon(1968). Family LAFOEIDAE Hincks, 1868 DIAGNOSIS. Colony stolonal or erect and branching ; hydrothecae tubular, evenrimmed, without operculum, with or without diaphragm, pedicellate or sessile,adnate in some species ; hydranth with conical hypostome ; gonothecae aggregatedas scapus or coppinia ; sexual generation a fixed sporosac. TYPE GENUS. Lafoea Lamouroux, 1821. REMARKS. Naumov (1960, 1969) has proposed two sub-families, diagnosed asfollows : Subfamily EULAFOEINAE. Lafoeidae lacking hydrothecal diaphragm ;gonosome in form of coppinia (type-genus Lafoea Lamouroux, 1821). 37 P. F. S. CORNELIUS Subfamily LICTORELLINAE. Lafoeidae with hydrothecal diaphragm ; gonosome in form of scapus (type-genus Lictorella Allman, 1888).In the present area only the first subfamily is represented. 1 Sub-family EULAFOEINAE Naumov, 1960With the characters discussed immediately above. Genus FILELLUM Hincks, 1868 Campanularia : Hassall, 1848 : 2223. Capsularia : Gray, 1848 : 151 (part) ; Naumov, 1960 : 280 ; Naumov, 1969 : 303 ; [nonCapsularia Cuvier, 1797 : 665 (= Coryne Gaertner, in Pallas, 1774) (Lamouroux, Saint- Vincent& Deslongchamps, 1824 : 224) ; Gray, 1848 : 88 (= Lafoea Lamouroux, 1821) ; see Remarks]. Reticularia Thomson, 1853 : 443 ; Rees & Thursfield, 1965 : 85 (part). Filellum Hincks, 1868 : 214 [nom. nov. pro Capsularia : Gray, 1848 (part)] ; Vervoort, 1972 : 50. Grammaria : Vervoort, 1946 : 194 (part). DIAGNOSIS. Colony comprising a creeping, irregularly-branched stolon bearinghydrothecae without pedicels. Hydrothecae tubular, curved centrally, withoutoperculum. Gonothecae borne in hermaphrodite coppiniae typical of the sub-family. TYPE SPECIES. Campanularia serpens Hassall, 1848 (Hincks, 1868, by monotypy ;Naumov, 1960 : 280 ; Naumov, 1969 : 303). REMARKS. The genus Capsularia Cuvier, 1797, was considered by Naumov (1960,1969) to be identical with Filellum Hincks, 1868. This seems unlikely, however,since Cuvier stated that the hydranths in Capsularia were non-retractile. Capsulariawas referred to Coryne Gaertner, in Pallas, 1774, by Lamouroux, Saint- Vincent andDeslongchamps (1824 : 220), and this synonymy was discussed and accepted byBedot (1901 : 437). Gray (1848 : 61) also included Capsularia in the synonymy ofCoryne, but elsewhere in the same work (pp. 85-88, 151) used the name for speciesnow included in other genera and also (p. 151) for the generic name of the specieshere called Filellum serpens (Hassall, 1848). Vervoort (1972 : 50) showed the name Reticularia Thomson, 1853, to be preoccupiedby the brachiopod name Reticularia McCoy, in Griffith, 1844. Filellum serpens (Hassall, 1848)(Fig. 2) Campanularia serpens Hassall, 1848 : 2223 ; Hassall & Coppin, 1852 : 163, pi. 21, fig. 4. Capsularia serpens : Gray, 1848 : 151. Reticularia immersa Thomson, 1853 : 443, pi. i6a, figs 2-3. 1 The species Zygophylax pinnata (Sars, 1874), from the second sub-family, has yet to be recordedfrom the area and appears seldom to occur within Continental Shelf depths in Europe. Although Kramp(1935) suggested the species might occur in northern Danish waters there are apparently still no records(K. W. Petersen, pers. comm.). It has, however, been recorded from a depth of 90 m in HardangerFjord, W Norway (Broch, 1918) and from N Biscay at 186 m depth (Billard, 1923). A statement thatthe species occurs along 'the entire coast of Europe from [the] Bay of Biscay' northwards (Naumov,1969) seems erroneous. BRITISH LAFOEIDAE AND HALECIIDAE 379 FIG. 2. Filellum serpens. Part of colony, W Scotland (1967.11.10.5). Scale = 500 [im. Filellum serpens : Hincks, 1868 : 214-215, pi. 41, fig. 4 (syn. Reticularia immersa Thomson) ; Naumov, 1960 : 281, figs 47, 170, 171 ; Naumov, 1969 : 303, figs 47, 170, 171 ; Vervoort, 1972 : 49-50- Lafoea abietina : Billard, 1904 : 164 (see Remarks).Grammaria serpens : Vervoort, 1946 : 194-196, fig. 82.Reticularia serpens : Rees & Thursfield, 1965 : 87-88. TYPE MATERIAL. Holotype : Dublin, 1842, infertile colony on herbariumspecimen of Abietinaria abietina (Linnaeus, 1758), coll. A. H. Hassall, 1973.10.8.4 ;mentioned Gray, 1848 : 151. OTHER MATERIAL EXAMINED. Vattlestraumen, Espegrend, nr Bergen, Norway,30-40 m, 15 Aug. 1962, colony on Abietinaria abietina, infertile fragments in spiritand coppinia on microslide, coll. W. J. Rees, 1962.11.7.10. Vadero Islands, Sweden,i Oct. 1964, colony teased from substrate, on microslide, coll. W. J. Rees,1967.11.10.6. Gaso Ranna, Gullmarfjord, Sweden, 20-30 m, infertile colony onAbietinaria abietina, in spirit and on microslide, coll. W. J. Rees, 1962.11.8.14.Newhaven, nr Leith, Edinburgh, Scotland, infertile colony on Abietinaria abietina,dried material, holotype of Reticularia immersa Thomson, 1853, 1922.6.19.6. OffMillport, Great Cumbrae Island, Bute, Scotland, 40 m, infertile colonies on Kirchen-paueria pinnata (Linnaeus, 1758) and Nemertesia ramosa (Lamarck, 1816), 10 Sep.1970, coll. C. Edwards, 1971.5.11.33. Off lighthouse, Little Cumbrae Island,31 Jul. 1966, infertile colony on unidentified substrate, microslide, coll. W. J. Rees,1967.11.10.5 (Fig. 2). 'Ireland', 13 Jan. 1902, infertile colony on Abietinariaabietina, microslide, coll. E. T. Browne, 1959.9.17.119. False Bay, Republic ofSouth Africa, fertile colony on Amphisbetia operculata (Linnaeus, 1758), spirit, coll.N. A. H. Millard, 1957.4.26.37 (mentioned Millard, 1957 : 203). DESCRIPTION. Colony stolonal, tortuous ; branching frequent and irregular withhydrothecae at irregular intervals. Hydrothecae tubular, wider than stolons, bentupwards at 60-90 approximately in centre ; rim even, sometimes flared, withoutoperculum ; renovations frequent ; basal - of hydrotheca adnate, narrower(Naumov, 1960, 1969), wider (Vervoort, 1972) or same width as upper part. Hydro-thecae more or less spaced out depending on amount of substrate available (Hincks,1868). Hydranth greenish (Hincks, 1868) or lemon-yellow (Hamond, 1957) ; P. F. S. CORNELIUS tubular, with conical hypostome ; 9-12 tentacles, alternately elevated and de-pressed. Gonothecae borne in hermaphrodite coppiniae typical of the sub-family. MEASUREMENTS. See Table i. TABLE i F Helium serpens. Measurements in (Jim HYDROTHECADiameter at rimLength of free partLength of attached part HYDRORHIZA Diameter of stolon COPPINIA (young)Overall lengthOverall width * Dried material. E SCOTLAND (holotype of R. immersa Thomson) 90-140 60* W SCOTLAND(1967.11.10.5) 100-175200-900200-300 100 IRELAND(I959.9.I7-II9) H5-I35 80-170 220-300 100 NORWAY(1962.11.7.10) c. 1500c. 1000 VARIATIONS. Hydrothecae in this species are longer in some specimens than inothers, long hydrothecae usually being those with one or more renovations. Thehydrothecal rim appears to be flared in some specimens and simply tubular in others,with intermediate conditions. The angle at which the distal portion of the hydro-theca is raised varies approximately between 60 and 90. The proportion of thehydrotheca which is adnate varies between and f of the total length. Hydrothecaemay be either narrower (Naumov, 1960, 1969) or wider (Vervoort, 1972) basallythan distally, or of uniform diameter. Hydranth colour has been recorded asgreenish (Hincks, 1868) and lemon yellow (Hamond, 1957). Colonies growing in alimited space - as on another hydroid colony - have the hydrothecae closer spacedthan in colonies with ample substrate - as on a bivalve shell (Hincks, 1868). REPRODUCTIVE SEASON. Fertile material recorded April-August off Norfolk(Hamond, 1957), April off NW France (Teissier, 1965), mid August off Norway(1962.11.7.10). DISTRIBUTION. Common throughout the area. HABITAT. Sublittoral to edge of continental shelf ; apparently not recordedintertidally. Recorded growing most frequently on sertularian hydroids, especiallyAbietinaria dbietina, but also Hydrallmania falcata (Linnaeus, 1758) and Sertulariacupressina Linnaeus, 1758. Also occurs on non-living substrates such as bivalveshells. Hincks' (1868) statement that the species is parasitic almost certainlyreferred to the growth of the species over other hydroids, and was probably notintended to imply an association of living tissues. REMARKS. Many authors have considered the two nominal species F Heliumserpens (Hassall, 1848) and Reticularia immersa Thomson, 1853, to be conspecificand the present examination of type material of both species supports this view. BRITISH LAFOEIDAE AND HALECIIDAE 381 The material recorded by Billard (1904 : 164) from Cherbourg as Lafoea abietinais here identified as Filellum serpens. The specimen was a lafoeid growing over acolony of a sertularian hydroid, A bietinaria abietina. Billard stated that it resembleda colony of F. serpens except that the hydrothecal aperture was not flared (elargie),and hence concluded that the specimen was a stoloniferous growth of Grammariaabietina. Such colony habit has not otherwise been recorded from G. abietina, andit is now known (present paper) that hydrothecae of F. serpens are not invariablyflared. G. abietina is, therefore, unrecorded from the English Channel (see p. 384). Genus GRAMMARIA Stimpson, 1854 Grammaria Stimpson, 1854 : 9 '> Broch, 1918 : 18 ; Totton, 1930 : 161 ; Vervoort, 1946 : 194 (part) ; Naumov, 1960 : 283 ; Vervoort, 1972 : 56.Salacia : Hincks, 1868 : 211 ; [non Salacia Lamouroux, 1816 : 212 ; Stechow, 1922 : 150 ; Rees & Thursfield, 1965 : 149 ; =Thuiaria Fleming, 1828 (see Remarks)].Reticularia : Rees & Thursfield, 1965 : 85 (part). TYPE SPECIES. Grammaria robusta Stimpson, 1854 (= Campanularia abietinaSars, 1850 ; by designation by Totton, 1930). DIAGNOSIS. Hydrocaulus polysiphonic, branched ; hydrothecae tubular, out-ward-curving, even-rimmed, lacking operculum and diaphragm, arranged in longi-tudinal rows ; with hermaphrodite coppiniae. REMARKS. The name Salacia Lamouroux, 1816, was used for the present genusby Hincks (1868). The type species of Salacia is 5. tetracythara Lamouroux, i8i6 2(by monotypy) . The type material was examined by Deslongchamps (in Lamouroux,Saint- Vincent & Deslongchamps, 1824) who criticized Lamouroux' description asinaccurate and commented that the vase-shaped gonotheca of the specimen, shownclearly in the original illustration, suggested its affinities were with the genusSertularia Linnaeus, 1758. Later Bale (1884) examined the same specimen andreferred it to Thuiaria Fleming, 1828. Bale also provided a new trivial name,calling the species Thuiaria fenestrata. Like Deslongchamps, Bale consideredLamouroux' description to be inaccurate and, somewhat illogically, regarded Salaciaas unavailable. In this he was followed by Bedot (1901), who did not see the speci-men. Billard (1909) did see the specimen and confirmed that it had the charactersof the genus Thuiaria which name he employed although it was junior to Salacia.Stechow (1922, 1923) seems to have been the first to have included Thuiaria in thesynonymy of Salacia, his usage being followed by Rees & Thursfield (1965). Un-fortunately the type material of Salacia tetracythara was destroyed along with thebulk of the Lamouroux collection at Caen by a bomb on 7 July 1944 (Redier, 1967),but there would seem little doubt from the available evidence that Salacia is asenior synonym of Thuiaria. Hincks (1868) thus seems to have been unjustifiedin including the genus Grammaria Stimpson, 1854, in the synonymy of Salacia.The earliest available name for the present genus is thus Grammaria Totton (1930 : 161) nominated G. robusta Stimpson, 1854, as genotype. It appearsto be conspecific with G. abietina (Sars, 1850). 2 Various subsequent authors have spelt the specific name tetracyttara. 382 P. F. S. CORNELIUS Vervoort (1946) and Rees & Thursfield (1965) regarded as congeneric the twogenera here called Grammaria and Filellum. The distinction adopted here, however,follows Vervoort's more recent opinion (1972) that colony habit is of generic value.Nevertheless further information on the life-cycles of the species concerned and ofother species of Lafoeidae may show this generic distinction to be inappropriate. Grammaria abietina (Sars, 1850)(Fig- 3) Campanularia abietina Sars, 1850 : 139. Grammaria robusta Stimpson, 1854 : g, pi. 3, fig. I. Grammaria ramosa Alder, 1856 : 361-362, pi. 14, figs 1-4. Salacia abietina : Hincks, 1868 : 212-213, pi. 41, fig. 3 (syn. G. robusta Stimpson). Grammaria stentor Allman, 1888 : 48, pi. 23, figs i, la ; Hartlaub, 1905 : 599-600, fig. V 2 (syn.G. intermedia Pfeffer) ; Jaderholm, 1905 : 22-23, pi- 8, figs 4-5 (syn. G. intermedia Pfeffer)(syn. nov.) ; [? non Grammaria stentor: Linko, 1911 : 140-143, fig. 23 ; Naumov, 1960 : 282-283, fig. 173 ; Naumov, 1969 : 305-306, fig. 173 ; (see Remarks)] ; (syn. nov.). Grammaria magellanica Allman, 1888 : 48-49, pi. 23, figs 2, 2a-b ; Vervoort, 1972 : 58-60,fig. 166 ; (syn. nov.). Grammaria insignis Allman, 1888 : 49, pi. 23, figs 3, 2a-b ; (syn. nov.). Grammaria intermedia Pfeffer, 1889 : 53-54. Grammaria abietina : Broch, 1917: i-i 6, figs A-C, pis 1-2 ; Broch, 1918: 18-21; Kramp,1935:127-128, fig. 55; Naumov, 1960:283-284, fig. 174; Naumov, 1969:306-307,fig. 174 ; Calder, 1970 : 1523, pi. 5, fig. i ; Vervoort, 1972 : 56 (syn. G. robusta Stimpson). Grammaria abietina var. brevicyatha Broch, 1918 : 20-21, fig. 5. Reticularia abietina : Rees & Thursfield, 1965 : 85-86 (syn. G. robusta Stimpson). TYPE MATERIAL AND LOCALITY. Colony 25 mm high, near Bergen, Norway,55-75 m, summer of 1849 (Sars, 1850) ; specimen not located. MATERIAL EXAMINED. 3 North Atlantic : Carl Island and Cape Torell, Spitzbergen,colony in spirit, coll. A. E. Eaton, 1874.4.4. 5ga. Unnamed locality, Spitzbergen,fragments of colonies in spirit, coll. A. E. Eaton, 1874.4.4.64. E of Faroes,62oi' N, 5i9' W, 210 m, 1869, two colonies in spirit, coll. H.M.S. 'Porcupine', viaA. M. Norman coll., 1912.12.21.303. Trondheim Fjord, Norway, 1893, fragments ofhydrocauli in spirit and i microslide (Fig. 3), via A. M. Norman coll.,1912.12.21.599. Brattholmen, Hjeltefjord, Espegrend, nr Bergen, Norway,40-90 m, 9 Apr. 1962, infertile part of colony in spirit and one coppinia on micro-slide, coll. W. J. Rees, 1962.10.7.23. Off Huglin, Hardanger Fjord, Norway,180 m, 1879, several colonies in spirit, via A. M. Norman coll., 1912.12.21.305.Shetland Isles, 1861, several colonies in spirit, coll. A. M. Norman, 1912.12.21.302.Northumberland coast, branched colony on herbarium sheet, coll. J. Alder, Han-cock Museum, Newcastle-upon-Tyne [syntype of Grammaria ramosa Alder, 1856 ;figured, Alder, 1856 : pi. 14, fig. i (J. B. Garfath, pers. comm.)]. Dried colony inglass tube, Northumberland coast, coll. J. Alder, 1857.8.3.52 [syntype of Grammariaramosa Alder, 1856]. Northumberland, branched hydrocaulus in spirit, coll. J.Alder, via A. M. Norman coll., 1912.12.21.301 (? mentioned, Hincks, 1868 : 213). 3 With the exception of 1962.10.7.23 all material listed is infertile. BRITISH LAFOEIDAE AND HALECIIDAE 383 FIG. 3. Grammaria abietina. Part of colony, Trondheim Fjord, Norway (1912.12.21.599).Scale = 500 [/.m. NE of Shetland Isles, 6i35' N, o47' E ('Goldseeker' sta. 10), 204 m, 3 Sept. 1908,two fragments of hydrocauli on microslide, via J. Ritchie coll., 1964.8.7.61 (men-tioned, Rees & Thursfield, 1965 : 85). Other areas: Barents Sea, 'Vitiaz' sta. 523,no m, 16 Aug. 1950, large colony in spirit, Leningrad Academy of Sciences ZoologicalMuseum, and microslide donated to British Museum (Natural History), 1975.7.11.1(mentioned as G. stentor Allman, Naumov, 1960 : 283 ; 1969 : 306 ; see Remarks).Near Falkland Islands, 5i35' S, 653g' W, 130 m ('Challenger' sta. 314), 21 Jan.1876, two branched fragments of colonies plus one microslide, coll. H.M.S. 'Chal-lenger', 1888.11.13.36 (syntypes of Grammaria magellanica Allman, 1888). RoyalSound, Kerguelen Island, 4928' S, 70i3' E, 37-110 m ('Challenger' sta. 1490),20 Jan. 1874, fragments of colonies in spirit plus one microslide, coll. H.M.S.'Challenger', 1888.11.13.35 (syntypes of Grammaria stentor Allman, 1888). OffMarion Island, S Indian Ocean, 4643' S, 3804' E, 90-135 m ('Challenger' sta. 145),27 Dec. 1873, colony and fragments in spirit plus one microslide, coll. H.M.S.'Challenger', 1888.11.13.37 ( svn types of Grammaria insignis Allman, 1888). DESCRIPTION. Colony erect, main stem and branches polysiphonic, branchingirregular or imperfectly pinnate, branches tapering near the base. Hydrothecaelong, tubular, variably outward-curving ; rims even, circular, often renovated,frequently slightly flared ; no apparent junction with pedicel ; arranged in 4-8longitudinal rows with hydrothecae of adjacent rows alternate ; if 4 rows, hydro-thecae often in decussate pairs. Hydranth yellow (Hincks, 1868), extensibleroughly one hydrotheca-length beyond aperture ; widest distally, hypostomeconical, 18-20 tentacles held alternately elevated and depressed. Gonothecaeborne in hermaphrodite coppiniae 'arranged on branches in small muffs. Surfaceof coppinia smooth owing to dense arrangement of the sterile tubes, their inwardly-curved ends forming a thick protective tangle above the gonothecal mouths'(Naumov, 1969). 384 P. F. S. CORNELIUS MEASUREMENTS. See Table 2. TABLE 2 Grammaria abietina. Measurements in (Jim NORTH SEA TRONDHEIM FJORD NR BERGEN, NORWAY (1964.8.7.61) (1912.12.21.599) (1962.10.7.23)HYDROTHECA Diameter at rim 240-280 280-360 280-490COPPINIA Overall length 3000 Overall width 1500 Diameter of sterile tubes 90-130 VARIATION. There are normally 4, 5 or 6 rows of hydrothecae on a hydrocaulus,and 8 rows have been recorded (Vervoort, 1972, as G. magellanica}. When in 4rows the hydrothecae are in opposite or almost opposite pairs. The hydrothecalrim is flared in some specimens and straight in others, this character having ap-parently no systematic value. The degree of outward curving of the hydrothecavaries, so that the plane of the hydrothecal aperture is parallel with the axis of thehydrocaulus in some specimens, while in specimens with shorter hydrothecae theplane of the aperture makes an angle of about 45 with the axis, and this charactermay vary even within a colony (Fig. 3). REPRODUCTIVE SEASON. Little information. Fertile specimen taken near Bergen,Norway, 9 Apr. 1962 (1962.10.7.23). DISTRIBUTION. Widely distributed in sub-arctic and arctic areas (Broch, 1918),extending south to much of the North Sea and some Danish waters. There seemsto be no valid record from western coasts south of the Shetlands. Southerlyrecords include Northumberland (Hincks, 1868 ; present material), Skagerrak(Kramp, 1935) and Oslo Fjord (Christiansen, 1972). A record from the north coastof France (Billard, 1904, repeated in Broch, 1918) is probably erroneous (see p. 381)and the statement of Christiansen (1972) that the species occurs on that coast lacksevidence. The weight of published opinion is that the species is absent from theEnglish Channel and southern North Sea (Hincks, 1868 ; Hartlaub, 1895 ; Broch,1927 ; Vervoort, 1946, 1949 ; Leloup, 1952 ; Hamond, 1957 ; Marine BiologicalAssociation, 1957 ; Teissier, 1965 ; Robins, 1969). HABITAT. Published records indicate that the normal depth-range of this speciesis approximately 50-1500 m in boreal waters, rising to 10-250 m in arctic regions(Linko, 1911 ; Naumov, 1960, 1969 ; Christiansen, 1972). The species has beenrecorded on both silty and rocky substrates (Naumov, 1960, 1969 ; Calder, 1970). REMARKS. There seems no doubt from the original description of Grammariaramosa Alder, 1856, that it should be regarded as a junior synonym of the presentspecies and examination of the type series confirms that they are conspecific. Vervoort (1972) has expressed doubt as to the distinctiveness of the three specieserected by Airman (1888), Grammaria insignis, G. magellanica and G. stentor, andfollowing examination of the type material they are here referred to the present BRITISH LAFOEIDAE AND HALECIIDAE 385 species. The characters on which they were based were differences in the branchingof the colony and of the number of rows of hydrothecae, and whethei or not thehydrothecal rim was flared. These characters are now known to be variable in G.abietina and the three species proposed by Allman appear invalid. The material assigned to G. stentor Allman, 1888, by Linko (1911) and Naumov(1960, 1969, examined here) has hydrothecae longer than in G. abietina as heredenned, and may represent a distinct species. If so, a new name will have to beprovided. G. stentor sensu Hartlaub, 1905, and Jaderholm, 1905, fall within theproposed limits of G. abietina. Jaderholm examined the type material of G.intermedia Pfeffer, 1889, and referred it to his own concept of G. stentor. His accountsuggests that G. intermedia also should be referred to G. abietina. G. abietina var brevicyatha Broch, 1818, was distinguished on the basis of short,slightly out-turned hydrothecae, but such variation seems normal for the speciesand no distinct variety need be recognized. The material from Cherbourg, N France, referred to the present species by Billard(1904 : 164) is here identified as F Helium serpens (see p. 381). Genus LAFOEA Lamouroux, 1821 Sertularia : Fleming, 1820 : 83 (part). Lafoea Lamouroux, 1821 : 8 ; Hincks, 1868 : 198 (part) ; Naumov, 1960 : 272 ; Naumov,1969 : 295. NOMENCLATURE. The genus was named after an amateur botanist, Professor deLafoye, of the University of Caen (Lamouroux, 1821 ; Lamouroux, Saint-Vincent &Deslongchamps, 1824). TYPE SPECIES AND MATERIAL. Lafoea cornuta Lamouroux, 1821 : 8, pi. 65,figs 12-14 (by monotypy) ; 44-44^ N, 52-53 W, 55-60 m ; branched colony100 mm long, coll. Capt. Laporte (Lamouroux, 1821) ; Botanical Institute collec-tions, Caen. Specimen destroyed by bomb, 7 July 1944 (Redier, 1967). DIAGNOSIS. Colony either stoloniferous or with stolons united to form erectpolysiphonic hydrocauli with monosiphonic terminal branches. Hydrothecaelacking both operculum and diaphragm ; tubular or bell-shaped, sometimes bentand thus bilaterally symmetrical ; rim even, circular, sometimes flared ; renovationsfrequent ; pedicel twisted if present ; gonothecae borne in hermaphrodite coppiniae. REMARKS. Billard (1909 : 311) examined the genotype material and referred thetype species L. cornuta Lamouroux, 1821, to L. dumosa (Fleming, 1820), supportingthe suggestion of Hincks (1868 : 199) that the two are conspecific. Lafoea dumosa (Fleming, 1820) (Fig. 4) Sertularia dumosa Fleming, 1820 : 83-84.Lafoea cornuta Lamouroux, 1821 : 8, pi. 65, figs 12-14. Campanularia dumosa: Fleming, 1828:548-549; Johnston, 1832:254, pi. n, fig. n ;Johnston, 1838 : 157, pi. 23, figs 2-5 ; Johnston, 1847 : 113-115, pi. 27, figs 2-5, text-figs 386 P. F. S. CORNELIUS Capsularia dumosa : Gray, 1848 : 88. Campanularia fruticosa Sars, 1850 : 138-139 ; (syn. nov.). Campanularia gracillima Alder, 1856 : 361, pi. 14, figs 5-6. Lafoea dumosa: Hincks, 1868:200-201, pi. 41, fig. i, xa (? syn. L. cornuta Lamouroux) ; Nutting, 1899:747-751, pi. 64; Rufford, 1902:62; Billard, 1909:311 (syn. L. cornuta Lamouroux) ; Broch, 1918 : 7-9 ; Totton, 1930 : 158, fig. 14 ; Kramp, 1935 : 123-124, figs 52a, 53 ; Fraser, 1944 : 221-222, pis 45-46, fig. 2O5a-e ; Ralph, 1958 : 310 ; Rees & Thursfield, 1965 : 79-80 ; Teissier, 1965 : 19 ; Calder, 1970 : 1524, pi. 5, fig. 3.Lafoea fruticosa : Hincks, 1868 : 202-203, pi. 41, figs 2, 2a-b (syn. C. gracillima Alder) ; Broch, 1918 : 12-15 (syn. L. pocillum Hincks) ; Totton, 1930 : 157-158, fig. 13 ; Kramp, 1935 : 124-125, fig. 52C-d (syn. L. pocillum Hincks) ; Fraser, 1944 : 223-224, pi. 46, fig. 206 ; Hodgson, 1950:11; Naumov, 1960:275-276, fig. 164; Millard, 1964:13-14, fig. 3a-f ; Rees & Thursfield, 1965 : 80 (syn. L. pocillum Hincks) ; Millard, 1967 : 175-176, fig. 2C ; Naumov, 1969 : 297-298, fig. 164 ; Calder, 1970 : 1524-1525, pi. 5, fig. 4 ; Vervoort, 1972 : 66-74, n s J 9- 21 ( s y n - L. gracillima Alder).Lafoea pocillum Hincks, 1868 : 204, pi. 40, fig. 2 ; Hincks, 1874!) : 147 ; Crawford, 1895 : 260 ; Rufford, 1902 : 62 ; Linko, 1911 : 114-116, fig. 20 ; Naumov, 1960 : 273-274, fig. 161 (syn. Campanularia parvula Hincks) ; Naumov, 1969 : 295-296, fig. 161 (syn. C. parvula Hincks).Hebella pocillum : Ritchie, 1911 : 33.Lafoea gracillima : Broch, 1918:9-11; Totton, 1930:158-159, fig. I5a-b ; Kramp, 1935: 125, fig. 526 ; Fraser, 1944 : 224-225, pi. 46, fig. 207 ; Ralph, 1958 : 310, figs ly, 2a-c ; Rees & Thursfield, 1965 : 80-81 ; Teissier, 1965 : 20 ; Calder, 1970 : 1525, pi. 5, fig. 5.Lafoea fruticosa forma pocillum : Vervoort, 1949 : 148-149. TYPE MATERIAL. Arbroath, Angus, Scotland, 1809 ; 4 location of specimensunknown. MATERIAL EXAMINED. Brattholmen, Hjeltefjord, nr Bergen, Norway, 40-90 m,9 Apr. 1962, part of a colony on microslide, coll. W. J. Rees, 1962.10.7.22 (Fig. 4).Kosterfjord, Sweden, 585o' N, noo' E, 80-100 m, 23 Sep. 1964, hydrocaulus onmicroslide, coll. W. J. Rees, 1965.1.14.73. Vadero Islands, Sweden, 5805' N,n04' E, 80 m, i Oct. 1964, several colonies in spirit, coll. W. J. Rees, 1965.1.14.136and 1966.1.4.65. Loken, Gaso Ranna, Gullmarfjord, Sweden, 25-30 m, 14 May1959, two fertile colonies on Abietinaria sp., in spirit, coll. W. J. Rees, 1959.6.11.36.Gaso, Gullmarfjord, Sweden, 20-30 m, 27 Aug. 1962, hydrocaulus on microslide,coll. W. J. Rees, 1962.11.8.13. Off Balta, Shetland Isles, Scotland, 100 m, colonyin spirit, coll. A. M. Norman, 1912.12.21.237. Oban, Argyll, Scotland, two hydro-cauli on microslide and colony on herbarium sheet, coll. T. Hincks, 1899.5.1.160,216. Oban, Argyll, Scotland, infertile colony on Phycodrys mbens (L.) Batt., 6 inspirit, 1899.5.1.162 (holotype of Lafoea pocillum Hincks, 1868). Off Millport,Great Cumbrae Island, Bute, Scotland, 40 m, 18 Sep. 1970, four colonies in spirit,coll. C. Edwards, 1971.5.11.31. Berwick Bay, Berwickshire, Scotland, andNorthumberland, England, several colonies on four herbarium sheets, coll. G.Johnston, 1847.9.24.68, 69, 70, 71 (mentioned, Gray, 1848 : 88 ; ? Johnston,1838 : 157). 'Deep water, Northumberland' (label with specimen), colony on her-barium sheet, coll. J. Alder, Hancock Museum, Newcastle-upon-Tyne [paratype of 4 Fleming (1820) based the original description on specimens from three Scottish localities: fromrefuse of oyster-boats, 'Newhaven', identified in a later work as being near Edinburgh (Fleming, 1828);'Aberbrothick', now called Arbroath, Angus; and 'Zetland'. The first and last being relatively imprecise,the type locality is here restricted to Arbroath. 8 Det. J. M. Price. BRITISH LAFOEIDAE AND HALECIIDAE 387 n L f FIG. 4. a-e, Lafoea dumosa. a, part of colony, near Bergen, Norway, 40-90 m(1962.10.7.22) ; b, part of colony, east of Straits of Magellan, Argentina, 100 m(1890.4.11.2(1) ; c-e, three adjacent hydrothecae, SW England (1959.9.17.108). f-h,three hydrothecae of holotype of Lafoea pocillum (1899.5.1.162), here referred to L. dumosa.Scale (a-h) = 500 [zm. Campanularia gracillima Alder, 1856 ; figured specimen, Alder, 1856 : pi. 14, fig. 5(J. B. Garfath, pers. comm.)]. Probably off Northumberland, 'in deep water'(Alder, 1856), colony in spirit, coll. J. Alder, 1857.8.3.51 [lectotype of Campanulariagracillima Alder, 1856 (Totton, 1930)]. Scarborough, Yorkshire, England, colonieson two herbarium sheets (one on Flustra sp.) coll. W. Bean, via G. Johnston coll.,1842.12.9.1, 1847.9.24.67 (mentioned, Gray, 1848:88). North Sea, 6oO2' N,3I3'W, 160 m, 19 June 1906 ('Goldseeker' sta. 2ia), two hydrocauli on microslide,coll. J. Ritchie, 1964.8.7.54 (mentioned, Rees & Thursfield, 1965 : 80). North Sea,592i' N, 5oo' W, 125 m, 6 July 1906 ('Goldseeker' sta. 50), hydrocaulus on micro-slide, coll. J. Ritchie, 1964.8.7.50 (mentioned, Rees & Thursfield, 1965 : 81). Ply-mouth, Devon, 8 Mar. 1898, hydrocauli on two microslides, coll. E. T. Browne,1959.9.17.108, 114. Eddystone Ground, NW English Channel, 8 Mar. 1898, sevenhydrocauli on microslide, coll. E. T. Browne, 1959.9.17.111. Eddystone ground,NW English Channel, 30 Mar. 1938, hydrocaulus on microslide, coll. W. J. Rees, P. F. S. CORNELIUS 1969.12.2.45. Probably British, two colonies on herbarium sheets, coll. G. Johnston,1847.9.24.69, 72 (mentioned, Gray, 1848 : 88). Davis Strait, Greenland, 6659' N, 5527' W, 120 m, fragments of hydrocaulus onmicroslide, coll. H.M.S. 'Valorous', 1878.3.26.3. E of Str. of Magellan, Argentina,522o' S, 67^9' W, 100 m, 20 Jan. 1876 ('Challenger' sta. 313), part of colony onmicroslide, coll. H.M.S. 'Challenger', 1890.4. n.2d (Fig. 4). Off Gates Land,Antarctic mainland, 6943' S, i6324' E, 329-366 m, 22 Feb. 1911 (Terra Nova'sta. 194), spirit material and two microslides, coll. British Antarctic (Terra Nova')Expedition, 1929.10.28.64 (mentioned, Totton, 1930 : 158). DESCRIPTION. Colonies stoloniferous or erect. Stoloniferous colonies compriseirregularly-branched monosiphonic hydrorhizae with hydrothecae borne at irregularintervals. Erect colonies consist of polysiphonic hydrocauli, loosely and irregularlybranched, arising from a straggling stolon network. Component perisarc tubes oferect colonies parallel, each bearing hydrothecae at irregular intervals givingbristly appearance. Hydrothecae long, tubular, even-rimmed, variably taperedbelow, sometimes asymmetrically ; pedicel more or less well defined, usually kinked,but sometimes lacking so that hydrotheca is sessile. Form of hydrothecal pedicelusually constant within a colony. Hydranth long, retractile, c. 20 tentacles heldalternately elevated and depressed (Kramp, 1935), hypostome conical. Hydranthand coenosarc often yellow. Gonothecae in hermaphrodite coppiniae. MEASUREMENTS. See Table 3. TABLE 3 Lafoea dumosa. Measurements in [Jim W ENGLISH CHANNEL (1969.12.2.45) HYDROTHECABase to rim (unrenovated) 500-650Diameter at rim 150-220Pedicel length o W SWEDEN(1965.1.14.73) 485-510 100 100-150 N SCOTLAND(1964.8.7.50) 770-830150-21050 W SCOTLAND(1899.5.1.160) 380-550120-140140-170 W SCOTLAND (1899.5.1.162) (holotype of Lafoea pocillum) 120-35090-19080-170 VARIATION. The hydrothecal pedicel varies in length between colonies (seeRemarks) and the degree of bilateral symmetry of the hydrothecae varies also.Neither character appears to be of systematic importance. Specimens from deepwater have fewer hydrothecae per unit length of stolon, but within continentalshelf depths this character seems constant. The so-called pedicel of the presentspecies grades into the base of the hydrotheca. Although a pedicel is readily dis-cernible if long, shorter pedicels are less distinct and it is possible to arrange specimensin a series in which the pedicel becomes progressively less distinct and is finallyabsent (Fig. 4). In each colony the range of pedicel length is small, but pedicellatehydrothecae can be found in colonies largely lacking pedicels, and sessile hydrothecaeoccur in predominantly pedicellate colonies. BRITISH LAFOEIDAE AND HALECIIDAE 389 REPRODUCTIVE SEASON. Fertile specimens recorded August in NW France(Teissier, 1965), 14 May 1959 in W Sweden (1959.6.11.36). The paucity of recordssuggests that in this species reproduction is usually vegetative. DISTRIBUTION. Found throughout the area but apparently not common in thesouthern North Sea and eastern English Channel (Broch, 1918 ; Vervoort, 1946 ;Marine Biological Association, 1957 ; Teissier, 1965 ; Naumov, 1969), not havingbeen found in a faunal survey oft the coasts of Belgium (Leloup, 1952). HABITAT. Occurs at all continental shelf depths, but not intertidally. On awide variety of animal, plant and inert substrates, commonly on other hydroids ;not on sand. REMARKS. The two nominal species Lafoea fruticosa (Sars, 1850) and L. gracillima(Alder, 1856) have been regarded as conspecific by some authors (e.g. Hincks, 1868 ;Hodgson, 1950 ; Naumov, 1960, 1969 ; Millard, 1967 ; Vervoort, 1972) and asvalid species by others (e.g. Broch, 1918 ; Kramp, 1935 ; Fraser, 1944 ; Rees &Thursfield, 1965 ; Calder, 1970). Vervoort's detailed appraisal is particularlyconvincing and his view that the two are conspecific is accepted here. In contrast, it appears that the distinctness of L. fruticosa from L. dumosa hasbeen questioned only by Millard (1964), who described intermediate material. Thetwo species have been distinguished by Hincks (1868) and apparently all subsequentauthors (except Rufford, 1902, and Ralph, I958 6 ) by the presence of a hydrothecalpedicel in L. fruticosa and its absence in L. dumosa which has been held to havesessile hydrothecae. However, the original description of L. dumosa (Fleming, 1820,repeated in Fleming, 1828) referred to 'nearly sessile' hydrothecae, implying thatshort pedicels were present. Johnston (1832) similarly mentioned 'nearly sessile'hydrothecae, stating that they 'can rarely be observed to be twisted at their inser-tions' as shown in his illustration. His later descriptions refer to 'nearly sessile'and 'almost sessile' hydrothecae (Johnston, 1838, 1847). Gray (1848) did not pro-vide a description, but the ten colonies listed by him as L. dumosa and re-examinedduring the present work all have pedicellate hydrothecae. The material was fromJohnston's collection, some being labelled Berwick Bay and probably, therefore,the Berwick Bay material described by Johnston (1832, 1838). Whether or not thisis so it seems probable that Johnston, like Fleming, wished to imply that the pedicelswere short, and not that they were absent. Further, in describing L. gracillima,Alder (1856) stated that the pedicels were 'longer' than in L. dumosa, indicatingthat he too considered L. dumosa to be pedicellate. Thus, when Hincks (1868)later gave specific status to specimens without pedicels and distinguished L. fruticosaas having them he was wrong to refer non-pedicellate material to L. dumosa andshould have provided a new name. This seems never to have been done. Itappears probable, however, that specimens lacking hydrothecal pedicels do notconstitute a valid species and should nevertheless be referred to L. dumosa sensuFleming, Johnston and Alder (op. cit.), L. fruticosa (Sars, 1851) and L. gracillima(Alder, 1856) falling within its synonymy. 8 In a key to the two taxa L. dumosa and L. gracillima Ralph separated them on the basis of short andlong pedicels respectively, but she gave a description of L. gracillima only. Rufford ascribed a shortpedicel to L. dumosa without comment. 22 390 P. F. S. CORNELIUS The two nominal species Lafoea parvula (Hincks, 1853, as Campanularia) and L.pocillum Hincks, 1868, were thought to be conspecific by Naumov (1960, 1969).However, several authors (Broch, 1918 ; Kramp, 1935 ; Vervoort, 1949 ; Rees &Thursfield, 1965) have considered L. pocillum conspecific with L. dumosa (as L.fmticosa) and this opinion is supported by the present examination of the typematerial (Fig. 4). The material referred to Hebella pocillum (Hincks, 1868) byFraser (1937, 1944) and Yamada (1955) is at present indeterminate ; it may requirea new specific name. The type material of L. parvula could not be located, butspecimens labelled L. parvula (off Durham, England, coll. A. M. Norman,1912.12.21.240) seemed identical with infertile colonies of Calicella syringa (Linnaeus,1767) from which the hydrothecal operculae have been lost. It seems plausiblethat L. parvula was founded on such material. Lafoea pygmaea Alder, in Hincks (1868 : 205, pi. 40, fig. 3 ; type localities Tyne-mouth, Northumberland, and Sark, Channel Isles), was referred to Calicella in alater publication by Hincks [i874b : 147 (footnote)]. This was accepted by Jader-holm (1909 : 80), Bedot (1912 : 315), Broch (1918 : 9) and Stechow (1921 : 228).However, Kramp (1935 : 125) and Vervoort (1946 : 199) regarded L. Pygmaea asidentical with L. gracillima (= L. dumosa), while Naumov (1960, 1969) referred it toL. pocillum (= L. dumosa}. The type material in the Hancock Museum, Newcastle-upon-Tyne, and the description of Hincks (1868 : pi. 40, figs 3, 3a-b), showing awell-demarcated pedicel with transverse annulations, are clearly of Calicella and itseems Hincks was justified in placing L. pygmaea in that genus (Cornelius & Garfath,in prep.). L. pygmaea was regarded as conspecific with C. syringa (Linnaeus, 1767)by Jaderholm (1909) and Broch (1918). Family HALECIIDAE Hincks, 1868 DIAGNOSIS. Colonial Hydroida with short, cylindrical hydrothecae usuallywider than deep ; hydrothecal rims even, often flared, renovation common, desmo-cytes large, often birefringent. Hydrothecal pedicel often lacking. Hydranthslarger than hydrothecae. Gonothecae usually sexually dimorphic. Sexual genera-tion said to be a medusa in Campalecium Torrey, 1902 (Kossowska, 1911), eumedusoidin Hydranthea Hincks, 1868, and a sessile sporosac in Halecium Oken, 1815, andOphiodissa Stechow, 1919. TYPE GENUS. Halecium Oken, 1815. 7 REMARKS. The nature of the sexual generation was shown by Rees (1957) to bea valid generic character in only some families among the athecate capitate hydroids.In the Haleciidae it appears to be correlated with other generic characters and isincluded here in the generic diagnoses. A similar correlation has been shown incertain genera of the Campanulariidae (Cornelius, 1975). 7 Oken's book was placed on the list of rejected works by the International Commission on ZoologicalNomenclature (1956 : Opinion 417), but application is currently being made to the Commission forvalidation of the name Halecium (see Cornelius, 1976). BRITISH LAFOEIDAE AND HALECIIDAE 391 The small birefringent bodies found inside the hydrothecal wall throughout thisfamily (Fig. n) and widely referred to as punctae (e.g. Vervoort, 1972) are probablydesmocytes. They resemble those previously described in other classes of coelenter-ates, reviewed by Chapman (1969), but are much larger, being visible in some speciesat only 20 diameters' magnification. A preliminary account of their ultrastructureis to appear elsewhere. It is probable that the 'punctae' of the family Lafoeidaeare also desmocytes. Present knowledge suggests that they have little systematicvalue at species level in either family, but comparative studies of their ultrastructuremight prove differently. Genus HALECIUM Oken, i8i5 7 Halecium Oken, 1815 : 91 ; Johnston, 1847 : 58 ; Hincks, 1868 : 220 ; Fraser, 1944 : 183 ; Naumov, 1960 : 442. Thoa Lamouroux, 1816 : 210 ; Johnston, 1838 : 119.Haloikema Bourne, 1890 : 395. TYPE SPECIES. Sertularia halecina Linnaeus, 1758 : 809 (Oken, 1815 : 91 ; Bedot,1901 : 448). DIAGNOSIS. Haleciidae with branched, erect colonies ; lacking nematophores,large nematocysts, medusae and eumedusoid gonophores. Gonothecae usuallysexually dimorphic. REMARKS. Johnston (1847) drew attention to the seniority of Halecium to Thoaand his synonymy was followed by Hincks (1868) and all subsequent authors.Haloikema was first included in Halecium by Bedot (1911). Halecium torreyi Kossowska, 1911, said by that author to release a medusa, wasplaced in the synonymy of Campalecium medusiferum Torrey, 1902, by Huve (1954). Halecium beanii (Johnston, 1938) (Fig- 5) Thoa beanii Johnston, 1838 : 120-121, pi. 7, figs 1-2. Halecium beanii : Johnston, 1847 : 59-60, pi. 9, figs 1-2 ; Hincks, 1868 : 224-225, pi. 43, figs 2, 2a-c ; Vervoort, 1946 : 161 163, figs 65-66 ; Ralph, 1958 : 332-334, fig. loa-b, e-k ; Rees & Thursfield, 1965 : 105-106.Halecium scutum Clarke, 1876 : 14-15, pi. 4, figs 13-14.Halecium boreale Lorenz, 1886 : 26-27, pi. i, figs 1-2.Halecium beanei Stechow, 1919 : 33 (lapsus pro beanii).Halecium beani : Naumov, 1960 : 447449, fig. 336 (syn. H. scutum Clarke ; H. boreale Lorenz) ; Naumov, 1969 : 483-484, fig. 336. NOMENCLATURE. The species is named after W. Bean, who collected the holotype. TYPE MATERIAL AND LOCALITY. Holotype i Nr Scarborough, Yorkshire, England,in 'deep water', $ colony 42 mm high on herbarium sheet, plus one microslide of$ gonotheca from same colony, coll. W. Bean, 1847.9.18.181, i8ia (Johnston, 1838, 1847). 7 See footnote on opposite page. 392 P. F. S. CORNELIUS FIG. 5. Halecium beanii. a, part of colony, NE England (1956.2.2.27) ; b-c, part ofcolony and $ gonotheca, Isle of Man (1959.10.17.8) ; d, $ gonotheca, near Bergen,Norway (1962.11.7.38). Scale (a-d) = 500 (zm. OTHER MATERIAL EXAMINED. Bergen Fjord, Norway, 30-80 m, 1878, two coloniesin spirit, coll. A. M. Norman, 1912.12.21.191. Espegrend, nr Bergen, Norway,30 m, 9 Aug. 1962, $ colony on microslide, coll. W. J. Rees, 1962.11.7.38 (Fig. 5).NW of Bulbjerg, Denmark, 57 12' N, 8 54' E, 16 m, 3 Oct. 1922 ('Dana' sta. 2865),part of colony in spirit, coll. R. V. 'Dana', Copenhagen Zoological Museum (mentionedKramp, 1935 : 50, as Halecium articulosum) . Off Durham, England, Jul. 1874, colony in spirit, coll. A. M. Norman,1912.12.21.186. Filey, Yorkshire, $ colony in spirit, coll. T. Hincks, 1899.5.1.164.Bridlington Bay, Yorkshire, colony in spirit, coll. Ministry of Agriculture, Fisheriesand Food, S.S. 'George Bligh', 1956.2.2.27 (Fig. 5). Sheerness, Kent, 10 Oct. 1892,one slide, coll. E. T. Browne, 1973.6.17.1. Weymouth Bay, Portland, Dorset, 20 m,several colonies in spirit, coll. R. Kirkpatrick, 1897.8.9.16. Berry Head, Brixham,Devon, 25 m, four colonies in spirit, coll. R. Kirkpatrick, 1893.8.7.3, 1897.8.9.16.Eddystone ground, off Plymouth, Devon, 15 Sep. 1897, several <$ colonies onChaetopterus tubes in spirit and one slide, coll. E. J. Allen, det. E. T. Browne,1941.3.20.310. Plymouth, Devon, 1898, $ colony in spirit, coll. E. T. Browne,1941.3.20.331. Isle of Man, 25 Mar. 1894, three colonies in spirit and one microslide(Fig. 5), coll. E. T. Browne, 1959.10.17.8. DESCRIPTION. Colony erect, imperfectly pinnate, shrubby in habit. Main stempolysiphonic. Side-branches polysiphonic basally, component coenosarcs graduallybranching off, final branches monosiphonic and usually flexuose ; internodes equal,usually longer than broad, nodes oblique or transverse. Hydrothecae alternate,each on short hydrophore at distal end of internode ; short, tubular, rim slightlyflared, even. Usually 1-3 hydrothecal renovations, but up to 10 reported (Vervoort, BRITISH LAFOEIDAE AND HALECIIDAE 393 1972). Gonothecae on short pedicels below hydrothecae. <$ club-shaped, apertureterminal ; $ approximately bean-shaped, aperture tubular, in centre of concaveside, c. 6 ova, i or 2 hydranths projecting through aperture.MEASUREMENTS. See Table 4. TABLE 4 Halecium beanii. Measurements in (jim unless otherwise stated HYDROTHECA Length Breadth at rimHYDROCAULUS DIAMETERLENGTH OF INTERNODESo* GONOTHECA Length Breadth (max.)$ GONOTHECA Length Breath (max.)MAXIMUM HEIGHT OF COLONY U.S.S.R. NEW ZEALAND SW ENGLAND SE ENGLAND(Naumov, 1969) (Ralph, 1958) (1959.9.17.3) (1973.6.17.1) 25-45110-140120-150500-530 IOOO-I2OO430-480 310 0-3 m VARIATION. Hydrocauli of this species are variably flexuose ; extremes areshown in Fig. 5. REPRODUCTIVE SEASON. Fertile specimens recorded from S Devon, Norfolk andRoscoff, France, most months from January to October (Hamond, 1957 ; MarineBiological Association, 1957 ; Teissier, 1965). DISTRIBUTION. Common throughout Britain and nearby seas. Apparentlyabsent from the Baltic and rare in the Kattegat, commoner in the Skagerrak andOslo Fjord (Broch, 1928 ; Christiansen, 1972). HABITAT. Rocky substrates ; 5-100 m depth, occasionally deeper. Halecium halecinum (Linnaeus, 1758)(Fig. 6) Corallina scruposa pennata, cauliculis crassiusculis rigidis Ray, 1724 : 36. Corallina erecta, tubulosa, pennata, halecis spinae facie Ellis, 1755 : 17-19, pi. 10. Sertularia halecina Linnaeus, 1758 : 809. Halecium halecinum: Oken, 1815:91; Johnston, 1847:58-59, pi. 8, figs 1-4; Hincks, 1868 : 221-223, pl- 4 2 . figs a-d ; Broch, 1918 : 36-38, fig. n ; Vervoort, 1946: 158-161, figs 6364; Leloup, 1952:140-141, fig. 74; Naumov, 1960:446-447, fig. 335, pi. 17, fig. 3 ; Naumov, 1969 : 482-483, fig. 335, pi. 17, fig. 3.Thoa halecina: Johnston, 1838 : 119-120, pi. 6, figs 1-4.Halecium geniculatum Norman, 1867 : 205 ; Hincks, 1868 : 229 ; [non Halecium geniculatum Nutting, 1899 (= H. tenellum Hincks, 1861 (Naumov, 1960, 1969) ; see also synonymy of H. tenellum, p. 409)]. P. F. S. CORNELIUS FIG. 6. Halecium halecinum. a-b, part of colony and $ gonotheca, Isle of Man( I 959-9-i7-5) ; c, cJ gonotheca, SW England (1898.5.7.86). Scale (a-c) = 500 fim. TYPE MATERIAL AND LOCALITY. Whitstable, Kent, England, on shell of Ostreaedulis Linnaeus, 1758 (Ellis, 1755 : pi. 10, figs A, a, B) ; location of specimensunknown. 8 MATERIAL EXAMINED. Oban, Argyll, 1877, three colonies in spirit, coll. A. M.Norman, 1912.12.21.198. Bridlington Bay, Yorkshire, 7 Nov. 1921, severalcolonies in spirit, coll. S.S. 'George Bligh', 1956.2.2.26. S edge of Smith's Knoll,off Norfolk, 19 Jan. 1905, part of $ colony on microslide, coll. M.V. 'Goldseeker',det. J. Ritchie, 1964.8.7.273 (mentioned, Rees & Thursfield, 1965 : 105). Isle ofMan, 5 Jul. 1894, part of colony on microslide, coll. E. T. Browne, 1959.9.17.5(Fig. 6). Menai Straits, Anglesey, Jul. 1964, part of <$ colony on microslide, coll.W. J. Rees, 1969.12.1.2. R. Deben estuary, Suffolk, several <$ colonies in spirit,coll. D. L. Serventy, 1933.7.1.12. East Solent, Hampshire, part of infertile colonyon microslide, coll. A. Dendy, 1886.10.4.6. Off Lulworth, Dorset, 7 m, fragments of 8 Linnaeus quoted the earlier designations of Ray (1724) and Ellis (1755) and probably saw no materialhimself since he did not give a locality. Although the collections of the Linnean Society of Londoninclude material labelled H. halecinum [catalogued 1298.7-8 by Savage (1945)], it was probably addedto the collection after 1758. Furthermore, none of it is the species currently recognized under this name.The specimens comprise two fertile colonies of H. beanii (sheet 7) and a colony of Sertularella sp.(sheet 8). Although some of Ray's and Ellis' material was later incorporated in the Hans Sloane herbarium,now in the British Museum (Natural History), there is no indication that the specimens of H. halecinum[H.S. 150 : 37 (R.H. 79) ; H.S. 114 : i (12)] in the Sloane herbarium are theirs since they have no collector'sname or locality. They cannot, therefore, be identified as type material. Some Ellis hydroid material, which might have included the originally illustrated specimens ofH. halecinum, was until recently preserved in the Hunterian Museum of the Royal College of Surgeonsof England. Unfortunately, the Ellis material was almost certainly destroyed by a bomb during theSecond World War and no Halecium material survives (see Cornelius, 1975: 267, footnote, for details). BRITISH LAFOEIDAE AND HALECIIDAE 395 $ colony on two microslides, coll. F. Beckford, 1889.7.27.7. Tor Bay, Devon, 1875,part of < colony on microslide, coll. A. M. Norman, 1898.5.7.86 (Fig. 6). StartPoint, Devon, 18 m, part of $ colony on microslide, coll. R. Kirkpatrick, 1893.8.7.4.Plymouth, Devon, 19 Dec. 1892, part of <$ colony on microslide, coll. E. T. Browne,1959.9.17.4. Manacle Rocks, Porthoustock, Lizard Peninsula, Cornwall, 18 m,29 Aug. 1965, part of $ colony on microslide, coll. R. Davis, pres. R. C. Vernon,1966.1.1.3. DESCRIPTION. Colony stiffly erect, regularly pinnate, with second and some thirdorder branching. Hydrocaulus and main branches polysiphonic basally, componentperisarc tubes branching off successively in groups ; final branches monosiphonic,straight or slightly flexuose. Internodes equal, with distal hydrophore. Hydro-thecae alternate, short, rim even, slightly flared, secondary, tertiary and somequaternary hydrothecae inside primary (Fig. 6) and longer than it. Hydranthlarger than hydrotheca, hypostome conical, 17-22 tentacles. <$ gonotheca club-shaped, tapering basally, rounded distally ; $ gonotheca oblong, tapering basally,subtruncate distally with lateral tubular aperture, 1-4 ova, with 1-3 hydranthsprojecting through aperture. Sexes on separate colonies. MEASUREMENTS. See Table 5. TABLE 5 Haleciiim halecinum. Measurements in unless otherwise stated HYDROTHECA Length (diaphragm to rim) Breadth at rim Hydrocaulus diameter (min.) Length of internodescj GONOTHECA Length Breadth (max.)$ GONOTHECA Length Breadth (max.)MAXIMUM HEIGHT OF COLONY ? U.S.S.R. NETHERLANDS SE ENGLAND SW ENGLAND (Naumov,1969) 90-150170-210 5000-25 m (Vervoort,1946) 1 00-200 800-1000c. 400 1000-1300 400-500 0-25 m (1964.8.7.273) (1889.7.27.7) 20-40140-165190-225350-380 1250-1400350-600 25-40 130-145100-150 280-400 700-800250-400 REPRODUCTIVE SEASON. Fertile specimens recorded December to July in EnglishChannel and off Norfolk (Hamond, 1957 ; Marine Biological Association, 1957 ;Teissier, 1965) ; one fertile $ specimen 29 August, S Cornwall (1966.1.1.3). DISTRIBUTION. Common throughout British Isles and adjacent continentalshelf areas. HABITAT. On stones, shells and other hard substrates, from sublittoral to edge ofcontinental shelf and sometimes deeper (Broch, 1918). Detached fragments fre-quently occur on strand-line. 396 P. F. S. CORNELIUS REMARKS. Halecium geniculatum Norman, 1867, appears from the originaldescription to be conspecific with H. halecinum. The long, tubular hydrothecaedescribed by Norman and regarded by him as diagnostic were probably just therenovated hydrophores which normally occur in H. halecinum. Halecium labrosum Alder, 1859(Fig. 7) ? Eudendrium pusillum Sars, 1857 : 154, pi. i, figs 14-16. Halecium labrosum Alder, 1859 : 354, pi. 13 ; Hincks, 1868 : 225-226, pi. 44, fig. i, text-fig. 27 ; Rees & Thursfield, 1965 : 107-108 (syn. H. crenatum Hincks, i874a) ; Naumov, 1969 : 489- 490, fig. 343 ; Calder, 1970 : 1506-1508, pi. i, figs 6-8.Halecium pusillum: Kossowska, 1911:347-350, figs 15-16; Broch, 1912:16-17, fig. 2; Bedot, 1916 : 115-116 ; [? non Teissier, 1965 : 21 ; see Remarks].Halecium annulatum Stechow, 1919 : 33 (nom. nov. pro H. pusillum: Kossowska).Halecium reflexum Stechow, 1919 : 37 -39, figs G-H ; Teissier, 1965 : 21.Halecium undulatum Billard, 1921 : 137-139, fig. 3 ; Leloup, 1952 : 144-145, fig. 79 ; Hamond, 1957 : 304-307, figs 12, 13 [syn. H. tenellum: Broch, 1918 (part) ; Kramp, 1929 (part) ; Kramp, 1932 (part) ; Kramp, 1938 (part)] ; Hamond, 1963 : 667 ; Calder, 1970 : 1510-1512, pi. 2, figs 7-9 ; (syn. nov.).Halecium tenellum : Fraser, 1944 : 201-203, pi. 37, fig. 179 (part) ; Leloup, 1952 : 144, fig. 77 (part).Halecium schneideri : Leloup, 1952 : 144, fig. 78 (? non H. schneideri Bonnevie, 1898). TYPE MATERIAL AND LOCALITIES. Syntypes : Large infertile colony on herbariumsheet and a fertile spirit specimen, probably male, in four pieces, both labelled'deep water, Northumberland coast', Hancock Museum, Newcastle-upon-Tyne.Other type localities : Moray Firth and Shetland (Alder, 1859). MATERIAL EXAMINED. Vatlestraumen channel, Hardanger Fjord, Norway,6o2o' N, 5i2' E, 15-25 m, 13 Apr. 1962, several colonies in spirit, three microslidesof fragments (2 $, i sterile), coll. W. J. Rees, 1962.10.7.13 (Fig. 7), 35, 66. NVadero Is., Skagerrak, 5835' N, n04' E, 80 m, i Oct. 1964, colony in spirit andone microslide, coll. W. J. Rees, 1965.1.14.135 (Fig. 7). Firth of Lorn, Argyll,Scotland, 120-140 m, part of colony on microslide, coll. J. Murray, 1888.6.9.6.Mull of Kintyre, Argyll, Scotland, 90 m, two ? colonies in spirit and two microslides,coll. J. Murray, 1888.1.24.19 (Fig. 7). Plymouth, Devon, England, 17 Aug. 1898,six fragments of $ colonies on microslide, coll. E. T. Browne, 1959.9.17.9. Eddy-stone ground, western English Channel, 15 Sep. 1897, several fragments of coloniesin spirit, coll. E. T. Browne, 1941.3.20.413. Eddystone ground, western EnglishChannel, 19 Nov. 1897, parts of colonies on two microslides, coll. E. T. Browne,1959.9.17.6, 123. Messina, Sicily, Italy, infertile colony on alga in spirit, holotypeof Halecium pusillum Sars, 1857, Oslo Zoological Museum Cat. No. 61156 (men-tioned, Broch, 1912 ; see Remarks, below). DESCRIPTION. Colony erect, up to c. 50 mm, imperfectly pinnate, all but finalbranches polysiphonic ; main branches slightly flexuose, side-branches insertedalternately with characteristic curve near base. Living tissues said to be purplish(Hincks, 1868). Internodes unequal, perisarc transversely wrinkled over much or BRITISH LAFOEIDAE AND HALECIIDAE 397 FIG. 7. Halecium labrosum. a, part of colony, Skagerrak, 80 m (1965.1.14.135) ; b, <$gonotheca, near Bergen, Norway, 15-25 m (1962.10.7.13) ; c, $ gonotheca, W Scotland,90 m (1888.1.24.19). Scale (a-c) = 500 (zm. all of length or, occasionally, smooth. Hydrothecae sessile, borne on prominenthydrophores, short, rims strongly recurved ; up to c. 12 renovations common. Gono-thecae on i- or 2-ringed pedicel ; gonothecal hydranths absent ; <$ ovate to linear,tapering basally and sometimes distally also, aperture simple, terminal ; $ ovate,larger than <, aperture simple, terminal ; sexes on separate colonies. MEASUREMENTS. See Table 6. REPRODUCTIVE SEASON. Incompletely known. Fertile specimens recorded earlyMay in southern North Sea, late May in Orkneys and Faroes, late June in Iceland(Hamond, 1957) ; 17 August 1898 at Plymouth (present material) ; September atRoscoff (Teissier, 1965). DISTRIBUTION. A northern species known from localities throughout Britishcoastal waters (Hincks, 1868 ; Hamond, 1957 ; Marine Biological Association, 1957).Although recorded from Roscoff, NW France (Teissier, 1965), the species appearsabsent from the Scilly Isles, Channel Islands, Belgium and Holland (Vervoort, 1946,1949 ; Leloup, 1952 ; Robins, 1969) and the English Channel is probably near itssouthern limit. The species is known from the Danish coast and the Skagerrak(Kramp, 1935) but has been reported absent from the Baltic (Broch, 1928). HYDROTHECA Length (diaphragm to rim) Breadth at rim Hydrocaulus diameter (min.) Length of internodeso* GONOTHECA Length Breadth (max.)$ GONOTHECA Length Breadth (max.) P. F. S. CORNELIUSTABLE 6 Halecium labrosum. Measurements in (Jim W ENGLISH ARGYLL, CHANNEL PLYMOUTH SCOTLAND (1959.9.17.123) (I959.9.I7-9) (1888.1.24.19) 30-50240-280 160-200950-1175 30-45150-165 90-110800-1700 640-1100470-530 40-50220-255 130-150550-700 1350-1580950-1090 W NORWAY(1962.10.7.13) 30-45(one 60) 140-155 (one 170) 90-110 600-820 700-1000300-900 HABITAT. Offshore, 5-200111 ; found epizoic on exoskeletons of various inver-tebrates (Hamond, 1957) and probably occurs also on inanimate substrates. REMARKS. The variation in transverse wrinkling of the internodal perisarc inthis species has caused systematic confusion. As noted by Hamond (1957), speci-mens with completely smooth internodes have probably been confused with H.tenellum Hincks, 1861, which is, however, smaller in all dimensions, seldom has awrinkled perisarc and lacks the curvature of the bases of the hydrocladia typical ofH. labrosum. Eudendrium pusillum Sars, 1857, type locality northern Sicily, remains problema-tical. It was recognized as Halecium by Kossowska (1911) who wrongly includedH. lankesteri in its synonymy. Broch (1912) redescribed and figured part of theholotype and regarded the species as valid. The type material, re-examined here,has a recurved hydrothecal rim, a feature noted by Broch but not by Sars, and thisand the curving bases of the branches are reminiscent of the present species. How-ever, identification is made difficult by the type material being sterile and stuntedin comparison with colonies of H. labrosum. It is not, therefore, possible to identifyit confidently with the present species, which in addition has not been recorded sofar south. Possibly the material from NW France identified as H. pusillum byTeissier (1965) was in fact H. labrosum as defined here ; his is apparently the onlysuggestion that H. pusillum occurs in the present area. Halecium reflexum Stechow, 1919, appears from the original description toresemble H. labrosum in its manner of branching, recurved hydrothecal rim andwrinkled perisarc, and the two taxa appear conspecific. Halecium undulatum Billard, 1921, seems to have been based merely on smallcolonies of H. labrosum. The two taxa appear similar in all characters exceptcolony size and their distributions coincide so that there seems no reason to keepthem distinct. Consequently the many H. tenellum records referred to H . undulatum BRITISH LAFOEDIAE AND HALECIIDAE 399 by Hamond (1957) are identified here as H. labrosum. The gonothecae describedby Calder (1970) under H. undulatum are similar to those of H, labrosum s. str. The material described by Leloup (1947, 1952) under the name H. schneideriBonnevie, 1898, is here referred to the present species. H. schneideri was a newname for H. nanum Alder, 1859, a species otherwise unrecorded from the presentarea. It was redescribed by Fraser (1944). Halecium lankesteri (Bourne, 1890)(Fig. 8) Halecium robustum Pieper, 1884 : 166-167 > Babic", 1913 : 470-473, figs 1-7 ; Bedot, 1914 : 82 ; Stechow, 1919 : 39-40 ; Teissier, 1965 : 21 ; Fey, 1969 : 397 ; [non Halecium robustum Verrill, J 873 : 9 ; (= Zygophylax sp. ; see p. 402)].Haloikema lankesterii Bourne, 1890 : 395-396, pi. 26.Halecium lankesteri : Bedot, 1911 : 213-217, pi. n, figs 1-5 (= H. sessile : Billard, 1904) ; Stechow, 1923 : 88 ; Prenant & Teissier, 1924 : 25 ; Broch, 1933 : 16-17, fig- 3 ' Vervoort, 1949 : 145 ; Hamond, 1957 : 302-304, figs 9-10 ; Marine Biological Association, 1957 : 4 6 : Vervoort, 1959 : 221-224, fig s 3~5 ' Millard, 1968 : 257-258, fig. i. NOMENCLATURE. The species was named after Professor E. R. Lankester. TYPE MATERIAL AND LOCALITY. Syntypes : Near Duke Rock Buoy, PlymouthSound, Devon, England, May 1889, several infertile colonies in spirit, coll. G. C.Bourne, 1974.2.28.1. Remaining type material recorded from Jennycliff Bay,Plymouth Sound, May 1890, not located. OTHER MATERIAL EXAMINED (All BM(NH) material except types). Studland Bay,Dorset, England, 6-8 m, $ colonies on six microslides, coll. R. Kirkpatrick, pres. F.Beckford via A. M. Norman, 1890.7.22.5 (Fig. 8). Mewstone, off Wembury, Devon,infertile colony on H. halecinum, in spirit, coll. E. T. Browne, 1948.10.1.125 (Fig. 8).Plymouth, Devon, 15 Jul. 1898, infertile fragments on microslide, coll. E. T. Browne,1959.9.17.1. Plymouth, Devon, 1892, several infertile colonies in spirit, coll. W.Garstang, pres. A. M. Norman, 1912.12.21.572. E of Duke Rock Buoy, Plymouth,on piece of wire, c. 10 m, 2 Jul. 1973, dense growth of <$ colony in spirit and hydro-caulus on microslide, coll. P. F. S. Cornelius, 1974.3.1.1 (Fig. 8 ; duplicate spiritmaterial deposited in the Museum of the Marine Biological Association, Plymouth). DESCRIPTION. Colony erect, up to c. 80 mm, usually unbranched but sometimesimperfectly pinnate ; monosiphonic. Internodes often irregular in length, o-io(usually i) annulations between hydrothecae ; bases of hydrocladia often curvedinwards sharply. Hydrotheca on prominent hydrophore at distal end of internode ;usually alternate ; slightly tapering basally, short ; rim not flared ; up to c. 5hydrothecal renovations. Hydranth large, 10-20 tentacles. Gonotheca on shortpedicel below a hydrotheca. $ cylindrical, aperture terminal ; $ kidney-shapedwith tubular aperture in centre of concave side, 3-20 ova, 1-2 protruding polyps.Vervoort (1959) records sexes on separate colonies, $ gonothecae 'exclusively onrenovated hydrophores' over entire colonies, $ on lower parts of colonies on secondaryor tertiary hydrophores. 400 P. F. S. CORNELIUS FIG. 8. Halecium lankesteri. a, part of colony, SW England (1948.10.1.125) ; b, $gonotheca, SW England, 6-8 m (1890.7.22.5) ; c, <$ gonotheca, SW England, c. 10 m(1974.3.1.1). Scale (a-c) = 500 [im. MEASUREMENTS. See Table 7. VARIATION. Hydrocladia may or may not appear to be sharply incurved basally,and Fig. 8 shows a specimen with both conditions. There may be o-io internodalannulations between hydrothecae, and successive internodes are not identical inshape or length. The degree of wrinkling of the perisarc is variable. Tendrildevelopment occurs (e.g. 1974.3.1.1) but is unusual. REPRODUCTIVE SEASON. June-July at Roscoff (Teissier, 1965) ; gonothecae ofboth sexes, Norfolk coast, 26 June 1952 (Hamond, 1957) ; $ gonothecae, Plymouth,Devon, 2 July 1973 (1974.3.1.1). Infertile material from Plymouth includes thetype specimens collected May 1889 and May 1890, and other material collected 15July 1898 (1959.9.17.1). DISTRIBUTION. There are published records from NW France, the ChannelIslands, SW England, SW Wales and Norfolk (Bourne, 1890 ; Vervoort, 1949 ; BRITISH LAFOEIDAE AND HALECIIDAETABLE 7 Halecium lankesteri. Measurements in [zm unless otherwise stated 401 HYDROTHECA Length (diaphragm to rim)Breadth at rim INTERNODESDiameter (min.)Length <J GONOTHECALengthBreadth (max.) $ GONOTHECA Length Breadth (max.)MAXIMUM HEIGHT OF COLONY SW ENGLAND (1974.2.28.1 ; SW ENGLAND(1974.3.1.1) 30-35130-140 90-110320-600 610-720200 type) 45-70200-230 150550-750 c. 3 mm c. 40 mm GUINEA (Vervoort, 1959) 20-25120-130 420-540 410-510200-240 450c. 8 mm MOZAMBIQUE (Millard, 1968) 30-40140-150 200-660 520-650380-450c. 5 mm Hamond, 1957 ; Teissier, 1965 ; Fey, 1969). The present records fall within thisdistribution and southern British waters appear to be the northern limit of thisspecies. It does not appear to have been recorded from Ireland. The distributionabroad was summarized by Stechow (1923), Broch (1933), Vervoort (1959) and Millard(1968). HABITAT. Recorded on stones (Bourne, 1890), other hydroids, crabs and Bryozoa(Hamond, 1957 ; Vervoort, 1959 ; present material), and on algae (Teissier, 1965 ;Fey, 1969). o-c. 50 m depth ; not recorded intertidally. REMARKS. The most useful descriptions of this species are those of Bourne(1890), Bedot (1911), Broch (1933), Vervoort (1949, 1959), Hamond (1957) andMillard (1968). The measurements of the type material given here appear high in comparisonwith those of the other measured material. There is, however, no doubt that thetype material is referable to the concept of the species both as first described byBourne (1890) and as understood by recent authors (Hamond, 1957 ; Vervoort,1959 ; Millard, 1968). The type specimens lack a flared hydrothecal rim whichmight otherwise identify them as H. labrosum Alder, 1959. They resemble closelythe illustrations of Bedot (1911). Although the hydranths of the type specimens were brown (Bourne, 1890),colourless and green hydranths have been recorded (Vervoort, 1949) and hydranthcolour cannot be regarded as a useful specific character. The confusion between Halecium lankesteri and the several species to which thecombination Halecium robustum has been applied was summarized by Hamond(1957). The earliest usage, H. robustum Verrill, 1873, referred to a species of 402 P. F. S. CORNELIUS FIG. 9. Part of neotype of Halecium robustum Verrill, 1873, Massachusetts (YalePeabody Museum No. 9137). Scale = 500 [zm. The specimen is here referred to thegenus Zygophylax. 9 Zygophylax so that the name robustum is not available for a Halecium species. 9 H.lankesteri (Bourne, 1890) is the next available name for the present species. Halecium muricatum (Ellis & Solander, 1786)(Fig. 10) Sertularia echinata Linnaeus, 1761 : 541 (part) ; Pallas, 1766 : 152 (part) ; Linnaeus, 1767 : 1310 (part).Sertularia muricata Ellis & Solander, 1786 : 59-60, pi. 7, figs 3-4. 9 The holotype of the nominal species Halecium robustum Verrill (1873 : 9; 786 m, St George's Bank,E Canada, 15 Sep. 1872, Yale Peabody Museum No. 3651) was never illustrated and is now lost (W. D.Hartman, pers. comm.) but a non-type specimen, labelled H. robustum in Yen-ill's hand, is extant andis here made neotype of H. robustum (off Cape Ann, Massachusetts, 17 Sep. 1878, branched hydrocauluson microsclide, YPM No. 9137). The specimen (Fig. 9) has the characters of Zygophylax sensu Totton(1930). Thus the specific name robustum is not available for a Halecium species; and the species describedby Verrill should be known as Zygophylax robustum. It predates other species of Zygophylax, but as specificlimits within that genus are at present unclear no new synonymy is proposed. However, it may benoted that in lacking nematothecae and in having campanulate, straight or only slightly curved hydro-thecae the neotype specimen resembles closely the original illustrations of Z. pinnata (Sars, 1874). BRITISH LAFOEIDAE AND HALECIIDAE 43 FIG. 10. Halecium muricatum. a, part of colony, E Scotland, 55 m (1969.12.5.9) ;b, $ gonotheca (identified from contents ; $ is externally similar), E Scotland, 20 rn(1964.8.7.87). Scale (a-b) = 500 pun. Halecium filiforme Alder, i862a : 315 (part) ; Alder, i862b : 236 (part) ; Hincks, 1868 : 228(part). Halecium muricatum : Hincks, 1868 : 223-224, pi. 43, figs i, la-b ; Broch, 1918 : 43-45, fig. 17 ;Eraser, 1944 : 197-198, pi. 37, fig. 176 ; Vervoort, 1946 : 163-164, fig. 67 ; Naumov, 1960 :45 6 -457. n g- 347 ; Naumov, 1969 : 492-493, fig. 347 ; Vervoort, 1972 : 27, fig. 3b-d. Halecium murigatum Williams, 1954 : 4& (lapsus pro muricatum). TYPE LOCALITY AND MATERIAL. The original description was based on materialsupplied by a Dr David Skene of Aberdeen, but no locality was given and the speci-men has not been located (see also footnote 8, p. 394). The type locality is hererestricted to near Aberdeen, Scotland. MATERIAL EXAMINED. Firth of Lorn, Argyll, Scotland, 130-150 m, part ofinfertile colony on microslide, coll. J. Ritchie, 1974.3.7.1. Off SE coast of Isle ofMull, Argyll, 10 Apr. 1971, two colonies in spirit, coll. P. F. S. Cornelius, 1971.5.11.30.Moray Firth, E Scotland, 55 m, 8 Dec. 1965, fertile fragments on two microslides,coll. F. W. E. Rowe, 1969.12.5.9 (Fig. 10). Burghead Bay, Moray Firth, 20 m,22 Nov. 1905, fertile fragments on microslide, coll. J. Ritchie, 1964.8.7.87 (Fig. 10 ;mentioned, Rees & Thursfield, 1965:109). Gullmarfjord, Sweden, 20-30 m,27 Aug. 1962, fragment on microslide, coll. W. J. Rees, 1962.11.8.15. Probably Wcoast of Sweden, about 1758-60, 30 mm portion of fertile colony, Linnaeus colln.,Linnean Society of London Catalogue No. 1298.14 (not type material ; see Remarks). 44 P. F. S. CORNELIUS DESCRIPTION. Colony erect, up to 100 mm, rarely 200 mm, imperfectly pinnate ;main stem and larger branches polysiphonic ; hydrocladia alternate. Nodesequally spaced, annulations often sloping alternately left and right but sometimestransverse. Hydrothecae alternate, on the monosiphonic branches, distal hydro-phore usually demarcated by deep, irregular annulus ; hydrotheca short, rim even,usually flared but sometimes straight ; commonly up to c. 6 renovations. Gono-thecae ovoid, pedicellate, with numerous 100-200 |j.m spines, in 9-15 rows. Sexesgenerally reported similar (see Remarks). MEASUREMENTS. See Table 8. TABLE 8 Halecium muricatum. Measurements in HYDROTHECA Length (diaphragm to rim) Breadth at rimINTERNODE Length Minimum breadthGONOTHECA (cJ = $) Length Breadth SE SCOTLAND SW SCOTLAND W SWEDEN (1964.8.7.87 ; Fig. 10) (1974.3.7.1) (1962.11.8.15) 30-80130-160 480-60080- 1 10 1000-1500900-1300 35-50140-170 600-800100-180 40-60240-280 850-1100130-150 REPRODUCTIVE SEASON. Little information. Fertile material recorded fromNorfolk, 16 Nov. 1960 (Hamond, 1963) and in the British Museum (Natural History)collections from the Moray Firth, Scotland, 22 Nov. 1905 (1964.8.7.87) and 8 Dec.1965 (1969.12.5.9). The apparent autumn breeding of this species in British watersis perhaps related to its northerly distribution. In Oslofjord specimens with gono-thecae have been recorded in December and February and also in June (Christiansen,1972) but it is not clear if the June material was living when collected. DISTRIBUTION. An arctic and northern boreal species, extending south to thecoasts of Scotland, NE England, Denmark and Sweden with occasional records inthe central North Sea, Dogger Bank and off the Norfolk coast (Hincks, 1868 ; Broch,1928 ; Kramp, 1935 ; Hamond, 1957, 1963 ; Rees & Rowe, 1969 ; Christiansen,1972). Other records include SW Scotland (Vervoort, 1946 ; present material),Isle of Man (Bruce et al., 1966), Co. Antrim and Co. Down, Northern Ireland (Hincks,1868 ; Stephens, 1905 ; Williams, 1954, in 1857). The species has not been includedin recent faunal surveys of the following areas : Roscoff (Teissier, 1965), Scilly Isles(Robins, 1969), Plymouth (Marine Biological Association, 1957), Pembrokeshire(Crothers, 1966) and Anglesey (K. Hiscock, pers. comm.) ; and further eastwards wasnot reported from Belgium (Leloup, 1952), Holland (Vervoort, 1946) and the coastof Germany (Broch, 1927). There is apparently no record from the English Channelsince Hincks (1868) recorded the species from Cornwall and South Devon, and norecord this century from Irish waters. BRITISH LAFOEIDAE AND HALECIIDAE 405 HABITAT. Mainly 10-200 m in Russian seas (Naumov, 1969), occurring down to1350 m in the North Atlantic (Broch, 1918) ; not intertidal. Recorded on bothhard substrates such as shells and rocks (Hincks, 1868) and on red algae (Christiansen,1972). REMARKS. The combination Sertularia echinata Linnaeus, 1761, was used inseveral works between 1761 and 1802 (Bedot, 1901), but the specific name wasapparently not used between then and 1910 (Bedot, 1905, 1910, 1912, 1916, 1918,1925) and has apparently not been used since. The taxon was tentatively referredto the present species by Johnston (1838 : 121 ; 1847 : 61) an d later by Vervoort(1946 : 163) but it appears that this action was only partly correct. The typematerial of S. echinata is preserved in the collections of the Linnean Society ofLondon, on herbarium sheet No. 1298.14 (Savage, 1945). The sheet bears two speci-mens. That on the left comprises two infertile colonies arising from a commonsubstrate, each referable to Sertularia cupressina Linnaeus, 1758. That on theright, however, is a fertile fragment 30 mm long referable to H. muricatum as definedabove. The original diagnosis of 5. echinata was probably made from both thespecimens, as it includes characters of both 5. cupressina ('denticulis opposite ob-tusiusculis') and of H. muricatum ('calycibus [gonotheca] obovatis compressisreticulatis muricatis'). The specimen on the left of the sheet is here nominated aslectotype, so that the name S. echinata Linnaeus, 1761, falls into the synonymy ofS. cupressina Linnaeus, 1758. The later descriptions of 5. echinata provided byPallas (1766) and Linnaeus (1767) perpetuated Linnaeus' (1761) confusion of thetwo species. The first available name for the present species is thus Sertulariamuricata Ellis & Solander, 1786. Halecium filiforme Alder, i862a, was described only briefly by Alder withoutillustration (i862a, b) and the description was repeated by Hincks (1868). Hincksconsidered the species closest to 'H. plumosum' ( = H. sessile ; see p. 409) but quotedAlder as later regarding H, filiforme as 'probably a mere variety or an immaturestate of some other species'. The original description agrees with 'H. plumosum'in the following characters : hydrocaulus 'very slender, flexible', monosiphonic,little branched ; length 115 mm (4^ inches) ; 'branchlets' arising from the sideof a hydrotheca ; and differs in that the hydrothecae are said to be 'ratherslender, tubular, with a slightly everted margin'. The type material of H. filiformein the Hancock Museum, Newcastle-upon-Tyne, is a mixed series comprisingspecimens referrable to both H. muricatum and H. sessile Norman, 1867. The her-barium specimen labelled 'Halecium filiforme, deep water, Northumberland' ishere designated lectotype of H. filiforme. It is a large infertile colony of H.muricatum, of which name H. filiforme thus becomes a junior synonym. TheNewcastle-upon-Tyne material will be more full} 7 described elsewhere (Cornelius &Garfath, in prep). Most authorities have considered the gonothecae of the two sexes identical(e.g. Hincks, 1868 ; Fraser, 1944 ; Vervoort, 1946), but Rees & Thursfield(1965 : 109) implied that there is a difference without providing details of how theydiffer. The weight of opinion at present suggests that there is no difference,however. 2.1 406 P. F. S. CORNELIUS Halecium sessile Norman, 1867(Fig. u) Halecium filiforme Alder, 18623. : 315 (part) ; Alder, i862b : 236 (part) ; Hincks, 1868 : 228 (part) ; (see p. 405).Halecium sessile Norman, 1867 : 205 ; Hincks, 1868 : 229-230, pi. 44, fig. 2 ; Storm, 1882 : 19 (? syn. H. plumosum) ; Billard, 1904:52-53, 157-160; pi. 3, figs 8-9, pi. 6, figs 1-14; Kuhn, 1913:139, fig. 58; Vervoort, 1941:195-196; Ralph, 1958:331-332, figs 9h-i, loc-d ; Teissier, 1965 : 21 ; Vervoort, 1966 : 100-102, fig. i (syn. H. kofoidi Torrey, 1902 ; H. lighti : Nutting, 1927) ; Christiansen, 1972 : 298.Halecium plumosum Hincks, 1868 : 227-228, pi. 64, fig. i ; Kuhn, 1913 : 70, 112, fig. 46 ; Christiansen, 1972 : 299 ; (syn. nov.).Halecium articulosom Clarke, 1875 : 63, pi. 10, fig. 6 ; Jaderholm, 1909 : 58, pi. 5, fig. 7 ; Kramp, 1935 : I 49- I 5 fig- 62c ; Kramp, 1938 : 32 ; Fraser, 1944 : 185-186, pi. 33, fig. 159 ; Leloup, 1952 : 143-144, fig. 76 ; Christiansen, 1972 : 299 ; (syn. nov.). TYPE MATERIAL AND LOCALITY. 'Deep water in the Minch', between OuterHebrides and mainland Scotland ; specimen not located. MATERIAL EXAMINED. 10 Gullmarfjord, W Sweden, c. 30 m, 13 Jan. 1910,colony in spirit, coll. T. Mortensen, Copenhagen Zoological Museum (CZM). Gull-marfjord, W Sweden, 15 Oct. 1918, colony in spirit with epizoic Halecium beanii,coll. Kristineberg Zoological Station, CZM. 22 km N of Hirtshals, Denmark, no m(Thor' sta. 1576), 27 Jun. 1911, colony in spirit, CZM. Nr Fredriskhavn, Denmark,17 m, 13 Aug. 1930, coll. P. L. Kramp, CZM. Oresund, Denmark, c. 25 m, 27 Aug.1928, CZM. Great Cumbrae Island, R. Clyde, W Scotland, 15-30 m, 18 May 1955, colony inspirit plus one microslide, coll. W. J. Rees, 1956.1.1.16. Salcombe, Devon, onNemertesia sp., infertile colony in spirit, coll. T. Hincks, 1899.5.1.167. Near Eddy-stone Rock, off SW coast of England, 10 Sep. 1897, infertile colony in spirit andfragments on microslide, coll. E. T. Browne, 1941.3.20.414. 3 km E of EddystoneRock, 6 Jul. 1971, two infertile colonies in spirit plus one microslide, coll. P. F. S.Cornelius, 1971.7.10.1. 4 km NNW of Eddystone Rock, c. 40 m, 20 Jul. 1972,infertile colony in spirit, coll. P. F. S. Cornelius, 1973.11.22.7. Bay of Biscay,7 Mar. 1912, fragments of J colony on microslide, coll. E. T. Browne, 1959.9.17.8.Off R. Sado, Portugal, infertile colony in spirit, coll. W. Saville Kent, 1872.2.3.137.South of Terceira Island, Azores, 610 m, Aug. 1959, infertile colony and fragmentson microslide, coll. Imperial College (University of London) Azores Expedition,1962. 1. 15. 12. 11 Bay of Fundy, New Brunswick, Canada, 1872, 30 mm infertilecolony in two parts, coll. U.S. Fisheries Commission (U.S.F.C.), Yale PeabodyMuseum (YPM) 3478. Casco Bay, Maine, U.S.A. 1873, five fragments of colonies(four infertile, one $), coll. U.S.F.C., YPM 5460 (syntypes of Halecium articulosumClarke, 1875). Eastport, Maine, U.S.A. 1868, eight fragments of colonies (fiveinfertile, three $), coll. A. E. Verrill, YPM 3663-4. Eastport, Maine, U.S.A. 1868,40 mm colony, coll. A. E. Verrill & S. I. Smith, YPM 3477. Jervis Bay, NewSouth Wales, Australia, c, 20 m, 1898, infertile fragment on microslide, coll. 10 The Danish and Swedish material is that mentioned by Kramp (1935 : 150) as Halecium articulosum. 11 The microslide specimen has an epizoic colony of Lafoeina sp., a genus not previously recorded fromthe Azores (Rees & White, 1966). BRITISH LAFOEIDAE AND HALECIIDAE 407 des FIG. ii. Halecium sessile, a-b, part of colony and one hydrotheca, SW England(1971.7.10.1) ; c, $ gonotheca, N France (redrawn after Billard, 1904 : pi. 6, fig. 14) ;d, <J gonotheca, Bay of Biscay (1959.9.17.8). Scale i (a, c, d) = 500 (Jim ; scale ii (b) =50 [Am. des. = desmocyte ('puncta'). H.M.S. Thetis', ex J. Ritchie collection, 1964.8.7.88 (mentioned, Rees & Thursfield,1965 : 109). DESCRIPTION. Colonies monosiphonic, up to 50 mm, or polysiphonic up to300 mm ; all sizes branching, the larger colonies branching irregularly, and beinglimp when out of water. Hydrocaulus variably flexuose, nodal constrictions trans-verse or sloping in alternate directions. Internodes approximately equal, becomingshorter distally, widely variable in length to breadth ratio between colonies ; hydro-phore distal, long, usually parallel with internode but sometimes shorter and slopingoutwards. Base of hydrotheca usually at approximately 90 to internodal axisalthough this angle may be greater ; hydrotheca very reduced, walls straight,divergent, rim even, not flared ; characteristic annulus of perisarc thickening belowbase of hydrothecal wall, wedge-shaped in optical section (Fig. ii). Hydrothecalrenovation frequent, secondary hydrothecae usually forming directly on diaphragmresulting in very short hydrothecal chains (Fig. ii). Hydrocladia inserted 408 P. F. S. CORNELIUS alternately, arising from sides of hydrophores, associated hydrotheca consequentlyaxillary. Hydranth long, 18-25 tentacles (Vervoort, 1941) ; citron-yellow colourrecorded (Storm, 1882) but this perhaps unusual. $ gonotheca tubular, slightlycurved, aperture terminal ; $ gonotheca kidney-shaped, with tubular aperture onconcave side, similar to that of H. beanii, with two projecting hydranths (Billard,1904 ; Ralph, 1958). MEASUREMENTS. See Table 9. TABLE 9 Halecium sessile. Measurements in [xm NEW BAY OF W ENGLISH NEW SOUTH ZEALAND BISCAY CHANNEL WALES AZORES (Ralph, (1959.9.17.8) (1971.7.10.1) (1964.8.7.88) (1962.1.15.12) 1958)HYDROTHECA Length (diaphragm to rim ) 25-35 45-55 4 -50 25-35 15-20* Breadth at rim 130-170 160-185 150-170 120-150 c. 125 INTERNODE Breadth (min.) 90-130 200 130-170 Length 550-700 550-650 340-500 500-680 o* GONOTHECA Length 980-1035 620-750 Breadth (max.) 185-220 210 $ GONOTHECA Length 750 Breadth (max.) 400 * Rim to desmocytes ('punctae'). REPRODUCTIVE SEASON. August to end October in NW France (Teissier, 1965 ;Fey, 1969) ; possibly also June -July, Cherbourg (Billard, 1904). (The fertile Bayof Biscay material is dated 7 March 1912 in E. T. Browne's hand, but this maysimply be the date on which he prepared the slide.) DISTRIBUTION. Widespread in the North Atlantic including the whole of theBritish Isles, North Sea and English Channel (Hincks, 1868 ; Broch, 1918, Kramp,1935) including the Skagerrak and Kattegat but not the Baltic (Broch, 1928 ;Kramp, 1935, as H. articulosum ; Christiansen, 1972). Records are neverthelessscattered. As H. sessile the species has been recorded from the Hebrides (Norman,1867 ; Hincks, 1868), Dorset and Devon (present material), Denmark (Kramp,1935) and NW France (Teissier, 1965 ; Fey, 1969) ; and as H. plumosum, probablyindicating records of larger colonies, from Aberdeen (Forbes, 1872), Firth of Forth(Leslie & Herdman, 1881), the Clyde (Chopin, 1894) and 'Ireland' (Hincks, 1868 ;Duerden, 1897). HABITAT. Apparently all depths to edge of continental shelf. Recorded onalgae and worm-tubes (Sabellaria sp.) (Billard, 1904). REMARKS. This species was last redescribed by Ralph (1958). BRITISH LAFOEIDAE AND HALECIIDAE 409 The species Halecium filiforme Alder, i862a, is discussed under H. muricatum (P- 405)- Halecium plumosum Hincks, 1868, was tentatively identified with the presentspecies by Storm (1882) ; Christiansen (1972) also commented on the similarity.The present material includes specimens referable to Hincks' description, but intheir mode of branching, structure of hydrotheca, manner of hydrothecal renovationand possession of an annular thickening below the diaphragm they resemble speci-mens referable to H. sessile sensu Norman, i.e. smaller colonies. It seems that H.plumosum was founded merely on large colonies of the present species and is hereregarded conspecific. Halecium articulosum Clarke, 1875, can be reduced to a synonym of the presentspecies. Of the type material only that from Casco Bay, Maine, could be located(W. D. Hartman, Yale Peabody Museum, pers. comm.). It agrees with H. sessilematerial in having very short chains of hydrothecae, in having the annular thickeningbelow each hydrotheca, in having a hydrotheca axillary to each side-branch, in theshape of the female gonotheca and in general colony habit, so that it appears thatthe two taxa are conspecific. Most of the material assigned to H. articulosum by Kramp (1935 : 150) was ex-amined and similarly found to be identical with H. sessile. Apparently the onlyother European records as H. articulosum are those of Jaderholm (1909), Vervoort(1942), Leloup (1952) and Christiansen (1972), respectively from W Sweden, NFrance, Belgium and Oslofjord. The Belgian material is well described, and isalmost certainly H. sessile. The W Swedish, French and Oslofjord material, identi-fied from the descriptions of Clarke (1875), Jaderholm (1909) and Kramp (1935),which are here referred to H. sessile, are also probably the present species. Halecium lighti Hargitt, 1924, was referred to the present species by Vervoort(1941) who later, however, accepted the species as valid (Vervoort, 1966). Nutting(1927) had previously doubted its validity. Halecium tenellum Hincks, 1861(Fig. 12) Halecium tenellum Hincks, 1861 : 252, pi. 6, figs 1-4 ; Hincks, 1868 : 226-227, pi. 45, figs i,za-c ; Broch, 1912 : 17-18, fig. 3 ; Broch, 1918 : 46-50, fig. 20 (part) ; Kramp, 1935 : I 45~146, fig. 6oA ; Eraser, 1944:201-203, pi. 37, fig. 179 (part); Vervoort, 1946:164-165,fig. 68 ; Leloup, 1952 : 144, fig. 77 (part) ; Hamond, 1957 : 307, fig. 14 ; Millard, 1957 : T 93>fig. 5 ; Vervoort, 1959 : 229-231, fig. 8 ; Naumov, 1960 : 454, fig. 344 (syn. H. geniculatumNutting, 1899) ; Millard, ig66a : 471, fig. n ; Vervoort, 1966 : 102, fig. 2 ; Naumov, 1969 :490-491, fig. 344 (syn. H. geniculatum Nutting, 1899). Halecium geniculatum Nutting, 1899 : 744-745, pi. 63, figs la-d ; [non H. geniculatum Norman,1867 = H. halecinum (Linnaeus, 1758)]. Halecium washingtoni Nutting, 1901 : 789 (nom. nov. pro H. geniculatum Nutting, 1899). TYPE MATERIAL AND LOCALITY. Syntypes : Salcombe Bay, Devon, England,cJ colony, on Cellaria fistulosa (Linnaeus, 1758) (Bryozoa), 12 in spirit plus threemicroslides, 1899.5.1.168 (Fig. 12). 12 Syn. Salicornaria farciminioides : Hincks, 1861 (International Commission on Zoological Nomen-clature, 1971: Opinion 949). 410 P. F. S. CORNELIUS FIG. 12. Halecium tenellum. a, part of colony, SW England (syntype, 1899.5.1.168) b, $ gonotheca, Republic of South Africa, 46 m (redrawn after Millard, ig66a : fig. HE) c, (J gonotheca, McMurdo Bay, Antarctica (1964.8.7.90). Scale (a-c) = 500 (Am. OTHER MATERIAL EXAMINED. 13 Hardanger Fjord, Norway, 200 m, fragments inspirit, coll. A. M. Norman, 1912.12.21.208. Oban, Argyll, Scotland, 1877, severalcolonies on Diphasia rosacea (Linnaeus, 1758), coll. A. M. Norman, 1912.12.21.209.Monte Brazil West, Terceira Island, Azores, 20 m, Aug. 1959, fragments of colonieson sertularian hydroid, coll. Imperial College (University of London) Azores Expedi-tion, 1962.1.15.24. Porto Santo Island, Madeira, 120 m, several colonies onEudendrium sp., in spirit and microslide, coll. R. Kirkpatrick, 1919.8.14.7. Simon'sBay, Simon's Town, Republic of South Africa, 'shallow water', colonies on fragmentsof type of Halecium dichotomum Allman, 1888, two microslides, coll. H.M.S. 'Chal-lenger', 1888.11.13.96. False Bay, Republic of South Africa, several colonies inspirit, pres. South African Museum, sample of material mentioned by Millard,1957 : 193, station FAL 29oC, BM(NH) 1957.4.26.18. Bay to east of Cape Royds,Antarctica, May 1908, 20-40 m, fragments on microslide, coll. Shackleton AntarcticExpedition, 1964.8.7.90 (mentioned, Rees & Thursfield, 1965 : no). Off cable,McMurdo Bay, Antarctica, 17 Feb. 1904, on Halecium arboreum Allman, 1888,several colonies in spirit, coll. National Antarctic ('Discovery') Expedition,1907.8.20.42 (mentioned, Hickson & Gravely, 1907 : 28). DESCRIPTION. Colony erect, up to 20 mm, delicate, monosiphonic, irregularlybranched. Hydrocaulus zig-zag in general appearance, internodes usually straight 13 This section includes all material of this species in the British Museum (Natural History) collectionexcept the type series. BRITISH LAFOEIDAE AND HALECIIDAE 411 and narrow, angle between them about 140. Usually 1-3 annular wrinkles ateach end of an internode, but some internodes wrinkled throughout, with inter-mediate conditions. Hydrotheca borne on prominent distal hydrophore, short,rim flared. <$ gonotheca pedicellate, ovoid, flattened in one plane ; $ similar, slightlylarger, no protruding hydranths (Millard, ig66a) ; sexes on separate colonies(Millard). MEASUREMENTS. See Table 10. TABLE 10 Halecium tenellum. Measurements in pun unless otherwise stated SOUTH AFRICA SOUTH AFRICA ? U.S.S.R. SW ENGLAND (Millard, HYDROTHECA Length Breadth at rimHYDROCAULUS DIAMETER (min.) LENGTH OF INTERNODES$ GONOTHECA Length Maximum breadth9 GONOTHECA Length Maximum breadthMAXIMUM HEIGHT OF COLONY (syntypes) 25-30100-130 50500-850 540-720200 c. 5 mm ig66a) (Vervoort,1966) 30-65120-135 875-1300 (Naumov,1969) 40-80120-190 1070 (max.)600 1070 (max.)5io4 mm 15-20 mm REPRODUCTIVE SEASON. Not recorded. DISTRIBUTION. Poorly known, as many published records are dubious (Hamond,1957 ; see Remarks below). British material in the British Museum (NaturalHistory) collection comes from only two localities, Salcombe, Devon (syntypes), andOban, Argyll. However, the wide distribution suggested by apparently valid pub-lished records implies that the species occurs throughout British and nearby waters. HABITAT. Frequently recorded on other thecate hydroids and on Bryozoa,occurring from sublittoral at least to 495 m (Vervoort, 1966). Recorded on Bryozoaby Hincks (1861, Cellaria fistulosa), Hamond (1963, Bugula plumosa) and Robins(1969, Cellaria sp.). Its apparent association with the hydroid Idiella pristis(Lamouroux, 1816) on the West African coast was considered by Vervoort (1959)simply to reflect substrate availability. REMARKS. The small size and straight, smooth internodes of this species serveto distinguish infertile specimens from young specimens of H . labrosum. In additionthe curving bases of the hydrocladia in H. labrosum contrast with the straighterbases in the present species. Published records should be treated with caution. Asummary of records in which the species was confused with H. labrosum and itsjunior synonym H. undulatum Billard, 1921, was given by Hamond (1957). Seealso the Remarks section under H. labrosum (p. 398). 4 i2 P. F. S. CORNELIUS Genus HYDRANTHEA Hincks, 1868 Atractylis Wright, 1858 : 447 (part) ; Hincks, 1862 : 461. Hydranthea Hincks, 1868 : 99 [nom. nov. pro Atractylis Wright (part)]. TYPE SPECIES. Hydranthea margarica (Hincks, 1862), by monotypy. DIAGNOSIS. Colony hydrorhizal ; hydrotheca pedicellate, borne singly, too shortto contain hydranths which are large ; gonophore eumedusoid ; large nematocystspresent. Hydranthea margarica (Hincks, 1862)(Fig. 13) Atractylis margarica Hincks, 1862 : 461, pi. 9, figs 4, 4a-e, x ; Hincks, 1863 : 45-46. Hydranthea margarica : Hincks, 1868 : 100-101, pi. 19, figs i, la-c ; Marine Biological Associa-tion, 1957 : 46; Huv6, 1954 : 178-182, pis 3-6 (= Halecium billardii Kossowska) ; White,1956 : 39-41 ; Millard & Bouillon, 1973 : 45-46, fig. 6a. Halecium margaricum : Kossowska, 1911 : 327-328, fig. i. Halecium billardii Kossowska, 1911 : 328-331, figs 2-3. Halecium billardii var. exigum Kossowska, 1911 : 331, fig. 4. TYPE MATERIAL AND LOCALITY. Ilfracombe, Devon, England, on Flustra foliacea(Bryozoa), c. 20 m (Hincks, 1863) ; probably collected summer, 1861 (Hincks,1862) ; present location unknown. MATERIAL EXAMINED. Off Sanda Island, Argyll, Scotland, 20-30 m, on Flustrasp. (Bryozoa), colony in spirit and two microslides, coll. R. B. Pike, 1955.11.15.11(mentioned, White, 1956). Off Oxwich Point, Gower Peninsula, Glamorgan, Wales,c. 20 m, 15 Jun. 1971, on Flustra sp., colony in spirit, coll. D. N. Huxtable, 1973.3.3.1.Off Poole, Dorset, England, c. 15 m, on Flustra sp., colony in spirit, coll. F. Beckford,1889. 7.27.5. Outside Swanage Bay, Dorset, c. 20 m, on Flustra sp., colony in spirit,coll. R. Kirkpatrick, 1897.8.9.2. Plymouth, Devon, England, 24 Feb. 1914, colonyin spirit, coll. E. T. Browne, 1954.8.3.42. Plymouth, 23 Oct. 1913, fertile colony onHalecium halecinum, in spirit and one microslide, coll. E. T. Browne, 1954.8.3.34.Stoke Point, Plymouth, 23 Oct. 1913, colony on fragments of gastropod shell, inspirit, coll. E. T. Browne, 1954.8.3.44. Roscoff, NW France, 12 Sep. 1956, colonyon bivalve shell, in spirit, coll. W. J. Rees, 1956.10.24.1-2. DESCRIPTION. Colony reptant, stolon bearing single hydranths at irregularintervals. Hydranths yellowish-white (Hincks, 1868), on short unringed pedicels,narrow at base, widest just below hypostome, 20-30 tentacles held alternatelyelevated and depressed (Hincks, 1868 ; Browne, in White, 1956), with small basalweb. Hydrotheca short, walls slightly divergent, rim even, diaphragm domeddownwards ; c. 20 desmocytes ('punctae'). Large, conspicuous banana-shaped orstraight nematocysts (microbasic eury teles, Huve, 1954), 30-40 (xm long, throughouttissues of hydranth and coenosarc, scattered or in clumps, particularly conspicuouson tentacular web. Gonophores on short pedicel tapering basally, attached tostolon ; gonotheca short (Fig. 13) or absent ; pedicel base said to be enclosed in smallchitinous cup (Hincks, 1868) but this not present in all specimens (e.g. BM(NH) BRITISH LAFOEIDAE AND HALECIIDAE 413 FIG. 13. Hydranthea margarica. a, reconstruction of hydranth and hydrotheca, WScotland, 20-30 m (1955.11.15.11); b, developing gonophore, two-cell stage, SWEngland (1954.8.3.43) ; c, mature $ eumedusoid, E Mediterranean, lower shore (redrawnafter Kossowska, 1911 : fig. 2, as H. billardii) ; d, undischarged nematocysts, SWEngland (1954.8.3.43). Scales in a-c = 500 [im ; scale in d = 25 (zm. 1954.8.3.43). Fully developed <$ and ? gonophores similar, inverted, medusiform,retained ; four branched gastrovascular canals said to be orange in colour.gametes develop in gonads borne on radial canals ; mature $ gonophore containsc. 300 ova. (Description of gonophores based on Hincks, 1868, Kossowska, 1911,and Huve, 1954.) Developing eggs reported in hydrorhiza in a colony apparentlylacking gonophores (Millard & Bouillon, 1973). MEASUREMENTS. See Table n. TABLE n Hydranthea margarica. Measurements in fim STOLON DiameterHYDRANTH Height (base to hypostome) Tentacle lengthHYDROTHECA Height (diaphragm to rim) Breadth at base Breadth at rim Hydrothecal pedicelNEMATOCYSTS Length * Contracted during fixation. S WALES(IQ73-3-3-I) 60200-400* 100 100100 W SCOTLAND (White, 1956 ; S FRANCE = 1955.11.15.11) (Huv6, 1954) 60-80 400-1000100-170 80-9080-90 40 90 700 180 3580 IOO 15030 REPRODUCTIVE SEASON. Fertile specimens recorded August (Roscoff, NW France,Teissier, 1965) and October (Plymouth, SW England, White, 1956). 24 4 i4 P. F. S. CORNELIUS DISTRIBUTION. Recorded from Devon and Dorset, Liverpool Bay, Norfolk andArgyll (White, 1956), mainland Shetland (D. N. Huxtable, pers. comm.), Anglesey(Herdman, 1891), S Wales (present material) and Roscoff, NW France (Teissier,1965). Elsewhere recorded from the Mediterranean Sea (Kossowska, 1911 ; Huve,1954) and the Seychelles (Millard & Bouillon, 1973). HABITAT. Most frequently recorded epizoic on Flustra spp. and other Bryozoa,but also found on hydroids (Tubularia sp. and Halecium spp.), mollusc shells andbare rock [Hincks, 1868 ; Kossowska, 1911 ; Huve, 1954 ; White, 1956 ; Teissier,1965 ; present material (1954.8.3.44.)] ; also on Laminaria holdfasts (D. N.Huxtable, pers. comm.). Genus OPHIODISSA Stechow, 1919 Ophiodes Hincks, 1866 : 421 [non Ophiodes Wagler, 1830 = Reptilia ; nee Ophiodes Guenee,1841 = Lepidoptera ; nee Ophiodes Hartig, 1847 = Hymenoptera ; nee Ophiodes Murray,1877 = Arachnida (Marschall, 1873 : 266 ; Stechow, 1919 : 41 ; Neave, 1940 : 434)]. Ophiodissa Stechow, 1919 : 41 (part) (nom. nov. pro Ophiodes Hincks). Hydrodendron : Millard, 1957 ' J 86 (part). TYPE SPECIES. Ophiodes mirabilis Hincks, 1866, by monotypy. DIAGNOSIS. Haleciidae with long capitate nematophore and short nematotheca ;hydranth large ; hydrotheca wider than deep, rim even ; desmocytes conspicuous ;gonotheca barrel-shaped, variably rugose, aperture broad, < = <; no medusageneration ; large nematocysts present. REMARKS. This generic diagnosis agrees with the restriction of Vervoort (1959,1972) in excluding species lacking a nematotheca. Following Vervoort, the onespecies concerned is retained in the genus Hydrodendron Hincks, i874a ; but asnoted by Watson (1969) a review of Ophiodissa and related genera is needed andpresent generic limits are somewhat arbitrary. Ophiodissa mirabilis (Hincks, 1866)(Fig. 14) Ophiodes mirabilis Hincks, 1866 : 422-423, pi. 14, figs 1-5 ; Hincks, 1868 : 231-233, pi. 45, fig. 2 ; Teissier, 1965 : 20.Ophiodes caciniformis Ritchie, 1907 : 500-501, pi. 23, figs 11-12, pi. 24, fig. i, pi. 25, fig. 5 ; Babi6, 1913 : 473, fig. 7 ; (syn. nov.). Ophiodissa mirabilis : Stechow, 1919 : 42 ; Vervoort, 1959 : 220.Ophidissa caciniformis : Stechow, 1919 : 42 ; Vervoort, 1959 : 218-221.Hydrodendron caciniformis : Millard, 1957 : 186-187, n g- 3 ' Ralph, 1958 : 342-344, figs i3b-c, i4a ; Millard, ig66b : 490-491, fig. i. TYPE MATERIAL AND LOCALITY. Holotype i 'on weed', 10-15 m, autumn, 1866,Swanage Bay, Dorset, England ; location of material unknown. Neotype : Lowerledges of Capstone, Ilfracombe, Devon, England, lower shore, 1867, infertile colonyon four fragments of holdfast of Laminaria sp., 1899.5.1.169. The neotype materialis labelled 'Ophiodes mirabilis, Ilfracombe, 1867' in Hincks' hand, and is almostcertainly the material he cited later (Hincks, 1868 : 233). BRITISH LAFOEIDAE AND HALECIIDAE 415 FIG. 14. Ophiodissa mirabilis. a, part of colony with hydranth and nematophore, CapeVerde Islands, 20 m (paralectotype of O. caciniformis, 1964.8.7.92) ; b, gonotheca,probably SW England (redrawn after Hincks, 1868 : pi. 45, fig. 2d). Scale (a) = 500 [/.m ;(b) unknown. OTHER MATERIAL EXAMINED. Aran Isles, Galway Bay, Eire, 8 Mar. 1899, colonieson bryozoan, in spirit, coll. E. T. Browne, 1954. 8. 3. 40-41. 14 Freshwater West,near Castlemartin, Pembrokeshire, Wales, lower shore, 15 Sep. 1974, young colonyon Dynamena pumila (Linnaeus, 1758), microslide, coll. P. F. S. Cornelius,1974.12.17.1. Porto Praya, Santiago, Cape Verde Islands, 20 m, 12 Aug. 1904, twofragments on microslide, coll. J. Ritchie, 1964.8.7.92, paralectotype of 0. caciniformis(mentioned, Rees & Thursfield, 1965). False Bay, Republic of South Africa,infertile colony on alga, in spirit, coll. N. A. H. Millard, 1957.4.26.23 (mentioned,Millard, 1957, as 0. caciniformis). Dalebrook, False Bay, Republic of South Africa,LWST, 16 Jan. 1961, infertile colony on brown alga, in spirit, coll. N. A. H. Millard,1961.6.26.4. DESCRIPTION. Established colonies irregularly pinnate, loosely polysiphonicbasally, up to c. 50 mm but usually less than 20 mm ; younger colonies stoloniferous.Hydrorhiza irregularly branched, perisarc thick, with internally-projecting spines ;hydrocauli arising erratically. Internodes smooth to slightly wrinkled, hydrophoredistal, sloping outwards at c. 45. Greatest width of hydrotheca c. i| times depth ;rim even, reflexed ; desmocytes conspicuous ; annular thickening between desmo-cytes and diaphragm. New hydrothecae arise from side of hydrophore (Millard, 14 The material bears the locality 'Aran'. Browne's ms diaries in the British Museum (Natural History)show that on the date given his collectors were in the Aran Isles, Galway Bay, and not the Isle of Aranin Donegal. 416 P. F. S. CORNELIUS o on co o ro IO 00 o o S5 W ai 3 9 en ^ <o-2 S Q ^^ ffi BRITISH LAFOEIDAE AND HALECIIDAE 417 1957 ; Ralph, 1958). Hydranth large, tapering basally, 18-24 tentacles, heldalternately raised and lowered. Nematothecae borne irregularly on internodes,hydrophores and stolon ; curved cone-shaped, narrowest basally, sometimes withconstriction below rim, both varieties occurring on single colonies ; nematophorelong, axial cells large, terminal knob with numerous nematocysts including largemicrobasic mastigophores. Gonothecae apparently recorded only twice. Describedas 'ovate, ringed transversely with wide tubular aperture, subpedicellate, borne onstolon', probably <$ (Hincks, 1866, 1868) ; similar, but aperture wider, less rugose,$ slightly smaller than $, no medusa (Millard, i966b). MEASUREMENTS. See Table 12. DISTRIBUTION. Recorded infrequently. From the British Isles, known fromIlfracombe, Devon ; Swanage, Dorset (Hincks, 1868 ; Garstang, 1900) ; Pembroke-shire (Crothers, 1966 ; present material) and Aran Isles, Galway Bay (presentmaterial). A record from Aberdeen, Scotland (Forbes, 1872), was regarded asdubious by its author. Apparently the only other European record is from Roscoff,NW France (Teissier, 1965). Other records, as 0. caciniformis, include the CapeVerde Islands (Ritchie, 1907), South Africa (Millard, 1957), SW Indian Ocean(Millard, I966b), New Zealand (Ralph, 1958), mid South Atlantic (Vervoort, 1959)and the West Indies (Vervoort, 1968). HABITAT. Most frequently recorded on algae, particularly laminarian holdfasts,but also on other thecate hydroids. Recorded from Laminaria zone of intertidal(Hincks, 1868 ; present material) down to 65 m (Vervoort, 1959). REMARKS. Ophiodissa caciniformis (Ritchie, 1907) is here reduced to a synonymof the present species. 0. caciniformis was proposed to include colonies larger thanthose previously referred to 0. mirabilis by Hincks (1868). 0. mirabilis has notpreviously been redescribed ; and the larger specimens described by various authorsthis century have been referred to 0. caciniformis. Characters ascribed to 0.caciniformis since the original description include an annular thickening of the hydro-thecal perisarc below the ring of desmocytes, a goblet-shaped nematotheca andinternal projections of the hydrorhizal perisarc (Millard, 1957, 1967 ; Ralph, 1958 ;Vervoort, 1959), all of which characters are present in the neotype material of 0.mirabilis. Further, the paralectotype specimen of 0. caciniformis has all thecharacters of the older species, and the dimensions of the type specimens of the twotaxa are similar. It seems that 0. mirabilis was founded on small colonies, and thatlarger colonies found later were wrongly given specific status. ACKNOWLEDGEMENTS I am grateful to Dr C. Edwards, Scottish Marine Biological Association, for generaladvice on hydroid systematics and to the following for kindly providing unpublishedlocality records and for loaning material : Dr M. E. Christiansen, University of OsloZoological Museum ; Dr W. D. Hartman , Yale Peabody Museum ; K. Hiscock,University College of North Wales, Bangor ; the late D. N. Huxtable, UniversityCollege of South Wales, Swansea ; Dr A. G. Long, Hancock Museum, Newcastle-upon-Tyne ; Dr D. V. Naumov, Zoological Institute, Academy of Sciences, 4i8 P. F. S. CORNELIUS Leningrad and Dr K. W. Petersen, University Zoological Museum, Copenhagen. DrN. A. H. 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CORNELIUS beanii, Halecium 391-393, 394 beanii, Thoa 391 billardii, Halecium 412, 413 boreale, Halecium 391 brevicyatha, Grammaria abietina var. 382, 385 caciniformis, Hydrodendron 414 caciniformis, Ophiodes 414 caciniformis, Ophiodissa 414, 415, 417 Calicella 390 C. syringa 390 Campalecium 390 C. medusiferum 391 Campanularia 378 C. abietina 381, 382 C. dumosa 385 C. fruticosa 386 C. gracillima 386-387 C. parvula 386, 390 C. serpens 378 CAMPANULARIIDAE 377 Capsularia 378 C. dumosa 386 C. serpens 378 cornuta, Lafoea 385, 386 Coryne 378 crenatum, Halecium 396 cupressina, Sertularia 380, 405 dichotomum, Halecium 410Diphasia rosacea 410dumosa, Campanularia 385dumosa, Capsularia 386dumosa, Lafoea 385-391dumosa, Sertularia 385Dynamena pumila 415 echinata, Sertularia 402, 405Eudendrium 410 E. pusillum 396, 398 EULAFOEINAE 377, 378 exigum, Halecium billardii var. 412 falcata, Hydrallmania 380fenestrata, Thuiaria 381Filellum 378, 381 F. serpens 379 filiforme, Halecium 403, 405, 406, 409fruticosa, Campanularia 386fruticosa, Lafoea 386, 389 geniculatum, Halecium 393, 396, 409gracillima, Campanularia 386-387gracillima, Lafoea 386, 389 Grammaria 378, 381 Grammaria 381-382 G. abietina 382-385 G. abietina var. brevicyatha 382, 385 G. insignis 382, 383, 384 G. intermedia 382, 385 G. magellanica 382, 383, 384 G. ramosa 382, 384 G. robusta 381, 382 G. serpens 379 G. stentor 382, 383, 384-385 HALECIIDAE 390-391 halecina, Sertularia 391, 393 halecina, Thoa 393 halecinum, Halecium 393-396, 399 Halecium 390-391, 414 H. annulatum 396 H. arboreum 410 H. articulosum 392, 406, 408, 409 H. beanii 391-393, 394 //. billardii 412, 413 //. billardii var. exigum 412 H. boreale 391 H. crenatum 396 H. dichotomum 410 //. filiforme 403, 405, 406, 409 //. geniculatum 393, 596, 409 H. halecinum 393-396, 399 //. kofoidi 406 H. labrosum 396-399, 401, 411 H. lankesteri 398, 399-402 H. lighti 406, 409 H. margaricum 412 H. muricatum 402-405 H. murigatum 403 H. nanum 399 H. plumosum 405, 406, 409 H. pusillum 396, 398 H. reflexum 396, 398 //. robustum 399, 401402 H. schneideri 396, 399 //. scutum 391 H. sessile 405, 406-409 H. sessile 399 H. tenellum 393, 409-411 H. tenellum 396, 398 //. torreyi 391 /f. undulatum 396, 398, 411 //. washingtoni 409 Haloikema 391 H. lankesteri 399 Hebella 377 H. pocillum 386, 390 INDEX 425 HEBELLIDAE 377Hydrallmania falcata 380Hydranthea 390, 412H. margarica 412-414Hydrodendron 414H. caciniformis 414 Idiella pristis 411immersa, Reticularia 378, 380insignis, Grammaria 382, 383, 384intermedia, Grammaria 382, 385 Kirchenpaueria pinnata 379kofoidi, Halecium 406 labrosum, Halecium 596-599, 401, 411 Lafoea 377, 385 L. abietina 379, 381 L. cornuta 385, 386 L. dumosa 385~3gi L. fruticosa 386, 389 L. fruticosa var. pocillum 386 L. gracillima 386, 389 L. parvula 390 L. pocillum 386, 390 L. pygmaea 390 LAFOEIDAE 377 Lafoeina 406 (footnote) lankesteri, Halecium 398, 599-402 lankesterii, Haloikema 399 Lictorella 378 LlCTORELLINAE 378 lighti, Halecium 406, 409 magellanica, Grammaria 382, 383, 384margarica, Atractylis 412margarica, Hydranthea 412-414margaricum, Halecium 412medusiferum, Campalecium 391mirabilis, Ophiodes 414mirabilis, Ophiodissa 414-417muricata, Sertularia 402, 405muricatum, Halecium 402-405murigatum, Halecium 403 nanum, Halecium 399Nemertesia 406N. ramosa 379 operculata, Amphisbetia 379Ophiodes 414 O. caciniformis 414, 415, 417O. mirabilis 414Ophiodissa 390, 414 O. caciniformis 414, 415, 417O. mirabilis 414-417 parvula, Campanularia 386, 390parvula, Lafoea 390pinnata, Kirchenpaueria 379pinnata, Zygophylax 402plumosum, Halecium 405, 406, 409pocillum, Hebella 386, 390pocillum, Lafoea 386, 390pocillum, Lafoea fruticosa forma 386pristis, Idiella 411pumila, Dynamena 415pusillum, Eudendrium 396, 398pusillum, Halecium 396, 398pygmaea, Lafoea 390 ramosa, Grammaria 382, 384ramosa, Nemertesia 379reflexum, Halecium 396, 398Reticularia [BRACHIOPODA] 378Reticularia [HYDROZOA] 378, 381R. abietina 382R. immersa 378-379, 380R. serpens 379robusta, Grammaria 381, 382robustum, Halecium 401-402robustum, Zygophylax 402rosacea, Diphasia 410 Salacia 381 Salacia 381 S. abietina 382 S. tetracythara 381 S. tetracyttara 381 Scandia 377 schneideri, Halecium 396, 399 scutum, Halecium 391 serpens, Campanularia 378 serpens, Capsularia 378 serpens, Filellum 378-381 serpens, Grammaria 379 serpens, Reticularia 379 Sertularia 381 Sertularia 385 S. cupressina 380, 405 S. dumosa 385 S. echinata 402, 405 S. halecina 391, 393 5. muricata 402, 405 sessile, Halecium 399, 406-409 sessile, Halecium 405 stentor, Grammaria 382, 383, 384-385 syringa, Calicella 390 426 tenellum, Halecium 396tenellum, Halecium 393, 398,tetracythara, Salacia 381tetracythara, Thuiaria 381tetracyttara, Thuiaria 381Thoa 391T. beanii 391T. halecina 393Thuiaria 381T. fenestrata 381T. tetracythara 381 P. F. S. CORNELIUS T. tetracyttara 381torreyi, Halecium 391Tubularia 414 undulatum, Halecium 396, 398, 411washingtoni, Halecium 409 Zygophylax 399, 402Z. pinnata 402Z. robustum 402 Dr P. F. S. CORNELIUS 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.) 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. & HAL-, 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 byfohn Wright and Sons Ltd. at The Stonebridge Press, Bristol 884 yNU