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 the collections 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 in the 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 within the 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 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 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, 380 abietina, Campanularia . . . .381 abietina, Grammaria .381, 382, 383, 384, 385 abietina, Lafoea .... 379, 381 abietina, Reticularia .... 382 abietina, Salacia . . . . .382 Abietinaria .... 379, 380, 386 abyssalis, Spinocalanus . 297, 298-300, 304 abyssalis pygmaeus, Spinocalanus . . 298 acuta, Charybdis .... 146, 147 adelungi, Obelia ..... 266 adonis, Hyloniscus . . . .71 adriatica, Halophiloscia .... 82 adriatica rupium, Halophiloscia . . 82 aediculatus, Euplotes . 6, 8, 16, 17-i8, 26, 28 Aetideopsis ..... 305, 306 Aetideus ..... 304-305 Aetiteidae ..... 304-311 affinis, Euplotes . . . 5, 16, 18-19, 24 affinis, Philoscia . . . . .87 affinis, Philoscia muscorum ... 87 affinis tricciratus, Euplotes . . .18 agilis, Xanthocalanus . . . 311, 312 alata, Metridia . . . 342, 343, 345 alatus, Euplotes . . . . 16, 19, 20 alatus, Euplotes patella . . . -41 alauda, Ennea .... 158, 160 alba, Pherusa ..... 73 albicincta, Bilawrencia ... 80 albus, Titanethes ..... 73 aldabrae, Buliminus . . . 171 aldabrae, Gulella gwendolinae 158-164, I7I» 172, 173 aldabrae, 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, 336 alternata, Eucope . . . . .273 alternata, Obelia ..... 267 Amallophora . 330, 332-333, 336, 339, 341 Amallothrix . 318, 320, 325-335, 341, 365, 367 amieti, Euplotes . . . -17, 19-21 Amphisbetia . . . . -379 Amphiura ..... 188-189 Anasterias . . . . . .190 Anchiphiloscia ..... 79 andersoni, Obelia . . . 260, 264, 265 Androniscus . angulosa, Obelia . angusta, Niambia . angusticauda, Philoscia . angusticauda, Setaphora angusticeps, Haloptilus . angusticeps, Spinocalanus angustissima, Pseudophiloscia annulata, Charybdis annulata, Eucope . annulatum, Helecium annulipes, Uca anomala, Philoscia anomalus, Phalloniscus antarcticus, Euplotes antennarius, Cancer anthonyi, Cancer . antiquorum, Equus burchelli . 70 . 266 77 91 91 • 345, 347 297, 298, 3°3, 3°4 . 89 146, 147 . 280 • 396 • 245 . 87 . 87 . 16, 21-22 • 239 238, 239, 246 224 Aphiloscia . . . . -79, 88, 98 apsheronicus, Euplotes . . . 38-39 apterycis, Ascaris . . . . .199 apterycis, Cyrnea . 201-205, Pis. 1-2 apuanus, Haplophthalmus . . .71 Arborcinia . . . . . .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, 409 Ascaris .... 199 aspinosa, Scolecithricella . 321, 322, 323 Assiminea . . . . . .166 Asteroidea . . . . . .190 Atelecylus 240 atlanticus, Euaugaptilus . . 349-351 Atoxon 207, 215 Atractylis 412 attenuata, Obelia . . 260 Augaptilidae . . . 346-36 1 aurantonotus, Centropyge . . 227, 230 auropecten, Amallothrix. 33O-333, 334~336 auropecten, Scolecithricella (Amallothrix) 330, 333 auropecten, Scolecithrix . . .321, 333 australis, Obelia . . . 266, 271, 272 australis, Styloniscus .... 68 428 INDEX australis, Trichoniscus austriacus, Trichoniscus . austroafricanus, Styloniscus austroafricanus, Trichoniscus austrogeorgiae, Laomedea austrogeorgiae, Obelia 68 73 . . . .68 68 . 280 260, 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, 264 260 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 . . .412 Bilawrencia ...... 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 ... 84 Brachyura . . 101-120, i PI., 237-247 brasiliensis, Campanularia . . . 265 brasiliensis, Obelia .... 267 braziliensis, Obelia. .... 267 brembana, Tendosphaera ... 76 brentanus, Androniscus .... 70 breuili, Iberoniscus .... 72 brevicaudatus, Spinocalanus 297, 298, 299, 300 brevicornis, Pseudophiloscia ... 89 brevicornis, Valdiviella . . 309, 310, 311 brevicyatha, Grammaria abietina . 382, 385 brevis, Alloniscus ..... 78 briani, Philoscia squamuligera . . 93 briani, Tiroloscia squamuligera . . 93 bruggeni, 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 . . . 224 burchelli burchelli, Equus . . .224 Bureschia ...... 70 Burmoniscus ..... 80 caciniformis, Hydrodendron . . .414 caciniformis, Ophiodes . . . .414 caciniformis, Ophidissa . . . .414 caciniformis, Ophiodissa . . 415, 417 calcivagus, Androniscus .... 70 calcuttensis, Quickia . . 180, 183, 184 Calicella 390 callianassa, Charybdis . . . 146, 147 Callinectes . . . . . 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, 150 caribbeanensis, Centropages . . 344~345 carinata, Aetideopsis . . . 305, 306 carniolense, Lepidoniscus germanicus . 84 carynthiacus, Androniscus ... 70 cassivelaunus, Corystes. . . 103, 239 castellana, Campanularia . . . 280 castellata, 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 • 78 265, 272 • 234 70 70 73 265, 272, 273 . 265 409, 411 81 81 . 344 . 344 204 227-231, i PI. 68 68 230 . 81, 88, 98 17, 18, 22, 24, 49 146, 147, 148 71 . 266 306-307 191-197, 233-235 . 1-61 74 91 91 . 69 29 187-igo 66-67 Clytia 253, 254, 260, 264, 265, 279, 280, 281 coeca, Philoscia . . . . .91 coeca, Setaphora . . . . .91 Coelenterata .... 249-294, 375 commensalis, Trichoniscus commissuralis, Obelia comorensis, Gulella compar, Alloniscus compta, Philoscia . comta, Setaphora . concisa, Quickia concisa, Succinea . congdoni, Laomedea congdoni, Obelia . contractus, Hiatoniscus Copepoda Coptodon Corallina Cordioniscus . Cornelius, P. F. S. . cornuta, Lafoea cornutus, Alloniscus corona, Obelia Corsica, Philoscia . 73 265, 271, 272 169 . 78 91 91 180, 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 . . 72 corsicus, Trichoniscus .... 72 corsicus corsicus, Nesiotoniscus . . 72 coruscans, Campanularia . . .278 Corystes ..... 103, 239 costata, Bathytropa .... 79 costata, Bathytropa meinerti ... 79 Costigulella . . . . . .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, 245 cristatus, Euplotes . . n, 12-13, 34 Crustacea 63-101, 101-120, i PL, 121-151, 237-247, 295-372 cubensis, Rhyscotoides .... 76 cubensis, Rhyscotus .... 76 Cucumaria ..... 187, 188 cunningtoni, Anchiphiloscia . . . 79 cupressina, Sertularia . . . 380, 405 Curds, C. R. . . . . l-6i cursorium, Ligidium 75 Cyphonetes ...... 73 Cyphoniscellus . . . . • 71 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 . . • 71 dalmatinus, Armadilloniscus . . 79 damae, Hora . • 83 danicus, Haplophthalmus . 71, 79 debilis, Ischioscia ... -83 debilis, Philoscia ... -83 Decapoda . . • 237-247 deliculata, Obelia . • 280 demarcata, Philoscia . 91 demarcata, Setaphora . • 9* dentata, Charybdis six- . . 146, 147 dentata, Scolecithricella . • 341 denticulata, Campanularia . . 279, 280 denticulata, Scolecithricella . . 327 denticulatus, Lepidoniscus pruinosus . 84 dentiens, Gulella ... • 173 dentiger, Androniscus . • 7° dentiger ligulif er, Androniscus . . 7° dentipes, Ligia 75 430 INDEX depressa, Kogmania .... 69 depressifrons, Portunus . . . 146, 147 depressus, Macrophthalmus . 237, 242-246 Deto 81-82,98 Detonella ...... 98 Diacara ...... 82 Diaixidae ...... 367 diaphana, Eucope . . . . .273 diaphana, Thaumantias . . . .273 dichotoma, Campanularia . . . 265 dichotoma, Laomedea . . . 265, 267 dichotoma, Obelia . 251, 253, 254, 255, 256, 257, 258, 259, 266-272, 273, 277, 278, 279, 281 dichotoma, Sertularia . . 253, 254, 265 dichotomum, Halecium . . . .410 Didima ...... 82 difficilis, Scaphocalanus . 815-317, 333 dilatatus, Macrophthalmus . . . 245 dilectum, Philoscia .... 88 dimidiatus, Nanochromis . . . 235 diminuta, Philoscia .... 88 Diphasia ...... 410 divaricata, Laomedea . . . 265, 266 divaricata, Obelia ..... 266 dogieli, Euplotes . . . .16, 25-26 dolinensis, Haplophthalmus fiumaranus . 71 dolomiticus, Oroniscus .... 84 dorsalis, Chaetophiloscia . . . .81 dorsispinosus, Spinocalanus . . . 303 Dotilla ..... 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 I 339, 341-342 • 339 266, 272 333, 336, 338, 341 • 385 . 386 377,385-390 ... • • 385 • 415 273, 405 • 318 187-190 23,37 • 394 93 . 76 73 . 82 • 344 . 82 8, 47, 48 . 48 • 279 63-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 patella everta, Obelia exigua exigua, Tiroloscia exigua, Tiroloscia exigua exigum, billardi, Halecium exigum, Halecium billardi 81 • 361 158, 160, 162 217-226, 4 Pis. 267 217-226, 4 Pis. 93 93 93 349,351,353-361 266, 271, 273, 280 398, 410 . 378 204 204 • 74 6, 8, 10, 16, 17, 19, 26-29, 41 26, 27 267 93 93 . 412 . 412 fagorum, Stylohylea .... 72 fagorum, Trichoniscus .... 72 falcata, Hydrallmania . . . .380 falcifer, Amallothrix . . 325, 327, 341 falcifer, Scolecithrix . . . -325 farciminoides, Salicornaria . . . 409 fasciata, Philoscia . . . . .91 fasciata, Setaphora . . . .91 fasciatum, Atoxon . . . 207, 215 fecundu, Euaugaptilus . . . 351-352 fenestrata, Thuiaria . . . .381 ferox, Gaetanus ..... 306 ferrani, Scopalatum . . 336, 338, 342 Filellum . . . 378, 379, 380, 381, 385 filiforme, Halecium . 403, 405, 406, 409 fistulosa, Cellaria .... 409,411 fiumaranus, Haplophthalmus . . .71 fiumaranus dolinensis, Haplophthalmus . 71 flabellata, Campanularia . . . 266 flabellata, Obelia . . . 266, 270, 271 flava, Philoscia 88 flavescens, Niambia .... 77 flavus, Oritoniscus . . . 72, 73 flavus, Trichoniscus .... 72 flexuosa, Campanularia . . 256, 257, 281 flexuosa, Sertularia . . . .272 Flustra .... 387,412,414 foliacea, Flustra . . . . .412 formicarum, Niambia .... 77 formosana, Chaetophiloscia . . .81 Formososcia ...... 82 foveolatus, Trichoniscus .... 74 fragilis, Allocentrotus .... 37 INDEX 431 fragilis, Allocentrus . . . .22 fragilis, Alpioniscus .... 69 fragilis, Obelia ..... 266 fragilis, Pseudophiloscia .... 89 fragilis rharelbazi, Trichoniscus . . 73 fransiscanus, Strongylocentrotus . . 23 frigidana, Philoscia muscorum . . 88 fruticosa, Campanularia .... 386 fruticosa, Lafoea .... 386, 389 fucorum, Halophiloscia .... 82 fulva, Benthanops ..... 88 fusiformis, Encope . . . 271, 273 fusiformis, Obelia . . . . .271 gaboniscus, Nanochromis . . 233-235 Gaetanus . . . . . . 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, 281 265 265 272 272 251, 253, 254, 255, 256, 257, 258, 259, 271, 272-278 geniculata, Sertularia . . . 265, 272 geniculata subsessilis, Obelia . . .275 geniculatum, Halecium . . 393, 396, 409 Gentry, A. W. . . 217-226, 4 Pis. Geologia ...... 75 georgensis, Styloniscus .... 68 georgensis, Trichoniscus .... 68 germanicus carniolense, Lepidoniscus . 84 gibbera, Scopalatum . 335, 386-338, 342 gibbosulus. Cancer .... 239 glarearum, Stenophiloscia ... 92 globiceps, Rhyscotus .... 76 globosus, Scopimera . . . .245 Gonotha ..... 260, 264 Gonothyrea . . . 253, 260, 265, 266, 281 gelatinosa, Laomedea genicolata, Sertolare geniculata, Campanularia geniculata, Laomedea geniculata, Obelia gottscheensis, Cyphoniscellus gracilicauda, Heterakis gracilicornis, Halophiloscia gracilipes, Ligia gracilis, Campanularia gracilis, Euplotes . gracilis, Laomedea gracilis, Obelia gracillima, Campanularia gracillima, Lafoea . graecus, Labyrinthasius . Grammaria . granulata, Bathytropa . granulatus, Tylos . 71 199-201 82 • 75 . 281 8, 47, 48-49 . 266 266, 281 . 386, 387 386, 389, 390 . 83 378, 379, 381-385 79, 83 . 65 granuliferus, Tylos .... 65 graphophasiani, Cyrnea .... 204 grarosensis, Philoscia .... 88 gravieri, Clytia . . . . .281 grevyi, Equus . . . . .218 grimni, Obelia ..... 266 guinasana, Tilapia . . . .196 griseoflavus, Niambia .... 77 griseus, Hiatoniscus .... 83 guernei, Philoscia ..... 88 Gulella ..... 158-173 gwendolinae, Ennea . . . 158, 162 gwendolinae, Gulella 158, 162, 163, 164, 171 gwendolinae, Gulella gwendolinae . .163 gwendolinae aldabrae, Gulella 168-164, 171, 172, 173 gwendolinae gwendolinae, Gulella . .163 gwendolinae mkusiensis, Gulella . 162, 163 gwendolinae porrecta, Gulella . . 162, 163 gwendolinae scissidens, Gulella . 162, 163 gwendolinae tsadiensis, Gulella 158, 162, 163 gymnothalma, Obelia . . . .273 375-376, 390-4I5 • 393 . 393-396, 399, 4°9. 412 391-4U, 412, 414 391, 399 82, 88 72 345, 347-349 Haleciidae halecina, Sertularia halecinum, Halecium, Halecium Haloikema .... Halophiloscia halophilus, Miktoniscus . Haloptilus .... hamuligerus, Androniscus roseus . . 70 Hanoniscus ..... 82, 98 Haplophthalmus . . . . 71- 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 . . • .281 hercegowinensis, Cyphonetes ... 73 heroldi, 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, Hemiplax hispana, Bathytropa Holothurioidea holsatus, Macropipus holthuisi, Papuasoniscus hoplites, Spinocalanus Hora .... horae, Styloniscus . horae, Trichoniscus horridus, Spinocalanus . hottentoti, Styloniscus . hottentoti, Trichoniscus . humilis, Didima hyalina, Gonothyrea hyalina, Obelia Hydrallmania Hydranthea . Hydrodendron Hydroida Hydrozoa Hyloniscus . hyperboreus, Euaugaptilus hypnorum, Ligidium Hypotrichida 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. 304 68 68 82 . 266 265, 266, 267 380 390, 412-414 • 4*4 • 375 249-293 71-72, 73, 97 359, 360-361 75 . 1-61 72 . 411 . 310 70 244—246 378, 379, 380 . 310 8, 45-46 . 67 . 183 71 insulanus, Trichoniscus noricus insulare, Pembatoxon insulincola, Gulella intermedia, Bathypontia intermedia, Grammaria . intermedius, Scyphax invalidus, Scaphocalanus irregularis, Obelia . Isabella, Bilawrencia Isabelloscia . Ischioscia . . , isolonche, Heterakis Isopoda 101-120, I PI., 121-151 . 17, 29-30 382, 383, 385 309, 310 jacquelinae, Gulella Japanoniscus 74 . 207-216, i PI. 161, 168-169, 171, 172 • 363 . 382. 384 91 • 317 267 80 . 83 . 83 2OI 63-1006 1 66 83 japonica, Charybdis japonicum, Nippoligidium japonicus, Charybdis japonicus, Macrophthalmus karongae, Anchiphiloscia kempi, Burmoniscus kenepurensis, Oniscus kermadecensis, Styloniscus kermadecensis, Trichoniscus Kerona kincaidi, Campanularia . Kogmania Komatia Krantzia 149 75 147 245 79 80 84 68 68 41 280 69 89 83 396-399, 401, 411 • 83 . 187 187, 188 187-188 377. 379, 381, 385-390 375-376, 377-390 406 272, 273, 278 . 320 • 4*4 312, 320, 325 . 167 399-402 • 399 labrosum, Halecium Labyrinthasius laevigata, Cucumaria laevigata, Pentactella laevigatus, Pseudocnus Lafoea . Lafoeidae Lafoeina lairii, Laomedea lamellifer, Scolecithricella Laminaria laminata, Scolecithricella langi, Gulella lankesteri, Halecium lankesterii, Haloikema . Laomedea 251, 252, 253, 254, 260, 261, 264, 265, 266, 267, 272, 273, 278, 280, 281 lata, Manibia . . . . -77 lateralis, Paraphiloscia .... 85 lateralis, Pseudophiloscia ... 85 latifrons, Euaugaptilus .... 353 latipes, Lophothrix . . . 313-315 latipes, Portumnus . . . .145 latissima, Euryligia .... 74 latreillis, Macrophthalmus . . . 245 latum, Ligidium . . . . -75 latus, Euplotes .... 16, 30-3 1 latus, Euplotes patella . . . 30, 31, 41 latus, 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. . . 70 Lincoln, R. J. . . . . 63-iooe linearis, Obelia . . . . .280 INDEX 433 linearis, Rhyscotoides linearis, Rhyscotus Lithotis littoralis, Armadilloniscus littoralis, Euplotes elegans lobophora, Amallothrix . longa, Obelia longicarpus, Mictyris longicauda, Niambia longicirrhus, Euaugaptilus longicirrus, Haloptilus longicornis, Haloptilus . longicornis, Philoscia longicyatha, Clytia longicyatha, Gonotha longicyatha, Gonothyrea longicyatha, Laomedea . longicyatha, Obelia longifurca, Scaphocalanus longipes, Bathynectes longipes, Spinocalanus longiseta, Euaugaptilus . longissima, Laomedea longissima, Obelia . longissima, Sertularia longitheca, Clytia . longitheca, Obelia . Lophothrix . loveni, Gonothyrea Lubbock, R. lubricata, Philoscia lubricata, Setaphora lucifera, Charybdis lucifera, Obelia lymani, Amphiura . 76 . 76 . 184 79 . 48 329, 335, 336 260, 263 245, 246 77 355-357 • 348 • 348 88 260, 261, 262, 265 260, 264 260, 265 261 260, 261, 264, 280 316-317 . 142 . 301 • 35i 265 256, 258, 271, 278 • 265 260 260 313, 321, 323, 325 . 281 227-231, i PI. 92 92 146, 147 251, 272, 278, 279 188 macchiae, Tiroloscia .... 93 macrocephala, Rennelloscia ... 89 macrodactyla, Chiridiella . . . 306 Macrophthalminae . . . 245, 246 Macropipinae . . . . .127 Macropipus . . . . 118, 121-151 Macrothalmus . . . 237, 242-246 maculatus, Pinnotheres . . . 240, 242 maenus, Carcinus . 101-I2O, i PL, 123, 145 maenus mediterranea, Carcinus . . 118 maenus septentrionalis, Carcinus . . 118 magellanica, Grammaria . 382, 383, 384 magister, Cancer . . . . .239 magnicirratus, Euplotes . . .16, 31-32 magnus, Spinocalanus . . . 297, 298 maior, Campanularia .... 265 Mammalia .... 217-226, 4 Pis. Manibia ...... 77 margarica, Atractylis . . . .412 margarica, Hydranthea . . . 412-414 margaricum, Halecium . . . .412 marginata, Komatia . . . .89 marginata, Obelia ..... 280 margine papillosa, Niambia . . -77 mariae, Hyloniscus . . . .71 Marina ...... 25^ marina, Deto ..... 82 marina, Obelia .... 253, 279 marina, Philougria .... 82 marionensis, Cucumaria serrata . 187, 188 marionis, Nullamphiura . . . .188 marionis, Uca ..... 245 Marioniscus ...... 84 marmoreus, Macropipus . 124, 125, 126, 135-142 martensi, Ophioglypha . . . .189 martensi, Ophiurolepis .... 189 mascarenensis, Succinea . . .182 mauritiensis, Styloniscus ... 68 mauritiensis, Trichoniscus ... 68 maxillaris, Euaugaptilus. . . 353-355 mediterraneus, Carcinus 101-I2O, i PI., 145 mediterranea, Carcinus maenus . . 118 medius, Androniscus subterraneus . . 70 medius, Spelaeonethes .... 73 Medusa .... 252, 253, 254 medusa, Marina ..... 253 medusiferum, Campalecium . . 391 meeusei, Chaetophiloscia . . .81 meinerti, Bathytropa .... 79 meinerti costata, Bathytropa ... 79 melanocephala, Ligia .... 75 mendanai, Paraphiloscia ... 86 meridionale, Atoxon . . . 207, 215 methueni, 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 . .261 minor, Laomedea spinulosa . 260, 261, 264 minor, Valdiviella .... 309-3 n minuscula, Pupa ..... 166 minuta, Euplotes . . . 9, n, 13-14 minutissima, Trichorhina ... 78 mirabilis, Ophiodes . . . .414 mirabilis, Ophiodissa . . . 414-417 Mirigulella . . . . . .167 mixtus, Euaugaptilus . . . 358-360 mixtus, Trichoniscoides .... 73 mixtus, Trichoniscus 73 mkusiensis, Gulella gwendolinae . 162, 163 modesta, Niambia ..... 77 modestus, Trichoniscoides . • • 73 434 INDEX modestus, Trichoniscus .... 73 moebiusi, Euplotes 5, 8, 24, 42, 43, 47, 49-5O Mollusca 157-175; 177-186, i PL; 207-216, i PI. monocellatus, Microniscus monocellatus, Styloniscus Monosklera . montana, Plymophiloscia montanus, Paranotoniscus montanus, Trichoniscus . montanus, Trichoniscus vividus moruliceps, Styloniscus . moruliceps, Trichoniscus multidentata, Obelia muricata, Sertularia muricatum, Halecium murigatum, Halecium murrayi, Styloniscus murrayi, Trichoniscus muscivagus, Trichoniscus muscicola, Euplotes muscorum affinis, Philoscia muscorum biellensis, Philoscia muscorum dalmatica, Philoscia muscorum, Euplotes muscorum frigidana, Philoscia muscorum, Pogonoligia . muscorum triangulifera, Philoscia mussaui, Clavigeroniscus 68 68 252, 254, 273, 276 . 89 68 73 73 68 . 68 . 260 402 402-405, 409 • 403 . 69 . 69 74 - 8, 45, 46 . 87 . 87 47, 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, 33 nematodes .... 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 . . 74 noricus sassanus, Trichoniscus . . 74 noricus sturanus, Trichoniscus . . 74 nova brittanica, Rennellscia ... 89 novemcarinata, Euplotes nudiceps, Nanochromis . nudulus, Tylos Nullamphiuria nyiroensis, 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, Macrophthalmus pagurus, Cancer pallida, Chaetophiloscia . pallida, Niambia . 51-53 • 235 . 65 188, 189 . 167 . 252, 254 252, 254 249-293 252, 254 318-319 • 309 266, 272 267, 272 272 267 82 127, 142, 143, 144 204 . 38, 39-41 . 16, 32-33 • 245 • 245 242, 244, 245 . 84 . 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, 73 68 91 . 84 . 76 . 76 • 394 240, 242 . 69 . 69 145, 147, 148, 149 306-307 92 260, 263 • 177 • 245 237-240, 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, Styloniscus phormianus, Trichoniscus Phymatoniscus picteti, Laomedea bicuspidata pigmentata Ligia . pigmentatus, Olibrinus . pilosa, Setaphora . pinnata, Zygophylax Pinnotheres . 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 • 390 26, 30, 38, 41-42 41 41 . 41 41 26, 27 . 30. 3L 4i 41 41 41 88 177-186, 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, 369 87, 98 73 . 280 . 281 23 87-88, 91, 92, 93, 98,99 82, 88 . 69 . 69 72 260, 264 75 . 84 92 378, 402 237, 240—242 pinnotheres, Pinnotheres . . . 242 Pinnotheridae . . . . .242 piriformis, Obelia . . . . .266 Pisces .... 191-197, 233-235 pisum, Pinnotheres . . 237, 240-242 placunae, Pinnotheres . . . 240, 242 plana, Obelia ..... 266 plana, Thaumantias . . . .271 planctonicus, Euplotes patella . . 41 Platyarthrus .... 78, 97 platycephala, Ligia .... 75 platycephala, Pogonoligia • • • 75 platytarsis, Octypode .... 245 pleonalis, Stenoniscus . . . .76 plicata, Obelia ..... 266 Ploesconia . . . . n, 29, 41 plumipes, Euplotes polygena, Eucope . polystyla, Eucope . plumosa, Blugula . plumosum, Halecium Plymophiloscia pocillum, Hebella . pocillum, Lafoea . pocillum, Lafoea fruticosa Podophthalminae . poecila, Krantzia . Pogonoligia . poljanskyi, Euplotes 5, 26, 27 • 273 . 271 . 411 405, 406, 408, 409 . 89 • 390 386, 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 . . . 309 Pomacanthidae . . . 227-231, i PI. Pomacanthus .... 227, 230 ponticus, Tylos ..... 66 pooensis, Gulella .... 166, 167 porcellioides, Alloniscus .... 78 porcellioides, Arhina .... 78 porrecta, Gulella gwendolinae . . 162, 163 Portumnus Portunidae Portuninae Portunus poutrini, Gulella pretiosa, Gulella pretiosa pretiosa nyiroensis, Gulella pretiosa pretiosa, Gulella princeps, Metridia . pristis, Idiella . Procyrnea . . . . . productus, Cancer . profunda, Amallothrix . prolifera, Campanularia . prolifera, Sertularia propinqua, Paraphiloscia propinquus, Euaugaptilus provisorius, Trichoniscus pusillus pruinosus denticulatus, Lepidoniscus Pseudocnus . 145 IOI-I2O, I PI., I2I-I5I 127, 142, 144, 146, 147, 148, 149, 150 123, 125, 142, 146, 147 169 167 . I67 . I67 • 344 . 411 204, 205 • 239 320, 321 272 272, 273 86 358, 360 74 . 84 187-188 436 INDEX Pseudochirella Pseudophiloscia pteronus, Spinocalanus Ptychotrema puber, Macropipus pugilator, Uca pugnax, Uca pulchella, Philoscia Pulmonata 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 . 307-309 85,89 298, 303 . i66 . 121-152 • 245 • 245 88 177-186, i PL, 207-216, i PL • 415 127, 144, 147 . 166 23, 37 273, 276 77 • 398 • 396 • 245 74 74 • 390 . 266 . 87 . 87 . 298 74 72 93 93 . 266 266 quadrata, Octypode .... 245 quagga, 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, 384 ramosum, Schizocladium . 266, 272, 273 rariseta, Euplotes . . . 38, 42—43, 49 reflexum, Halecium . . . 396, 398 refugiorum, Hyloniscus .... 72 rendalli, Coptodon .... 196 rendalli, Tilapia ..... 196 Rennelloscia ..... 89-91 resplendens, Centropyge . 227-231, i PL Reticularia . . . 378, 380, 381, 382 Rhachis ...... 171 rharelbazi, Trichoniscus fragilis . . 73 rhunicola, Obelia ..... 266 Rhyscotidae ...... 76 Rhyscotoides ..... 76 Rhyscotus ...... 76 Rice, A. L. 101-I2O, i PL, 121-151, 237-247 rigidus, Euaugaptilus . . . 351, 357 riparius, Hyloniscus .... 73 riversdalei, Styloniscus .... 69 riversdalei, Trichoniscus ... 69 robusta, Deto ..... 82 robusta, Grammaria . . . 381, 382 robustipes, Amallothrix . . . 365, 367 robustum, Halecium . . 399, 401, 402 Roe, H. S. J 295-372 rosacea, Diphasia . . . . .410 roscoffensis, Euplotes .... 53 roseus, Androniscus .... 70 roseus hamuligerus, Androniscus . . 70 rotunda, Euplotes .... $3-54 rotundatus, Atelecyclus .... 240 Rowe, F. W. E 187-I9O rupicola, Anasterias . . . .190 rupicola, Lithotis . . . . .184 rupium, Halophiloscia adriatica . . 82 ruweti, Tilapia .... 194, 195 Sabellaria ...... 408 Sagitta ...... 279 Salacia ..... 381, 382 Salicornaria ...... 409 sancristobali, Paraphiloscia ... 86 sanguinolentus, Portunus . . 146, 147 Sankey, R. D. . . . 227-231, i PL santaisabellae, Paraphiloscia . . .87 sapidus, 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 . . 70 Scandia ...... 377 Scaphocalanus 312, 317~3i8, 320, 321, 325, 329, 333. 339 Schiodtia . . . . . . 97 Schizocladium . 252, 254, 266, 272, 273 schneideri, Halecium . . . 396, 399 schobli, Platyarthrus .... 78 Schoblia ...... 69 Schobliidae . . . . . 65, 69 scissidens, Gulella gwendolinae . 162, 163 Scolecithricella 312, 318, 320, 321, 323, 329-341 Scolecithricidae . . 311-342, 367, 369 Scolecithrix . . .321, 325, 333, 336, 338 Scopalatum . 333, 335, 336, 338, 341, 342 scoparum, Trichoniscoides ... 73 Scopimera .... 244, 245, 246 Scopimerinae ..... 245 scotti, Scolecithrix . . . -338 Scottocolanus . . . . -313 scutum, Helecium ..... 391 Scylla ..... 146, 147, 148 INDEX 437 Scyphax .... Scyphoniscus securifrons, Scottocalanus septentrionalis, Carcinus maenus seriepunctata, Philoscia . serpens, Capsularia serpens, Filellum . serpens, Grammaria serpens, Rictularia serrata, Cucumaria serrata, Scylla serrata marionensis, Cucumaria serratula, Obelia . serrulata, Campanularia Sertolare .... Sertularia 252, 253, 254, 265, 272, 385, 391, seruposa, Corallina sessile, Halecium . Setaphora .... setosa, Sagitta silicus, Haplophthalmus sima, Thalamita simoni, Oniscus sinensis, Centropus singhi, Cyrnea six-dentata, Charybdis . Slabberia .... smithae, Amallophora . Snelliaetideus solowetzkiana, Obelia sparrmanii, Tilapia . 191, spatium, Gulella spatulifrons, Marioniscus Spelaeonethes sphaerocephalus, Rhyscotus . sphaerulina, Obelia spinicarpus, Portunus spinifera, Bathypontia . Spinocalanidae Spinocalanus spinosa, Sertularia spinosus, Cordioniscus . spinosus, Spinocalanus . spinosus, Trichoniscus spinulosa, Campanularia spinulosa, Laomedea spinulosa, Obelia . spinulosa minor, Laomedea Spiruridae .... spurca, Quickia . . 177, 178, spurca, Succinea squamata, Niambia squamatus, Leptotrichus squamatus, Platyarthrus candatus squamiceps, Nanochromis Squamiferidae squamuligera bargensis, Tiroloscia squamuligera briani, Philoscia squamuligera briani, Tiroloscia 91 91 • 313 . 118 88 • 378 378-381 379, 385 • 379 . 188 146, 147 187, 188 . 280 . 280 265 273. 380-381, 393- 394. 4°5 • 393 405, 406-409 79, 91-92, 99 • 279 71 146 . 84 204 204 146, 147 . 252 332, 336, 342 304, 305 . 266 194, 195, 196 166, 167 . 84 73 . 76 253, 265, 266 146 . 364 297-304 297-304 • 253 . 67 297, 298 . 67 260, 263 260 260, 265 260, 261, 264 201-205 180, 182, 183 . 180 77 • 77 • 78 • 235 65, 77-78, 97 93 93 93 squamuligera tendana, Tiroloscia stammeri, Trichoniscus . stebbingi, Cordioniscus . stebbingi, Trichoniscus . stellatus, Spinocalanus . Stenoniscidae Stenoniscus .... Stenophiloscia stentor, Grammaria Stewart, D. J. 93 74 . 67 . 67 • 303 65, 76 . 76 92 382, 383, 384, 385 191-197 strasseri, Androniscus cavernarum . . 70 strasseri, Illyrionethes .... 70 strelkovi, Euplotes . 5, 7, 38, 43-44, 51 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, Philoscia suarezia, Setaphora subbrevicornis, Scaphocalanus subsessilis, 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 92 79,92 • 317 273, 277 • 275 88 • 70 • 70 70 177, 180, 182, 184 177-186, i PI. 244, 245 • 245 142, 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 252 thermophila, Bathytropa . 69 . 69 240, 242 8, 9. 45, 46-47 227-231, i PI. 93 - 76 65, 76 >, 398,409-411 . 346 260, 264 267 54, 55 . 381 146, 147 • 367 271, 273 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 . . . 188-189 tomentosa, Trichorhina .... 79 torreyi, Halecium . . . 391 toticostata, Gulella . . . .167 tottoni, Laomedea . . . .281 Trewavas, E. . . 191-197, 233~235 triangularis, Uca ..... 245 triangulifera, Philoscia muscorum . . 88 tricciratus, Euplotes affinis . . .18 Trichoniscidae ... 65, 69-74, 97 Trichoniscoides .... 72, 73 Trichoniscus . . 67, 68, 69, 72, 73-74, 97 Trichorhina . . . . . 78, 79 Trichoda . . . . . .41 trisulcatus, Euplotes . . .16, 86-37 trituberculatus, Portunus . . 146, 147 truncata, Niambia .... 77 truncatella, Philoscia .... 92 truncatella, Setaphora . . . .92 tsadiensis, Gulella gwendolinae 158, 162, 163 tuberculata, Pseudochirella . . 307-309 tuberculatus, Hanoniscus ... 82 tuberculatus, Paranotoniscus ... 68 tuberculatus, Phymatoniscus ... 72 tuberculatus, Trichoniscoides ... 72 Tubularia ...... 414 tuffraui, Euplotes . . 4, 6, 8, 16, 17, 37-38 Tylidae 65-66, 96 Tylos 65-66, 96 Tympanomerus ..... 244 typica, Amallophora . . . 335, 338 typicus, Euplotes patella . . .41 typicus, Xanthocalanus . . 332, 336, 338 tyrrhena, Halophiloscia .... 82 Urocyclidae .... usitatus, Spinocalanus Valdiviella .... valens, Amallothrix Van Bruggen, A. C. Van Goethem vannus, Euplotes . vannus bolticus, Euplotes variabilis, Euplotes variabilis, Euplotes patella ventosus, Styloniscus ventosus, Trichoniscus . verhoeffi, Trichoniscus . verrucosa, Tendosphaera verrucosus, Styloniscus . verrucosus, Trichoniscus verticillata, Sertularia Verticillina .... Vesicularia .... veterum, Pinnotheres vilis, Aphiloscia vilis, Philoscia villosa, Benthana . villosa, Philoscia . violaceus, Euplotes vittata, Philoscia . vividus, Trichoniscus vividus montanus, Trichoniscus vivipara, Heterorhabdus vivipara, Ophiacantha . volubis, Sertularia vorax, Scolecithrix waitatensis, Scyphoniscus warreni, 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, 29 41 . 69 . 69 • 74 . 76 . 69 . 69 • 254 • 254 • 253 . 240 79, 88 79 80 80 ii . 88 72 73 • 345, 346 . 189 •/ 254 332, 336, 342 88 409 56 Xanthocalanus 311-312, 332, 335, 338, 864-367 Xavia ....... 145 Uca . undotheca, Obelia . undulatum, Halecium 244, 245 . 266 396, 398, 399, 4" Zebra., Equus zenkewitchi, Euplotes zosterae, Trichoniscus Zygophylax . 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series, Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28, No. i, of the Zoology series. The abbreviated titles of periodicals cited follow those of 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 OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 19 May 1975 Price £3.80 A GUIDE TO THE SPECIES OF THE GENUS EUPLOTES (HYPOTRICHIDA, CILIATEA) By COLIN R. CURDS INTRODUCTION ALTHOUGH species of the genus Euplotes Ehrenberg, 1830 are frequently observed in both marine and freshwater samples it is often not possible for the protozoologist to make specific identifications. This difficulty is due to at least three major factors ; firstly, many of the features used for distinguishing species within the genus are known to vary considerably even within clonal cultures, secondly, there is a large amount of confusion in the literature concerning the identity of certain common species and thirdly, there is the added difficulty of having to search through a considerable body of literature before an identification can be attempted. It is hoped that the present paper will help solve some of these problems by gathering together descriptions and diagrams of all those organisms considered to be distinct Euplotes species. Naturally the serious worker will still wish to refer to the original material but the volume of literature needed to be examined should be significantly reduced. 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 knowledge no attempt has been made to devise a key to the species of Euplotes since that of Kahl (1932). FEATURES OF TAXONOMIC IMPORTANCE In the past 200 years over 80 species and varieties of the genus Euplotes have been described. Until relatively recently the species were separated simply on a basis of body size and shape, on the dorsal and ventral ridges and on the arrangement of 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 might be of taxonomic importance for species determination in the genus Euplotes. Tuffrau (1954) first suggested that the numbers of dorsolateral rows of cilia, the geometry of the argyrome, the number of frontoventral cirri and the form of the macronucleus could be used to distinguish species and later (Tuffrau, 1960) he presented his revision of the genus based on these ideas. Since that revision more species have been redescribed and confirmed on the basis of silver preparations (principally Borror 1962, 1963, igGSa) and several new species have been introduced. Borror (1972) published a list of species he found acceptable based on modern criteria together with their probable synonyms. In general the present author's findings confirm those of Borror (1972) although there are several instances where changes have been suggested. Carter (1972) added four new species to the list of Borror (1972) and redescribed 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 within the genus. It is unfortunate that of the taxonomic characters mentioned above only one - the gross geometry of the dorsal argyrome - remains constant. All other features have been reported to vary to a greater or lesser extent and an account of some of these variations follows. However, the application of these modern criteria now makes it possible to examine and reappraise the species and varieties within the genus. (a) General morphological and ecological features Although Kahl (1932) used the habitat of Euplotes species as a strict taxonomic character 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 be useful for species determination. To date only two species have been recorded in urchins, E. balteatus (Dujardin, 1841) which may be found both as a commensal and free-living and E. tuffraui Berger, 1965 which seems to be limited to a commensal mode of life. The size of a Euplotes species can be used as a taxonomic feature but must be applied with extreme caution. It is true that the smallest species never attain the size of the largest species and vice versa but it is also known that the size 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) and kind of food (Giese, 1938 ; Tuffrau, 1964). The overall shape of a Euplotes species is perhaps a rather more stable feature but is of limited taxonomic value since few species have really characteristic outline shapes. Dorsal and ventral ridges are sometimes of value in species determination particularly when comparing modern preparations with early descriptions where ridges were often clearly figured. Borror (igGSa) discussed cortical sculpturing and suggested that the use of nigrosin-HgCla-formalin method (Borror, i_968b) might help create a union of old and new species within the genus. Coloured species of Euplotes have been encountered for many years ; both Ehrenberg (1840) and Stein (1859) described green species which were likely to be forms of E. patella (Muller, 1773). Later Kahl (1932) listed three 'formae' of E. patella which contained zoochlorellae. It should be emphasized that one must distinguish 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, a species closely related to E. patella, as continually containing zoochlorellae and this perhaps 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 have been 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 of faith on the numbers of caudal cirri as a taxonomic character. Kahl (1932), for example, differentiated between varieties of E. moebiusi Kahl, 1932 and E. affinis (Dujardin, 1841) based on the numbers of caudal cirri and there are many other similar examples. However, modern evidence has clearly shown that variation in caudal cirri numbers is a common feature, and indeed seems to be the rule rather than the exception in certain species. Hufnagel and Torch (1967), for example, demonstrated in clones of E. vannus (Muller, 1786) that on division the proter always received five and the opisthe four caudal cirri no matter how many caudals the mother cell originally had ; thus these two morphological variants were found in equal proportions in the total population. Hufnagel and Torch (1967) further Frontals Fronloventrals Ventrals Anals or Transversals Caudals FIG. i. Ventral cirri of Euplotes. a. Wallengren (1900) system of cirrus numeration and older method of naming cirri groups, b. Modern method of naming groups of cirri. showed that this variation was due to the normal morphogenetic events involved in the development of the right caudal cirri from the right dorsal kinetics. In addition Hufnagel and Torch (1967) observed transient abnormal caudal cirrus development 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 evident from this type of information that the number of caudal cirri is of little, if any, value in the identification of species of Euplotes. The reader is recommended to consult the 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 reported to vary in their number which is remarkably constant throughout the genus. With the exception of E. strelkovi Agamaliev, 1967 which has 6 transversals all other species have 5 transverse cirri. The number of frontoventral cirri is of greater use for species determination than the other groups of cirri since although they vary from 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 frontoventral cirri, far fewer have 8 while only E. raikovi Agamaliev, 1966 has been reported to have 7 frontoventrals. With the exception of E. tuffraui the variation in fronto- ventral numbers seems to depend upon streak V (Wallengren, 1900 system, see Fig. i). Euplotes raikovi was reported (Agamaliev, 1967) to exhibit intraspecific polymorphism in the number of frontoventrals ; he noted that there were 7 or 8 frontoventral cirri and indicated that cirrus Vz (Fig. i) was that which did not develop in some specimens. Washburn and Borror (1972) described a strain of E. raikovi from America in which an eighth cirrus (¥2) never developed although they did observe a barren plaque in each case. Curds (1974) recently reported a similar case in his description of E. parkei Curds, 1974 where cirrus V2 was present in some and absent from other specimens even within the same clone. The frontoventral cirri of E. tuffraui show a wider variation than reported in other species and furthermore the 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 urchins possess 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,IIl3 and IV 2. A further two specimens were found with 9 frontoventral cirri where only cirrus bases III 3 and IV 2 were subdivided. With the exception of the examples listed above the number and positioning of the frontoventral cirri nevertheless remain very valuable taxonomic characters that have been widely used in the past and will do so in the future. (c) The dorsal argyrome The overall geometrical pattern of the dorsal argyrome is one feature of taxonomic importance 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 which he called 'muscicola', 'eurystomus' and 'vannus' after the species in which he first found them. Since that time many more silver-line systems of Euplotes species have been described in the literature and it is now possible to divide the dorsal argyrome patterns into five or six types as shown in Fig. 2. The simplest type (Fig. 2a) includes six species whose dorsal interkinetal argyrome pattern is composed of longitudinal kinetics with simple transverse connections between them so that there is a single row of polygons between the dorsal cilia or bristles. This group corresponds to the 'vannus' type of Tuffrau (1960) although it is suggested that the term 'single-vannus' is more descriptive and consistent with the terms to be used later. The first complication in the dorsal argyrome pattern is shown in Fig. 2b, where the 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 dorsal cilia. 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 and since '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 of 24 species from another group of 8 species which also have double but unequal rows of polygons between the dorsal cilia. Perhaps the best known species in the latter group 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 dorsal cilia, 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. strelkovi and E. zenkewitchi Burkovsky, 1970 the wide rows are on the right and the narrow rows 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 this does not seem warranted unless more species whose argyrome patterns conform to the second sub-type are found. In the present author's opinion the 'muscicola' type of Tuffrau (1960) included a heterogeneous assemblage of dorsal argyrome patterns that should now be divided into 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 seen that (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 of polygons 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. muscicola Kahl, 1932 respectively. It is suggested that argyrome patterns such as these be called 'multiple' types. The final complication in dorsal argyrome patterns is that the polygons become so subdivided that an irregular network or mesh is formed between the dorsal cilia so that distinct rows of polygons cannot be distinguished. It is suggested that this pattern be called the 'complex' type (Fig. 2f). The latter pattern is illustrated well by the species E. gracilis Kahl, 1932, E. muscorum Dragesco, 1970 and E. elegans Kahl, 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 polygons interspersed with an irregular mesh work. With two exceptions the dorsal argyrome patterns of the species of Euplotes that have been described to date fit neatly into the types outlined above. One of these exceptions, E. moebiusi has already been mentioned and can readily be fitted into the 'complex' group by the presence of an irregular meshwork between the dorsal cilia. The other exception, E. tegulatus Tuffrau, 1960, which was previously placed in the 'eurystomus' type (Tuffrau, 1960) is more difficult since it could almost equally well 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 rows of large polygons between the dorsal cilia as in E. eurystomus but here there is also a central regular line of narrow elongate polygons. Euplotes tegulatus therefore displays 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 species with two equal rows of polygons between the kinetics. Whereas the overall geometrical pattern of the dorsal argyrome appears to be a completely stable feature, the number of dorsolateral kinetics has been reported to vary in several species including the following ; E. polycarinatus Carter, 1972 (20-21 kinetics), E. variabilis Stokes, 1887 (a synonym of E. eurystomus Kahl, 1932, 8-12 kinetics), 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. mutabilis Tuffrau, 1960 (11-13 kineties, see Tuffrau, 1960), E. eurystomus (8-9 kineties, see Bonner, 1954) and E. tuffraui (9-10 kineties, see Berger, 1965). However, in the majority of species the number of dorsolateral kineties remains constant. In the case of E. mutabilis, Tuffrau (1960) concluded that the variation in the number of kineties was accidental and exceptional, although even accidental variations seem to be widespread in nature. In single clonal cultures Bonner (1954), Carter (1972) and Heckmann (1963) have all reported instances of variation in the number of dorsolateral kineties. Furthermore, Carter (1972) found in three strains of E. eurystomus (a synonym of E. aediculatus) that the number of kineties was constant within the clone but different from clone to clone. In spite of variations such as these 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 those of kinetics, even so variation within the clone is frequently much less than from species to species. Although several authors have recorded the number of dorsal cilia and by doing so have inferred that they are of taxonomic value no one has yet suggested this number to be of real taxonomic importance. In the key that follows the number of dorsal cilia has been used frequently but only when the numbers are sufficiently different to warrant their use. In each case the approximate range in the mid-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 since distinctive patterns are not immediately recognizable. However, some authors have used the general size of the constituent polygons on a comparative basis and this may be given as additional information. Silver-line preparations of the ventral surface display the infraciliary network, the cirri plaques and the adoral zone of membranelles (AZM) which is of kinetosomal origin. Whereas in the past the size and shape of the AZM have often been used as taxonomic features, the number of membranelles therein has only recently been introduced. Carter (1972) found, in n species that he studied, that the AZM features were even more stable than those characters suggested by Tuffrau (1960). However, all the features of the AZM are known 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 that both the size and shape of the AZM of E. balteatus was dependent upon the size of the cell and the number of membranelles in the AZM varied from 25 to 30 in small cells and from 70 to 80 membranelles in the case of giant individuals. Fortunately variation of this magnitude seems to be confined to that species, nevertheless variation in 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 to cases when there is a considerable difference in the numbers between two species, for example 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 in taxonomic schemes for many years and the dimorphic nuclei of ciliates have played an important part in ciliate taxonomy. The macronucleus of Euplotes is elongate and takes a variety of shapes which differ from species to species. Perhaps the most common form is the simple inverted C-shape while in others this has become modified to 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 is usually small and round but it does vary in size and shape and in E. tegulatus and E. aberrans Dragesco, 1960 the micronucleus is particularly large. The use of the shape of the macronucleus as a diagnostic feature in the genus Euplotes 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 say that the macronucleus of Euplotes takes a variety of forms during reorganization and 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 opinion seems to be that if the process of reorganization is understood and provided it is possible for the taxonomist to recognize the interphase macronucleus from one that is in another divisional state, then the interphase macronucleus is constant and charac- teristic of the species. The interphase macronucleus can be recognized by the presence of replication bands (Gall, 1959 ; Kluss, 1962 ; Prescott, Kimball and Carrier, 1962) which represent bands of DNA synthesis travelling from the two nuclear extremities towards the centre of the macronucleus. Prescott et al. (1962) showed that the macronucleus of E. eurystomus is in interphase when the replication bands are 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 and described for all species of Euplotes. The key that follows relies heavily upon features displayed by the silver-line technique but a series of descriptions of undesignated species follows the key (Section F, p. 50) and this includes all those species whose silver-line systems have not yet been described. Diagrams of silver preparations are given for all species where available and these have been obtained directly from original descriptions. Unless otherwise specified all scales given on diagrams indicate io jam. The characters for the key were selected and used in order of least variation. Therefore the first division of species into groups is made on a basis of the overall pattern of the dorsal argyrome and further subdivisions are made using 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 argyrome or 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 of the 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. balticus b 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. vannus b 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. crassus b ii dorsolateral kineties with 11-15 dorsal cilia in central kineties . E. mutabilis SECTION A. DESCRIPTIONS OF SPECIES Euplotes balticus (Kahl, 1932) Dragesco, 1966 Kahl (1932) first described this species as E. vannus var. balticus while later Tuffrau (1960) considered it to be a synonym of E. crassus. However, the silver preparations 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) ovoid marine species. The peristome is long and narrow extending down to about f of the body length with the AZM containing about 50 membranelles. There are 10 frontoventral, 5 transverse and 4 or 5 caudal cirri. The ventral argyrome consists of a few very large irregular polygons. The dorsal argyrome is of the single- vannus type 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 the macronucleus. 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 Ploesconia crassa 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 Chatton and Seguela (1940) and fully described by Tuffrau (1960). DIAGNOSIS. Euplotes crassus (Fig. 4) is a large (100-130 /mi long) elongate oval marine species. The dorsal surface is strongly sculptured by 8 longitudinal ridges. The peristome is long and narrow while the AZM consists of about 50 membranelles and extends f down the body length. The dorsal argyrome is essentially simple of the single type and there are 10 dorsolateral kineties with the central ones bearing about 26 dorsal cilia. There are 10 frontoventral, 5 transverse and 5 or 6 caudal cirri. 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 dorsal argyrome 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 the macronucleus. Euplotes cristatus Kahl, 1932 This species was originally briefly described by Kahl (1932) and subsequently by Tuffrau (1960) and Carter (1972). THE GENUS EUPLOTES 13 DIAGNOSIS. Eiiplotes cristatus (Fig. 5) is a medium (60 /tin long, 45 /mi wide) oval marine 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 dorsal surface is convex and there are 6 prominent ridges. The dorsal argyrome is of a simple single type with 8 dorsolateral kinetics bearing 11-15 dorsal cilia in the central rows. There are 10 frontoventral, 5 transverse and 4 caudal cirri. In the original description 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 with a heel-like extension on the posterior arm. The small, compact micronucleus is situated 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 marine species. The right margin of the peristome is almost straight and extends f down the length of the body. The AZM consists of 30-40 membranelles. The dorsal argyrome is of the single type with 9 dorsolateral kinetics bearing 12-13 cilia in the central 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 left side and a foot-like extension (Fig. 6). The micronucleus is situated anteriorly on the 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 revisionary monograph. Although the dorsal argyrome is of the single type, Tuffrau (1960) generally observed it as is shown in Fig. 7 where reorganization of the kinetosomal network is in progress. Tuffrau (1960) rarely found specimens in the quiescent state which 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 sections of the key since it would depend on the state of the organism into which group it would appear to fall. DIAGNOSIS. Euplotes mutabilis (Fig. 7) is a medium (95 /u,m long) oval marine species that is widest at its posterior extremity. The peristome is quite large and the right margin extends unevenly down the body to terminate in a spike-like projection. The AZM contains about 60 membranelles which extend down almost f of the body length. The dorsal argyrome is single in the quiescent state but with several irregular ramifications when undergoing reorganization. There are n dorso- lateral kinetics which bear 12 or 13 dorsal cilia in the central rows. There are 10 frontoventral, 5 transverse and 4 or 5 caudal cirri. The macronucleus is C-shaped with a pointed foot-like structure at the end of the posterior arm. The micronucleus is situated anteriorly. Euplotes vannus (Muller, 1786) Minkjewicz, 1901 This species has a long history which is given in detail by Tuffrau (1960), and the list 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 argyrome in state of reorganization. (After Tuffrau, 1960.) past but with the description of the silver-line system by Tuffrau (1960) most of the identification difficulties were eradicated. DIAGNOSIS. Euplotes vannus (Fig. 8) is a medium-sized (75-100 /*m long) marine species. 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 of the 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. There are 10 strong frontoventral, 5 transverse and 4 fine but rigid caudal cirri. The macronucleus is an open C-shape with a twisted foot-like extension to the posterior arm. The micronucleus is compact and lies close to and sometimes overlapping the 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 ....£. octocirratus b 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. parkei b 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. quinquecarinatus b 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. tujfraui b 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 echinoids b 21-25 dorsal cilia in central kineties, free-living ii or 12 dorsolateral kineties . 33 a E. eurystomus E. amieti 32 33 . E. tuffraui E. 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. neapolitanus b 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. harpa b 20-21 dorsolateral kineties with 20-23 dorsal cilia in central rows . E. polycarinatus SECTION B. DESCRIPTIONS OF SPECIES Euplotes aediculatus Pierson, 1943 Euplotes aediculatus was first described by Pierson (1943). Later both Tuffrau (1960) and Carter (1972) described organisms which were obviously E. aediculatus but 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 hypotrich 105-160 /Ltm long. The peristome is medium sized, triangular and has two depressions in the median border. One depression is located anteriorly whilst the other is more prominent and is situated midway along the peristomial border. The AZM collar is 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. Dorsal argyrome. (After Tuffrau, 1960.) and contains about 40 membranelles. The macronucleus is C-shaped with an arched or flattened back and the micronucleus is distinctly separate from the macronucleus. The dorsal argyrome is of the double-eurystomus type and there are typically 8 dorsolateral 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 forma tricirratus Kahl, 1932 had not been described using modern techniques. Curds (1974) redescribed a freshwater species which closely resembles the latter form and gave diagrams of the silver-line system. Tuffrau (1960) considered E. affinis to be a synonym of E. charon (Muller, 1773). DIAGNOSIS. Euplotes affinis (Fig. n) is a small (38 ^m long, 26 /mi wide) ovoid freshwater hypotrich with 9 frontoventral, 5 transverse and 3-4 caudal cirri. One of the caudal cirri is larger than the others and is held stiffly out to the right. The ventral surface is sculptured with 3 prominent ridges and the dorsal surface with 5 longitudinal ridges. The AZM has 18-20 membranelles and extends f the length of the cell. There is a small undulating membrane. The dorsal argyrome is of the double-eurystomus type with 7 dorsolateral kinetics and a maximum of 9 dorsal cilia in the central kinetics. The macronucleus is a definite 3-shape and there is a small 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 data concerning the silver-line system were added by Borror (igGSa). DIAGNOSIS. Euplotes alatus (Fig. 12) is a small (40 pm long, 30 /urn wide) oval marine species. The dorsal surface has several inconspicuous low ridges but those on the ventral surface are far more conspicuous. The AZM extends just halfway down the length of the body and is composed of approximately 26 membranelles. The dorsal argyrome is of the double-eurystomus type with 8 dorsolateral kinetics bearing 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 indentation in 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 in Africa. It has many similarities with E. eurystomus including the presence of a anterior peristomial pouch, a similar nucleus, a sigmoidal AZM which makes the present author doubt whether this is a true species or simply a geographical variety of E. eurystomus. However, because of the very large size, and more particularly the presence of 14 dorsolateral kinetics, this organism has been treated here as a separate species until more information concerning E. eurystomus and E. amieti has been 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 the ventral aspect is concave. The peristome is large, open and triangular in appearance. There is a well-developed peristomial collar and anterior pouch present. The AZM consists of 52-62 membranelles and winds sigmoidally down towards the cytostome. The dorsal argyrome is of the double-eurystomus type with 14 dorsolateral kinetics bearing 28-30 cilia in the mid-dorsal rows. There are 9 fronto ventral, 5 transverse and 4 caudal cirri. The macronucleus is an irregular 3-shape with a compact micronucleus 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 by Fenchel and Lee (1972) based on material collected in Antarctica. DIAGNOSIS. Euplotes antarcticus (Fig. 15) is a medium-sized (85 /xm long, 30 /urn wide) marine species. The shape is unlike that of other species in being very elongate 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 § of the 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 be of 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. The macronucleus is an elongate 3-shape. 22 C. R. CURDS FIG. 15. Euplotes antarcticus. a. Ventral argyrome. b. Macronucleus. c. Dorsal argyrome. (After Fenchel & Lee, 1972.) Euplotes bait cat us (Dujardin, 1841) Kahl, 1932 This species was first described by Dujardin (1841) and although the descriptions given were brief and incomplete they were just sufficient to enable Kahl (1932) to identify the species and present better diagrams. The morphology of the silver-line systems were not available until Tuffrau (1964) published his work on polymorphism in 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) in size 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 been frequently recorded in the intestinal tract of certain sea urchins (Allocentrus fragilis, FIG. 16. Euplotes balteatus. a. Ventral argyrome. b. Macronucleus. c. Dorsal argyrome. (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 AZM size which enlarges considerably when feeding upon ciliates such as Philaster sp. When feeding on bacteria there are 25-30 small membranelles but when feeding on other 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 is usually found. Tuffrau (1964) reported that there are 8 dorsolateral kinetics with up to ii cilia in the mid-dorsal rows. There are 10 frontoventral, 5 transverse and 4 or 5 caudal cirri. The macronucleus is an open C-shape when feeding upon bacteria but more horse-shoe shaped when feeding on ciliates. Euplotes bisulcatus Kahl, 1932 This species has been reported from marine sponges by Wenzel (1961) since its first description by Kahl (1932). Since that time Borror (1963) isolated it from algal growths in tidal marsh ponds and described its silver-line system. DIAGNOSIS. Euplotes bisulcatus (Fig. 17) is a small (40 /urn long, 30 /urn wide) oval marine hypotrich. The dorsal surface has prominent double-edged ridges separated by shallow grooves parallel to the dorsal ciliary rows 4, 5 and 6. The ventral surface is also ridged and the central ridge is very conspicuous. The AZM extends almost § down the length of the cell and is composed of about 17 membranelles. The dorsal argyrome is of the double-eurystomus type with 8 dorsolateral kinetics bearing 5-7 cilia in the mid-dorsal rows. There are 9 frontoventrals, 5 transverse and 3 caudal cirri. 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 descriptions vary considerably and it was not until almost a century later that Kahl (1932) gave a 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) has demonstrated 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 marine species 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 body length. The AZM bears approximately 70 strong membranelles while the AZM as a whole extends down to about f of the body length. The dorsal argyrome is of the double-eurystomus type with 12 dorsolateral kinetics carrying 35-40 dorsal cilia in the 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. The macronucleus is horseshoe shaped and there is a small compact micronucleus situated anteriorly. Euplotes crenosus Tuffrau, 1960 This species was first discovered and described by Tuffrau (1960) and has remained unmentioned since that time. THE GENUS EUPLOTES 25 DIAGNOSIS. Euplotes crenosus (Fig. 19) is a small (50-70 /Am long) freshwater oval hypotrich. There is a prominent notch at the anterior end of the cell which is coincident with a longitudinal depression on the ventral surface of the cell. The peristome is quite small and extends just over halfway down the body length. There are 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. The macronucleus is C-shaped with the micronucleus situated half-way down the left border 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 in sand samples taken from the Caspian Sea. DIAGNOSIS. Euplotes dogieli (Fig. 20) is a small (60 /mi long) marine species whose 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 the cell. There is a definite curved indentation almost midway down the left peristomial margin. The AZM is composed of 35-38 membranelles and is rather narrow. The dorsal argyrome is of the double-eurystomus type with 7 dorsolateral kinetics carrying about 13 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and 3 caudal cirri. The macronucleus is C-shaped with the anterior curve being angular and acute. The compact micronucleus is situated to the left of this anterior angular bend. 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 several other 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 comparative study 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 that was named E. aediculatus Pierson, 1943. It was not until Tuffrau (1960) introduced silver impregnation methods to Euplotes taxonomy that E. patella could readily be distinguished from E. eurystomus and E. aediculatus; unfortunately Tuffrau (1960) also introduced a considerable amount of confusion since he described E. aediculatus but called it E. eurystomus and described E. eurystomus but called it E. plumipes Stokes, 1884. Even though Pierson, Gierke and Fisher (1968) pointed out the errors of Tuffrau (1960), Carter (1972) disregarded the evidence and followed the scheme of Tuffrau (1960). Thus Carter (1972) also described E. aediculatus under the name of E. eurystomus and E. eurystomus under the name E. plumipes ; furthermore, this latter author reintroduced the species E. variabilis Stokes, 1887 which in the opinion of 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 particular emphasis upon this shape as being a distinctive and immediately visible feature. A comparison of the original diagrams in Figs. 2ia and 2ib of E. plumipes drawn by Stokes (1884) and E. patella var. eurystomus drawn by Wrzesniowski (1870) shows that 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 by Carter (1972). In the descriptions that follow E. eurystomus (Figs. 23, 24) is regarded as 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 macronucleus of 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 and prominent. The AZM is markedly sigmoid in shape and contains 50-65 membran- elles. There are 9 frontoventral, 5 transverse and commonly 4 caudal cirri although Carter (1972) found 5 caudals in one specimen. The arrangement of cirri is shown in Figs. 22, 23 and 24. The dorsal argyrome is typical of the double-eurystomus type with 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 are shown 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 concave notch which contains the micronucleus. Carter (1972) believed that the nuclear pattern, the sigmoidal AZM, the possession of an anterior peristomial pouch and a wide variation in the numbers of dorsolateral kinetics were sufficient to distinguish two strains of Euplotes from E. eurystomus and proposed the restoration of the species E. variabilis Stokes, 1884. However, a similar nuclear pattern was observed by Pierson (1943) in E. eurystomus', the sigmoidal AZM has already been stated to be a characteristic feature of E. eurystomus which species also has an anterior peristomial pouch. 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 conform well 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 with new specimens collected in 1955. DIAGNOSIS. Euplotes harpa (Fig. 25) is a large (150-160 /zm long) marine species with a more or less oval shape, although the left side is always more curved than the other. The peristome is large and open and there is a conspicuous lip on the right margin. 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 is of the double-eurystomus type with 13 dorsolateral kinetics bearing 40-45 dorsal cilia in the central rows. There are 10 fronto ventral, 5 transverse and 4 caudal cirri. 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 described by him but preparations that he made were described by Tuffrau (1960). DIAGNOSIS. Euplotes inkystans (Fig. 26) is a medium (70-80 /urn long) freshwater oval species. There is a prominent notch on the right of the peristomial collar which is narrow but rounded. The peristome is long and extends almost f down the length of the cell. The AZM is composed of approximately 40 membranelles. The dorsal argyrome is of the double-eurystomus type and there are 10 dorsolateral kineties 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 open C-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 forma latus Kahl, 1932. However, it is evident from the silver-line system that this species cannot be a form of E. patella for several reasons (compare Figs. 27 and 40) ; for example E. latus has a double-eurystomus type of dorsal argyrome whereas E. patella has 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 to date and the present author considers it sufficiently different to warrant elevating it to a species in its own right. DIAGNOSIS. Euplotes latus (Fig. 27) is a medium (70 /am long) marine species that is broadly rounded posteriorly but narrows anteriorly. The peristome is large and extends just over | down the length of the cell. The AZM is broadly curved and is composed of 35-40 membranelles. The dorsal argyrome is of the double-eurystomus type with only 6 dorsolateral kinetics bearing up to 15 dorsal cilia in the central rows. There are 9 fronto ventral, 5 transverse and 4 caudal cirri. The two centrally positioned frontoventral cirri (V2 and VI 2) are situated very close together. The macronucleus is an open angular C-shape with a micronucleus situated close to the left 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 is relatively straight for the majority of its length and then sharply curves in towards the cytostome. The dorsal surface is convex and prominently ridged. The dorsal argyrome is of the double-eurystomus type but the polygons are rather more square than is usual. There are 8 dorsolateral kinetics each containing 13-17 dorsal cilia except the left lateral one which is short and contains only 5-8 cilia. The transverse cirri 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. Dorsal argyrome. (After Carter, 1972.) cirri. The macronucleus is an irregular hoop-shape and is highly characteristic of the species, since both ends are pointed. The micronucleus is situated in a small depression 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 the Bay 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 and more truncated than the posterior. The peristome is conspicuous and extends approximately 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 n dorsolateral 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 both arms point posteriorly and the small spherical micronucleus lies in the middle of the body on the left of the macronucleus. Euplotes octocirratus Agamaliev, 1967 Agamaliev (1967) described this species from samples of sand collected from the Caspian Sea. DIAGNOSIS. Euplotes octocirratus (Fig. 30) is a small (55-60 /xm long) marine hypotrich 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 the cell. The AZM is regularly curved and is composed of about 30 membranelles. The dorsal argyrome is of the double-eurystomus type with 7 dorsolateral kinetics carrying 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 compact micronucleus in an anterior position. 30um FIG. 29. Euplotes neapolitanus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Wichterman, 1964.) FIG. 30. Euplotes octocirratus. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (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 inconspicuous low longitudinal ridges and the ventral surface has 7 minor ridges. The AZM is about f of the body length and composed of 18 membranelles. A deep pocket near the 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 the double-eurystomus type with 8 dorsolateral kinetics bearing a maximum of n cilia in the mid-dorsal rows. The ventral argyrome consists of a series of few but large polygons and resembles that of E. cristatus (see Tuffrau, 1960). The macronucleus is 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) from samples of sand collected from the Caspian Sea. DIAGNOSIS. Euplotes poljanskyi (Fig. 32) is a small to medium-sized (55-70 /mi long) marine ciliate. It has an elongated ellipsoidal shape with a medium-sized peristome 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 kinetics bearing 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 bluntly narrower 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 /mi wide) almost triangular shaped marine hypotrich. The shape of the AZM is one of its 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 the double-eurystomus type with 20 narrowly spaced dorsolateral kinetics bearing up to 23 dorsal cilia. A few specimens have been seen with 21 dorsolateral kinetics. The ventral surface is flat and there are 10 frontoventral, 5 transverse and 5-9 caudal cirri. The macronucleus is a highly irregular 3-shape with a deep involution on the upper 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 /mi wide) marine species whose general outline shape tends to be oval although there may be conspicuous wing-like extensions to the ridges associated with dorsal cilium rows i, 3 and 7. The AZM is small and extends halfway down the body length and contains 25-30 membranelles. The dorsal argyrome is of the double-eurystomus type with 9 dorsolateral kinetics carrying 13-15 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and 4 caudal cirri. The macronucleus is C-shaped and the micronucleus lies in an anterior position. C. R. CURDS FIG. 33. Euplotes polycarinatus . a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Carter, 1972.) FIG. 34. Euplotes quinquecarinatus. a. Ventral aspect, b. Nuclei, c. Dorsal aspect. (After Borror, i968a.) Euplotes trisulcatus Kahl, 1932 Kahl (1932) originally found this species in marine aquariums and gave sufficiently precise diagrams and descriptions to enable Tuffrau (1960), Borror (1963) and Carter (1972) to identify this species in other marine samples. The descriptions of the silver-line system of this species given both by Tuffrau (1960) and Carter (1972) are identical. THE GENUS EUPLOTES 37 DIAGNOSIS. Euplotes trisulcatus (Fig. 35) is a small (40 pm long, 30 pm wide) form that has been only recorded in marine habitats. The shape of the body is distinctive in that it is prominently narrower at its posterior end. There is a pronounced extension 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 is evenly curved along the outer border of the peristome and contains 25-36 membran- elles. The dorsal surface is deeply ridged showing three prominent furrows. The dorsal argyrome is of the double-eurystomus type and there are 7 dorsolateral kinetics 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-shape with angular rather than rounded ends. The micronucleus is compact and is situated anteriorly 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 the posterior 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 the body 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 dorsal argyrome 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 frontoventral cirri depending upon the degree of fusion of the infraciliary bases in rows III and IV. There are 5 transverse and 4 caudal cirri. The macronucleus is C-shaped with a 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 a b 37 a b 38 a b 39 a b 40 a b 41 a b 42 a b 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 SPECIES Euplotes apsheronicus Agamaliev, 1966 This organism was originally described by Agamaliev (1966) from specimens collected from sand samples from the Caspian Sea. A year later the same author redescribed the species (Agamaliev, 1967) with a few amendments to the original description. DIAGNOSIS. Euplotes apsheronicus (Fig. 37) is a small (50-60 /urn long) marine species 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 is composed of 30-35 membranelles. The dorsal argyrome is of the double-patella type 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, 5 transverse and 4 caudal cirri. The macronucleus is an angular and very open C-shape with a micronucleus located anteriorly. FIG. 37. Euplotes apsheronicus . a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (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 generally flattened and oval in outline. The peristome extends slightly beyond the midline of the body. The AZM consists of about 40-45 membranelles that curve smoothly to 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-shape with 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 description available is that by Carter (1972). 4o C. R. CURDS DIAGNOSIS. Euplotes octocarinatus (Fig. 39) is a medium (80 pm long, 50 wide) freshwater ellipsoid species. The peristome is triangular in shape and it extends about halfway down the length of the body. The AZM is narrow as it emerges 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 of the 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. Dorsal argyrome. (After Carter, 1972.) THE GENUS EUPLOTES 41 argyrome is of the double-patella type with 8 dorsolateral kinetics containing 18-21 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and 4 caudal cirri, 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 first described by Miiller (1773) under the name Trichoda patella and subsequently by Miiller (1786) as Kerona patella. It was called E. patella by Ehrenberg (1838) but was then named Ploesconia patella by Dujardin (1841). Several other early authors also refer to this species and much later Kahl (1932) found it necessary to subdivide the species into five forms which he named formae typicus, latus, alatus, planctonicus and variabilis (Stokes, 1887). These subdivisions were criticized by Pierson (1943) who observed that the first four of the above-mentioned forms were simply temporary varieties 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 readily be distinguished from E. eurystomus but with silver-line preparations this is now quite a simple task. DIAGNOSIS. Euplotes patella (Fig. 40) is a large (no /urn long, 65 /mi wide) oval freshwater species that has a pronounced blunt posterior end. The peristome is large, wide and almost triangular in appearance extending just over halfway down the length of the body. The AZM is narrow, evenly curved and contains 44-50 membranelles. 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 noted that Carter (1972) observed rather fewer cilia (13-19). There are 9 frontoventral, 5 strong transverse and 4 caudal cirri. The macronucleus is an open C-shape with slightly pointed extremities. The upper arm points down slightly and bears several indentations. 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 ciliate whose outline body shape is broadly rounded. The peristome is of medium size and extends about f down the body length. The AZM is regularly curved and contains 30-35 membranelles. The dorsal argyrome is of the double-patella type with 7 dorsolateral kinetics bearing up to n cilia in the mid-dorsal rows. There are 7 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 his later description (Agamaliev, 1967) (see Fig. 41). The micronucleus is small and situated anteriorly. FIG. 41. Euplotes raikovi. a. Ventral argyrome. b. Dorsal argyrome. (After Agamaliev, 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 their opinion 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 precisely with 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 by Curds et al. (1974) and that the E. moebiusi described by Curds (1974) conforms to the descriptions 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 cirrus below the AZM is stout. The ventral surface is heavily sculptured with 6 posteriorly projecting ridges. The dorsal surface has an argyrome of the double-patella type with 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 from the 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 of medium size and the right margin is essentially straight. The AZM extends about f down the body and is composed of 33-38 membranelles. The dorsal argyrome is of the double-patella type with 6 dorsolateral kinetics bearing up to 10 cilia in the mid-dorsal rows. The small polygons of the dorsal argyrome are on the left of the kinetics. There are 8 frontoventral, a unique 6 transverse and 3 caudal cirri. The macronucleus is an irregular open C-shape with a compact micronucleus situated anteriorly. 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 by Agamaliev (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 extending just over halfway down the body. The AZM contains 50-55 membranelles. The dorsal argyrome is of the double-patella type except that the narrow polygons lie FIG. 44. Euplotes zenkewitchi. a. Ventral argyrome. b. Nuclei, c. Dorsal argyrome. (After Burkovsky, 1970.) THE GENUS EUPLOTES 45 on the left of the kinetics. There are 10 dorsolateral kinetics with up to 18 dorsal cilia in the central rows. There are 9 frontoventral, 5 transverse and 3 or 4 caudal cirri. 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 cilia and 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. indentatus b 9 frontoventral cirri, 10 dorsolateral kinetics with over 30 dorsal cilia in central rows E. muscicola SECTION D. DESCRIPTIONS OF SPECIES Euplotes indentatus Carter, 1972 This species is a recent addition to the species of the genus and its description relies 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 prominent anterior notch in the upper border of the dorsal surface. The AZM is composed of 42-48 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.) 46 C. R. CURDS The dorsal argyrome is of the multiple type with three regular rows of polygons between the kinetics. There are 9 dorsolateral kinetics bearing a maximum of 15-21 cilia in the mid-dorsal rows. The macronucleus is a closed C-shape with the anterior micronucleus situated in a slight invagination. Euplotes muscicola Kahl, 1932 Although this species was intially described by Kahl (1932) and its encystment by Faure-Fremiet, Gauchery and Tuffrau (1954) a complete description was not available until Tuffrau (1960) published its silver-line system. DIAGNOSIS. Euplotes muscicola (Fig. 46) is a medium (60-70 /um long) elongate oval freshwater hypotrich. The peristome is long and occupies about f of the anterior left side of the ventral surface. The AZM contains about 35 membranelles. There are 9 frontoventral, 5 transverse and 4 caudal cirri. The dorsal argyrome is of the multiple 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 about halfway 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 species that has an elongate ellipsoidal shape. The dorsal surface is prominently sculptured with 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 short but projects forward beyond the margin of the body. The peristome is long and curved 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 there are two longitudinal fibrils between the dorsolateral fibrils. There are 6 dorsolateral kinetics with up to 15 cilia in the mid-dorsal rows. There are 9 fronto ventral, 5 transverse and 4 caudal cirri. The macronucleus is highly angular in appearance ; it is composed of two arms joined anteriorly at an acute angle. The micronucleus is very large for the genus (7-8 ^m diameter) and is situated anteriorly on the left of 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 Section A, p. 14.) 47 a 7 dorsolateral kinetics with 11-15 cma m mid-dorsal rows . . . E. gracilis b 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. muscorum b 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 SPECIES Euplotes elegans Kahl, 1932 This species was originally described briefly by Kahl (1932) and then redescribed in 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 this differs 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 extends about | 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. The original description by Kahl (1932) appears to be the only observation of the presence of 3 caudal cirri. The dorsal argyrome is of the complex type consisting of many irregular polygons. There are 8 dorsolateral kinetics with a maximum of 30-46 dorsal cilia in the central kinetics. The left lateral kinety is short with only 4-8 cilia 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 by Tuffrau (1960). DIAGNOSIS. Euplotes gracilis (Fig. 49) is a small (37-50 /xm long) freshwater species 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 cirri are very long and styliform whereas the caudals are thin. The dorsal argyrome is of a complex nature consisting of an irregular assemblage of polygons between the 7 dorsolateral kinetics. There are 10-13 large kinetosomes in the mid-dorsal kine- tics. The macronucleus is C-shaped and the micronucleus is situated approximately £ of 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 incomplete descriptions of Kahl (1932). Borror (1963) described an organism as E. moebiusi but 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 not sufficiently comprehensive for identification purposes they do conform with the silver-line systems described by Curds (1974). Tuffrau (1960) considered E. moebiusi to 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 with 5 longitudinal ridges. The AZM is composed of 35-40 membranelles which extend | the length of the cell. The dorsal argyrome is unique amongst those species described to date ; there are 5 longitudinal rows of narrow polygons interspersed with an irregular network of larger polygons. The presence of the irregular network has 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 bearing a maximum of n cilia in the mid-dorsal rows. The macronucleus is 3-shaped and the 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 membranelles and extends about f down the body. There are 9 frontoventral, 5 transverse and 4 long caudal cirri. The dorsal argyrome is of the complex type with many polygons arranged irregularly between the kinetics. There are 8 dorsolateral kinetics carrying 22-28 cilia in the mid-dorsal rows on the peaks of the longitudinal ridges. The macronucleus is C-shaped with the pointed right arm projecting posteriorly over halfway down the length of the body. The micronucleus is situated anteriorly almost 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 yet been 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) from samples of marine sands but for several reasons may be regarded as a distinct species until further data are gathered. DIAGNOSIS. Euplotes aberrans (Fig. 52) is a medium (70-80 /mi long) marine ciliate whose shape is highly elongate. There is a distinct spike-like projection on the right of the peristomial collar. The dorsal surface bears 4 pronounced ridges and there are 2 major ones, amongst others, on the ventral surface which travel slightly trans versally almost the entire length of the body. The peristome is long and narrow 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 shaped macronuclei) . There are 5 transverse cirri in two groups and 4 caudals. There is a very large subequatorial vacuole which has not yet been seen to contract. The macronucleus 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 of the 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. Dorsal aspect. (After Wang, 1930.) THE GENUS EUPLOTES 53 3 less prominent ones are restricted to the ventral surface. The anterior end of the body 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 is C-shaped. Euplotes roscoffensis Dragesco, 1966 Although this species was not described until 1966 it was in fact first studied in 1950 long before the importance of silver preparations for Euplotes taxonomy was realized. However, because the right-hand border of the peristome is so distinctive this 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 reaches the level of the undulating membrane just below the middle of the cell, here the peristome border bears a distinctive deep invagination or pocket (Fig. 53) which appears 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 contractile vacuole which is situated in the proximity of transverse cirrus VI i often has several small satellite vacuoles surrounding it. The macronucleus is C-shaped with a club- like structure at the posterior end. The micronucleus is small, round and situated close 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 description of Gelei (1950). 54 C. R. CURDS DIAGNOSIS. Euplotes rotunda (Fig. 55) is a small (50 /mi long, 40 /urn wide) round freshwater species. The peristome extends down to about the centre of the body and the AZM carries approximately 25 membranelles. There are 8 dorsolateral kinetics with about 10 cilia in the mid-dorsal rows. There are 9 frontoventral, 5 transverse and 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 in samples of moss. Tuffrau (1960) was unconvinced that this organism is in fact a member 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°ng) freshwater species that is generally oval in outline but tends to narrow posteriorly to a blunt point. The peristome is relatively short and extends just to the centre of the body and there appear to be somewhere in the order of 20 membranelles in the AZM. The dorsal surface seems to have 6 ridges and there are 9 frontoventral and 5 transverse cirri. According to Kahl (1932), there are 10 caudal cirri although Penard (1922) originally described these as being very fine marginal cirri. The macronucleus is a simple hoop-shape but the micronucleus is illustrated as lying inside the curve of the macronucleus which is an unusual feature for the genus. It is 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 by Dragesco (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 projecting peristomial collar. The dorsal surface carries 12 longitudinal furrows and there appear 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 the peristome border beneath the peristomial collar. There are 5 transverse and 4 caudal cirri. A group of 3 contractile vacuoles is positioned close to the transverse cirri. The macronucleus is an open C-shape with the micronucleus in an anterior position. 0 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 macronucleus enables it to be easily recognized. DIAGNOSIS. Euplotes woodruffi (Fig. 58) is a large (145 p,m long) euryhaline oval species with a well-defined peristomial collar. The peristome is large, triangular in shape and extends about f of the way down the body. The AZM is composed of 60-70 membranelles. The dorsal surface is sculptured with 8-10 grooves and there are 8 dorsolateral kinetics with about 60 closely set cilia in the central rows. There are 9 frontoventral, 5 transverse and 4 caudal cirri. The unique macronucleus is T- 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, 1957 Euplotes eurystomus Tuffrau, 1960 Euplotes eurystomus Carter, 1972 3. Euplotes affinis (Dujardin, 1841) Kahl, 1932 18 Ploesconia affinis Dujardin, 1841 Ploesconia subrotundus Dujardin, 1841 Euplotes subrotundus Perty, 1852 Euplotes 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, 1773 Ploesconia charon (Muller, 1773) Bory, 1826 Euploea charon Ehrenberg, 1830 Euplotes appendiculatus Ehrenberg, 1838 Ploesconia charon Dujardin, 1841 Ploesconia radiosa Dujardin, 1841 Ploesconia longiremus Dujardin, 1841 Euplotes balteatus Burkovsky, 1970 12. Euplotes crassus (Dujardin, 1841) Kahl, 1932 ii Ploesconia crassa Dujardin, 1841 Euplotes taylori Garnjobst, 1928 Euplotes violaceus Kahl, 1928 Euplotes salina Yocum, 1930 Euplotes 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, 1786 Euplotes plumipes Stokes, 1884 Euplotes variabilis Stokes, 1887 Euplotes patella var. eurystomus Wrzes- niowski, 1870 Uronychia paupera Daday, 1907 Euplotes patella forma variabilis Kahl, 1932 Euplotes plumipes Tuffrau, 1960 Euplotes plumipes Carter, 1972 Euplotes 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, 1773 Kerona patella Muller, 1786 Coccudina keronina Bory, 1826 Himantopus charon Ehrenberg, 1833 Ploesconia patella Dujardin, 1841 Euplotes charon var. marina Quenner- stedt, 1867 Euplotes paradoxa Kent, 1880 Euplotes carinatus Stokes, 1885 Euplotes patella var. alatus Kahl, 1932 Euplotes 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 42 Euplotes 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, 1786 Ploesconia vannus (Muller, 1786) Bory, 1826 Euplotes striatus Ehrenberg, 1838 Euplotes longipes Claparede and Lach- mann, 1858 Euplotes extensus Fresenius, 1865 Euplotes gabrieli Gourret and Roeser, 1886 Euplotes Worcester i Griffin, 1910 Euplotes caudatus Meunier, 1910 Euplotes truncatus Meunier, 1910 Euplotes 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 a silver-line preparation of E. woodruffi has been published (Magagnini & Nobili, 1964). It is of the double-eurystomus type with 8 dorsolateral kinetics and 25 middorsal cilia. In the existing key these characters would lead to E. aediculalus and N. eurystomus. All three species may be easily distinguished by their nuclear features. REFERENCES AGAMALIEV, F. G. 1966. New species of psammobiotic ciliates of the western part of the Caspian Sea. Acta Protozool. 4 : 169-183. — 1967. Faune des cilies mesopsammiques de la cote ouest de la Mer Caspienne. Cahiers de Biol Mar. 8 : 359-402. — 1972. Ciliates from microbenthos of the islands of ApSeroskij and Bakinskij archipelagos of the Caspian Sea. Acta Protozool. 10 : 1-27. BERGER, J. 1965. The infraciliary morphology of Euplotes tuffraui n. sp., commensal in Strongylocentrotid echinoids, with comments on echinophilous populations of Euplotes balteatus (Dujardin) Ciliata, Hypotrichida. Protistologica 1 : 17-31. BONNER, J. T. 1954. The development of cirri and bristles during binary fission in the ciliate Euplotes eurystomus. J . Morph. 95 : 95-108. BORROR, A. C. 1962. Euplotes minuta Yocum (Ciliophora, Hypotrichida). /. Protozool. 9 : 271-273. — 1963. Morphology and ecology of the benthic ciliated protozoa of Alligator Harbour, Florida. Arch. Protistenk. 106 : 465-534. — ig68a. Systematics of Euplotes (Ciliophora, Hypotrichia) ; toward union of the old and the new. /. Protozool. 15 : 802-808. — ig68b. Nigrosin-HgCl2-formalin ; a stain fixative for ciliates (Protozoa, Ciliophora). Stain Technol. 43 : 293-294. — 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Protozool. 19 : 1-23. BORY, DE ST VINCENT 1826. Essai d'une classification des animaux microscopiques. Paris. io4pp. BOVEE, E. C. 1957. Euplotes leticiensis n. sp. from the Letician drainage into the Amazon river. /. Protozool. 4 : 124-128. THE GENUS EUPLOTES 59 BURKOVSKY, I. V. 1970. The ciliates of the mesopsammon of the Kandalaksha Gulf (White Sea). A eta Protozool. 7 : 475-489. CARTER, H. P. 1972. Infraciliature of eleven species of the genus Euplotes. Trans. Amer. micros. Soc. 91 : 466-492. CHATTON, E. & LWOFF, A. 1930. Impregnation, par diffusion argentique, de 1'infraciliature des dies marins et d'eau douce, apres fixation cytologique et sans dessication. C. r. Seanc. Soc. Biol. 104 : 834-836. — & SEGUELA, J. 1940. La continuite genetique des formations ciliaires chez les cilies hypotiches. Le cinetome et 1'argyrome au cours de la division. Bull. biol. Fr. Belg. 74 : 349-442. CLAPAREDE, E. & LACHMANN, J. 1858. Etudes sur les infusoires et les rhizopodes. Mem. Inst. nat. Genevois, 5 : 1-260 ; 7 : 1-291. CURDS, C. R. 1974. Description of three species of Euplotes (Protozoa, Ciliatea). Bull. Br. Mus. nat. Hist. (Zool.), 27, 113-125. & COCKBURN, A. 1971. Continuous monoxenic culture of Tetrahymena pyriformis. J. gen. Microbiol. 66 : 95-108. WEST, B. J. & DORAHY, J. E. 1974. Euplotes rariseta sp. n. (Protozoa, Ciliatea), a new small marine hypotrich. Bull. Br. Mus. nat. Hist. (Zool.), 27 : 95-102. DADAY, J. 1907. Adatok Nemet-Kelet-Afrika edesvizi mikrofaunajanak ismeretehez. Mat. termeszettud. Ert. 25 : 402-420. DILLER, W. F. & KOUNARIS, D. 1966. Description of a zoochlorella-bearing form of Euplotes, E. diadaleos n . sp. (Ciliophora, Hypotrichida). Biol. Bull. mar. biol. Lab. Woods Hole, 131 : 437-445- DRAGESCO, J. 1960. Cilies mesopsammiques littoraux. Trav. Stn biol. Roscoff, 12 : 1-356. 1966. Observations sur quelques cilies libres. Arch. Protistenk. 109 : 155-206. — 1970. Cites libres du Cameroun. Annales de la Faculte des Sciences de Yaounde. Numero Hors-Serie. Annls Fac. Sa. Univ. Cameroun 1970, 1-141. (Hors-serie.) DUJARDIN, F. 1841. Histoire Naturelle des Zoophytes. Infusoires. Paris. EHREXBERG, C. G. 1830. Beitrage zur Kenntniss der Organisation der Infusorien und ihrer geographischen Verbreitung, besonders in Sibirien. Abh. dt. Akad. Wiss. Berl. yr 1830 : 1-89. - 1833. Dritter Beitrage zur Erkenntniss grosser Organisation in der Richtung des kleinsten Raumes. Abh. dt. Akad. Wiss. Berl. yr 1833 : 145-336. 1838. Die Infusionsthierchen als vollkommene Organismen. L. Voss, Leipzig. 6i2pp. — 1840. Diagnosen von 274 neuen Infusorien. Mber. K.preuss. Akad. Wiss. yriS^o : 197-219. FAURE-FREMIET, E., GAUCHERY, M. & TUFFRAU, M. 1954. Les processus d'encystement chez Euplotes muscicola Kahl. Bull. biol. Fr. Belg. 88 : 154-167. FENCHEL, T. & LEE, C. C. 1972. Studies on ciliates associated with the sea ice from Antarctica. I. The nature of the fauna. Arch. Protistenk. 114 : 231-236. FRANKEL, J. 1973- The positioning of ciliary organelles in hypotrich ciliates. /. Protozool. 20:8-i8. FRESENIUS, G. 1865. Die Infusiorien des Seewasseraquariums. Zool. Gart., Frank/. 6 : 81-89, 121-129. GALL, J. G. 1959. Macronuclear duplication in the ciliated protozoan Euplotes. J. biophys. biochem. Cytol. 5 : 295-308. GARNJOBST, L. 1928. Induced encystment and excystment in Euplotes taylori sp. nov. Physiol. Zool. 1 : 561-575. GAW, H. Z. 1939. Euplotes woodruffi sp. nov. Arch. Protistenk. 93 : 1-5. GELEI, J. 1950. Uj Euplotesek a tisza vizrenszerebol. Annls biol. Univ. szeged, 1 : 241-247. GIESE, A. C. 1938. Cannibalism and gigantism in Blepharisma. Trans. Am. microsc. Soc. 57: 245-255. GOURRET, P. & ROESER, P. 1 886. Les protozoaires du vieux-port de Marseille. Archs Zool. exp. gen. 4 : 443-534- 60 C. R. CURDS GRIFFIN, L. E. 1910. Euplotes worcesteri sp. nov. i. Structure. Philipp.J. Sci. 5 : 291-312. HECKMANN, K. 1963. Paarungssystem und genabhangige Paarungstypdifferenzierung bei dem hypotrichen Ciliaten Euplotes vannus O. F. Muller. Arch. Protistenk. 106 : 393-421. HUFNAGEL, L. A. & TORCH, R. 1967. Intraclonal dimorphism of caudal cirri in Euplotes vannus : cortical determination. /. Protozool. 14 : 429-439. KAHL, A. 1928. Die Infusorien der Oldesloer Salzwasserstellen. Arch. Hydrobiol. 19 : 50-123, 189-246. 1932. Urtiere oder Protozoa. I : Wimpertiere oder Ciliata (Infusoria), eine Bearbietung der freilebenden und ectocommensalen Infusorien der Erde, unter Ausschluss der marinen Tintinnidae. In Dahl, F., Die Tierwelt Deutschlands, Teil 25, 399-650. G. Fischer, Jena. KENT, W. S. 1880-1882. A Manual of the Infusoria. Vols I-III. David Bogue, London. 9I3PP- KLEIN, B. M. 1958. The 'dry' silver method and its proper use. /. Protozool. 5 : 99-103. KLUSS, B. C. 1962. Electron microscopy of the macronucleus of Euplotes eurystomus. J . Cell. Biol. 13 : 462-465. MAGAGNINI, G. & NOBILI, R. 1964. Euplotes woodruffi Gaw e su Euplotidium arenarium n. sp. (Ciliatea, hypotrichida) . Monitore Zool. ital. 72 : 178-202. MEUNIER, A. 1910. Microplancton des mers de Berents et de Kara. In Due d' Orleans Campagne Ar clique de 1907. Bulen, Brussels. MINKJEWICZ, R. 1901. fitude sur les Protozoaires de la Mer Noire ; I. L'organisation, la multiplication et la position systematique du genre Euplotes Ehrbg. Trudy Obshch. Estest. imp. kazan. Univ. 35 : 1-67. MULLER, O. F. 1773. Vermium Terrestrum et Fluviatilum, seu Animalium Infusorium, Helminthicorum et Testaceorum, non Marinorum, Succincta Historia. Havniae et Lipsiae. I35PP- 1786. Animalcula Infusoria Fluviatilia et Marina. Havniae et Lipsiae. 367pp. PENARD, E. 1922. Etudes sur les infusoires d'eau douce. George et Cie, Geneve. 33ipp. PERTY, M. 1852. Zur Kentniss kleinster Lebenformen nach Bau, Funktionen, Systematik, mil Specialverzeichniss der in der Schweiz beobachteten. Jent und Reinert, Bern. 228pp. PIERSON, B. F. 1943. A comparative morphological study of several species of Euplotes closely related to Euplotes patella. J. Morph. 72 : 125-151. — GIERKE, R. & FISHER, A. L. 1968. Clarification of the taxonomic identification of Euplotes eurystomus Kahl and E. aediculatus Pierson. Trans. Am. microsc. Soc. 37 : 306-316. PRESCOTT, D. M., KIMBALL, R. F. & CARRIER, R. F. 1962. Comparison between the timing of micronuclear and macronuclear DNA synthesis in Euplotes eurystomus. J. Cell. Biol. 13 : 175-176. PROWAZEK, S. VON. 1902. Protozoenstudien. Ill, Euplotes harpa. Arb. zool. Inst. Univ. Wien. 14:8i-88. QUENNERSTEDT, A. 1867. Bidag till Sveriges infusorie-fauna. Ada Univ. lund. 4 : 1-47. RUINEN, J. 1938. Notizen iiber Ciliaten aus konzentrierten Salzgewassern. Zool. Meded. 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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9-70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28, No. 2, of the Zoological series. The abbreviated titles of periodicals cited follow those of 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 OF THE 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 Museum collection of types with the exception of the pseudotracheate group which are listed in Part I of the type-catalogue (Lincoln & Ellis, 1974). Part II comprises 429 separate 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 the manuscript names referred to in the Appendix has been deposited in the library of the British Museum (Natural History). In total, this impressive collection of woodlice type material has over noo registered entries representing 767 species in 153 different genera, and is thus of considerable significance in a group which has only about 300 genera and rather more 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 by C. 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. December 1897 '• '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. Collected by 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 STYLONISCIDAE CLAVIGERONISCUS 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'. Collected by 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 by P. 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. Lawrence and 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 by the Royal Society. SYNTYPES : four females. Reg. no. 1970:347:4. Fulakora Pt, Raja, Santa Isabel, Solomon Is. 30.9.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by the Royal Society. 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 the Royal 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. of Santa 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 the Royal 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. Mt Gallago, 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, nr Mt Gallego, c. 6 miles S.W. Tamboko, Guadalcanal, Solomon Is. 4.7.1965 ; 'disturbed forest litter'. 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 the Royal 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 by P. 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. Norman Collection (ex Patience Collection). stebbingi (Patience) (i9O7b : 42) [Trichoniscus stebbingi] SYNTYPES: two males; three females. Reg. no. 1907:5:29:1-5. Springburn Public Park, 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. Collected by 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 by P. 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 by the 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, Cape Province, 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, Cape Province, 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. Hottentots Holland 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. Collected and 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, Cape Province, 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. Jonkershoek Mts, 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. Presented by 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 by K. 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 TITANIIDAE KOGMANIA 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 SCHOBLIIDAE SCHOBLIA Budde-Lund circular is Budde-Lund (1909 : 66) SYNTYPE : few fragments only. Reg. no. 1921:10:18:451. East Africa. Budde-Lund Collection. Family TRIGHONISGIDAE ALPIONISCUS Racovitza fragilis Budde-Lund (1909 : 68) SYNTYPES : two males. Reg. no. 1921:10:18:571-572. Sardinia. Budde-Lund Collection. 7o 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. Collected by 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. Verhoeff Collection. 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. Verhoeff Collection. dentiger Verhoeff (igoSc : 139, 143) SYNTYPES : one male ; one female. Reg. no. 1908:6:1:33-34. Italian Riviera. Verhoeff Collection. 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. Verhoeff Collection. 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. Verhoeff Collection. 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. Verhoeff Collection. 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 Collection danicus 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 Dollfus Collection) . SYNTYPES : two females. Reg. no. 1921:10:18:393-395. Lake Lentini, Sicily. Collected by 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. Verhoeff Collection. 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. Verhoeff Collection. 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. Old St Michael's Cave, Gibraltar. 6.3.1951 ; 'rotten wood; 650 ft from entry'. Collected by T. 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 vividus Budde-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-Lund Collection (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 : 246) [Now Spelaeonethes medius (Carl)] SYNTYPES: three males; five females. Reg. no. 1921:10:18:580-587. Pyrenees, 'in caves'. 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. Verhoeff Collection. 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. Norman Collection (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 (ex Verhoeff Collection). Family BUDDELUNDIELLIDAE BUDDELUNDIELLA Silvestri cater actae Verhoeff (i93ob : 30) SYNTYPE : one specimen. Collection. Reg. no. 1930:5:26:41. Dalmatia, Yugoslavia. Verhoeff Family LIGIIDAE EURYLIGIA 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-Lund Collection. 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. September 1896. 2000 ft. Collected by Perkins. Presented by the Joint Committee of the Royal Society and British Association for Investigating the Fauna of the Sandwich Is. per Dr David Sharp. 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 the Joint Committee of the Royal Society and British Association for Investigating the Fauna of the Sandwich Is. per Dr David Sharp. pigtnentata Jackson (1922 : 699) SYNTYPES: one male (?) ; one female. Reg. no. 1921:10:18:131. Suez. Budde-Lund Collection. 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. Verhoeff Collection. latum Jackson (1923 : 834) SYNTYPES: five females. Reg. no. 1921:10:18:168-174. San Francisco, USA. Collected by 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 STENONISCIDAE PARASTENONISCUS 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. Verhoeff Collection. 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'). Verhoeff Collection. Family RHYSCOTIDAE RHYSCOTOIDES 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 by K. 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. Possession I. 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, Cape Province, 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. Collected by R. H. Stevenson. Barnard Collection. microps Barnard (1932 : 271) SYNTYPES: four females. Reg. no. 1933:1:25:192-194. Maxixe, Mosambique. Collected by 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. Verhoeff Collection. costulatus Verhoeff (igoSb : 179) SYNTYPES: one male ; two females. Reg. no. 1908:6:1:19-21. Italian Riviera. Verhoeff Collection. schobli Budde-Lund (1885 : 200) SYNTYPES : three females. Reg. no. 1911:11:8:10837-39. Bona, Algeria. Collected by Meinert. 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., Cargados Carajos, 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., Cargados Carajos, 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 ONISCIDAE ALLONISCUS 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 : I24) 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 of Madagascar. 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-Lund Collection. 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, Lake Tanganyika, 'under stones'. Collected by W. A. Cunnington. Presented by the Tanganyika Exploration 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 Tanganyika Exploration 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 by Drege. 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. Larwood Collection (ex Verhoeff Collection). littoralis Budde-Lund (1885 : 237) SYNTYPES: one male; one female. Reg. no. 1921:10:18:1057-1058. Venetias. Collected by Schaufuss. Budde-Lund Collection. BATHYTROPA Budde-Lund granulata Aubert & Dollfus (1890 : 9) SYNTYPE : female. Reg. no. 1911:11:8:10549. Marseille, France. Norman Collection (ex Dollfus Collection). hispana Dollfus (1893 : 5°) [Now Haplophthalmus danicus Budde-Lund] SYNTYPE: female. Reg. no. 1911:11:8:10546. Valencia, Spain. Norman Collection (ex Dollfus 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 by Meinert. 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 the Royal Society. SYNTYPES : one male ; one female. Reg. no. 1970:415:1. Fulakora Pt, Raja, Santa Isabel, Solomon Is. 30.9.1965; 'forest litter'. Collected by P. N. Lawrence. Presented by the 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 the Royal Society. SYNTYPES : four males ; five females. Reg. no. 1970:417:13 (part). Tatamba, Santa Isabel, Solomon Is. 4.10.1965 ; 'mangrove litter'. Collected by P. N. Lawrence. Presented by 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 the Royal 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 the Royal 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 the Royal Society. SYNTYPES : one male ; four females. Reg. no. 1970:419:4. E. Central San Jorge, Santa Isabel, 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. Lunga Beach, 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'. Collected by 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, East Madagascar. 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. Botanic Gardens, 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. Verhoeff Collection. 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. Norman Collection (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., Indian Ocean. 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 South Wales. 30.12.1883 ; 'in rock pools'. Collected by C. Chilton. Budde-Lund Collection (ex Dundee 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. Collected by 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. Verhoeff Collection. SYNTYPES : two males ; three females. Reg. no. 1937:7:6:100-104. Riviera. Verhoeff Collection. 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, Langeberg Range, 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, opposite Lilihinia 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. 84 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. Presented by 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. Verhoeff Collection. SYNTYPES : three females. Reg. no. 1937:7:6:121-123. Dolomite Mts. Verhoeff Collection. 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'. Collected by 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 the Royal 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 the Royal 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 by the 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 Royal Society. 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 Royal Society. 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 by P. N. Lawrence. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:411:1. Mt Austern, nr Honiara, Guadalcanal, Solomon Is. 24.8.1965 ; 'forest litter'. Collected by P. N. Lawrence. Presented by the Royal Society. 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. Presented by the Royal Society. SYNTYPES : two males ; one female. Reg. no. 1970:413:3. 7 mis S. of Wainoni, San Cristobal, 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. Collected by 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. Stanley Gardiner. 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, River Umasami, Guadalcanal, Solomon Is. 7.7.1965. 1000 ft ; 'ridge, forest litter'. Collected by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : three males ; two females. Reg. no. 1970:400:20 (part). S.W. end of Monitor Creek, River Umasami, Guadalcanal, Solomon Is. 5.7.1965 ; 'forest litter in a hollow'. 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 Royal Society. 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 Royal Society. 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 Royal Society. SYNTYPES : one male ; three females. Reg. no. 1970:387:22 (part). 4 miles up Warahito from River Pagato, San Cristobal, Solomon Is. 4.8.1965 ; 'forest litter'. Collected by Leone and 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 by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : two females. Reg. no. 1970:388:6 (part), c. 6-75 miles S. Wainoni, San Cristobal, 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 Royal Society. SYNTYPES : one male; two females. Reg. no. 1970:388:6 (part), nr Wainoni, San Cristobal, 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 by P. 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 by P. 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 the Royal 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 the Royal 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 the Royal 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 the Royal 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 Fernandez I., 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. Verhoeff Collection. 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. Verhoeff Collection. 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, nr Pietermaritzburg, 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, August 1891. Collected by F. Meinert. Budde-Lund Collection. flava Budde-Lund (igisa : 70) HOLOTYPE : female. Reg. no. 1921:10:18:1878. Victoria, Australia. Collected by Hauschild. 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-Lund Collection. 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, nr Durban, 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. Stanley Gardiner. 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. October 1894. 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. Presented by 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, nr Honiara, 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 by P. 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 ; 'forest litter'. 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 the Royal 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 Royal Society. 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. Presented by 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 the Royal 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 miles S.W. Tamboko, Guadalcanal, Solomon Is. 1.7.1965 ; 'forest litter, sandy soil'. Collected by P. 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 by P. N. Lawrence and Isiah. Presented by the Royal Society. SYNTYPES : five males ; six females ; six juveniles. Reg. no. 1970:375:18. Huni River Estuary, N.E. Wainoni, San Cristobal, Solomon Is. 9-12.8.1965 ; 'forest litter on coral limestone'. 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. Presented by the Royal Society. SYNTYPE : female. Reg. no. 1970:377:29 (part). 4 miles up Warahito River, from Pagato River, 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, San Cristobal, Solomon Is. 1.8.1965 ; 'stream litter'. Collected by P. N. Lawrence. Presented by 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 ; 'rotten wood 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 by P. N. Lawrence. Presented by the Royal Society. SYNTYPES : twenty specimens : males, females and juveniles. Reg. no. 1970:379:22. nr Kuzi, 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. Presented by 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. January 1887. 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., Chagos Archipelago, 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-Lund Collection. 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 Dollfus Collection) . truncatella (Budde-Lund) (1902 : 379) [Philoscia truncatella] SYNTYPE : few fragments only. Reg. no. 1921:10:18:2383. Malay Peninsula. Collected by 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. Verhoeff Collection. 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. Collected by 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. Verhoeff Collection. 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: 34J-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. 379-381- 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 Diagnosen verwandter 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 Ergebnisse einer 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 the leadership of Mr. J. Stanley Gardiner. IV. No. XXII. Terrestrial Isopoda, particularly considered 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 a la 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 the Royal 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, a sub-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). Archs Zool. 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 Labyrinth Kretas. 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, mostly from the Budde-Lund Collection, for which no reference can be found in published literature. It seems possible that these are specimens with manuscript names which have been incorporated into personal collections but which have never been published. A list of this material comprising some 68 items is given below, and a list of 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 by Dr Plate. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 97 Styloniscus sp. Two specimens. Reg. no. 1921:10:18:420-421. Auckland, New Zealand. Budde-Lund Collection. 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 i Species 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 2 Species a One specimen. Reg. no. 1931:4:27:71. Georgia. Verhoeff Collection. Genus 3 Species a One specimen. Reg. no. 1921:10:18:793. No locality. Budde-Lund Collection. Genus 4 Species a Two specimens. Reg. no. 1921:10:18:5753-5754. Ralum. 27.2.1897. Budde-Lund Collec- tion. INDEX abbreviatus (Haplophthalmus) 71 adonis (Hyloniscus) 71 adriatica (Halophiloscia) 82 adriatica rupium (Halophiloscia) 82 affinis (Philoscia) 87 affinis (Philoscia muscorum) 87 alba (Pherusa) 73 albicincta (Bilawrencia) 80 albus (Titanethes) 73 Alloniscus 78, 97 alpinus (Androniscus) 70 Alpioniscus 69 Anchiphiloscia 79 Androniscus 70 angusta (Niambia) 77 angusticauda (Philoscia) 91 angusticauda (Setaphora) 91 angustissima (Pseudophiloscia) 89 anomala (Philoscia) 87 anomalus (Phalloniscus) 87 Aphiloscia 79, 98 apuanus (Haplophthalmus) 71 Armadilloniscus 79, 98 armata (Deto) 81 armata (Paraphiloscia) 85 australis (Styloniscus) 68 australis (Trichoniscus) 68 austriacus (Trichoniscus) 73 austroafricanus (Styloniscus) 68 austroafricanus (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) 93 brunnea (Niambia) 77 Buddelundiella 74 BUDDELUNDIELLIDAE 74 bulgarica (Bureschia) 70 Bureschia 70 Burmoniscus 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) 75 Cyphonetes 73 Cyphoniscellus 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) 81 Formososcia 82 foveolatus (Trichoniscus) 74 fragilis (Alpioniscus) 69 fragilis (Pseudophiloscia) 89 fragilis rharelbazi (Trichoniscus) 73 frigidana (Philoscia muscorum) 88 fucorum (Halophiloscia) 82 fulva (Philoscia (Benthanops)) 88 georgensis (Styloniscus) 68 georgensis (Trichoniscus) 68 germanicus carniolense (Lepidoniscus) 84 glarearum (Stenophiloscia) 92 globiceps (Rhyscotus) 76 gottscheensis (Cyphoniscellus) 71 gracilicornis (Halophiloscia) 82 gracilipes (Ligia) 75 graecus (Labyrinthasius) 83 granulata (Bathytropa) 79, 83 granulatus (Tylos) 65 granuliferus (Tylos) 65 gravosensis (Philoscia) 88 griseoflavus (Niambia) 77 griseus (Hiatoniscus) 83 guernei (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)) 70 heroldi (Isabelloscia) 83 herzegowinense (Ligidium) 75 Hiatoniscus 83 hirsuta (Halophiloscia) 82 hirsuta (Nahia) 84, 88 hirsuta (Niambia) 77 hirsuta (Philoscia) 84 hispana (Bathytropa) 79 holthuisi (Papuasoniscus) 85 Hora 83 horae (Styloniscus) 68 horae (Trichoniscus) 68 hottentoti (Styloniscus) 68 hottentoti (Trichoniscus) 68 humilis (Didima) 82 Hyloniscus 71, 97 hypnorum (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) 79 kempi (Burmoniscus) 80 kenepurensis (Oniscus) 84 kermadecensis (Styloniscus) 68 kermadecensis (Trichoniscus) 68 Kogmania 69 Komatia 89 Krantzia 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) 74 novabritannica (Rennelloscia) 89 nudulus (Tylos) 65 ocellata (Formososcia) 82 Olibrinus 84 ONISCIDAE 78, 97 Oniscus 84 opercularis (Tylos) 66 orientalis (Indoniscus) 67 Oritoniscus 72 ornatus (Paranotoniscus) 68 ornatus (Scyphax) 91 Oroniscus 84 ortonedae (Rhyscotoides) 76 ortonedae (Rhyscotus) 76 otakensis (Styloniscus) 69 otakensis (Trichoniscus) 69 ovata (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, 97 squamuligera bargensis (Tiroloscia) 93 squamuligera briani (Philoscia (Tiroloscia)) 93 squamuligera briani (Tiroloscia) 93 squamuligera tendana (Tiroloscia) 93 stammeri (Trichoniscus) 74 stebbingi (Cordioniscus) 67 stebbingi (Trichoniscus) 67 STENONISCIDAE 76 Stenophiloscia 92 strasseri (Alpioniscus (Illyrionethes)) 70 strasseri (Androniscus cavernarum) 70 sturanus (Trichoniscus noricus) 74 Stylohylea 72 STYLONISCIDAE 66, 97 Styloniscus 68, 97 styricus (Cyphoniscellus) 71 suarezia (Philoscia) 92 suarezia (Setaphora) 79, 92 subterranea (Philoscia) 88 subterraneus medius (Androniscus) 70 subterraneus noduliger (Androniscus) 70 subterraneus scaber (Androniscus) 70 swellendami (Styloniscus) 69 swellendami (Trichoniscus) 69 tabulae (Styloniscus) 69 tabulae (Trichoniscus) 69 tendana (Tiroloscia squamuligera) 93 Tendosphaera 76 TENDOSPHAERIDAE 76 thermophila (Bathytropa) 79 Tiroloscia 92 Titanethes 73 TITANIIDAE 69 tomentosa (Trichorhina) 79 triangulifera (Philoscia muscorum) 88 TRICHONISCIDAE 69, 97 Trichoniscoides 73 Trichoniscus 73, 97 Trichorhina 78 truncata (Niambia) 77 truncatella (Philoscia) 92 truncatella (Setaphora) 92 tuberculatus (Hanoniscus) 82 tuberculatus (Paranotoniscus) 68 tuberculatus (Phymatoniscus) 72 tuberculatus (Trichoniscoides) 72 TYLIDAE 65, 96 Tylos 65, 96 tyrrhena (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) 91 warreni (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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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^^iw C. 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, BRACHYI PORTUNIDAE) REARED IN THE LABORATORY BY A. L. RICE - -£_ Institute of Oceonographic Sciences AND 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28, No. 3, of the Zoological series. The abbreviated titles of periodicals cited follow those of 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 OF THE 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 waters are 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 Carcinus have recently been separated as distinct species on the basis of adult characters, the name C. maenas (L.) being applied to the Atlantic form and C. mediterraneus Czerniavsky to the Mediterranean form. Although the larval differences support the possibility that the two populations are genetically distinct, if, as presently seems to be the case, they are found to be totally allopatric with no chance of interbreeding in areas of overlap, it is suggested that they should 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-western Europe were better known than those of any other region, mainly as a result of Lebour'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 detailed accounts 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 and providing detailed descriptions of all their developmental stages. As a result of this programme descriptions of the sponge crab Dromia personata (L.), and of the masked crab Corystes cassivelaunus (Pennant), have already been published (Rice, Ingle & Allen, 1970 ; Ingle & Rice, 1971) and although crabs of any of the fourteen families represented in British waters will be reared and described as and when ovigerous females become available, efforts are now being concentrated on the swimming crab family Portunidae and the spider crab family Majidae and the programme has been expanded to include the rearing of Mediterranean species belonging to these families. Apart from the spider crabs (Majidae) the portunids are better represented around British coasts than any other family of crabs, fourteen species and five genera having been recorded. This paper deals with the commonest of these species, Carcinus maenas (L.) and with the closely related Mediterranean form, C. mediterraneus Czerniavsky. The common shore crab or green crab, C. maenas, is found in the Atlantic on all types of shore and sublittoraly to depths of 200 m, from northern Norway to 6* io4 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 its development investigated, the first zoea having been hatched and described by Couch in 1840. Since that time there have been many accounts of the various larval stages of Carcinus, those by Williamson (1900, 1903) being by far the most detailed and complete (see Lebour, 1928, for earlier references). Williamson hatched the first zoea which moulted to the second stage, but he obtained the later zoeae and the megalopa from the plankton and it was not until 1967 that the species was reared in the laboratory with any degree of success (Williams, 1967). Williams, however, reared Carcinus mainly for behavioural studies and did not describe the larval stages which 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 crab stages of Carcinus maenas from laboratory-reared material. At that time Carcinus was 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 with which ovigerous females could be obtained. Compared with many other British crabs the larvae of C. maenas were well known and a new account did not seem to be warranted, even though the description could now be based on a complete series of reared stages. However, Williamson's papers were published in a journal which is not now readily accessible to many workers and when one of us (R. W. I.) reared the larval stages of C. mediterraneus from a female collected in Tunisian waters, it seemed worth while to publish a comparative account of the development of these two closely 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, in May 1973 (B.M. reg. no. 1974:332), and from one female C. mediterraneus collected at 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 and in 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 ooo units/litre) and reduced glutathione (o-ooi M), the last additive acting as a feeding stimulant. Every other day the culture water was changed and at the same time the 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 rearing penaeid larvae. In these cases 300-400 larvae were placed into each container (see Fig. 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 drain tube. level was reduced to the 'drain' line at each water change. A gentle air stream induced sufficient water movement in the jars to keep the larvae circulating. Survivals of both the tray-reared and mass-cultured C. maenas larvae were very high, up to 68 per cent of the stage i zoeae reaching the first crab stage. During the C. mediterraneus culture period, however, an air-conditioner breakdown resulted in several days of oscillating water temperatures reaching 25 °C. Survivals were there- fore poor, only two animals reaching the first young crab stage and neither surviving to the next moult. The larvae and moults were preserved in 70 per cent ethanol, and cleared and dissected in lactic acid. Drawings and measurements were made with the aid of a camera lucida. The measurements taken were (a) the distance between the tips of the dorsal and rostral spines (T.T.), (b) the rostral spine length (R.S.) from the tip of 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 carapace margin. RESULTS Adults Until relatively recently the genus Carcinus was considered to contain only the single species, C. maenas, which was recorded from the Atlantic, the Mediterranean and, 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 and Mediterranean 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-lateral carapace teeth, the denser setation of the anterio-lateral carapace margins, the sharper carpal tooth on the cheliped and the more pronounced anterio-external angle of the merus of the third maxilliped, are not apparent in the females from which the larvae reported in this paper were obtained. There are, nevertheless, good mor- phological distinctions between the females and these are listed in Table I and illustrated in Plate i. TABLE i Differences between the female C. maenas and C. mediterraneus from which the larvae 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 rough to touch 3. 5th (posterior) pair of anterio-lateral teeth directed forwards 4. Front does not protrude and is not setose (Plate lA) 5. Carapace regions not strongly elevated and, when viewed from behind, carapace relatively 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 carapace vaulted (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 where the only morphological distinctions noted were the relative lengths of the dorsal and rostral carapace spines. With the exception of the dimensions, therefore, the following descriptions of the zoeae apply both to C. maenas and C. mediterraneus. FIRST ZOEA Dimensions 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-lateral angles 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 ZOEA Dimensions 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 from C. maenas specimens and the remainder from C. mediterraneus. Bar scale represents 0-2 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 ZOEA Dimensions 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 ZOEA Dimensions 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 the zoeae. However, the differences noted involve only the pleopods, uropods and telson and with the exception of these features the following description, like those of 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 with 3 or 4 aesthetascs. Terminal segment with 2 setae, penultimate segment with a lateral seta and with or without a medial seta. Ventral flagellum unsegmented with 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 row of 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 setae on the surfaces of the blade. Endopod unarmed or with a single short seta. Lobes of 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 proximal segment and 3-5 on the distal segment. Unsegmented endopod with 4 or 5 marginal setae. Basal segment with 14-16 marginal and sub-marginal setae. Coxal endite with 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 scales represent 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. Endopod of 5 segments, armed with numerous spines and setae and with the ischium expanded and 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 of carapace in C. maenas (b) and C. mediterraneus (c) ; (d and e) ventral views of telson and uropods in C, maenas (d) and C. mediterraneus (e) ; (g and h) dactyl of fifth pereiopod in C. 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. Legs 2-5 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 dorsal setae, 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 the dorsal and ventral surfaces, somewhat variable in shape, but with consistent differences between the two forms ; in C. maenas the telson narrows posteriorly and n6 A. L. RICE & R. W. INGLE d 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 parallel or even diverging slightly, the posterio-lateral angles are more abrupt and the posterior 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 the setation of the exopods of these appendages and of the uropods was consistently different in the examples of the two forms examined and afforded the clearest means of separation. In general C. mediterraneus had fewer setae on the pleopods than did C. maenas, though since there was a good deal of overlap the setation of no single appendage would separate the two forms (see Table 2). When, however, the total Range Mean Range Mean IO-I2 II-O 11-13 u-i 11-12 11-9 11-12 n-8 12-13 12-4 11-12 «'3 ii ii 9-II 9-8 5 5 4-6 4'5 LARVAL DEVELOPMENT OF CARCINUS 117 number of setae on the pleopods of the 5th abdominal somite and on the uropods was combined, a clear distinction was apparent, C. maenas megalopae always having a total of 32 setae on these appendages, while C. mediterraneus larvae never had more than 30. TABLE 2 Setation of the pleopods and uropods in the megalopa larvae of C. maenas and C. mediterraneus, based on counts of five specimens in each case C. maenas C. mediterraneus Rai Pleopod, somite 2 somite 3 somite 4 somite 5 Uropod FIRST CRAB STAGE (Fig. 8) Carcinus maenas : Carapace length 1-49-1-52 mm ; carapace width (across tips of 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 so that little reliance can be placed on their comparison with the corresponding stage in C. maenas. However, in the material examined the two forms were extremely similar, the only differences being a slightly more prominent rostrum and dorsal carapace tubercles and a squarer telson in C. mediterraneus than in C. maenas. The other characters distinguishing the adults were not apparent in the first crab stage although mediterraneus has a slightly lighter build, the carapace being relatively a little narrower, than maenas. DISCUSSION Williamson's (1903) account of the development of Carcinus maenas includes a great deal of detail and, in this respect, has rarely been equalled by any subsequent description of decapod larvae. His larvae were generally a little larger than those reared from the Plymouth and Brighton females reported here, but otherwise the two accounts agree almost without exception. This agreement confirms, if such confirmation was needed, firstly the accuracy of Williamson's observations and secondly 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 the loss in the late zoeae of two of the three telson fork spines which are present in the early stages, whereas a second spine was almost always present in our stage four larvae. However, this second spine is extremely small and difficult to see, so that it was probably simply missed by Williamson and Lebour. The earlier accounts also agree with the present description of the British Carcinus larvae 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, and ii respectively, while both he and Lebour report 5 setae on the uropods as opposed to the 4 setae usually found on our Mediterranean larvae. The account of these Mediterranean larvae is based on material obtained from only one brood so that the distinctions noted might simply reflect individual variation. But the agreement between all the available descriptions of British larvae argues against this and suggests that definite genetic differences exist between the Atlantic and Mediterranean Carcinus populations. Whether these differences are sufficient to warrant the two populations being accorded full specific status is, however, debatable, for they appear to be allopatric, C. mediterraneus never having been reported from outside the Mediterranean, while C. maenas is not known from within it. In these circumstances the ultimate criterion of the absence of interbreeding between overlapping populations cannot be applied and any opinion about their taxonomic status must be somewhat sub- jective. Comparing the degree of difference between the two allopatric forms with that between undoubtedly distinct species in the same group does not provide very conclusive evidence. For although portunid crabs are often quite difficult to separate as juveniles, there are usually more distinct differences between the mature forms than those noted between Carcinus maenas and C. mediterraneus, and in the larval stages the situation is similar. In terms of the number of species of which larvae have been described, the best known portunid genus is Macropipus, and although the known larval stages of this genus are all very similar, detailed examina- tion has generally revealed better distinctions between the species than those between 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 separation of maenas and mediterraneus as distinct species, despite the existence of consistent differences between the adults. A study of both adults and larvae from the vicinity of the Straits of Gibraltar, both in the Mediterranean and in the Atlantic, would be of considerable interest, though if no area was found in which the two forms co-exist, or intergrade, the problem would still be unresolved. Since the adults are so readily distinguishable, however, it seems worth while at present to maintain their specific status, but if they are ultimately shown to be totally allopatric the most sensible course would probably be to consider them to be subspecies, as was apparently the intention of Czerniavsky (1884) when he divided C. maenas into the two varieties mediterranea and septentrionalis (see Holthuis & Gottlieb, 1958). For as Mayr, Linsley & Usinger (1953) point out, the use of trinominals has the advantage in such situations of conveying the important information that the two forms are closely related and geographically separated. ACKNOWLEDGEMENTS We thank Dr R. B. Manning who sponsored a visit by one of us (R. W. I.) to Tunisia under the Smithsonian Institution Foreign Currency Program, partly for LARVAL DEVELOPMENT OF CARCINUS 119 the purpose of obtaining ovigerous material for comparative larval studies. We also thank members of the West Wickham Branch of the British Subaqua Club for 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 the female 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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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 CRABS MACROPIPUS 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*44 MORPHOLOGY OF THE BRITISH PORTUIS CRABS MACROPIPUS PUBER (L.) AND M. HOLSATUS (FABRICIUS), WITH A DISCUSSION OF GENERIC AND SUB-FAMILIAL LARVAL CHARACTERS WITHIN THE PORTUNIDAE 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28, No. 4, of the Zoological series. The abbreviated titles of periodicals cited follow those of 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 OF THE 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, are described from material reared in the laboratory. Comparison of these larvae with one another and with those of M. holsatus, the only other species of the genus described in detail, indicates that while the larvae of the Macropipus species are all very similar there are clear distinctions between them. The larval information available for the family as a whole generally supports the currently accepted sub-divisions within the Portunidae based on adult characters, but further detailed larval descriptions, particularly of those sub-families of which the larval stages are at present unknown, would probably help in the interpretation of phylogenetic relationships both within the 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 larval development 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. This second paper deals with two of the eight species of the genus Macropipus which have 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 of chromatophore pattern. Since that time the larvae of only one species of Macropipus have been reared and described in detail (Goldstein, 1971), but with a further two species described here it is now possible to discuss the nature of the specific larval characters 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 on 27 February 1973 and both broods were reared partly in mass culture and partly individually in compartmented plastic boxes (see Rice and Ingle, 1975). All of the larvae 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 was somewhat 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 individually reared M. puber only 17 (about 9 per cent) reached this stage. Although there was considerable individual variation in the number of days spent in 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 hatching while M. puber reached this moult in 46-56 (mean 50) days. The reared larvae and the females from which they were obtained are deposited in the British Museum (Natural History) under registration numbers 1974 : 333 (M. holsatus) and 1974 : 334 (M. puber}. MORPHOLOGY OF THE LARVAL STAGES AND SPECIFIC DISTINCTIONS WITHIN THE GENUS MACROPIPUS The larvae of Macropipus puber and M. holsatus are illustrated in Figs. 2-9. The illustrations are supplemented in Tables 1-6 where these two species are compared with 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 very similar, but although no single character separates them at all stages there are clear distinctions between them. Thus the zoeae of puber differ from the other species at all stages in having straight rather 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 later stages but at this time, that is in the fourth and fifth stages, an obvious distinction is the loss in puber of one of the spines on the telson forks. The larvae of puber are also 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 first stage, specimens of which are deposited in the British Museum (Natural History).* As in the adults (see Christiansen, 1969), the larvae of holsatus and marmoreus are more difficult to separate on morphological grounds, both of them having curved carapace spines and being similar in size. However, there are a number of fairly easily seen differences. First, the posterio-lateral processes on the abdominal somites are much better developed in marmoreus than in either holsatus or puber, particularly in the late zoeal stages. Secondly, with the possible exception of stage III, 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 between puber and holsatus in this respect. LARVAL DEVELOPMENT OF MACROPIPUS SPECIES 125 of the antenna much shorter relative to the rostral spine. Finally, the lateral knobs on the third abdominal somite are lost in stage II in marmoreus, that is one stage earlier than in either holsatus or puber. In addition to these characters there are less obvious differences, particularly between 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 megalopae of puber tending to be smaller than those of either holsatus or marmoreus, the size ranges overlap a good deal. Similarly, while the megalopae of holsatus seem to have a rather lighter build than those of the other two species, this distinction is not very clear and in any case is obscured by the distortion resulting from preservation. There are, however, good distinctions between the species based on the antennae and telson. Thus, while puber has an antennal flagellum of seven segments and a telson with three pairs of dorsal setae, holsatus has eight segments in the flagellum and only two pairs of dorsal telson setae, and marmoreus has six segments in the antennal flagellum and four pairs of dorsal telson setae. Finally, the slender dactyl of 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 Macropipus puber, M. holsatus and M. marmoreus to be differentiated, it is not possible to anticipate which of these characters, if any, will be of use in distinguishing between the other species of the genus. However, the fact that such consistent differences exist, particularly between the larval stages of holsatus and marmoreus which are so similar as adults, indicates that when the larvae of these other species are described in sufficient detail similarly clear distinctions will be found between them. GENERIC AND SUB-FAMILIAL LARVAL DISTINCTIONS WITHIN THE FAMILY PORTUNIDAE Previous attempts to define generic and sub-familial distinctions amongst portunid larvae have been clouded by the unfortunate nomenclatural confusion surrounding this family of crabs (see Opin. Decl. Int. Commn zool. Nom. 1956, vol. 12, Opinion 394, pp. 317-336). This confusion was primarily due to the use of the name Portunus for two distinct genera, one mainly confined to European waters, and the other widely distributed in the tropics and sub-tropics. The International Com- mission's decision was to retain the name Portunus Weber, 1795, for the widely distributed genus, which had also been called Neptunus in the Mediterranean and Indo-West-Pacific, and to adopt the name Macropipus Prestandrea, 1833, for the European genus. These changes have had repercussions in the nomenclature of the sub-familial divisions within the Portunidae, but the situation was clarified by Stephenson and Campbell (1960) whose interpretation and nomenclature, based on adult characters, are followed here except that the sub-family name Polybiinae Ortmann, 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) distance from the tip of the dorsal spine to the tip of the rostrum, (C) distance between the tips of 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), but larvae of only the first three of these are known. Apart from Macropipus, larvae of only 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, (_;') endopod and 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 second maxillipeds, stage V. The bar scale represents o-i mm for h, j and k, and 0-5 mm for the 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 scales represent 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, second and third maxillipeds, M. puber ; (g) cheliped, M. holsatus ; (h, j) second and fourth pereiopods, 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 tip of rostral spines (a) : 1-9-2-2 mm Width including lateral spines (b) 0-67-0-78 mm Ratio a/b : 2-68-2-80 Length of dorsal spine : 0-9 mm Carapace : Dorsal spine stout, straight or slightly curved at tip, much longer than carapace. Rostral spine straight or slightly curved anteriorly, longer than carapace. Lateral spines well developed. Carapace dorsal surface minutely rugose. Eyes sessile (Fig. 2a, b) M. holsatus SIZE 0-4-0-6 mm 1-1-1-3 mm 0-61-0-74 mm i -80 0-4-0-5 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 curved posteriorly and equal to or only slightly longer than carapace. Rostral spine straight or curved backwards, equal to or shorter than carapace. Lateral spines well developed. Carapace smooth. Eyes sessile (Fig. -zd, e) As puber Similar to holsatus As puber Spinous process about as long as rostral spine. Otherwise as puber As puber As puber As puber Exopod with 3 terminal setae. Otherwise as puber As puber As puber Scaphognathite with 6 setae ; otherwise as puber Basipodite with only 6 setae ; 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. marmoreus M. puber M. holsatus M. marmoreus SIZE Carapace length : 0-85-0-90 mm 0-60-0-70 mm 0-60 mm Tip of dorsal to tip of 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 I Maxillule : Endopod as in stage I. Endites with additional setae. Exopod seta present Maxilla : Endopod as in stage I. Endites with additional setae. Scaphognathite with 1 1 setae, posterior one no longer significantly different First maxilliped : Exopod with 6 natatory setae ; otherwise as in stage I Second maxilliped : Endopod terminal segment with 4 setae. Exopod with 6 natatory setae Pereiopods : Limb buds prominent M. holsatus As puber As puber As puber As puber As puber Limb 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 telson forks well developed As puber M. marmoreus As puber As puber As puber Basipodite with 8 setae ; otherwise as puber Endopod terminal segment with 5 setae. Otherwise as puber As 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 mm Tip of dorsal to tip of rostral spine (a) : 3-0-3-3 mm Width including lateral spines (6) i -00-1-04 mm Ratio a/b : 2-95-3-17 Length of dorsal spine : 1-40 mm Carapace : As in stage II (Fig. 5a, b) M. holsatus SIZE 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 equal to or slightly greater than carapace length (Fig. $d, e) 138 A. L. RICE & R. W. INGLE M. puber Antennule : As in stage II Antenna : Endopod bud almost J length of exopod. Spinous process c \ length of rostral spine. Exopod with 2 terminal aesthetascs (Fig. 30) Maxillule : Additional setae on endites. Otherwise as in stage II Maxilla : Scaphognathite with 13-14 setae First maxilliped : Exopod with 8 natatory setae. Otherwise as in previous stages Second maxilliped : Exopod with 8 natatory setae. Endopod terminal segment with 4 setae M. holsatus SIZE As in stage II As puber but spinous process > £ length of rostral spine As puber As puber As puber Endopod with 3 terminal setae. Otherwise as puber M. marmoreus As in stage II Endopod bud almost £ length of exopod. Exopod with 4 terminal setae. Otherwise as holsatus As puber As puber As puber Endopod with 5 terminal 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 puber As puber More prominent, those on somite 3 about £ length of somite 4 Telson forks each with 3 spines 4 or 5 pairs of setae As puber As 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 mm Tip of dorsal to tip of rostral spine (a) : 3-40-3-60 mm Width including lateral spines (b) c i -20 mm Ratio a/b : 2-86-2-96 Length of dorsal spine : 1-30-1-40 mm M. holsatus SIZE M. marmoreus 1-10-1-20 mm 2-60-2-90 mm 1-20-1-25 mm 2-10-2-29 1-10-1-20 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 and i or 2 setae. Ventral flagellum may be present as small bud Antenna : Spinous process about £ length of rostral spine. Exopod f length of spinous process, with 2 terminal setae. Endopod J exopod (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 clearly chelate 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 spinous process, with 2 terminal setae. Endopod f exopod As puber As puber As puber 10 natatory setae. Endopod with 4 terminal setae, i.e. as in puber As puber Posterio-lateral processes of somites 3-5 longer than in stage III, those of 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 as buds ABDOMEN (Fig. 6c, f) More prominent than in puber, those of somite 3 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 puber As puber 10 natatory setae. Endopod with 5 terminal setae As puber More prominent than in holsatus, those of somite 3 > £ 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 mm Tip of dorsal to tip of rostral spine (a) : 3-8-4-1 mm Width including lateral spines (b) 1-60-1-70 mm Ratio a/b : 2-4-2-5 M. holsatus SIZE M. marmoreus 1-30-1-48 mm 3-00-3-50 mm 1-50-1-60 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. holsatus SIZE 1-3 mm M. marmoreus Carapace : As in previous stages (Fig. 7a, 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 > f length of rostral spine. Otherwise as puber Posterio-lateral processes of somites 3 and 4 about J length of succeeding somites Telson forks with 2 spines Posterior margin with 10 or, more usually, 12 setae Pleopods as long as succeeding somites As puber As puber As puber As puber Endopod terminal segment with 4 or 5 setae. Exopod with 12 natatory setae As puber As puber ABDOMEN (Fig. 7c, f) As puber 3 spines 10 setae As puber 1-40 mm As in previous stages 13 aesthetascs plus terminal seta. Ventral flagellum well developed Spinous process < f rostral spine. Exopod < £ spinous process ; 4 terminal setae. Endopod > exopod As puber As puber Scaphognathite with about 40 setae As puber As puber As puber As puber ? Posterio-lateral process of somite 3 > J length of somite 4 3 spines 10 setae As puber TABLE 6 A comparison of the megalopa stage in Macropipus puber, M. holsatus and M. marmoreus M. 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 (see length/width ratio above). Rostrum directed ventrally and not prominent in dorsal view. Dorsal surface with two medial tubercles and two pairs of lateral ones. These tubercles, and particularly the posterior medial one, vary considerably in prominence. Front about £ maximum carapace width. Posterior margin almost straight and meeting lateral margins in abrupt angles (Fig. 8a, b) Antennule : Three-segmented peduncle. Dorsal flagellum of 4 segments with 2 terminal setae and 3 groups of aesthetascs (Fig. gc) Antenna : Peduncle of 3 segments and flagellum of 7 segments, the fifth bearing 2 long setae (Fig. go) Mandible : Well-developed 2- segmented palp with 8-10 setae Maxillule : Endopod of 2 segments carrying 2 and 4 setae respectively. Endites with many marginal spines and setae Maxilla : Endopod unarmed or with a single seta. Schaphognathite with about 50 marginal setae First maxilliped : Exopod with 3-5 setae on the proximal segment and 4 or 5 on the distal one. Endopod unsegmented and reduced. Well- developed epipod of fairly constant width throughout its length (Fig. gd) Second maxilliped : Exopod 2-segmented, proximal with i small seta, distal with 4 or 5 terminal setae. Endopod 4-segmented, proximal naked. Epipod small (Fig. ge) Third maxilliped. Exopod with 6 terminal setae. Endopod of 5 segments, all armed. Epipod long (Fig. 9/) Pereiopods. Cheliped with prominent curved spine on ischio-basis. Pereiopods 2-4 with straight spines on coxae (Fig. gg, h, j) M. holsatus CEPHALOTHORAX Relatively narrow. Rostrum generally directed more anteriorly than in puber and therefore more prominent in dorsal view. Posterior margin arched and not meeting the lateral margins in abrupt angles. Otherwise as puber (Fig. 8e,f) As puber Flagellum of 8 segments (Fig. gb) As puber As puber As puber As puber As puber As puber Pereiopods 1-4 as in puber M. marmoreus Relatively broad. Rostrum prominent in dorsal view. Posterio- lateral tubercles absent. Front about £ maximum carapace width As puber Flagellum of 6 segments, the long setae being on the third As puber As puber As puber Epipod expanded basally. Otherwise as puber As puber As puber Pereiopods 1-4 as in puber I42 M. puber A. L. RICE & R. W. INGLE M. holsatus D actyl of pereiopod 5 slender, length more than 5 times maximum width. Sensory setae clearly sub-terminal. Coxal spine minute or absent (Fig. gk) CEPHALOTHORAX Dactyl of pereiopod 5 broad, length about 4 times maximum width. Sensory setae virtually terminal (Fig. gl, m) M. marmoreus As holsatus Somites 2-5 with posterio-lateral expansions rounded Exopods of pleopods on somites 2-5 with 17-19, 16-19, 17-18 and 13-15 setae respectively Endopods of pleopods with 3, or rarely 4, coupling hooks Exopods of uropods with 8-10 setae Telson usually narrows posteriorly, but may be almost square. Posterior margin usually straight or slightly concave, but may be markedly so Telson dorsal surface with 2 pairs of posterio-medial setae arranged in a rectangle elongated longitudinally. Additional pair of anterio-lateral setae (Fig. 3d) ABDOMEN (Fig. 8c) As puber Exopods of pleopods with 17-20, 19-20, 18-19 and 14-16 setae respectively As puber 9 or 10 setae Telson normally square, but may narrow posteriorly. Shape therefore not consistently distinguishable from puber Telson dorsal surface with 2 pairs of posterio- medial setae arranged in a rectangle elongated transversely. No anterio-lateral setae (Fig. 8g) As holsatus As puber 8 setae Telson dorsal surface with 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 with the exception of an inadequate account of the first zoeal stage of Bathynectes longipes Risso (Lebour, 1931) their description of the development of Ovalipes ocellatus seemed to be the first of a true polybiinid. Their discussion of possible sub-familial larval characters within the Portunidae was therefore largely invalid, but they nevertheless recognized the possession of sternal cornua in the megalopa stage as a portuninid character (see below). Roberts was similarly confused by the generic nomenclature when he compared his reared Bathynectes larvae with those of other species in the family and consequently did not recognize some larval differences which seem to be generic or even sub-familial. On the other hand, Goldstein (1971) did not mix polybiinid and non-polybiinid species in her comparison with the larvae of Macropipus marmoreus, and was able to point out several differences between M. marmoreus, Ovalipes ocellatus and Bathynectes superba. The additional information presented here on the larvae of M. 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 larval Polybiinae. The three genera Macropipus, Ovalipes and Bathynectes are therefore compared in Tables 7 and 8, revealing differences which at the moment appear to be generic. TABLE 7 A 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 5 Posterio-lateral processes, 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 from stage 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 described by 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 forwards Present Absent 2 setae 3 setae Unarmed or with a single seta Unsegmented, expanded at tip with 3-6 setae Not markedly flattened ; L: W>4: i Rounded 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, not expanded, about 17 setae Markedly flattened L:W<3-5:i 8-10 marginal setae ? Directed forwards Absent 2 setae 6 setae Unsegmented, with about 17 setae As O. ocellatus Acute 7 hooks 31-40 marginal setae 3 setae About 22 marginal setae In the zoeal phase Ovalipes can be distinguished from the other two genera by the possession of dorso-lateral processes on the fifth abdominal somite in all stages and on the fourth somite up to stage III, and in lacking a dorsal spine on the telson forks. 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 the zoeal stages Bathynectes has much longer posterio-lateral processes on the fourth somite and also a much more prominent anterior spine on the telson fork than either Macropipus or Ovalipes. Unfortunately Roberts was unable to obtain the megalopa of Bathynectes so that in this stage Macropipus can be compared only with Ovalipes, and even here much of the information which would be useful in such a comparison is not available (see Table 8). Nevertheless, the megalopae of Ovalipes seem to differ from all described Macropipus megalopae in having more setose appendages, particularly the abdominal ones, 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 relation to the Carcininae and the Portuninae, for which at least some larval information has been 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 the larval stages of this species, they have only recently been described in detail from reared material and compared with the closely related C. mediterraneus Czerniavsky (see Rice and Ingle, 1975). The only other carcininid genus of which larvae are known is Portumnus, Lebour (i944a) having hatched and reared the first two stages of P. latipes and attributed to the same species a plankton-caught terminal zoea which 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 ascribed to 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-familial characters, but in addition there are a number of much more obvious differences which appear to enable larval carcininids to be distinguished readily from poly- biinids. Firstly, the two Carcinus species, and probably also Portumnus, pass through only four zoeal stages, whereas all the known polybiinids have five zoeae. The consequent differences in developmental rates result in distinct combinations of characters in the two sub-families ; pleopod buds, for instance, occur on zoeae with 8 natatory setae on the exopods of the maxillipeds in the Carcininae, that is in the third stage, whereas the pleopods do not appear in the Polybiinae until the fourth zoeae in which the maxillipeds carry 10 setae. This character would not, of course, 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 these early stages, and from those of Bathynectes and Ovalipes at all stages, in having no dorso-lateral processes on the third abdominal somite. Carcininid zoeae also differ from the polybiinids, certainly from the third stage, in having only three pairs of setae 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. But the most obvious carcininid zoeal character, which distinguishes them from all other known portunids, both polybiinids and portuninids (see below), is the total absence of lateral carapace spines. In the megalopa stage Carcinus and Portumnus differ from Macropipus in having no coxal spines on pereiopods 2-4, and from Ovalipes in having rounded posterio- lateral margins to the fifth abdominal somite. Both polybiinid genera differ from Carcinus in having more setae on the pleopods and uropods, the difference being more 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 there are published accounts of one or more stages of five portuninid genera, many of these accounts are inadequate, several are based on material collected from the plankton and 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 allow some 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* SPECIES Callinectes sapidus Scylla serrata Scylla serrata Charybdis acuta Charybdis japonica Charybdis lucifera Charybdis annulata Charybdis callianassa Charybdis orientalis Charybdis 6-dentata Charybdis bimaculata Portunus trituberculatus Portunus pelagicus Portunus pelagicus Portunus pelagicus Portunus pelagicus Portunus sayi Portunus depressifrons Thalamita crenata Thalamita sima Portunus sanguinoleutus Portunus pelagicus Charybdis orientalis Thalamita crenata STAGE(S) SOURCE OF MATERIJ Zoeae I -VII, Hatched and reared megalopa Zoeae I-V, megalopa Zoea I Hatched and reared Hatched Zoeae I -VI, Hatched and reared megalopa Zoeae I -VI, Hatched and reared megalopa Zoea I Hatched Zoea I Hatched Zoea I Hatched Zoea I Hatched Zoea I Plankton Zoea IV, megalopa Zoea I Plankton Hatched Zoea I, megalopa Zoea I Plankton Hatched Zoeae I -IV, megalopa Zoeae I, II and III ? Zoea I ? Zoea I ? Zoea I Megalopa Zoea I Zoea I Zoea I Zoea I Hatched and reared Hatched and plankton Plankton Plankton Hatched Plankton Plankton Plankton Plankton Plankton AUTHOR Costlow and Bookhout, 1959 Ong, Kah Sin, 1964 Naidu, 1955 Kurata and Omi, 1969 Yatsuzuka, 1952 Hashmi, 1970 Hashmi, 1970 Hashmi, 1970 Hashmi, 1970 Aikawa, 1937 Aikawa, 1937 Aikawa, 1937 Aikawa, 1937 Delsman and De Man, 1925 Yatsuzuka, 1962 Prasad and Tampi, 1953 Lebour, 19440 Lebour, i944b Prasad and Tampi, 1953 Muraoka, 1969 Chhapgar, 1956 Chhapgar, 1956 Chhapgar, 1956 Chhapgar, 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 by subsequent 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 this manuscript had been submitted. Fortunately, P. spinicarpus is a typical portuninid and its omission does not affect the discussion of sub-familial larval characters in this paper. Firstly, there is considerably more variation in the rate of development in the portuninids than in either the polybiinids or carcininids, the number of zoeal stages ranging 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, according to Prasad and Tampi, 1953, although the sizes given for their zoeae indicate that they may have missed at least one stage). This heterogeneity extends also to some morphological features, such as the telson fork armature, for while there is a tendency to a 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 and P. sayi) retain all three spines on each fork, one of the lateral spines is reduced to a fine hair in Charybdis callianassa, C. lucifer and C. annulata, and totally disappears in 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 of Charybdis orientalis, Thalamita crenata, Portunus sanguinolentus and P. pelagicus. This variability allows the zoeae of individual portuninid species to be distinguished fairly easily from all known polybiinids, but there does not seem to be a single feature which will separate the two sub-families at all zoeal stages. For those characters which are common to all portuninid zoeae, such as the possession of well- developed dorsal, rostral and lateral carapace spines, dorso-lateral processes on abdominal somites 2 and 3, posterio-lateral processes on somites 3-5 in the later stages, telson forks usually with at least two spines of which one is dorsal, and the addition of at least one extra pair of setae on the posterior margin of the telson from stage 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 portuninid zoeae is the absence of any setae on the middle segment of the endopod of the first maxilliped in the first zoeal stage.* [Hashmi (1970) figures a seta in this position in the first zoea of Charybdis callianassa, but we have been unable to confirm its existence in material deposited by him in the collections of the British Museum (Natural History).] Even this fine distinction disappears in the later stages of Scylla serrata and Charybdis acuta, and as early as the second zoea in Callinectes sapidus, and would hardly seem sufficient to support the sub-familial separation of the Portuninae and Polybiinae based on adult characters. But in the megalopa stage 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 somite which 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 on the uropods beyond that found in the other two families, again with the exception of Ovalipes, 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 striking portuninid megalopal character, however, is the possession of a pair of prominent posterio-ventrally directed spines, the so-called sternal cornua, arising from the sternum between the last pair of legs. This feature at once distinguishes the por- tuninids, not only from the carcininids and polybiinids, including Ovalipes, but also from all other brachyuran megalopae so far described. These differences between the larvae of the Carcininae, the Polybiinae and the Portuninae are summarized in the following diagnoses. * One other possibly unique portuninid zoeal character is the position of the pair of fine setae on the dorsal surface of the carapace. In the Carcininae and Polybiinae these setae are anterior to the dorsal spine and close together, whereas in the Portuninae they are between the dorsal and lateral carapace spines. However, they may not always be present, for Costlow and Bookhout, 1959, specifically looked 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 on abdominal somite 2 only ; posterio-lateral processes of abdominal somites 3 and 4 less than half length of succeeding somites in all stages ; telson fork armature reduced to a single spine in late stages ; telson posterior margin with 3 + 3 setae in all stages ; middle segment of endopod of first maxilliped armed with a single seta in stage I. MEGALOPA Rostrum directed forwards or downwards ; no coxal spines on pereiopods ; no sternal cornua ; pleopods with 9-12 (rarely 13) marginal setae ; uropods with 4-10 marginal setae ; posterio-lateral spines on abdominal somite 5 absent. Polybiinae ZOEAE 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-lateral processes of abdominal somites 3 and 4 usually less than half length of succeeding somites in later stages ; telson forks with at least 2 spines in all stages ; telson posterior margin with at least 4 + 4 setae in late stages ; middle segment of endopod of first maxilliped armed in stage I. MEGALOPA Rostrum directed forwards or downwards ; coxal spines on pereiopods 2-4 present (Macropipus) or absent (Ovalipes) ; no sternal cornua ; pleopods with 14-20 (rarely 13) marginal setae (Macropipus) or 31-40 (Ovalipes) ; uropods with 8-10 marginal setae (Macropipus) or c 22 (Ovalipes) ; posterio-lateral spines on abdominal somite 5 absent (Macropipus) or present (Ovalipes) . Portuninae ZOEAE Four to seven zoeal stages ; well-developed lateral carapace spines ; dorso-lateral projections on abdominal somites 2 and 3 in all stages ; posterio-lateral processes of abdominal somites 3 and 4 more than half length of succeeding segments in late stages ; 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 segment of endopod of first maxilliped unarmed in stage I. MEGALOPA Rostrum directed forwards ; coxal spines absent from pereiopods 3 and 4 but may be 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-reaching somite 6. THE BEARING OF LARVAL CHARACTERS ON PORTUNID CLASSIFICATION The above distinctions between the larval stages of the Carcininae, Polybiinae and Portuninae generally support the current divisions within the family which are based largely on the degree to which the adults show adaptations to the swimming habit. These adaptations include a tendency to lighten the integument, to flatten the 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 fringe them with setae to produce effective paddles (Hartnoll, 1971). There is a good deal of variation in the extent of these adaptations within the sub-families, but in general the Carcininae and Portuninae represent the extreme conditions, the Carcininae showing the least and the Portuninae the most modification from the unspecialized brachyuran form. The Polybiinae are somewhat intermediate between the other two sub-families and even within the genus Macropipus there is a considerable range in swimming adaptations, and particularly in the degree of flattening of the legs (see Palmer, 1927). This situation is reflected in the larvae, for while the Carcininae and Portuninae exhibit major differences in both the zoea and megalopa stages, the Polybiinae have zoeae which resemble the Portuninae while the megalopae are more similar to those of the Carcininae. This does not, of course, necessarily mean that the Polybiinae stand phylogenetically between the Carcininae and the Portuninae, and the presence in the zoeae of Ovalipes of characters such as the dorso-lateral processes on abdominal somites 4 and 5, which are apparently unique amongst the portunids, indicates that this genus, at least, is well away from any such route. It has been generally accepted that the swimming adaptations of adult portunids are secondary acquisitions and that the relative absence of such adaptations in the Carcininae is therefore a primitive condition. There have, however, been suggestions that the morphological series into which extant swimming crabs can be arranged might be read in the opposite direction, so that the loss of swimming adaptations becomes an advanced feature (Palmer, 1927; Lebour, 1928). The information available from the larval stages does not provide any clear evidence for one or other of these views. For the main character distinguishing carcininid larvae from all other 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 to be of any particular phylogenetic importance. On the other hand, the presence of sternal horns on all portuninid megalopae, a feature which is unique not only amongst the Portunidae but also amongst the Brachyura generally, indicates that this sub- family probably represents the end of a portunid evolutionary line rather than an intermediate 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 parallel evolutionary tendencies within the Portunidae, one involving a loss of swimming adaptations and a return to the relatively unspecialized brachyuran condition of the Carcininae, and the other leading to the increased specialization of the Portuninae. Morphological details of the larvae of more swimming crab species, and particularly those 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 the swimming crabs and other brachyuran families. But the pelagic larvae are generally much more similar than the adults since they show none of the specializations for such habits as swimming, burrowing or commensalism which characterize the benthic adult phase. A study of larval systematics will therefore probably require a numerical approach, making use of a much greater variety of features than has usually been used in the past. Such an investigation is underway at the moment using the available published information for all brachyuran larvae, and it is hoped that this will not only supplement the systematics based on adult characters but will also help in the identification of unknown plankton-caught larvae. ACKNOWLEDGEMENTS We wish to thank Trevor Davies and members of the Guildford Branch of the British 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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. These papers form Vol. 28, No. 5 of the Zoology series. The abbreviated titles of periodicals cited follow 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 OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 17 September, 1975 Price £6.20 CONTENTS Page 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 & 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 circumtropical distribution. 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 (9°24' S 46°2o' E) is situated in the southwestern Indian Ocean (fig. 5) ; the nearest land of any magnitude is the island of Madagascar at a distance of about 420 km to the southeast and the African mainland at about 640 km to the west. Apart from very small atolls the nearest high islands are the Comoros at a distance of about 400 km to the southwest. The Seychelles are further away than both the African continent and Madagascar, viz. about 1200 km to the northeast. Maxwell Smith (1909) was the first to record the presence of streptaxids on the island of Aldabra. Messrs J. F. Peake and J. D. Taylor have been participating in various phases of the Royal Society Expedition to Aldabra. In the course of their work on the atoll and its satellite island Assumption they have collected extensive series of streptaxid shells, the study of which they have entrusted to the present author. Aldabra and Assumption appear to harbour four species of Streptaxidae, three of which are extinct. All are described below followed by a discussion on their relationships 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 may not 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 staff of the Mollusca Section of the British Museum (Natural History) for assistance in various respects. I am also indebted to The Royal Society, who have fostered and encouraged research on Aldabra Island. Thanks are due to Dr A. Zilch of the Senckenberg-Museum, Frankfurt am Main, for hospitality for comparative studies at 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 of Leiden University. Localities have been pinpointed by means of their coordinates on the Royal Society grid as shown on the map by Stoddart in Westoll & Stoddart (1971) (between pp. 632 and 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 between the typical form and G. g. tsadiensis Blume, but with the aperture smaller than in the 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, obtusely conical. Whorls six to seven, slightly convex, almost completely smooth, only with 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 : a reasonably well-developed oblique angular lamella touching the tip of the labrum ; two subequal mid-labral denticles on a slightly raised common base, labral complex corresponding to shallow but extensive external pit ; a mid-basal denticle, also corresponding to a little and shallow outside depression, which is sometimes hardly noticeable ; bipartite columellar process consisting of two prominent subequal denticles on a raised common base - the bifurcation of the columellar process may vary 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 width 1-1-1-6 x 1-0-1-4 rnm, 6-7 whorls. Table i details the measurements of 56 adult shells ; 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 Cedres Aldabra, Sykes Aldabra, Sykes He Michel Aldabra, Thomasset Anse Cedres Aldabra, Sykes Assumption, Sykes Anse Cedres Aldabra, Thomasset He Michel Aldabra, Sykes He Michel Aldabra (a), Sykes Aldabra, Sykes Anse Cedres Aldabra, Thomasset Assumption, Sykes Platin Aldabra (a), Sykes Assumption, Sykes Anse Cedres He Michel He Michel He Michel Aldabra, Sykes Anse Cedres Aldabra, Thomasset Assumption, Sykes Assumption, Sykes Anse Cedres He Michel Anse Cedres Anse Cedres Aldabra, Thomasset Aldabra, Sykes Aldabra (a), Sykes He Michel Aldabra, Thomasset Aldabra, Thomasset Anse Cedres Aldabra (a), Sykes Assumption, Sykes Aldabra, Sykes Aldabra (a), Sykes Aldabra (a), Sykes Assumption, Sykes I 3-6 x 1-4 mm 2.48 1-8 mm 1-2 x I-I mm 6J 2 3-6 X 1-5 mm 2-35 1-8 mm i-i X I-I mm 6 3 3-6 X 1-5 mm 2*37 1-9 mm 1-2 x I-I mm 6 4 3-6 X 1-6 mm 2-28 1-9 mm 1-2 x I-I mm 6 5 3'7 x 1-4 mm 2-68 1-8 mm 1-2 X i-o mm 6J 6 3'7 X i -4 mm 2-61 1-9 mm 1-2 x i-i mm 6 7 3'7 X 1-4 mm 2-60 1-9 mm 1-2 X i-i mm 6J 8 3'7 X i -6 mm 2-40 1-9 mm 1-2 X 1-2 mm 6 9 3-8 x 1-5 mm 2-54 1-9 mm 1-2 X I-I mm 6i 10 3-8 X 1-6 mm 2-44 1-9 mm 1-2 x I-I mm 6i ii 3-8 X 1-6 mm 2-44 2-O mm 1-2 x I-I mm 6 12 3-8 X 1-6 mm 2-44 2-0 mm 1-2 x I-I mm 6£ 13 3-8 X 1-6 mm 2-35 2-0 mm 1-2 x I-I mm 6 14 3-8 X 1-7 mm 2-26 2-1 mm 1-2 x 1-2 mm 6 15 3'9 x 1-4 mm 2-69 2-O mm 1-2 x I-I mm 6i 16 3'9 X 1-6 mm 2-48 1-9 mm 1-2 X I-I mm 6J 17 3'9 X 1-6 mm 2-48 1-9 mm 1-2 X I-I mm 6i 18 3'9 X 1-6 mm 2-48 2-1 mm 1-3 X I-I mm 6 19 3'9 x 1-6 mm 2-35 1-9 mm 1-4 X I-I mm 6i 20 3'9 x 1-7 mm 2-30 2-1 mm 1-2 X 1-2 mm 6 21 3'9 X 1-5 mm 2-63 1-9 mm 1-2 X 1-2 mm 6 22 3'9 x 1-5 mm 2-63 1-9 mm 1-2 x i-o mm 6f 23 3'9 X 1-6 mm 2-52 2-1 mm 1-2 x 1-2 mm 6J 24 3'9 x 1-6 mm 2-42 2-0 mm 1-2 X 1-2 mm 6j 25 3-9 X 1-6 mm 2-42 2-1 mm 1-2 X 1-2 mm 6i 26 4-0 X 1-5 mm 2-67 1-9 mm I-I X I-I mm 6| 27 4-0 X 1-6 mm 2-56 2-0 mm 1-2 x 1-2 mm 6f 28 4-0 X 1-6 mm 2-56 2-0 mm 1-2 X 1-2 mm 6i 29 4-0 X 1-6 mm 2-46 2-0 mm 1-4 X 1-2 mm 6 30 4'1 X 1-5 mm 2-70 2-1 mm 1-3 x 1-2 mm 6J 31 4-1 X 1-6 mm 2-60 1-9 mm 1-3 X 1-2 mm 7 32 4'1 X 1-6 mm 2-52 2-1 mm 1-2 x 1-2 mm 6J 33 4-1 x 1-5 mm 2'75 2-1 mm 1-2 x I-I mm 6f 34 4'1 X 1-5 mm 2-75 2-1 mm 1-4 x I-I mm 7 35 4'1 X 1-6 mm 2-64 2-0 mm 1-2 x I-I mm 6£ 36 4-1 x 1-6 mm 2-64 2-0 mm 1-2 X I-I mm 6| 37 4'1 x i-6 mm 2'54 2-O mm 1-2 X 1-2 mm 6i 38 4'1 .X 1-7 mm 2-44 2-2 mm 1-3 X 1-2 mm 6i 39 4-2 X 1-6 mm 2-68 2-0 mm 1-3 X I-I mm 6J 40 4-2 X 1-6 mm 2-62 2-O mm 1-4 x 1-2 mm 6J 41 4-3 x 1-6 mm 2-76 2-O mm I-3 x I-I mm 7 42 4'3 X i -9 mm 2-23 2-3 mm 1-6 x 1-4 mm 6 43 4'4 X 1-6 mm 2-80 2-1 mm 1-4 x 1-2 mm 6f 44 4'4 X i -6 mm 2-69 2-1 mm 1-4 x 1-2 mm 7 45 4'4 X 1-6 mm 2-69 2-2 mm 1-3 x 1-2 mm 6* 46 4'4 X 1-7 mm 2-50 2-1 mm 1-4 X 1-3 mm 64 47 4'4 X 1-6 mm 2-84 2-2 mm I-4 X I-I mm 7 i6o A. C. VAN BRUGGEN 48 4'4 x 1-6 mm 2 •84 2-2 mm 1-4 X 1-2 mm 6f 49 4-6 x 1-9 mm 2 •43 2-4 mm 1-6 x 1-3 mm 6* 50 4'7 X 1-7 mm 2 •78 2-4 mm 1'5 X 1-4 mm 6£ 51 4'7 X 1-8 mm 2 •59 2-2 mm J>3 x 1-2 mm 7 52 4'7 X 1-8 mm 2 '59 2-2 mm 1-4 x 1-3 mm 6J 53 4'9 X 1-7 mm 2 •82 2-2 mm 1-5 x 1-3 mm 7 54 4'9 X 1-9 mm 2 •63 2-4 mm 1-6 X i -3 mm 7 55 5'° X 1-8 mm 2' 76 2-4 mm 1-6 X 1-2 mm 7 56 5'1 X 1-9 mm 2 73 2-4 mm I-5 X 1-4 mm 7 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 size taken 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 is the 'type' of Ennea alauda Sykes nom. nud. Specimens nos. 5, 17, 18, and 48 show (very) faint traces of apical spiral sculpture. MATERIAL EXAMINED : RECENT : Holotype, ALDABRA ISLAND : South Island (Grande Terre), Anse Cedres, c. 400 m inland (Grid ref. 359112) under stones in open grassy area amongst mature bush, ii.ix.ig67, (Peake, RSE) BMNH. No. 197424. Paratypes include all material listed below, similar locality, habitat and collecting data as above, 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 around Casuarina trees, 7.ix.i967 (Peake, RSE), BMNH No. 197427 ; He Michel (Coconut Island, 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 cavity fill cut into Takamaka Limestone buff 'soil' with abundant rootlets and associated fauna of Tropidophora and Rachis, age probably last glacial, that is post 125 ooo years 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 length 2-0 mm ; 3, G. peakei n. sp., juvenile shell from below, to show the angular lamella in the 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 columellar complex, which may vary in the depth of bifurcation. There is no trace of apertural dentition in juvenile shells. The above numerical data are for all material examined. The population on Assumption Island is presumably genetically separated from those on Aldabra Island proper. Assumption is situated roughly 30 km to the southeast of Aldabra. However, the number of shells available for Assumption amounts to only nine, so that no conclusions may be drawn. Measurements are in mm and the following data 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, 9 Mean l\d, 2-53, average Ijd 2-54 (Aldabra) ; do., 2-62, 2-66 (Assumption). In both cases the shells have six to seven whorls. The range of measurements is on the whole smaller on Assumption, but this may reflect limitations imposed by the small sample. The only local population on Aldabra which may enjoy a certain degree of spatial and 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, 8 In this case mean and average lid have values of 2-40 and 2-42 respectively and the shells have only 6-6| whorls. However, drawing of conclusions based on eight specimens again seems unwarranted. lie Michel is only about i km from the nearest land, South Island (Grande Terre), so that there is really no question of effective isolation. Smith (1909) recorded this species as 'Ennea sp . . . may prove to be new'. A year later Preston (1910) described Ennea gwendolinae from Kenya, with which Connolly (1925) identified his Aldabra material. Gulella gwendolinae is known from the 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 at first somewhat far-fetched to identify an Aldabra snail with a species of the African mainland. The gap between the coast of East Africa and Aldabra atoll consists of roughly 640 km of ocean, a seemingly unsurmountable barrier for a land snail. On the other hand, the character of the fauna of many islands in the western Indian Ocean is unmistakably African. The following are two quotations from Peake in Westoll & Stoddart (1971 : 581-610) : The affinities of the vertebrate fauna on all islands in the western Indian Ocean are predominantly orientated towards Africa' (p. 586) ; 'The fauna exhibits a wide range of affinities depending on the taxa and taxonomic levels considered ; those with Africa are dominant, . . .' (p. 606). Cogan, ALDABRA STREPTAX1DAE 163 Hutson & Shaffer in Westoll & Stoddart (1971 : 315-325) write : To summarize it may 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, although some may have reached the atoll via Madagascar. All this stresses African origin and derivation, but so far no non-marine molluscan species on Aldabra has actually been 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 modern summary is available, it appears from a scrutiny of a series of papers by Morelet (1860-1885) and material in various collections that African species indeed do occur here. 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 the Frankfurt and Leiden museums. Data shown under porrecta are from the holotype in the Frankfurt museum and those for tsadiensis have been extracted from Blume (1959). In Table 2 the various forms of G. gwendolinae are compared ; the subspecies or varieties 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 typical form (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 a wide 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 a small overlap ; (d) has a smaller aperture than both the typical form and tsadiensis, with small overlaps ; (e) has somewhat fewer whorls than the typical form, but somewhat more than tsadiensis. This shows on the whole that at least three recognizable units may be distinguished, viz. the Aldabra-Assumption populations, the typical form, and the subspecies tsadiensis. 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 the isolated location of the Aldabra-Assumption populations, it is proposed here to separate these as the subspecies aldabrae. Unfortunately anatomical studies are not as yet possible because of the lack of preserved soft parts. Gulella gwendolinae aldabrae is the only recent streptaxid on Aldabra atoll. It is reasonable to expect it to have been transported from the African mainland to the island. Aldabra has always been a remote place and in view of the noticeable differences found and fossil occurrence it is unlikely that it has arrived in the wake of Man. Also, there are no records of the species from either the Comoros or Madagascar (cf. Fischer-Piette & Bedoucha, 1964^). Dispersal by tropical storms is 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 populate Aldabra. Rafting may merit consideration, but there are no suitable surface currents to effect a dispersal from Africa to Aldabra. Obviously G. gwendolinae has good 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 smooth interstices 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 width about the middle, with open umbilicus, costulate, creamy white. Spire produced, sides slightly to markedly convex, subparallel, apex somewhat flattened, obtusely conical. Whorls six to six-and-a-half, convex and sculptured with comparatively prominent, regular, straight and perpendicular, widely distant, costulae, interstices much wider than riblets, smooth, under high magnification very finely granulate. In front view the holotype shell shows only about eleven costulae on the part of the whorl above the aperture ; the last whorl has a total of about fifteen riblets. Initial two whorls smooth, very fine granulate under high magnification ; penultimate and last whorls comparatively small. Sutures shallow, simple to subcrenellate, somewhat impressed. Aperture somewhat oblique, invertedly triangular with smoothly rounded base, about as high as wide, peristome fairly thick, expanded and somewhat reflected, dentition more or less four-fold. To the right of the middle of paries a fairly large, obliquely perpendicular, angular lamella, which is nothing but a V-shaped pleat in the peristome, connected with labrum ; about half-way down the labrum a superficial swelling may be interpreted as a labral process ; somewhat below this, but much deeper inside the aperture, and at a slight distance from the tip of the angular lamella, a blunt inner labral process is seen, slightly above which there 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 shallow depression on the outside of the aperture ; the columellar lamella corresponds to a shallow furrow on the left of the outside bottom of the aperture, thus adjoining the umbilicus. 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 damaged apex damaged aperture rather oblique holotype, fig. 2 aperture damaged aperture obscured by matrix matter I 1-7 x i [-3 mm i •30 0-8 mm 0-6 X 0-6 mm 6 2 1-8 x •3 mm i •43 0-8 mm 0-6 X 0-6 mm - 3 1-8 x •3mm i •36 0-8 mm 0-6 X 0-6 mm 6i 4 1-8 x •3 mm i •36 0-9 mm 0-6 X 0-6 mm 6 5 1-8 x •3 mm i •39 0-8 mm 0-7 X 0-6 mm 6 6 1-8 x •3 mm i 35 0-8 mm 0-6 X 0-6 mm 6i 7 1-9 x i •3 mm i 48 0-8 mm 0-6 X 0-7 mm 6i 8 1-9x1 •3 mm i 48 0-8 mm 0-6 X 0-7 mm 6 9 1-9x1 •3 mm i 47 0-8 mm 0-7 X 0-6 mm 61 10 1-9x1 •3 mm i 42 0-9 mm 0-7 X 0-6 mm 6* ii 2-0 X 1 •3 mm i 57 0-9 mm 0-7 X 0-7 mm 64 12 2'O X 1 •3 mm i 51 0-8 mm 0-6 X 0-6 mm 61 13 2-1 X 1 •3 mm i 55 0-9 mm - 61 14 2-1 X 1 •5 mm i 45 0-8 mm - 6£ 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 adult shells ; the holotype is no. n. MATERIAL EXAMINED : FOSSIL : Holotype, ALDABRA ISLAND : Middle Island (He Malabar), Stn 34F (Grid ref. 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 paratypes in poor condition or still covered by the matrix BMNH No. GG2I204, numerous juvenile 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), age inferred as 27 ooo BP (Taylor et al, RSE), BMNH No. GG2I206, some shell fragments. 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 following details on the localities where Guletta peakei has been obtained. Stn 26F : 'Brown cavity-fill deposit in Takamaka Limestone. Many small gastropods. Age un- certain, possibly pre-125 °°° yrs-' Stn 34F : 'Cavity-fill deposit cut into the Takamaka Limestone, buff 'soil' containing abundant large ribbed Tropidophora (only site for this species). Abundant Assiminea, Gulella within cavities of the Tropidophora and in the matrix. Almost certain last glacial, post 125 ooo yrs BP.' Stn 3gA : 'Solution cavity-fill cut into Takamaka Limestone. Buff 'soil' with abundant rootlets and associated fauna of Tropidophora and Rachis. Age probably last glacial, post 125 ooo yrs BP.' The species has been named after Mr J. F. Peake, Deputy Keeper of Zoology and Head of the Mollusca Section of the British Museum (Natural History), as a token of friendship and admiration for his island research. The angular lamella is present in all juvenile shells in the form of a long and conspicuous, 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 course of growth while being added to at the actual aperture. No other dental processes are present in the juvenile shells. Juvenile dentition is rare among species of the genus Gulella and much more common among representatives of allied genera, such as Ptychotrema (vide, e.g., van Bruggen, 1971, fig. 2, p. 249). Among the about 125 species of Guletta in Southern Africa there are about three species in which this phenomenon has been described (Burnup, 1925 ; Connolly, 1939), although juvenile shells are as yet unknown for a number of species. At least three of the species which will be considered below when trying to assess the relationships of the new species 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 as interpretation of Blume, 1965). Guletta peakei has no allies on Europa Island (so far no streptaxids have been reported 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 the species enumerated by Morelet for the Comoros, such as Pupa minuscula Morelet (1877 : 340, pi. 12, fig. 5), which may be a streptaxid, superficially resemble Guletta peakei, but never show the peculiar spaced lamellae on the whorls. As regards Pupa minuscula Morelet writes 'obsolete costulata', and 'Le test est orne de cotes fines, 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 shell except 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 each other. 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 also Zilch, 1961). The subgenera Mirigulella (monotypic) and Aenigmigulella (with two species, cf. Adam, 1965 : 40) are very probably products of a long and separate development ; the subgenus Costiguletta with perhaps about ten species altogether is rather an assemblage of diverse elements. A preliminary assessment of the species of 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 and spiral 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 the whorls (see also Adam, 1965 : 46, who states that all three may also have costulate early whorls), and (c) the West African species G. hedwigae Degner, with smooth interstices between the lamellae on the whorls. G. microtaenia Pilsbry & Cockerell most probably does not belong to Costiguletta s. lat. because of the 'delicate riblets' (Pilsbry & Cockerell, 1933 : 372). For the time being the present author refrains from naming the subdivisions of Costiguletta s. lat., particularly because of the differences in dentition of the shells and the ab- sence of anatomical data. However, zoogeographically the above three groups at first sight seem to be fairly natural : group (a) inhabits the West and Central African equatorial forest, group (b) the East African forests and group (c) the western parts of the West African equatorial forest. The subgenera Aenigmigulella and Mirigulella, both from East African forests on elevated country, may have ancestors 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 outside the forest. Therefore one may be tempted to consider all above taxa (all forest dwellers 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 these species in the museums in London, Frankfurt am Main and Leiden. G. peakei is not to be confused with any of the species under discussion. By virtue of the absence of sculpture on the interstices between the lamellae on the whorls the new species belongs to group (c) of Costiguletta s. lat. Zoogeographically this is a somewhat surprising conclusion, because one hardly expects a species from Liberia and one from Aldabra to have common ancestors. Of course, the minute terrestrial snails of continental Africa are still very incompletely known and G. hedwigae or allied species 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 African origin. 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 to help solve this question ; the fact that G. peakei is extinct will, however, frustrate such 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 available on Aldabra, indeed the overall impression is that of a 'semi-arid island' (Stoddart in Westoll & Stoddart, 1971 : 8). Aldabra is an elevated atoll consisting of an elevated reef situated on the summit of a mountain rising from the sea floor. There is evidence that 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), which must have resulted in a moister climate and consequently also the presence of a type of forest suitable for species such as G. peakei. Some of the islands in the Western Indian Ocean still have a fair amount of forest, but so far no apparent allies of G. peakei have been obtained here. Gulella insulincola n. sp. Figs. 4, 8 DIAGNOSIS. A small species of Gulella with smooth whorls, open umbilicus, and dentition consisting of angular lamella, two labral processes, a basal denticle, and columellar lamella. DESCRIPTION OF SHELL. Shell (fig. 4) small, subcylindriform, with open umbilicus, smooth, creamy white. Spire produced, sides subparallel, apex flattened, obtusely conical. 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, somewhat impressed, 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 denticles on a slightly raised common base, labral complex corresponding to very shallow external pit ; a small mid-basal denticle, which usually is situated slightly to the left 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 lamella small 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 adult shells ; the holotype is no. 4. MATERIAL EXAMINED : FOSSIL : Holotype, ALDABRA ISLAND : Middle Island (He Malabar), Stn 34F (Grid ref. 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 of locality see data under 'Distribution' for that species. All specimens are in poor condition, the shells being worn and very fragile. Distribution (fig. 8). So far only known from the Quaternary of He Malabar or Middle 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 repaired The holotype shell, no. 4, was broken, but has been repaired. The specific name insulincola is a noun derived from insula (Lat. : island) and incola (Lat. : inhabitant). The new species represents a common pattern in the genus Gulella. Taxa with a smooth shell and a five-fold dentition have been brought together for Southern Africa by Connolly (1939 : 20) as group 4 (ii) (10 species) and the East African ones are treated by Verdcourt (1962 : 20) in part of Key 4 (9 species). Many of these species show a dental pattern similar to that of G. insulincola. Among the island dwellers 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 has failed to provide satisfactory identification, so that we may conclude that G. in- sulincola represents indeed a hitherto undescribed species. Its closest allies are perhaps the above species from Mauritius and the Comoros ; these differ from the new species in being much larger (sometimes twice as large), having more whorls and being less slender than G. insulincola. Perhaps the group with a smooth shell and a dental pattern consisting of angular lamella, two labral processes, a basal denticle, 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 Ocean as well. The possibility that G. poutrini and G. comorensis may be the components of a superspecies or represent the subspecies of a widely-dispersed species may be merely food for thought in this respect. On the other hand, a shell type such as has been discussed here occurs throughout the genus Gulella and may well be the result of convergent evolution. G. insulincola is obviously extinct on Aldabra, which may also be due to the fact that suitable habitat has disappeared in the course of the process of the island becoming progressively drier. 1 7o A. C. VAN BRUGGEN Gulella spec. There are two specimens (BMNH) which represent a fourth species of the genus Gulella. 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 are smooth 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 (6-8 Mascarenes) . ALDABRA STREPTAXIDAE 171 specimen is a much younger juvenile shell of probably the same species ; it measures only 2 -3 mm and the aperture is filled with matrix matter. This shell was obtained not 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 as 3gA and D'. (Notes supplied by Peake and Taylor ; Stns 3gA and 390 have been described above under the other species.) No opinion as regards identity of the present 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 presence of four species of Streptaxidae, viz. Gulella gwendolinae aldabrae n. subsp., the only Recent representative of the family on the island, and three extinct taxa, G. peakei n. sp., G. insulincola n. sp. and an incompletely known species, G. spec. The family Streptaxidae has excellent dispersal abilities, at least in the Western Indian Ocean. Streptaxids have reached both the Seychelles and the Mascarene Islands at distances of 1400 and 2500 km from the African mainland respectively. The Mascarene Islands are also situated betweeen 700 and 1300 km from Madagascar from 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 (if any) with the streptaxids of that continent. G. gwendolinae is a very widely dispersed African species with a tendency to become locally separated into reasonably recognizable subspecies. Its dispersal abilities are sufficiently illustrated by its wide distribution on Aldabra and Assumption (fig. 6) ; moreover, it is also locally abundant - sufficient reason to consider it a successful 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. Fossil occurrence shows that it has been on the atoll for a long time, very probably having been 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 (van Bruggen, 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. There are 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 indicates that 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 relatives of at least G. peakei are restricted to forest habitats such as are no longer available on 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 gwendolinae aldabrae n. subsp. (the arrow points to Assumption) ; 7, G. peakei n. sp. ; 8, G. insulincola n. sp. (dot) and G. spec, (asterisks). which may be the nearest allies of G. insulincola. This species or its ancestors may therefore have arrived secondarily from these islands, although Aldabra is closer to the 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 with forest components and that a progressive drying out of conditions has caused these to disappear together with the forest dwellers among the animals that led a sheltered life in the then available leaf mould. This has caused some of the predators, the three 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 Aldabra streptaxids have strong links with those from that continent. Dispersal through human agency, by tropical storms, by birds, and by rafting have all been ruled out when G. gwendolinae aldabrae was discussed above. The same applies, mutatis mutandis, for the extinct species. Yet, much of the present flora and fauna of Aldabra atoll or their ancestors have obviously come from Africa. Perhaps dispersal along 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 comprehensive paper 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 family Streptaxidae appears to occupy a dominant position with 46 species. Gulella gwendolinae aldabrae 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 in Innerafrika 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 description of 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 lies Sechelles. 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 den Sammlungen 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, ainsi que 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 Grande Comore. /. 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 German East 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 FROM ALDABRA ATOLL, WESTERN INDIAN OCEAN By C. M. PATTERSON INTRODUCTION THE Succineidae, a rather diverse land snail family, currently includes 12 recent genera (Patterson, 1971, I973a). Succinea, Oxyloma and Catinella have received the most attention by researchers, while representatives of the remaining genera have scarcely been studied. Quickia is distinguished from other succineids by the position of the right tentacular assembly (situated entirely to the inside of the terminal genitalia) combined with the lack of a penial sheath and penial appendix. Some unique features of Quickia were first reported by H. E. Quick (1936), but the genus was not named formally until 14 years later by Odhner (1950). The distribution of Quickia given by Odhner was '. . . from Liberia to the Cameroons and Gabon, on Prince 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 cytological study of two Indian species (Patterson, 1970). In addition to its unique genital morphology, Quickia is of cytological interest because all five species studied have 25 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 the Indian 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 the biology 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 successive generations were produced during the following two years. Methods of laboratory culture are described in Patterson (1971, 1972). The following locality data and habitat description were provided by J. F. Peake (personal communication) also of the British Museum (Natural History). The specimens were collected by J. D. Taylor while participating in the Royal Society Expedition (1967-68) to Aldabra Atoll. 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 somewhat brackish water. They were also found hidden under rock slabs, in cracks or other places of probable protection from desiccation. The climate of Aldabra Atoll is dry for most of the year with a short wet period which does not necessarily occur regularly. Dissections of relaxed, alcohol-preserved specimens were made under 120 x or 250 x magnification using a Wild M5 stereoscopic microscope. All drawings were prepared with the aid of a camera lucida attachment. For details of specimen preparation and dissection see Patterson (1971). SPECIES DESCRIPTION AND OBSERVATIONS Quickia aldabraensis, sp. n. Shell Shells from both field-collected and laboratory-reared Quickia aldabraensis vary from attenuate to somewhat more ovate (Plate i and Figs. 3, 4). The same kind of variation was noted among Q. spurca shells (Patterson, 1968). The attenuate form of adult laboratory-reared Q. aldabraensis has three whorls and measures 8-5-9 mm in height and 4-5-5 mm in width. The teardrop-shaped aperture is 575-6 mm high and 4 mm wide. More ovate shells measure 8-5-9 mm m height, 5-5-6 mm in width and have an aperture 6 mm high and 4 mm wide. The largest shells may reach a height of 10 mm. Large adult shells collected from the field (Fig. 3) were comparable in size to those reared in the laboratory. Shells cleansed with sodium hypochlorite have a translucent light amber colour. Mud was adherent to most shells obtained from the field and appeared to be arranged in three spiral ridges on some specimens while no definite arrangement was discernible on others. Mud ridges were not present on shells of laboratory reared snails. Fine growth lines provide a minimal amount of shell sculpture. The outer shell surface is dull but the inside is very glossy. There is a well-defined columellar plait and a weakly developed, untwisted columellar fold. The peripheral margin of the shells is rounded with moderately impressed sutures. The holotype (Plate i) is deposited in the mollusc 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) and the University of Michigan, Museum of Zoology. Jaw The jaw is small but relatively strong. Jaw colour varies from a medium brown to a translucent amber, with darker brown markings on the cutting piece and basal accessory 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 extremities which bend toward the central longitudinal body axis. The anterior margin of the cutting piece is deeply convex with a median prominence varying from scarcely noticeable to conspicuous. The sides of the accessory plate slant slightly inward and 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. There are 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 a strikingly 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 and anterior mantle border varies from almost none to a maximum amount shown in Fig. 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 foot of 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 is shown in Fig. 7. The ovotestis is of moderate size and consists of 70-80 acini. It has a dull, light beige coloration. The ovotestis duct is very narrow as it departs from the ovotestis and has a white colour. It soon becomes distended to serve as the seminal vesicle along most of its length. The seminal vesicle portion is usually coiled once and folded three times along its length. The duct again becomes a narrow channel as it opens into the upper portion of the spermoviduct. There are two subequal, narrow receptacula seminis and a well-developed fecundation pouch (Fig. 8). Both have a translucent white colour. The cream-coloured albumen gland is of moderate size, has a linguiform shape and follicular appearance. The tubules 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 along their length. The inner folds of tissue along the uteral portion of the oviduct are visible through the duct wall. The spermatheca is nearly spherical, of moderate size and cream coloured. Its duct passes along the proximal ventral wall of the uteral 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 is quite short in this species. The walls of the anterior portion of the oviduct and vagina become thicker and somewhat muscular in appearance. The prostate gland is of moderate size and white in colour. It is composed of a rather loose collection of 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 the prostate gland. It then passes dorsally over the oviduct and inserts at the apex of 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, thin penial retractor muscle originates adjacent to the vas deferens, passes under the right tentacle assembly and courses posteriorly to its insertion in the tissue covering the 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. It has a muscular appearance and opens to the exterior via the oval-shaped genital aperture (Fig. 9). Chromosome number The haploid chromosome number of Quickia aldabraensis is n = 25, the same number observed in Q. spurca, Q. bensoni and Q. calcuttensis , as well as in members of two other unidentified Quickia from Tamilnadu, India. Fig. 2 shows a camera lucida drawing of a cell in late meiotic diakinesis. There were no chromosomal anomalies observed in any cells and chromosome pairing appeared to be completely normal. FIG. 2. Meiotic chromosomes of Quickia aldabraensis. DISCUSSION AND TAXONOMY Prior to 1968, the genus was considered to be monotypic, containing only the type species, Quickia concisa (Morelet). Subsequently, 'Succinea spurca Gould, 'Succinea bensoni Pfeiffer and a recently discovered Indian species (Q. calcuttensis Patterson) have also been found to belong to the genus Quickia. Q. aldabraensis is now the fifth species included in the genus. The Liberian Q. spurca differs in several respects QUICKIA ALDABRAENSIS SP. N. 181 ovotestis common reproductive channel cutting piece basal ccessory plate ovotestis duct receptacula seminis fecundation pouch prostate gland ovotestis duct receptacula 3£j-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 to either. For comparative purposes, Table I gives a short summary description of various distinct morphological and antomical characters of these three species.1 TABLE i Morphological characters of Quickia Q. aldabraensis Q. spurca non-granulate granulate white character shell sculpture shell colour radula formula head -foot colour prostate gland receptacula seminis penis position (ventral view) penial retractor muscle insertion amber to reddish amber Q. concisa granulate white 14-15:8-9:1:8-9: 12-14:9-10:1:9-10: ii : 10 : i : 10 : i 14-15 12-14 reddish white ; branching tubules ; relatively large highly subequal right of vagina posteriorly translucent white white ; loosely branching tubules small slight subequal to equal usually left of vagina posteriorly translucent white yellow ; small slightly subequal to equal left of vagina anteriorly Connolly (1925) mentioned the collection of Succinea mascarenensis Nevill (More- let) Nevill from Grand Terre, Aldabra Island. Apparently this is the only occasion of assigning a name to the Aldabran succineid. Madge (1938) correctly established that 'S.' nevellei Crosse and 'S.' mascarensis Nevill are synonyms of 'S.' concisa. He further stated that 'S.' mascarenensis Nevill (Morelet) is probably also a synonym of 'S.' concisa. However, the name mascarenensis used by Morelet (1882) does not have validity since his name was an alteration of Nevill's name mascarensis and, according to Madge (1938), was based on a mis-identification because the specimens were not comparable to those of Nevill. Quickia aldabraensis is thus the second valid species of the genus known to inhabit Indian Ocean islands. The presence of mud ridges on shells of Quickia concisa has, in the past, been a key character used in its identification. However, the presence of mud ridges on shells of 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 anterior insertion of the penial retractor muscle in Q. concisa, but the specimens were otherwise unsuitable for a more complete anatomical study. QUICKIA ALDABRAENSIS SP. N. 183 specific character of Q. concisa. From my examination of a specimen of Q. concisa from Rodriguez and published results of Quick's (1936) observations of Q. concisa from various localities, the most reliable species specific character for Q. concisa is the anterior insertion of a short penial retractor muscle. Q. concisa appears to be the only succineid species, studied to date, with such a condition. Based on present information, 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. spurca from Liberia, while being the most widely separated geographically, appear to be the most closely related (morphologically) of the species. There is an indication that Q. concisa is rather distantly related to Q. spurca and Q. bensoni. Q. calcuttensis and Q. aldabraensis have evolved to be most different from the other species. It is difficult 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 chromosome numbers are associated with snail taxa which are considered morphologically more advanced by systematists (see Burch, 1965 ; Patterson, 1969). Odhner (1950) stated that 'Quickia and Indosuccinea evidently both represent a more primitive stage 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 Succineidae hitherto examined'. One is now forced to consider whether simplification of the terminal reproductive system is instead a morphologically more advanced stage of evolutionary development which was accompanied by an increase in chromosome number derived through aneuploidy over a long period of time. Indosuccinea has a more simplified terminal male genital development but a lower chromosome number (n = 24) than Quickia. Indosuccinea could have experienced an aneuploid reduction of one (or more) bivalent (s) from a higher chromosome number or evolved in its own direction from a predecessor with a lower chromosome number. The question of the evolutionary relationship of the various succineid genera is unsettled, especially in the 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 the broadest distribution. However, anatomical validation of the species from many areas is necessary to substantiate such a vast distribution determined largely from i84 C. M. PATTERSON identifications often based only on shell characters. There are no records of the occurrence of Q. aldabraensis on any island in the Indian Ocean other than Aldabra Atoll. It would be interesting to know if additional Quickia species have evolved in other 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 of permanent fresh water. Two species (Lithotis rupicola Blandford and Succinea bernardii Recluz) are actually found in running water of falls (Patterson, 1973^. Some tropical succineids are arboreal. Quickia is interesting because some of its species are adapted to areas devoid of permanent standing or running water. These species depend only on seasonal rains for moisture. Usually the snails are found on 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) in the absence of monsoon rains. Q. calcuttensis from Calcutta and two other popula- tions of Quickia located in Tamilnadu (Madras), India do inhabit moist areas adjacent to sources of permanent fresh water. Because of its adaptability, Quickia has been able 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 the Australian 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 are due to John Taylor and C. A. Wright of the British Museum (Natural History) for collecting and sending the specimens of Q. aldabraensis to me. I would also like to express my appreciation to John Peake also of the British Museum (Natural History) and J. B. Burch of the Museum of Zoology, University of Michigan for critically reading the manuscript and for their many kindnesses during the study. SUMMARY (1) A morphological-anatomical description of Quickia aldabraensis, a new species from 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 particularly closely related to each other. (5) Members of the genus Quickia occupy a variety of habitats including those which are extremely dry while other species are found near areas of permanent fresh 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 FROM MARION ISLAND By F. W. E. ROWE & A. M. CLARK THE specimens which form the basis of the present report were collected at Marion Island in the Southern Ocean (approx. 47° S, 37° E) by Mr A. F. de Villiers during the 1972/73 South African Expedition sponsored by the Department of Transport at Pretoria and sent to the British Museum for identification. Marion and the adjacent Prince Edward Islands were the subject of another South African expedition in 1965/66, the holothurians from which were reported on by Pawson and the other echinoderms by Bernasconi in 1971. Both these authors remark on the zoogeographical affinities of the fauna with that of Kerguelen to 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 and the remaining echinoderms by A. M. Clark. HOLOTHURIOIDEA Pseudocnus 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 at 4 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 ; 2 specimens. 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 body wall increases posteriorly. The spicules are more or less cone-shaped, ranging generally 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 size is no jum x 50 /nm. Unfortunately in many cases the spicules have been eroded by initial storage in formalin. However, the complete spicules compare closely with those figured by Pawson (1968) from what he considers to be the type specimen of Pentactella laevigata Verrill from Kerguelen, though the present specimens are all much smaller. Similarly, direct comparison of the spicules of these Marion Island specimens 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 treat Theel's C. serrata with its varieties as conspecific with P. laevigata Verrill. Dissection of several specimens shows that hermaphrodite gonads are present in specimens over 15 mm length (partly contracted), though the egg follicles are empty in the smaller individuals and I doubt whether even the larger specimens are fully mature. No brood pouches are present and only in the largest syntype of C. 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 in his 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. OPHIUROIDEA Amphiura 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 that cannot be attributed to the larger size (d.d. 6-5 mm) of the type, the largest Marion Island specimen at d.d. 3-5 mm being slightly larger than Bernasconi's holotype of Nullamphiura marionis. The discrepancies in the published descriptions of the two nominal species, such as the contiguity of the adoral shields (said to be touching in A . tomentosa and separate in N. marionis - appearing abnormally so in the pair figured by Bernasconi in 1968) are attributable to variation. Indeed Bernasconi (1971) notes that the adorals are 'rarely joined'. The distinctive widely separated radial shields of larger specimens (d.d. 3 mm or more) agree with Bernasconi's and Lyman '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 the present Marion Island specimens as well as in the holotype of A. tomentosa. I think that the lack of imbrication in the disc scales of the latter is due to the somewhat distended condition of the disc. The shape of the oral shields is rather variable but 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 tentacle scales, as Bernasconi notes. Her supposed difference that A. lymani has smaller distal oral papillae than proximal (infradental) ones, whereas the reverse is the case 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 infradental papillae. In the Marion Island specimens the distal papillae are variable not only in size but also in shape, being either pointed or rounded at the tip. The dorsal arm plates have their distal edges flattened medially in these Marion Island speci- mens and can better be described as fan-shaped than rhombic, as Bernasconi describes them. The ventral arm plates are pentagonal. As for the generic position of this species, in 1970 I rejected Nullamphiura Fell on the grounds that the number of tentacle scales unsupported by other characters is inadequate for a generic distinction from Amphiura, being variable in several species and resulting in artificial grouping of otherwise morphologically diverse species (Clark, 1970). In fact, the holotype of Amphiura tomentosa does have a few pores showing a rudimentary scale, though these are quite lacking in the smaller Marion Island specimens. It is surprising that A. tomentosa has not been reported again from Kerguelen despite 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 ; 14 specimens. 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 plating irregular. In contrast, out of a total of 168 specimens from South Georgia - the type 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 Ross Sea, 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 no obvious morphological difference between them and those specimens from other localities which share their irregular disc plating. One of these specimens has a number of ? loricates in the grooves between the disc 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 with several emerging and loose young. This species was evidently not taken by the 1965/66 expedition, though collected by the 'Discovery' Investigations at 90 or more metres off Marion Island. igo F. W. E. ROWE & A. M. CLARK ASTEROIDEA Anasterias 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 record for the species, although one of Bernasconi's had R 49 mm. REFERENCES BERNASCONI, I. 1968. Equinodermos de las Islas Marion y Principe Eduardo, con descricion de 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 fig8- 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 modal number of vertebrae (28, cf. 27) and a slightly longer caudal peduncle and has relatively shorter pectoral fins, features suggesting adaptation to flowing waters. The new species also appears to exhibit sexual dichromatism in the dorsal fin, an unusual feature in substrate-spawning Tilapia. INTRODUCTION THE new Tilapia described herein was collected by Dr Eugene K. Balon* while he was stationed at the Central Fisheries Research Institute (UNDP/FAO), Chilanga, Zambia, in 1970. Fish samples were taken in the Luongo River as part of Dr Balon's stock and production assessment programme for Lake Mweru and its drainage area. Analysis of the Luongo fish collections is continuing ; a complete taxonomic 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 enters the Luapula River about 20 km upstream from Johnston Falls. On the Luongo about 20 km upstream from its mouth Musonde Falls may have historically formed a barrier to upstream movement of some fish species. Recently a dam was built in the vicinity of Musonde Falls, completely isolating upstream areas. The new Tilapia was collected at two localities above Musonde Falls : (a) Nsenga (or Insenga) stream, a shallow, clear forest stream, 2 -6m wide with banks overgrown with brush, (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 Museum SL = Standard length The upper jaw is measured from the anterior point of the upper lip to the posterior end of the maxilla, the lower from the anterior point of the lower lip to the posterior end of the jaw, here the position of the retroarticular. The preorbital bone is measured from the middle of its orbital rim along a line continuing 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 the penultimate at its base, even if it is smaller than the penultimate. A simple ray fused basally to the penultimate is not counted. Tilapia baloni n. sp. HOLOTYPE. ROM 28120 : male, 136 + 30 mm, Nsenga stream, tributary of Upper Luongo. PARATYPES. ROM 28120 : 5 males, 86 + 23-5 to 114 + 28 mm, 6 females, 83 + 22 to 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 above Musonde Falls. BMNH 1974.4.23.1 : male, 138 + 36-5 mm from Luongo River above Musonde Falls (formerly ROM 28071). BMNH 1974.4.23.2-4: female, 101 + 27 mm and 2 males, 100 + 26 and 85-5 + 22 mm from Nsenga stream (formerly ROM 28120). DESCRIPTION. Meristic characters from holotype and all paratypes, proportions from 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) ; length of head 30-4-32-0 ; length of pectoral fin 23-0-29-0 (over 28 in only 2) ; length of caudal peduncle 13-3-17-2, 14 or more in all but i specimen (0-96-1-25 times its depth, less than i-o in only 2 specimens). Proportions as % length of head. Length of snout 33-9-38-4 ; diameter of eye 24-6-29-0 ; depth of preorbital 18-0-21-6 ; interorbital width 33-0-36-6 ; length of 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 4 in lower, 40-50 in outer row of upper jaw. Outer teeth bicuspid with the main cusp 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 the joint often bluntly bifid. Microbranchiospines present on outer sides of 2nd, 3rd and 4th arches. Lower pharyngeal bone (Fig. 2) with a median length of 24-3-29-4% length of head, 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, between origin 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) ; last spine 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 rounded corners, 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 tube with scalloped rim. Loose ovarian eggs with long diameter about i -9 mm in a fish of 100-5 mm. Intestine in a specimen of SL 95 mm a little over 3 times SL, this and the stomach containing fine dark debris including parts of vascular plants, sparse fragments of filamentous 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 ; flanks more diffuse green with 9 or 10 dark vertical bars and 2 horizontal bands present or absent (present in all preserved fish) ; scales of lower half of flanks and caudal peduncle each with a dark spot either centrally or at base. In the larger male main 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, distally with 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 dorsal and postero-dorsal corner of caudal the same colour. In females lappets dark or dusky, 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. A search through the British Museum and University of Michigan collections of T. sparrmanii from the Luapula basin and the Bangweulu Region revealed no overlooked specimens of the new species and none was encountered during the survey of the Kafue Flats in which one of us (D. J. S.) took part in 1969/70. We have both examined catches in the Mweru basin without finding it. Dr Balon informs us that there is topographical and geological evidence that the upper Luongo once drained to the Kalungwishi, an eastern tributary of Lake Mweru. Some headwater streams of the Kalungwishi River rise in the swamps of the Kawambwe plateau, whose waters also feed the Luongo. The fish fauna of the upper Kalungwishi is not well known. (The 'undescribed species from the Lake Mweru area' related to T. sparrmanii mentioned by Thys (1968 : 369) was T. ruweti and not T. baloni.} AFFINITIES. T. baloni most resembles T. sparrmanii A. Smith, within whose general area of distribution it is found, but T. baloni is a more attenuate fish with a more acute head. The different shape of the body is expressed in the lower ratio of depth to length and the shape of the caudal peduncle as well as the number of vertebrae (Table i). There is also a higher modal number of scales, but since this is true not only of the lateral line series but also the number between lateral line and origin of dorsal fin and rows on the cheek it may not be related to the presence of an additional metamere, the basic difference being the smaller size of the scales. The pectoral fin is usually relatively shorter, a feature which with the slender form probably indicates a stronger swimmer, with less use of the pectorals as paddles and balancers and more use of the tail for propulsion. At the sizes of our examples some individuals of T. sparrmanii have an additional row of teeth, but this is not constant. The long intestine and its contents in the one specimen examined do not suggest 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 upper Zambezi, with organic debris an important supplement. The two species differ also in colour. In T. sparrmanii the red colour on the flanks takes the form of red edges to the otherwise green scales around and behind the pectoral. In the biggest photographed (male) T. baloni it is a red flush involving the greater part of the operculum and the abdominal region at and below the level of the pectoral nearly to the ventral surface. It is more metallic in appearance than the 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 I3'3-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 29 5 I5j 30 2 6£ D-l.l. 3 f.i9 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 Kawambwa plateau in waters including tributaries of the Luongo (specimens in Chilanga Fisheries Research Station, examined by E. T., others collected by Dr Balon examined by D. J. S.). Its modal dorsal formula is XIV 10, nearly approached by XIV u, and the 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 than T. baloni, with an even shorter pectoral fin. Distinctive elements of the colour pattern are the tricolour band edging the dorsal fin and the light blue and purple-red spots 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 the Upper Luongo. T. ruweti is sympatric with T. baloni in the Upper Luongo, just as elsewhere (e.g. in the Mweru lagoons) it is caught together with T. sparrmanii. I96 E. TREWAVAS & D. J. STEWART The fact that to this extent T. baloni occupies the niche elsewhere belonging to T. sparrmanii suggests that it is either a vicariating species or a subspecies of the latter. We have no evidence so far that the differences are less than specific, and the probability that there are colour and size differences between the sexes in T, baloni is further support for its specific status. Although T. sparrmanii is reported from rivers, it is not a fish of the main streams, but occupies lagoons and backwaters rich in vegetation. T. baloni is probably derived from T. sparrmanii or a sparrmanii-like common ancestor in response to life in a river-bed of steeper gradient than that favourable to T. sparrmanii, with all that it means in terms of locomotion and food. Sexual dichromatism in the dorsal fin of T. baloni is a significant difference between it and other species of Tilapia (as distinct from Sarolherodon) and it suggests that the 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 species with 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 subgenus Coptodon), low total numbers of dorsal rays and the relatively narrow preorbital bone. But there are two features in which T. baloni resembles the species included in Coptodon - the modal number of vertebrae, also 28 in Coptodon, and the presence of 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 one most unlike these species, having a modal 29 vertebrae, and is more advanced in some other ways, and this is another reason for relating T. baloni to the species of T. (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 his notes 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, Ann Arbor, Michigan, for providing D. J. S. with facilities for work on the Luongo River collections ; the authorities of the British Museum (Natural History) for the facilities enjoyed by E. T. including the making of radiographs (by M. McLellan) for vertebral counts and use of the drawing by Sharon Chambers ; Mr Graham Bell-Cross for introducing 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. STEWART Laboratory of Limnology UNIVERSITY OF WISCONSIN MADISON, WISCONSIN U.S.A. TWO NEW NEMATODES PARASITIC IN THE KIWI IN NEW ZEALAND By EILEEN A. HARRIS CONTENTS Page SYNOPSIS ........... 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) apterycis sp. nov., are described from the kiwi (Apteryx sp.) in New Zealand. INTRODUCTION THROUGH the kindness of Mr D. M. Rutherford of the Whangarei Animal Health Laboratory, Whangarei, South Island, New Zealand, a small collection of nematodes obtained from a kiwi (Apteryx sp.) was received for study. So far as the writer is aware, the only record of nematodes occurring in Apteryx appears to be that of Chatin (1884, 1885), who gave a very brief and inadequate description of a form that he called Ascaris apterycis. This species does not seem to have been met with again, and it is therefore not surprising that the present material has proved to be very interesting, consisting as it does of two new species which are described below. DESCRIPTION OF SPECIES Family HETERAKIDAE Railliet & Henry, 1914 Heterakis gracilicauda sp. nov. (Figs i & 2) DESCRIPTION. This description is based upon specimens from the caecum of Apteryx sp. in South Island, New Zealand. The male measures 3-3-4-4 mm in length and 0-17-0-33 mm in maximum thickness and the female 5-6-4 mm and 0-19-0-3 mm, respectively. The head bears 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 this leads 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 the pharynx together with the oesophagus, including the bulb, is 0-52-0-7 mm in the 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 just posteriorly to the nerve-ring at 0-26 mm from the anterior end. No cervical papillae have been observed. The caudal end of the male is long, slender and tapering, ending in a long, thin process. 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 /Ltm in 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 of pedunculate papillae around the sucker, two pairs of sessile papillae lateral to the cloaca and two large pairs of pedunculate papillae lateral to these. The other four pairs are distributed along the tail, and it is just posterior to the hindmost pair that the tail begins to narrow gradually. The spicules are short, alate and constantly subequal, measuring 0-24-0-29 mm for 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. The left spicule is constantly longer than the right and each spicule bears a small process at the tip, as shown in Fig. 2b. The vulva is situated anteriorly to the middle of the body, at approximately 2 mm from the anterior end. This leads into a muscular vagina which runs posteriorly for a short distance before opening into the 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 Heterakis into species-groups, based upon the structure and the relative lengths of the spicules. The present species in having subequal alate spicules falls into their 'Heterakis gallinarum group. Of the species in this group, H. gracilicauda is most similar to H. isolonche Linstow, 1906, but differs from it in the length of the spicules, those of 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 spicule and i : 7 for the right, in H. gracilicauda it is I : 20 for the right spicule and i : 15 for the left. It appears that in all the male specimens of H. gracilicauda examined the left spicule is longer than the right. This is somewhat unusual, because in other members of the 'H. gallinarum' group, the relationship of the left spicule to the right is reversed. It is interesting to note that in the collections of the British Museum (Natural History) there are specimens of H. gallinarum and of H. isolonche from Australia and of H. beramporia from New Guinea*. It would seem, therefore, that the 'H. gallinarum' species-group as denned by Inglis, Schmidt & Kuntz is quite widespread in 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 of Apteryx 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 of 30-40 /mi. The dorsal and ventral lips are well developed and rather deeply bilobed, each bearing a median process. Both the lobes of each lip carry two n 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) (Pis i & 2). A large papilla-like structure is visible on each of the lateral lips and posteriorly to this there is a small amphid. The oesophagus is, as usual, in two parts, a short anterior portion and a longer posterior portion, the total length being 1-1-1-8 mm in the male and 1-6-2-3 mm in the female. The nerve-ring is situated anteriorly to the junction of the two oesophageal 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 male are exceedingly difficult to make out, as the tail of each male specimen is, as so often happens in spiruroid nematodes, coiled in a tight, double spiral. Nevertheless, the caudal alae are wide and well developed, and the tail is covered over a wide area of its ventral surface with cuticular elevations, which are arranged in longitudinal rows, 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 found in 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 are distributed at the tip of the tail. They are arranged in two rows, each of four papillae, with the fifth pair lying centrally between the two rows. The spicules are 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 shorter and 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, the gubernaculum 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, flattened mucron at the tip (Fig. 5). The vulva is situated at 2-6-4-6 mm from the anterior end. 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 anteriorly for a short distance before bending posteriorly to run parallel with the other branch of the uterus (Fig. 6). The eggs measure 40-50 p.m by 24-28 | 0-3 mm FIG. 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 is regarded by these authors as the only valid species known hitherto in this group, for both C. (C.) graphophasiani Yamaguti, 1935, and C. (C.) euplocami Maplestone, 1930, are considered to be synonymous with C. (C.) eurycerca. C. (C.) apterycis differs from the latter species in having shorter spicules, and in the more anterior position of the vulva. It may also be distinguished by the number and distribution of the caudal papillae on the male tail. In C. (C.} apterycis there are six pairs of large, pedunculated papillae plus five pairs of small, sessile papillae, whilst in C. (C.) eurycerca there are only five pairs of large papillae, one pair of adanal and four pairs of small sessile papillae. The new species also possesses a barb-like tip to the left spicule. Of the species described since Chabaud & Rousselot presented their key, only C. (C.) ochotensis Belogurov & Zueva, 1967, and C. (C.) singhi Ali, 1961, can be referred to the C. (C.) eurycerca group. C. (C.} ochotensis, described from Falco subbuteo in the U.S.S.R., differs from the present form in the greater length of its spicules. C. (C.) singhi, from Centropus sinensis in India, can be distinguished from C. (C.} apterycis by its much longer right spicule, by the number of cloacal papillae and by the apparent possession of only two pairs of teeth on the 'pseudolips'. C. (C.} casuarii (Maplestone, 1932) has been placed by Chabaud and Rousselot in a separate group of its own. This species is mentioned here because it appears to be the only member of the subgenus to have been described hitherto from an Australian 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 in the 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 to the 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 its death, but it is more probable that the infestation was a natural one rather than one 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 Unit of the British Museum (Natural History) for the electron-micrographs. Thanks are also 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 birds of the Far East. Zool. Zh. 46 : 999-1008. CHABAUD, A. G. 1958. Essai de classification des nematodes Habronematinae. Annls Parasit. 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 i Cyrnea (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 a stimulatory organ (diverticulum atrii, stylophorus or sarcobelum) in different species cannot be the peremptory argument for uniting them in a single genus. For example, in a recent revision of the group (Van Goethem, in press) it was clearly demonstrated that three species formerly united in a single genus, Atoxon Simroth 1888, could be divided into two groups on the basis of a wide variety of morphological characters. These groups have been recognized as separate genera ; the first includes Atoxon fasciatum Verdcourt 1965, while the second is represented by Atoxon meridionals Forcart 1967 and Atoxon bruggeni Forcart 1967. The specimens described below certainly represent a new species and though it lacks 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 posterior end. Mantle posteriorly completely fused to the cephalopodium ; covering an internal shell ; mantle aperture present, forming a small, medio-dorsal slit at the posterior end of the mantle. Shell unguiform, presenting a more or less bilateral symmetry ; nucleus at the posterior end, medially ; protoconch oval. Jaw strongly arcuate, with a median projection on the ventral margin. Radula with median and laterals tricuspid ; marginals lacking the endocone, bicuspid (some main marginals unicuspid, the ectocone being reduced or lacking), very slender, much longer than the laterals ; mesocone of the marginals extending far beyond the posterior margin of the basal tooth plate. Pulmonary cavity little vascularized ; heart transverse ; aorta divided in two branches immediately after leaving the ventricle. Reproductive system : epiphallus with a small bursa calcifera and a long tubular caecum ; hermaphrodite gland situated well behind the albumen gland, 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 the penial tube ; penial papilla simple, very small ; praeputium very short ; vagina absent ; epiphallus 2, very short. Spermatophore helicoid, without an angle ; presenting scaly spines over about its whole length ; top blunt ; filiform part very short. Alimentary tract forming 3 loops (intestine with one forwardly directed loop) 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. Material in British Museum (Natural History) : holotype (Reg. No. 1909.5.10.52) and 5 paratypes (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 10-5 Distance between respiratory orifice and anterior mantle margin 6-5 6-5 6-5 Length of respiratory orifice 0-5 0-7 0-5 Distance 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 0-2 Distance 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 situated just behind and beneath the right lower tentacle, above the right anterior angle of 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 fine longitudinal grooves (irregular on the dorsal part) limiting small polygonal tubercles and 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 the mantle, 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, suprapedal groove ; ut, upper tentacle) pedal groove by fine vertical grooves. Caudal aperture running to the posterior end of the sole. Mantle finely and very distinctly granulated. Mantle aperture small, 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 stripe and 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 lateral stripe. 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 rather distinct transition between laterals and marginals ; only 3 transitional teeth ; median tooth slightly asymmetrical. Median and laterals tricuspid. Marginals without endocones, very slender, with a high mesoconal supporting ridge ; main marginals (teeth 25-40) very long, with generally a reduced ectocone (posterior margin of the basal tooth plate not reaching half the tooth length. Only c. 4 irregular teeth, with a blunt mesocone and several ectocones ; generally i rudimental tooth. 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 and 42nd without an ectocone (unicuspid). Only c. 4 irregular teeth ; generally I rudimental tooth. Size: 3-4 x 1-5 mm; c. 39 transversal rows/mm; length of M: 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 and 43rd without an ectocone (unicuspid). Mesoconal supporting ridge of marginals illustrated 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 ; the anterior 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 a level 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, columellar retractor muscle ; per, pericardium ; pn, pneumostome ; re, rectum ; rp, penial retractor muscle ; ur i, primary ureter ; ur 2, secondary ureter ; ve, ventricle) voluminous. Vas deferens short, passing under the spermathecal duct. Epiphallus with 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 around the 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 a thick 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 of the diaphragm near the posterior margin of the kidney (Fig. 9), slightly adhered to the inner side of the penis windings and ending at about the distal end of the penial 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 with numerous high, narrow folds ; containing 2 spermatophores. In situ, the end of the spermathecal sac situated ventrally, on the same level as the insertion of the colu- mellar retractor muscle. Atrium (atrium genitale) well developed, with numerous retractor muscles : at the issue of the oviduct gland, 3 long and strong ventral retractor 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 between penis 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 a terminal lobe on the left side. Vas deferens wider at its distal end. The first section of the epiphallus, 14-5 mm long. Caecum, 9-0 mm long. Penis: distal end of penial sheath (indicated by a small arrow, Fig. I3A) is coincident with the insertion point of penial retractor muscle. Penial papilla very small, smaller than in 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 indicated by a small arrow on Fig. 12. Penial papilla not distinct ; however, a small lobe can 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 nearly i mm) smooth. Outer side of the whorls with 4-5 irregular rows of forwardly directed scaly spines, each spine about o-i mm long. Posteriorly the number of rows 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: Ventral view 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, glandula albuminalis (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 (penial sheath) ; 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 of spines at E in Fig. I4A ; F : Detail of spines at F in Fig. I4A ; G : Detail of denticles at G in 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 mm in 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 part measuring c. i -5 mm) . Visceral organs Visceral cavity extending to the posterior end of the tail. Alimentary tract forming 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 digestive gland extending to the inner side of the forwardly directed intestinal loop. Rectum passing above the basal part of the columellar retractor muscle (Fig. 9, ventral view). 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 a stimulatory organ, but differs from that genus in : 1. the spermatophore having a very short filiform section ; a blunt, rounded apex and numerous forwardly directed spines over almost the whole length of the fusiform 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 and at 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 the number 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 Melderen for 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 make the 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, just before 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, is identified by its skull characters as Equus quagga, the extinct quagga of South Africa. An equid 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, in the museum of the Department of Zoology, University College, London. No written information existed about its history, but the number 41 had been written on the mandible a long time ago, and the late Professor D. M. S. Watson had once informed Dr K. A. Kermack that it was a quagga. Z58i lacks canines and is therefore a female. It is adult ; all the permanent cheek teeth are in wear and their occlusal surfaces much flattened. Infundibula, otherwise known as marks or central cavities, are still present in the upper and first lower incisors, so that the animal could have been 10-15 years old, were its ageing at all comparable with that of horses. Upper first premolars are present on both sides. 7581 is not a horse, hemione or ass. Horse is ruled out by the relatively great width 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 the more dorsal part of the premaxilla, and deep V-shaped internal sulci of the lower molars. The zygomatic width and the normal instead of diminutive size of the occipital rule out the hemione. It cannot be an ass because of the great zygomatic width, insufficient narrowing of the premaxilla, the lack of any marked backward deflection 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 doubt that Z^8i is a zebra, of which four species are known : Grevy's zebra, Equus grevyi Oustalet, 1882, of parts of Somalia, eastern and southern Ethiopia and northern Kenya. Mountain zebra, E. zebra Linnaeus, 1758, of the coastal region of the Cape Province of South Africa and southern South West Africa. Burchell's zebra, E. burchelli Gray, 1824, with a range extending from the southern parts of the Sudan, Ethiopia and Somalia southwards to the Orange Free State and northernmost Cape Province. Quagga, E. quagga Gmelin, 1788, an incompletely striped zebra formerly found in South Africa to the south of the Vaal River, in the Orange Free State and the southern Cape Province. It was heavily slaughtered in the 18505 and i86os and became extinct in the 18705. Specimens in European zoos survived until 1872 (London), 1875 (Berlin) and 12 August 1883 (Amsterdam). A number 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 it was 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 London Zoo from 1858 until 1864 (Sclater, 1901). Its tooth wear is similar to Z$8i. The results of the comparison will be discussed character by character. The positions of all characters except the first are shown on a quagga skull in Fig. I and the skull of Z58i is illustrated in Plates 1-4. 1. The skull is smaller than in Equus zebra. Many E. burchelli approach it more closely, while the E. quagga skull is about the same size. It should be remembered that the quagga is a zoo specimen, that in the hemione and wild ass zoo specimens have 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 have been a zoo specimen itself. Thus the size of Z58i is appropriate for either burchelli or quagga. The back of M3 is more posterior relative to the front of the orbit in Z58i and the quagga skull than in zebra, while burchelli is intermediate. Hence E. quagga and Z58i are rather short faced, but this character appears to be linked with overall skull size in living equids, and both the large Grevy's zebra and large domestic horses have long faces. 2. The interorbital area of the f rentals shows a slight doming in burchelli but is flatter 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 in the 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 marked in specimens from South West Africa. Furthermore Dr D. A. Hooijer has told me 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, so that its affinities are with quagga or zebra. 3. The level of fusion of the two temporal lines to form a median sagittal line is more anterior in burchelli than in zebra. In 36 out of 50 burchelli it was forward of the rearmost level of the zygomatic arch in dorsal view ; in the remainder it was level or behind. In 2 zebra it was anterior, and in 8 level or behind. The quagga skull and Z$8i have the fusion in the more posterior position, thus being unlike most burchelli but not unlike zebra. 4. The external auditory meatus is slanted upwards as it passes laterally in burchelli but emerges horizontally in zebra in which most individuals also have a larger meatal orifice. Both Z$8i and the quagga skull resemble burchelli. 5. The maxilla-nasal suture dips anteriorly as it approaches the top of the premaxilla in both zebra and burchelli, but the phenomenon is more pronounced in zebra in which the top of the premaxilla suture is also more rounded so that a flange of the nasal bone descends a short way behind the premaxilla. Unfortunately the sutures become obliterated in older individuals. They are no longer visible with complete certainty in the quagga skull, but in 7581 they resemble the burchelli pattern. 6. The alisphenoid area on either side of the basisphenoid antero-medially to the articulation for the lower jaw is more nearly horizontal in ventral view in zebra than in burchelli. 7581 and the quagga skull are like burchelli. 7. E. zebra has a longer styloid process between the auditory bulla and the meatus than 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 not nearly so much in Z$8i or the quagga. Among the sample of burchelli 17 resemble zebra, 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 zebra and 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 burchelli it is narrow in 37 and wide in 18 skulls. The flange of the squamosal between the mastoid and the external auditory meatus is also sometimes wide and sometimes slim, but may well be associated with the mastoid width. Of the 37 burchelli with narrow mastoids only n also have a slender squamosal flange, but among the 18 with 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 view and remains at more or less the same level. In the quagga and Z58i the top edge is more strongly curved and ascends posteriorly. Of the burchelli sample 27 skulls are like zebra and 25 like quagga. 11. On the upper molars the protocone or inner pillar is elongated anteriorly in 40 out of 50 burchelli individuals but not in zebra. The quagga skull and Z58i both resemble the majority of burchelli. 12. Cement is better developed around the exposed sides of the cheek teeth of zebra than of burchelli. A crude assessment of 'more' or 'less' cement showed that in only 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 be more like zebra than burchelli. 13. A small P1 is present in 5 out of 10 zebra, these being mostly the younger individuals. It is present on at least one side in 30 of 51 burchelli, is often larger than in zebra, and there are indications that it is less frequent among the older individuals ; 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 present in burchelli it is often accompanied by a concavity in the front part of the medial wall of P2. A relatively large deciduous P1 is present in Z58i, with only a slight concavity on P2, but it is absent on the quagga skull. For this character Z58i thus resembles 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 of zebra although less marked than those appearing in the uppers. This condition agrees with horses (Grossman, 1953 : 400). In old individuals the infundibula disappear from upper and lower incisors. However in burchelli infundibula were present in the lower incisors of only 6 among the 42 individuals in which they occurred in the upper incisors, and in 2 of the 6 they were present in I1s alone. The London sample of burchelli again gives a hint of regional variation in that those from South West Africa appear to have lower incisor infundibula more frequently. In Z58i 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 and burchelli. 15. The posterior edge of the mandibular symphysis in the median line is high and more or less upright in burchelli. In zebra the top edge dips gently as it passes backwards and so curves round at a lower level to become the posterior edge. The height of the posterior edge is thus less than in burchelli. This character is difficult to assess except in sectioned mandibles, but is real enough as can be seen from Lundholm's illustration (1951, fig. 2). Both the quagga skull and Zs8i resemble burchelli. A. W. GENTRY 16. The top surface of the mandibular symphysis is noticeably flat behind the incisors 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 with most burchelli and zebra, even when the latter have very well worn teeth. It does not appear to be linked with a regression of bone around the incisor roots such as has taken place in both the quagga skull and DISCUSSION It seems from the 16 characters considered above that Z$8i can be taken quite safely as E. quagga. The characters can be summarized as in Table i, from which it is seen that Z$8i differs from zebra in 13 of the 16 characters, from burchelli in 5 of the 16, but from quagga in only 2 of 15 characters, one of the two being the scarcely significant presence of Pls. Accepting Z$8i as quagga, one can suggest that skulls of quagga are like burchelli but differ in flatter frontals, a shorter united sagittal line on 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 P1, 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' V x zebra x V v absent - V X slanted v' V X short x V X more upright V intermediate X narrow \ V X curved, high V intermediate X elongated V V X much V x V present X V X in Iis only \' X X tall V V X \/ — = character not visible. A QUAGGA AT UNIVERSITY COLLEGE 223 in the lower incisors and a flatter mandibular symphyseal surface behind the lower incisors. The poor extent to which the occipital top is drawn out backwardly and the strong curvature of the zygomatic arch are also characters which appear to be more distinctive of quagga than of burchelli. These findings are in agreement with the photographs of quagga skulls in Hilzheimer (1912, pi. 7, fig. 4, pi. 8, pi. 9, figs 3-4). The anterior elongation of the protocone is poorly developed in the Berlin skull 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 quagga but 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 the nasals. Z$8i has a longer diastema than the London quagga skull, as seen in the table 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 well as a juvenile at 55 mm. Lydekker (1904 : 428) observed the preorbital depression in the 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 no preorbital depression in £581. The front part of the nasals is bent slightly upwards in the London quagga skull but not in Z^Si. Such an outline is present in about a third of the burchelli sample, although perhaps less frequent in specimens from South West Africa. Hilzheimer (1912 : 93) attributed the strength of this character in the London quagga skull to its having come from an animal which had been captive for a long period. EQUUS BURCHELLI " ZEBRA " QUAGGA FIG. 2. Map of southern Africa to show the approximate original distributions of three zebra species. The original distributions of E. quagga and E. zebra are very uncertain, and their overlap 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. burchelli burchelli, were available for use in the comparisons and it might be alleged that their skulls could have resembled quagga skulls more closely than do those of other burchelli races. As far as skin characters go, it is known that southern populations of burchelli tended to lose the posterior striping, and the beginnings of this trend can be seen even in surviving populations of E. burchelli antiquorum, the next southern- most race to burchelli burchelli. However, this resemblance to quagga may be more apparent than real ; quagga had a brown body colour posteriorly, its posterior striping assumed a pattern unlike burchelli as can be seen on mounted skins in Tring and Vienna (Rau, 1974, figs 19, 21), it had no lighter 'shadow' stripes between the more posterior black stripes as seen in southern burchelli, and its dark stripes were wider 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 arid country lying to the south-west of the range of burchelli (Fig. 2), and that there is no need to suppose that skulls of burchelli burchelli would have approached those of quagga 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 in the latter half of 1927 and thought to be of a quagga. A pencilled note in the museum 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 a quagga or any other species of zebra. It showed the following differences from zebras : 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), and narrows 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 skull in 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 small canines agree well with the Shetland pony, Osteology register 1952.4.1.3, and raise the 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 M3 at occlusal surface 209 208 174 Back M3 at occlusal surface to front pre- maxilla 274 273 254 Diastema length, I3 to P1/2 (right side) 73-5 92 65 Occlusal length, M1 to M3 73 69-8 71 Occlusal length, P1/2 to P4 88-2 87-6 89 ACKNOWLEDGEMENTS I thank Dr Pamela Robinson for allowing me to study the University College quagga 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 read the 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 quagga quagga (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's menagerie. 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 in this 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 mandible just above the number '£581'. Bull. Br. Mus. nat. Hist. (Zool.) 28, 5 PLATE 2 PLATE 3 Palatal 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 pomacanthid to be recorded from Ascension, and the third Centropyge to be recorded from the tropical Atlantic ; it appears to be derived from tropical western Atlantic forms. INTRODUCTION To DATE only one species of angelfish (family Pomacanthidae) has been recorded from Ascension Island in the south Atlantic, namely the widespread Pomacanthus paru (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 specimens of P. paru from Ascension and various West Indian localities, and on coloration and superficial morphology, they clearly appear to be conspecific. We now report a second species of angelfish from Ascension, belonging to the genus Centropyge. Two species of Centropyge have previously been recorded from the tropical western Atlantic, C. argi Woods & Kanazawa, 1951 and C. aurantonotus Burgess, 1974 ; the present 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 in parentheses by those of the larger and smaller paratypes respectively where these differ from the holotype. Dorsal fin rays XIV 16 (last ray divided to base) ; anal fin rays III 17 (last ray divided to near base in holotype) ; pelvic fin rays I 5 ; pectoral fin rays 16, first ray rather closely applied to second ray, all rays branched except for upper 2 and lower 2 rays ; 17 principal caudal fin rays, branched with the exception of the uppermost and lowermost rays, which do not quite reach posterior margin ; three small supplementary rays above and below principal caudal fin rays ; pored lateral line scales 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 below lateral line to origin of anal fin 14 (14, 16) ; circumpeduncular scales 16 ; gill-rakers on 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. Total length 123-8 (123-8, 122-9) '> nead length (measured from front of upper lip to end of opercular membrane) 30-0 (30-7, 31-3) ; snout length 9-9 (9-0, 10-4) ; orbit diameter 11-9 (12-1, 13-3) ; bony interorbital width 7-1 (7-6, 6-4) ; predorsal length 36-3 (37-2, 40-2) ; depth at first dorsal spine 43-1 (44-1, 47-4) ; body width behind 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 dorsal fin 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 base length 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 lip approximately as broad (vertically at front) as lower, upper lip height about a third of orbit diameter. Teeth slender and elongate, close-set, flexible, tricuspid (the central cusp notably longer and broader than the lateral ones), in two principal rows in jaws, the teeth of the 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 separated by a space about a third to a half of length of posterior nostril ; posterior nostril oval, slightly larger than round anterior nostril, separated from eye by a space about equal to its width ; anterior nostril with a short membraneous tube with an elevated posterior flap. A stout spine at corner of preoperculum, its length about three-quarters of orbit diameter ; 20 (25, 23) small spines of somewhat irregular size along upper margin of preoperculum and one spine on lower edge of preoperculum, about a third (about a 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 near upper 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 portion ridged ; from fourth dorsal spine onwards, dorsal fin with scaly base, scalation extending over three-quarters if not virtually all of fin on median and posterior portion ; anal fin with scaly base, scalation extending over three-quarters or more of fin on median and posterior portion ; scales on basal half of caudal fin, with small scales extending out on fin rays for about a third more of length of fin ; pectoral fins scaled only basally ; pelvic fins with small scales extending out on fin rays for up 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 and third, and third and fourth spines, deeply incised, while those between posterior spines and rays only slightly if at all indented. Membranes of anterior portion of anal 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. Origin of pelvic fins below base of pectoral fins ; first pelvic soft ray prolonged, reaching spinous 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 from snout between eyes along dorsal body contour onto upper part of caudal peduncle. Orbit ringed with bright blue ; iris orange. Dorsal fin bright yellow-orange to orange with dark blue distal margin ; anal fin bright to dark blue, becoming bluish orange distally and posteriorly, with bright blue distal margin ; pelvic fins dusky bluish orange with bright blue anterior margin ; pectoral fins hyaline ; caudal fin bright 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 of Centropyge are very similar in body form and counts, but they are distinctive in colour 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 be expected, C entropy ge resplendens is closest to C. argi Woods & Kanazawa and to C. aurantonotus Burgess, both restricted to the tropical western Atlantic. Centropyge argi has a dark blue body, but only the head and chest are yellow to orange (see colour illustrations in Burgess, 1974, and Bohlke & Chaplin, 1968). Centropyge aurantonotus also has a dark blue body but with orange head, chest, dorsal body contour 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 the dorsal margin of the caudal peduncle, which are dark blue in C. aurantonotus but yellow-orange to orange in C. resplendens ; also, the head of C. aurantonotus is orange, while that of C. resplendens is bright to dark blue, with only the snout and dorsal contour yellow-orange to orange. Briggs (1974) stated that much of Ascension Island's shore fauna was apparently derived from the western Atlantic, and that some justification existed for placing Ascension in the West Indian Province of the Western Atlantic Region. Only two Pomacanthidae are known from Ascension : Centropyge resplendens, which is closely related to tropical western Atlantic forms (no Centropyge is known as yet from the eastern Atlantic), and Pomacanthus paru, which is otherwise recorded only from the tropical western Atlantic. (The only record of P. paru from the eastern Atlantic is that of Fowler, 1936, who considered that Chaetodon leachii Bowdich from 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.} The derivation of Ascension Pomacanthidae appears prima facie to be in accordance with Briggs' statement, although it should be emphasized that the pomacanthid fauna of the eastern Atlantic is still relatively poorly known, making genuine comparison difficult. 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 the text and offered suggestions. We are also grateful to George Earnshaw, who collected the specimens, and to Gordon Howes of the British Museum (Natural History), 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 Zoology UNIVERSITY OF CAMBRIDGE DOWNING STREET CAMBRIDGE R. D. SANKEY 24 HILL CLOSE STANMORE MIDDLESEX 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 a single specimen of a cichlid of an undescribed species. Unwilling to describe it from 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. Recently Thys 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 Gabonese fish as to require close comparison with it. Nanochromis gabonicus sp. nov. HOLOTYPE: $, 47 + 13 mm, from a pool by the road from Mitzic to Medouneu, probably in the drainage basin of R. Okano, tributary of the Ogowe. BMNH 1967.10.12.57. CHARACTERS OF GENERIC IMPORTANCE. Pad on roof of pharynx with a groove around its antero-lateral edge and covered with tubercles resembling, under low magnification, the batteries of taste-buds of the pad of Chromidotilapia (see Trewavas, 1974, pi. 4). Microbranchiospines absent. Outer ceratobranchial gill-rakers with two subequal tubercles. At least in female, first and second soft pelvic rays approximately equal, not produced, but coming to a blunt point. Last scale of upper lateral line separated from dorsal fin by only a half-scale. Scales cycloid, the circuli parallel to the edge of the scale (roman), except a few with a gothic arrangement. Although the base of the skull was not examined in this unique specimen, the other characters indicate that it belongs to the group in which the apophysis for the pharyngeals is formed by the parasphenoid alone. DISTINGUISHING SPECIFIC CHARACTERS. A black spot at the tip of each pelvic fin ($). 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 of upper 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 rather few, 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, the lower 4 on the left side, 8 on the right, the last of which is on the caudal fin ; 3 between origin of dorsal and lateral line ; 3 between bases of pectoral and pelvic fins ; 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, not produced, just reaching vent when laid back. Caudal rounded at the corners, the straight 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 or six places ; traces of three broad blotches from dorsum, tapering towards lateral band. Dorsal lappets and upper edge of soft dorsal black. Postero-dorsal edge of caudal black with a white submarginal band ; a black spot at the tip of each pelvic fin. The ovaries of this small fish are partly discharged, still containing a few eggs of about 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 that species. It also has a wider interorbital region, more teeth in the upper jaw (cf. 42 in N. cavalliensis) and a slightly shorter pectoral fin. No black marginal and white submarginal bands were described on the caudal of N. cavalliensis and in that species ripe eggs were 2-0 x 1-5 mm. Those in the type of N. gabonicus may of course 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, namely scaleless chest, middle soft rays of pelvic fin longer than the others. In it scales are also absent from the cheek, top of head and a narrow strip along the back on each side 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) from R. Lindi, Congo system, and finds that both fall within the range of variation of specimens 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 separate occasions show that the incomplete squamation is maintained, at least in that population. Thys, in an addendum to his paper of 1968, claims to have evidence from living specimens that they are distinct. Thys also refers to an unnamed (new) species from R. Ivindo, tributary of the Ogowe, which he relates to N. squamiceps. From his brief characterization of this I believe it must be a species of which Dr Gery has sent me a few specimens. I mention it here to state that it differs from N. gabonicus in having the pelvic fin a little more acute in the female (much more in the male), with a black streak along the first soft rays but no black spot at its tip, a broad naked area on the chest on either 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 except at the edges ; in the male they bear conspicuous series of spots. REFERENCES BOULENGER, G. A. 1899. Materiaux pour la faune du Congo. Poissons nouveaux. Cinquieme partie. 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. Paris 1900 : 98-101. — 1904. Contribution a 1'etude anatomique, biologique et taxonomique des poissons de la famille 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 systematic revision 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 CANCER PAGURUS 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 are described and larval characters within their respective families are discussed. INTRODUCTION ALTHOUGH there are published accounts of one or more larval stages of several hundreds of brachyuran species, less than one hundred of these descriptions include a 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 of crabs reared in the laboratory, there is still a need for detailed re-descriptions, even of single zoeal stages, where the previous accounts are clearly inadequate. The purpose of this paper is to provide such details of the first zoeal stages of three crabs from 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 twice as long as the carapace and slightly longer than the straight rostral spine. Rostral spine with minute spinules. Lateral spines about half carapace length. Carapace with a low anterio-median papilla, a pair of setae at the base of the dorsal spine and posterio-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, with spinules on the distal two-thirds increasing in size towards the tip. Exopod about one-third length of spinous process, with i long terminal seta and 2 shorter ones, of which 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 second maxilliped ; 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, i and 6 setae. Abdomen (Fig. la) : Somite 2 with forwardly directed dorso-lateral knobs ; somites 2-5 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 one dorsal spine, and minute spinules distally ; inner posterior margin with a deep, rounded median notch and three pairs of processes armed with short setules and spinules, those of the distal one-third of the outer process being particularly stout and tooth-like. DISCUSSION. The larvae of Cancer pagurus were first hatched by Thompson in 1829 and have subsequently been described many times (see Lebour, ig28a). However, with the exception 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 known than are those of C. pagurus. Thus, Aikawa (1937) published details, including appendage setation, of the first stage of C. gibbosulus (de Haan), Mir (1961) compared the 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 these accounts, including the details of C. pagurus given here, it is now possible to define generic, and possibly familial, zoeal characters. All of the Cancer zoeae so far known possess well-developed dorsal, rostral and lateral carapace spines, antennae with spinous processes more than half as long as the 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 distinct groups, lateral knobs on only the second abdominal somite and the telson furcae with one lateral and one dorsal spine. In all the species, with the exception of C. anthonyi, the proximal segment of the endopod of the first maxilliped is described as carrying three setae and this, together with the characters listed above, dis- tinguishes these Cancer zoeae from those of all other known brachyurans. Mir also recorded 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 seems generally to be rather conservative amongst brachyuran zoeae, at least within genera* and often within whole families, so that it seemed at least possible that Trask was 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. But this species possesses the strong tooth-like spines on the outer posterior telson process (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 not two 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. Lateral spines arise close to the posterio-lateral carapace margins, directed downwards and slightly backwards. No dorsal spine, but with a pair of setae close to the mid-dorsal line of the carapace. Antennule (Fig. 2d) : Reduced to small hemispherical buds carrying 2 aesthetascs and 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 3 setae, of which 2 are more or less terminal ; scaphognathites with 3 marginal setae and a long plumose posterior process. First maxilliped (Fig. 2g) : Basis with 2, 2, 3, 3 setae ; endopod segments with 2, 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 and 4 or 5 setae ; exopod with 4 natatory setae. Abdomen (Fig. 2c) : Somites 2 and 3 with small dorso-lateral knobs, those of the second somite joined by a slight ridge over the dorsal surface. Somites 4 and 5 widening to the tri-lobed telson ; rounded median lobe not protruding beyond the acute lateral lobes ; three setose processes between the median and lateral lobes on each side. DISCUSSION. Lebour (i928a, b) described the first stage of P. pisum hatched from the egg and attributed to the same species a second-stage zoea taken in the plankton. Her description is very inadequate, but where comparison is possible it agrees with that given here except that Lebour was able to detect the rudimentary antenna and found the median telson lobe to overreach the lateral lobes. The available descriptions of the larvae of other species of Pinnotheres indicate that there is a good deal of morphological variation within the genus. Thus, while the 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 (Hornell and Southwell) (Lebour, i928a, b ; Hashmi, 1970 ; this paper), the laterals are absent in P. taylori Rathbun (Hart, 1935), while none of the carapace spines are present in P. ostreiim (Say) (Hyman, 1925 ; Sandoz & Hopkins, 1947). Similarly, 24I 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. maculatus and P. taylori the telson is the much more typical brachyuran forked type. The degree of development of the antennules and antennae is also very variable, these appendages being greatly reduced or even absent, at least in the early stages, in P. ostreum, P. placunae and P. pisum, moderately developed in P. taylori, and quite normal 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 probably having three, P. ostreum four and P. maculatus five. One result of this variation in development rate is that the pleopods make their appearance at different stages in the 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 stage in 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 far more typically 'brachyuran' development than any of the other described species, since it is apparently the only one in which the sixth abdominal somite is separated from the telson in the late zoeal stages. Clearly, then, there is no difficulty in distinguishing between the known zoeae larvae of Pinnotheres species. A more difficult problem may be the recognition of characters 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 lateral carapace spines are present. Firstly, in all those species for which the information is available the endopod of the maxilla carries only three setae and these are arranged in a single more or less terminal group, or at least not clearly divided into two distinct groups. This character at once distinguishes the Pinnotheridae from almost all other brachyuran larvae, for such a setal armature has been recorded outside this family only in the Leucosiidae and the ocypodid sub-family Ocypodinae (see also below). The three-segmented endopod of the second maxilliped readily distinguishes these ocypodids from both the Leucosiids and the pinnotherids in which this endopod never has more than two joints. Finally, while there are a number of differences between the pinnotherids and the leucosiids in the detailed morphology of the ap- pendages, the most obvious distinction is the simple triangular telson with the closely spaced row of six processes on the relatively straight posterior margin which has been found in every leucosiid zoea so far described ; this contrasts strongly with both the 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 the foreshore 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 ; lateral spines 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, first maxilliped ; j, second maxilliped. Bar scale represents 0-5 mm for a-c, 0-25 mm for e-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 that given 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 larvae of 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 the second maxilliped whereas I was unable to find a seta in this position in my material and it seems to be unusual in the family as a whole (Table i). Using characters of the carapace spines, antennae, telson, maxillae and maxillipeds Aikawa (1937) was able to separate zoeae of the ocypodid genera Macrophthalmus, Tympanomerus (= Ilyoplax), Scopimera and Uca into groups corresponding to the three sub-families based an adult taxonomy. Thirty years later Wear (1968) re- examined the larval situation within the Ocypodidae, information on the larvae of fourteen species belonging to seven genera by then being available, but was unable to obtain any support for the adult classification. However, in separating the larvae into groups Wear gave greatest significance to the presence or absence of lateral carapace spines, and somewhat lesser significance to the form of the abdomen and the degree of development of the antennal exopod. There is, however, a good deal of evidence to suggest that the setation of the mouthparts may reflect taxonomic divisions between larvae more effectively than these more 'obvious' characters. Certainly, on the basis of the setation of the endopods of the maxillules, maxillae and of the second maxillipeds the described ocypodid larvae fall into distinct groups, 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 the presence or absence of lateral carapace spines in described ocypodid zoeae MACROPHTHALMINAE Macrophthalmus depressus Macrophthalmus depressus Macrophthalmus depressus Macrophthalmus dilatatus Macrophthalmus japonicus Macrophthalmus sulcatus Macrophthalmus latreillis Macrophthalmus pacificus Macrophthalmus crinitus Hemiplax hirtipes OCYPODINAE Ocypode quadrata Ocypode platytarsis Ocypode gaudichaudii Uca annulipes Uca annulipes Uca marionis Uca pugilator Uca pugnax Uca minax Uca triangularis SCOPIMERINAE Scopimera globosus Dotilla sulcata Dotilla sulcata Dotilla blanfordi Ilyoplax pusillus Ilyoplax gangetica MICTYRIDAE ? Mictyris longicarpus Maxillule Maxilla Maxilliped 2 endopod endopod endopod i.5 2, 2 (4) o, i, 5 i, 5 2, 2 (4) i, i, 5 i, 6 2, 2 (4) i, i, 6 i,5 2, 2 (4) i, i, 6 i.5 2, 2 (4) i, i, 6 i, 5 2, 2 (4) i, i, 6 i, 5 2, 2 (4) o, i, 6 J, 5 2, 2 (4) o, i,5 L4 2, 3 (5) o, i, 6 L5 2, 2 (4) o, i, 6 , 5 o, 5 0,4 o, 4 o, 4 o, 4 o,4 2 (3) 2, 3 (5) 2, 3 (5) 2, 2 (4) 2, 3 (5) 2, 3 (5) (4-5) , o, 5 — i. 2 (3) o, o, 4 0,4 i. 2 (3) o, o, 5 0,4 I, 2 (3) o, o, 4 o, 4 i, 2 (3) o, o, 4 o, 4 i, 2 (3) o, o, 5 o. 4 i. 2 (3) o, o, 4 o, 4 i. 2 (3) o, o, 4 0,4 i, 2 (3) o, o, 4 o.4 i, 2 (3) o, o, 4 o, i, 6 0, 2, 4 1, 1,4 i, i, 5 o, 5 i, 5 2, 2 (4) i, i, 6 Telson Lateral fork carapace spines spines Source — This paper — Hashmi, 1969 Aikawa, 1929 Aikawa, 1929 Aikawa, 1929 — Hashmi, 1969 Hashmi, 1969 Hashmi, 1969 + Hashmi, 1969 + Wear, 1968 + Diaz & Costlow, 1972 + Rajabai, 1951 + Crane, 1940 Feest, 1969 — Hashmi, 1968 Hashmi, 1968 — Hyman, 1920 Hyman, 1920 Hyman, 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 a single seta, the endopod of the maxilla armed with a total of four or five setae, and the 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 the basal segment of the endopod of the maxillule and the middle segment of the endopod of the second maxilliped unarmed, while the endopod of the maxilla carries a total of 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, the first zoea of Mictyris longicarpus Latreille has setal characters similar to those of the Macrophthalminae 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 the telson, 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 or absence of lateral caparace spines, showing that these characters are not correlated with 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 the British Museum (Natural History) available for study, to divers of the Guildford Branch of the British Sub-Aqua Club who collected the adult Cancer pagurus, and to 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 material of 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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa: the Biology 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, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28 No. 6 of the Zoology series. The abbreviated titles of periodicals cited follow those of 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 533,7-4-. 1 TRUSTEES OF THE 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 Page SYNOPSIS. ........... 251 INTRODUCTION ........... 251 GENERIC DIAGNOSIS AND IDENTIFICATION OF THE HYDROID STAGE . . 252 MORPHOLOGICAL 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 described from 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 information the many species described from the medusa stage cannot yet be evaluated or related to the hydroids. The affinities of those species of hydroids no longer assigned to Obelia are discussed briefly. The genus Laomedea Lamouroux, 1812, is reduced to a synonym of Obelia but the genus Campanularia Lamarck, 1816, is regarded as valid. INTRODUCTION IN many genera of hydromedusae there is specific diversity of the medusa generation while 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 the hydroid species Obelia dichotoma (Linnaeus, 1758) and 0. geniculata (Linnaeus, 1758) and reared to maturity appear morphologically identical, both being referable to 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 from the hydroid, some seventy having been proposed between 1830 and 1948. It seemed 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 species described 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 species previously assigned to Obelia are removed to allied genera. Although an attempt has been made to consider all described hydroid species, only the more important usages of the name of each have been cited. Full reference lists are already available in the works of Bedot and in abstracting journals. The taxonomic status of all of the numerous species described from the medusa generation remains problematical. The difficulties outlined by Russell (1953) and Kramp (1961) still remain, and further rearing work may be necessary before the hydroid and medusa species can be related. References to species described from the medusa before 1910 were provided by Mayer (1910) and Bedot, while those described between then and 1959 were listed by Kramp (1961). The material examined during the present study was drawn mainly from the collections of the British Museum (Natural History). The figures were prepared with the aid of a camera lucida. GENERIC DIAGNOSIS AND IDENTIFICATION OF THE HYDROID STAGE Genus OBELIA Peron & Lesueur, iSioa1 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, iSioa1 : 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, iSioa1 (nom. nov. pro Medusa marina Slabber, 1769) ; by monotypy ; for nomenclatural purposes taken as conspecific with the hydroid 0. dichotoma (Linnaeus, 1758) (van der Hoeven, 1862 : 280 ; Russell, 1953 : 297), not 0. geniculata (Linnaeus, 1758) as proposed by Naumov (1960 : 260 ; 1969 : 281). DIAGNOSIS. Colonial Campanulariidae (sensu Russell, 1953) with free medusae. Polyp generation forming upright colonies, branched or unbranched, variably flexuose ; internodes annulated proximally, supporting hydrothecal pedicel on distal lateral process. Hydrotheca bell-shaped, hydranth with prominent spherical hypostome. 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) in excluding from the genus species with no medusa generation (accommodated in the genera Campanularia Lamarck, 1816, and Gonothyrea Airman, 1864) and also those in 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 Laomedea Lamouroux, 1812, with Obelia as has been proposed by Naumov (1960, 1969). The appearance 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 above diagnosis. The polyphyletic genus Medusa Linnaeus (1758 : 659) was disbanded by Peron & Lesueur (iSioa) who assigned the originally included species to other genera. No species of Obelia was included in the original scope of Medusa, however, and Peron & Lesueur were justified in forming the genus Obelia to accommodate Medusa marina 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, while Peron's 1804 papers (Peron, i8o4a-f) contain no mention of Obelia. It seems that d'Orbigny was mistaken in giving these dates and that the name Obelia was not introduced until 1810, by Peron & Lesueur. The genus Laomedea Lamouroux, 1812 : 184 was proposed to accommodate two species, Sertularia dichotoma Linnaeus, 1758, and 5. spinosa Linnaeus, 1758. The first-named had, however, previously been removed from Sertularia under the name Obelia sphaerulina Peron & Lesueur, iSioa, while the second species is currently referred 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 was largely a reprint of part of the earlier paper (Pe"ron & Lesueur, iSiob). A footnote in the first paper states 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 the genus Laomedea which can thus be considered a junior synonym of Obelia. The next available 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. dichotoma Linnaeus, 1758. Although Nutting (1915 : 28) made Sertularia verticillata genotype of Campanularia, Naumov (1960 : 249) later nominated S. volubilis Linnaeus, 1758 (not sensu Ellis & Solander, 1786), as type-species of Campanularia and made Sertularia verticillata genotype of a new genus, Verticillina Naumov, 1960 : 269. Millard (1966 : 477) commented on this confusion and nominated S. volubilis sensu Ellis & Solander, 1786 : 51 as type-species of Clytia Lamouroux, 1812 : 184. She showed the correct identification of this type-species to be Medusa hemisphaerica1 Linnaeus, 1767 : 1098. However, this species had previously been nominated type-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 reference to the International Commission on Zoological Nomenclature. The availability of the generic name Eucope Gegenbauer (1856) was discussed by Rees (1939)- The genus Schizocladium Allman (1871) was regarded as a junior synonym of Obelia by Bedot (1910 : 470). The sole included species is reduced to a synonym of Obelia dichotoma in the present review (p. 272) . The genus Obelaria Haeckel (1879) was proposed as a nom. nov. for the hydroid stage 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 present seem superfluous and can be regarded as junior synonyms of Obelia. The genus Monosklera von Lendenfeld, i885b : 910, was synonymized under Obelia by Bedot (1916 : 152). The sole included species is regarded as a synonym of 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 can usually be identified using the characters shown in Table I, but occasional specimens occur with characters apparently intermediate between 0. dichotoma and 0. genicu- lata. Usually such specimens are examples of 0. dichotoma with a slight thickening of the internodal perisarc, and can nevertheless be identified from the characters shown. 1 Although this species name has been attributed to Gronovius (1760 : 38), his usage was not strictly binominal (Millard, 1966 '.477). HYDROID SPECIES OF OBELIA 255 SUBSTRATE* •g _c 0 13 tn 03 43 solid substrates and T) 1 tn 1 IN O 13 6 'S 3 c o 13 3 X substrates, less often on algae g ^bc 13 2 43 O 13 3 tn P i rarely on animal or inen substrates ._> i 0 H w hH •S o "o 0) bo tn 1 43 (LI 43 GROWTH HABIT Mature colonies polysiphonic a numerous monosiphonic side g s o ro 1 3 tn (LI CJ 1 Usually monosiphonic, much branched, up to c. 350 mm Monosiphonic ; usually unbrar S e 0 u o •*-> PH H "8 o O tn « 4-1 HH 2 en i 5 --. *m 1o E M H 3 43 CJ O 5 H PU be E _bo £, ntification fMMETRIC INTERNOD Absent ( Absent ( Present o »•" tn h i— i < 0 1 f ^ IN _bc j_, O fr. ^ O bb tn •~-* o tn rt CJ CD W W W a -2 i 1 s u s o g •« s 256 P. F. S. CORNELIUS MORPHOLOGICAL AND OTHER VARIATIONS THE large number of species of Obelia described from the hydroid stage reflects the high degree of morphological variation present. Although between-colony variation is readily apparent and many characters are inconstant in expression, nevertheless such characters have been freely used in diagnosing new species. That much variation is phenotypic has long been suspected (Mayer, 1910 ; Hammett, 1943) but does not seem to have been given due attention in systematic accounts of the genus. All of the new synonymies proposed here result from the adopting of wider specific limits to take account of this variation. The characters used in defining the specific limits are now discussed. COLONY SIZE. Crowell & Wyttenbach (1957) indicated that there are probably no 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. It follows that colony-size is probably of little or no systematic value. They suggested that colonies of C. flexuosa continue their apical growth until natural breakage occurs. 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], while shorter colonies hitherto regarded as more typical of 0. dichotoma occur intertidally and sublittorally where wave-action is greater. Colonies of intermediate lengths occur and on present evidence it appears that colony-size in 0. dichotoma s. lat. is related to wave-action. The status of 0. longissima is discussed further below (p. 271). 0. geniculata colonies at a single locality in Massachusetts were usually shorter than 25 mm (Hammett, 1943). Ralph (1956), however, demonstrated a correlation between length and latitude in populations between 35° S and 41° S around New Zealand. Cold-water specimens were up to 40 mm long, while those from the warmest places measured only 5 mm. Later Ralph & Thomson (1968) showed that in Wellington harbour colonies formed in the austral winter were longer than those formed in the summer. Nevertheless, it appears that variationin colony-size in 0. geniculata is less than in 0. dichotoma. Information on 0. bidentata is apparently lacking. MONOSIPHONIC VERSUS POLYSiPHONic STEMS. Old stems of 0. bidentata are invariably polysiphonic, those of 0. dichotoma are usually monosiphonic and those of 0. geniculata always so. BRANCHING OF COLONY. Several species of Obelia have been proposed on the basis of the pattern of branching of the colonies. However, this appears not to be correlated with other morphological features. Similarly the length of the branches seems unimportant. The influence of habitat on branching is poorly documented, but Ralph & Thomson (1968) reported that low temperatures induced branching in 0. 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 internode length, breadth, length : breadth ratio, curvature, amount of asymmetric internal perisarc 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 and asymmetric (although variable) in 0. geniculata and usually slight and symmetrical in the other two species. Occasional specimens of 0. dichotoma occur in which the internodal perisarc is slightly thickened on one side, but such specimens can be identified by their hydrothecae. Asymmetric internodal thickening is further discussed below (pp. 271, 277). DARKENING OF PERISARC. In large colonies of 0. dichotoma darkening of the internodal perisarc proceeds with age, older parts being dark brown, grading to brown, horn-coloured or transparent in younger parts of the colony. Colonies of 0. geniculata do not become so dark, while the limited material of 0. bidentata available suggests that no darkening occurs. Histochemical studies by Knight (1970) showed that in the normal perisarc tanning process of Campanulariaflexuosa several of the biochemical precursors of melanin are formed and it is possible that this 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 consequent formation of the hydrothecal rim are only partly known (Knight, 1965) and it would seem unwise to attach systematic importance to rim-variation until more is known of the factors responsible. The shape of the hydrotheca appears to be variable in length : breadth ratio, in the angle between the sides as seen in median vertical section and in the degree to which the basal region is curved inwards. Hydrothecae of 0. bidentata are tubular with a curved basal region while those of 0. geniculata usually have walls curved throughout their length (Figs 2, 5). Those of 0. dichotoma are more variable, being tubular to conical (Fig. 4), with or without a curved basal region. A variable amount 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 delimit species otherwise resembling 0. dichotoma, but it appears to be a variable character and 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 always oblique. 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 and in 0. bidentata between 3-26 (Mammen, 1965). In both there is often a smooth central portion. There seems to be no indication at present whether this variation is phenotypic or genotypic. 258 P. F. S. CORNELIUS GONOTHECA. Gonotheca shape is similar in the three species and with the exception of the terminal region is approximately constant. In 0. bidentata and to a lesser extent in 0. dichotoma gonothecae with apparently truncate ends have been 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 that nematocysts do not provide useful specific criteria in Obelia. a. 0. dichotoma and 0. geniculata. The lengths of fresh, undischarged microbasic mastigophores from living hydranth tentacle-tips of the two species are shown in Table 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) 22 6-8 0-0826 All 2-0 Fresh 2O 6-9 0-0444 All 2-0 Fresh 2O 2O 7'3 6-7 0-0857 0-0466 All 2-0 All 2-0 Fresh Fresh, O. bidentata Norfolk (1953.11.16.1) 10 5-6 0-0897 specimen resembled O. longissima sensu Alder All 1-5 Formalin-preserved n= number in sample, ~x= mean, S.E. = standard error. 'Fresh' nematocysts are those taken from living hydranths. from one species fell within the range recorded from the other. Application of Student's /-test showed nematocyst-length to be similar in the two 0. geniculata populations (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 /mix 1-3-1-5 /mi undischarged) from the hydranths, while Weill (1934^ identified basitrichous isorhizas (length 5 /mi) from the tentacles of both hydranth and medusa. Only the latter type has been recorded from 0. dichotoma, from the medusa (Westfall, 1966, as 0. longissima ; measurements not stated). Thus the present recording of microbasic mastigophores in 0. dichotoma is new. However, the identification of the two kinds of nematocyst depends on the presence or absence of 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 in Obelia 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 (I973-9-24-2). (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 being unobtainable. The preserved nematocysts at least were similar in size to those from live hydranths of the other two species. Identification was not attempted as no discharged nematocysts were seen, but the undischarged ones closely resembled those of the other species and may prove to be of the same kind. BIOLUMINESCENCE. Although Hincks (1868) reported bioluminescence only in 0. 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 laminarian and fucoid algae, whereas 0. dichotoma occurs usually on animal and inert substrates and less frequently on algae (Hincks, 1868 ; Hammett, 1943 ; Barrett & Yonge, 1958 ; Fey, 1969). Manton (1942) examined the attachment of the hydrorhiza of Obelia 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 between hydroid and alga. It seems likely, therefore, that the growth-form of O. geniculata is 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 influence morphology. The substrate-preference of 0. bidentata is less well known but is said to be for inert solid substrates and also sand (Vervoort, I946a) to which neither of the other^two species attaches directly. MEDUSA GENERATION. The medusae of the three hydroid species, although variable, 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. spinulosa var. 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. spinulosa Bale ; G. longicyatha Thornely ; Obelia sp. Clarke ; O. oxydentata Stechow ; L. bicuspidata var. 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, J959 ' 3I5 '• 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 perisarc and not usually strongly tanned ; rim of hydrotheca with a variable number of cusps, 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 ripe gonotheca 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. Pedicel ringed throughout or with central smooth portion ; hydrotheca 1-3 times long as broad, bell-shaped, slightly asymmetric, sometimes having folds in the hydrothecal wall between cusps running proximally from the rim ; diaphragm oblique (can appear transverse in optical section) ; rim with 10-20 bimucronate cusps (rarely, simply cusped ; Fig. 2a-c) ; hydranth undescribed. Gonotheca usually an inverted cone with raised aperture ; occasionally truncated with broad aperture, lacking raised 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 Peabody Museum of Natural History no. 3119. Thimble Island, Branford, Connecticut, U.S.A., 23 September 1874, three fragments in spirit, syntypes of 0. bicuspidata Clarke, Yale Peabody Museum of Natural History no. 7265. Hunstanton, Norfolk, England, 26 September 1953, several large colonies in spirit, 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, Leiden Rijksmuseum 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, unless otherwise 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, 20°57' N, ii5°23' E, c. 200 m, 21 February 1929, one microslide of fragments, coll. British East India Company, 1929.4.18.20. OTHER MATERIAL RECORDED. North Atlantic - West European waters from Helgoland (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 ; Vannucci Mendes, 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 some temperate seas in both northern and southern hemispheres. Records furthest from the 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 Zealand and the Chatham Islands (Ralph, 1957, 1961), south and west coasts of the Republic of South Africa (Millard, 1957). REMARKS. The number of bimucronate cusps on the hydrothecal rim may vary between 10 and 20 (Nutting, 1915 ; Mammen, 1965), although at any one locality the range of variation is less. Most specimens have alternating deep and shallow notches around the hydrothecal rim but in some the gaps are of almost equal depth and the bimucronate condition is obscured (Fig. 2b-c). Two kinds of gonothecae have been recorded. The more usual kind is typical of 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 from normal gonothecae by Hamond, while a BM(NH) specimen shows medusae developing within an apparently truncate gonotheca (Fig. 2a). The two nominal species 0. bicuspidata and 0. bidentata were first described on the same page by Clarke (1875). They were synonymized under 0. bidentata by Jaderholm (i904a, b, 19053.), whose usage as first reviser was followed by Mayer (1910), Billard (1912) and Babic (1913). Nutting (1915), however, used 0. bicuspidata as the senior synonym, although including Jaderholm's I904a paper in his synonymy. Since then bicuspidata has been used more frequently than bidentata but Jaderholm's usage 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 height of the colony. Vervoort (i972a), however, placed var. minor Leloup together with L. bicuspidata var. tenuis Vervoort, I946b (a new name for Leloup's variety), in the synonymy of 0. bicuspidata. The second variety, 0. bicuspidata var picteti Leloup, 1932, was distinguished on specimens with long gonothecae. As such variation is common and has no apparent taxonomic value, the separation is not upheld here. There is some confusion concerning Obelia longicyatha Allman, 1877, and another nominal species of the same name, 0. longicyatha (Thornely, 1899). The type specimens of neither species could be located. Allman's species, described from infertile material, was said to have long pointed cusps on the hydrothecal rim reminiscent 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 by Allman equally suggests 0. bidentata, and the affinities of the species remain unclear. Pictet (1893) referred to Allman's species material with developing medusae clearly not of Obelia type and referred his material to Clytia. Vervoort (i946b) described under Allman's species material identical with 0. bidentata except that the hydro- thecae were unusually long (900-950 /urn). The gonothecae were typical for the species. Vervoort (1968) later described other, infertile, material with hydrothecae nearer in size (580-620 /mi) to the measurements given here (320-490 /u,m) and illustrated an oblique hydrothecal diaphragm characteristic of 0. bidentata. Vervoort (ig72a) subsequently referred another infertile specimen to Allman's species, distinguishing it from 0. bidentata on the basis of hydrothecal length. However, it seems unclear at present whether or not specimens of such dimensions fall within the range of variation of 0. bidentata, and the identity of Vervoort 's material is problematical. In contrast the material identified with Allman's species by Rees & White (1966) and re-examined here seems identical with 0. bidentata, having characteristic bimucronate hydrothecal rims and falling within the normal size-range. The other species, 0. longicyatha (Thornely, 1899), was stated in the original description to have such hydrothecal rims and Stechow (1919) and Hummel- inck (1936) justifiably referred it to 0. bidentata. Jarvis (1922) referred to Gonotha longicyatha specimens with bimucronate hydrothecal cusps and truncate gonothecae which seem also to have been typical 0. bidentata. Obelia andersoni Hincks, 1889, from the Mergui Archipelago, can be referred to 0. 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 the species 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, as Gonothyrea longicyatha ; Billard, 1927, as 0. spinulosa ; Fraser, 1944, as Clytia longicyatha 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 resembling those of the other Obelia species until Billard (1927) identified them in the gonotheca. Subsequently Hamond (1957) and Mammen (1965) confirmed the resemblance by observing newly liberated medusae. Adult specimens have not been described and the best available description, of the young stage, is that by Mammen. The young stage 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> ngs 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, monosiphonic hydrocauli ; internodes long, straight or slightly curved, without asymmetric perisarc 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, frequently with minute longitudinal folds in the hydrothecal wall. DESCRIPTION. Main stems of colonies usually monosiphonic, flexuose, I0~35° mm > racemose hydrocauli with shorter lateral branches ; occasional specimens apparently dichotomous (see p. 256). Main hydrocauli frequently tanned 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 greatest breadth ; rim usually even but often sinuous or castellate (Fig. 4), frequently with minute 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. It shows 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 denticulis obsoletis, calycibus [= gonotheca] obovatis axillaribus, pedunculis intortis, caule dichotomo geniculato'). Some of Ellis' hydroid material was eventually incorporated in Sir Hans Sloane's herbarium which formed the basis of the biological collections of the British Museum; but there are no specimens of O. dichotoma in that herbarium. Other Ellis hydroid material, some of it probably figured and hence eligible for typification, was until recently preserved in the Hunterian Museum of the Royal College of Surgeons of England (Royal College of Surgeons of England, 1830, 1860; Harmer, 1931). Fragments of O. 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 was destroyed during the Second World War, and it is virtually certain that only a single specimen, of Nemertesia Lamouroux, 1812, survived (Dobson, 1971; Miss E. Allen, personal communication). The exquisite 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 recorded it 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 rim Breadth at rim HYDROTHECAL PEDICELS Length INTERNODES Length Maximum breadth Length/breadth ratio GONOTHECA Length Maximum breadth MASSACHUSETTS, U.S.A. (1915.3.6.37 ; Fig. 4d) 270-360 290-340 SW FRANCE (I959.II.I7.4) 320-350 22O-3IO IRELAND (I959.9.I7-35) 250-340 250-330 ISLE OF MAN, U.K. (I959.9-I7-37) 2OO-25O 210-230 280-790 750-1200 080-150 C. IO 200-730 930-1150 070-110 C. IO 750-800 (obscure) 210-790 830-930 120-190 c. 7 870-1000 260-320 120-450 430-500 080-110 c. 4 500-590 210-270 HYDROID SPECIES OF OBELIA 269 MATERIAL EXAMINED. Atlantic Ocean - Oban, Argyll, Scotland, 1877, several colonies 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. Caol Scotnish, 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, numerous colonies 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 and microslide, coll. E. T. Browne, 1959.10.17.1-2. Clyde Sea, Scotland, 35m, on stem 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 and angle 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 on microslide, 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 of wood, in spirit, coll. E. T. Browne, 1954.8.3.71. Plymouth, England, 14 March 1898, 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 on microslide, 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). Southend Pier, 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). 'Probably British', 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, Republic of 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 and microslide, pres. Smithsonian Institution, Washington, D.C., 1880.9.27.93. W'oods Hole, 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 on wood in spirit and microslide, coll. Cambridge University Expedition to the Suez Canal, 1928.5.31.47, 49. Pacific Ocean - Port Jackson, New South Wales, Australia, several colonies on algal 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 Suez Canal (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 frequently recorded south of the equator. Most northerly record probably 77° N, 138° E, north of New Siberian Islands (Linko, 1911, as 0. longissima) ; most southerly record 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 perisarc consequently appear intermediate between this species and 0. geniculata. Constant differences between the hydroid stages are given in Table i. Colonies of 0. dichotoma showing variations in the length and nature of branching of the hydrocaulus and in the shape of the hydrothecal rim have been given specific status by some authors (see also p. 256). Alder (1857), Hincks (1868) and some subsequent authors assigned specimens with long, strongly tanned, dark main hydrocauli to 0. longissima (Pallas, 1766), retaining only specimens with shorter, less tanned main hydrocauli in 0. dichotoma. However, although specimens resembling 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 distinguished from hydrocauli of 0. dichotoma, and the two taxa appear inseparable. Thus specimens hitherto assigned to longissima are probably simply older colonies of dichotoma s. str. Originally Pallas (1766) introduced longissima as an alternative, perhaps more appropriate, name for Linnaeus' species but Linnaeus (1767) gave his own name priority. Johnston (1838, 1847) also regarded longissima as the junior synonym. Obelia flabellata (Hincks, 1866) represents an intermediate point in this series and can be regarded as conspecific with 0. dichotoma.1 Obelia commissuralis McCrady, 1857, was founded on a medusa of which the hydroid was imperfectly known. Agassiz (1862) provided detailed descriptions of both stages and subsequent descriptions of the hydroid alone were given by Nutting (1915) and Fraser (1944). Berrill (1949), although describing the develop- ment of stolons, hydranths and medusae, did not describe systematic characters of his 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 the confusion surrounding the validity of the medusa species it is probably best to regard these synonymies with caution. Thus, Sars (1835) described the medusa Thaumantias plana which Nutting (1915) and Stechow (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. longissima hydroid (= 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 per internode and the arrangement of the side-branches. These characters are variable in 0. dichotoma and on present evidence 0. commissuralis appears invalid. The original description of 0. australis von Lendenfeld, i885a, is inadequate for identification, but the subsequent restriction by Bale (1888) is clearly referable to 0. dichotoma, as also is the non-type material identified as 0. australis by von Lendenfeld (1886.6.8.102). Recently the species was distinguished by Ralph (1957) solely on the presence of an oblique hydrothecal diaphragm, a character noted by Bale and present in von Lendenfeld's material. However, this is a common variation in specimens of 0. dichotoma from British localities (Fig. 4e-f) and its occurrence in southern populations cannot be considered grounds for maintaining 0. 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 originally likened the species to 0. dichotoma and his illustration, showing a much-branched campanularian 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, is assigned to 0. dichotoma. Bedot (1925 : 302) had previously placed it in 0. dubia Nutting, 1901, which Millard also assigned to 0. dichotoma. The original description did not mention gonothecae, and those described by Fraser (1944) differed from those of 0. dichotoma only in being stolonal. It seems probable that Fraser's material was simply a growth-form of 0. dichotoma. Campanularia obtusidentata is a name ascribed to Vanhoffen (1910 : 272) by Bedot (1925 : 302) in the latter's synonymy of 0. dubia Nutting. However, the name does not occur in Vanhoffen's paper and seems to have been used by Bedot alone. It is clearly a lapsus for obtusidens. MEDUSAE. Those reared from both this species and from 0. geniculata have been found to resemble 0. lucifera (Forbes, 1848) (summary in Russell, 1953). Their relation with the hydroid is discussed below (p. 278). They were first recorded by Baster (1762) who saw their release on 3 June 1757. The subsequent history of the elucidation of the life-cycle was reviewed by van Beneden (1844) and briefly by Hincks(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 & Edwards1 ; C. caulini Chiaje1). 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 & Edwards1 ; Schizocladium ramosum Allman1). 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 Allman1 ; 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. b2 ; Dover, Kent, England ; intertidal, on fucoid alga. Present location of specimen 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 illustration shows a specimen having typical Obelia gonothecae but lacking hydrothecae. Linnaeus' designation 'Sertularia denticulis obsoletis, calycibus [= gonotheca] obovatis subrostratis, caule geniculato flexuoso simplici' similarly describes the gonotheca, but omits reference to the hydrotheca. It is likely that the specimens in the Linnean herbarium of the Linnean Society of London (Savage, 1945) were not before Linnaeus 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 Ellis stated: '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 to have come from the earlier locality, Dover, as the Harwich material possibly came to Ellis too late for an illustration to be prepared. He recorded it on 'podded Fucus' , which could refer either to Fucus vesiculosus 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 Sertularia echinata Linnaeus, 1761, to be described in a later paper (Cornelius, 1975: in press], indicates that it at least had reached Linnaeus by or during 1761. This is evident since the original designation of S. echinata includes 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' hydroid material at the end of 1766. Unfortunately the correspondence between Linnaeus and Ellis during 1763 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 to Linnaeus 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 of shortening and thickening. Scale = 500 [xm throughout. DIAGNOSIS OF SPECIES. Obelia hydroid usually with unbranched, erect, flexuose, monosiphonic hydrocauli ; internodes short, curved, with prominent internal thickening of perisarc below origin of pedicel ; hydrotheca even-rimmed. DESCRIPTION. Colony comprises attached stolons from which arise vertical monosiphonic hydrocauli, usually unbranched, up to 40 mm (Fig. 5). Hydrocaulus flexuose ; internodes short, curved, internally thickened below origin of each pedicel, usually with one to five proximal annulations. Pedicel variable in length, attached to short lateral process near distal end of internode, annulated throughout or with smooth central portion. Hydrotheca broad, even-rimmed, bell-shaped, length about equal to width at rim. Gonotheca situated on short annulated pedicel in axil of hydrotheca ; conical, wider distally, apex domed with narrow tubular aperture. Measurements - see Table 5. MATERIAL EXAMINED. Atlantic Ocean -No locality (? Baltic), probably after 1758, 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 comprising about 20 erect hydrocauli. All the hydrothecae are missing, and only two gonothecae remain. Sheet 1298.20 bears four specimens, one at the top of the sheet, two side by side in the centre and one at the bottom. The top specimen is a piece of Laminaria sp. Lamour. to which a colony of O. geniculata is attached. It has many hydrothecae but no gonothecae. Of the two centre specimens that on the left is a piece of alga with no hydroids attached, and that on the right is another piece of alga, probably Fucus 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 identified as it is decayed. It appears from an attached label to have been added later. These specimens are probably 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 270-390 250-310 230-320 Maximum 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. 3 Maximum 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, on Fucus 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 Laminaria saccharina (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 colonies in 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 September 1898, several hydrocauli on microslide, coll. E. T. Browne, 1959.9.17.30. Great Britain Rock, St Mary's, Scilly Isles, on Laminaria sp., 10 m, 22 July 1964, several colonies 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, spirit material and microslide, coll. W. J. Rees, 1962.11.7.47. Nez de Joburg, NW. of Cherbourg Peninsula, France, on Laminaria sp., 3 May 1965, 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 and microslide, pres. Smithsonian Institution, 1890.8.23.14. Potts Point, South Harspowell, 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 Town Ecological 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. von Lendenfeld, 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), Jan Mayen Island, Iceland, coast of Norway at least to 68° N (Broch, 1918) ; Barents Sea, European coast and Mediterranean Sea (Naumov, 1969) ; fjords of West Greenland (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 (Vannucci Mendes, 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 absent only from the northern Indian Ocean and the tropical West Atlantic (although subsequently recorded from the Caribbean). Apart from South Georgia and Macquarie Island there appear to be no records also from the Southern Ocean, and the species appears unrecorded from much of the Pacific, including northern parts of 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 Greenland by Broch (1918) and was not found by Kramp (1932) in some west Greenland collections, Calder (1970) nevertheless recorded the species from both east and west Greenland, but did not cite material. REMARKS. Variation in the hydroid stage of this species occurs mainly in the angle of flexure between internodes, the asymmetric thickening of the internodal perisarc, 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 a series ranging from specimens with shallow angles between internodes, little inter- nodal thickening and proportionately long internodes to more flexuous specimens with greater internodal thickening and shorter internodes (Fig. 4a-c). The occurrence of these variations has prompted authors to recognize several varieties and formae. Vanhoffen (1910), however, was of the opinion that the following 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 described by Ralph (1956) based on variations in length of hydrocaulus, number of annulations of 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 present not to regard them as systematically valid. The species appears to be variable within the above limits over the whole of its range. Thus, specimens showing the extreme of shortening and internodal thickening have been reported from Chile, Kerguelen Island and the Republic of South Africa (Jaderholm, igosb ; Vanhoffen, 1910 ; Fig. 5c), while specimens with both minimal and maximal thickening are known from the English Channel (1959.9.17.30). Almost the full range of variation is represented in British specimens in the BM(NH) collection, and the naming of distinct varieties seems unnecessary. Hammett & Hammett (1945) followed the seasonal morphological changes in colonies of 0. geniculata during several summers in Massachusetts but variations in features of accepted systematic importance were not reported. More recently detailed accounts of variation in New Zealand populations by Ralph (1956) and Ralph & 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 varied with 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 specimen was subsequently destroyed in the Second World War (Redier, 1967). Campanularia coruscans Schneider, 1897, was originally stated to release a medusa and was subsequently assigned to Obelia by Stechow (i92ib, 1923^. The wide distal 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 found to resemble 0. lucifera (Forbes, 1848) (summary in Russell, 1953). Their relation with 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), who commented 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 having been 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 are conspecific (Kramp, 1961). As noted by Russell (1953), it is at present impossible to relate them to the hydroid species, and it may remain so until further rearing work has been done and the characters of the medusae reassessed. Although all three species recognized from the hydroid stage occur in British seas, only two nominal species of medusae are known. The five taxa recorded are as 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 plankton 0. 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, this cannot be the whole answer if, as is shown above, 0. longissima is conspecific with 0. dichotoma from which 0. lucifera has been reared. The occurrence of the hydroid 0. bidentata in the North Sea and English Channel, and the fact that the appearance of 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 the two species have different distributions in western Europe. Thus, the hydroid occurs no further north than the coasts of France, Belgium and Holland and the southern North Sea (Vervoort, ig^6a. ; Leloup, 1952 ; Hamond, 1957 ; Kramp, 1961 ; Teissier, 1965), while the medusa is known from as far north as western Scotland, Iceland and Bergen, Norway (Browne, 1905 ; Thiel, 1932 ; Kramp, 1939 ; Rees, 1953). Sir Frederick Russell (personal communication) is of the opinion that in the seas around the British Isles 0. nigra is associated with mixed oceanic and coastal water typified by the presence of Sagitta elegans (Chaetognatha), while 0. lucifera occurs in coastal water of which S. setosa is characteristic (Meek, 1928). 0. nigra is said to be larger than 0. lucifera and to have darker pigmentation around the tentacle bases (Russell, 1953). It has long been suspected that the mixed water is more productive than British coastal water (Russell, 1939), and as suggested by Browne (in Kramp, 1927) the larger size characteristic of 0. nigra might simply reflect better feeding opportunities. Thus it is possible that medusae from all three hydroid species develop 0. lucifera characters in coastal water and those of 0. nigra in more productive areas. It is noteworthy that although Browne recorded both species of medusae off Plymouth between 1897 and 1899, published records suggest that he never obtained them together (Marine Biological Association, 1957). This perhaps indicates that the furthest extent of mixed water into the English Channel then varied in position around Plymouth. Although mixed oceanic and coastal water frequently reached Plymouth in the 1920*3, it did not do so for some decades after the early i93o's (Russell et al., 1971). During that time 0. lucifera was the prevalent species (Russell, personal communication), further suggesting a correlation with water mass and, possibly, feeding opportunity. Although it might be inferred from the available evidence that the two medusa species are conspecific, the possibility that they are genetically distinct nevertheless remains open, while another possibility is that the hydroid of 0. nigra is as yet undescribed. SPECIES TRANSFERRED TO OTHER GENERA Although Campanularia gelatinosa (Pallas, 1766) was reported to release a medusa by van Beneden (1844), his illustrations identify his material as Obelia dichotoma. Hincks (1868) was aware of this misidentification, but independently described a medusa stage in C. gelatinosa and placed the species in Obelia. Later Maitland (1876 : 13) identified the medusa Obelia marina (Slabber, 1769) with Campanularia gelatinosa auct. (? sensu Hincks). A medusa has not been reported since in the species, and recent opinion (Vervoort, ig46a ; Naumov, 1969 ; J. Clare, personal communication) is that one is not released. It is possible that Hincks mistook the large ova characteristic of C. gelatinosa for developing medusae. Campanularia denticulata Clarke (1876 : 9, pi. i, fig. 4) has usually been referred to Obelia (references in Bedot, 1912-25) although Pictet (1893) assigned it to Clytia noliformis 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 species should nevertheless be referred to Clytia. Obelia marginata Allman (1877 : 9-10, pi. 6, figs 1-2) is now accepted as a species of Cnidoscypus Splettstosser, 1929 (Vervoort, 1968). The two nominal species Obelia longicyatha Allman, 1877, and 0. longicyatha Thornely, 1899, are discussed above under 0. bidentata (p. 264). Eucope annulata von Lendenfeld (i885a : 602-603, pi. 28, figs 53-57) was placed in Obelia by Bedot (1925 : 298), but von Lendenf eld's illustration shows an approxi- mately hemispherical medusa quite unlike Obelia and Kramp (1961) referred the species to Phialella Browne, 1902. Campanularia serrulata Bale (1888 : 757, pi. 12, fig. 4) was referred to Obelia by Thornely (1899) and Mayer (1910, mis-spelt as 0. serratula) but later transferred to Clytia by Bedot (1918, 1925). Although the triangular cusps on the hydrothecal rim shown by both Bale and Mayer are typical of Clytia, the gonotheca described by Thornely was unlike that of Clytia in having a rounded, truncate top. However, as the gonotheca may have been immature, it is still possible that the three authors were describing the same species. Nutting (1927), however, provided the new name Obelia thornelyi for Thornely's material. Nevertheless, the nature of the hydro- thecal rim described by both Bale and Mayer suggests that their material at least should 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 it had a castellated hydrothecal rim. Bedot (1918 : 196 ; 1925 : 300) mis-spelt the name as castellana. Campanularia kincaidi Nutting (1899 : 743-744, pi. 62, figs 2a-c) was referred to Laomedea by Leloup (1940 : 21) and to Obelia by Rees & White (1966 : 277). Its creeping 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 probably best excluded from Obelia. Obelia linearis Thornely (1899 : 453, pi. 44, fig. 6), although retained in Obelia by Mayer (1910 : 257), was transferred to Clytia sp. by Mammen (1965 : 21). The presence of a deeply cleft hydrothecal rim and a reflexed gonothecal aperture support Mammen's opinion. Obelia delicatula Thornely (1899 : 453, pi. 44, fig. 7) was assigned to Clytia by Stechow (i923b) and Blackburn (1942). Although the medusa is unknown, its stolonal gonotheca and sharply-cusped hydrothecal rim are features typical of Clytia 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 not been shown to produce a medusa and is here provisionally referred to the genus Campanularia. The species has also been recorded by Nutting (1915) and Vervoort (i972b, as Laomedea (Obelia} austrogeorgiae). As reported by Jaderholm, gonothecae are absent from the schizoholotype material (1960.8.29.34) which was examined during 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 assigning the species to 0. bidentata. Obelia striata Clarke (1907 : 9-10, pis 6-7) is a distinctive hydroid known since the 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 species to a particular genus with confidence, but the four tentacles of the young medusa suggest that it is not an Obelia species. Vervoort (1968 : 19) considered that Laomedea tottoni Leloup, 1935, was very similar to 0. striata Clarke. Laomedea (Obelia} bistriata Leloup (1931 : 4-6, figs 8-n) was referred to Clytia hendersonae Torrey, 1904, by Mammen (1965) and to C. gravieri (Billard, 1904^ by Vervoort (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 are otherwise unreported from Clytia, Vervoort's observation that the developing medusa has a 'strongly convex umbrella', and Millard & Bouillon's that it has four tentacles, 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 genus Obelia by Naumov (1960, 1969) are here referred to the genera Campanularia and Gonothyrea. The species involved are Campanularia gelatinosa (Pallas, 1766), C. gracilis1 Sars, 1850 (non 0. gracilis Calkins, 1899, = 0. dichotoma), C. flexuosa (Hincks, in Alder, 1856) and G. loveni (Allman, 1859), none of which releases a medusa. ACKNOWLEDGEMENTS I am grateful to R. W. Sims and Dr W. Vervoort for detailed criticism of the manuscript, 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 unpublished observations 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 Falkland Islands. 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 with literature. I was fortunate in being permitted to work at that Laboratory and to make use of its research vessels, and similarly fortunate in being able to work for a short 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 kindly allowed me to study Linnaeus' hydroid material and correspondence with John Ellis, and Miss Elizabeth Allen of the Hunterian Museum of the Royal College of Surgeons 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. pelagica van Breemen, 1905. 28a P. F. S. CORNELIUS Specimens were lent by Dr W. D. Hartman of the Peabody Museum of Natural History, Yale, and Dr W. Vervoort of the Rijksmuseum van Natuurlijke Historic, Leiden. Fig. 3a was drawn by Miss J. B. Garfath and Figs 2a, 5a and 5b by Miss J. M. Paul. REFERENCES AGASSIZ, A. 1865. Illustrated catalogue of the Museum of Comparative Zoology, at Harvard College. No. II. North American Acalephae. Cambridge, Massachusetts. AGASSIZ, L. 1862. Hydroidae. Contributions to the natural history of the United States of America. Second monograph, 4 : 181-380. Boston. AHRLING, E. 1885. Carl von Linnes brefvexling. Stockholm. ALARD, M. i8na. F-loge historique de Francois Peron. Paris. — i8nb. £loge historique de Francois Peron. 2nd edition. Paris. ALDER, J. 1856. Descriptions of three new British zoophytes. Ann. Mag. nat. Hist. (2), 18 : 439-441. — 1857. A catalogue of the zoophytes of Northumberland and Durham. Trans. Tyneside Nat. Fid Cl. 3 : 93-162. ALLMAN, G. J. 1859. Notes on the hydroid zoophytes. Ann. Mag. nat. Hist. (3), 4 : 137-144. - 1864. On the construction and limitation of genera among the Hydroida. Ann. Mag. nat. Hist. (3), 13 : 345-380. — 1871. On a mode of reproduction by spontaneous fission in the Hydroida. Q. Jl microsc. Sci. (N.S.) 11 : 18-21. — 1877. Report on the Hydroida collected during the exploration of the Gulf Stream by L. F. de Pourtales, assistant United States Coast Survey. Mem. Mus. comp. Zool. Harv. 5 (2) : 1-66. AUDIAT, L. 1855. F. Peron (de Cdrilly). Sa vie, ses voyages et ses ouvrages. Moulins. BASIC, K. 1913. Bemerkungen zu den zwei in der Adria vorkommenden thecaphoren Hydroiden. Zool. Am. 43 : 284-288. BALE, W. M. 1888. On some new and rare Hydroida in the Australian Museum collection. Proc. Linn. Soc. N.S.W. (2), 3 : 745-799. — 1924. Reports on some hydroids from the New Zealand coast, with notes on New Zealand Hydroida generally, supplementing Farquhar's list. Trans. Proc. N.Z. Inst. 55 : 225-268. BARRETT, J. H. & YONGE, C. M. 1958. Collins pocket guide to the sea shore. London. BASTER, J. 1762. Opuscula subseciva, observationes miscellaneas de animalculis et plantis. Volume i. Haarlem, Holland. BEDOT, M. 1901. Materiaux pour servir a 1'histoire des hydro'ides. ler periode. Revue suisse Zool. 9 : 379-515. - 1905. Materiaux pour servir a 1'histoire des hydro'ides. 2me periode (1821 a 1850). Revue suisse Zool. 13 : 1-183. — 1910. Materiaux pour servir a 1'histoire des hydro'ides. 3me periode (1851 a 1871). Revue suisse Zool. 18 : 189-490. — 1912. Materiaux pour servir a 1'histoire des hydro'ides. 4me periode (1872 a 1880). Revue suisse Zool. 20 : 213-469. — 1916. Materiaux pour servir a 1'histoire des hydro'ides. 5me periode (1881 a 1890). Revue suisse Zool. 24 : 1-349. — 1918. Materiaux pour servir a 1'histoire des hydro'ides. 6me periode (1891 a 1900). Revue suisse Zool. 26 (Suppl.) : 1—376. - 1925. Materiaux pour servir a 1'histoire des hydro'ides. 7me periode (1901 a 1910). Revue suisse Zool. 32 (Suppl.) : 1-657. BENEDEN, P.-J. VAN 1844. Memoire sur les campanulaires de la cote d'Ostende. Nouv. Mem. Acad. R. Sci. Bruxelles, 17 (5) : 1-42. HYDROID SPECIES OF OBELIA 283 BENTHEM JUTTING, W. S. S. VAN 1970. Martinus Slabber (1740-1835). Amateur-zoolog in Zealand. Archf zeeuw. Genoot. Wet. (1970) : 45-66. BERRILL, N. J. 1949. The polymorphic transformations of Obelia. Q. Jl microsc. Sci. 90 : 235-264. BILLARD, A. igoia. De la scissiparite chez les hydroi'des. C. r. hebd. Seanc. Acad. Sci., Paris, 133 : 441-443. - igoib. De la stolonisation chez les hydroi'des. C. r. hebd. Seanc. Acad. Sci., Paris, 133 : 521-524- — i9O4a. Contribution a 1'etude des hydro'ides. (Multiplication, regeneration, greffes, variations.) Annls Sci. nat., Zoo/. (8) 20 : 1-251. — 1904^ Hydroi'des recoltes par M. Ch. Gravier dans le Golfe de Tadjourah. Bull. Mus. Hist. nat. Paris, 10 : 480-485. — 1909. Revision des especes types d'hydroides de la collection Lamouroux conservee a 1'Institut Botanique de Caen. Annls Sci. nat., Zoo/. (9), 9 : 307-336. — 1912. Hydro'ides de Roscoflf. Archs Zoo/, exp. gen. 51 : 459-478. — 1927. Les hydro'ides de la cote Atlantique de France. C. r. Congr. Socs sav. Paris, Sect. Sci. (1926) : 326-346. BLACKBURN, A. 1942. A systematic list of the Hydroida of South Australia with a summary of their distribution in other seas. Trans. R. Soc. S. Aust. 66 : 104-118. BLANCO, O. M. 1967. Nueva distribution para 'Obelia gracilis' Calkins (un caso de epibiosis). Revta Mus. La Plata, N.S. (Zool.), 10 : 129-136. BREEMEN, P. J. VAN 1905. Plankton van Noord- en Zuiderzee. Tijdschr. ned. dierk. Vereen. (2), 9: 145-324- BRIGGS, E. A. 1939. Hydroida. Scient. Rep. Australas. antarct. Exped. (C), 9 (4) : 1-46. BROCH, H. 1914. Hydrozoa benthonica. Beitr. Kennt. Meeresfauna Westafr. 1 : 19-50. — 1918. Hydroida. (Part II). Dan. Ingolf Exped. 5 (7) : 1-206. BROWNE, E. T. 1900. The fauna and flora of Valencia harbour on the west coast of Ireland. Part i. The pelagic fauna. II. Report on the medusae. Proc. R. Ir. Acad. (3), 5 : 694- 736. - 1902. A preliminary report on Hydromedusae from the Falkland Islands. Ann. Mag. nat. Hist. (7), 9 : 272-284. — 1905. A report on the medusae found in the Firth of Clyde (1901-1902). Proc. R. Soc. Edinb. 25 : 738-778. BUCHANAN, J. B. 1957. The hydroid fauna of the Gold Coast. Revue Zool. Bot. afr. 56 : 349- 372. BUGE, E. 1951. Note sur la synonymic de trois anciennes especes de Bryozaires : Diastopora latornarginata d'Orbigny, 1852, Eschara andegavensis Michelin, 1847 et Obelia disticha Michelin, 1847. Bull. Soc. geol. Fr. (5), 20 : 459-465. CALDER, D. R. 1970. Thecate hydroids from the shelf waters of northern Canada. /. Fish. Res. Bd Can. 27 : 1501-1547. CALKINS, G. N. 1899. Some hydroids from Puget Sound. Proc. Boston Soc. nat. Hist. 28 : 333-368. CAVOLINI, F. 1785. Memorie per servire alia storia de' polipi marini. Naples. CHIAJE, S. DELLE, 1841. Polipi sertolarici. Animale senza vertebre del regno di Napoli (5) : 1-165. Naples. CLARKE, S. F. 1875. Descriptions of new and rare species of hydroids from the New England coast. Trans. Conn. Acad. Arts Sci. 3 : 58-66. - 1876. Report on the hydroids collected on the coast of Alaska and the Aleutian Islands, by W. H. Dall, U.S. Coast Survey, and party, from 1871 to 1874 inclusive. In W. H. Dall, ed., Scientific results of the exploration of Alaska, by the parties under the charge of W . H. Dall, during the years 1865-1874, 1 : 5-34. Washington. - 1879. Report on the Hydroida collected during the exploration of the Gulf Stream and Gulf of Mexico by Alexander Agassiz, 1877-78. Bull. Mus. comp. Zool. Harv. 5 : 239- 252. 284 P. F. S. CORNELIUS CLARKE, S. F. 1894. Reports on the dredging operations off the west coast of central America to the Galapagos, to the west of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer 'Albatross', during 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. XI. The hydroids. Bull. Mus. comp. Zool. Harv. 25 : 71-77. 1907. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer 'Albatross', from October, 1904, to March, 1905, Lieut. -Commander L. M. Garrett, U.S.N., commanding. VIII. The hydroids. Mem. Mus. comp. Zool. Harv. 35 (i) : 1-18. CORNELIUS, P. F. S. 1975. A revision of the species of Lafoeidae and Haleciidae (Coelenterata : Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. Hist. (Zool.) 28 (in press}. COUGHTREY, M. 1876. Critical notes on the New-Zealand Hydroida, suborder Thecaphora. Ann. Mag. nat. Hist. (4) 17 : 22-32. CROWELL, S. & WYTTENBACH, C. 1957. Factors affecting terminal growth in the hydroid Campanularia. Biol. Bull. mar. biol. Lab., Woods Hole, 113 : 233-244. DALYELL, J. G. i836a. Farther illustrations of the propagation of Scottish zoophytes. Edinb. New phil. J. 21 : 88-94. i836b. Fernere Beobachtungen iiber die Fortpflanzung der scottischen Zoophyten. Froriep's Notizen (i) 50 (6) : 81-90. DEEVEY, E. S. 1950. Hydroids from Louisiana and Texas, with remarks on the Pleistocene biogeography of the western Gulf of Mexico. Ecology, 31 : 334-367. DESK AYES, G. P. & EDWARDS, H. M. (Eds), 1836. Histoire naturelle des animaux sans vertebres par J. B. P. A. de Lamarck. Volume 2. 2nd edition. Paris. (Eds) 1840. Histoire naturelle des animaux sans vertebres par J. B. P. A. de Lamarck, Volume 3. 2nd edition. Paris. DOBSON, J. 1971. Descriptive catalogue of the physiological series in the Hunterian Museum of the Royal College of Surgeons of England. Part II. Hunterian specimens demonstrating the products of generation together with surviving Hunterian specimens from other sections. Edinburgh and London. ELLIS, J. 1755. An essay towards a natural history of the corallines, and other marine productions of the like kind, commonly found on the coasts of Great Britian and Ireland. London. & SOLANDER, D. 1786. The natural history of many curious and uncommon zoophytes, collected from various parts of the globe. Ed. M. Watt. London. ESCHSCHOLTZ, F. 1829. System der Acalephen. Berlin. FEY, A. 1969. Peuplements sessiles de 1'archipel de G16nan. i. Inventaire : Hydraires. Vie Milieu (B) 20 : 387-413. FORBES, E. 1848. A monograph of the British naked-eyed medusae: with figures of all the species. London. FRASER, C. M. 1914. Some hydroids of the Vancouver Island region. Trans. R. Soc. Can. (3) 8 (4) : 99-216. 1937. Hydroids of the Pacific coast of Canada and the United States. Toronto. — 1938. Hydroids of the 1934 Allan Hancock Pacific Expedition. Allan Hancock Pacif. Exped. 4 : 1-105. 1941. New species of hydroids, mostly from the Atlantic Ocean, in the United States National Museum. Proc. U.S. natn. Mus. 91 : 77-89. — 1943. Distribution records of some hydroids in the collection of the Museum of Comparative Zoology at Harvard College, with description of new genera and new species. Proc. New Engl. zool. Club, 22 : 75-98. 1944- Hydroids of the Atlantic coast of North America. Toronto. — 1946. Distribution and relationship in American hydroids. Toronto. — 1948. Hydroids of the Allan Hancock Pacific Expeditions since March, 1938. Allan Hancock Pacif. Exped. 4 : 179-335. HYDROID SPECIES OF OBELIA 285 GEGENBAUER, C. 1856. Versuch eines Systemes der Medusen, mit Beschreibung neuer oder wenig gekannter Formen ; zugleich ein Beitrag zur Kenntniss der Fauna des Mittelmeeres. Z. wiss. Zool. (1857) 8 : 202-273. GIRARD, M. 1857. F. Peron naturaliste, voyageur aux terres australes. Sa vie, appreciation de ses travaux. Paris. GOSSE, P. H. 1853. A naturalist's rambles on the Devonshire coast. London. GRANT, R. E. 1826. Observations on the spontaneous motions of the ova of the Campanularia dichotoma, Gorgonia verrucosa, Caryophyllea calycularis, Spongia panicea, Sp. papillaris, cristata, tomentosa, and Plumularia falcata. Edinb. New phil. J. 1 : 150-156. GREGORY, J. W. 1909. Catalogue of the fossil Bryozoa in the Department of Geology, British Museum (Natural History). The Cretaceous Bryozoa. Volume II. London. GRONOVIUS, L. T. 1760. Observationes de animalculis aliquot marinae aquae in-natantibus atque in littoribus belgicus obviis. Acta helv. 4 : 35-40. HAECKEL, E. 1879. Das System der Medusen. Erster Theil einer Monographic der Medusen. Denkschr. med.-naturw. Ges. Jena, 1 : 1—360. HAMMETT, F. S. 1943. The role of the amino acids and nucleic acid components in develop- mental growth. Part one. The growth of an Obelia hydranth. Chapter one. Description of Obelia and its growth. Growth, 7 : 331-399. & HAMMETT, D. W. 1945. Seasonal changes in Obelia colony composition. Growth, 9:55-144- HAMOND, R. 1957. Notes on the Hydrozoa of the Norfolk coast. /. Linn. Soc., Zool. 43 : 294- 324- HARGITT, C. W. 1909. New and little known hydroids of Woods Hole. Biol. Bull. mar. biol. Lab., Woods Hole, 17 : 369-385. 1924. Hydroids of the Philippine islands. Philipp. J. Sci. 24 : 467-505. 1927. Some hydroids of south China. Bull. Mus. comp. Zool. Harv. 67 : 489-520. HARMER, S. F. 1930. Presidential address, 24th May, 1929. Proc. Linn. Soc. Lond. (Nov. 1928-May 1929) 141 : 68-118. 1931. Presidential address. Recent work on Polyzoa. Proc. Linn. Soc. Lond. (Oct. i930-May 1931) 143 : 113-168. HARTLAUB, C. 1884. Beobachtungen iiber die Entstehung der Sexualzellen bei Obelia. Z. wiss. Zool. 41 : 159-185. HILGENDORF, F. W. 1898. On the hydroids of the neighbourhood of Dunedin. Trans. Proc. N.Z. Inst. (1897), 30 : 200-218. HINCKS, T. 1861. A catalogue of the zoophytes of south Devon and south Cornwall. Ann. Mag. nat. Hist. (3) 8 : 251-262. 1866. On new British Hydroida. Ann. Mag. nat. Hist. (3) 18 : 296-299, 1868. A history of the British hydroid zoophytes. 2 volumes. London. 1889. On the Polyzoa and Hydroida of the Mergui Archipelago collected for the Trustees of the Indian Museum, Calcutta, by Dr J. Anderson, Superintendent of the Museum. /. Linn. Soc. 21 : 121-135. HIROHITO. 1969. Some hydroids of the Amakusa Islands. Publ. Biol. Lab. Imp. Household, Tokyo (1969), (9) : 1-32. HODGSON, M. M. 1950. A revision of the Tasmanian Hydroida. Pap. Proc. R. Soc. Tasm. (1949) : 1-65. HOEVEN, J. VAN DER 1 862. Eenige aanteekeningen over Martinus Slabber's Natuurkundige Verlustigingen ; benevens opgave der systematische namen van de daarin afgebeelde diersoorten. Versl. Meded. K. Akad. wet. Amst. (Afd. Nat.) 14 : 270-285. HUMMELINCK, P. W. 1936. Hydropoliepen. Flora Fauna Zuiderzee (Suppl.) : 41-64. ITO, T. & INOUE, K. 1962. Systematic studies on the nematocysts of Cnidaria. I. Nemato- cysts of Gymnoblastea and Calyptoblastea. Mem. Ehime Univ. (2, B) 4 : 445-460. JADERHOLM, E. igo4a. Aussereuropaische Hydroiden im schwedischen Reichsmuseum. Ark. Zool. (1903) 1 : 259-312. 286 P. F. S. CORNELIUS JADERHOLM, E. 19045. Mitteilungen ueber einige von der schwedischen Antarctic-Expedition 1901-1903 eingesammelte Hydroiden. Arch. Zool. exp. gin. (4) 3, Notes et Revue : I- XIV. — i9O5a. Hydroiden aus den Kiisten von Chile. Ark. Zool. (i) 2 (3) : 1-7. — 19055. Hydroiden aus antarktischen und subantarktischen Meeren, gesammelt von der schwedischen Sudpolarexpedition. Wiss. Ergebn. schwed. Sudpolarexped. 5 (8) : 1-41. — 1909. Northern and arctic invertebrates in the collection of the Swedish State Museum. IV. Hydroiden. K. svenska VetenskAkad. Handl. 45 (i) : 1-124. JARVIS, F. E. 1922. The hydroids from the Chagos, Seychelles and other islands and from the coasts of British East Africa and Zanzibar. Trans. Linn. Soc. Lond. (Zool.) 18 : 331-360. JOHNSTON, G. 1838. A history of the British zoophytes. London. 1847. A history of the British zoophytes. 2nd edition, 2 volumes. London. KNIGHT, D. P. 1965. Behavioural aspects of emergence in the hydranth of Campanularia flexuosa (Hincks). Nature, Lond. 206 : 1170-1171. — 1970. Sclerotization of the perisarc of the calyptoblastic hydroid, Laomedea flexuosa. i. The identification and localization of dopamine in the hydroid. Tissue Cell, 2 : 467- 477- KOLLIKER, A. 1843. Ueber die Randkorper der Squallen, Polypen und Strahlthiere. Neue Notizen Geb. Nat. Heilk. 25 : 81-84. KRAMP, P. L. 1927. The hydromedusae of the Danish waters. K. dansk. Vidensk. Selsk. Skr. 8 : 1-291. 1932. Hydroids collected in West-Greenland fjords in 1911 and 1912. Meddr. Gronland, 91 (3): i -35- — 1935. Polypdyr (Coelenterata) . I. Ferskvandspolypper og goplepolypper. Danm. Fauna, 41 : 1-208. — 1939. Medusae, Siphonophora, and Ctenophora. Zoology Iceland, 2 (5b) : 1-37. 1961. Synopsis of the medusae of the world. /. mar. biol. Ass. U.K. 40 : 1-469. LAMARCK, J. B. P. A. DE 1816. Histoire naturelle des animaux sans vertebres. Volume 2. Paris. LAMOUROUX, J. V. F. 1812. Extrait d'une memoire sur la classification des polypiers coral- ligenes non entierement pierreux. Nouv. Bull. Soc. philom. Paris, 3 : 181-188. — 1816. Histoire des polypiers coralligenes flexibles, vulgairement nommes zoophytes. Caen. — 1821. Exposition mdthodique des genres de I'ordre des polypiers. Paris. LELOUP, E. 1931. Trois nouvelles especes d'hydropolypes. Bull. Mus. r. Hist. nat. Belg. 7(25): i -6. 1932. Une collection d'hydropolypes appartenant 1'Indian Museum de Calcutta. Rec. Indian Mus. 34 : 131-170. — 1935. Hydraires calyptoblastiques des Indes Occidentales. Me"m. Mus. r. nat. Hist. Belg. (2) 2: 1-73. — 1939. Hydropolypes marins et dulcicoles du Congo Beige. Revue Zool. Bot. afr. 32 : 418- 423- — 1940. Hydropolypes provenant des croisieres du Prince Albert icr de Monaco. Result. Camp, scient. Prince Albert I, 104 : 1-38. — 1952. Coelentere's. Faune Belg. pp. 1-283. LENDENFELD, R. VON i885a. The Australian Hydromedusae. Part V. Conclusion. The Hydromedusinae, Hydrocorallinae and Trachymedusae. Proc. Linn. Soc. N.S.W. (1884) (i) 9: 581-634. — i885b. Addenda to the Australian Hydromedusae. Proc. Linn. Soc. N.S.W. (1884) (i) 9 : 908-924. LESUEUR, C. A. 1811. Voyage de decouvertes aux terres australes. Histoire naturelle. Histoire generale et particuliere de tons les animaux qui composent la famille des meduses, et de quelques autres radiares molasses. Paris. LING, S. 1938. Studies on Chinese Hydrozoa. II. Report on some common hydroids from the East Saddle Island. Lingnan Sci. J. 17 : 175-184. HYDROID SPECIES OF OBELIA 287 LINKO, A. K. 1911. Hydraires (Hydroidea). Volume I. Haleciidae, Lafoeidae, Bonneviel- lidae et Campanulariidae. Fauna Rossii, pp. 1-250. LINNAEUS, C. 1758. Systema naturae. loth edition. Holmiae. 1761. Fauna svecica, sistens animalia Sveciae regni. 2nd edition. Stockholm. 1767. Systema naturae. I2th edition. Tom I, Pars II. Holmiae. MAITLAND, R. T. 1876. Determinatie der dieren, beschreven en afgebeeld in de werken van Job Baster en Martinus Slabber. Tijdschr. ned. dierk. Vereen. 2 : 7-15. MAMMEN, T. A. 1965. On a collection of hydroids from south India. II. Suborder Thecata (excluding family Plumulariidae) . /. mar. biol. Ass. India, 7 : 1-57. MANTON, S. M. 1942. On the hydrorhiza and claspers of the hydroid Myriothela cocksi (Vigurs) . /. mar. biol. Ass. U.K. 25 : 143-150. MARATTI, J. F. 1776. Plantis zoophytis et lithophytis in mari mediterraneo viventibus. Rome. MARINE BIOLOGICAL ASSOCIATION, 1957. Plymouth marine fauna. 3rd edition. Plymouth, England. MARKTANNER-TURNERETSCHER, G. 1890. Die Hydroiden des k. k. naturhistorischen Hof- museums. Annln naturh. Mus. Wien, 5 : 195-286. MAYER, A. G. 1910. Medusae of the world. Volume II. The Hydromedusae. Washington. McCRADY, J. 1857. Gymnopthalmata of Charleston Harbor. Proc. Elliott Soc. nat. Hist. 1 : 103-221. MEEK, A. 1928. On Sagitta elegans and Sagitta setosa from the Northumbrian plankton, with a note on a trematode parasite. Proc. zool. Soc. Lond. (1928) : 743-776. MEYEN, F. J. F. 1834. t)ber das Leuchten des Meeres und Beschreibung einiger Polypen und anderer niederer Thiere. Nova Acta Acad. Caesar. Leop. Carol. 16 (Suppl. i) : 125-216. MICHELIN, J. L. H. 1847. Iconographie zoophytologique, description par localites et terrains des polypiers fossiles de France et pays environnants. Paris. MILLARD, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43 : 173-243. — 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part i. Calypto- blastea. Ann. S. Afr. Mus. 44 : 165-226. - 1966. The Hydrozoa of the south and west coasts of South Africa. Part III. The Gymnoblastea and small families of Calyptoblastea. Ann. S. Afr. Mus. 48 : 427-487. — & BOUILLON, J. 1973. Hydroids from the Seychelles (Coelenterata) . Annls Mus. r. Afr. cent. 206 : 1-106. 1974. A collection of hydroids from Mo9ambique, east Africa. Ann. S. Afr. Mus. 65 : 1-40. MODEER, A. 1791. Tentamen systematis medusarum stabiliendi. Nova Acta physico-med. 8 (Appendix) : 19-34. MORIN, J. G. & COOKE, I. M. 1971. Behavioural physiology of the colonial hydroid Obelia. II. Stimulus-initiated electrical activity and bioluminescence. /. exp. Biol. 54 : 707-721. NAUMOV, D. V. 1960. Gidroidy i gidromeduzy morskikh, solonovatovodnykh i presnovodnykh basseinov S.S.S.R. Fauna S.S.S.R. 70 : 1-626. - 1969. Hydroids and Hydromedusae of the U.S.S.R. Fauna S.S.S.R. 70. Israel Program for Scientific Translation. NORTON, G. W. 1896. The development of the free-swimming medusae of Obelia commissuralis. Am. mon. microsc. J. 18 : 291-296. NUTTING, C. C. 1899. Hydroida from Alaska and Puget Sound. Proc. U.S. natn. Mus. 21 : 741-753- -1901. Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proc. Wash. Acad. Sci. 3 : 157-216. - 1915. American hydroids. Part III. The Campanulariidae and the Bonneviellidae. Spec. Bull. U.S. natn. Mus. pp. 1-126. - 1927. Report on the Hydroida collected by the United States Fisheries steamer 'Albatross' in the Philippine region, 1907-1910. Bull. U.S. nat. Mus. (100) 6 : 195-242. OKEN, L. 1815. Okens Lehrbuch der Naturgeschichte. Dritte Theil. Zoologie. Jena. 288 P. F. S. CORNELIUS ORBIGNY, A. D' 1853. Paleontologie franfaise. Description zoologique et geologique de tous les animaux mollusques et rayonnes fossiles de France. Terrains Cretaces. 5. Bryozoa. Paris. PALLAS, P. S. 1766. Elenchus zoophytorum. The Hague. PARKE, M. & DIXON, P. S. 1968. Check-list of British marine algae - second revision. /. mar. biol. Ass. U.K. 48 : 783-832. PERON, F. i8o4a. Memoire sur le nouveau genre Pyrosoma. Annls Mus. Hist. nat. Paris, 4 : 437-446- i8o4b. Precis d'un me'moire lu a 1'Institut National, sur la temperature de la mer soit a sa surface, soit a diverses profondeurs. Annls Mus. Hist. nat. Paris, 5 : 123-148. i8o4C. Memoire sur le nouveau genre Pyrosoma. ] . Phys. Chim. Hist. nat. 59 : 207-213. i8o4d. Observations sur la dyssenterie des pays chauds, et sur 1'usage du betel. /. phys. Chim. Hist. nat. 59 : 290-299. 18046. Extrait d'un memoire sur la temperature des eaux de la mer, soit a sa surface, soit a diverses profondeurs, le long des rivages et en pleine mer. /. Phys. Chim. Hist. nat. 59:361-366. 1804! Memoire sur quelques faits zoologiques applicables a la theorie du globe, lu a la classe des sciences physiques et math6matiques de 1'Institut National. /. Phys. Chim. Hist. nat. 59 : 463-479. & LESUEUR, C. A. iSioa. Tableau des caracteres generiques et specifiques de toutes les especes de meduses connues jusqu'a ce jour. Annls Mus. Hist. nat. Paris (1809) 14 : 325- 366. iSiob. Suite de 1'histoire generate et particuliere de tous les animaux qui composent la famille des meduses. /. Phys. Chim. Hist. nat. 70 : 357-371. PICARD, J. 1955. Hydraires des environs de Castiglione (Algerie). Bull. Stn Aquic. Peche Castiglione (N.S.), 7 : 177-199. PICTET, C. 1893. Etude sur les hydraires de la Baie d'Amboine. Revue suisse Zool. 1 : 1-64. PRENANT, M. & BOBIN, G. 1956. Bryozaires. Premiere partie. Entoproctes, phylactolemes, ctenostomes. Faune Fr. 60 : 1-398. RALPH, P. M. 1956. Variation in Obelia geniculata (Linnaeus, 1758) and Silicularia bilabiata (Coughtrey, 1875) (Hydroida, F. Campanulariidae) . Trans. R. Soc. N.Z. 84 : 279-296. 1957- New Zealand thecate hydroids. Part I. Campanulariidae and Campanulinidae. Trans. R. Soc. N.Z. 84 : 811-854. 1961. A checklist of the hydroid fauna of the Chatham Islands. N.Z. Dept. Sci. Ind. Res. Bull. 139 : 235-238. & THOMSON, H. G. 1968. Seasonal changes in growth in the erect stem of Obelia geniculata in Wellington Harbour, New Zealand. Zool. Pub. Victoria Univ. Wellington, 44 : 1-21. REDIER, L. 1966. Hydraires et bryozaires. Cah. pacif. 9 : 77-122. - 1967. Revision de la collection du Museum des hydraires de Lamouroux. Bull. Mus. natn. Hist. nat. Paris. (2) 39 : 381-410. REES, A. 1819. Sertularia. The Cyclopaedia: or, Universal Dictionary of Arts, Sciences and Literature. Volume 32. (Unpaginated.) London. REES, W. J. 1939. A revision of the genus Campanulina van Beneden, 1847. Ann. Mag. nat. Hist, (i i) 3:433-447. 1953- Records of hydroids and medusae taken at Herdla, Bergen in 1937. Univ. Bergen Arb. (1952), (16) : 1-8. — & THURSFIELD, S. 1965. The hydroid collections of James Ritchie. Proc. R. Soc. Edinb. (B) 69 : 34-220. — & WHITE, E. 1966. New records and fauna list of hydroids from the Azores. Ann. Mag. nat. Hist. (13) 9 : 271-284. RITCHIE, J. 1909. Supplementary report on the hydroids of the Scottish National Antarctic Expedition. Trans. R. Soc. Edinb. 47 : 65-101. 1910. The marine fauna of the Mergui archipelago, lower Burma, collected by Jas. J. Simpson, M.A., B.Sc., and R. N. Rudmose-Brown, D.Sc., University of Aberdeen, February to May 1907. -The hydroids. Proc. zool. Soc. Lond. (1910) : 799-825. HYDROID SPECIES OF OBELIA 289 ROYAL COLLEGE OF SURGEONS OF ENGLAND, 1830. Catalogue of the contents of the Museum of the Royal College of Surgeons in London. Part 4, fasciculus i, comprehending the first division of the preparation of natural history in spirit. London. [Anonymous ; compiled by Richard Owen.] — 1860. Catalogue of the contents of the Museum of the Royal College of Surgeons of England. Part i. Plants and invertebrate animals in the dried state. London. [Anonymous.] RUSSELL, F. S. 1939. Hydrographical and biological conditions in the North Sea as indicated by plankton organisms. /. Cons. perm. int. Explor. Mer, 14 : 171-192. 1953. The medusae of the British Isles. Anthomedusae, Leptomedusae , Limnomedusae , Trachymedusae and Narcomedusae. Cambridge, England. — -, SOUTHWARD, A. J., BOALCH, G. T. & BUTLER, E. I. 1971. Changes in biological conditions in the English Channel off Plymouth during the last half century. Nature, Lond. 234 : 468-470. SARS, M. 1835. Beskrivelser og iagttagelser over nogle maerkelige eller nye i havet ved den Bergenske kyst levende dyr af polypernes, acalephernes , radiaternes, annelidernes og mollusk- ernes classer, med en kort oversigt over de hidtil af for fatter en sammesteds fundne arter og deres forekommen. Bergen. 1850. Beretning om en i Sommeren 1849 foretargen zoologisk Reise i Lofoten og Finmarken. Nyt Mag. Naturvid. 6 : 121-211. SAVAGE, S. 1945. A catalogue of the Linnaean herbarium. London. — 1948. Catalogue of the manuscripts in the library of the Linnean Society of London. Part I V. - Calendar of the Ellis manuscripts. London. SCHMIDT, H. 1972. Some new records of hydroids from the Gulf of Aqaba with zoogeographical remarks on the Red Sea area. /. mar. biol. Ass. India (1971) 13 : 27-51. SCHNEIDER, K. C. 1897. Hydropolypen von Rovigno, nebst Uebersicht iiber das System der Hydropolypen im Allgemeinen. Zool. Jb., Syst. 10 : 472-555. SCHYDLOWSKY, A. 1902. Materiaux relatifs a la faune des polypes hydraires des mers arctiques. I. Les hydraires de la Mer Blanche le long du littoral des Jsles Solowetzky. Trudy khar'kov Obshch. Ispyt. Prir. (1901) 36 (i) : 1-276. SHERBORN, C. D. 1929. Index animalium sive index nominum quae ab A.D. MDCCLVIII generibus et speciebus animalium imposita sunt. Letters O, P. London. SLABBER, M. 1769—1778. N atuurkundige verlustigingen, behelzende microscopise waarneem- ingen van in- en uitlandse water- en land-dieren. Haarlem, Holland.1 1775-1781. Physicalische Belustigungen oder microscopische W ahrnehmungen in- und auslandischer Wasser- und Landthierchen. Translated by P. L. St. Miiller. Niirnberg.1 SMITH, J. E. 1821. A selection of the correspondence of Linnaeus, and other naturalists, from the original manuscripts. 2 volumes. London. SPAGNOLINI, A. 1871. Catalogo degli acalefi discofori del golfo di Napoli. Atti Soc. ital. Sci. nat. 14 : 144-223. SPLETTSTOSSER, W. 1929. Beitrage zur Kenntnis der Sertulariiden. Zool. Jb., Syst. 58 : i- 134- SPRENGEL, W. 1813. Philip Cavolini's Abhandlungen iiber Pflanzen-Thiere des Mittelmeers. Nurnberg. STECHOW, E. 1914. Zur Kenntnis neuer oder seltener Hydroidpolypen, meist Campanulariden, aus Amerika und Norwegen. Zool. Anz. 45 : 120-136. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete, nebst Angaben iiber einige Kirchenpauer'sche Typen von Plumulariden. Zool. Jb., Syst. 42 : 1-172. i92ia. Symbiosen zwischen Isopoden und Hydroiden. Zool. Anz. 53 : 221-223. 1 Both editions of Slabber's work were issued in parts which were not dated individually (Benthem Jutting, 1970). Title-pages of the German version dated 1781 carried a slightly amended title, Physika- lische Belustigungen oder mikroskopische W ahrnehmungen von drey und vierzig in- und auslandchen Wasser- und Landthierchen, while later title-pages of the Dutch version retained the original wording. 29o P. F. S. CORNELIUS STECHOW, E. 192 ib. Neue Genera und Species von Hydrozoen und anderen Evertebraten. Arch. Naturgesch. 87 (Abt. A, Heft 3) : 248-265. — iQ23a. Neue Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen iiber einige andere Formen. Zool. Anz. 56 : 1-20. — iQ23b. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete. II Tiel. Zool. Jb., Syst. 47 : 29-270. — 1925. Hydroiden der Deutschen Tiefsee-Expedition. Wiss. Ergebn. dt. Tiefsee-Exped. 'Valdivia' , 17 : 383-546. TEISSIER, G. 1965. Inventaire de la faune marine de Roscoff. Cnidaires - ctenaires. Roscoff, France. THIEL, M. E. 1932. Die Hydromedusenfauna des Nordlichen Eismeeres in tiergeographischer Betrachtung. Arch. Naturgesch. (N.F.) 1 : 435-514. THORNELY, L. R. 1899. The hydroid zoophytes collected by Dr Willey in the southern seas. In : Willey, A., 1898-1902, Zoological results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere. Collected during 1895, 1896 and 1897, 4 : 451-457. Cambridge. - 1904. 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 collected by 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 Historic and 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-Doherty Geological 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. Valeur taxonomique 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 are printed in italic type. Main page references also are in italics. adelungi, Obelia 266 alternata, Eucope 273 alternata, Obelia 267 andersoni, Obelia 260, 264, 265 angulosa, Obelia 266 annulata, Eucope 280 arruensis, Obelia 266 articulata, Eucope 266 articulata, Obelia 266 attenuata, Obelia 260 australis, Obelia 266, 271, 272 austrogeorgiae (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 260 bifurcata, Obelia 260 biserialis, Obelia 267 bistriata, Laomedea (Obelia) 281 borealis, Obelia 266 brasiliensis, Campanularia 265 braziliensis, 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} 280 denticulata, Campanularia 279-2^0 diaphana (Eucope & Thaumantias) 273 dichotoma (Campanularia & Laomedea) 265 dichotoma, Obelia 251, 253, 254, 255, 256- 259 (variation), 265-272 (main section), 275-279 dichotoma, Sertularia 253-254, 265 divaricata, Laomedea 265, 266 dubia, Obelia 266, 272 echinata, Sertularia 273 equilateralis, Obelia 267 Eucope 252, 254 everta, Obelia 267 flabellata, Campanularia 266 flabellata, Obelia 266, 270, 271 (text and footnote) flexuosa, Campanularia 256, 257, 281 flexuosa, Laomedea 265 flexuosa, Sertularia 272 fragilis, Obelia 266 fusiformis, 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, 272 Gonotha 260 Gonothyrea 253 Gonothyrea 281 gracilis, Campanularia 281 gracilis, Laomedea 266 gracilis, Obelia 266, 281 gravieri, Clytia 281 griffini, Obelia 266 gymnopthalma, 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, 277 irregularis, 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 278-279 nigrocaulus, Obelia 266 nodosa, Obelia 266 noliformis, Clytia 279 Obelaria 252, 254 Obeletta 252, 254 Obelia 257-259 (introduction, generic syn- onymy, variation and identification) Obelia sp. 260 Obelia spp. See under specific name Obelissa 252, 254 obtusidens, Campanularia 266, 267, 272 obtusidens, Obelia 267 obtusidentata, Obelia 267, 272 oxydentata, Obelia 260 parasitica, Eucope 265 pelagica, Campanularia 266, 281 Phialella 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, 277 subtropica, var. of Obelia geniculata 273, 277 surcularis, Obelia 266 syringa, 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 254 Verticillina 254 Vesicularia 253 volubilis, 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 SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9-70- 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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, is issued in five series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28 No. 7 of the Zoology series. The abbreviated titles of periodicals cited follow those of 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 OF THE 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 calanoid copepods are described. A further eighteen rare or poorly known species are described or commented upon. Five existing species are transferred to the new genus ; new synonymies are proposed for six species, and two species are recorded as incertae sedis. All the specimens were 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 the vertical distributions and migrations of zooplankton and nekton in the northeastern Atlantic. These investigations have been based largely upon a series of vertical hauls made with horizontally towed opening and closing net systems. The calanoid copepods 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 Fuerteventura in the Canary Islands. Details of the sampling methods used in this series are given by Foxton (1969), Angel (1969) and Roe (19720). The second series was made in 1969 in a position centred on i8°N 25°W near the Cape Verde Islands. A day and a night series of hauls was made with an acoustically controlled net system. This comprises two rectangular midwater trawls, one of 8 m2 mouth area and mesh of 4-5 mm (RMT 8), the other of i m2 mouth area and mesh 0-32 mm (RMT i), combined within the same frame. This net system, designated the RMTi + 8, is described by Baker, Clarke & Harris (1973). The series was taken at 1250-0 m depth and was fished in the manner described by Baker et al. Both these series contained a number of very rare calanoid copepods, several undescribed sexes of known species and some new species. These specimens are all described 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. usttatus Park, 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 are more 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 known species. Since the males of many species of Spinocalanus are unknown and as the present 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 m depth in a position i8°N 25°W, between 12 and 16 November 1969. 44 specimens deposited 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 of i -14 mm (35 specimens). The cephalothorax is stout and 1-7 times as long as the 5-segmented abdomen. In lateral view the cephalothorax is arched dorsally. The head and ist thoracic segment are completely fused and thoracic segments 4 and 5 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 4th abdominal segment. The ist antenna has 19 free segments and reaches to about halfway along the 2nd abdominal segment ; the basal eight segments have large flattened sensory filaments ; the ist segment has a small group of spines. The remaining mouthparts and legs are as shown. A satisfactory preparation of the ist 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 small feeble setae. The swimming legs of all specimens are damaged. The 5th pair of legs reaches back to the hind edge of the 2nd abdominal segment ; the right endopod and 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 of S. 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 is unknown, although Vervoort (1946) mentioned a male of 5. abyssalis var. pygmaeus measuring 1-14 mm. Presumably the males listed by Vervoort (1957) and that described by Bradford (19710;) are too large (1-7-2-2 mm) to be 5. abyssalis. The males 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 - the females 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, 4th teg I j» 5th leg ; k-u, Spinocalanus ? brevicaudatus figs 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 depth off Fuerteventura (28°N I4°W) on 24 November 1963. 3 females from 2 RMT i hauls fished at 600-410 m depth in a position i8°N 25°W on 15 and 16 November 1969. 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 of 2-09 mm (4 specimens). The cephalothorax is 4-1 times as long as the 4-segmented abdomen. The head and ist thoracic segment are fused ; thoracic segments 4 and 5 are 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, mandibular palp ; 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. 332 H. S. J. ROE basally. The last thoracic segment protrudes laterally. The genital segment is slightly longer than the combined length of the three subsequent segments. The furcal 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. The remaining mouthparts are as shown. The endopodite of the 2nd maxilla has five brush-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 as shown. The endopodite of the ist leg has no lobe and no surface spines. The 5th leg has three segments on each side. REMARKS. The present females have been compared with the type specimens of Amallophora 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 the holotype has two setae on the basipodite of the mandibular palp and no apparent segmentation 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 here and have two large brush-like setae and three smaller ones. There is no doubt that the present females are conspecific with Amallophora alter a Farran. The identity of this species is confused and it has been mixed up with Amallothrix auropecten (Giesbrecht, 1892). For comparison the latter species is described below (p. 333). Bradford (1973) described as Amallophora altera a species having one very large and four small brush-like setae on its 2nd maxilla. This arrangement is conspicuously different from both the present material and Farran's types. Bradford's specimen is incompletely described but it also differs from Amallothrix altera in the structure of its rostrum, its ist maxilla and in its ist and 5th legs. It is clearly a different species. Bradford used her specimen to typify a group containing Amallophora smithae Grice, 1962, Scolecithrix vorax Esterley, 1911 and Xanthocalanus typicus Farran, 1908. This group of species is considered here to comprise a new genus which is described later on p. 335. The true Amallothrix altera does not belong in this new genus. Vervoort's (1957) description is also incomplete but his specimens differ from the present species by (a) a short outer edge spine on the ist exopodite segment of the 2nd leg, (b) an incompletely segmented 5th leg, (c) differences in the setation of the 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. Similarly the 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 later material. In 1965 Vervoort described a further specimen which may be Amallothrix altera, but the rostrum appears to be smaller and the endopod of the 2nd maxilla is different, although this may be due to the difficulty in seeing the small brush-like heads. A . altera is generally similar to A . auropecten (Fig. 19) but differs in the following respects, (a) the rostrum of A . altera is larger and of a different shape, (b) the setation of 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 in A. auropecten, (d) the ist basipodite of the maxilliped has a large brush-like seta in A. altera but not in A. auropecten, (e) the endopod of the ist leg has a comb of surface spines in A. auropecten but not in A. altera, and (f) the 5th leg has three segments in A. altera but is incompletely segmented in A. auropecten. This last difference may be variable but all the examined specimens of A. altera have a completely segmented 5th pair of legs whereas those of A. auropecten are normally 2-segmented or incompletely divided into three (Fig. 19 and see Giesbrecht, 1892 ; Rose, 1942 ; Wilson, 1942, 1950 ; Park, 1968 ; Bradford, 1973). One of the present specimens of A. auropecten has an abnormal 5th pair of legs similar to that figured by Park (1968, pi. 9, fig. 3). In 1965 Vervoort transferred Farran's Amallophora altera to Amallothrix. The main 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 inner lobe. 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 enlargement of two of the brush-like setae in the present species is not, by itself, a generic character and it is shown by several other species and genera within the Scolecithricidae (see, for example, A. falcifer, p. 325, Scolecithricella obscura, p. 318, Scaphocalanus difficilis, 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 made at 950-475 m depth off Fuerteventura (28°N I4°W), between 24 and 28 November 1965. 6 females and 6 males taken in 7 RMT I hauls made at 785-300 m depth in a position i8°N 25°W, between 13 and 16 November 1969. DESCRIPTION (Fig. 19) : Body length of the females is 1-9-2-05 mm with a mean of 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 has five brush-like setae, all with small heads, but only four could be seen in the specimen illustrated. The males agree with Rose's account and redescription is unnecessary. The male described by Wilson (1950) as Scolecithricella auropecten is not of this species. REMARKS. Rose (1942) transferred Scolecithrix auropecten to Amallothrix. Bradford (1973) omitted it from her redefined Amallothrix and considered that it should 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, mandibular palp ; 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} laminata differs in many respects from the present species (see p. 320) and is not, in my opinion, closely related. A. altera and A. lobophora, however, do seem to be related to the present species. All three differ from Bradford's Amallothrix in having a Scaphocalanus-type ist maxilla (p. 329). The female 5th legs of all three species and 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 spines on the ist basipodite of the 3rd leg. The rostrum of A . lobophora is similar to that of A . alter a but the structure of its 5th legs (both sexes) , the spiny lobe on the endopod of the ist leg, and the uniformly small brush-like setae on the female 2nd maxilla are all more like A. auropecten. Whether or not these three species are sufficiently alike and 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 of the acceptable generic limits, especially regarding the ist maxilla, will clarify their position. Meanwhile all three species can conveniently remain within Amallothrix. SCOPALATUM gen. nov. DIAGNOSIS. Head and ist thoracic segment fused ; thoracic segments 4 and 5 fused or separate. Rostrum with two filaments, ist antenna with 23 segments in the female, 20 in the male ; there are large sensory filaments on the proximal segments of the male ist antenna. 2nd antenna exopod slightly longer than the endopod. ist maxilla with two posterior surface setae on the ist inner lobe, two setae on the 2nd inner lobe and four on the 3rd inner lobe ; in the female the endopod is ornamented with spinules. 2nd maxilla endopod with five brush-like setae and three flattened worm-like setae ; in the female one of the brush-like setae is greatly enlarged ; in the male two are enlarged. MaxiUiped with one brush-like seta on the ist basipodite. The male mouthparts are weakly chitinized and slightly reduced. ist leg has a 3-segmented exopod and a single endopodite segment : all three exopodite segments have an outer spine ; the endopod has a lobe carrying small spines. 2nd leg has a 3-segmented exopod and a 2-segmented endopod ; the outer edge spines on all exopodite segments are large ; the 2nd and 3rd exopodite segments and the 2nd endopodite segment carry prominent surface spines. 3rd leg has a 3-segmented exopodite and endopodite : the ist basipodite segment has a group of long spines ; the 2nd and 3rd exopodite and endopodite segments have surface spines. 4th leg has a 3-segmented exopodite and endopodite : the ist basipodite may have a group of long spines ; the 2nd and 3rd endopodite segments have very few small surface spines ; the exopodite segments have no surface spines. Female 5th 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 similar to that of Scaphocalanus : left exopod is shorter than the endopod ; right exopod is much 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 male copepod, A. typica, having a peculiar enlarged sensory seta on its 2nd maxilla. Giesbrecht (1892) had previously described a similar structure in the male of Xanthocalanus and considered that Amallophora was a junior synonym of Xantho- calanus (Giesbrecht & Schmeil, 1898). Farran (1908) named a female which he believed 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's male is a xanthocalanid but transferring Farran's female to the Scolecithricidae. She placed this, as X. typicus Farran, into a group together with Amallophora smithae 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 to an entirely different group. Nevertheless, I believe she was correct in collecting together her specimen of 'A . altera' and the other three species into a distinct group within 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 (see below) 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 status within the Scolecithricidae. This new genus is denned above (p. 335). The genus Scopalatum 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 also includes the species 'Amallophora altera' wrongly identified by Bradford (1973) which should be further described and named. Grice's (1962) account of S. smithae is 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 to be modified when these latter two species are fully described. Details of S. gibbera sp. 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 the Scolecithricidae. The combination of characters given on p. 335, especially those of the ist and 2nd maxillae and the legs, distinguishes this new genus from all others in the family. It is related to A . altera, A . auropecten and A . lobophora, see p. 334. These three species may have to be removed from Amallothrix but they differ 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, and both this species and A. altera are further distinguished by having no basipodite spines on either their 3rd or 4th legs. A. lobophora has a similar 2nd maxilla in the male to Scopalatum and both sexes have a group of spines on the ist basipodite of the 3rd leg. They do not, however, have an external spine on all exopodite segments of 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 off Fuerteventura (28°N I4°W) on 13 November 1965. 2 females in 2 RMT i hauls fished in a position i8°N 25°W ; one (the holotype) caught at 300-210 m depth on 12 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, exopodite segments 2-3 (paratype) ; n, 3rd leg ; o, 4th leg ; p, 5th leg. Bar scale o-i mm unless indicated. 338 H. S. J. ROE DESCRIPTION. Female (Fig. 20) : Body length 2-58-2-81 mm with a mean of 2-73 mm (3 specimens). The holotype measures 2-81 mm. The cephalothorax is robust and 3-9 times as long as the 5-segmented abdomen. The head and ist thoracic segment are fused, as are thoracic segments 4 and 5. The mid-dorsal region of the head has a very characteristic protuberance which is particularly conspicuous when seen from the side. The rostrum has two filaments with thickened bases. The hind margin of the cephalothorax is deeply indented laterally. The genital segment has a slight ventral protrusion and is longer than the combined length 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 the ist maxilla is covered with small spines. The endopodite of the 2nd maxilla has five brush-like and three flattened worm-like setae, one of the brush-like setae is greatly 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. The terminal spines on legs 2-4 have laminae. The 5th pair of legs is uniramous with two fused segments on each side attached to a basal segment ; the line of fusion is clearly visible. The terminal segment on each side has one long and one shorter spine. There is an additional small spine on one side only, but this is clearly variable since in one of the paratypes it is about halfway down the outside edge of the fused 2nd 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 belongs to a new species which I have named Scopalatum farrani. The female specimens of S. 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 off Fuerteventura (28°N I4°W) on 26 November 1965. 8 males taken in 5 RMT i hauls made at 900-305 m depth in a position i8°N 25°W 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 of 2-48 mm (9 specimens). The cephalothorax is 2-7 times as long as the 5-segmented abdomen. The head and ist thoracic segment are fused ; thoracic segments 4 and 5 are separate. The rostrum has two filaments attached to a common base. The hind margin of the cephalothorax has a slight lateral indentation. The ist antenna reaches to the hind edge of the 2nd abdominal segment ; both ist antennae have 20 segments and the proximal 12 segments on each side carry large flattened sensory filaments. The mouthparts and legs are as shown. The endopod of the 2nd maxilla has five brush-like setae and three flattened worm-like setae ; the heads of three of the brush-like setae are small whereas two are greatly enlarged - one of them especially so. All the 'brush' heads consist of fine threads ending in rounded swellings. 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 end of 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 exopodite segment 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 (Natural History). The type material consists of several slides on which are mounted the mixed appendages and abdomens of two specimens. There is only one example of most appendages - including the 2nd maxilla, and there is no specimen of the ist maxilla. Neither of the two cephalosomes is in the Museum's collection. The material is very faded and it is impossible to ascertain the detailed structure of some of the appendages. An attempt to remount some of the specimens failed because the mountant would not dissolve. The only observable difference between the present 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 wrongly described as the 4th leg (Scott, 1894 : PL 4, fig. 16). According to Scott the right ist antenna has 18 joints and is indistinctly geniculate and the left ist antenna has 23 joints. There are three ist antennae in the type collection ; one has 19 segments but 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 (different specimen) ; 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 segment left leg; u, 5th leg, last endopodite segment left leg (different specimen). Bar scale o-i mm unless indicated. NEW AND RARE CALANOID COPEPODS 341 better with the present male's antennae, which have 20 segments on each side, than with Scott's description. The type material of the 2nd maxilla is in very poor condition and it is impossible to see any structure except the general presence of two large brush-like setae. Despite the possible difference in the 3rd leg I believe that the present males are conspecific with Scott's Amallophora dubia. Sars (1907) erected a genus, Heteramalla, for a female which he believed was conspecific with Scott's male. In 1925, however, he expressed doubts as to the correctness of his assumption, and in 1965 Vervoort also mentioned the possibility that Sars' females and Scott's male were not the same species. Further male specimens have been recorded by Wolfenden (1911) and by Rose (1929) but they were not described. Bradford (1973) transferred Heteramalla from the Phaennidae to the Scolecithridae. The present males are clearly scolecithricids according to her redefinition of the family. I have examined female specimens of H. dubia Sars, 1907 taken at 1020- 800 m depth in a position i8°N 25°W. These specimens agree with Sars' (1924, 1925) description except that they have no 5th pair of legs (thereby conforming with Grice & Hulsemann's (1965) specimen), and there is no seta on the 2nd inner lobe of the ist maxilla. The endopod of the 3rd leg has two segments only. These females also agree with Bradford's definition of the Scolecithricidae. I believe, however, that Sars was mistaken in identifying his female with Scott's male. Apart from the normal sexual differences they differ in the following respects : (a) the rostrum 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 number of setae in the female are i, o and 3, (c) the ist leg of $ H. dubia has a 2-segmented exopod with no external spine on the ist exopodite segment ; the endopod has no lobe and has a group of large spines ; in the male the ist leg has three exopodite segments, all with outer spines, and the endopod has a lobe with a comb of small spines 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. dubia has a 2-segmented endopod and has no spines on the ist basipodite segment ; the 3rd leg of the male has three endopodite segments and two groups of spines on the ist basipodite. I agree with Bradford (1973) that the genus Heteramalla Sars, 1907 belongs to the Scolecithricidae. It is at present a monospecific genus known only from the female (see p. 342). Scott's male is here considered to belong to the new genus Scopalatum described on p. 335. The most striking difference between S. dubia and the females of Scopalatum is in the development of the sensory setae of the 2nd maxilla. The females have one brush-like seta much enlarged but the male has two. Enlargement of one or two of these setae is, however, common to the males of many species and genera of the Scolecithricidae (see, for example, Amallothrix falcifer, p. 325, and Scolecithricella dentata (Rose, 1942 : 55). I believe that the increased size of one seta in the present males is a sexual character. Apart from the normal sexual differences in the 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. gibbera that these two species may be conspecific. They differ in the length of the outer edge spines of the exopodite of the ist leg, and the males do not have the curious dorsal swelling of the female. In addition to these differences 5. farrani has groups of spines on the ist basipodite of the 4th leg. 5. smithae and S. vorax are much smaller species. Heteramalla sarsi sp. nov. Heteramalla dubia ; Sars, 1907 : 17 (part) ; A. Scott, 1909 : 86-87 (Part)> 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 a new species which I have named H. sarsi. The validity of this species is considered earlier together with the discussion of Scopalatum dubia (T. Scott). Family METRIDIIDAE Metridia alata sp. nov. MATERIAL EXAMINED : 6 females and 6 males taken in 6 RMT i hauls made at 1220-700 m depth in a position i8°N 25°W, between 12 and 18 November 1969. The holotype (female) and allotype (male) were caught on 18 November 1969 at i22O-iooom. Deposited at the British Museum (Natural History), holotype reg. no. 1974 : 630, paratypes reg. no. 1974 : 631-641. DESCRIPTION. Female (Fig. 22) : Body length 8-64-10-24 mm with a mean of 9-09 mm (5 specimens). The holotype measures 9-12 mm. The cephalothorax is 1-2 times as long as the 3-segmented abdomen. It is very robust and in lateral view the anterior part of the head protrudes sharply from the hind part. The right lateral margin of the head is produced into a very conspicuous wing ; this wing protrudes both laterally and ventrally and is present on the right side only. The head and ist thoracic segment are separate ; thoracic segments 4 and 5 are fused. The rostrum has two strong spikes pointing ventrally. The hind edge of the cephalothorax is smooth and rather abruptly rounded. The genital segment is swollen both laterally and 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 than wide. 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 has three hooks and the 2nd, 4th and 5th segments each have a single hook. The remaining mouthparts and legs are as shown. The endopodite of the mandibular palp 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, basal segments ; 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 legs is 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 terminates with three plumose setae. Male (Fig. 23a-b) : Body length 8-00-8-64 mm wrth a mean of 8-51 mm (6 specimens) . The allotype measures 8-64 mm. The appearance of the cephalothorax is as in the female ; the head has a conspicuous wing on the right side only. The abdomen has five segments. The left ist antenna is geniculate ; both ist antennae are slightly longer than the body. The remaining mouthparts and swimming legs are as in the female. The 5th pair of legs is asymmetrical : on the left side, the basal segment 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 2nd bearing a long curved process and the 5th ending in three small spines. REMARKS. The appendages of the present species are very similar to those of Metridia princeps Giesbrecht, 1892, but it can be distinguished easily from this and from all other species in the genus by the peculiar wing-like development of the right side of the head. Family CENTROPAGIIDAE Centropages caribbeanensis Park, 1970 MATERIAL EXAMINED : 43 females and 21 males from 6 RMT i hauls made at 194-0 m depth in a position i8°N 25°W, between 15 and 18 November 1969. 5 males 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 of 1-95 mm (15 specimens). The general appearance of the cephalothorax is as for the female ; it is 2-2 times as long as the 5-segmented abdomen. The furcal rami are about 3 times longer than wide. The ist antenna exceeds the body length by three or four segments. The right ist antenna is geniculate and has 22 segments, segments 17, 18 and the proximal part of 19 have a row of fine teeth, the hinged joint is between segments 18 and 19. The remaining mouthparts and swimming legs are as in the female. The 5th pair of legs is asymmetrical. On the right side the terminal exopodite segment forms a large chela ; on the left side the last exopodite segment has two large spine-like processes. REMARKS. The present males were found in the same hauls as females which agree in every respect with Park's (1970) description. The male is similar to that of C. elegans Giesbrecht, 1895 described by Park (1968). It can be distinguished from this latter species by the 5th pair of legs. In C. caribbeanensis the spines on the terminal segment of the left exopodite are longer than in C. elegans and are not pitted 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 second record of this species and the first from the eastern north Atlantic. NEW AND RARE CALANOID COPEPODS 345 FIG. 23. a-b, Metridia alata <$. a, body, dorsal ; b, 5th leg. c-e, Centropages carib- beanensis 3- c, body, dorsal ; d, last thoracic segment and abdomen, lateral ; e, 5th legs, f, Heterorhabdus vipera <$ 5th legs, g, Heterorhabdus tenuis <$ 5th legs, h-k, Haloptilus angusticeps <$. h, body, dorsal ; i, forehead and rostrum, lateral ; j, last thoracic segment and abdomen, lateral ; k, 5th legs. Bar scale o-i mm unless indicated. 346 H. S. J. ROE Family HETERORHABDIDAE Heterorhabdus vipera (Giesbrecht, 1892) MATERIAL EXAMINED : 14 females and 4 males from 12 Nii3H hauls fished at 590-90 m depth off Fuerteventura (28°N i4°W), between n and 26 November 1965. 22 females and 3 males from 12 RMT i hauls made at 1250-110 m depth in a position i8°N 25°W between 12 and 17 November 1969. REMARKS. The males of H. vipera and H. tenuis Tanaka, 1964 have been confused. Park (1970) described the male of H. tenuis as H. vipera (Roe, 1972^), and since the present material includes both species it seems advisable to illustrate the differences in their fifth pair of legs. H. vipera (Fig. 23f) has a strongly asymmetric fifth pair of legs. The endopodites consist of three segments on both sides and have been omitted for clarity. The 3rd exopodite segment of the right leg is abruptly terminated and about as long as the combined length of the two previous segments. The present specimens differ slightly from that of Giesbrecht (1892) by having a spine on the 2nd basipodite segment of both legs, and by having an outer edge spine on the 2nd exopodite of the right leg. The similarity is such that I have no doubt that they are conspecific. Heterorhabdus tenuis Tanaka, 1964 Heterorhabdus vipera ; Park, 1970 : 523-525, figs 250-252 (male only). MATERIAL EXAMINED : 7 males from 5 Nii3H hauls made at 660-460 m depth off Fuerteventura (28°N 14° W), between u and 26 November 1965. 14 males from 6 RMT i hauls made at 600-210 m depth in a position i8°N 25°W, between 12 and 16 November 1969. REMARKS. The present males agree with the descriptions of Tanaka (1964) and Park (1970 - as H. vipera <$}. 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 3rd exopodite 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 recently been reviewed by Matthews (1972). Because of the high degree of intraspecific variation within this genus he used a system of numerical taxonomy, based upon 35 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 in this genus but retained a normal diagnostic key for their identification. Several species of Euaugaptilus are described here, and all of them show marked intraspecific variations. This variability emphasizes the difficulties in recognizing or 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 discovery and description of more specimens is needed before firm conclusions can be reached regarding their validity. Haloptilus angusticeps Sars, 1907 MATERIAL EXAMINED : 2 males and 7 females from 5 Nii3H hauls made at 625-0 m depth off Fuerteventura (28°N I4°W), between 17 and 26 November 1965. 12 females from 5 RMT I hauls made at 700-305 m depth in a position i8°N 25°W, 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-segmented abdomen. The head and ist thoracic segment are separate ; thoracic segments 4 and 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 antennae reach to the end of the furcal rami ; the left ist antenna is geniculate and has 20 segments, 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 which is larger on the left, and the 2nd basipodite has a patch of hairs. Both endopods have three segments and are omitted here for clarity. The 2nd and 3rd exopodite segments are almost fused and only a faint line of separation remains. On the left side the 3rd exopodite segment has three small spines (the outer one is missing on the illustrated specimen) ; on the right the 3rd exopodite has one long curved spine and two of medium length. REMARKS. Except for sexual differences these males agree so closely with the female 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 at 4°°~55 m depth in a position i8°N 25°W, between 12 and 16 November 1969. 5 males 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 of i -87 mm (6 specimens). The shape of the cephalothorax is similar to that of the female. The head and ist thoracic segment are fused, as are thoracic segments 4 and 5. In dorsal view the forehead is shaped like a rounded triangle, and in some specimens it is more rounded than shown here. The rostral filaments are fairly long and have swollen bases. The hind margin of the cephalothorax is smoothly rounded. The abdomen has five segments, the furcal rami are about twice as long as wide. The ist antennae are longer than the body. The left ist antenna is geniculate and has 21 segments ; the proximal 12-13 segments have long sensory setae, segments 16, 17 and 18 have an undulating lamella, and segment 18 contains H. S. J. ROE FIG. 24. Haloptilus paralongicirrus same specimen j. * . , . ' no. 17 J 10125 I7 (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 be distinguished, and in one specimen there is some variation between the left and right side. 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 depth off Fuerteventura (28°N I4°W), between 24 and 28 November 1965. 3 females and 2 males from 3 RMT i hauls made at 785-610 m depth in a position i8°N 25°W, between 13 and 14 November 1969. 5 females and the males are deposited at the British Museum (Natural History), reg. no. 1974 : 664-670. DESCRIPTION. Female (Fig. 27a-g) : Body length 3-27-5-76 mm with a mean of 5-11 mm (6 specimens). The cephalothorax is long and narrow and just over 3-5 times as long as the 3-segmented abdomen. The head and ist thoracic segment are 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 subsequent segments. The ist antenna has 25 segments and exceeds the body length by 4-5 segments. Except for the ist maxilla the remaining mouthparts are as figured by 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. Both Sars' and Tanaka & Omori's specimens had a single endopodite segment with five setae. 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. Bar scale 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 a single segment with six and seven setae respectively. The endopodite of the ist maxilla of the smallest female (3-27 mm, Fig. 27f) agrees with Sars' account but the 2nd and 3rd endites have two and one setae respectively - the converse of the arrangement in both previous accounts and in the other specimens here. The swimming legs are as described by Sars. Male (Fig. 27!!, i) : Body length 5-12 mm (both specimens). The cephalothorax is 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 segments and exceeds the body length by five segments. The remaining mouthparts are identical 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 slightly asymmetrical. The external exopodite spines on the left leg are missing in the illustrated specimen but present in the other. The 2nd basipodite segment of both legs has a small patch of spines. The endopodites are 3-segmented on both sides and are omitted here for clarity. REMARKS. The female of E. maxillaris has only been described and figured by Sars (1924, 1925) and by Tanaka & Omori (1974). The male has hitherto been unknown. The present female specimens agree with these descriptions and I have no doubt that they are conspecific. The variation in structure of the ist maxilla accords with the intraspecific variations noted in Euaugaptilm by Matthews (1972) and by Tanaka & Omori (1974). Similar differences are also present in several of the species described here. Euaugaptilus ? longicirrhus (Sars, 1905) MATERIAL EXAMINED : i female taken in an NiiaH fished at 800-680 m depth off Fuerteventura (28°N I4°W) on 26 November 1965. 3 females and I male from 2 RMT i hauls made at 500-410 m depth in a position i8°N 25°W, between 22 and 24 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 mean of 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 the mouthparts 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 the exopod having four and the endopod two setae ; the blade has two more teeth than either Sars' or Sewell's specimens. The ist maxilla is reduced and agrees with Sewell's description. The 2nd maxilla has 3 (2 long and i spike), i, 2, 3, 2 and 3 setae on lobes 1-6 respectively, whereas Sars' specimen had — , i, 2, 3, 3, 3 setae and 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 in shape to the female but has slightly more conspicuous shoulders. It is 3-7 times as long as the 5-segmented abdomen. Thoracic segments 4 and 5 are fused and the hind 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 21 segments. The right ist antenna is broken. The mouthparts are as shown and are identical 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, ist leg. d-h, Euaugaptilus longicirrhus J99» 2OO> 248-250, figs 3h, 26. MATERIAL EXAMINED : 2 females in 2 Nii3H hauls made at 960-750 m depth off Fuerteventura (28°N I4°W), between 25 and 28 November 1965. 5 females taken in 5 RMT I hauls made at 900-410 m depth in a position i8°N 25°W, between 12 and 16 November 1969. 6 females are deposited at the British Museum (Natural History), reg. no. 1974 : 676-681. DESCRIPTION. Female (Fig. 2911-0) : Body length 3-50-3-80 mm with a mean of 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 is 2-6 times as long as the 3-segmented abdomen. The head and ist thoracic segment are separate ; thoracic segments 4 and 5 are fused. The forehead is rounded and the 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. The genital 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 has 25 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 antenna and ist maxilla, and in the number of mandibular teeth. The 2nd antenna was examined in three specimens, two of which had nine large setae on the endopodite and the 3rd (Fig. 29]) had eight. The ist outer lobe of the ist maxilla has six large and one small setae in three specimens, six large in one, five ? large in one, and five large and one small in another (Fig. 29m). The mandibular teeth are in groups of 2, 2 and 3 in four specimens (Fig. 29!), but in two of these four examples the second tooth 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 maxilla and maxilliped have well-developed 'buttons'. The legs agree with previous descriptions. 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 this station 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 in 1920 and more fully in 1924 and 1925. It was recorded by Vives (1970, 1972) and a further description has recently been given by Tanaka & Omori (1974). Sars listed his specimens as coming from Trincesse Alice' stations 1781, 2738 and 3021. I have borrowed a female specimen of E, propinquus from the Oceanographic Museum, Monaco, which was caught at station 1768 in 1904. This specimen is not mentioned by 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. propinquus and 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 , 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, is issued in Jive series corresponding to the Scientific Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 28, No. 8, of the Zoology series. The abbreviated titles of periodicals cited follow those of 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 OF THE 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 . Page 375 375 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 recorded from Britain and neighbouring seas are revised. Three species of Lafoeidae and nine species of Haleciidae are accepted. INTRODUCTION IDENTIFICATION guides covering the Hydroida faunas of the majority of countries bordering 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 of Hincks (1868) and Allman (1871) . Perhaps owing to the excellence of its illustrations Hincks' somewhat outdated work still often serves as an identification guide to the British thecate hydroids. The large amount of European work produced during the 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 recorded from 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 than hitherto, and some features previously regarded as good specific characters are shown 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 unclear affinities. It has variously been included in the families Campanulariidae (by Allman, 1888) and Lafoeidae (by Ralph, 1958 ; Vervoort, 1968, 1972), and removed to a separate family, the Hebellidae (by Nutting, in Fraser, 1912 ; Nutting, 1927 ; Fraser, 1937, 1944, 1946). The Hebellidae was established to accommodate genera with separate gonothecae, a hydrothecal diaphragm and a conical hypostome to the hydranth (said to be spherical in Campanulariidae). The two originally included genera, Hebella and Scandia Fraser (1912 : 371), were separated as having respec- tively free medusae and fixed sporosacs. It seems logical at present to retain the family Hebellidae to accommodate these two genera, which are implicitly excluded from the Lafoeidae in Naumov's diagnosis also. The British records of Hebella are discussed below under Lafoea dumosa. None appears valid. The faunal area outlined in Fig. i corresponds approximately with the continental shelf. Unavoidably it is arbitrary, some species being limital and none restricted to it. It was found that to include the whole of the English Channel, southern North Sea and all Danish waters would add only a few species to the list of all thecate hydroids known from Britain, whereas to include the coast of Norway would add sub-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, so that the whole of the English Channel is included ; while to the west, north and north-east the 183 m (600 ft) depth contour provides the arbitrary limit. Taking the thecate hydroid fauna as a whole there appears to be only a few species confined to the 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. The numbers reflect the approximate dates, in reverse, on which the specimens were registered and not the dates of collection or deposition in the Museum. Specimens loaned 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, even rimmed, without operculum, with or without diaphragm, pedicellate or sessile, adnate in some species ; hydranth with conical hypostome ; gonothecae aggregated as scapus or coppinia ; sexual generation a fixed sporosac. TYPE GENUS. Lafoea Lamouroux, 1821. REMARKS. Naumov (1960, 1969) has proposed two sub-families, diagnosed as follows : 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, 1960 With the characters discussed immediately above. Genus FILELLUM Hincks, 1868 Campanularia : Hassall, 1848 : 2223. Capsularia : Gray, 1848 : 151 (part) ; Naumov, 1960 : 280 ; Naumov, 1969 : 303 ; [non Capsularia 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 bearing hydrothecae without pedicels. Hydrothecae tubular, curved centrally, without operculum. 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. Capsularia was referred to Coryne Gaertner, in Pallas, 1774, by Lamouroux, Saint- Vincent and Deslongchamps (1824 : 220), and this synonymy was discussed and accepted by Bedot (1901 : 437). Gray (1848 : 61) also included Capsularia in the synonymy of Coryne, but elsewhere in the same work (pp. 85-88, 151) used the name for species now included in other genera and also (p. 151) for the generic name of the species here called Filellum serpens (Hassall, 1848). Vervoort (1972 : 50) showed the name Reticularia Thomson, 1853, to be preoccupied by 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 recorded from 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 Hardanger Fjord, W Norway (Broch, 1918) and from N Biscay at 186 m depth (Billard, 1923). A statement that the 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 herbarium specimen 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 spirit and 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 on Abietinaria 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. Off Millport, 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 Abietinaria abietina, microslide, coll. E. T. Browne, 1959.9.17.119. False Bay, Republic of South 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 with hydrothecae at irregular intervals. Hydrothecae tubular, wider than stolons, bent upwards at 60-90° approximately in centre ; rim even, sometimes flared, without operculum ; 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 HYDROTHECA Diameter at rim Length of free part Length of attached part HYDRORHIZA Diameter of stolon COPPINIA (young) Overall length Overall width * Dried material. E SCOTLAND (holotype of R. immersa Thomson) 90-140 60* W SCOTLAND (1967.11.10.5) 100-175 200-900 200-300 100 IRELAND (I959.9.I7-II9) H5-I35 80-170 220-300 100 NORWAY (1962.11.7.10) c. 1500 c. 1000 VARIATIONS. Hydrothecae in this species are longer in some specimens than in others, long hydrothecae usually being those with one or more renovations. The hydrothecal 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 the hydrotheca which is adnate varies between £ and f of the total length. Hydrothecae may be either narrower (Naumov, 1960, 1969) or wider (Vervoort, 1972) basally than distally, or of uniform diameter. Hydranth colour has been recorded as greenish (Hincks, 1868) and lemon yellow (Hamond, 1957). Colonies growing in a limited space - as on another hydroid colony - have the hydrothecae closer spaced than 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 recorded intertidally. Recorded growing most frequently on sertularian hydroids, especially Abietinaria dbietina, but also Hydrallmania falcata (Linnaeus, 1758) and Sertularia cupressina Linnaeus, 1758. Also occurs on non-living substrates such as bivalve shells. Hincks' (1868) statement that the species is parasitic almost certainly referred to the growth of the species over other hydroids, and was probably not intended to imply an association of living tissues. REMARKS. Many authors have considered the two nominal species F Helium serpens (Hassall, 1848) and Reticularia immersa Thomson, 1853, to be conspecific and 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 abietina is here identified as Filellum serpens. The specimen was a lafoeid growing over a colony of a sertularian hydroid, A bietinaria abietina. Billard stated that it resembled a colony of F. serpens except that the hydrothecal aperture was not flared (elargie), and hence concluded that the specimen was a stoloniferous growth of Grammaria abietina. Such colony habit has not otherwise been recorded from G. abietina, and it is now known (present paper) that hydrothecae of F. serpens are not invariably flared. 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 abietina Sars, 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 genus by Hincks (1868). The type species of Salacia is 5. tetracythara Lamouroux, i8i62 (by monotypy) . The type material was examined by Deslongchamps (in Lamouroux, Saint- Vincent & Deslongchamps, 1824) who criticized Lamouroux' description as inaccurate and commented that the vase-shaped gonotheca of the specimen, shown clearly in the original illustration, suggested its affinities were with the genus Sertularia Linnaeus, 1758. Later Bale (1884) examined the same specimen and referred it to Thuiaria Fleming, 1828. Bale also provided a new trivial name, calling the species Thuiaria fenestrata. Like Deslongchamps, Bale considered Lamouroux' description to be inaccurate and, somewhat illogically, regarded Salacia as 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 characters of 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 the synonymy of Salacia, his usage being followed by Rees & Thursfield (1965). Un- fortunately the type material of Salacia tetracythara was destroyed along with the bulk 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 a senior synonym of Thuiaria. Hincks (1868) thus seems to have been unjustified in 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 appears to 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 two genera 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 of other 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. V2 (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, 62°oi' N, 5°i9' W, 210 m, 1869, two colonies in spirit, coll. H.M.S. 'Porcupine', via A. M. Norman coll., 1912.12.21.303. Trondheim Fjord, Norway, 1893, fragments of hydrocauli 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 in glass tube, Northumberland coast, coll. J. Alder, 1857.8.3.52 [syntype of Grammaria ramosa 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, 6i°35' N, o°47' 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 Zoological Museum, 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, 5i°35' S, 65°3g' 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). Royal Sound, Kerguelen Island, 49°28' S, 70°i3' 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). Off Marion Island, S Indian Ocean, 46°43' S, 38°04' 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 (svntypes of Grammaria insignis Allman, 1888). DESCRIPTION. Colony erect, main stem and branches polysiphonic, branching irregular or imperfectly pinnate, branches tapering near the base. Hydrothecae long, tubular, variably outward-curving ; rims even, circular, often renovated, frequently slightly flared ; no apparent junction with pedicel ; arranged in 4-8 longitudinal rows with hydrothecae of adjacent rows alternate ; if 4 rows, hydro- thecae often in decussate pairs. Hydranth yellow (Hincks, 1868), extensible roughly one hydrotheca-length beyond aperture ; widest distally, hypostome conical, 18-20 tentacles held alternately elevated and depressed. Gonothecae borne in hermaphrodite coppiniae 'arranged on branches in small muffs. Surface of 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 240-280 280-360 280-490 COPPINIA 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 4 rows the hydrothecae are in opposite or almost opposite pairs. The hydrothecal rim is flared in some specimens and straight in others, this character having ap- parently no systematic value. The degree of outward curving of the hydrotheca varies, so that the plane of the hydrothecal aperture is parallel with the axis of the hydrocaulus in some specimens, while in specimens with shorter hydrothecae the plane of the aperture makes an angle of about 45° with the axis, and this character may 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 seems to be no valid record from western coasts south of the Shetlands. Southerly records include Northumberland (Hincks, 1868 ; present material), Skagerrak (Kramp, 1935) and Oslo Fjord (Christiansen, 1972). A record from the north coast of 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 lacks evidence. The weight of published opinion is that the species is absent from the English Channel and southern North Sea (Hincks, 1868 ; Hartlaub, 1895 ; Broch, 1927 ; Vervoort, 1946, 1949 ; Leloup, 1952 ; Hamond, 1957 ; Marine Biological Association, 1957 ; Teissier, 1965 ; Robins, 1969). HABITAT. Published records indicate that the normal depth-range of this species is 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 been recorded on both silty and rocky substrates (Naumov, 1960, 1969 ; Calder, 1970). REMARKS. There seems no doubt from the original description of Grammaria ramosa Alder, 1856, that it should be regarded as a junior synonym of the present species and examination of the type series confirms that they are conspecific. Vervoort (1972) has expressed doubt as to the distinctiveness of the three species erected by Airman (1888), Grammaria insignis, G. magellanica and G. stentor, and following 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 branching of the colony and of the number of rows of hydrothecae, and whethei or not the hydrothecal 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 here denned, and may represent a distinct species. If so, a new name will have to be provided. G. stentor sensu Hartlaub, 1905, and Jaderholm, 1905, fall within the proposed 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 account suggests 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 species and 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 de Lafoye, 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 colony 100 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 erect polysiphonic hydrocauli with monosiphonic terminal branches. Hydrothecae lacking both operculum and diaphragm ; tubular or bell-shaped, sometimes bent and thus bilaterally symmetrical ; rim even, circular, sometimes flared ; renovations frequent ; pedicel twisted if present ; gonothecae borne in hermaphrodite coppiniae. REMARKS. Billard (1909 : 311) examined the genotype material and referred the type species L. cornuta Lamouroux, 1821, to L. dumosa (Fleming, 1820), supporting the 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 J9-21 (syn- 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 specimens unknown. 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, 58°5o' N, n°oo' E, 80-100 m, 23 Sep. 1964, hydrocaulus on microslide, coll. W. J. Rees, 1965.1.14.73. Vadero Islands, Sweden, 58°05' N, n°04' E, 80 m, i Oct. 1964, several colonies in spirit, coll. W. J. Rees, 1965.1.14.136 and 1966.1.4.65. Loken, Gaso Ranna, Gullmarfjord, Sweden, 25-30 m, 14 May 1959, 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, colony in 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 in spirit, 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, and Northumberland, 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: from refuse 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 Campanularia gracillima Alder, 1856 (Totton, 1930)]. Scarborough, Yorkshire, England, colonies on 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, 6o°O2' N, 3°I3'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, 59°2i' N, 5°oo' 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, seven hydrocauli 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, 66°59' N, 55°27' W, 120 m, fragments of hydrocaulus on microslide, coll. H.M.S. 'Valorous', 1878.3.26.3. E of Str. of Magellan, Argentina, 52°2o' S, 67^9' W, 100 m, 20 Jan. 1876 ('Challenger' sta. 313), part of colony on microslide, coll. H.M.S. 'Challenger', 1890.4. n.2d (Fig. 4). Off Gates Land, Antarctic mainland, 69°43' S, i63°24' 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 comprise irregularly-branched monosiphonic hydrorhizae with hydrothecae borne at irregular intervals. Erect colonies consist of polysiphonic hydrocauli, loosely and irregularly branched, arising from a straggling stolon network. Component perisarc tubes of erect colonies parallel, each bearing hydrothecae at irregular intervals giving bristly appearance. Hydrothecae long, tubular, even-rimmed, variably tapered below, sometimes asymmetrically ; pedicel more or less well defined, usually kinked, but sometimes lacking so that hydrotheca is sessile. Form of hydrothecal pedicel usually constant within a colony. Hydranth long, retractile, c. 20 tentacles held alternately elevated and depressed (Kramp, 1935), hypostome conical. Hydranth and 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) HYDROTHECA Base to rim (unrenovated) 500-650 Diameter at rim 150-220 Pedicel length o W SWEDEN (1965.1.14.73) 485-510 100 100-150 N SCOTLAND (1964.8.7.50) 770-830 150-210 50 W SCOTLAND (1899.5.1.160) 380-550 120-140 140-170 W SCOTLAND (1899.5.1.162) (holotype of Lafoea pocillum) 120-350 90-190 80-170 VARIATION. The hydrothecal pedicel varies in length between colonies (see Remarks) and the degree of bilateral symmetry of the hydrothecae varies also. Neither character appears to be of systematic importance. Specimens from deep water have fewer hydrothecae per unit length of stolon, but within continental shelf depths this character seems constant. The so-called pedicel of the present species 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 specimens in a series in which the pedicel becomes progressively less distinct and is finally absent (Fig. 4). In each colony the range of pedicel length is small, but pedicellate hydrothecae can be found in colonies largely lacking pedicels, and sessile hydrothecae occur 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 records suggests that in this species reproduction is usually vegetative. DISTRIBUTION. Found throughout the area but apparently not common in the southern North Sea and eastern English Channel (Broch, 1918 ; Vervoort, 1946 ; Marine Biological Association, 1957 ; Teissier, 1965 ; Naumov, 1969), not having been found in a faunal survey oft the coasts of Belgium (Leloup, 1952). HABITAT. Occurs at all continental shelf depths, but not intertidally. On a wide 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 as valid species by others (e.g. Broch, 1918 ; Kramp, 1935 ; Fraser, 1944 ; Rees & Thursfield, 1965 ; Calder, 1970). Vervoort's detailed appraisal is particularly convincing and his view that the two are conspecific is accepted here. In contrast, it appears that the distinctness of L. fruticosa from L. dumosa has been questioned only by Millard (1964), who described intermediate material. The two species have been distinguished by Hincks (1868) and apparently all subsequent authors (except Rufford, 1902, and Ralph, I9586) by the presence of a hydrothecal pedicel in L. fruticosa and its absence in L. dumosa which has been held to have sessile hydrothecae. However, the original description of L. dumosa (Fleming, 1820, repeated in Fleming, 1828) referred to 'nearly sessile' hydrothecae, implying that short 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-examined during the present work all have pedicellate hydrothecae. The material was from Johnston's collection, some being labelled Berwick Bay and probably, therefore, the Berwick Bay material described by Johnston (1832, 1838). Whether or not this is so it seems probable that Johnston, like Fleming, wished to imply that the pedicels were short, and not that they were absent. Further, in describing L. gracillima, Alder (1856) stated that the pedicels were 'longer' than in L. dumosa, indicating that he too considered L. dumosa to be pedicellate. Thus, when Hincks (1868) later gave specific status to specimens without pedicels and distinguished L. fruticosa as having them he was wrong to refer non-pedicellate material to L. dumosa and should have provided a new name. This seems never to have been done. It appears probable, however, that specimens lacking hydrothecal pedicels do not constitute a valid species and should nevertheless be referred to L. dumosa sensu Fleming, 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 and long pedicels respectively, but she gave a description of L. gracillima only. Rufford ascribed a short pedicel 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 type material (Fig. 4). The material referred to Hebella pocillum (Hincks, 1868) by Fraser (1937, 1944) and Yamada (1955) is at present indeterminate ; it may require a new specific name. The type material of L. parvula could not be located, but specimens 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 plausible that 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 a later 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 as identical with L. gracillima (= L. dumosa), while Naumov (1960, 1969) referred it to L. 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 a well-demarcated pedicel with transverse annulations, are clearly of Calicella and it seems 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 usually wider than deep ; hydrothecal rims even, often flared, renovation common, desmo- cytes large, often birefringent. Hydrothecal pedicel often lacking. Hydranths larger than hydrothecae. Gonothecae usually sexually dimorphic. Sexual genera- tion said to be a medusa in Campalecium Torrey, 1902 (Kossowska, 1911), eumedusoid in Hydranthea Hincks, 1868, and a sessile sporosac in Halecium Oken, 1815, and Ophiodissa Stechow, 1919. TYPE GENUS. Halecium Oken, 1815. 7 REMARKS. The nature of the sexual generation was shown by Rees (1957) to be a 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 is included here in the generic diagnoses. A similar correlation has been shown in certain genera of the Campanulariidae (Cornelius, 1975). 7 Oken's book was placed on the list of rejected works by the International Commission on Zoological Nomenclature (1956 : Opinion 417), but application is currently being made to the Commission for validation of the name Halecium (see Cornelius, 1976). BRITISH LAFOEIDAE AND HALECIIDAE 391 The small birefringent bodies found inside the hydrothecal wall throughout this family (Fig. n) and widely referred to as punctae (e.g. Vervoort, 1972) are probably desmocytes. They resemble those previously described in other classes of coelenter- ates, reviewed by Chapman (1969), but are much larger, being visible in some species at only 20 diameters' magnification. A preliminary account of their ultrastructure is to appear elsewhere. It is probable that the 'punctae' of the family Lafoeidae are also desmocytes. Present knowledge suggests that they have little systematic value at species level in either family, but comparative studies of their ultrastructure might prove differently. Genus HALECIUM Oken, i8i57 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 usually sexually dimorphic. REMARKS. Johnston (1847) drew attention to the seniority of Halecium to Thoa and 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, was placed 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 : 447—449, 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 of colony 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 colonies in 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 (mentioned Kramp, 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, Fisheries and 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 on Chaetopterus 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 stem polysiphonic. Side-branches polysiphonic basally, component coenosarcs gradually branching 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 slightly flared, 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, aperture terminal ; $ approximately bean-shaped, aperture tubular, in centre of concave side, 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 rim HYDROCAULUS DIAMETER LENGTH OF INTERNODES o* 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) 25-45 110-140 120-150 500-530 IOOO-I2OO 430-480 50-70 200-250 60 I2O-I8O 20-25 H5-I50 120-lSo — 250-750 250-700 Up to 2 2OO 62O-IOOO 250 700-850 I2O-I8O Up to 22OO II20-II25 310 0-3 m VARIATION. Hydrocauli of this species are variably flexuose ; extremes are shown in Fig. 5. REPRODUCTIVE SEASON. Fertile specimens recorded from S Devon, Norfolk and Roscoff, France, most months from January to October (Hamond, 1957 ; Marine Biological Association, 1957 ; Teissier, 1965). DISTRIBUTION. Common throughout Britain and nearby seas. Apparently absent from the Baltic and rare in the Kattegat, commoner in the Skagerrak and Oslo 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- 42. figs a-d ; Broch, 1918 : 36-38, fig. n ; Vervoort, 1946: 158-161, figs 63—64; 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 (I959-9-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 Ostrea edulis Linnaeus, 1758 (Ellis, 1755 : pi. 10, figs A, a, B) ; location of specimens unknown.8 MATERIAL EXAMINED. Oban, Argyll, 1877, three colonies in spirit, coll. A. M. Norman, 1912.12.21.198. Bridlington Bay, Yorkshire, 7 Nov. 1921, several colonies 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 of Man, 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 colony on 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 material himself since he did not give a locality. Although the collections of the Linnean Society of London include material labelled H. halecinum [catalogued 1298.7-8 by Savage (1945)], it was probably added to 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's name or locality. They cannot, therefore, be identified as type material. Some Ellis hydroid material, which might have included the originally illustrated specimens of H. halecinum, was until recently preserved in the Hunterian Museum of the Royal College of Surgeons of England. Unfortunately, the Ellis material was almost certainly destroyed by a bomb during the Second 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). Start Point, 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 third order branching. Hydrocaulus and main branches polysiphonic basally, component perisarc 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 some quaternary hydrothecae inside primary (Fig. 6) and longer than it. Hydranth larger 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 hydranths projecting 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 internodes cj GONOTHECA Length Breadth (max.) $ GONOTHECA Length Breadth (max.) MAXIMUM HEIGHT OF COLONY ? U.S.S.R. NETHERLANDS SE ENGLAND SW ENGLAND (Naumov, 1969) 90-150 170-210 500 0-25 m (Vervoort, 1946) 1 00-200 800-1000 c. 400 1000-1300 400-500 0-25 m (1964.8.7.273) (1889.7.27.7) 20-40 140-165 190-225 350-380 1250-1400 350-600 25-40 130-145 100-150 280-400 700-800 250-400 REPRODUCTIVE SEASON. Fertile specimens recorded December to July in English Channel 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 continental shelf areas. HABITAT. On stones, shells and other hard substrates, from sublittoral to edge of continental 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 original description to be conspecific with H. halecinum. The long, tubular hydrothecae described by Norman and regarded by him as diagnostic were probably just the renovated 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 herbarium sheet 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, 6o°2o' N, 5°i2' E, 15-25 m, 13 Apr. 1962, several colonies in spirit, three microslides of fragments (2 $, i sterile), coll. W. J. Rees, 1962.10.7.13 (Fig. 7), 35, 66. N Vadero Is., Skagerrak, 58°35' N, n°04' E, 80 m, i Oct. 1964, colony in spirit and one 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 colonies in spirit, coll. E. T. Browne, 1941.3.20.413. Eddystone ground, western English Channel, 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, holotype of 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 final branches polysiphonic ; main branches slightly flexuose, side-branches inserted alternately 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 prominent hydrophores, 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 early May in southern North Sea, late May in Orkneys and Faroes, late June in Iceland (Hamond, 1957) ; 17 August 1898 at Plymouth (present material) ; September at Roscoff (Teissier, 1965). DISTRIBUTION. A northern species known from localities throughout British coastal waters (Hincks, 1868 ; Hamond, 1957 ; Marine Biological Association, 1957). Although recorded from Roscoff, NW France (Teissier, 1965), the species appears absent from the Scilly Isles, Channel Islands, Belgium and Holland (Vervoort, 1946, 1949 ; Leloup, 1952 ; Robins, 1969) and the English Channel is probably near its southern 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 internodes o* GONOTHECA Length Breadth (max.) $ GONOTHECA Length Breadth (max.) P. F. S. CORNELIUS TABLE 6 Halecium labrosum. Measurements in (Jim W ENGLISH ARGYLL, CHANNEL PLYMOUTH SCOTLAND (1959.9.17.123) (I959.9.I7-9) (1888.1.24.19) 30-50 240-280 160-200 950-1175 30-45 150-165 90-110 800-1700 640-1100 470-530 40-50 220-255 130-150 550-700 1350-1580 950-1090 W NORWAY (1962.10.7.13) 30-45 (one 60) 140-155 (one 170) 90-110 600-820 700-1000 300-900 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 in this 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 a wrinkled perisarc and lacks the curvature of the bases of the hydrocladia typical of H. labrosum. Eudendrium pusillum Sars, 1857, type locality northern Sicily, remains problema- tical. It was recognized as Halecium by Kossowska (1911) who wrongly included H. lankesteri in its synonymy. Broch (1912) redescribed and figured part of the holotype 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 this and 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 stunted in comparison with colonies of H. labrosum. It is not, therefore, possible to identify it confidently with the present species, which in addition has not been recorded so far south. Possibly the material from NW France identified as H. pusillum by Teissier (1965) was in fact H. labrosum as defined here ; his is apparently the only suggestion that H. pusillum occurs in the present area. Halecium reflexum Stechow, 1919, appears from the original description to resemble H. labrosum in its manner of branching, recurved hydrothecal rim and wrinkled perisarc, and the two taxa appear conspecific. Halecium undulatum Billard, 1921, seems to have been based merely on small colonies of H. labrosum. The two taxa appear similar in all characters except colony size and their distributions coincide so that there seems no reason to keep them 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 described by 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. schneideri Bonnevie, 1898, is here referred to the present species. H. schneideri was a new name for H. nanum Alder, 1859, a species otherwise unrecorded from the present area. 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, J873 : 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 : 46 : Vervoort, 1959 : 221-224, figs 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, Plymouth Sound, 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 spirit material deposited in the Museum of the Marine Biological Association, Plymouth). DESCRIPTION. Colony erect, up to c. 80 mm, usually unbranched but sometimes imperfectly pinnate ; monosiphonic. Internodes often irregular in length, o-io (usually i) annulations between hydrothecae ; bases of hydrocladia often curved inwards sharply. Hydrotheca on prominent hydrophore at distal end of internode ; usually alternate ; slightly tapering basally, short ; rim not flared ; up to c. 5 hydrothecal renovations. Hydranth large, 10-20 tentacles. Gonotheca on short pedicel below a hydrotheca. $ cylindrical, aperture terminal ; $ kidney-shaped with tubular aperture in centre of concave side, 3-20 ova, 1-2 protruding polyps. Vervoort (1959) records sexes on separate colonies, $ gonothecae 'exclusively on renovated hydrophores' over entire colonies, $ on lower parts of colonies on secondary or 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 internodal annulations between hydrothecae, and successive internodes are not identical in shape or length. The degree of wrinkling of the perisarc is variable. Tendril development occurs (e.g. 1974.3.1.1) but is unusual. REPRODUCTIVE SEASON. June-July at Roscoff (Teissier, 1965) ; gonothecae of both sexes, Norfolk coast, 26 June 1952 (Hamond, 1957) ; $ gonothecae, Plymouth, Devon, 2 July 1973 (1974.3.1.1). Infertile material from Plymouth includes the type specimens collected May 1889 and May 1890, and other material collected 15 July 1898 (1959.9.17.1). DISTRIBUTION. There are published records from NW France, the Channel Islands, SW England, SW Wales and Norfolk (Bourne, 1890 ; Vervoort, 1949 ; BRITISH LAFOEIDAE AND HALECIIDAE TABLE 7 Halecium lankesteri. Measurements in [zm unless otherwise stated 401 HYDROTHECA Length (diaphragm to rim) Breadth at rim INTERNODES Diameter (min.) Length fig. 5 ; Vervoort, 1959 : 229-231, fig. 8 ; Naumov, 1960 : 454, fig. 344 (syn. H. geniculatum Nutting, 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 three microslides, 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 in spirit, coll. A. M. Norman, 1912.12.21.208. Oban, Argyll, Scotland, 1877, several colonies 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 colonies on sertularian hydroid, coll. Imperial College (University of London) Azores Expedi- tion, 1962.1.15.24. Porto Santo Island, Madeira, 120 m, several colonies on Eudendrium sp., in spirit and microslide, coll. R. Kirkpatrick, 1919.8.14.7. Simon's Bay, Simon's Town, Republic of South Africa, 'shallow water', colonies on fragments of 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 in spirit, 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 Antarctic Expedition, 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, irregularly branched. Hydrocaulus zig-zag in general appearance, internodes usually straight 13 This section includes all material of this species in the British Museum (Natural History) collection except the type series. BRITISH LAFOEIDAE AND HALECIIDAE 411 and narrow, angle between them about 140°. Usually 1-3 annular wrinkles at each 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, slightly larger, 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 rim HYDROCAULUS DIAMETER (min.) LENGTH OF INTERNODES $ GONOTHECA Length Maximum breadth 9 GONOTHECA Length Maximum breadth MAXIMUM HEIGHT OF COLONY (syntypes) 25-30 100-130 50 500-850 540-720 200 c. 5 mm ig66a) (Vervoort, 1966) 30-65 120-135 875-1300 (Naumov, 1969) 40-80 120-190 1070 (max.) 600 1070 (max.) 5io 4 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 (Natural History) collection comes from only two localities, Salcombe, Devon (syntypes), and Oban, 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 Bryozoa by 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 serve to distinguish infertile specimens from young specimens of H . labrosum. In addition the curving bases of the hydrocladia in H. labrosum contrast with the straighter bases in the present species. Published records should be treated with caution. A summary of records in which the species was confused with H. labrosum and its junior synonym H. undulatum Billard, 1921, was given by Hamond (1957). See also the Remarks section under H. labrosum (p. 398). 4i2 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 short to contain hydranths which are large ; gonophore eumedusoid ; large nematocysts present. 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 Flustra sp. (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, colony in spirit, coll. E. T. Browne, 1954.8.3.42. Plymouth, 23 Oct. 1913, fertile colony on Halecium 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, in spirit, coll. E. T. Browne, 1954.8.3.44. Roscoff, NW France, 12 Sep. 1956, colony on bivalve shell, in spirit, coll. W. J. Rees, 1956.10.24.1-2. DESCRIPTION. Colony reptant, stolon bearing single hydranths at irregular intervals. Hydranths yellowish-white (Hincks, 1868), on short unringed pedicels, narrow at base, widest just below hypostome, 20-30 tentacles held alternately elevated and depressed (Hincks, 1868 ; Browne, in White, 1956), with small basal web. Hydrotheca short, walls slightly divergent, rim even, diaphragm domed downwards ; c. 20 desmocytes ('punctae'). Large, conspicuous banana-shaped or straight nematocysts (microbasic eury teles, Huve, 1954), 30-40 (xm long, throughout tissues of hydranth and coenosarc, scattered or in clumps, particularly conspicuous on tentacular web. Gonophores on short pedicel tapering basally, attached to stolon ; gonotheca short (Fig. 13) or absent ; pedicel base said to be enclosed in small chitinous 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, W Scotland, 20-30 m (1955.11.15.11); b, developing gonophore, two-cell stage, SW England (1954.8.3.43) ; c, mature $ eumedusoid, E Mediterranean, lower shore (redrawn after Kossowska, 1911 : fig. 2, as H. billardii) ; d, undischarged nematocysts, SW England (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 contains c. 300 ova. (Description of gonophores based on Hincks, 1868, Kossowska, 1911, and Huve, 1954.) Developing eggs reported in hydrorhiza in a colony apparently lacking gonophores (Millard & Bouillon, 1973). MEASUREMENTS. See Table n. TABLE n Hydranthea margarica. Measurements in fim STOLON Diameter HYDRANTH Height (base to hypostome) Tentacle length HYDROTHECA Height (diaphragm to rim) Breadth at base Breadth at rim Hydrothecal pedicel NEMATOCYSTS Length * Contracted during fixation. S WALES (IQ73-3-3-I) 60 200-400* 100 100 100 W SCOTLAND (White, 1956 ; S FRANCE = 1955.11.15.11) (Huv6, 1954) 60-80 400-1000 100-170 80-90 80-90 40 90 700 180 35 80 IOO 150 30 REPRODUCTIVE SEASON. Fertile specimens recorded August (Roscoff, NW France, Teissier, 1965) and October (Plymouth, SW England, White, 1956). 24 4i4 P. F. S. CORNELIUS DISTRIBUTION. Recorded from Devon and Dorset, Liverpool Bay, Norfolk and Argyll (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 and bare 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 '• J86 (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 medusa generation ; large nematocysts present. REMARKS. This generic diagnosis agrees with the restriction of Vervoort (1959, 1972) in excluding species lacking a nematotheca. Following Vervoort, the one species concerned is retained in the genus Hydrodendron Hincks, i874a ; but as noted by Watson (1969) a review of Ophiodissa and related genera is needed and present 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, ng- 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 : Lower ledges of Capstone, Ilfracombe, Devon, England, lower shore, 1867, infertile colony on four fragments of holdfast of Laminaria sp., 1899.5.1.169. The neotype material is labelled 'Ophiodes mirabilis, Ilfracombe, 1867' in Hincks' hand, and is almost certainly 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, Cape Verde 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, colonies on 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 colony on 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, two fragments 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 polysiphonic basally, 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, hydrophore distal, 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 Aran in Donegal. 416 P. F. S. CORNELIUS o o •n co o ro I O 00 o o « « £ S5 W ai 3 9 en ^ - q o~^^ 75 £ ££ 0 £o § 000 0 £• 2 00° 00 0 ° NO O O «i C -i w t_ II ^^ "•^ *-* 1 1 1 -4-* - C 0 i •£ -£ ooo o S S S r(- in O^ O OOO H uS o o o o o o ro vO O^ ^O *O CO ^G^ &^£> 8 t**«» 0 OOO r^ o o o PO o MO CO M HI 00 o o o O ro O O IT) M IO M i (N | | 1 1 o O ,.• O OOO O o o o o o in m N ^ CO t^-CO PI O M MM PO Tt- oo N oo in o iaphragm istome) B d -^ -C 2 ffl i ll» £.- 43 6 Of a n - J2 S .5 iv^ > "o ^ o 'C 'S — - c x~ ?S M 'O ° ^-y-S2!:ij0O'Ti^i-tM0 ucg«--^:2^«j;g wgw ^jw ci<;c DMh^ frt^T *~"fi^ ^liefiiofila* | a £ g ^og i-aJ g^ 2 1 43 S g 32 •§ S 3 5 -i is s<5S B53 §s p>-M MI— 4P*-HM^iGil ^ w : i- j -i- i£ ;£ ffi BRITISH LAFOEIDAE AND HALECIIDAE 417 1957 ; Ralph, 1958). Hydranth large, tapering basally, 18-24 tentacles, held alternately raised and lowered. Nematothecae borne irregularly on internodes, hydrophores and stolon ; curved cone-shaped, narrowest basally, sometimes with constriction below rim, both varieties occurring on single colonies ; nematophore long, axial cells large, terminal knob with numerous nematocysts including large microbasic mastigophores. Gonothecae apparently recorded only twice. Described as 'ovate, ringed transversely with wide tubular aperture, subpedicellate, borne on stolon', 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 from Ilfracombe, Devon ; Swanage, Dorset (Hincks, 1868 ; Garstang, 1900) ; Pembroke- shire (Crothers, 1966 ; present material) and Aran Isles, Galway Bay (present material). A record from Aberdeen, Scotland (Forbes, 1872), was regarded as dubious by its author. Apparently the only other European record is from Roscoff, NW France (Teissier, 1965). Other records, as 0. caciniformis, include the Cape Verde 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 synonym of the present species. 0. caciniformis was proposed to include colonies larger than those previously referred to 0. mirabilis by Hincks (1868). 0. mirabilis has not previously been redescribed ; and the larger specimens described by various authors this 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 and internal 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 the characters of the older species, and the dimensions of the type specimens of the two taxa are similar. It seems that 0. mirabilis was founded on small colonies, and that larger colonies found later were wrongly given specific status. ACKNOWLEDGEMENTS I am grateful to Dr C. Edwards, Scottish Marine Biological Association, for general advice on hydroid systematics and to the following for kindly providing unpublished locality records and for loaning material : Dr M. E. Christiansen, University of Oslo Zoological Museum ; Dr W. D. Hartman , Yale Peabody Museum ; K. Hiscock, University College of North Wales, Bangor ; the late D. N. Huxtable, University College 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. Dr N. A. H. Millard, South African Museum (Natural History), kindly gave permission for one of her figures to be redrawn and published here. I am grateful also to J. M. Price, British Museum (Natural History), for identifying algae, and to Dr W. Vervoort of the Rijksmuseum van Natuurlijke Historic, Leiden, for commenting on this paper in the proof stage. Lastly I am much indebted to D. J. McGrail for redrawing the figures. REFERENCES ALDER, J. 1856. A notice of some new genera and species of British hydroid zoophytes. Ann. Mag. nat. Hist. (2) 18 : 353-362. 1859. Descriptions of three new species of sertularian zoophytes. Ann. Mag. nat. Hist. (3) 3 : 353-356. i862a. Descriptions of some new and rare zoophytes found on the coast of Northumber- land. Ann. Mag. nat. Hist. (3) 9 : 311-317. 18620. Supplement to a catalogue of the zoophytes of Northumberland and Durham. Trans. Tyneside Nat. Fid Cl. 5 : 225-247. ALLMAN, G. J. 1871. A monograph of the gymnoblastic or tubularian hydroids. London. 1888. Report on the Hydroida dredged by H.M.S. 'Challenger' during the years 1873-76. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and Thalamo- phora. Rept scient. Results Voy. Challenger, 23 : i-go + i-lxix. BABIC, K. 1913. Uber einige Haleciiden. Zool. Anz. 41 : 468-474. BALE, W. M. 1884. Catalogue of the Australian hydroid zoophytes. Sydney. BEDOT, M. 1901. Materiaux pour servir a 1'histoire des hydroi'des. zer periode. Revue suisse Zool. 9 : 379-515. - 1905. Materiaux pour servir a 1'histoire des hydro'ides. 2me periode (1821 a 1850). Revue suisse Zool. 13 : 1-183. — 1910. Materiaux pour servir a 1'histoire des hydro'ides. 3me periode (1851 a 1871). Revue suisse Zool. 18 : 189-490. - 1911. Notes sur les hydro'ides de Roscoff. Archs Zool. exp. gen. (5) 6 : 201-228. 1912. Materiaux pour servir a 1'histoire des hydroides. 4me periode (1872 a 1880). Revue suisse Zool. 20 : 213-469. — 1914. Nouvelles notes sur les hydroides de Roscoff. Archs Zool. exp. gen. 54 : 79-98. — 1916. Materiaux pour servir a 1'histoire des hydroides. 56 periode (1881 a 1890). Revue suisse Zool. 24 : 1-349. — 1918. Materiaux pour servir a 1'histoire des hydro'ides. 6e periode (1891 a 1900). Revue suisse Zool. 26 (suppl.) : 1-376. — 1925. Materiaux pour servir a 1'histoire des hydroides. 76 periode (1901 a 1910). Revue suisse Zool. 32 (suppl.) : 1-657. BILLARD, A. 1904. Contribution a 1'etude des hydroides. (Multiplication, regeneration, greffes, variations.) Annls Sci. nat., Zool. (8) 20 : 1-251. — 1909. Revision des especes types d'hydroides de la collection Lamouroux conservee a 1'Institut Botanique de Caen. Annls Sci. nat., Zool. (9) 9 : 307-336. 1921. Note sur deux especes d'hydroides du littoral d'Ostende. Annls Soc. r. zool. malacol. Belg. 52 : 135-139. — 1923. Note sur quelques hydroides des cotes de France. Bull. Soc. zool. Fr. 48 : 13-20. BONNEVIE, K. 1898. Neue norwegische Hydroiden. Bergens Mus. Arb. (1898) (5) : 1-16. BOURNE, G. C. 1890. Notes on the hydroids of Plymouth. /. mar. biol. Ass. U.K. (N.S.) 1 : 391-398. BROCH, H. 1912. Hydroidenuntersuchungen. III. Vergleichende Studien an adriatischen Hydroiden. K. norske Vidensk. Selsk. Skr. (1911) (i) : 1-65. BRITISH LAFOEIDAE AND HALECIIDAE 419 BROCH, H. 1917. Hydroidenuntersuchungen. V. Studien iiber die Coppinia von Grammaria abietina. K. norske Vidensk. Selsk. Skr. (1916) (i) : 1-16. 1918. Hydroida. (Part II.) Dan. Ingolf Exped. 5 (7) : 1-206. 1927. 15 Hydrozoen. Tierwelt Dtl. 4 : 95-160. — 1928. Hydrozoa I. Tierwelt N.- u. Ostsee 3(b) : i-ioo. 1933- Zur Kenntnis der adriatischen Hydroidenfauna von Split. Skr. norske Vidensk- Akad. mat.-nat. Kl. (1933) 4 : 1-115. BRUCE, J. R., COLMAN, J. S. & JONES, N. S. 1963. Marine fauna of the Isle of Man. L.M.B.C. Mem. typ. Br. mar. PL Anim. 36 : i-3O7 + i-ix. CALDER, D. R. 1970. Thecate hydroids from the shelf waters of northern Canada. /. fish. Res. Bd Can. 27 : 1501-1547. CHAPMAN, D. M. 1969. The nature of cnidarian desmocytes. Tissue Cell, 1 : 619-632. CHOPIN, A. 1894. A visit to Cumbrae : The veteran naturalist : The Millport Biological station. Trans, a. Rep. Manchr micr. Soc. (1894) '• 43~54- CHRISTIANSEN, B. O. 1972. The hydroid fauna of the Oslo fiord in Norway. Norw. J. Zool. 20 : 279-310. CLARKE, S. F. 1875. Descriptions of new and rare species of hydroids from the New England coast. Trans. Conn. Acad. Arts Sci. 3 : 58-66. — 1876. Report on the hydroids collected on the coast of Alaska and the Aleutian Islands, by W. H. Dall, U.S. Coast Survey, and party, from 1871 to 1874 inclusive. In : W. H. Dall (Ed.), Scientific results of the exploration of Alaska, by the parties under the charge of W. H. Dall, during the years 1865-1874. 1 : 5-34. Washington. CORNELIUS, P. F. S. 1975. The hydroid species of Obelia (Coelenterata, Hydrozoa : Cam- panulariidae) , with notes on the medusa stage. Bull. Br. Mus. nat. Hist. (Zool.) 28 : 249-293. — 1976. Proposed validation of Halecium Oken, 1815 (Coelenterata : Hydroida), and suppression of Thoa Lamouroux, 1816 (Hydroida), under the plenary powers. Bull. zool. Nom. 32. (In press.) & GARFATH, J. B. Revisionary notes on type specimens of hydroids (Coelenterata : Hydroida) in the collections of Joshua Alder (1792-1867). (In prep.) CRAWFORD, J. H. 1895. The hydroids of St. Andrews Bay. Ann. Mag. nat. Hist. (6) 16 : 256-262. CROTHERS, J. H. (Ed.). 1966. Dale Fort marine fauna. 2nd edn. Fid Std. 2 (suppl.) : i-i69 + i-xxiv. CUVIER, G. L. C. F. D. 1797. Tableau elementaire de I'histoire naturelle des animaux. Paris. [The dating of this work follows /. Typogr. Biblphique (1797) 1 : 81-82.] DUERDEN, J. E. 1897. The hydroids of the Irish coast. Scient. Proc. R. Dubl. Soc. (N.S.) 8 : 405-420. ELLIS, J. 1755. An essay towards a natural history of the corallines, and other marine productions of the like kind, commonly found on the coasts of Great Britain and Ireland. London. & SOLANDER, D. 1786. The natural history of many curious and uncommon zoophytes, collected from various parts of the globe. London. FEY, A. 1969. Peuplements sessiles de 1'archipel de Glenan. i. Inventaire : Hydraires. Vie Milieu (B) 20 : 387-413. FLEMING, J. 1820. Observations on the natural history of the Sertularia gelatinosa of Pallas. Edinb. phil. J. 2 : 82-89. 1828. A history of British animals. Edinburgh. FORBES, H. O. 1872. List of hydroid zoophytes from the north-east coast of Scotland. Scott. Nat. 1 : 190-191. ERASER, C. M. 1912. Some hydroids of Beaufort, North Carolina. Bull. Bur. Fish., Wash. (1910) 30 : 337-387. I937- Hydroids of the Pacific coast of Canada and the United States. Toronto. 15 Although dated 1928 this publication was received in the Zoology Library, British Museum (Natural History), on 20 December 1927. 420 P. F. S. CORNELIUS FRASER, C. M. 1944. Hydroids of the Atlantic coast of North America. Toronto. — 1946. Distribution and relationship in American hydroids. Toronto. GARSTANG, W. 1900. Marine zoology. In : Doubleday, H. A. (Ed.), Victoria history of the counties of England : a history of Hampshire and the Isle of Wight, pp. 82-102. London. GRAY, J. E. 1848. List of the specimens of British animals in the collection of the British Museum. Part I. Centroniae or radiated animals. London. GRIFFITH, R. 1844. A synopsis of the characters of the Carboniferous limestone fossils of Ireland. Dublin. HAMOND, R. 1957. Notes on the Hydrozoa of the Norfolk coast. J. Linn. Soc. Zool. 43 : 294-324. 1963. Further notes on the Hydrozoa of the Norfolk coast. Ann. Mag. nat. Hist. (13) 6 : 659-670. HARGITT, C. W. 1924. Hydroids of the Philippine islands. Philipp. J. Sci. 24 : 467-505. HARTLAUB, C. 1895. Die Coelenteraten helgolands. Wiss. Meeresunters. Kiel, 1 : 161-206. 1905. Die Hydroiden der magalhaensischen Region und chilenischen Kiiste. Zool. Jb. (Suppl. 6) : 497-714. HASSALL, A. H. 1841. Supplement to a catalogue of Irish zoophytes. Ann. Mag. nat. Hist. (i) 7 : 276-287, 363-374- — 1848. Definitions of three new British zoophytes. Zoologist, 6 : 2223. & COPPIN, J. 1852. Descriptions of three species of marine zoophytes. Trans, microsc. Soc. Lond. 3 : 160-164. HERDMAN, W. A. 1891. Fourth annual report of the Liverpool Marine Biological Station on Puffin Island. Proc. Trans. Lpool biol. Soc. (1890) 5 : 19-72. HICKSON, S. J. & GRAVELY, F. H. 1907. Coelenterata. II. Hydroid zoophytes. Nat. Antarct. Exped. 1901-1904, Nat. Hist. 3 : 1-34. HINCKS, T. 1853. Further notes on British zoophytes, with descriptions of new species. Ann. Mag. nat. Hist. (2) 11 : 178-185. 1861. A catalogue of the zoophytes of south Devon and south Cornwall. Ann. Mag. nat. Hist. (3) 8 : 152-161, 251-262, 290-297, 360-366 ; 9 : 22-30. 1862. On the production of similar gonozoids by hydroid polypes belonging to different genera. Ann. Mag. nat. Hist. (3), 10 : 459-461. 1863. On some new British hydroids. Ann. Mag. nat. Hist. (3) 11 : 45-47. 1866. On Ophiodes, a new genus of Hydroida. Ann. Mag. nat. Hist. (3), 18 : 421-423. 1868. A history of the British hydroid zoophytes. 2 vols. London. i874a. Notes on Norwegian Hydroida from deep water. Ann. Mag. nat. Hist. (4) 13 : 125-137. • i874b. On deep-water Hydroida from Iceland. Ann. Mag. nat. Hist. (4) 13 : 146-153. HODGSON, M. M. 1950. A revision of the Tasmanian Hydroida. Pap. Proc. R. Soc. Tasm. (1949) : 1-65. HUVE, P. 1954. Hydranthea et Campalecium. Genres mediterraneens aberrants d'hydroides de la famille des haleciides. Reel Trav. Stn mar. Endoume, 13 : 173-192. JADERHOLM, E. 1905. Hydroiden aus antarktischen und subantarktischen Meeren, gesammelt von der schwedischen Sudpolarexpedition. Wiss. Ergebn. Schwed. Sudpolarexped. 5 (8) : 1-41. 1909. Northern and arctic invertebrates in the collection of the Swedish State Museum. IV. Hydroiden. K. svenska VetenskAkad. Handl. 45 (i) : 1-124. JOHNSTON, G. 1832. A descriptive catalogue of the recent zoophytes found on the coast of north Durham. Trans, nat. Hist. Soc. Northumb. 2 : 239-272. — 1838. A history of the British zoophytes. London. 1847. A history of the British zoophytes. 2nd edn. 2 vols. London. KOSSOWSKA, S. M. 1911. Contribution a la connaissance des hydraires de la Mediterranee occidentale. II. Hydraires calyptoblastiques. Archs Zool. exp. gen. (5) 6 : 325-352. KRAMP, P. L. 1929. Marine Hydrozoa. Zoology Faroes, 1 (5) : 1-59. BRITISH LAFOEIDAE AND HALECIIDAE 421 KRAMP, P. L. 1932. Hydroids collected in West-Greenland fjords in 1911 and 1912. Meddr. Gr0nland, 91 (3) : 1-35. 1935- Polypdyr (Coelenterata) . I. Ferskvandspolypper og Goplepolypper. Damn. Fauna, 41 : 1-208. 1938. Marine Hydrozoa. a. Hydroida. Zoology Iceland, 2 (^a) : 1-82. KUHN, A. 1913- Entwicklungsgeschichte und Verwandtschaftsbeziehungen der Hydrozoen. i Teil : Die Hydroiden. Ergebn. Fortschr. Zool. 4 (1-2) : 1-284. LAMARCK, J. B. P. A. DE 1816. Histoire naturelle des animaux sans vertebres. Volume 2. Paris. LAMOUROUX, J. V. F. 1816. Histoire des polypiers coralligenes flexibles, vulgairement nommes zoophytes. Caen. 1821. Exposition methodique des genres de I'ordre des polypiers. Paris. , BORY DE SAINT- VINCENT, J. B. G. M. & DESLONGCHAMPS, E. 1824. Histoire naturelle des zoophytes, ou animaux rayonnes, faisant suite a I'histoire naturelle des vers, de Bruguiere. In: Encyclopedic methodique (suppl.). Paris. LELOUP, E. 1947. Les ccelentere's de la faune Beige. Leur bibliographic et leur distribution. Mem. Mus. r. Hist. nat. Belg. 107 : 1-73. 1952. Coelenteres. Fauna Belg. pp. 1-283. LESLIE, G. & HERDMAN, W. A. 1881. The invertebrate fauna of the Firth of Forth. Edinburgh. LINKO, A. K. 1911. Hydraires (Hydroidea). Volume I. Haleciidae, Lafoeidae, Bonneviel- lidae et Campanulariidae. Fauna Rossii, pp. 1-250. LINNAEUS, C. 1758. Systema naturae. loth edn. Holmiae. 1761. Fauna svecica, sistens animalia Sveciae regni. 2nd edn. Stockholm. 1767. Systema naturae. I2th edn. Tom. I, Pars II. Holmiae. LORENZ, L. VON 1886. Polypomedusen von Jan Mayen. Beob.-Ergebn. Int. Polarforsch. 3 (6) : 25-28. MARINE BIOLOGICAL ASSOCIATION, 1957. Plymouth marine fauna. 3rd edn. Plymouth, England. MARSCHALL, A. F. 1873. Nomenclator zoologicus continens nomina systematica generum animalium tarn viventium quam fossilium, secundum ordinem alphabeticum disposita. Vindobonae. MILLARD, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43 : 173-243- 1964. The Hydrozoa of the south and west coasts of South Africa. Part II. The Lafoeidae, Syntheciidae and Sertulariidae. Ann. S. Afr. Mus. 48 : 1-56. ig66a. The Hydrozoa of the south and west coasts of South Africa. Part III. The Gymnoblastea and small families of Calyptoblastea. Ann. S. Afr. Mus. 48 : 427-487. ig66b. Hydroids of the Vema seamount. Ann. S. Afr. Mus. 48 : 489-496. 1967. Hydroids from the south-west Indian Ocean. Ann. S. Afr. Mus. 50 : 169-194. 1968. South African hydroids from Dr Th. Mortensen's Java - South Africa Expedition, 1929-1930. Vidensk. Meddr dansk. naturh. Foren. 131 : 251-288. & BOUILLON, J. 1973. Hydroids from the Seychelles (Coelenterata). Annls Mus. r. Afr. cent. 206 : 1-106. NAUMOV, D. V. 1960. Gidroidy i gidromeduzy morskikh, solonovatovodnykh i presnovodnykh basseinov S.S.S.R. Fauna S.S.S.R. 70 : 1-626. 1969. Hydroids and hydromedusae of the U.S. S.R. Fauna S.S.S.R. 70 : 1-660. Israel Program for Scientific Translations. NEAVE, S. A. 1940. A list of the names of genera and subgenera in zoology from the tenth edition of Linnaeus 1758 to the end of 1935. M-P. Nomencl. zool. 3 : 1-1065. NORMAN, A. M. 1867. Report of the committee appointed for the purpose of exploring the coasts of the Hebrides by means of the dredge. Part II. On the Crustacea, Echinoder- mata, Polyzoa, Actinozoa, and Hydrozoa. Rep. Br. Ass. Advmt Sci. (1866) : 193-206. NUTTING, C. C. 1899. Hydroida from Alaska and Puget Sound. Proc. U.S. natn. Mus. 21 : 741-753- 422 P. F. S. CORNELIUS NUTTING, C. C. 1901. No title. [Letter to the editor re nomenclatural revision of certain preoccupied names.] Am. Nat. 35 : 789. — 1927. Report on the Hydroida collected by the United States Fisheries steamer 'Alba- tross' in the Philippine region. Bull. U.S. nat. Mus. (100) 6 : 195-242. OKEN, L. 1815. Okens Lehrbuch der Naturgeschichte. Dritte Theil. Zoologie. Jena. PALLAS, P. S. 1766. Elenchus zoophytorum. The Hague. — 1774. Spicilegia zoologica. Tom. I, fasc. 10 : 1-42. Berolini. PARKE, M. & DIXON, P. S. 1968. Check-list of British marine Algae - second revision. /. mar. Uol. Ass. U.K. 48 : 783-832. PENNINGTON, A. S. 1885. British zoophytes : an introduction to the Hydroida, Actinozoa, and Polyzoa found in Great Britain, Ireland, and the Channel Islands. London. PFEFFER, G. 1889. Zur Fauna von Sud-Georgian. Jb. hamb. wiss. Anst. 6 : 37-55. PIEPER, F. W. 1884. Erganzungen zu 'Heller's Zoophyten etc. des adriatischen Meeres'. Zoo/. Anz. 7 : 148-152, 164-169, 185-188, 216-221. PRENANT, M. & TEISSIER, G. 1924. Notes 6thologiques sur la faune marine sessile des environs de Roscoff. Cirripedes, bryozaires, hydraires. Trav. Sin biol. Roscoff, 2 : 1-49. RALPH, P. M. 1958. New Zealand thecate hydroids. Part II. Families Lafoeidae, Lineo- lariidae, Haleciidae and Syntheciidae. Trans. R. Soc. N.Z. 85 : 301-356. RAY, J. 1724. Synopsis methodica stirpium Britannicarum. Edn. 3. London. REDIER, L. 1967. Revision de la collection du Museum des hydraires de Lamouroux. Bull. Mus. natn. Hist. nat. Paris (2) 39 : 381-410. REES, W. J. 1957. Evolutionary trends in the classification of capitate hydroids and medusae. Bull. Br. Mus. nat. Hist. (Zool.) 4 : 453-534. — & ROWE, M. 1969. Hydroids of the Swedish west coast. Acta R. Soc. scient. lift, gothoburg. 3 : 1-24. — & THURSFIELD, S. 1965. The hydroid collections of James Ritchie. Proc. R. Soc. Edinb. (B) 69 : 34-220. & WHITE, E. 1966. New records and fauna list of hydroids from the Azores. Ann. Mag. nat. Hist. (13) 9 : 271-284. RITCHIE, J. 1907. On collections of the Cape Verde Islands marine fauna made by Cyril Crossland, M.A. (Cantab.), B.Sc. (Lond.), F.Z.S., of St. Andrews University, July to September, 1904. The hydroids. Proc. zool. Soc. Lond. (1907) : 488-514. — 1911. Contribution to our knowledge of the hydroid fauna of the west of Scotland. Ann. Scot. nat. Hist. 77 : 29-34. ROBINS, M. W. 1969. The marine flora and fauna of the Isles of Scilly. Cnidaria and Cteno- phora. /. nat. Hist. 3 : 329-343. RUFFORD, P. J. 1902. Notes on British hydroid zoophytes and other subjects (polyzoan, con- chological & geological). Hastings, England. SARS, G. O. 1874. Bidrag til Kundskaben om Norges Hydroider. Forh. VidenskSelsk. Krist. (1873) : 91-150. SARS, M. 1850. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmarken. Nyt. Mag. Naturvid. 6: 121-211. - 1857. Bidrag til Kundskaben om Middelhavets Littoral-Fauna, Reisebemaerkninger fra Italien. Nyt. Mag. Naturvid. 9 : 110-164. SAVAGE, S. 1945. A catalogue of the Linnaean herbarium. London. STECHOW, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete, nebst Angaben iiber einige Kirchenpauer'sche Typen von Plumulariden. Zool. Jb., Syst. 42 : 1-172. — 1921. Uber Hydroiden der Deutschen Tief see-Expedition, nebst Bemerkungen iiber einige andre Formen. Zool. Anz. 53 : 223-236. — 1922. Zur Systematik der Hydrozoen, Stromatoporen, Siphonophoren, Anthozoen und Ctenophoren. Arch. Naturgesch. 88 (A) (Heft 3) : 141-155. — 1923. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete. II Teil. Zool. Jb., Syst. 47 : 29-270. BRITISH LAFOEIDAE AND HALECIIDAE 423 STEPHENS, J. 1905. A list of Irish Coelenterata, including the Ctenophora. Proc. R. Ir. Acad. 25 (B) : 25-92. STIMPSON, W. 1854. Synopsis of the marine Invertebrata of Grand Manan : or the region about the mouth of the Bay of Fundy, New Brunswick. Smithson. Contr. Knowl. 6 (5) : 1-67- STORM, V. 1882. Bidrag til Kundskab om Trondhejmsfjordens Fauna. IV. Om de i Fjorden forekomne hydroide Zoophyter. K. norske Vidensk. Selsk. Skr. (1881) (i) : 1-24. TEISSIER, G. 1965. Inventaire de la faune marine de Roscoff. Cnidaires-Ctenaires. Roscoff, France. THOMSON, W. T. C. 1853. Notes on some British zoophytes. Ann. Mag. nat. Hist. (2) 11 : 442-444. TORREY, H. B. 1902. The Hydroida of the Pacific coast of North America, with especial reference to the species in the collection of the University of California. Univ. Calif. Publs Zool. 1 : 1-104. TOTTON, A. K. 1930. Coelenterata. Part V. Hydroida. Nat. Hist. Rep. Br. antarct. Terra Nova Exped. 5 : 131-252. VERRILL, A. E. 1873. Results of recent dredging expeditions on the coast of New England. Am. J. Sci. (3) 5 : 1-16, 98-106. VERVOORT, W. 1941. Biological results of the Snellius expedition. XI. The Hydroida of the Snellius expedition (Milleporidae and Stylasteridae excluded) . Temminckia, 6 : 1 86 - 240. — 1942. Northern Hydroida in the collections of the Rijksmuseum van Natuurlijke Historic and the Zoological Museum at Amsterdam, with notes on their distribution. Zool. Meded. Leiden, 23 : 275-312. — 1946. Hydrozoa (C i). A. Hydropolypen. Fauna Ned. 14 : 1-336. - 1949. Notes on a small collection of hydroids from Jersey (Channel Islands). Zool. Meded. Leiden, 30 : 133-162. - 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. 1968. Report on a collection of Hydroida from the Caribbean region, including an annotated checklist of Caribbean hydroids. Zool. Verh. Leiden, 92 : 1-124. - 1972. Hydroids from the Theta, Vema and Yelcho cruises of the Lament -Doherty Geological Observatory. Zool. Verh. Leiden, 120 : 1-247. WATSON, J.E. 1969. Scoresbia. A new hydroid genus from South Australian waters. Trans. R. Soc. S. Aust. 93 : 111-116. WHITE, E. 1956. The hydroid Hydranthea margarica (Hincks) in Scottish waters. Scott. Nat. 68 : 39-42. WILLIAMS, G. 1954. Fauna of Strangford Lough and neighbouring coasts. Proc. R. Ir. Acad . 56 (B) : 29-133- WRIGHT, T. S. 1858. Observations on British zoophytes. Proc. R. phys. Soc., Edinb. 1 : 447-455- YAMADA, M. 1955. Some hydroids from Agattu, in the Aleutian Islands. Annotnes zool. jap. 28 : 121-125. INDEX Accepted generic and specific names are in Roman type, synonymous and invalid names in italic type. Names of higher taxa are in SMALL CAPITALS. Main page references are in italics. abietina, Abietinaria 379, 380, 381 abietina, Campanularia 381, 382 abietina, Grammaria 381, 382-385 abietina, Lafoea 379, 381 abietina, Reticularia 382 abietina, Salacia 382 Abietinaria 386 A. abietina 379, 380 Amphisbetia operculata 379 annulatum, Halecium 396 arboreum, Halecium 410 articulosum, Halecium 392, 406, 408, 409 A tvn.r,tMl,i.s AT2. Atractylis 412 A. margarica 412 424 P. F. S. 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 410 Diphasia rosacea 410 dumosa, Campanularia 385 dumosa, Capsularia 386 dumosa, Lafoea 385-391 dumosa, Sertularia 385 Dynamena pumila 415 echinata, Sertularia 402, 405 Eudendrium 410 E. pusillum 396, 398 EULAFOEINAE 377, 378 exigum, Halecium billardii var. 412 falcata, Hydrallmania 380 fenestrata, Thuiaria 381 Filellum 378, 381 F. serpens 379 filiforme, Halecium 403, 405, 406, 409 fruticosa, Campanularia 386 fruticosa, Lafoea 386, 389 geniculatum, Halecium 393, 396, 409 gracillima, Campanularia 386-387 gracillima, 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, 401—402 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 377 Hydrallmania falcata 380 Hydranthea 390, 412 H. margarica 412-414 Hydrodendron 414 H. caciniformis 414 Idiella pristis 411 immersa, Reticularia 378, 380 insignis, Grammaria 382, 383, 384 intermedia, Grammaria 382, 385 Kirchenpaueria pinnata 379 kofoidi, 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, 384 margarica, Atractylis 412 margarica, Hydranthea 412-414 margaricum, Halecium 412 medusiferum, Campalecium 391 mirabilis, Ophiodes 414 mirabilis, Ophiodissa 414-417 muricata, Sertularia 402, 405 muricatum, Halecium 402-405 murigatum, Halecium 403 nanum, Halecium 399 Nemertesia 406 N. ramosa 379 operculata, Amphisbetia 379 Ophiodes 414 O. caciniformis 414, 415, 417 O. mirabilis 414 Ophiodissa 390, 414 O. caciniformis 414, 415, 417 O. mirabilis 414-417 parvula, Campanularia 386, 390 parvula, Lafoea 390 pinnata, Kirchenpaueria 379 pinnata, Zygophylax 402 plumosum, Halecium 405, 406, 409 pocillum, Hebella 386, 390 pocillum, Lafoea 386, 390 pocillum, Lafoea fruticosa forma 386 pristis, Idiella 411 pumila, Dynamena 415 pusillum, Eudendrium 396, 398 pusillum, Halecium 396, 398 pygmaea, Lafoea 390 ramosa, Grammaria 382, 384 ramosa, Nemertesia 379 reflexum, Halecium 396, 398 Reticularia [BRACHIOPODA] 378 Reticularia [HYDROZOA] 378, 381 R. abietina 382 R. immersa 378-379, 380 R. serpens 379 robusta, Grammaria 381, 382 robustum, Halecium 401-402 robustum, Zygophylax 402 rosacea, 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 396 tenellum, Halecium 393, 398, tetracythara, Salacia 381 tetracythara, Thuiaria 381 tetracyttara, Thuiaria 381 Thoa 391 T. beanii 391 T. halecina 393 Thuiaria 381 T. fenestrata 381 T. tetracythara 381 P. F. S. CORNELIUS T. tetracyttara 381 torreyi, Halecium 391 Tubularia 414 undulatum, Halecium 396, 398, 411 washingtoni, Halecium 409 Zygophylax 399, 402 Z. pinnata 402 Z. robustum 402 Dr P. F. S. CORNELIUS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD ••' A LIST OF SUPPLEMENTS TO THE ZOOLOGICAL SERIES OF THE BULLETIN OF THE 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 and Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4. 3. TAYLOR, J. D., KENNEDY, W. J. & HAL£-, A. The Shell Structure and Mineralogy 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 ; 72 Text-figures. 1973. £9.70. 6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the Biology 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