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, I 7 I 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, 33, 34 . 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 4 12 attenuata, Obelia . . 260 Augaptilidae . . . 346-36 1 aurantonotus, Centropyge . . 227, 230 auropecten, Amallothrix. 33O-333, 334~33 6 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 . . . . 7 1 Cyrnea . . . 201-205, Pis. 1-2 dahli, Titanethes . . -73 daidaleos, Euplotes . . 4, 38, 39, 40 dalmatica, Philoscia . . -87 dalmatica, Philoscia muscorum . . 87 dalmatica, Stenophiloscia . 92 dalmaticus, Hyloniscus . . 7 1 dalmatinus, Armadilloniscus . . 79 damae, Hora . 83 danicus, Haplophthalmus . 71, 79 debilis, Ischioscia ... -83 debilis, Philoscia ... -83 Decapoda . . 237-247 deliculata, Obelia . 280 demarcata, Philoscia . 9 1 demarcata, Setaphora . 9* dentata, Charybdis six- . . 146, 147 dentata, Scolecithricella . 34 1 denticulata, Campanularia . . 279, 280 denticulata, Scolecithricella . . 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- 3 8 5 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- 4 I5 393 . 393-396, 399, 49. 4 12 391-4U, 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 . . . . 7 1 - 79, 97 harpa, Euplotes . . . . 17, 29, 30 Harris, E. A. . . . 199-2O5, 2 Pis. hastata, Chaetophiloscia . . .81 Hebella 377, 39<> Hebellidae . . -377 hedwigae, Gulella . . . . .167 helecina, Sertularia . . . 39 X 393 helgolandica, Obelia . . .266 Hemiplax ...... 245 hemisphaerica, Medusa .... 254 hendersonae, Clytia . . .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 244246 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 4 1 280 69 89 83 396-399, 401, 411 83 . 187 187, 188 187-188 377. 379, 381, 385-390 375-37 6 , 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 . 8 4 . 310 65, 78-93, 97-99 , . 63-iooe . 84 66 379 . 189 . 414 414 390, 414-417 . 189 188-189 \ . 189 146, H7 . 67 72, 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, 240242 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 . I6 7 . I6 7 344 . 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, 4243, 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. 45 393 405, 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 . 6 7 . 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, 5 1 Streptaxidae .... 157-175 striata, Obelia . . . . .281 Strongylocentrotus . . . 23, 37 sturanus, Trichoniscus noricus . . 74 Stylohylea ...... 72 Styloniscidae . . .65, 66-69, 96-97 Styloniscus .... 68-69, 96, 97 styricus, Cyphoniscellus . . . .71 suarezia, 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 2 5 2 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, 2 73 79 254, 438 INDEX Thoa . 391, 393 thomasi, Scottocalanus . . -313 thononensis, Euplotes . . . 54-55 thorn elyi, Obelia . . . . .280 Thuiaria . . . . . .381 Tilapia ..... 191-197 Tiroloscia ..... 92-93 Titanethes ...... 73 Titaniidae . . . . . 65, 69 tomentosa, Amphiura . . . 188-189 tomentosa, Trichorhina .... 79 torreyi, Halecium . . . 391 toticostata, Gulella . . . .167 tottoni, Laomedea . . . .281 Trewavas, E. . . 191-197, 2 33~ 2 35 triangularis, Uca ..... 245 triangulifera, Philoscia muscorum . . 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-HgCl a -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 (V 2 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). 4 o 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.) 4 6 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 n g) 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. 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. 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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. 7 o J. P. ELLIS AND R. J. LINCOLN Subgenus ILLYRIONETHES Verhoeff heroldi Verhoeff (19315 : 22) HOLOTYPE : male. Reg. no. 1931:4:27:73. Hercegovini, Yugoslavia. Verhoeff Collec- tion. strasseri (Verhoeff) (19275 : 270) SYNTYPE : male. Reg. no. 1928:7:4:55. Istrien, Italy. Collected by K. Strasser. Verhoeff Collection. SYNTYPES : three males ; one female. Reg. no. 1930:5:26:76-79. Istrien, Italy. Col- lected by K. Strasser. Verhoeff Collection. SYNTYPES: three males; one female. Reg. no. 1937:7:6:132-135. Istrien, Italy. Collected by K. Strasser. Verhoeff Collection. ANDRONISCUS Verhoeff alpinus Verhoeff (igoSc : 140, 143) SYNTYPE : female. Reg. no. 1908:6:1:30. Lugano, Italy. Verhoeff Collection. brentanus Verhoeff (1932 : 22) SYNTYPES : one male ; two females. Reg. no. 1937:7:6:57-59. Brenta, Italy. 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 : 2 4 6 ) [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 : I2 4) SYNTYPES : one male ; two females ; one other specimen in fragments. Reg. no. 1921:10: 18:2137-2139. Caracas. Collected by Meinert. Budde-Lund Collection. cornutus Budde-Lund (1885 : 228) SYNTYPES: one male; one female. Reg. no. 1921:10:18:2140-2141. California, USA. Budde-Lund Collection. nacreus Collinge (1922 : 108) SYNTYPES : one male; five females. Reg. no. 1922:11:10:1-5. Tamatave, east coast 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. 8 4 J. P. ELLIS AND R. J. LINCOLN LEPIDONISCUS Verhoeff germanicus carniolense Verhoeff (19280 : 127) SYNTYPE : female. Reg. no. 1928:7:4:28. Krain. Verhoeff Collection. pruinosus denticulatus Verhoeff (19280 : 126) SYNTYPES : two males ; four females. Reg. no. 1928:7:4:23-27. Italy. Collection. Verhoeff MARIONISCUS Barnard spatulifrons Barnard (1932 : 234) SYNTYPES : ten males ; twelve females. Reg. no. 1933:1:25:67-74. Peninsula, S. Africa. Collected and presented by K. H. Barnard. Hout Bay, Cape NAHIA Budde-Lund hirsuta (Budde-Lund) (1906 : 89) [Philoscia hirsuta] SYNTYPES : mixed with other (non-type) specimens collected by Schultze in Cape Town. Reg. no. 1921:10:18:2022-2033. Simonstown, S. Africa. 19.6.1903. Budde-Lund Collec- tion. OLIBRINUS Budde-Lund pigmentatus Budde-Lund (i9i3b : 390) SYNTYPES : two females. Reg. no. 1913:1:8:154-155. Coin, Peros, Chagos Archipelago, Indian Ocean. Collected by the Percy Sladen Trust Expedition. Presented by J . S. Gardiner. ONISCUS Linnaeus kenepurensis Chilton (1901 : 135) SYNTYPES: four males. Reg. no. 1900:11:1:43-46. Kenepuru, New Zealand. 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: 34 J -348- BUDDE-LUND, G. 1885. Crustacea Isopoda Terrestria, per familias et genera et species descripta. 3i9pp. Hauniae. - 1893. Landisopoder fra Venezuela, insamlede af Dr. Fr. Meinert. Ent. Meddr. 4 : m- 129. 1895. Viaggio di Leonardo Fea in Birmania e regioni vicine. 64. Isopodi terrestri. Ann. Mus. Genova, 34 : 602-612. 1902. A list of the terrestrial isopods. In : Lanchester, W. F. On the Crustacea collected during the 'Skeat' Expedition to the Malay Peninsula. Pt II. Proc. zool. Soc. Lond., pp. 379-3 81 - 1904. A Revision of 'Crustacea Isopoda Terrestria' with additions and illustrations. 2. Spherilloninae. 3. Armadillo, pp. 33-144. Kj0benhaven. - 1906. Die Landisopoden der Deutschen Siidpolar-Expedition 1901-1903 mit 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* io 4 A. L. RICE & R. W. INGLE Mauritiana in the east and from Nova Scotia to Brazil in the west (Christiansen, 1969). It is therefore not surprising that it was one of the first crabs to have 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, an d 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. 5 a, 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. 7 a, b) Antennule : 8-10 aesthetascs in 2 or 3 groups, plus a terminal seta. Ventral flagellum well developed (Fig. 3b) Antenna : Spinous process about length of rostral spine. Exopod about length of spinous process ; 2 terminal setae. Endopod > exopod (Fig. $g) Mandible : Palp now present Maxillule : Similar to stage IV Maxilla : Schaphognathite with 30-36 marginal setae First maxilliped : As in stage IV but with 12 natatory setae (Fig. 3>z) Second maxilliped : Endopod terminal segment with 5 setae. Exopod with 12 natatory setae (Fig. 30) Third maxilliped : Well developed, biramous but unarmed Pereiopods : Limbs clearly segmented CEPHALOTHORAX As in previous stages (Fig. rf, e) As puber Spinous process > 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. 7 c, 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 I 4 2 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 (924' S 462o' 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 6 i 10 3-8 X 1-6 mm 2-44 1-9 mm 1-2 x I-I mm 6 i 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 6 i 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 6 i 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 6 i 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 2 4 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 6 i 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 6 i 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 6 i 38 4' 1 .X 1-7 mm 2-44 2-2 mm 1-3 X 1-2 mm 6 i 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 4 8 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) an d 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 y rs -' 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 7 o 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 3j-fecundation {( pouch 8 lumen vas deferens penis FIGS 3-11. 3, Shell drawing ; 4, Shell shape variability ; 5, Jaw ; 6, Body pigmentation ; 7, Ventral view of the reproductive tract ; 8, Receptacula complex ; 9, Terminal genitalia ; 10, Prostate gland ; n, Cross section of the penis. 182 C. M. PATTERSON from Q. concisa (Morelet) (Patterson, 1968, 1971). However, Q. spurca and Q. concisa appear to be more closely related to each other than Q. aldabraensis is 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 i8 4 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 fig 8 - STUDER, T. 1885. Die Seesterne Siid-Georgiens nach der Ausbeute der deutschen Polar- station in 1882 u. 1883. Jb. hamb. wiss. Anst. 11 : 143-166, 2 pis. THEEL, H. 1886. Report on the Holothurioidea dredged by H.M.S. "Challenger" during the years 1873-1876. Part 2. Rep. scient. Results Voy. "Challenger" (Zool.) 39 11-290, 1 6 pis. VERRILL, A. E. 1876. Echinoderms. In : Kidder, J. H. Contribution to the Natural History of Kerguelen Island. Washington, pp. 68-75. [Also in : Bull. U.S. natn. Mus. 3.] F. W. E. ROWE Department of Marine Invertebrates THE AUSTRALIAN MUSEUM 6-8 COLLEGE STREET SYDNEY, N.S.W. 2000 AUSTRALIA AILSA M. CLARK Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 560 A NEW SPECIES OF TILAPIA (PISCES, CICHLIDAE) IN THE ZAMBIAN ZAIRE SYSTEM By ETHELWYNN TREWAVAS & DONALD J. STEWART SYNOPSIS A new species of the cichlid genus Tilapia A. Smith, 1840, is described from the Luongo River, a tributary of the Luapula in northern Zambia. It is assigned to the subgenus Tilapia. Com- pared to its near relative, T. sparrmanii, the new species is more elongate, with higher 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 an d 2 males, 100 + 26 and 85-5 + 22 mm from Nsenga stream (formerly ROM 28120). DESCRIPTION. Meristic characters from holotype and all paratypes, 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 I 3'3-i7'2 divided by its depth 0-66-0-99 0-96-1-25 (only 2 < i-o) Scales : 1.1. series 26 f . 2 f. o 27 18 o 28 17 o 2 9 5 I5j 30 2 6 D-l.l. 3 f.i 9 f. o 3i 24 6 4 5 15 4i i i cheek rows 2 f.37 f. o 3 7 22 Dorsal spines XIII f. 2 f. o XIV 38 3 XV 9 19 Total dorsal rays 23 f. i o 24 ii 3 25 3 17 26 7 2 Modal formula XIV n XV 10 Vertebrae 26 f. 2 f. o 27 22 3 28 8 17 29 o i T. ruweti (Poll & Thys van den Audenaerde) is also found on the 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. I 9 6 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 M 3 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 P 1 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 P 2 . A relatively large deciduous P 1 is present in Z58i, with only a slight concavity on P 2 , 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 I 1 s 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 P l s. 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 P 1 , present or absent 14. Infundibula of lower incisors, present or absent 15. Shape of mandibular symphysis pos- teriorly, tall or low 16. Surface behind lower incisors, flat or concave flat V = agreement with Z58i ; x = difference from burchell' 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