'/ -> 1 I . BULLETIN OF /& j GttRAL THE BRITISH MUSEUM^ i 6 ju NI976 (NATURAL HISTORY) ZOOLOGY Vol. 27 1974 BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1976 DATES OF PUBLICATION OF THE PARTS No. i . . . . . .13 June 1974 No. 2 -"i . . . . .18 July 1974 No. 3 19 July 1974 No. 4 . . . . .20 September 1974 No. 5 . . . . .23 December 1974 No. 6 . . . . .23 December 1974 No. 7 . . . . .24 December 1974 ISSN 0007-1498 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU CONTENTS ZOOLOGY VOLUME 27 PAGE No. I. Anatomy of head and neck in the Huia (Heteralocha acutirostris) with comparative notes on other Callaeidae. By P. J. K. BURTON (i PI.) . . i No. 2. Miscellanea .......... 49 A new skink (Reptilia : Scincidae : genus Emoia) from the New Hebrides. By LORD MEDWAY ...... 53 Scutocyamus parvus, a new genus and species of whale-louse (Amphipoda : Cyamidae) ectoparasitic on the North Atlantic white-beaked dolphin. By R. J. LINCOLN and D. E. HURLEY (i PL) . . 59 Catalogue of the whale-lice (Crustacea : Amphipoda : Cyamidae) in the collections of the British Museum (Natural History). By R. J. LINCOLN and D. E. HURLEY 65 A review of Laephotis Thomas, 1901 (Chiroptera : Vespertilioni- dae). By J. E. HILL 73 The genus Finmarchinella Swain, 1963 (Crustacea : Ostracoda) and its species. By J. W. NEALE (2 Pis) .... 83 Euplotes rariseta sp. n. (Protozoa : Ciliatea). A new small marine hypotrich. By C. R. CURDS, B. J. WEST and J. E. DORAHY (i PL) 95 Observations on clonal cultures of Eugfypha acanthophora and Euglypha strigosa (Testacea : Protozoa). By R. H. HEDLEY, C. G. OGDEN and J. I. KRAFFT (2 Pis) 103 Descriptions of three species of Euplotes (Protozoa : Ciliatea). By C. R. CURDS (i PL) 113 New records of bats from Southeastern Asia, with taxonomic notes. By J. E. HILL 127 No. 3. The Haplochromis species (Pisces : Cichlidae) of Lake Rudolf, East Africa. By P. H. GREENWOOD 139 No. 4. A review of Scotoecus Thomas, 1901 (Chiroptera : Vespertilionidae) . By J. E. HILL 167 No. 5. Catalogue of the types of terrestrial isopods (Oniscoidea) in the collections of the British Museum (Natural History) I. Superfamily Pseudotracheata. By R. J. LINCOLN and J. P. ELLIS . . 189 No. 6. Campylaspis species (Crustacea : Cumacea) from the deep Atlantic. By N. S. JONES -247 No. 7. A new family, genus and species of bat (Mammalia : Chiroptera) from Thailand. By J. E. HILL 301 Index to Volume 27 . . . . . . 337 INDEX TO VOLUME 27 The page numbers of the principal references and the new taxonomic names are printed in bold type. abbreviatus, Cyamus abnormis, Gelsana . abyssinicus, Microcercus . Acaeroplastes . Acanthoniscus acanthophora, Euglypha . Actoecia Actoeciidae aculeata, Campy laspis acuminatus, Rhinolophus . . 65 . 214 . 215 193 . 219 . 103-112, PI. I . 218 . 191, 218 250, 253, 268-270 . 131 acuminatus, Rhinolophus acuminatus 131-132, 137 acuminatus acuminatus, Rhinolophus 131-132, 137 acuminatus audax, Rhinolophus . . 132 acuminatus calypso, Rhinolophus . -132 acuminatus circe, Rhinolophus . . .132 acuminatus sumatranus, Rhinolophus . 132 acustiserra, Porcellio .... 197 acutirostris, Heteralocha . . 1-48, i PI. Adinda ....... 244 aediculatus, Euplotes . . . .124 aegaeum, Armadillidium .... 208 aegaeus, Porcellio . . . . . 197 aenigma, Diploexochus .... 223 aerarius, Diploexochus .... 224 affinis, Armadillo . . . . .219 affinis, Campylaspis. . 250, 254, 263, 277 affinis, Cubaris . . . . .219 affinis, Euplotes 113, 114, 116, 117-1 19, 120, 123 affinis tricirratus, Euplotes . 113, 119, 123 Agabiforius . . . . . -237 Agabiformius ...... 193 Agnara ....... 205 Akermania . . . . . .219 alassiense, Armadillidium . . 208, 212 alatus, Euplotes .... 97, 101 Alauda . . . . . . -41 alba, Campylaspis . . 250, 251, 256, 258 albanicum, Armadillidium . . . 208 alberti, Diploexochus . . . .224 albertianus, Haplochromis . . 150, 159 albescens, Diploexochus .... 224 albicornis, Lucasius .... 191, 197 albicornis, Mesarmadillo . . . -215 albicornis, Porcellio . . . . 197 albifrons, Armadillidium .... 208 albigula, Scotoecus . . 1 80, 1 8 1, 182, 183 albigula, Scotoecus hindei 180, 181, 182-i83 albinus, Porcellio . . . . .198 albipennis, Tura ..... 204 albirostris, Lagenorhynchus . . 59, 71 albofuscus, Scotoecus 169, 171, 172, 174-175, 176, 177, 186 albofuscus, Scotoecus albofuscus . 1 75-176 albofuscus, Scotophilus . . . 169, 175 albofuscus albofuscus, Scotoecus . 175-176 albofuscus cinnamomeus, Scotoecus . . 175 albofuscus woodi, Scotoecus . 175, 176-177 albomaculatus, Buddelundia . . .222 albomarginatus, Armadillo . . .219 albomarginatus, Porcellio . . . 198 albospinosus, Armadillo . . . .219 album, Armadillidium .... 208 alloniscus ...... 205 alluaudi, Porcellio . . . . .198 alticola, Diploexochus . . . .224 altimontis, Saidjahus .... 229 alveolata, Euglypha .... 104 ambitiosus, Armadillo . . . .229 ambitiosus, Spherillo .... 229 amblyoda, Campylaspis . . . 250, 251 Amblyrhamphus . . . . -37 americanum, Ethelum . . . -213 americanus, Mesarmadillo . . .213 amieti, Euplotes . . . . .124 amoenus, Porcellio ..... 198 Amphipoda . . . 59-64, I PI.; 65-72 ampullatus, Hyperoodon . . . -71 Anchicubaris . . . . . .219 anconanum, Armadillidium . . . 208 anconanus, Metoponorthus pruinosus . 197 aneityumensis, Emoia . . . 53-55, 56 Angara ....... 193 angolensis, Laephotis 73, 76-77, 7 8 - 79, 80, 81 angolensis, Laephotis wintoni . . -73 angularis, Campylaspis . . . 250, 251 angulata, Barentsovia . . 86, 87, 88-89, 91, 92, Pis. 1-2 angulata, Cythere .... 87, 88 angulata, Cythereis . . .83, 88 angulata, Finmarchinella . . 86, 87, 88-89, 90, 91, 92, Pis. 1-2 angulata, Hemicythere . . 86, 87, 88, 89 angulata, "Hemicytherinae" . . .88 angulata, Nereina . . 86, 87, 88, 90, 91 angusta, Tura . .204 angustulus, Porcellio .... 198 anna, Ciridops . 25, 28 annectans, Myotis . . 33 annulicornis, Cylisticus .... 192 antarctica, Campylaspis . . 250, 256, 299 Anthochaera . 34 apenninorum, Armadillidium . . . 208 338 INDEX apenninorum, Trachelipus . . . 207 apenninorum, Tracheoniscus . . . 207 aperta, Campylaspis . . . 250, 253 apfelbecki, Armadillidium . . . 208 approximates, Metoponorthus . . .196 apulicus, Trachelipus .... 207 apulicus, Tracheoniscus .... 207 Arachnothera . . . . . .42 arcuata, Campylaspis . 250, 253, 265-266 arcuatus, Porcellio ..... 202 argentarium, Armadillidium . . . 208 argentarius, Acaeroplastes . . . 193 argentarius, Metoponorthus . . 193 Armadillidae . . 197, 219-232, 244-245 Armadillidiidae . 191, 208-213, 240-242 Armadillidium 208-212, 213, 221, 240, 241, 242 Armadillo . 208, 219-221, 222, 224, 225, 226, 227, 228, 229, 230, 231, 232, 244, 245 armatus, Benechinus . . . .213 artinii, Scotoecus . . . 180, 181, 182 artinii, Scotoecus hindei . . . .182 asifensis, Metoponorthus .... 197 asifensis, Metoponorthus sexfasciatus . 197 aspera, Campylaspis . . . 250, 255 aspheronicus, Euplotes . . . .124 assimile, Armadillidium .... 209 astriger, Armadillidium .... 208 astriger, Armadillo . . . .208 ater, Porcellio . . . . .198 aternanus, Trachelipus larii . . . 207 aternanus, Tracheoniscus larii . . . 207 aterrimus, Phoeniculus . . . 37, 45 Atlantidiidae . . . . .191 atrox, Phoniscus . . . 135-136, 137 audax, Rhinolophus aciminatus . . 132 aulacoeis, Campylaspis . . . 250, 252 Aulonocara . . . . . .160 auritus, Porcellio ..... 198 ausseli, Armadillo . . . . .219 australis, Eubalaena glaciolis . 67, 68, 69, 71 avium, Haplachromis . . . .159 badium, Armadillidium .... 208 badium siculorum, Armadillidium . . 208 Balaena . . . . . 67, 71 Balaenoptera . . . . 65, 66, 71 balaenopterae, Cyamus . . . 65-66, 71 Balantiopteryx . . . . -3*7 Balionycteris . . . . .128 Balloniscidae .... 191, 205 Balloniscus ...... 205 balstoni, Scoteinus . . . . . 173 balteatus, Euplotes ..... 101 balticus, Euplotes . . . . . 101 balticus, Porcellio ..... 207 balticus, Trachelipus .... 207 balticus burzenlandicus, Porcellio . . 207 balticus burzenlandicus, Trachelipus . . 207 barbertoni, Bethalus . . . .221 Barentsovia . .83, 84, 86, 87-92, Pis. 1-2 barentzovoensis, Barentsovia . 83, 86-88, 89, 90, 91, Pis. 1-2 barentzovoensis, Finmarchinella 83, 86-88, 89, 90, 91, Pis. 1-2 barentzovoensis, Nereina . . . 83, 86 barnardi, Bethalus ..... 222 barnardi, Cubaris . . . . .222 barroisi, Porcellio ..... 198 batesoni, Porcellio . . . . .198 benaci, Metoponorthus . . . .196 Benechinus . . . . . .213 bergomatius, Cylisticus plumbeus . .192 besi, Periscyphis . . . . .216 Bethalus .... 221-222, 245 bicarinata, Campylaspis . 250, 254, 287-28g bicarinata, Merulana .... 228 bicarinata, Spherillo .... 228 bicolor, Exzaes . . . . .212 bicoloratus, Armadillo .... 229 bicoloratus, Periscyphis . . . .216 bicoloratus, Spherillo .... 229 bicornis, Mahehia . . . . . 195 biellensis, Cylisticus. .... 192 bifrons, Armadillo ..... 229 bifrons, Spherillo . . . . .229 bimaculatus, Hemichromis . . .141 binotatus, Buddelundia . . . .223 bipartitus, Buddelundia .... 223 bistriatus, Porcellio . . . .198 bisulcatus, Euplotes . . . 99, 101 bituberculatus, Armadillo . . .219 bituberculatus, Diploexochus . . .224 bizonatus, Periscyphops . . . .217 blattarius, Porcellio .... 198 bloyeti, Haplochromis . . 145, 162, 163 blythi, Rhinolophus . . . 131, 137 blythi, Rhinolophus blythi . . -131 blythi blythi, Rhinolophus . . -131 blythi calidus, Rhinolophus . . .131 blythi szechwanus, Rhinolophus . -131 bocki, Nesodillo ..... 228 bodenheimeri, Hemilepistus . . -194 bonetti, Campylaspis . . 250, 251, 261 boopis, Cyamus .... 66-67, 71 borelli, Plataoniscus .... 205 borellii, Alloniscus ..... 205 botswanae, Laephotis . 73, 76, 77-79, 80, 81 brachycephalus, Spherillo. . . .229 brachypterus, Philetor . . . . 134 brachypterus, Philetor brachypterus . . 135 brachypterus, Vespertilio . . -134 brachypterus brachypterus, Philetor . .135 brachypterus rohui, Philetor . . . 135 brachypterus verecundus, Philetor 134-135, 137 bredanensis, Steno .... 70, 71 brentanus, Trachelipus .... 207 brentanus, Tracheoniscus .... 207 brevicornis, Armadillo . . . .219 brevicornis, Campylaspis 250, 254, 270-271 brevipennis, Porcellio . . . .198 INDEX 339 brevis, Ignamba brevis, Phalaba buddelundi, Porcellio Buddelundia . bulbosa, Campylaspis . 215 . 206 . 198 222-223 250, 254, 273-274, 289 bulgaricus, Trachelipus .... 207 bulgaricus, Tracheoniscus . . . . 207 bullatus, Haplochromis . . 159, 160, 161 Burton, P. J. K. . . . l- 4 8, i PI. burzenlandicus, Porcellio balticus . . 207 burzenlandicus, Trochelipus balticus . . 207 Cacicus ..... 36, 37 cadornae, Pipistrellus .... 303 caelatum, Armadillidium .... 208 Caeroplastes . . . . . . 193 calidus, Rhinolophus blythi . . -131 caligans, Spherillo . . . . .229 Callaeas . . 3, 4, 6, 7, n, 12, 14, 22, 23. 26, 35-36, 44 Callaeidae .... 1-48, i PI. callosus, Buddelundia .... 223 calmani, Porcellio ..... 198 calypso, Rhinolophus acuminatus . .132 Campy laspides . . . . .270 Campylaspis ..... 247-3 oo Campylorhamphus . . . . -42 canaliculata, Campylaspis . 250, 252, 263 canaliculata, Merulana . . . .228 canaliculatus, Spherillo .... 228 canariensis, Armadillo .... 224 canariensis, Diploexochus. . . .224 canariensis, Porcellio . . . .198 caperata, Campylaspis . . . 250, 254, 279-280, 282 carinatus, Armadillo .... 222 carinatus, Bethalus ..... 222 carinatus, Hemiporcellio . . . -195 carniolense, Armadillidium . . . 208 Carolliinae . . . . . .319 carpathicus, Protracheoniscus saxonicus . 207 carthaginensis, Porcellio . . . .198 carunculata, Anthochaera . . -34 carunculatus, Creadion . 3, 4, 5, 6, 10, n, 34 cassida, Armadillo ..... 222 cassida, Bethalus . . . . .222 castor, Diploexochus . . . .224 catodon, Physeter . . . 66, 67, 70, 71 catodontis, Cyamus . . . 67, 71 caucasius, Cylisticus . . . .192 caudatus, Ourachaerus .... 232 cavernicola, Periscyphis . . .216 cayennensis, Porcellio . . . . 199 cela, Cacicus . . . . . -37 celsicauda, Diploexochus .... 224 ceti, Cyamus ..... 67, 71 charon, Euplotes . . 97, 113, 115, 123 chindeensis, Periscyphis . . .216 Chironax .... 127-129, 137 Chiroptera 73-82; 128-138; 167-188; 301-336 Cichlidae ..... 139-165 ciliata, Euglypha . . . . .104 Ciliatea . . 95-102, i PI.; 113-126, i PI. cilicius, Metoponorthus . . . .196 cinctus, Armadillo ..... 220 cinctutus, Pyrgoniscus . . . .232 cinerascens, Buddelundia . . . 223 cinerea, Callaeas . . . 3, 4, 6, n, 12, 22, 23, 26, 35-36 cinerea, Creatophora . . . -37 cingendum, Armadillidium . . . 208 cingendum, Armadillidium maculatum . 208 cingulatus, Diploexochus .... 224 cinnamomomeus, Scotoecus albofuscus . 175 circe, Rhinolophus acuminatus . . . 132 Circoniscus . . . . . .218 circumdatus, Pipistrellus . . 132-134, *37 Ciridops ..... 25, 28 civilis, Periscyphis . . . . .216 clathrata nuda, Cythere . . . 86, 87, 88 clausus, Synarmadillo . . . .231 clavata, Campylaspis . . 250, 255, 292 Coelops ...... 320 cognata, Campylaspis 250, 254, 275, 277-278 cognatus, Porcellio . . . .199 Coleura ....... 323 collicolus, Porcellidium .... 206 collicolus, Porcellio ..... 206 collicolus, Porcellium .... 206 collinus, Armadillo . . . .220 coloratus, Diploexochus .... 224 compressa, Euglypha . . . .104 concinella, Normanicythere . . -85 concinnus, Scleropactes . . . .218 congener, Armadillo . . . .224 congener, Diploexochus . . . .224 conglobator, Armadillo . . . ^ . 220 conisaleus, Diploexochus . . . 224 contra, Sturnus . . . . -37 Contractus, Porcellio . . . . 199 Corcorax . . . . . .29 corcyraeum, Armadillidium . . . 209 cornutus, Rhinolophus . . . -131 cornutus pumilus, Rhinolophus . . -131 Corvus ....... 38 costata, Campylaspis . . 250, 253, 266 costata speciosa, Campylaspis . . . 266 coxalis, Metoponorthus .... 196 Graseonycteridae .... 301-336 Craseonycteris .... 301-336 crassus, Armadillo . . . . .224 crassus, Diploexochus .... 224 Creadion . . . 3, 4, 5, 6, 7, 10, n, 14. 34- 35. 36. 38. 44 Creatophora . . . . . -37 crenosus, Euplotes ..... 101 creper, Saidjahus ..... 229 crispa, Campylaspis. . . . 250, 253 cristatus, Euplotes . . . 99, 101, 121 340 INDEX cristatus, Hemilepistus .... 194 cruentatus, Porcellio . . . . 199 Crustacea . 59-64, i PL; 65-72; 83-94, 2 Pis.; 189-246; 247-300 Cubaris . 219, 220, 221, 222, 225, 226, 231 Cumacea ..... 247-300 Curds, C. R. . 95-102, i PL; 113-126, i PL curvicosta, Barentsovia curvicosta, Finmarchinella Cushmanidea Cyamidae Cyamus .... cyanomelas, Rhinopomastus Cylisticidae Cylisticus Cythere .... Cythereis . 83, 86, 87, 88, 89, 90-91, Pis. 1-2 . 83, 86, 87, 88, 89, 90-91, Pis. 1-2 . 86 59-64, i PL ; 65-72 59, 65-70, 71 . 42 191, 192 . 192 86, 87, 88 . 83, 84, 88 dalmatinus frascatensis, Orthometopon . 206 danae, Armadillo ..... 230 danae, Spherillo . . . . .220 davidi, Armadillidium .... 209 debueni, Porcellio ..... 199 decoratus, Spherillo . . . .229 decumanus, Psarocolius . . . -37 delphini, Isocyamus . . . 70, 71 dentifrons, Cylisticus . . . .192 depressus, Armadillo .... 222 depressus, Bethalus .... 222 Desertoniscus . . . . . .193 desertorum, Protracheoniscus . . . 206 Desmodontidae diadaleos, Euplotes . Diclidurinae . Diclidurus dimorphus, Porcellionides Diploexochus . disjunctus, Diploexochus . dispar, Porcellio dispersus, Spherillo . . dollfusi, Eubelum Dolphin, white-beaked Dorahy, J. E. dugesi, Armadillo dugesi, Diploexochus dumorum, Armadillo dumorum, Venezillo ecaudatus, Diploexochus echinata, Campylaspis egens, Armadillo elbanum, Armadillidium Ellis, J. P. elongatus, Hemilepistus Eluma . emarginatus, Bethalus 319, 320, 322, 330 . 124 319 323, 325. 326, 333 . 204 223-227, 245 . 224 . 199 . 229 . 214 59-64, i PL . 95-102, i PL 225 225 232 232 225 250, 253, 270 . 220 . 209 . 189-2 4 6 194 212, 242 . 222 emarginatus, Scotophilus . Emballonuridae . 317, 322, 323, 328, 329, Emballonuroidea Emoa .... Emoia .... emunitus, Armadillo emunitus, Bethalus . Ennurensis episimus, Hekelus . epops, Upupa Eptesicus erinaceus, Armadillo erinaceus, Spherillo . erraticus, Cyamus . erythroleucus, Armadillo . Eschrichtius . esterelanum, Armadillidium esterelanus, Cylisticus Ethelum Eubalaena Eubelidae . . 191 Eubelum Euglypha Euglyphidae . Euplotes . 95-102, i Euporcellio eurystomus, Euplotes 118, evansi, Porcellio exarata, Campylaspis 250, Exzaes . 173 318, 319, 320, 321, 324. 325, 326, 327, 330. 332, 333, 334 329, 334 . 56 53-58 . 222 . 222 . . . 193 213 37. 42 74. 134 . 229 . 229 . 67-68, 71 . 220 70. 71 . 2O9 . 192 213-214 67, 68, 69, 71 , 213-218, 242-244 214, 215, 242, 243 103-112, 2 Pis. . IO3-II2, 2 Pis. PL; 113-126, i PL . 207 119, I2O, 121, 124 . 199 254, 266, 284-285 . 212 falabae, Scotoecus . . 169, 178, 183, 184 falabae, Scotoecus hindei . . . 183-184 Falculea. . . . . . -42 famosa, Nectarinia . . . .42 feae, Armadillo . . . . .228 feae, Pericephalus ..... 228 festivus, Armadillo ..... 225 festivus, Diploexochus .... 225 ficorum, Porcellio . . . . .199 ficulneus, Porcellio . . . . . 199 finmarchica, Cythereis . . . 83, 84 finmarchica, Finmarchinella . 83, 84-85, 86, Pis. 1-2 finmarchica, Hemicythere . . .84 Finmarchinella . . . 83-94, 2 Pis. fissifrons, Pagana . . . . .206 flavocinctus, Porcellio . . . . 199 flavoscutatum, Armadillidium . . . 209 flavovittata, Porcellio . . . . 199 flavus, Armadillo ..... 220 fongosiensis, Anchicuboris . . .219 fontium, Protracheoniscus . . . 206 formosana, Nagara .... 205 formosanus, Nagurus .... 205 fragilis, Agnara . . . . .205 frascatensis, Orthometopon dalmatinus . 206 INDEX frigida, Campylaspis . . . 250, 255 fritschei, Reductoniscus .... 229 frontalis, Sphaeroniscus . . . .218 frontalis, Spherillo . . . . .230 frontexcavatum, Armadillidium . . 209 frontirostre, Armadillidium . . . 209 frontosus, Buddelundia .... 223 frontosus, Metoponorthus . . . 196 furcatum, Armadillidium .... 209 furcatus, Diploexochus . . . .225 Furipteridae . 318, 319, 320, 326, 327, 328, 330 fuscomarmoratus, Metoponorthus . .196 fusiformis, Campylaspis . . . 250, 252 grisescens, Spherillo. grossus, Armadillo . grossus, Spherillo guttata, Campylaspis Gymnastinops 230 230 . 230 250, 251, 258-259 39 galapagoensis, Cubans . galapagoensis, Diploexochus galeatus, Armadillo . galeatus, Pericephalus . gallicus, Porcellio . . Gelsana . . . . gemmulatus, Porcellio . Gerufa .... Gerutha ... gestroi, Armadillidium . gibbosus, Eschrichtius . gigas, Armadillidium simoni gigas, Cubaris .. gigas, Diploexochus . glabra, Campylaspis . glacialis, Eubolaena. . glacialis, Eubolaena glacialis glacialis australis, Eubalaena glacialis glacialis, Eubalaena glacialis japonica, Eubalaena glebulosa, Campylaspis . Globicephala ... globosa, Campylaspis globus, Synarmadillo . Glomerulus ... glomus, Armadillo . . glomus, Diploexochus . Glossophaga . . . gordoniensis, Diploexochus gracilior, Merulana translucida gracilipennis, Cylisticus . gracilis, Cyamus . . grandinatum, Armadillidium graniger, Porcellio . . granulata, Campylaspis . granulatum, Armadillidium . . granulatum peloponnesiaca, Armadillidium granuliferus, Porcellio ... gravei, Metoponorthus . . . ... 225 . . . 225 ... 228 ... 228 . . . 199 . . .214 ... 199 ... 194 .. 214-215 . . . 209 . . 70, 71 . . .211 ... 225 . . .225 . 250, 252, 261 . . .69 . 68, 69, 71 . 67, 68, 69, 71 . 68, 69, 71 . . 69, 71 . 250, 255, 293-295 ... 70, 71 250, 255, 291, 295, 297 . . . .231 .... 228 . . . .225 .... 225 . . . -325 . 225 . 228 .192 68, 71 . 209 . 199 250, 252 .210 209 200 196 Greenwood, P. H grenadensis, Armadillo grenadensis, Venezillo greyii, Scoteinus . griseoalbus, Armadillo griseoalbus, Bethalus 139-i65 . 232 .232 . 173 .222 .222 Haloporcellio .. Haplarmadillo . . Haplochromis ... harpa, Euplotes . . harsadiensis, Armadillo . hartae, Campylaspis . hebridarum, Spherillo . Hedley, R. H. . . Hekelus . . . Hemichromis . . Hemicythere ... "Hemicytherinae" . . Hemignathus . . . . Hemilepistus .... Hemiporcellio . . . hermagorensis, Protracheoniscus Hermanites . . . . herzegovinensis, Porcellio . . herzegovinensis, Porcellium . Heteralocha .... Hiallum . . . . hilgendorfi, Eubelum . . hilgendorfi, Hiallum . . ... 194 . . -215 .. 139-I&5 . . 101, 119 ... 220 . . 250, 253 ... 230 103-1 12, 2 Pis. . . .213 . . .141 84, 86, 87, 88, 89 . . .88 . . 5, 42 .. 194, 237 . . .195 206 86 206 206 i PI. 215 215 215 1- Hill, J. E. 73-82; 127-138; 167-188; 301-336 hindei, Nycticeius hirundo . . .182 hindei, Scotoecus . 169, 171, 172, 175, 178, 179-i8o, 181, 182, 183, 184, 185, 186 hindei, Scotoecus hindei 179, 180-i82, 183, 184 hindei, Scotoecus hirundo . 181, 182, 183 hindei albigula, Scotoecus 180, 181, 182-i83 hindei artinii, Scotoecus . . . .182 hindei falabae, Scotoecus . . . 183-1 84 hindei hindei, Scotoecus 179, 180-i82, 183, 184 Hipposideridae 317, 318, 319, 320, 322, 323, 325, 326, 327, 328, 329, 330, 332, 333 Hipposideros ...... 303 hirsutulus, Protracheoniscus . . . 206 hirticornis, Gerufa . . . . . 194 hirtum, Armadillidium .... 209 hirtus, Agabiformius . . . . 193 hirtus, Lucasius ..... 193 hirundo, Scotoecus . . 169, 171, 172, 175, 177-179, 181, 183, 184, 185, 186 hirundo, Scotophilus . . . . 177 hirundo hindei, Nycticeius . . .182 hirundo hindei, Scotoecus . 181, 182, 183 hispida, Metoponorthus . . . .196 hispida, Porcellio . . . . .196 hispida, Porcellionides . . . 196 hispidus, Ennurensis . . . . 193 holosericeus, Amblyrhamphus . . -37 horrida, Campylaspis . . 250, 255, 292 342 INDEX horridoides, Campylaspis . horsfieldii, Myotis . horsfieldii, Myotis horsfieldii horsfieldii horsfieldii, Myotis horsfieldii lepidus, Myotis. hughscotti, Periscyphis Huia .... Hurley, D. E. hybridum, Armadillidium hybridum, Schizidium Hyperoodon . hypotoreus, Spherillo Hypotrichida . hypselos, Diploexochus hypsinephes, Diploexochus 250, 255, 292-293 . 132 . 132, 137 . 132, 137 132 . 216 1-48, i PI. 59-64, i PI. ; 65-72 . 213 . 213 . 71 230 95-102, i PI.; 113 . 225 . 225 Ignamba . . . . . .215 ignavum, Eubelum . . . . .214 imbutus, Porcellio ..... 200 imbutus pellegrinensis, Porcellio . . 200 immotus, Armadillo .... 220 immsi, Hemiporcellio . . . . 195 immsi, Porcellio . . . . .195 inaequalis, Buddelundia .... 223 incanus, Porcellio ..... 200 Inchanga ...... 195 incicus, Scleropactes . . . .218 incisa, Nagara ..... 205 incisus, Nagurus ..... 205 inconspicuus, Armadillo .... 220 indentatus, Euplotes .... 101 ingens, Spherillo ..... 230 iniqua, Merulana ..... 228 iniquus, Spherillo ..... 228 inkystans, Euplotes . . . . 101 inornata, Campylaspis . . . 250, 251 instrenuum, Eubelum . . . .214 integer, Armadillo .... 220 intercalarius, Porcellio .... 200 intermedia, Campylaspis 250, 255, 277, 289, 297 intermixtus, Armadillo .... 220 interpolator, Porcellio .... 200 Isocyamus ..... 59, 70, 71 japonica, Barentsovia . . . 91-92 japonica, Eubalaena glacialis . . 69, 71 japonica, Finmarchinella . . . 91-92 japonica, Nereina . . . . 83, 91 jelskii, Porcellio ..... 200 johnstoni, Campylaspis . . . 250, 253 johorensis, Tadarida . . . 136-137 Jones, N. S 247-30O jonesi, Campylaspis . . . 250, 253, 263 kaokoensis, Diploexochus . . . 225 kiiensis, Campylaspis . . . 250, 252 Kerivoula Kerivoulinae . Kisuma . kogmani, Diploexochus Krafft, J. I. . kuhlii, Scotophilus . kunenensis, Periscyphis kunenensis, Periscyphops . 328 319, 320, 325 232 225 103-112, 2 PIS. 174 . 216 . 216 labiatus, Buddelundia .... 223 lacteolus, Metoponorthus .... 196 Laephotis ..... 73-82 laevigata, Campylaspis 250, 251, 258, 260-26i laevigatus, Buddelundia .... 223 laevis, Gerutha . . . . .214 laevis, Porcellio . . . . .201 Lagenorhynchus .... 59, 71 lamellatus, Porcellio . . 191, 194, 201 laminigerum, Armadillidium . . .210 larii, Trachelipus ..... 207 larii, Tracheoniscus ..... 207 larii aternanus, Trachelipus . . . 207 larii aternanus, Tracheoniscus . . . 207 Lasiurus ...... 328 lateralis, Buddelundia .... 223 laticarpa, Campylaspis . . . 250, 254 laticauda, Mahehia .... 195 laticauda, Tura ..... 204 latidactyla, Campylaspis . . . 250, 253 Latidens . . . . . -303 latifrons, Armadillo . . . .222 latifrons, Bethalus . . . . .222 latissimus, Periscyphis . . . .216 latissimus, Porcellio. .... 200 latus, Euplotes patella .... 124 latus, Protracheoniscus .... 206 legendrei, Campylaspis . . . 250, 253 lekaguli, Hipposideros .... 303 lentus, Spherillo ..... 230 lepidus, Myotis . . . . .132 lepidus, Myotis horsfieldii. . . .132 lepidus group, Rhinolophus . . .129 lepidus subgroup, Rhinolophus . .129 lepineyi, Porcellio ..... 200 Leptotrichus . . . . . 195, 237 leucocephalus, Microcercus . . -215 leucocephalus, Periscyphis . . -215 leucorhamphus, Cacicus . . . -37 leucostictus, Sarotherodon . . .163 lifuensis, Armadillo ..... 220 lifuensis, Cubaris . . . . .220 limbata, Periscyphis . . . .216 limbatus, Bethalus . . . .222 limenites, Diploexochus . . . .225 Lincoln, R. J.. 59-64, I p l-.' 66-72; 189-246 lineare, Trinema . . . . -103 Liponycteris . . . . . -323 littoralis, Metoponorthus . . . .196 INDEX 343 loati, Haplochromis . . . . .162 Lonchorhina ...... 326 longicauda, Porcellio .... 200 longipennis, Porcellio .... 200 longipes, Armadillo .... 225 longipes, Diploexochus .... 225 loyaltiensis, Emoia samoensis . . 53-58 lubricum, Eubelum . . . . .214 Lucasius . 191, 193, 195, 197, 201, 237 lugubris orarum, Porcellio . . .201 lugubris vizzavonensis, Porcellio . .201 lusitanus, Porcellio .... 202 lusitanus, Porcellio scaber . . . 202 lutshniki, Trachelipus .... 207 lutshniki, Tracheoniscus .... 207 Lygosoma . . . . . -56 Lyprobius ...... 205 macconneli, Haplochromis . 141, 149, 153, 154-i6i, 162, 164 macmahoni, Armadillo .... 220 macrocneme, Miniopterus medius . 135, 137 macrodens, Bethalus .... 222 macrophthalma, Campylaspis . 250, 253, 264 macrops, Gerufa ..... 194 maculata, Balloniscus .... 205 maculata, Campylaspis . . . 250, 255 maculata, Mahehia . . . . 195 maculata, Philoscia .... 205 maculatum cingendum, Armadillidium . 208 maculipes, Porcellio. .... 200 maculosa, Pagana ..... 206 maculosus, Spherillo .... 230 madagascariensis, Agnara . . . 205 madagascariensis, Metoponorthus . . 205 magna, Sturnella . . . . -37 magnicirratus, Euplotes . . . . 101 magnificus, Porcellio .... 200 mahagiensis, Haplochromis . . .162 Mahehia. ...... 195 makuae, Diploexochus .... 226 Mammalia . 127-138; 167-188; 301-336 mammillata, Sunniva . . . -213 mansa, Campylaspis 250, 254, 283, 286-287 marcidus, Armadillo . . . .228 marcidus, Pericephalus .... 228 mareoticum, Armadillidium . . .210 marginalis, Uramba. .... 205 marginatus, Spherillo . . . -230 marginenotatus, Porcellio. . . . 200 marginepilosa, Gerutha . . . -215 marinensium, Armadillidium . . .210 marmorata, Gerufa . . . . .194 marmoratus, Synarmadillo . . .231 marquesarum, Spherillo .... 230 marquesarum, Xestodillo .... 230 marshalli, Rhinolophus .... 303 mauritanica, Campylaspis . 250, 251, 261 mayeti, Armadillo . . . . .220 medius, Miniopterus . . . -135 medius macrocneme, Miniopterus . 135, 137 Medway, Lord .... 53-58 Megadermatidae . . 318, 319, 320, 322, 325, 326, 327, 329, 330, 332 Megaptera . . . .66, 67, 70, 71 mehelyi, Protracheoniscus . . . 206 mehelyi, Protracheoniscus politus . . 206 meiringi, Diploexochus . . . .226 melaena, Globicephala . . . 70, 71 melanocephalus, Chironax . 127-129, 137 melanurus, Acaeroplastes . . -193 melanurus, Metoponorthus . . . 193 meleagris, Metoponorthus . . . 197 meleagris, Metoponorthus pruinosus . 197 Merulana ...... 228 Mesarmadillo . . 213, 214, 215, 243 Metoponorthus . . 193, 196-197, 199, 200, 204, 205, 206, 237, 238 197 215, 243 318, 322, 324, 325 325 228 . 228 . 215 . 201 319, 327 135. 137. 328 250, 253 . 129, 131 . 213 . 129 193 . IOI 213 193 . 22O . 2I 4 . 214 113, 114, 115-117, Il8, 119, I2O, 121, 123 . 201 moebiusi quadricirratus, Euplotes . 113, 123 mohamedanicum, Armadillidium . .210 Molossidae . . 318, 319, 320, 322, 324, 325, 326, 327, 328, 329, 330 monocellatus, Haplarmadillo . . -215 monoceros, Monodon . . 68, 69, 71 Monodon .... 68, 69, 71 monodontis, Cyamus . . . 68, 71 montagui, Diploexochus .... 226 montana, Gerufa . . . . . 194 montanus, Armadillo .... 220 montanus, Porcellio. .... 203 montanus, Porcellio spinipennis . . 203 monticola, Buddelundia .... 223 montivagus, Armadillo .... 230 montivagus, Spherillo .... 230 Mica Microcercus Microchiroptera Micronycteris . microps, Armadillo . microps, Glomerulus microps, Ignamba . mildei, Porcellio Miniopterinae . Miniopterus minor, Campylaspis minor, Rhinolophus. minor, Sunniva minor subgroup, Rhinolophus minuta, Angara minuta, Euplotes minuta, Pareluma . minutus, Agabiformius miser, Armadillo modestum, Ethelum modestus, Mesarmadillo . moebiusi, Euplotes moebiusi, Porcellio 344 INDEX Mormoopidae . 319. 320, 323. 325. 326. 327, 328, 329, 330, 332, 333 325 . 222 250, 256, 297-2QQ . 2IO . 320 205 . 205 65, 71 100, 117 132, 137. 303 Mormoops tnucidus, Bethalus . multinodosa, Campylaspis muricatum, Armadillidium Murininae mus, Lyprobius mus, Uramba musculus, Balaenoptera . mutabilis, Euplotes Myotis .... myrmecophilus, Metoponorthus myrmecophilus 196 myrmecophilus, Myrmeconiscus myrmecophilus 196 myrmecophilus myrmecophilus, Metoponorthus 196 myrmecophilus myrmecophilus, Myrmeconiscus 196 myrmicidarum, Metoponorthus . .196 Mystacinidae . ' . mystica, Sunniva Mysticeti mysticetus, Balaena Myzopodidae . 318, 319, 320, 324, 325. 327. 329, 33 . 213 . 71 67, 71 318, 320, 326, 327, 330 Nagara . Nagurus . namibensis, Laephotis nana, Nagara . Nannastacidae nanus, Diploexochus nanus, Nagurus narentanus, Porcellio nasatum, Armadillidium . 205 . 205, 239 73, 75-76, 77, 79, 80 . 205 . 249 . 226 205 . 201 . 212 nasatum sorrentinum, Armadillidium . 210 nasatus, Cylisticus ..... 192 natalensis, Inchanga .... 195 Natalidae . 318, 320, 326, 327, 328, 330 naupliensis, Porcellio Neale, J. W. . nebulosus, Diploexochus Nectarinia Neocyamus Nereina . Nesodillo . nicklesi, Porcellio nicobaricus, Armadillo nicobaricus, Spherillo . nigra, Emoia . nigra, Porcellio nigricans, Periscyphis nigripes, Buddelundia . nigrobrunneus, Metoponorthus nigromarginatus, Armadillo nigropunctatus, Periscyphis nigropunctatus, Synarmadillo . 2OI 83-94, 2 Pis . 226 . 42 59, 70, 71 83, 86, 87, 88, 90, 91 . 228 f . . 201 230 230 53 . 201 . 217 . 223 . 196 . 221 231 . 231 niloticus, Sarotherodon nitens, Campylaspis. nitidulus, Armadillidium . nobilis, Spherillo . . Noctilio . Noctilionidae . . 318, 325, 326 noctulinus, Nycticejus noctulinus, Vespertilio nodosus, Cyamus nodosus, Hemilepistus nodulosa, Campylaspis normani, Lucasius . normani, Porcellio . Normanicythere novaeangliae, Megaptera . nuda, Campylaspis . nuda, Cythere clathrata . Nyctalus Nycteridae . 318, 319, 325. 326, 327 Nycticeius Nyctophilinae 140, 149 250, 251, 256, 258 . 210 230 329 319, 320, 323, 324, 327, 329, 330, 332 i?4 . 172, 174 68-69, 7i 194 . 250, 255 . 2OI . . 201 8 5 66, 67, 70, 71 250, 251, 256-258 86, 87, 88 . 171 320, 322, 323, 324, 329, 330, 332, 333 169, 170, 171, 173, 174, 175, 182, 186 317, 322 obliquidens, Diploexochus . . . 226 obliquipes, Spherillo . . . -230 obsoletus, Porcellio ..... 201 obtusa, Angara . . . . . 193 obtusifrons, Porcellio . . . .201 obtusiserra, Porcellio .... 201 obtusus, Agabiformius .... 193 occidentals, Protracheoniscus . . .196 ocellatus, Porcellio . . . . .201 octocarinatus, Euplotes . . . .124 octocirratus, Euplotes . . . 100, 101 odherni, Armadillidium . . . .210 odherni, Armadillidium peraccai . .210 Odontoceti . . . . . -71 oertzeni, Armadillidium . . . .213 oertzeni, Schizidium . . . .213 officinalis syriaca, Armadillo . . .221 Ogden, C. G 103-1 12, 2 Pis. oliveti, Armadillidium . . . .210 olivieri, Porcellio . . . . .198 Oniscoidea ..... 189-246 opacus, Buddelundia .... 223 opihensis, Actoecia . . . .218 orarum, Porcellio lugubris . . . 201 orarum, Porcellio orarum . . .201 orarum orarum, Porcellio . . .201 orarum vizzavonensis, Porcellio . . 201 orientalis, Campylaspis . . . 250, 251 ormeanum, Armadillidium . . .210 ornatus, Scotomanes . . . . 173 Orodillo 228 orphanus, Diploexochus .... 226 Orthometopon ..... 206 Ostracoda .... 83-94, 2 Pis. INDEX 345 Ourachaerus . ovalis, Campylaspis . ovalis, Cyamus ovampoensis, Cubaris ovampoensis, Diploexochus oxyzomus, Armadillo oxyzomus, Cubaris 232 250, 253 69, 71 . 226 . 226 . 221 . 221 pachytos, Diploexochus pacifica, Campylaspis pacificus, Cyamus paeneglabra, Campylaspis Pagana . palaestinus, Hemilepistus pallasi, Armadillidium palliata, Falculea pallidus, Cylisticus . pallidus, Lucasius pallidus, Porcellio pallidus, Scotoecus . pallidus, Scotophilus Pandanus panurus, Armadillo . panurus, Bethalus . papillata, Campylaspis papillosa, Kisuma Paracubaris paradoxolophus, Rhinolophus Paraperiscyphis Pareluma parietinus, Porcellio parkei, Euplotes parvus, Armadillo parvus, Porcellio parvus, Scutocyamus . parvus, Spherillo patella, Euplotes patella latus, Euplotes paucinodosa, Campylaspis paucispina, Campylaspis . pauper, Porcellio pauperculus, Diploexochus pellegrinense, Armadillidium pellegrinensis, Porcellio imbutus . . 200 Pelmatochromis . . . . .141 peloponnesiaca, Armadillidium granulatum 209 peltatus, Spherillo . . . . -230 penicilliger, Haloporcellio . . . -194 peraccae, Armadillidium . . . .210 peraccai, odherni, Armadillidium . .210 Pericephalus ...... 228 Periscyphis . . 215,216-217,218,231,243 Periscyphops . . .216,217-218,243,244 perkinsi, Armadillo . . . . .231 perkinsi, Spherillo . . . . .231 petraeum, Armadillidium . . . .210 petronius, Haplochromis . . . 161, 163 phaeacorum, Porcellio rathkei . . . 207 . 226 . 250, 251 70, 71 250, 251, 258, 261 . 206 194 . 2IO . 4 2 . 192 . 195 195 171, 172-174, 175, 186 169, 170, 172, 173 195 . 222 . 222 250, 255, 296 232 . 218 303 . 216 213 . 2OI 119-121, 122, 123, 124, PI. I . 230 . 201 59-64, i PL; 71 230 97. 99. IO . 117. 124 . 124 2 5. 255, 291-292 250, 254, 275-277 . 201 . 226 . 210 phaeacorum, Trachelipus . Phalaba .... phaleronensis, Metoponorthus . phaleronensis, Orthometopon phaleronensis, Porcellio pharyngalis, Haplochromis Philetor Philoscia .... philoscoides, Metoponorthus Phoeniculus .... Phoniscus .... Phyllostomatidae 317, 319, 320, 327. Phyllostomatoidea . physalus, Balaenoptera Physeter .... physeteris, Neocyamus pictus, Rotungus piger, Armadillo pila, Gerutha .... pileus, Campylaspis . pilosa, Campylaspis . 250, pilosa, Gerutha pilularis, Armadillo . pilum, Eubelum Pipistrellus . . . 132- Pisces ..... pisum, Armadillo pisum, Diploexochus planarius, Porcellio . Plataoniscus .... Platycyamus .... platyuropus, Campylaspis . plicata, Campylaspis 250, 254, plicata, Tadarida plumbeus, Cylisticus plumbeus bergomatius, Cylisticus plumbeus umbricus, Cylisticus . plumipes, Euplotes . politus, Leptotrichus politus mehelyi, Protracheoniscus pollex, Diploexochus Polyacanthus .... polythele, Diploexochus . pontremolensis, Cylisticus . porcata, Campylaspis . 250, 252, Porcellidium .... Porcellio 191, 194, 195, 196, . 207 206, 239, 240 . 206 . 206 . 206 161, 163 134-135. 137 205 . 196 37-45 135-136, 137 322, 325, 326, 329, 330. 332 329 . 65, 66, 71 66, 67, 70, 71 70, 71 . 218 . 221 . 215 250, 252 254, 271-272 215 . 221 . 215 134. 137. 303 . 139-165 . 226 . 226 . 202 . 205 59.70-71 250, 252, 259 280-282, 289 136, 137 . 192 . 192 . 192 117, 119, 124 195 . 206 . 226 . 229 . 226 Porcellionidae Porcellionides . Porcellium . porphyrivagus, Caeroplastes porphyrivagus, Metoponorthus . portofinense, Armadillidium praeustus, Porcellio . pretoriensis, Armadillo pretoriensis, Bethalus Prionoplus . procerus, Hemignathus . 192 261-2&3, 265 . 206 197-204, 206, 207, 238, 239 I9L 193-205, 237-239 196, 204 . 206 193 193 210 2O2 222 222 4 42 346 INDEX Protozoa 95-102, i PI.; 103-112, 2 Pis.; 113-126, i PI. Protracheoniscus . . . 196, 206-207 provincialis, Porcellio .... 202 proximatus, Armadillo . . . .221 pruinosus, Metoponorthus . 199, 200, 204 pruinosus anconanus, Metoponorthus . . 197 pruinosus meleagris, Metoponorthus . . 197 Psarocolius ..... 37, 39 pseudoratzeburgi, Euporcellio . . . 207 pseudoratzeburgi, Porcellio . . . 207 pseudoratzeburgi, Trachelipus . . . 207 pseudorcae, Syncyamus . . . .62 Pseudotracheata .... 189-246 Pteropodidae . . . . . -325 pujetanum, Armadillidium . . .211 pujetanus, Porcellio .... 202 pulchella, Campylaspis . . . 250, 251 pulcher, Periscyphis . . . ..217 pumila, Campylaspis . . 250, 252, 263 pumilus, Armadillo ..... 232 pumilus, Rhinolophus cornutus . . -131 pumilus, Venezillo ..... 232 purpurascens, Eluma . . . .212 purpurascens, Spherillo . . . .231 purpureus, Phoeniculus . . . -37 purpureus, Porcellio ..... 202 pusillus, Rhinolophus . . 129-131, 137 pusillus group, Rhinolophus . . 129,131 pustulosa, Campylaspis . . . 250, 255 pygmaeus, Armadillo . . . .221 pygmaeus, Periscyphis . . . -231 pygmaeus, Synarmadillo . . . -231 pyrenaeus, Porcellio ..... 202 Pyrgoniscus ...... 232 pyriformis, Tetrahymena . . . -97 Pyrrhocorax . . . . . -37 pyrrhocorax, Pyrrhocorax . . -37 quadricirratus, Euplotes moebiusi . 113,123 quadrimaculatus, Diploexochus . . .226 quadrimaculatus, Mesarmadillo . -215 quadrimaculatus, Periscyphis . . .217 quadriplicata, Campylaspis . . 250, 254 quadriserriatum, Armadillidium . .211 quadritracheata, Buddelundia . . . 223 quietum, Eubelum . . . . .214 quinquecarinatus, Euplotes . . 97, 101 quinquepustulatum, Armadillidium . .211 ragusae, Porcellio ..... 202 raikovi, Euplotes . . .99, 100, 101, 123 rariseta, Euplotes . 95-IO2, i PL; 115, 118 rathkei phaeacorum, Porcellio . . . 207 razae, Alauda . reaumuri, Hemilepistus redacta, Campylaspis . 250, Reductoniscus reflexum, Ethelum . reflexus, Mesarmadillo rehobotense, Armadillidium Reptilia .... reticularis, Prionoplus reticulata, Campylaspis Rhinolophidae 318,319,320, 327. 328, Rhinolophoidea Rhinolophus . Rhinopoma . . 304, 316, Rhinopomastus Rhinopomatidae3i6, 317, 318, 323, 324, 325, rhodesiae, Bethalus . rhodesiensis, Diploexochus ribauti, Sphaerobathytropa riparium, Armadillidium . robusta, Arachnothera rohui, Philetar rohui, Philetar brachypterus rosai, Armadillidium roscida, Campylaspis roscoffensis, Euplotes rostellata, Campylaspis 250 rostrata, Campylaspis rotunda, Euglypha . Rotungus Rousettus rubicunda, Campylaspis . 250 rubromaculata, Campylaspis, rudolfianus, Haplochromis 153, rufa, Campylaspis rufescens, Diploexochus ruficauda, Periscyphis rufobrunneus, Agabiformius rufomarginatus, Spherillo . rugifrons, Buddelundia rugulosus, Cubaris . rugulosus, Spherillo . rupta, Campylospis . sabuleti, Metoponorthus sabulifer, Porcellio . Saccolaimus . Saccopteryx . sagamiensis, Campylaspis Saidjahus saldanhae, Diploexochus salimalii, Latidens . . 41 194 254, 274-275, 278 . 229 . 214 . 214 . 211 53-58 4 . 250, 253 323, 324, 325, 326, 329, 33, 332, 333 332 129-132, 137, 303 ,321,323,332,333 . 38,42,45 319, 320, 321, 322, 326, 328, 329, 330, 332, 333. 334 . 222 . 227 . 218 . 211 . 4 2 134. 135 135 . 211 . 250, 254 . IOI , 253, 267-268, 270 250, 255, 268, 289 103, 108 . 218 . 129 ,251, 259-26o, 261 250, 255 141, 142-150, 152, 155. 156, 157. 158. 159, 160, 161, 162 . 250, 252 . 227 . 217 193 . 231 . 223 . 231 231 . 250, 252 . 197 . 202 323 323 250, 255 . 229 . 227 303 INDEX 347 salisburyensis, Diploexochus . . .227 saltuum, Porcellio ..... 202 samoense, Emoa . . . . .56 samoense, Lygosoma . . . -56 samoensis, Emoia .... 54, 55, 56 samoensis loyaltiensis, Emoia . . 53-58 sanctum, Armadillidium . . . .211 sanfordi, Emoia .... 55, 56 sarajevensis, Porcellio .... 202 sarasini, Nesodillo . . . . .228 sarculatus, Porcellio ..... 202 Sarotherodon .... 148, 149, 163 sauteri, Orodillo . . . . .228 savonense, Armadillidium . . . .211 saxicola, Haplochromis . . . -157 saxonicus, Protracheoniscus . . . 207 saxonicus carpathicus, Protracheoniscus . 207 scaber, Porcellio .... 199, 201 scaber lusitanus, Porcellio . . . 202 scaberrimum, Armadillidium . . .211 scabrum, Armadillidium . . . .211 scammoni, Cyamus .... 70, 7 1 schellenbergi, Nesodillo .... 228 schirasi, Hemilepistus .... 194 Schizidium . . . . . -213 schlieffenii, Nycticeius . . . . 175 schlieffenii, Scoteinus . . 173, 1 8 1, 182 Scincidae ..... 53-58 Scleropactes . . . . .218, 244 Scoteinus . . 170, 171, 173, 181, 182, 186 Scotoecus ..... 167-i88 Scotomanes . . . . . . 173 Scotophilus . . . 169, 170, 171, 172, 173, 174, 175, 177 Scotozous . . . . . .169 Scutocyamus . . 59-64, l Pl- 7 1 serratipes, Campylaspis . . . 250, 254 serratum, Armadillidium . . . .211 setaceus, Spherillo . . . . .231 sexfasciatus, Metoponorthus . . . 197 sexfasciatus asifensis, Metoponorthus . . 197 sharpi, Armadillo . . . . .221 siculorum, Armadillidium badium . . 208 silvanus, Periscyphops . . . .217 silvarum, Armadillo .... 227 silvarum, Diploexochus . . . .227 similis, Campylaspis . . . . 250,252 simoni, Armadillidium . . . .211 simoni gigas, Armadillidium . . .211 simplex, Synarmadillo .... 232 simulator, Porcellio ..... 202 sinuosa, Campylaspis . . . 250, 253 Skink . .... 53-58 sociablis, Porcellio ..... 203 societatis, Pipistrellus . . . 133, 134 solitarius, Cacicus .... 36, 37 sollers, Spherillo . . . . .231 sordidus, Porcellio ..... 203 sorrentinum, Armadillidium nasatum . 210 spatulata, Porcellio . . . . .203 spatulatus, Porcellio .... 200 speciosa, Campylaspis . . . 250, 253 speciosa, Campylaspis costata . . .266 speiseri, Emoia .... 54, 55 Sperillo ....... 245 speyeri, Armadillidium . . . .212 Sphaerobathytropa . . . . .218 Sphaeroniscidae . . . 191, 218, 244 Sphaeroniscus . .... 218,244 Spherillo . . .220,228,229-231,245 sphinx, Haloporcellio . . . 191, 194 spicatus, Spherillo . . . . .231 spinicornis, Porcellio .... 203 spinifera, Campy laspides . . . .270 spiniger, Acanthoniscus . . . .219 spinipennis, Porcellio .... 203 spinipennis, Porcellio spinipennis . 202, 203 spinipennis montanus, Porcellio . . . 203 spinipennis spinipennis, Porcellio . 202, 203 spinipes, Porcellio ..... 203 spinosa, Akermania . . . . .219 spinosa, Campylaspis . 250, 253, 267, 270 spinosus, Circoniscus . . . .218 spinosus, Paracubaris . . . .218 spretus, Porcellio ..... 203 squamatus, Periscyphops . . . .217 squamifera, Campylaspis . 250, 255, 293, 295, 296 squamosus, Periscyphops . . . .217 stebbingi, Paraperiscyphis . . .216 steenbrasi, Diploexochus . . . .227 Steno . . . . . . 70, 71 Stenoderminae . . . . .319 sticta, Campylaspis . . 250,253,264-265 stipulatum, Eubelum . . . .214 stolikanum, Armadillidium . . .212 straeleni, Haplochromis . . . .162 strelkovi, Euplotes ..... 101 striata, Campylaspis . . . 250, 252 stricticauda, Armadillo .... 222 stricticauda, Bethalus . . . .222 strigosa, Euglypha . . . 103-1 12, PI. 2 Sturnella . . . . . -37 Sturnus ....... 37 subbadius subgroup, Rhinolophus . .129 subdentatum, Armadillidium . . .212 suberorum, Cylisticus .... 192 subjaponica, Cushmanidea . . .86 submersa, Campylaspis . 250, 255, 290-291 subterraneus, Desertoniscus . . . 193 subtransversus, Periscyphis . . .217 succinctus, Porcellio .... 203 sulcata, Campylaspis . . 250, 253, 275 sulcatus, Buddelundia .... 223 sumatranus, Rhinolophus acuminatus . 132 Sunniva ...... 213 sylvatica, Exzaes . . . . .212 Synarmadillo ..... 231-232 Syncyamus ..... 59, 62 syriaca, Armadillo omcinalis . . .221 szechwanus, Rhinolophus blythi . -131 INDEX tabularis, Diploexochus . . . .227 Tadarida . . . 136-137 tamei, Periscyphis . . . . .217 Taphozous . . 323, 326, 328, 333 tarangensis, Spherillo . . . .231 tardus, Mica ...... 197 tardus, Porcellio ..... 197 taschkententis, Protracheoniscus . . 207 tegulatus, Euplotes .... 101, 124 temminckii, Scotophilus . . . . 174 tendanum, Armadillidium . . .212 tenuipunctatus, Armadillo . . .221 Testacea .... 103-112, 2 Pis. testacea, Tura ..... 204 Tetrahymena . . . . . -97 thetidis, Campylaspis . . . 250, 255 thompsoni, Campylaspis . . . 250, 252 thompsoni, Platycyamus . . . 70-7 1 thomsoni, Cyamus . . . . .70 thonglongyai, Craseonycteris . 301-336 Thyropteridae . 318,319,320,326,327,330 Tura .... turkanae, Haplochromis . . 204 141, 149, 150-153, 155. i 60 tigris, Armadillidium Tilapia . tirolense, Armadillidium tomentosus, Buddelundia Toradjia torulosa, Campylaspis tosaensis, Hermanites trachealis, Porcellio . Trachelipidae . Trachelipus .... Tracheoniscus tradouwi, Bethalus . translucida, Merulana translucida gracilior, Merulana . translucidus. Armadillo transmutatus, Porcellio transsilvaticus, Cylisticus . transvaalensis, Polyacanthus travancoria, Nagara . travancorius, Nagurus Trematocranus triangulifera, Uramba triarticulatus, Periscyphis . triarticulatus, Periscyphops tricirratus, Euplotes affinis trifolium, Armadillo. trilobatus, Porcellio . Trinema. .... triplicata, Campylaspis trisulcatus, Euplotes trivialis, Periscyphis trochilirostris, Campylorhamphus truncorum, Armadillo truncorum, Venezillo . 212 148, 149 . 212 223 . 242 250, 255, 295-296 203 191, 205-207, 239-240 . 207 . 207 . 222 . 228 . 228 . 228 203 . 192 . 229 205 205 . 160 205 . 218 . 218 113, 119, 123 . 221 2O3 . 103 250, 253 . IOI . 217 . 42 232 232 tuberculatus, Mesarmadillo tubulata, Campylaspis tuffraui, Euplotes tugelae, Diploexochus tunetanum, Armadillidium 215 250, 252 101, 124 . 227 . 212 umbensis, Campylaspis . . . 250, 253 umbricus, Cylisticus plumbeus . . .192 undata, Campylaspis .250, 253, 283, 287, 289 undulata, Periscyphis . . . .217 uniformis, Sunniva . . . . .213 uniplicata, Campylaspis . . . 250, 252 unisulcata, Campylaspis . . 250, 252, 263 Upupa . 37, 42 Uramba ..... 205, 239 uraniponnica, Finmarchinella . . 83, 86 valleculata, Campylaspis. 250, 254, 282-284 vallombrosae, Armadillidium . . .212 vanderhorsti, Haplochromis . . .162 vannus, Euplotes . . . . . 101 variabilis, Euplotes . . . . .124 variabilis, Porcellionides .... 204 velifer, Haplochromis . . . -143 venetus, Protracheoniscus . . . 207 Venezillo ...... 232 venustus, Armadillo . . . . .232 venustus, Venezillo .... 232 verecundus, Eptesicus . . . -134 verecundus, Philetar brachypterus 134-135, T 37 verrucosa, Campylaspis . . 250, 255, 297 verrucosus, Armadillo . . . .227 verrucosus, Diploexochus . . . .227 versicolor, Armadillidium . . . .212 Vespertilio . . . . 134, 172, 174 vespertilio, Porcellio .... 203 Vespertilionidae 73-82, 167-188, 317, 318, 319, 320, 322, 324, 325, 326, 327, 328, 329, 330 violaceus, Porcellio .... 198, 204 virgatus, Metoponorthus .... 197 viridis, Metoponorthus .... 197 viticola, Armadillo . . . . .221 vitrea, Campylaspis . . 250, 252, 263, 285 vittatus, Periscyphis . . . .217 vizzavonensis, Porcellio lugubris . . 201 vizzavonensis, Porcellio orarum . .201 vulcani, Tilapia .... 148, 149 vulgare, Armadillidium . . 210,212,221 vulgaris, Sturnus . . . . -37 wagleri, Psarocolius . weberi, Armadillo weberi, Spherillo West, B. J. . Whale-lice White-beaked dolphin 37 . 231 231 95-102, i PI. 59-64, i PI. ; 65-72 59-64, i PI. INDEX wilsmorei, Armadillo wilsmorei, Cubaris . wingatii, Haplochromis wintoni, Laephotis . wintoni, Laephotis wintoni wintoni angolensis, Laephotis wintoni wintoni, Laephotis woodi, Scotoecus woodi, Scotoecus albofuscus Xestadillo . 221 . 221 162, 163 73. 74-75, 76, 77, 78, 79, 80, 81 73 73 73 176, 182, 183 175, 176-177 230 yemenensis, Porcellio zachvatkini, Hemilepistus zealandicus, Porcellio zebricolor, Armadillo zebricolor, Cubaris . zenkewitchi, Euplotes zigzag, Armadillo zigzag, Diploexochus zwartbergensis, Diploexochus 349 204 194 204 221 221 I2 4 227 227 227 17 ANATOMY OF HEAD AND NECK IN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITH COMPARATIVE NOTES ON OTHER CALLAEIDAE P. J. K. BURTON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. i LONDON: 1974 FRONTISPIECE. Male and female Huias (Heteralocha acutirostris) showing probable feeding methods. The male is depicted excavating decayed wood by the 'gaping' technique described in the text. 17 JULI9I ANATOMY OF HEAD AND NECK IN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITH COMPARATIVE NOTES ON OTHER CALLAEIDAE BY PHILIP JOHN KENNEDY BURTON Pp 1-48 ; i Plate, 26 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. I LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 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. 27, No. i, 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.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 13 June, 1974 Price 2-30 ANATOMY OF HEAD AND NECK IN THE HUIA (HETERALOCHA ACUTIROSTRIS) WITH COMPARATIVE NOTES ON OTHER CALLAEIDAE By PHILIP J. K. BURTON CONTENTS Page I. INTRODUCTION ......... 3 II. METHODS AND MATERIAL ....... 5 III. SKULL AND LIGAMENTS ........ 6 IV. JAW MUSCULATURE ........ l6 V. BUCCAL CAVITY AND GLANDS ....... 24 VI. TONGUE APPARATUS ........ 24 VII. NECK AND NECK MUSCULATURE ...... 28 VIII. OTHER CALLAEIDAE ........ 34 IX. FUNCTIONAL ASPECTS ........ 36 X. SEXUAL DIMORPHISM ........ 40 XI. CONCLUDING REMARKS ........ 44 XII. ACKNOWLEDGEMENTS ........ 46 XIII. REFERENCES ......... 46 SYNOPSIS Spirit specimens and skeletons of the Huia (Heteralocha acutirostris) have been used in a detailed study of feeding adaptations in this extinct species. The Huia showed remarkable sexual dimorphism in bill form, and particular attention is paid throughout to differences between male and female. Comparison is made with the other members of the Callaeidae - Creadion carunculatus and Callaeas cinerea. The features studied indicate that the Huia was highly specialized for feeding by 'gaping' in decayed timber, in order to gain access to beetle grubs. The full extent of sexual dimorphism revealed by this study is reviewed, and its sig- nificance is discussed. I. INTRODUCTION THE extinct Huia (Heteralocha acutirostris) of New Zealand is frequently quoted as an example of extreme sexual dimorphism among birds. This dimorphism is shown most conspicuously by the bill, which in the male was typically fairly straight, moderately long, and tapered evenly to a point ; while in the female it was much longer, more slender, and strongly downcurved. There is little information on the extent of dimorphism beyond this. Phillipps (1963) mentions plumage differences and a tendency towards stronger claws in older males, and Selander (1966) gives bill, wing and tarsus measurements for a small sample of each sex. The anatomical study of the Huia by Garrod (1872) concentrated principally on establishing the affinities of the species, and did not inquire closely into its sexual dimorphism. The problem of its relationships is still not fully resolved beyond the fact that the Huia is closely related to two other New Zealand species -the Saddleback (Creadion 4 P. J. K. BURTON camnculatus) and the Kokako (Callaeas cinerea) (Stonor, 1942). The three together form the family Callaeidae, placed between the Dicruridae and Grallinidae by Mayr and Green way (1956) ; close affinity with the Sturnidae has been suggested in the past (Garrod, 1872 ; Gray, 1870). The present investigation has been concerned with the anatomical basis of the Huia's feeding specializations. In particular, I have attempted to discover the full extent of sexual dimorphism in feeding structures, and to correlate this where possible with bill form and feeding methods. It is thus a study of adaptation, unusual in that the principal forms to be compared are the two sexes of a single species. The material on which the investigation is based is contained in the avian anatomical collections of the British Museum (Natural History). These consist of six spirit and one osteological specimens of the Huia, one spirit specimen of the Saddleback and four spirit specimens of the Kokako. It is thought that the Huia spirit specimens may be the only fluid-preserved examples of this species in existence, although there is at least one skeleton in New Zealand (Phillipps, 1963) and another in the U.S.A. (Bock, igGoa.). The history of the Huia's disappearance, and many aspects of its life and habits, are summarized by Phillipps. Last reported alive in 1907, it is probable that some survived after this date, and Phillipps entertains the possibility that a few may still exist. Its range was restricted to the mountainous Wellington Province of North Island, New Zealand. Here, the combination of a mild climate and heavy rainfall has produced densely forested country with a high incidence of timber decay and epiphytic growth. The Huia fed principally on insects, varied with some vegetable matter, but by far the most important prey appears to have been the larvae of the Huhu beetle, Prionoplus reticularis (Cerambycidae) . This large beetle is common in many parts of New Zealand, and its larvae, found in decaying timber, may reach a length of 70 mm and a width of 20 mm. The most detailed account of the Huia's manner of extracting these is provided by Buller (1888). This account has been often quoted, but because of its relevance to the present study, it seems essential to do so again. Buller's description refers to a pair of captive Huias which at first he fed on individual huhu grubs : 'On offering one of these to the Huia he would seize it in the middle, and, at once transferring it to his perch and placing one foot firmly upon it, he would tear off the hard parts, and then throwing the grub upwards to secure it lengthwise in his bill, would swallow it whole . . . They seemed never to tire of probing and chiselling with their beaks. Having discovered that the canvas lining of the room was pervious, they were incessantly piercing it, and tearing off large strips of paper, till, in the course of a few days, the walls were completely defaced. But what interested me most of all was the manner in which the birds assisted each other in their search for food, because it appeared to explain the use, in the economy of nature, of the differently formed bills in the two sexes. To divert the birds, I introduced a log of decayed wood infested with the huhu grub. They at once attacked it, carefully probing the softer parts with their bills, and then vigorously assailing them, scooping out the decayed wood till the larva or pupa was visible, when it was carefully drawn from its cell, treated in the way described above, and then swallowed. The very different development of the mandibles in the two sexes enabled them to perform separate offices. The male always attacked the more decayed portions of the wood, chiselling out his prey after the manner of some Woodpeckers, while the female probed with her long pliant bill the other cells, where hardness of the surrounding parts resisted the ANATOMY OF HEAD AND NECK IN THE HUIA 5 chisel of her mate. Sometimes I observed the male remove the decayed portion without being able to reach the grub, when the female would at once come to his aid, and accomplish with her long slender bill what he had failed to do. I noticed, however, that the female always appro- priated to her own use the morsels thus obtained.' An additional reference to the different feeding techniques of male and female is the note by J. M. Wright quoted by Oliver (1955): 'I have watched them in pairs hunting for wetas.* The male would tear away at the outer part of a green sapling. The female then tried to retrieve the weta with her long slender bill. If not successful she would stand back while the male tried to enlarge the hole. Unless the tree was a maire the birds would generally succeed and would then fly away with the weta. 1 Phillipps (1963) also provides the most detailed information available on the extent of dimorphism ascertainable from skins, and on the range of variation in the two sexes, based on 119 specimens in New Zealand museums. Young birds show little difference in size and shape of the bill. Some male bills were more curved than usual, but were normally deeper than those of females. The male bills reached an extreme length of 59-60 mm (from feathers), and females up to 104 mm. Phillipps mentions two doubtful cases of birds with bills 63 mm long which he has treated as males, though they may be females. The skin collection of the British Museum (Natural History) includes two doubtful individuals ; one, with a bill length of 76-4 mm is labelled as a male ; the other, labelled as a female, has a bill of 71-8 mm. Excluding these, bill length data for other specimens in the collection may be summarized as follows : Males : Mean 59-7 ; Min. 54-7 ; Max. 65-0 (16 specimens). Females : Mean 96-3 ; Min. 87-1 ; Max. 108-0 (7 specimens). Buller (1878) also gives details on the range of bill form in the Huia and figures several specimens, including a female with the upper jaw much longer than the lower, recalling some species of Hemignathus (Drepanididae) . II. METHODS AND MATERIAL Most of the observations reported here were obtained simply by dissection using a stereomicroscope at powers mainly in the range X2| to X20. The main com- plicating factor was the irreplaceable nature of the Huia specimens, and because of this, it was thought prudent to keep dissections of this species to a minimum. In general, dissections have been limited to the left side, and as little structure as possible has been removed. Because of its history, the single specimen of Creadion carunculatus was regarded as of similar value, and no dissections were attempted which would have required removing or severing structures. For convenient reference, the Huia specimens were given a simple individual coding in addition to their British Museum (Natural History) Registration numbers. * Orthoptera, Gryllacridoidea. 6 P. J. K. BURTON Details of the specimens of Callaeidae used, with their conditions at commencement of the study, are as follows : Heteralocha acutirostris $ i. Reg. No. A 1973.1.3. Intact spirit specimen. o N O P H y> b Jo s , wt?773 (M , w a b >o !t c- p 'ro M -^ p ^o" &ot it t j'8 1 1 ro ro l l Tj- 10 cb ^ UOlpVUQ N 2- J^ ^2- SU9UII09ds JO UB9J\[ O b 00 o in ^t- v ^N (N 'fO >O M V1{OOlV49ia H ^3 ^~ >ti o o IH ^^ ^ro H ^ X T3 ! SU9UIID9ds JO tTE9J^ b iO ^ yi 00 ro oo" O Tf b oo 04 ro IH M co" co b 9 M "t^ P ^ 00 M ^ "0? ^ 01 IN {H O o ot Q\ P b OI f^ ro JM o M VilOOJV^SJSpl 00 ^ro "^ ^ O M ^^ ^ro M ^. O 1 P! ^oo 'b" iO "ro 1) a. O 1 1 1 ob N 'J- ro ro i *? 5 miooivwH M ^ Jo IH - 1 ~ "ro J*" - - M 1H OI p 'ro 13 V O CO r^ oo op i l b ^ i 1 u vqooivwfdfi Ok Jo t- M 00 Jo H ^ H "o w t/5 P3 i 7 o O^ 'cT' ro SET 0) o" M s ^ 9Mj9 f, 1 00 ? i l ob op 'o u-) 1 H 'o o ^^ - ' ' 8- c p ro ^ o 0* o\ ^ "r^ r 5 SET ^ 'ro Q o * b OO O o O f^ M u^ vO 01 S mpojVMtaH HI T *" *. O 00 ^rOJ9?3 ff M -? ^1 o ^ ro f> M o 6 * 1 " 1 ^ 1 C* 03 c 4 fe o 1 o o ., fl ex ig r/; M .^ o o en H B "Pn "o ft g tr C3 ! g i E CH 73 a re oi C O 1) IH S ^ "c ^ fc "2 0) i ^ ^ ^ 3 o o o IH 03 IH O CM & P removed Index of O rtj t? | 9 (U o removed Retroarti Retroarti relative t 0) ^! CO ^B u Q W 0_ ST ANATOMY OF HEAD AND NECK IN THE HUIA 15 00 CO oo* f t^ fO 1o OO* vO oo* 00 o ^ 'o^ ^ f^ roiOON^ONONV^HVpbPl I CO V~> I I I t^ I f> , M WQfOM "* b\ N CO irj^bvO N p M ^lO "~> M vO u-) P 1 I t^TfOO " Mb Hl b f M^b M b^b t>o ^' H b IH b N b ro o "> u-> vo g p't^p x ''5 : 7*"'o^i o oo"r*"'oNi o '?rc oo c r> '?rr*"''^ ?>""> p o ^y^vo7^ t ^ 1 ^o^ l or*"^r^^9t^^c^'^"l cb? 1 ! Nvot^ u b O N M^M b M b MhH b b M O ONt^ro^n vpf^Orort-roiopMbNr^oboOHro 2 ^"^ 4) 3 4-> en rt en ON(O^ ^'^OPN'P P " J ^f- 4^- |H VN f^ \*J H ^ ^Q UV Q*y y> Q W1 \} -I ^j- vy^l CN ^-> vO'pb T *"r r )VOcb M H^I't^'pioOiH^f| ^'NiO^bNO'^iO '2 & C IH o M b r *~M cv 'M M b'~'o^'i-i' H o coM^ooNOOt^- .S^ ^^^^,^^_^ ^ N N'5~'cp'^p'Q^r^'^t^o\t^'o N 7i-co? < 'M So p'?p M p CL.V I b^'wvpcxjpM^I-f^vpioptNTf'co^M^vb^l t^POOO cn-g Hl b T) "H^M H O w O' N M IH O M b rr) M< r >O ONONl> ^-5 |"o poo v op'!^^o'p1o?-'o^^'o'V 'f^t H 'f^^ r 5 : P w P i ^^ l oj r ^N > poopc > <^"oovpvopN^" i ^-y^bi H f^? SN cbtNNO M b T *"H^M M b'~'b N w iH b l ~ l b fr) M rr >oooNONt^ -Stn ^^, ^ x . ^"^ ^~^ ^, ^ *?r, *~- ^~' f M t-i M -*^ eu tuO VH N'^M^pN'o'pN'>-ry~ > o6"~''5" vi - > 'o~" v P'o~' jo fo 7f ^^ I N?tinvpoN M roy~>obvpr^pr' ) iOroy>| t^.^1 | IN 13-I- 1 MQ-^- M tN M M O l-tQN^M O M O fOO 00 Wlti O CD b\t^M^-Kf^MWroTi-bN9f^Pl I MMcbt^f^^ib -2-g MOiH O -J yy v^ Q[J ^ w 1 s -. ^ . _ M M m'^"covOi^Ofn T i"i ON < 2o' s Pf^u-)N< i o ^"'s; rO M M O M O iN M M O NM"^O>OONCOOO ^_ft 42 'g ^ O _O(y,Op^-NQipQfO o o >O 1o ^) CO 00 rO o -4-> _____ _ O.CM ^ i (-1 S a rt "2 "o3 p 5 S g bo to jd ^ .2 1> m H 3 be C > L CU -l-> bo a o 11 e~ '3 c3 u < 6 i) a ft H C/) $ O *Z H H pq ^^K^^^^^QCL Oi^ ft^ 16 P. J. K. BURTON sesamoids, one placed laterally and the other posterior to the quadrate. In common with all passerines so far studied, the lower jaw has no medial brace (see Bock, I96ob). IV. JAW MUSCULATURE A work of major importance for studies of avian jaw musculature is Lakjer's (1926) review of the trigeminal musculature. For passerines in particular, the paper by Fiedler (1951) provides valuable clarification. Investigations by Bock (i96oa and MS) have resolved many remaining problems of structure and ter- minology within the order, and his nomenclature for the subdivision of complex muscles is the one followed here. The jaw musculature of the Callaeidae remains undescribed beyond the comments of Garrod (1872) and Lowe (1938) on the large size of the 'digastric muscle' (= M. depressor mandibulae). In describing the structure of complex muscles it is often useful to designate their various aponeuroses by letters or numbers. It is far from clear to what extent these aponeuroses can be homologized between different groups, although the work of Starck and Barnikol (1954) has made progress in this respect for M. adductor mandibulae externus. In the present case, a simple numbering system has been used where necessary, but is not intended to imply homology with any numbered apo- neuroses described from other species, with the exception of Aps. i, 2 and 3 in M. adductor mandibulae externus. M. adductor mandibulae externus This important and complex muscle acts to raise the lower jaw and to maintain a grip on objects held in the bill. Its architecture is intricate, and the relationships of its aponeuroses can be better understood by the examination of cross sections (Fig. 18). The abbreviation 'M.a.m.e.' is used in parts of the description to shorten the otherwise unwieldy names of some subdivisions. The three major aponeuroses described by Starck and Barnikol (1954) in a wide range of birds of several orders can be recognized with little difficulty in Heteralocha, and their numbering here (Aps. i, 2 and 3) is the same as Starck and Barnikol's. M . adductor mandibulae externus rostralis The most dorsal part of M. adductor mandibulae externus, with an extensive fleshy origin on the cranium. Insertion is made principally via Ap. i, which nar- rows anteriorly to form a flat tendon attached to a crista on the surangular. Follow- ing Bock, three sections are recognized : M.a.m.e. rostralis medialis. Origin is from the postero-lateral border of the orbit, medial to the postorbital process, and from a vertically oriented aponeurosis (Ap. 4) attached to the orbit along the medial border of the muscle. Insertion is made via Ap. i. Medially, a group of fibres from the ventral part of the muscle overlaps part of Ap. i as a more or less distinct slip. M.a.m.e. rostralis temporalis. Origin is fleshy from the border of the temporal fossa ; a short aponeurosis (Ap. 5) runs forwards from the tip of the postorbital ANATOMY OF HEAD AND NECK IN THE HUIA Mdt Mdm L po FIG. ii. Heteralocha acutirostris, male (<$ i), jaw muscles and ligaments. For abbreviations see p. 47. process across the dorsal lateral surface, and in some specimens extends onto the dorsal surface of M.a.m.e. rostralis medialis. The fibres of this section converge anteriorly, falling into an ill-defined bipinnate arrangement, and are attached to the ventro-lateral surface of Ap. i ; a few fibres overlap its dorsal surface posteriorly, but it is exposed for some 8 mm from the insertion. M.a.m.e. rostralis lateralis. This section lies superficial to the dorsal part of M.a.m.e. ventralis. Its origin is from the lateral edge of the zygomatic process, and from the lateral edge of Ap. 2 in its narrow posterior region. Fibres run upward and medially from the origin and insert on the lateral surface of the mandible over a narrow zone dorsal to the insertion of M.a.m.e. ventralis. This insertion is fleshy, and via a superficial aponeurosis (Ap. 6) which merges medially with Ap. i. The anterior ventral part of this section lies medial to the jugal bar and is in consequence flattened into a relatively thin sheet of fibres. M. adductor mandibulae externus ventralis Origin is from the zygomatic process, by a strong aponeurosis (Ap. 2) which traverses the gap between the cranium and mandible as a fairly narrow tendon, then fans out over the lateral surface of the muscle. Fibres arising from this aponeurosis make an extensive fleshy insertion on the lateral surface of the mandible, spanning the fenestra in the posterior part of the mandible. M. adductor mandibulae externus caudalis The shortest section of M. adductor mandibulae externus. Origin is from the otic process of the quadrate and from the ventral surface of the posterior half of Ap. 2. The origin is fleshy, and by a weak aponeurosis (Ap. 7). Insertion is on the dorsal lateral edge of the surangular, posterior and ventral to that of Ap. i. The insertion is made by a short, narrow aponeurosis (Ap. 3), which bifurcates within i8 P. J. K. BURTON the body of the muscle to form a dorsal (Ap. 3a) and a ventral (Ap. 3b) branch. Ap. 7 passes between 3a and 3b before fading out. The muscle shows multipinnate structure, clearest in its middle third, and most conspicuous about Ap. 3a. Fibres run forwards from Ap. 2 to Ap. 3a, back from Ap. 3a to Ap. 7, and forwards again from Ap. 7 to Ap. 3b. M. pseudotemporalis superficialis A muscle of relatively simple structure, whose action is adduction of the lower jaw. The origin is a broad fleshy one on the posterior wall of the orbit, immediately medial to M.a.m.e. rostralis medialis. There is also a weak dorsal aponeurosis attached to the orbit. Insertion is made via a strong tendon attached to the medial side of the surangular, ventral to the ramus mandibularis of the trigeminal nerve. Mdt Md m L po FIG. 12. Heteralocha acutirostris, female ($ C), jaw muscles and ligaments. For abbreviations see p. 47. M. pseudotemporalis profundus This muscle is an adductor of the lower jaw, and by its action on the quadrate it also functions to lower the upper jaw. It takes its origin from the dorso-lateral surface of the orbital process of the quadrate, and from a strong aponeurosis attached to the expanded tip of the orbital process. It makes an extensive fleshy insertion on the medial surface of the mandible, overlapping slightly onto its dorsal edge, adjacent to Ap. i of M.a.m.e. rostralis. M. adductor mandibulae posterior The actions of this muscle are similar to those of M. pseudotemporalis profundus. Due to its situation closer to the mandible-quadrate articulation, it has a lower mechanical advantage, but its upward component may be of value in holding the mandible against the quadrate. M. adductor posterior is contiguous with M. ANATOMY OF HEAD AND NECK IN THE HUIA Mptvm Mptdm Mpsp Mpq1 Mamerm Mamert M a m ev M amec FIG. 13. Heteralocha acutirostris (<$ i), jaw muscles in lateral view. For abbreviations see p. 47. pseudotemporalis profundus at its origin, which is also a fleshy one on the quadrate, occupying the base of the orbital process and the dorsal surface of the quadrate body. The two muscles are separated (following the criterion used by Lakjer, 1926) by N. pterygoideus. The muscle inserts on the expanded region of the dorsal surface of the mandible just anterior to the base of the internal process, and posterior to the insertion of M.a.m.e. caudalis. The insertion is fleshy and via a weak dorsal apo- neurosis. M. pterygoideus This important muscle complex acts both to lower the upper jaw and to raise the lower. Its attachments to the palatine and pterygoid are arbitrarily regarded as the origin, and attachment to the lower jaw as the insertion. M. pterygoideus dorsalis later alis This section has an extensive fleshy origin on the dorsal surface of the palatine. It inserts on the medial surface of the mandible, posterior to M. pseudotemporalis profundus, fleshily, and by a superficial aponeurosis (Ap. i), which is strongest ventrally. M. pterygoideus ventralis lateralis Origin is from an aponeurosis (Ap. 2) which is attached to the postero-lateral tip of the palatine, and extends for some distance across the ventral surface of the muscle. The medial edge of this aponeurosis serves to demarcate it from M. pterygoideus 20 P. J. K. BURTON ventralis medialis seen in ventral view. M. pterygoideus ventralis lateralis is at- tached at the base of the internal process of the mandible, ventral to the insertion of M. pterygoideus dorsalis lateralis. There is no overlap (Venter externus') onto the ventral edge or lateral surface of the mandible as in many birds, since the ventral medial edge of the mandible in this region is occupied by a forward extension of M. depressor mandibulae. M pt v m M pt d m a M pt d m p Mpt r Mptdl Aplame Mpsp Mamerm Mamert 5mm FIG. 14. Heteralocha acutirostris ( i), jaw muscles in dorsal view. For abbreviations see p. 47. M. pterygoideus dorsalis medialis In dorsal view, this muscle is seen to lie immediately adjacent and posterior to M. pterygoideus dorsalis lateralis, separated from it by a narrow groove which widens medially, exposing a small area of the dorsal surface of M. pterygoideus ventralis medialis. Its origin is confined to the pterygoid, and occupies much of its surface. The muscle falls into anterior and posterior portions, whose fibres diverge in a posterior direction roughly along the lines of the pterygoid, so that it appears bi- pinnate in dorsal view. M. pterygoideus dorsalis medialis anterior originates more laterally on the ptery- goid, and inserts narrowly on the mandible, immediately caudal to M. pterygoideus dorsalis lateralis. The insertion is fleshy and aponeurotic, the aponeurosis (Ap. 3) being strongest ventrally. M. pterygoideus dorsalis medialis posterior originates medially on the pterygoid, and fans out to a wide fleshy insertion on the dorsal part of the internal process of the mandible, dorsal to that of M. pterygoideus ventralis medialis. ANATOMY OF HEAD AND NECK IN THE HUIA M pt v m Eus Mpt r Ap 3a d m M d m 5mm FIG. 15. Heteralocha acutirostris (<$ i). M. pterygoideus and M. depressor in ventral view. Left side of palate left intact to show papillae. For abbreviations see p. 47. M . pterygoideus ventralis medialis There is an extensive fleshy origin on the ventral surface of the palatine and insertion is made on the internal process of the mandible, fleshily, and by a strong dorsal aponeurosis (Ap. 4). The fibres of this section are orientated more nearly parallel to the skull axis than those of any other part of M. pterygoideus. On the medial side of the muscle is a slip (M. pterygoideus retractor) exclusively specialized for retracting the palatal apparatus, attached at one end to the medial caudal tip of the palatine, and at the other to the basitemporal plate of the skull. A few of its most medial fibres are attached to the edges of the opening of the Eustachian tube. M. protractor quadrati et pterygoidei This muscle acts to raise the upper jaw by moving the palatal framework and jugals forwards. It has a wide fleshy origin from the posterior part of the interorbital septum and the adjacent region of the posterior orbital wall. The lateral border of the muscle lies immediately adjacent to M. pseudotemporalis superficialis. 22 P. J- K. BURTON Two parts can be distinguished. The more anterior and medial part (M. protractor i), arising mainly on the interorbital septum, shows a bipinnate fibre arrangement. Its raphe is a strong aponeurosis attached to a spur on the posterior dorsal tip of the pterygoid, immediately adjacent to the quadrate. This raphe serves as the principal site of insertion for the fibres of M. protractor i. The lateral part (M. protractor 2) originating from the posterior wall of the orbit inserts fleshily, and by a weak dorsal aponeurosis on the caudal edge of the basal half of the orbital process. Mptdl MptdmaMpss Mpql Mamerm M pt d m p / Ma mert M d m Mamev Ap2ame Mamp Mamec 5mm FIG. 16. Callaeas cinerea, jaw muscles in lateral view. For abbreviations see p. 47. M. depressor mandibulae This muscle, which depresses the lower jaw, is of enormous bulk (and, presumably, fibre number) in Heteralocha relative to the other jaw muscles. There is a wide fleshy origin on the squamosal, parietal and exoccipital. Fibres originating high on the cranium converge to insert fleshily near the extremity of the long retroarticular process, mainly on its medial side. Those originating lower, on the exoccipital, fan out to insert fleshily on both surfaces of the retroarticular process. Fibres inserting medially extend far forward beyond the retroarticular process along the ventral half of the mandible, ventral to the insertion of M. pseudotemporalis profundus. This extension is here termed the pars anterior of M. depressor mandibulae. ANATOMY OF HEAD AND NECK IN THE HUIA Mptdl Mptdma Mptdmp Mpq1 Mpss M a merm M a m e v M a m e rt Mdm 5mm FIG. 17. Callaeas cinerea, jaw muscles in dorsal view. For abbreviations see p. 47. M. pseudotemporalis profundus is concealed by M. pseudotemporalis superficialis. 2mm FIG. 18. M. adductor mandibulae externus, TS at about the midpoint of the otic process of the quadrate, and at right angles to it. Numbering of aponeuroses as explained in text. A - Callaeas cinerea. B - Heteralocha acutirostris (<$ i). The left side is lateral in both diagrams. The following aponeuroses can be distinguished : Ap. i. This is attached to the cranium around the dorsal edge of the origin on the squamosal and parietal, and extends some way across the lateral surface of the muscle. Ap. 2. A strong aponeurosis with a narrow attachment at about the middle of the posterior border of the auditory meatus, fanning out across the lateral surface of the muscle, and acting as a surface of origin for fibres inserting on the anterior half of the lateral surface of the retroarticular process. 24 P. J- K. BURTON Ap. 3. A strong aponeurosis attached to the occipital crest, which marks the postero- ventral limit of the origin. Ap. 3 extends some way across the medial surface of the muscle, and becomes stronger on the exoccipital process, whence it is produced as a strong raphe (Ap. 3a) across the ventral region of the muscle, and on into the pars anterior. Fibres arising from its medial side insert mainly on the medial surface of the internal process of the mandible. Those running from its lateral side insert on the medial surface of the basal half of the long retroarticular process, on the posterior face of the internal process at its base, and on the ventral medial surface of the mandible. Ap. 4. An aponeurosis attached to the dorsal edge of the internal process of the mandible, extending across much of the dorsal surface of the short part of the muscle between this and the exoccipital. This aponeurosis is branched and infolded within the muscle. Ap. 5. An aponeurosis attached to the dorsal edge of the retroarticular process, but concealed from lateral view by the thin sheet of fibres originating from Ap. 2, and inserted on the lateral surface of the process. V. BUCCAL CAVITY AND GLANDS Backwardly directed horny papillae are abundant on the surface of the palate. They are longest where they fringe the choanae, and in a densely clustered transverse row posterior to the internal opening of the Eustachian tubes. The surface of the larynx is also papillate, the papillae being longest at its posterior margin. Three pairs of salivary glands are present. The Gl. angularis oris lies on the side of the head immediately below the skin and just ventral to the jugal bar ; its duct opens at the angle of the gape. A second gland, which may be termed a Gl. palatinae (see Antony, 1920), lies between M. depressor mandibulae, pars anterior, and M. pterygoideus ventralis lateralis, immediately below the mucosa of the palate. The third gland is a sublingual one. It lies immediately lateral to M. genioglossus, and extends from the level of the anterior border of M. mylohyoideus to the region of the tongue base. Its duct runs alongside M. genioglossus, but diverges from it near the mandibular symphysis to open in a lateral position. VI. TONGUE APPARATUS The tongue is narrow, tapering evenly towards its anterior tip which is somewhat frayed and brush like (Fig. 19). The lateral edge bears a few papillae posteriorly, and the posterior edge is crowded with pointed papillae, longest laterally. Tongue length for the specimens examined is shown in Table i. The tongue skeleton is similar to that of many passerines. The paraglossalia which provide support for the tongue itself have long posterior processes, and anteriorly they meet and run side by side to a point about one-third of the way from the tip of the tongue. The remainder of the tongue is purely corneous. The basihyal ANATOMY OF HEAD AND NECK IN THE HUIA 5mm FIG. 19. Heteralocha acutirostris (<$ i), tongue and larynx in dorsal view. is a slender rod of roughly triangular section, with the apex of the triangle upper- most. The flattened urohyal, expanded at its posterior tip, and the hyoid horns (each consisting of the ceratobranchial posteriorly and epibranchial distally) are of similar form to those of many passerines. The tongue muscles in Heteralocha show few unusual features, and no significant sexual dimorphism was encountered in the dissections. The descriptions given are therefore brief ones ; illustrations are given in Figs. 21 and 22. More detailed accounts and background information are given in the general review by George and Berger (1966) and the paper on passerine tongue muscles by Engels (1938). A very full description of the tongue muscles of a single passerine species is that by Bock (1972) for the extinct Ciridops anna (Drepanididae) . For discussions on function, reference may also be made to the account of wader tongue muscles by Burton (I974)- M. mylohyoideus A thin muscular sheet with a long narrow origin on the medial side of the man- dibular ramus, inserting on a median raphe. M. mylohyoideus lies ventral to the tongue and all the muscles attached to it. M . serpihyoideus Origin is on the occipital plate, medial to the exoccipital process (Bock, igGob, p. 38) and insertion is on a median raphe continuous with that of M. mylohyoideus. 26 P. J. K. BURTON M . genioglossus A narrow, strap-like muscle whose origin is from the posterior edge of the mandi- bular symphysis. The left and right muscles lie side by side near the origin and then diverge, running along the ventral side of the mucosa of the floor of the buccal cavity, on either side of the position of the tongue. In the region of the basihyal, the fibres of the muscle fan out to insert on the connective tissue and mucosa over- lying the basihyal and its musculature, and, anteriorly, on the posterior process of the paraglossa. FIG. 20. Callaeas cinerea, tongue and larynx in dorsal view. M. stylohyoideus A long, narrow muscle originating on the ventral edge of the base of the retro- articular process, and running along the anterior edge of M. serpihyoideus, to insert on the lateral surface of the basihyal, just anterior to M. thyreohyoideus. M. branchiomandibularis (= M. geniohyoideus of many authors) A bulky muscle whose origin is on the medial surface of the mandible, ventral to that of M. mylohyoideus. A broad anterior and narrow posterior position can be distinguished. The two run parallel to insert on the hyoid horn. The anterior portion meets the horn from the ventral side, and is twisted around it for some dis- tance before inserting on the epibranchial. The posterior portion meets the anterior on the dorso-medial side, and merges with it. M. ceratohyoideus A thin, weakly developed muscle which was found only in <$ i. It originates on the hyoid horn, on the ventral lateral surface of the distal tip of the ceratobranchial, ANATOMY OF HEAD AND NECK IN THE HUIA Apm Mthh Mseh FIG. 21. Heteralocha acutirostris ($ B), tongue muscles in ventral view. For abbreviations see p. 47. deep to M. ceratoglossus. The insertion is on a median raphe continuous with that of M. serpihyoideus and M. mylohyoideus, near the posterior end of the urohyal. M. ceratoglossus A unipinnate muscle whose fibres arise on the surface of the ceratobranchial, and the anterior end of the epibranchial, and insert on a long lateral tendon which is itself inserted on the ventral surface of the paraglossal, just level with the anterior tip of the basihyal. No fibres insert on the tendon over the region lying alongside the basihyal, but just anterior to its attachment to the paraglossal, a small fleshy slip arises. It merges with that from the other side, and together they insert on a strong medial aponeurosis which runs along the ventral side of the paraglossalia, and inserts on the corneous anterior part of the tongue. This slip is referred to by Bock (1972 and MS) as M. hypoglossus anterior. A similar slip in shorebirds was described by Burton (1974) as M. ceratoglossus anterior ; the term 'M. hypoglossus anterior' is used by Burton (1974) to refer to a quite distinct muscle, apparently absent from 28 P. J. K. BURTON passerines, arising on the posterior tip of the paraglossals and also inserting on the median aponeurosis. M. hypoglossus obliquus Origin is on the postero-lateral process of the paraglossa. The right and left muscles are merged, the whole forming a bulky loop passing ventral to the anterior third of the basihyal. Mthh Mtrh M th h M tr I FIG. 22. Heteralocha acutirostris (? C), ventro-lateral view of tongue muscles attached to the cricoid cartilage. Cricoid and urohyal shown stippled. Both left and right Mm. tracheolateralis are visible. For abbreviations see p. 47. M. tracheohyoideus Origin is from the skin of the neck, and insertion on the lateral surface of the cricoid, just below the dorsal origin of M. thyreohyoideus. M. tracheolateralis The muscle originates on the syrinx, and passes along the side of the trachea, broadening anteriorly to insert by two heads on the lateral surface of the cricoid. M. thyreohyoideus This muscle originates from the lateral surface of the cricoid, by a dorsal and a ventral slip. The dorsal one is slender, and is attached just below the dorsal edge of the cricoid. The ventral slip is broader, and arises between the two heads of insertion of M. tracheolateralis. These slips unite anterior to the larynx and insert on the anterior lateral surface of the basihyal posterior to the insertion of the more slender M. stylohyoideus. M. ceratoglossus superior (Bock, 1972) was not found in Heteralocha. The glottal muscles (M. thyreoarytenoideus and M. constrictor glottidis) conform to Bock's description for Ciridops anna. VII. NECK AND NECK MUSCULATURE The thorough review by Boas (1929) has provided the basis for most subsequent studies of the avian neck and its musculature. Boas concentrated on non-passerines, principally large species, but detailed information on several small passerines is given by Palmgren (1949). There is, however, little information on the cervical ANATOMY OF HEAD AND NECK IN THE HU1A Mrcl M co M s co M sp Mil Mrcv Mfcb Mas 6-3 Mas 7-3 Mas 8-5 10mm FIG. 23. Heteralocha acutirostris ($ C), lateral view of superficial neck muscles. For abbreviations see p. 47. muscles of larger passerines comparable in size with the Callaeidae. The majority of neck muscles perform several actions which interact in a complex way, and their individual functions are omitted from the present account in the interest of brevity. Reference may be made to Boas and Palmgren, and to the excellent discussions on neck muscle function by Zusi (1962, 1969). There are 14 cervical vertebrae in Heteralocha, as in most passerines. This total includes the two cervico-dorsal vertebrae (13 and 14) which bear movable ribs not articulating with the sternum, but from a functional standpoint are best treated with the neck. The rib on 13 is extremely short, while that on 14 is long, almost reaching the sternum, but lacking an uncinate process. Six pairs of ribs actually articulate with the sternum, as in Cor cor ax, but unlike most passerines which only have 5 ribs attached to the sternum (Beddard, 1898). Boas showed that the neck of birds consists of three sections, distinguishable by both functional and morphological differences. Section I, the most anterior, can only be flexed downward, and Section II only upward. Section III can be flexed downward, and also upward at its anterior end. In Heteralocha, the constitution of the segments agrees with Palmgren's division for smaller passerines (although Palm- gren omitted the cervico-dorsal vertebrae from his count). The division is as follows : Section I, vertebrae i to 4 ; Section II, vertebrae 5 to 9 ; Section III, vertebrae 10 to 14. The vertebrae of Section I (except the atlas) have strong neural spines and hypapophyses. Those in Section II lack neural spines (except for a weak one on 5) and have no hypapophyses ; they are more elongated than the vertebrae of Sections I and III. The vertebrae of Section III have hypapophyses ; that on 10 is weak, but they increase in size posteriorly. The last, and largest, hypapophysis is on 15, the first dorsal vertebra. Vertebra 14 has a strong neural spine. The account below follows the same order and terminology as Palmgren, with modification in the case of Mm. splenii accessorii and Mm. intercristales. 30 P. J. K. BURTON M. biv enter This muscle arises from the dorsal surface of the aponeurosis of origin of M. spinalis in the region of 13, and inserts on the dorsomedial edge of the occipital deep to M. complexus. The muscle consists of two fleshy bellies linked by a flat tendon ex- tending approximately from 8 to 5. Little variation was encountered. M. spinalis The muscle consists of a series of fleshy slips from the ventral surface of an aponeurosis attached to the neural spines of vertebrae 14 to 18. These insert on the anapophyses of 2 and of 5 or 6 to 13 ; the most posterior of these is feeble and in- distinct in most of the specimens. Variations in the specimens of Heteralocha were as follows : Slips to both 5 and 6 present : $ A, $ B. Slip to 6 present, slip to 5 absent : ^ i, ^3, $ C. Slip to 5 present, slip to 6 absent : $2. Mm. splenii colli These muscles are a series of slips arising from the lateral surfaces of the neural spines of 3 or 4 successive vertebrae, and joining the most anterior slip of M. spinalis which inserts on 2. The vertebrae of origin in the Heteralocha specimens are : 4 to 7 : <$ i, $'s A, B and C. 4 to 6 : $ 2, ^ 3. Mm. splenii accessorii and Mm. inter cristales In order to clarify the relationships between these two rather similar groups of muscles it seems desirable to deal with them together. The muscles included under these headings in Heteralocha are as follows : a. A muscle running from the anterior surface of the neural spine of 14 to the transverse-oblique crest of 13, and a similar muscle connecting 13 and 12. b. A series of muscles connecting the transverse-oblique crests of successive verte- brae from 13-12 to 6-5. c. Muscles arising on the ventro-lateral surfaces of the neural spines of 5 to 3 and inserting on the transverse-oblique crests of 4 to 2. d. A muscle arising on the dorso-lateral surface of the neural spine of 3 and insert- ing on the anapophysis of 2, and a similar but weaker muscle arising on the neural spine and medial part of the neural arch of 2 and inserting on the anapophysis of i. That arising on 3 closely resembles Mm. splenii colli. e. A narrow, flat slip arising from the ventral part of the neural spine of 5 and inserting on the anapophysis of 3, somewhat resembling Mm. dorsales pygmaei. According to Palmgren's criteria, groups a and b only should be regarded as com- prising the Mm. intercristales, and groups c, d and e should be treated as Mm. splenii accessorii. (In the small species examined by Palmgren the two muscles running antero-laterally from the neural spine of 3 are inseparable.) However, it would seem more consistent with Boas's work to include group c also in Mm. intercristales, and to reserve the term Mm. splenii accessorii for groups d and e. ANATOMY OF HEAD AND NECK IN THE HUIA 31 No noteworthy variations between individuals were found. M. splenius capitis This muscle originates from the neural spine of 2 and inserts on the posterior surface of the skull deep to M. complexus and M. bi venter cervicis. No variations in siting were found, and there is little indication of cruciform structure (Burton, i97ia). 1 2 3 4 5 6 7 To 15 FIG. 24. Diagram to show arrangement of slips of some ventral muscles in the anterior part of the neck in Heteralocha acutirostris (? B). Heavy lines = M. flexor colli brevis. Broken lines = M. flexor colli profundus. Fine lines = anterior slips of M. longus colli ventralis. Mm. Pygmaei Origin is from the medial region of the neural arches of 12 to 8. Each muscle inserts on the lateral edges of the transverse-oblique crest of the second vertebra anterior to it except that from 12 which inserts on n. The latter muscle is weak in most of the specimens and absent in $ i. Mm. inter spinales These three muscles connect the well-developed neural spines of 2, 3, 4 and 5, and were present in all specimens examined. Mm. ascendentes cervicis These muscles arise on the diapophyses of cervical vertebrae up to and including 6. Most consist of two slips inserting on the anapophyses of the second and third vertebrae anterior. That from 7 consists of three slips inserting on 3, 4 and 5 in all specimens, and that from 8 sends an additional slip to 4 in ^'s i and 2. The short slips arising on 12 and posteriorly are relatively weak, and also showed some minor variations in siting. The series is continued posteriorly as Mm . ascendentes thoracicis, but those arising posterior to 14 were not dissected. Mm. intertransversarii These laterally situated muscles connect successive vertebrae. The most anterior are those from 3 to 2, and the most posterior those from 13 to 12. Each muscle arises from the anterior surface of the transverse process and inserts on the posterior surface of the transverse process of the vertebra in front ; that from 4 to 3 inserts on 32 P. J. K. BURTON the medial surface of the rib of 3, and that from 3 to 2 inserts on the lateral surface of the centrum of 2. As far as 6-5, the Mm. inclusi lie deep to Mm. intertransversarii, and closely associated with them. Palmgren regards the muscles anterior to this as continuing the series of Mm. inclusi ; Boas and Zusi treat them with Mm. intertransversarii as is done here. The Mm. intertransversarii are multipinnate muscles, traversed by interdigitating raphes from origin and insertion. The muscles are bulkiest and the number of raphes greatest in the region from 7 to 10. Up to 9 raphes have been detected in 9-8 or in 10-9, the number and situation of raphes showing small individual varia- tions. The muscle connecting 13 and 12 is reduced to a small dorsal slip, while those anterior to 5 are also of small size. Mm. inclusi These muscles are concealed by Mm. intertransversarii, and can only be separated from them with difficulty. Each one arises on the anterior surface of the transverse process medial to M. intertransversarius, and inserts on the lateral surfaces of the neural arch and centrum of the next vertebra in front. Most show division into dorsal and ventral bellies (Mm. inclusi superiores and inferiores). In the most posterior two (12 to n and n to 10), only inferiores can be distinguished. The most anterior are those connecting 5 and 6. M. longus colli ventralis This complex muscle consists of a series of fleshy slips arising on the sublateral processes, hypapophyses and anterior part of the centra of vertebrae 15 to 6. The main part of the muscle inserts by a series of 7 tendons on the ribs of n to 5. Each vertebra sends a slip to join each of the tendons traversing it ; there are thus 7 such slips from each of vertebrae 15 to 12 after which the number of slips decreases by one for each vertebra anterior to this. In the region 15 to 12 the slips are densely crowded and difficult to separate and some of the deepest fibres appear to attach to adjacent vertebrae. A smaller group of slips situated anteriorly is also included with M. longus colli. Three of these arise from a tendon attached to the sublateral process of 7, which also provides origin for part of M. flexor colli profundus. They insert on ribs 3 and 4 by short aponeuroses and on the long tendon attached to rib 5. Slips also arise on the ventral anterior surfaces of the lateral processes of 7, 6 and 5, the two former in close association with the Mm. intertransversarii, and immediately below them. These attach on rib 3, and those from 6 and 7 also on rib 4 ; minor variations occur. Two slips arise from the sublateral process of 6 and insert on ribs 3 and 4, and there is also a longer slip arising on 9 (8 in $ C) which inserts by a weak tendon on rib 4. This tendon also appears to receive some fibres from the posterior part of the muscle. M. flexor colli brevis Lateral and medial parts may be distinguished in this muscle. The lateral part constitutes the greater bulk of the muscle and originates from the ventral surface ANATOMY OF HEAD AND NECK IN THE HUIA 33 of the lateral strut of 3 ; and from the lateral processes of 4 and 5 in the $'s and 6 also in the g's. The medial portion is separated from the lateral by the anterior part of M. longus colli ; it takes origin from the sublateral processes of 3, 4 and 5. The lateral and medial portions join anterior to rib 3 and insert by a tendon on the postero-ventral processes of the centrum of the atlas. 10 12 13 14 15 FIG. 25. Diagram to show arrangements of slips of some neck muscles in Heteralocha acu- tirostris ($ A or $ B). Heavy lines = M. splenius capitis and Mm. splenii colli. Fine lines = M. spinalis (dorsal) and M. longus colli ventralis (ventral). Broken lines = Mm. splenii accessorii (as denned in text). Dotted lines = Mm. pygmaei. Tendons of M. spinalis and M. longus colli ventralis are represented by double lines. M. flexor colli profundus Origin is from the sublateral processes of 4, 5, 6 and 7. The slips from 4 and 5 arise immediately deep to the medial portion of M. flexor colli brevis and insert on the hypapophysis of 2. From 6, slips run to insert on the hypapophyses of 2, 3 and 4 (c? 2 > $ 3 $ B) or of 3 and 4 only ($ i, $ A, $ C). From 7 a long slip runs to the hypapophysis of 2 ; this slip shares a tendinous origin with the medial slip of M. longus colli from 7 to 3. In $ 2, $ 3 and $ B the slip bifurcates and a branch con- tiunes forwards to fuse with M. flexor colli brevis near its insertion. The slips arising on 6 show some fusion with M. rectus capitis ventralis at the origin. M. complexus Origin is from the lateral strut of 4, the diapophysis of 5, and from an aponeurosis attached to the diapophysis of 6. Insertion is on the dorsal edge of the occipitals. M. rectus capitis lateralis Origin is from the hypapophyses of 2, 3 and 4, and insertion on the lateral dorsal edge of the exoccipital. M. rectus capitis superior This muscle lies immediately superficial to M. flexor colli brevis. Origin is from the lateral surface of neural arch i, from the anterior surfaces of anapophyses 2 and 34 P. J- K. BURTON 3, from the lateral strut of 4, and from the transverse processes of 5 and 6. Origin from 6 is absent in all the female specimens and in < 2 ; origin from 5 is also absent in $ C. Insertion is aponeurotic and fleshy on a ridge at the posterior edge of the basitemporal plate. M. rectus capitis ventralis Origin is from the ventral surface of i, from the hyapophyses of 2, 3, 4 and 5, and from the sublateral process of 6 where there is some fusion with M. flexor colli profundus. The right and left Mm. recti capiti ventrales are fused in the midline and insert together on the basitemporal plate, anterior to M. rectus capitis superior. VIII. OTHER CALLAEIDAE Creadion carunculatus The description of the Huia's skull by Oliver (in Phillipps, 1963) refers to the general similarity of the skull of Creadion. Oliver notes the greater extent of ossi- fication in Heteralocha, manifested especially at the front of the orbit. The skull and skeleton of this species are figured by Shufeldt (1913), who includes some brief notes on its skull for comparison with Anthochaera carunculata (Meliphagidae) . The following additional points may be noted. The retroarticular process is highly developed in Creadion, but is nevertheless relatively shorter than that of the Huia ; there is no raised occipital crest, and little development of an exoccipital process. Otherwise, skull proportions are similar to Heteralocha, particularly female specimens, though the quadrate and pterygoid are relatively smaller. The skull is apparently larger relative to body size in the Saddleback if a rough index (sternum length) from a single specimen can be relied upon. Ligaments have been removed from the skull of the available specimen, and could not be studied. With the exception of M. depressor mandibulae, jaw muscles have also been removed from the specimen of Creadion. This remaining muscle is, however, of considerable interest, since it possesses a pars anterior as in Heteralocha - a feature which otherwise appears to be unique among birds so far studied. The rest of the muscle is also well developed, and similar in structure to that of the Huia, though relatively less massive. The tongue, like the jaws, is smaller relative to the skull in Creadion, but the hyoid musculature is closely similar in both Saddleback and Huia. Due to the historic interest of the specimen, only limited dissection of the neck muscles could be undertaken for Creadion. Five of the muscles exhibiting variation in the Huia or among other passerines were examined. These showed an overall reduction in the number of sites of attachment. M. spinalis lacks the slip inserting on 5 which is present in three of the Huia specimens, though a slip to 6 (absent in Huia <$ 2) is present. M. splenius colli lacks the slip to 7 found in four of the Huia specimens. M. flexor colli brevis and M. rectus capitis superior lack attachment to 6, a feature found in three and two of the male Huia specimens respectively. M. corn- plexus also lacks a slip to 6, although this is present in all six Huias dissected. ANATOMY OF HEAD AND NECK IN THE HUIA 35 Callaeas cinerea The skull and other features of the skeleton in the Kokako have been described in some detail by Stonor (1942). It is sufficient here to draw attention to the extensive differences in skull proportions and geometry between this bird, on the one hand, and Heteralocha and Creadion, on the other. The bill is shorter, but much deeper, and the jugal bar meets it at a greater angle. The quadrate is relatively much larger, and is rotated backwards by comparison with the other Callaeidae, so that its otic process meets the lower jaw more nearly at right angles. The cranium is shortened, and smaller relative to the orbits ; the jugal is very long relative to the skull. The lower jaw is deep, and bears strong cristae for the attachment of the adductor musculature. There is no retroarticular process, and the articular is extremely shallow at its articulation with the quadrate. The prominent sesamoids at this articulation were noted by Stonor, and their origin and functions are discussed in detail by Burton (1973). As in the Huia, the external jugomandibular ligament is absent. The occipitomandibular ligament is not ossified at all. The jaw musculature is notable for the highly developed adductor musculature, and the very small M. depressor by comparison with the other Callaeidae. M. adductor mandibulae externus is considerably more bulky in actual (as well as rela- tive) size than in Heteralocha, with increased number of fibres and greater com- plexity. Its aponeuroses are generally stouter, and more branched and subdivided internally (Fig. 18), providing extra surface for fibre attachment, and consequently a greater use of pinnate structure. M. pseudotemporalis superficialis is also much larger than in the Huia and Saddleback, and in dorsal view completely conceals M. pseudotemporalis profundus, which is much reduced - a consequence of the back- ward displacement of the quadrate. M. pterygoideus is bulky and the retractor palatini slip is especially well developed, and prominent in dorsal view. M. depressor mandibulae is not only smaller than in the other Callaeidae, but structurally simpler, and entirely lacks a pars anterior. It is unique, however, for the pulley arrangement between its aponeurosis and the internal jugo-mandibular ligament (Burton, 1973). The tongue is much broader relative to its length than in the Huia and Saddleback ; its shape is almost rectangular, and the tip brush-like. The paraglossa which sup- port it are widely separated. There is no median aponeurosis and M. ceratoglossus anterior is absent ; otherwise, the hyoid musculature resembles that of the other two genera of Callaeidae. The palate is provided with horny papillae similar to those of the other Callaeidae, but the horny lining of both jaws is developed near the bill tip into raised, papillate bosses. There is a pair of Gil. angularis oris, but a Gl. palatinae has not been found. Neck muscles showing variation in their points of attachment in Heteralocha or other passerines were examined in Callaeas. The two specimens dissected had the same number and sites of attachment points for the muscles examined. Slips or muscle components absent in some Huia specimens, but present in the Kokako specimens, were the insertions of M. spinalis cervicis on 5 and 6 ; M. splenius colli arising from 7 ; M. intercristalis from 5 to 3 ; the slip of M. ascendens from 8 to 4 ; and the slip of M. flexor colli brevis to 6. No Mm. pygmaei arise from 12, though this site is occupied in all but one male Huia specimen. However, an additional M. 36 P. J. K. BURTON pygmaeus from 7 to 6 is present in the Kokako specimens, though absent in all the Huias dissected. The Kokakos lacked the slip of M. flexor colli profundus from 6 to 2, present in three of the Huias, and the slip of M. complexus to 6, present in all the Huias. M. flexor colli brevis in the Callaeas specimens lacks the slip to 6 present in two male Huias, but has a slip to 5 (unlike Creadion and one female Huia specimen) . IX. FUNCTIONAL ASPECTS Jaw mechanism Next to the form and sexual dimorphism of the bill, the most striking features of the Huia's cranial morphology are the huge M. depressor mandibulae and associated skull modifications - the prominent occipital crest, providing extra surface for its origin, and the very long retroarticular process providing increased leverage for the muscle. Clearly these adaptations must permit the lower jaw to be depressed with great force. This can only be necessary if it is to be opened against considerable resistance by external forces, and it seems certain, therefore, that the Huia was highly specialized for feeding by 'gaping' or 'prying'. This feeding technique con- sists basically of thrusting the bill into a potentially food-bearing medium (earth, w r ood, fruit, etc.) and opening it, to widen the hole and so facilitate exploration or prey extraction. Gaping is a technique described from birds of several families. Among passerines, good examples are furnished by some Sturnidae and many Icteridae. Gaping behaviour and related modifications of M. depressor mandibulae have been studied in the Icteridae by Beecher (1951). It is interesting to compare the Huia with other birds specialized for gaping, and some figures for relative length of the retroarticular process in several examples, mainly of passerines, are given in Table 2. The nearest approach is shown by Cacicus solitarius in which the retroarticular process is slightly longer relative to the lower jaw than in the Huia. However, Cacicus solitarius is a shorter billed bird ; relative to skull length, its retroarticular process is shorter than in Heteralocha, a difference which would be even greater but for the Huia's elongated skull with enlarged occipital crest. It seems clear that the Huia was very highly specialized for feeding by gaping, perhaps more so than any bird now living. Buller's account of the Huia's feeding behaviour is an excellent one, but clearly gives an incomplete picture of the male's excavation methods. Buller remarks that the captive male used its bill to 'chisel out' pieces of decayed wood in a woodpecker- like manner. Almost certainly its precise technique must usually have been to drive the bill into the wood and 'gape' to split portions off ; this is actually quite different in principle from the methods of woodpeckers which depend purely on blows. Buller states that the female he observed fed in a quite different manner, by probing into relatively hard wood. However, M. depressor mandibulae and the retroarticular process, though somewhat smaller in the female, are still very large by comparison with other birds, and her capacity for forceful depression of the lower jaw must also have been very high. Due to the greater length of the female's bill, less force could be exerted at its tip than in the male ; and the more flexible nature of its anterior, purely rhamphothecal, portion, appears rather inefficient for gaping. ANATOMY OF HEAD AND NECK IN THE HUIA TABLE 2 Relative length of the retroarticular process in single specimens of various birds showing gaping adaptations, and in all intact Huia specimens used in this study 37 f S I (Si o .a K Upupa epops Phoeniculus purpureus Phoeniculus aterrimus Psarocolius decumanus Psarocolius wagleri Cacicus cela Cacicus leucorhamphus Cacicus solitarius Sturnella magna Amblyrhamphus holosericeus Creatophora cinerea Sturnus vulgaris Sturnus contra Heteralocha acutirostris, $ (mean of 3) Heteralocha acutirostris, $ (mean of 2) Pyrrhocorax pyrrhocorax 81-0 60-4 45'4 65-2 72-6 43'i 48-7 53'5 44'2 45'4 41-7 43'9 49-5 89-4 109-5 77-0 6-2 4'4 7-8 7.9 4-0 5'2 10-3 5'5 8-0 2-8 5'0 4'7 16-4 I3-5 5'4 25'4 26-3 20-5 36-9 37'0 29-1 29-8 31-1 31-3 28-0 28-8 29-4 27-8 46-8 40-9 0-07 o-io O'lO O'I2 O'll O'O9 O'll 0-19 0*12 0-18 0-07 O'll 0-09 0-18 0-07 0'22 0'24 O'2I 0'2I 0'2I 0-14 0-17 o'33 0-17 0-29 O'lO 0-17 0-17 0-31 0-13 Nevertheless, gaping must also have been an important part of her repertoire of feeding techniques in addition to exploratory probing (as described by Buller) for which the decurved bill shape is evidently adapted. In probing timber tunnelled by beetles, it seems feasible that the bill might occasionally be inserted through a crack into a larger cavity, so that gaping could take place with contact only in the more rigid basal part of the bill. However, the possibility remains that the female's apparent gaping adaptations reflect principally the shared genotype of the two sexes (see under Sexual Dimorphism). In addition to its large size, M. depressor mandibulae in the Huia also shows interesting structural complexities. A major contribution to the force of depression is obviously provided by the large mass arising posteriorly and dorsally on the cranium, since this is not only bulky, but has the longest moment arm due to its insertion near the posterior extremity of the retroarticular process. However, the 38 P. J. K. BURTON anterior parts of the muscle also exhibit various modifications. Several aponeuroses are present, serving as the basis for pinnate fibre arrangements ; this may be related to their short working distance, for which pinnate structure should provide more forceful contraction than parallel fibred muscles of the same physiological cross section (Gans and Bock, 1965). The pars anterior of the muscle (present also in C reaction) is of particular interest, since it appears not to have been described pre- viously in any other bird, and was certainly absent from gaping species of other families dissected during this study. Its functions are hard to surmise, and a satisfactory explanation will probably require a much more detailed knowledge of the jaw mechanics involved in gaping than is available at present. Except for fibres inserting on the internal process, the greater bulk of M. depressor mandibulae has a medial component in contraction which is greatest for the most posterior parts of the muscle originating near the midline of the cranium. The unusual prominence of the medial condyle of the quadrate may in part serve to resist the stress which this places on the quadrate/mandible articulation. Probably M. depressor also aids protraction of the upper jaw by the action of its upward force component on the quadrate (see Bock, 1964, 1968, and Zusi, 1962, 1967, for discussions of this mechanism), but M. protractor quadrati et pterygoidei is itself well developed. It is certainly evident that upper jaw action contributes substantially to 'gaping'. The unusual situation of the nasal-frontal hinge may be relevant in this respect. Its position, more dorsal and posterior than in other passerines, places it further from the line of action of the protractor force, acting through the palatines and jugals ; the moment arm of this force is consequently increased. The remainder of the jaw musculature, concerned with adduction of the lower jaw and depression of the upper jaw, calls for less comment. It is well developed, as seems necessary to provide sufficient grip while extracting huhu grubs, but not remarkably so ; in general features, the muscles other than M. depressor and M. protractor resemble those of Corvus described by Bock (MS). Skull architecture There is a need for detailed and comparative mechanical analyses of the skull in 'gaping' passerines of a comparable depth to those available for biting and seed crushing forms, e.g. Bock (i96oa, 1964^ 1966) and Bowman (1961). Here, some features of possible significance in the skull of Heteralocha will be briefly noted. The wedge-shaped bill of the male Huia is typical of the majority of 'gapers' and resembles that of many Icterids. The straight-sided culmen and gonys are more efficient than the blunter, arched bill of most passerines for the task of forcing the bill into the substrate prior to gaping. Decurved bills, such as those of the female's, or some wood hoopoes (Rhinopomastus spp.), depend for this on the presence of existing crevices or tunnels. In the case of the wedge-shaped bill, the narrower and more acutely conical its form, the more easily it will penetrate the substrate ; a limit is imposed by the danger of damage by forces directed across the bill axis, since this hazard is less in a more broadly based cone (see Bowman, 1961, p. 222-224). Evidently this danger was not exceptionally high in the Huia, since the male's bill ANATOMY OF HEAD AND NECK IN THE HUIA 39 is narrower and more acute than in many Icterids, including the wood-prying caci- ques ; and in both sexes, the angle between the jugal bars and mandibular rami (giving lateral support to the bill) is no greater than in many unspecialized passerines. The culmen of the bill and frontal bridge of the skull are virtually in line, again a feature of many 'gapers', but particularly well marked in the Huia, with the nasal- frontal hinge shifted to an unusual position nearly halfway across the orbital region. This profile eliminates the difficulties that might arise with a more conventional skull possessing a marked 'forehead' at the nasal-frontal hinge if the bill were thrust into the feeding substrate up to or past its base. Many Icterids in fact have the culmen raised above the level of the frontal bridge ; the horny plate extending onto the forehead of some members of this family (e.g. Gymnostinops, Psarocolius) is probably adaptive primarily to gaping in succulent fruits (Beecher, 1951). The stoutness of the quadrate-jugal articulation in Heteralocha is probably related to the great development of M. depressor mandibulae. The prominence of the medial condyle of the quadrate and its orientation are also unusual ; the long axis of the condyle is directed more nearly at right angles to the skull axis than in most passerines, a feature which may be adaptive to resisting backward disarticulation by strong external forces. The nasal-frontal hinge appears to be unusually ill defined in Heteralocha, and manipulation of skulls relaxed by boiling suggests that the bending zone is con- siderably longer and stiffer than in most holorhinal birds. This may have some significance as a safety factor, since a wood 'gaper' could be exposed to some danger of over-protraction in the event of timber splitting suddenly while a strong force was still being exerted by the upper jaw. However, there are no bony protraction stops comparable to the overhanging frontal 'brow' of woodpeckers - part of a completely distinct adaptive complex based on the use of M. protractor to distribute compression forces while hammering (Spring, 1965, Bock, 1966). Buccal cavity, tongue and hyoid musculature Although the Huia's manner of gaining access to its food is specialized, the food itself (Cerambycid larvae) and the process of swallowing it pose no unusual problems. It is consequently no surprise that the papillae of the upper jaw, tongue and larynx resemble those of many other passerines which feed on invertebrate prey. The hyoid musculature is similarly unspecialized. Cerambycid larvae are smooth skinned, and would not seem to require a great deal of lubrication. The limited complement of salivary glands is consistent with this supposition. Neck musculature By comparison with the other two species of Callaeidae, the neck musculature of Heteralocha appears better developed inasmuch as several of the muscles or muscle groups occupy additional sites of origin or insertion. It is difficult, however, to draw general inferences from this, owing to the scarcity of information on other passerines. Palmgren's (1949) study, though detailed, concentrated on small species of passerines ; Boas (1929) confined his work mainly to non-passerines. In the absence of such information for larger passerines, it is not possible to know to what 4 o P. J. K. BURTON extent this difference is due to the Huia's larger size. For the present it is only possible to remark that greater development of the neck musculature would be expected in a bird so highly adapted to the extraction of prey from timber by vigorous techniques. Other Callaeidae The Saddleback is known to excavate for insect food in decayed timber, bark, epiphytes, etc., and also takes some huhu grubs. It is clear from its skull that in this species, too, gaping must play a large part in its feeding activities. However, it is less specialized than the Huia, even apart from the lack of pronounced sexual dimorphism. Correlated with its smaller body size, it includes a larger proportion of small insects and other items in its diet, and has never reached the high degree of dependence on a single prey species seen in the Huia. The Kokako is primarily a fruit and leaf eater. In many features of its cranial morphology it shows convergence with finches or with parrots. These features are centred around the need for powerful adduction for biting hard or tough vegetable foods. In the skull, the jaws are shortened and deepened, and the quadrate/mandible articulation shifted posteriorly. The curious structure of the mandible/quadrate articulation is evidently a consequence of these trends (Burton, J-973a) . There is no retroarticular process, and it is doubtful if gaping adaptations were ever present in its ancestry. The adductor musculature is more bulky and complex than in the Huia and Saddleback, and the tongue is specialized, probably for dealing with fruits. X. SEXUAL DIMORPHISM Bill The great difference in size and shape of the bill between male and female Huias is of course well known. It does not appear to have been realized, however, to what a large extent this is due to differing development of the rhamphotheca. With the rhamphotheca removed, the bill length difference is reduced and the skulls of the two sexes resemble each other much more closely (Fig. 26) . The greatly elongated rhamphotheca of the female is remarkable not only by comparison with the male, but also when compared with other birds of similar bill form. Table 3 gives figures for upper jaw length with and without rhamphotheca in specimens of a variety of birds (mainly passerines) with long decurved bills. It can be seen that all of these fall well short of the female Huia in relative development of the rhamphotheca, and some bills of very extreme form are bony right to the tip. It is difficult to assess the significance of this feature. From a functional viewpoint, it might be suggested that the relatively flexible material of the rhamphotheca would be in less danger of breakage than bone when used for vigorous exploration of timber. However, developmental and genetic factors should also be taken into considera- tion. The greater development of the rhamphotheca in the female may be a special case of allometric growth ; this might be investigated by soft x-ray photography of the fairly extensive series of Huia skins in the world's museums. It is also possible ANATOMY OF HEAD AND NECK IN THE HUIA 20mm FIG. 26. Comparative development of the rhamphotheca in male and female of Heteralocha acutirostris (<$ i and $ C). Solid black areas represent the profile of the rhamphotheca. that the answer may lie in the differences between the genetic control of development of rhamphotheca and of bone ; it may simply be 'easier', genetically, to evolve extreme sexual dimorphism of bill length in this way. Other examples of this phenomenon may exist. For example, the lark Alauda razae shows marked sexual dimorphism in bill length (Burton, igyib) and soft x-ray photographs of three specimens of each sex show proportionately greater rhamphothecal development in the longer-billed males. Investigation of longer series of this and other sexually dimorphic species may shed further light on this problem. Close examination of Huia specimens showing features intermediate between male and female (e.g. some specimens mentioned by Phillipps, 1963) also seems desirable. It may be mentioned in passing that the male specimen figured by Garrod (1872) seems to have been unusual in this respect. Garrod mentions that its tongue was a third the length of the bill. Assuming the tongue was of normal length, this would mean a bill some 90 mm long, a length normally found only in females. The figures show other unusual features ; the jaw tips are attenuated and slightly decurved, rather as in female specimens, and the occipital crest is less prominent than in the males examined in this study. On the other hand, bill depth and stoutness of the jugal, palatines and pterygoids are characteristically male. The specimen was acquired by the Zoological Society of London nearly two years before its death 4 2 P. J. K. BURTON TABLE 3 Extension of rhamphotheca beyond upper jaw tip in single specimens of various birds with long downcurved bills, and in male and female Huia specimens used in this study Upupa epops Rhinopomastus cyanomelas Campy lor hamphus trochilirostris Falculea palliata Nectarinia famosa Arachnothera robusta Hemignathus procerus Heteralocha acutirostris, <$ (mean of 3) Heteralocha acutirostris, $ (mean of 2) C Si) -S O -E, 03 63-5 37-7 62-0 66-7 35'2 54'9 60-6 50-2 90-5 o n3 -i-> g i*i o 3 _^ oj 3 -g a s?- s 2 5 **- T3 O V-, 52-3 33'0 55'5 58-5 30-8 53'9 46-8 43'5 49'7 M ns 5 0-18 O'I2 O'lO O'I2 0-13 O'O2 0'23 o-io 0-45 (Zoological Society of London, 1870) ; possibly it was young when acquired, and developed abnormally in captivity. Jaw mechanism M. depressor mandibulae is of essentially similar structure in both sexes, but distinctly less massive in the female. Differences in skull features associated with this are the less prominent occipital crest (and consequently shorter skull), and shorter exoccipital and retroarticular processes of the female. Comparisons between male and female for the other jaw muscles are of very limited value, since these remain in only one female specimen ($ C) . In this speci- men, M. adductor mandibulae externus appears distinctly more bulky than in the three males, although the medial slip of M.a.m.e. rostralis median's is shorter. Greater development of this muscle would seem reasonable in view of the female's greater bill length, a consequence of which would be to reduce the mechanical advantage of the muscle for bill tip adduction - an action which might require considerable force when gripping and extracting huhu grubs. Skull Features related to differences in bill form and in development of M. depressor mandibulae have already been mentioned. Apart from these, the proportions of the ANATOMY OF HEAD AND NECK IN THE HUIA 43 skull are very similar in both sexes, as can be seen from Table i. The only significant difference in proportion is in the region of the bill base. In the female, the bill is relatively narrower and less deep at the base, and the skull is narrower anterior to the orbits. Probably her mode of feeling involved less exposure to forces directed across the bill axis than the more vigorous techniques of the male. Ratios of jugal length to sternum length give no indication of any difference between the sexes in relative head size. Buccal cavity, tongue and tongue musculature Despite the much greater length of the female's bill, the tongue is of similar size in both sexes. It would have been of little use for the extraction of prey by the female, and in both sexes was probably concerned simply with manoeuvring food during the process of swallowing. It is therefore not surprising that the hyoid musculature shows no obvious sign of sexual dimorphism. The same is true also of the buccal papillae and salivary glands, whose functions would have been similar in both sexes. Neck musculature Several of the neck muscles and muscle groups showed variations in the number and disposition of components. Table 4 compares the distribution of these variable components in the three spirit specimens of each sex, and a fairly clear pattern emerges. Variable slips and components of the dorsal muscles (concerned mainly TABLE 4 Distribution of neck muscle components present in some, but not all, of the Huia specimens Number of specimens in which present Dorsal components M. spinalis insertion on 5 M. spinalis insertion on 6 M. splenius colli origin on 7 M. pygmaeu? from 12 to II M. ascendens from 8 to 4 TOTAL Ventral components M. flexor colli profundus from 6 to 2 M. flexor colli profundus slip joining M. flexor colli brevis M. flexor colli brevis origin from 6 M. rectus capitis superior origin from 5 M. rectus capitis superior origin from 6 Male i 2 I 2 2 Female 2 3 3 3 o ii i i o 2 O TOTAL 3 5 4 5 2 19 3 3 3 5 2 TOTAL 12 16 44 P. J- K. BURTON with raising the neck) are slightly better represented in the female specimens. However, a much greater disparity exists among the variable components of the ventral muscles (concerned with lowering the head and neck) , which are much better represented in the male specimens. This discrepancy is entirely consistent with the difference between the feeding methods of male and female. Forceful downward movements of head and neck must have been essential to enable the male to insert his bill into timber before 'gaping' to split it ; this would have been much less important in the female who relied on the presence of existing tunnels and crevices into which to insert her bill. The preponderance of variable components of the dorsal musculature in the female is less extreme, but may also correspond to a difference in use. Extracting huhu grubs must have entailed exertion in lifting the head and neck for both sexes ; but this exertion may often have been greater for the female, since a larger proportion of her prey would have to be removed from more or less intact tunnels due to the harder wood in which she fed and her lower capacity for forceful 'gaping'. XI. CONCLUDING REMARKS Several studies of the Huia's morphology have been undertaken in attempts to clarify its systematic position. I would be reluctant, however, to draw any con- clusions about the Huia's affinities from the results of the present investigation. Most of the foregoing descriptions and discussions have been centred around the extremes of sexual dimorphism and adaptation for gaping shown by the Huia, but neither phenomenon is unique among passerines except in degree. Indeed several of the features to which Garrod (1872) gave greatest weight in allying Heteralocha with the Sturnidae were gaping adaptations and the genera of 'Sturnidae' which he examined included several New World forms now assigned to the quite distinct family Icteridae. Firstly, the size of M. depressor mandibulae (the 'digastric' muscle) which impressed Garrod and also Lowe (1938) with its similarity to various Sturnidae is an unsatisfac- tory character unless considered in conjunction with the structure of the muscle. In this respect, Heteralocha and Creadion both show a feature which is apparently unique - a pars anterior, extending forwards from the articulation with the quadrate. Secondly, Callaeas, an undoubted relative (Stonor, 1942) of these two genera, shows no gaping adaptations, and quite possibly did not evolve from a gaping ancestor. It seems reasonable to suggest that the gaping adaptations of the Huia and Saddle- back evolved independently of other families and in isolation, and if this is correct, they are poor evidence for affinity with other gaping forms. The details revealed by this study do not, either, affect the general conclusion of Selander (1966) that extreme sexual dimorphism in feeding structures is particularly frequent in insular bird species, since it provides a means of utilizing a wider range of food resources in conditions of reduced competition. However, the dimorphism of the bill in the Huia is the most extreme example known of this phenomenon, and it is natural to wonder how such a large difference between the sexes evolved. The gaping adaptations may well have played a crucial part. Gaping is an unusual means ANATOMY OF HEAD AND NECK IN THE HUIA 45 of exploiting the food resources of timber, and among passerines in general, probing is a much more common technique. Nevertheless, gaping was almost certainly the primary adaptation in the stock from which the Huia arose. The existence of similar adaptations in both sexes of the Saddleback lends strong support to this presumption. If so, the female Huia's bill form appears to be a secondary adaptation for probing, probably evolved from what was initially a slightly more slender (and very likely shorter) bill than that of the male. An early appearance of a sexual difference in feeding technique would be essential for this to take place. It should be stressed here that Buller's account gives no evidence for cooperation between the two sexes in feeding, although the terms in which it is couched appear to suggest this and have often been interpreted in this way. Almost certainly each simply fed independently in the manner for which it was adapted. The female probably derived occasional benefit from the excavations of the male, but the reasons for the Huia's regular association in pairs may primarily have been social or sexual. Assuming the Huia to be extinct, a full understanding of the significance of its dimorphism may never be achieved. However, there are still various lines of in- vestigation which might profitably be pursued. Further use could be made of the existing material of Heteralocha. In addition to the 119 New Zealand specimens examined by Phillipps (1963), several museums in other parts of the world hold series of skins. Examination of all this scattered material may prove difficult, but it would seem desirable to have the fullest available quantitative data on the extent of dimorphism and of variation within each sex. The data so far available suggest that the female's bill may have been more variable than the male's. If true, this might indicate that females obtained food in a greater variety of situations or by more versatile techniques than males (although Phillipps suggests that the male ate a greater proportion of insects other than huhu grubs) . It certainly appears that the pair bond was very strong in the Huia, and that both sexes participated in feeding the young. Studies of the Saddleback may provide information from which inferences about the Huia's behaviour can be drawn, and investigations on its feeding ecology and functional anatomy are currently in progress (Jenkins, pers. comm.). Finally, studies on other timber-feeding birds may shed further light on the Huia's adapta- tions. The closest parallels to these appear to be shown not by passerines, but by the Wood Hoopoes (Order Coraciif ormes, family Phoeniculidae) . Within this family, the genus Phoeniculus (particularly P. aterrimus) appears to parallel the male Huia, while Rhinopomastus spp. resemble the female, both genera showing well-marked gaping adaptations. A study of feeding behaviour and anatomy in this family, as well as being worth while in its own right, would make an interesting comparison with the Huia, in which similar bill forms and feeding methods have evolved in the two sexes of a single species. Yet, finally, it must be admitted that these indirect methods of investigation are a very poor substitute for observations on the living bird. A study of this species in life might have provided much information of general importance for evolutionary biology, and would certainly have been of intense interest ; its loss is a matter for the greatest regret. 46 P. J. K. BURTON XII. ACKNOWLEDGEMENTS I am much indebted to Dr Walter J. Bock for reading the first draft of this paper, and offering valuable criticism and advice. The excellent drawings of skulls (Figs. 1-9) are the work of Sharon Chambers, to whom I am deeply grateful. Mr P. F. Jenkins, of the University of Auckland, kindly let me have news of his work on the Saddleback. Thanks are due to the British Trans-Americas Expedition (1971-2) for enabling me to collect several of the specimens used for comparison with the Callaeidae. XIII. REFERENCES AMADON, D. 1943. Bird weights as an aid in taxonomy. Wilson Bull. 55 : 164-177. ANTONY, M. 1920. Uber die Speicheldriisen der Vogel. Zool. Jb. (Abt.Anat.) 41 : 545-660. BEECHER, W. J. 1951. Adaptations for food getting in the American Blackbirds. Auk 68 : 411-440. BOAS, J. E. V. 1929. Biologisch-anatomische Studien iiber den Hals der Vogel. K. dansk. Vidensk. Selsk. Skr. (Naturvidenskab. Math. Afdel.) Ser. 9, 1 : 102-222. BOCK, W. J. I96oa. The palatine process of the premaxilla in the Passeres. Bull. Mus. comp. Zool. Harv. 122 : 361-488. ig6ob. Secondary articulation of the avian mandible. Auk 77 : 19-55. Kinetics of the avian skull. /. Morph. 114 : 1-41. Bill shape as a generic character in the cardinals. Wilson Bull. 76 : 50-61. 1966. An approach to the functional analysis of bill shape. Auk 83 : 10-51. 1972. Morphology of the tongue apparatus of Ciridops anna (Drepanididae) . Ibis 114:61-78. MS : Drawings of jaw and tongue musculature in passerine birds. BOWMAN, R. I. 1961. Morphological differentiation and adaptations in the Galapagos finches. Univ. Calif. Publs Zool. 58 : vii + 326 pp. BULLER, W. L. 1878. Further descriptive notes of the Huia (Heteralocha acutirostris) . Trans. N.Z. Inst. 10: 211. 1888. A History of the Birds of New Zealand. 2nd edition. Vol. i. London. BURTON, P. J. K. 1969. Two bird specimens probably from Cook's voyages. Ibis 111 : 388- 390. I97ia. Some observations on the splenius capitis muscle of birds. Ibis 113 : 19-28. i97ib. Sexual size dimorphism in Alauda razae. Bull. Brit. Orn. Club 91 : 108-109. 1973- Structure of the depressor mandibulae muscle in the Kokako Callaeas cinerea. Ibis 115 : 138-140. 1974- Feeding and the Feeding Apparatus in Waders. Brit. Mus. (Nat. Hist.). ENGELS, W. L. 1938. Tongue musculature of passerine birds. Auk 55 : 642-650. FIEDLER, W. 1951. Beitrage zur Morphologic der Kiefermuskulatur der Oscines. Zool. Jb. (Abt.Anat.) 71 : 235-288. CANS, C. & BOCK, W. J. 1965. The functional significance of muscle architecture - a theoreti- cal analysis. Ergebn. Anat. EntwGesch. 38 : 115-142. GARROD, A. H. 1872. Notes on the anatomy of the huia bird (Heteralocha gouldi) . Proc. Zool. Soc. Lond. : 643-647. GEORGE, J. C. & BERGER, A. J. 1966. Avian Myology. London and New York. GRAY, G. R. 1870. Hand-list of Genera and Species of Birds distinguishing those contained in the British Museum. Part 2. London. LAKJER, T. 1926. Studien iiber die trigeminus-versorgte Kaumuskulatur der Sauropsiden. Copenhagen. LOWE, P. R. 1938. Some anatomical and other notes on the systematic position of the genus Picathartes, together with some remarks on the families Sturnidae and Eulabetidae. Ibis i4th Ser. 2 : 254-269. ANATOMY OF HEAD AND NECK IN THE HUIA 47 MAYR, E. & GREENWAY, J. C. 1956. Sequence of passerine families (Aves). Breviora No. 58 : i-n. OLIVER, W. R. B. 1955. New Zealand Birds. 2nd edition. Wellington. 1963. The skull of the Huia. In The Book of the Huia by W. J. Phillipps. Christchurch. PALMGREN, P. 1949- Zur biologischen Anatomic der Halsmuskulatur der Singvogel. In Ornithologie als biologische Wissenschaft, pp. 192-203. Heidelberg. PHILLIPPS, W. J. 1963. The Book of the Huia. Christchurch. SELANDER, R. K. 1966. Sexual dimorphism and differential niche utilization in birds. Condor 68 : 113-151. SHUFELDT, R. W. 1913. On the osteology of the Red Wattle-Bird (Anthochaera carunculata) . Emu 13 : 1-14. STARCK, D. & BARNIKOL, A. 1954- Beitrage zur Morphologic der Trigeminusmuskulatur der Vogel. Morphol. Jahrb. 94 : 1-64. STONOR, C. R. 1942. Anatomical notes on the New Zealand Wattled Crow (Callaeas) with especial reference to its powers of flight. Ibis nth Ser., 6 : 1-18. ZOOLOGICAL SOCIETY OF LONDON, 1870. Report on additions to the Society's Menagerie during the month of May. Proc. Zool. Soc. Lond. : 380-384. Zusi, R. L. 1962. Structural adaptations of the head and neck in the Black Skimmer Rynchops nigra Linnaeus. Publs Nuttall orn. Club No. 3 : vii + ioi pp. 1969. Osteology and myology of the head and neck of the Pied-billed Grebes (Podilvmbus). Misc. Publs Mus. Zool. Univ. Mich. No. 139 : 1-49. KEY TO ABBREVIATIONS IN FIGURES Ap i a m e Aponeurosis i, M. adductor mandibulae externus Ap 2 a m e Aponeurosis 2, M. adductor mandibulae externus Ap 3a d m Aponeurosis 3a, M. depressor mandibulae Ap m Median aponeurosis of tongue C o Occipital crest E u s Opening of Eustachian tube G a o Gl. angularis oris L i j m Internal jugomandibular ligament L p o Postorbital ligament M a m e c M. adductor mandibulae externus caudalis M a m e r 1 M. adductor mandibulae externus rostralis lateralis M a m e r m M. adductor mandibulae externus rostralis medialis M a m e r t M. adductor mandibulae externus rostralis temporalis M a m e v M. adductor mandibulae externus ventralis M a m p M. adductor mandibulae posterior M as M. ascendens cervicis. M b m M. branchiomandibularis Meg M. ceratoglossus M c g t Tendon of M. ceratoglossus M co M. complexus M d m M. depressor mandibulae M d m a M. depressor mandibulae pars anterior M dt M. dermotemporalis M f c b M. flexor colli brevis M hg a M. hypoglossus anterior M hg o M. hypoglossus obliquus M i t M. intertransversarius M p q i M. protractor quadrati et pterygoidei i M p q 2 M. protractor quadrati et pterygoidei 2 M ps p M. pseudotemporalis profundus 48 P. J. K. BURTON M ps s M. pseudotemporalis superficialis M pt d 1 M. pterygoideus dorsalis lateralis M pt d m a M. pterygoideus dorsalis medialis anterior M pt d m p M. pterygoideus dorsalis medialis posterior M pt r M. pterygoideus retractor M pt v 1 M. pterygoideus ventralis lateralis M pt v m M. pterygoideus ventralis medialis M r c 1 M. rectus capitis lateralis M r c v M. rectus capitis ventralis M s co M. splenius colli M se h M. serpihyoideus M sp M. spinalis M st h M. stylohyoideus M th h M. thyreohyoideus M tr h M. tracheohyoideus M tr 1 M. tracheolateralis N f h Nasal-frontal hinge P ex Exoccipital process P i Internal process of mandible P r Retroarticular process of mandible Dr P. J. K. BURTON Sub-department of Ornithology BRITISH MUSEUM (NATURAL HISTORY) TRING HERTFORDSHIRE 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. 1-134 > T Plate, 77 Text-figures. 1974- 375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU MISCELLANEA 19 AUG1974 v * .c^V BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 2 LONDON: 1974 MISCELLANEA Pp 49-138 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 2 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 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. 27, 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.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 1 8 July, 1974 Price 4.70 CONTENTS Page A new skink from the New Hebrides. By LORD MEDWAY ... 53 New genus of whale-louse. By R. J. LINCOLN & D. E. HURLEY . . 59 Catalogue of whale-lice. By R. J. LINCOLN & D. E. HURLEY ... 65 Review of Laephotis Thomas 1901. By J. E. HILL .... 73 The genus Finmarchinella Swain. By J. W. NEALE .... 83 Euplotes rariseta sp. n. By G. R. CURDS, B. J. WEST & J. E. DORAHY . 95 Clonal cultures of Euglypha (Protozoa). By R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT .......... 103 Three species of Euplotes. By C. R. CURDS . . . . . . 113 Bats from Southeastern Asia. By J. E. HILL . . . . . 127 A NEW SKINK (REPTILIA: SCINCIDAE: GENUS EMOIA) FROM THE NEW HEBRIDES WITH COMMENTS ON THE STATUS OF EMOIA SAMOENSIS LOYALTIENSIS (ROUX) By LORD MEDWAY DURING the Royal Society/Percy Sladen expedition to the New Hebrides, 1971, three immature skinks of an undescribed taxon were caught on Aneityum island. Subsequently three further specimens, taken on the same island by Miss Evelyn Cheesman, were found among the collections of the British Museum (Natural History) where they had been catalogued as Emoia nigra. Apart from a note that the period of collecting was 1954-55, Miss Cheesman's material lacks field data. It is likely, but not certain, that the skinks were caught near Red Crest, the camp three miles from the coast (at Anelgauhat) at about 1200 ft (366 m) elevation, occupied by Miss Cheesman from mid-March through July 1955 (see Cheesman, 1957 : 312-327). Of this site, Miss Cheesman has written : 'Plenty of lizards darted about all over the clearing on sunny days, overcast days made them inactive and then one found them under logs, in holes, or hollow trees . . . Individuals were inclined to be tame, a smooth blue-grey skink with a green mate would appear whenever I had meals outside on the clearing. She accepted cheese, bread, biscuit, boiled rice and banana but refused cooked taro, beating a retreat as if insulted. To catch lizards was another matter, but I did succeed in getting a good series.' During the 1971 expedition skinks were also collected on all other islands visited, i.e. in order northwards from Aneityum, Tanna, Erromanga, Efate, Malekula, Malo, Aore and Espiritu Santo. Previous collections from these islands, and from others in the group, have been reported by Boulenger (1887), Roux (1913), Schmidt & Hurt (1930), Burt & Burt (1932) and Angel (1935). Additional unpublished material from the New Hebrides, examined in the course of preparation of this note, is pre- served in the Museum National d'Histoire Naturelle, Paris (Frangois collection), the Natur-Museum Senckenberg, Frankfurt a. Main (Bregulla collection), and the British Museum (Natural History) (mainly collected by J. R. Baker and associates or by E. Cheesman). Other than the six examples from Aneityum, no other repre- sentatives of the undescribed skink have been found among any collections. Since it is evidently confined to the island of Aneityum, it is appropriate that the new taxon be named : Emoia aneityumensis n. sp. - the Aneityum skink HOLOTYPE. BM 1956.1.3.65, adult male, collected by Miss E. Cheesman on Aneityum island, New Hebrides, South West Pacific, 1954-55 ; for likely place and period of collection, see above. Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 54 LORD MEDWAY PARATYPES. BM 1956.1.3.63, adult male, and 1956.1.3.64, subadult female, also collected on Aneityum by Miss Cheesman during 1954-55 ; BM 1973.1534 (col- lectors' number RSNH 14-01), juvenile, apparently female, caught on 19 July 1971, BM 1973.1535 (RSNH 18-01), immature female, 21 July 1971, and BM 1973. 1536 (RSNH 19-03), juvenile male, 23 July 1971, all taken in disturbed and partially regenerated forest about i mile northeast of Anelgauhat, Aneityum, by Lord Medway and A. G. Marshall. DIAGNOSIS. In size and colour, and in the scalation of the head, similar to Emoia samoensis (Dumeril & Dumeril), differing in the greater number of longitudinal scale rows and the smaller number of subdigital lamellae. Similarly coloured to Emoia speiseri (Roux), but again differing in details of scalation, larger as a mature adult, and evidently producing a greater number of eggs at one time. DESCRIPTION. An Emoia , possessing the characters of that genus as defined by Gray (1845 : 95) and elaborated by Smith (1937). On the head, the prefrontals form a short medial suture, thus excluding the nasals from contact with the frontals. The interparietal is a separate shield, not fused to the parietals. The fifth or sixth upper labial is large, underlying the eye. The coloration of the holotype, after 18 years in alcohol, is dark brown on the upperparts, with a broken row of irregularly shaped black spots or flecks extending as an interrupted dorsolateral line from the ear orifice to the base of the tail, passing above the origins of the hindlimbs. The flanks are brown, barred and flecked by invasions of the buffy white ventral colour. In the three young animals, when freshly caught, the upperparts were grey-brown or olive-brown, marked with a broken row of irregular spots, flecks or blotches of black, intermixed with flecks of eau-de-nil, running from the ear to the base of the tail. The flanks were olive-brown, barred and flecked with the colour of the underparts, which were dirty white. The unbroken tail was the same brown as the ground colour of the upperparts, but regenerated portions were a lighter shade of brown, more or less unmarked. The specimen BM 1956.1.3.63 (again, after 18 years in preservative) is uniformly greyish brown above and buff below, without markings ; in my opinion, in life it could well have appeared 'smooth blue-grey' (cf. Cheesman 1957 : 325, quoted above). Details of scalation and measurements are given in Table i. By comparison, specimens of Emoia samoensis (two syntypes nos. 2764 & 7070 in the Paris museum, two from Samoa nos. 4632 & 4633 in the Basel museum, and two from Erromanga nos. 1860.3.18.8 & 1860.3.18.11 in the British Museum) have 32-36 scale rows at mid-body, 44-51 subdigital lamellae under the fourth toe, and measure snout-vent length 92-102 mm, total length 250-290 per cent. Twelve specimens of Emoia speiseri from all parts of this species' range in the New Hebrides have 28-34 scale rows at mid-body, 38-48 subdigital lamellae under the fourth toe, and measure 60-72 mm snouth-vent, total length about 270 per cent (cf. Roux, 1913). The female E. aneityumensis BM 1956.1.3.64 has four enlarged ova in the left ovary and at least one in the right, and BM 1973.1535 has four enlarging ova dis- cernible in the right ovary (left ovary not recognized). Of six undoubted female E. speiseri examined, two only had enlarging ova ; one had a single large follicle in A NEW SKINK FROM THE NEW HEBRIDES 55 the left ovary only, and the other had one large follicle in each ovary. Among E. samoensis, Roux (1913), has reported a gravid female containing four large ova or eggs ('gros oeufs'). There is no reason to believe that E. aneityumensis is confined to the vicinity of Anelgauhat on Aneityum. Retrospectively I attribute to this species a grey skink judged to be at least the size of the holotype (Table i), seen (but not caught) climbing the cliff above the sea-shore at Aegiptzav, on the north coast of the island, in July 1971. TABLE i Scale counts and measurements of the type series of Emoia aneityumensis Reg. no. Longitudinal Lamellae under Snout-vent Total length 1 BM scale rows fourth toe length (as % snout-vent) at mid-body (mm) Left Right 1956.1.3.652 40 42 42 92 (250) 1956.1.3.63 40 38 38 89 274 1956.1.3.64 40 36 36 83 (240) I973-I534 42 39 38 49 257 I973-I535 4 4 1 4 1 70 (203) I973-I536 4 1 38 39 66 (221) 1 Figures are given in parentheses if the tail showed signs of loss and subsequent regeneration. * Holotype. DISCUSSION The genus Emoia is represented in the New Hebrides by ten taxa (Medway & Marshall, in prep.), of which E. sanfordi, E. speiseri and E. aneityumensis are endemic to the archipelago. Only E. aneityumensis is confined, as far as known, to a single island. Of the two taxa most closely resembling E. aneityumensis, E. speiseri is sympatric with it on Aneityum island. Differences between the two, described above, establish that they are distinct species. E. samoensis, on the other hand, has been found in the New Hebrides only on Erromanga, some 180 km north-northwest of Aneityum and the next large island but one. The specimens (BM 1860.3.18.8 & 1860.3.18.11) were collected more than a century ago by Mr Cuming (Boulenger, 1887). In 1971 we did not find the species, but our stay on Erromanga was brief and collecting not intensive ; there are no grounds to doubt the record. The third of Cuming's speci- mens allocated to this species by Boulenger (BM 1860.3.18.12) is in fact E. sanfordi, an identification which confirms at least that the collection derived from the New Hebrides. Samples of E. samoensis from all parts of its range, from the Loyalty Islands to Fiji and the Samoan group, provide no evidence of geographical variation. Separa- tion of the Loyalty Islands population, described as a subspecies by Roux (1913), is not justified. The distinguishing character - adult size invariably less than that of examples from neighbouring archipelagoes ('archipels voisins') - was established by 56 LORD MEDWAY comparison with specimens from the New Hebrides (Roux 1913 : no). This material (in the Basel Museum of Natural History) was re-examined in the course of prepara- tion of this note. All New Hebrides specimens identified by Roux (1913 : 155) as Lygosoma (Emoa) samoense, a synonym of E. samoensis, prove to be E. sanfordi and thus provide a fallacious basis for comparison. In the scalation of the head, the number of longitudinal scale rows and of subdigital lamellae, specimens from the Loyalty Islands fall within the range of variation of E. samoensis from other parts of the South West Pacific. The absence of demonstrable geographical variation in E. samoensis over this wide area makes it unlikely that the species would differentiate markedly between Erromanga and Aneityum islands. E. samoensis and E. aneityumensis may replace each other ecologically on their respective islands in the New Hebrides, but there are no grounds for treating the two taxa as geographical races of one species. The characters that distinguish Emoia aneityumensis set the taxon apart from all other members of the genus, and it must be treated as a distinct species. ACKNOWLEDGEMENTS The Royal Society/Percy Sladen expedition was led by Dr K. E. Lee, on whom fell the chief burden of organization and administration. During field work, Dr A. G. Marshall was jointly responsible for the collection and curation of specimens ; we were assisted on Aneityum by Fred Boe, John Wycliffe and Hugo Tamata. The Royal Society subsequently provided a travel grant which enabled me to visit European museums containing collections of New Hebridean vertebrates ; for their kindness in making available specimens in their care and for other acts of hospitality, I am grateful to Professor J. Guibe, Professor U. Rahm and Dr K. Klemmer. All other work for this paper has been carried out at the British Museum (Natural History), and I am particularly grateful to the Curator of Herpetology, Miss A. G. C. Grandison, and staff for the facilities provided. Miss Grandison and Dr Marshall kindly read and criticized this paper in draft. REFERENCES ANGEL, F. 1935. Liste des reptiles recoltes par la Mission Aubert de la Rue aux Nouvelles Hebrides ou dans les iles voisines. Bull. Mus. Hist, nat., Paris (2) 7 : 54-56. BOULENGER, G. A. 1887. Catalogue of Lizards in the Collection of the British Museum. 2nd ed. London : British Museum. BURT, C. E. & BURT, M. D. 1932. Herpetological results of the Whitney South Sea expedition VI. Pacific island amphibians and reptiles in the collection of the American Museum of Natural Nistory. Bull. Am. Mus. nat. Hist. 63 : 462-597. CHEESMAN, E. 1957. Things Worth While. London : Hutchinson & Co. GRAY, J. E. 1845. Catalogue of Lizards in the Collection of the British Museum. London : British Museum. MEDWAY, LORD & MARSHALL, A. G. (in prep.). Observations on the vertebrate fauna of the New Hebrides. Roux, J. 1913. Les reptiles de la Nouvelle-CalMonie et des iles Loyalty. Appendice : Note sur quelques reptiles des Nouvelles Hebrides, des lies Banks et Santa Cruz. In : F. Sarasin & J. Roux (eds), Nova Caledonia, Zoologie 1 : 80-160, pis IV, V. A NEW SKINK FROM THE NEW HEBRIDES 57 SCHMIDT, K. P. & BURT, C. E. 1930. Herpetological results of the Whitney South Sea expedi- tion V. Description of Emoia sanfordi, a new lizard from the islands of the Western Pacific (Scincidae). Am. Mus. Novit. 436 : 1-3. SMITH, M. A. 1937. Review of the genus Lygosoma (Scincidae : Reptilia) and its allies. Rec. Indian Mus. 39 : 213-234. LORD MEDWAY GREAT GLEMHAM HOUSE SAXMUNDHAM SUFFOLK IPiy iLP SCUTOCYAMUS PARVUS, A NEW GENUS AND SPECIES OF WHALE-LOUSE (AMPHIPODA: CYAMIDAE) ECTOPARASITIC ON THE NORTH ATLANTIC WHITE-BEAKED DOLPHIN By ROGER J. LINCOLN & D. E. HURLEY 'The sea-lowse is an insect that is an enemy of all kinde of Whales, which by biting and tickling it puts into such a rage, that they are forced to run upon the sand, and hasten to dry land : I know nothing concerning the use of these creatures ; but I seriously exhort posterity to search out the use of them.' From : The Theater of Insects or Lesser Living Creatures Book n Chapter 38 p. 1126 by Tho. Mouffet, Doctor in Physick, London, 1658. INTRODUCTION It is rare to find a new species of cyamid, rarer still to discover a new genus. Since the first figure and description in the scientific literature of a 'Walfisches Lauss' by Martens in 1675, nineteen accepted species of cyamid have been described, only three of these in this century (Leung, 1967). Fifteen species are ascribed to the genus Cyamus and one each to Neocyamus, Platycyamus, Syncyamus and Isocyamus. Although the last two were established in 1955, only Sy ncyamus is based on a recently described species. Their rarity is not unexpected. Cyamids are host-specific to cetaceans which are a well-studied and numerically limited group. It is even more surprising, then, to discover in the collections of the British Museum (Natural History), unrecognized since their collection in 1933, specimens of a distinct and new species of cyamid from a common North Atlantic dolphin from which cyamids have never, to our knowledge, been recorded. These cyamids stood out for two reasons ; their generally small size in comparison with other species in the collection, and their distinctively rounded, saucer-shaped, low-profile outline which immediately suggested their novelty. These features indicated that their host would prove to be a fast-swimming species, which is indeed the case, and the subsequent examination has revealed further adaptation to life in an area of turbulent flow in the shape of the dactyls and cramp-iron teeth on the body and appendages. The cyamids were collected by Dr F. C. Fraser from a white-beaked dolphin, Lagenorhynchus albirostris Gray, and later presented to the Crustacea Section. The Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 60 R. J. LINCOLN & D. E. HURLEY dolphin was caught in the North Sea, off Peterhead in Aberdeenshire, on 21 July 1933. It was a young female measuring 5 ft 7! in. in length with a pronounced deformity in the curvature of the spine behind the dorsal fin (Fraser, pers. comm.). SCUTOCYAMUS gen. nov. DIAGNOSIS. Pereon segments 3-4 and 6-7 fused. Antenna i, 2-articulate ; antenna 2, i-articulate. Maxilla 2 with outer lobes absent. Maxillipeds fused and markedly reduced to form a small cleft flap. Pereopods i and 2 strongly dissimilar in shape and with i very much smaller than 2 ; pereopod 2 only 3-articulate. Male without accessory gills. Type species, Scutocyamus parvus sp. nov. Scutocyamus parvus sp. nov. (Text figs. la-g, 2a-h ; PI. la-e) DIAGNOSIS. With the characters of the genus given above. Body of small size, robust, strongly flattened dorsoventrally, head weakly immersed into anterior pereon segment. Pereopod i (Fig. ic) lacking unguis, dactylus armed with numerous small teeth (PI. le) ; pereopod 2 and pereopods 5-7 powerfully developed and held in characteristic posture giving a general oval symmetry to the shape of the animal (Fig. la ; PI. la). Gills simple, single. Pereon segments 5-7 each with a pair of large spines on the ventral surface. DESCRIPTION. Length of body from apex of head to end of pereon 1-7-2-4 mm in male and 2-6-3-1 mm in ovigerous female. Maximum width of body at the level of tergite 5 from 0-9 to 1-3 mm in male and from 1-4 to 1-8 mm in ovigerous female. No trace of pigmentation remaining in the alcohol preserved material. Pereon (Fig. la) somewhat oval in outline (pereon segment i fused with head), male rather more slender than female ; tergites 3-4 in female slightly shorter than other tergites, in male very much shorter and also much narrower than other tergites (Fig. ib) ; tergite 7 weakly immersed into pereon segment 6 (Fig. la, b) ; a pair of strong spines on the ventral side of segments 5-7. Head with sides convex, immersed slightly into anterior pereon segment, anterolateral angles expanded into large lobes giving a wide frontal margin to head ; eyes small, oval, mid-dorsal. Antenna i (Fig. 2c) small, 2-articulate, article 2 longer than i, apex with prominent group of sensory setae. Antenna 2 (Fig. 2d) extremely small, i-articulate, but constriction near tip gives impression of separate terminal article, apex with several large sensory setae. Upper lip (Fig. 2e) large, outer margin weakly concave and densely fringed with fine setae. Mandibles (Fig. 2g) with 2 incisor processes, the anterior of 1-2 teeth, the posterior incisor of several teeth ; right mandible with 2 penicils, left with only one ; molar process a rounded protuberance covered with very fine setae. Lower lip (Fig. 21) with inner lobes fused into a single elongate lobe, outer lobes slightly broader than inner, distal margins fringed with long setae. Maxilla i (Fig. 2h), apex with 3 pairs of curved spines each with 2-4 small teeth on inner margin ; palp i-articulate, reaching to about the apex of the outer lobe and with NEW GENUS AND SPECIES OF WHALE-LOUSE 61 g FIG. i. Scutocyamus parvus sp. nov. a, dorsal entire, female ; b, dorsal entire, male ; c, pereopod i ; d, pereopod 2, ventral ; e, pereopod 5, ventral ; f, pereopod 6, ventral ; g, pereopod 7, ventral ; bar scale a-b, i-o mm ; c, 0-2 mm ; d -g, 0-4 mm. 62 R. J. LINCOLN & D. E. HURLEY small group of sensory setae at tip. Maxilla 2 (Fig. 2b) a single elongate lobe with a small group of sensory setae at tip ; right and left maxilla 2 fused along mid-line. Maxillipeds (Fig. 2b) fused and reduced to a small cleft flap. Pereopod i (Fig. ic) extremely small, simple, 5-articulate, propodus elongate and about twice as long as wide ; dactylus broad with numerous combs of small teeth towards apex (PL le), unguis absent. Pereopod 2 (Fig. id) powerfully developed, only 3-articulate, proximal article large and robust with small marginal spine ; article 2 very broad and flattened, the outer margin with a deep indentation which probably marks the point of fusion of two articles ; article 2 with 2 small marginal spines ; dactylus robust strongly angled at its mid-point, and with a small but sharply pointed unguis. These cyamids have a quite characteristic posture with the expansive second pereopods held across the front of the head to form an effective shield. Pereopods 5-7 (Figs. le, f, g) powerful, 5-articulate, basal article (basis + ischium) short and stout with 3 well-developed ventral spines ; merus broad and flat with a single spine on distal margin ; carpus about equal to size of merus with a small spine on distal margin and an extremely large mid-ventral spine, the inner posterior angle produced into a small triangular process overlapping the dorsal surface of merus ; propodus elongate and extremely robust ; dactylus acutely angled and with sharply pointed unguis. Gills single, quite short, tapering somewhat, and held across ventral surface of pereon in forward direction ; accessory gills absent in male. Brood pouch rounded (Fig. 2a), containing only 7-10 eggs or young ; margins of oostegites fringed with many short setae ; genital valves well developed with inner margin also fringed with small setae. Pleon a small bilobed structure. MATERIAL EXAMINED. n$$ (ovigerous), 2-6-3-1 mm length, 1-4-1-8 mm width: 21$$ (immature), 2-0-2-5 mm length, 1-1-1-4 mm width : 28^, 1-7-2-4 mm length, 0-9-1-3 mm width : 37 juveniles. Holotype $ registration no. 1973 : 105 ; paratypes registration no. 1973 : 106, deposited in the collections of the British Museum (Natural History) . REMARKS. Scutocyamus parvus can be immediately recognized by the small body size, general symmetry given to the body posture by the expansive second pereopods, and by the fusion of pereon tergites 3-4 and tergites 6-7. Only one other species, Syncyamus pseudorcae Bowman, has a similar fusion of the pereon segments but it is restricted to tergites 6-7. While the amalgamation of tergites 3-4 is complete in Scutocyamus (PI. id) the posterior tergites retain a weak demarcation line which can be mistaken for a suture under a light microscope. Confirmation that pereon tergites 6-7 are in fact fused was obtained by examining material with a scanning electron microscope. The demarcation line was found to be a shallow depression which follows the line of fusion of the two segments. Of the five recognized genera of cyamids, Scutocyamus seems to be most closely allied to the monotypic Syncyamus. In addition to the fusion of the pereon tergites both have a similar reduction of the mouthparts, especially the reduction of the maxillipeds to a small flap, the fusion of the second maxillae, and the fusion of the inner lobes of the lower lip. Also, the absence of an unguis on the small pereopod I is a character shared by the genera. However, Scutocyamus retains a number of NEW GENUS AND SPECIES OF WHALE-LOUSE FIG. 2. Scutocyamus parvus sp. nov. a, ventral entire, female ; b, mouthparts, ventral ; c, antenna i ; d, antenna 2 ; e, upper lip ; f, lowerlip ; g, mandible with upper lip ; h, maxilla i ; bar scale a, i-o mm ; b-h, o-i mm. 64 R. J. LINCOLN & D. E. HURLEY unique characters such as the fusion of pereon tergites 3-4, extreme reduction of the antennae and pereopod i, a 3-articulate pereopod 2, and an absence of accessory gills in the male. The armature of comb-like teeth on the propodus of pereopod i has not previously been reported and appears to be an adaption to scrape the surface of the host. ACKNOWLEDGEMENTS The authors wish to thank Dr F. C. Fraser for his assistance with original data and for drawing attention to the opening quotation. The junior author wishes to thank the Trustees of the British Museum (Natural History) for facilities made available during Official Leave from New Zealand. REFERENCES BOWMAN, T. E. 1955. A new genus and species of whale-louse (Amphipoda, Cyamidae) from the false killer whale. Bull. mar. Sci. Gulf Car ibb. 5 (4) : 315-320. LEUNG, Y.-M. 1967. An illustrated key to the species of whale-lice (Amphipoda, Cyamidae), ectoparasites of Cetacea, with a guide to the literature. Crustaceana 12 (3) : 279-291. MARTENS, F. 1675. F. Martens. . . . Spitzbergische oder Groenlandische Reise Beschreibung gethan im Jahr 1671, etc. pp [viii] 132, 16 pis. Hamburg. Dr ROGER J. LINCOLN Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 Dr D. E. HURLEY NEW ZEALAND OCEANOGRAPHIC INSTITUTE P.O. Box 8009 WELLINGTON NEW ZEALAND It PLATE i Scutocyamus parvus sp. nov. a, entire ventral, male ( X 30) ; b, mouthparts, male (xi8o) ; c, pleon ventral, male ( X 120) ; d, fusion pereon tergites 3-4, dorsal female ( x 120) ; e, pereopod i, apex of propodus, male (x i-i k). Scanning electron microscope ; gold coating ; 20 kv. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i CATALOGUE OF THE WHALE-LICE (CRUSTACEA : AMPHIPODA : CYAMIDAE) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) By ROGER J. LINCOLN & D. E. HURLEY INTRODUCTION THE collections of the British Museum (Natural History), containing as they do much of the material taken during the Discovery Investigations (1925-27) on the biology of the southern whales and much incidentally collected Northern Hemisphere material, represent one of the more significant collections of cyamid material. Numerically this collection is not large but it contains some types and material which has been little studied. Furthermore the group is usually very poorly repre- sented in museum collections. The discovery of a new genus and species of cyamid (Lincoln & Hurley, 1974) has been one of the benefits resulting from a re-examination of the material in the collections of the Crustacea Section. Since all identifications have now been verified and labelling updated following the nomenclature given in the recent cyamid synopsis of Leung (1967), this is a suitable opportunity to present a published catalogue of holdings for the benefit of other workers. The nomenclature for the whales follows that used by Hershkovitz (1966). The assistance of Miss J. Ellis in collating the information is gratefully acknow- ledged. CYAMUS Latreille, 1796 Cyamus abbreviatus Say, 1818 (i) Type ? i specimen (dry collection). North America. Leach Collection. Cyamus balaenopterae Barnard, 1931 (1) Syntypes, 6^, 4$$ ; Ann. Mag. nat. Hist. (10) 7 : 425-430. Reg. no. 1936: 11:2:3495-3501. Saldanha Bay, South Africa ; on Balaenoptera physalis (L.) [blue whale according to label] whale no. W 1096. Discovery Collection. (2) 2&J, 8$$, i juv. Reg. no. 1936:11:2:3502-3531. Saldanha Bay, South Africa ; on Balaenoptera musculus (L.) whale no. W 867. Discovery Collection. (3) 9&?> 6$$. Reg. no. 1936:11:2:3502-3531. Durban, South Africa ; on Balaenoptera physalus (L.) whale no. W Dig. Discovery Collection. (4) *(?, 5?? (immature). Reg. no. 1936:11:2:3502-3531. Saldanha Bay, South Africa, 18.8.1926 ; on Balaenoptera musculus (L.) whale no. W 961. Discovery Collection. Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 66 R. J. LINCOLN & D. E. HURLEY (5) 3 6$$, i? ovig., i juv. Reg. no. 1936:11:2:3502-3531. Saldanha Bay, South Africa, 15.8.1926 ; on Balaenoptera physalus (L.) whale no. W 948. Discovery Collection. (6) 20(3$, 15??, 10 juvs. Reg. no. 1973:104:45. No locality data, 24.10.1948 ; on Balaenoptera physalus (L.). Discovery Collection. Cyamus boopis Lutken, 1870 (1) i$. Reg. no. 1853:68. Greenland. Collected by Holboll. (2) 10(3$, 10 juvs. Reg. no. 1910:8:9:1-6. Bransfield Straits, South Shetlands ; on Megaptera novaeangliae (Borowski). Collected by J. A. Morch. (3) 2 c?$> J ? (immature). Reg. no. 1911:11:8:23064-066. No locality data. A. M. Norman Collection. (4) 3$$. Reg. no. 1920:7:5:61-62. Deception Island, South Shetlands, 1914 ; on Megaptera novaeangliae (Borowski). Collected by A. G. Bennett. (5) iJc&J- Reg- no - 1924:7:10:1-5. Olna Firth, Shetland Islands, 19.6.1924 ; on Megaptera novaeangliae (Borowski). Collected by F. V. Morley. (6) 3(3$, 3?$ (immature), 2 juvs. Reg. no. 1928:12:1:2966-71. Durban, South Africa, 23.7.1908 ; on Megaptera novaeangliae (Borowski). T. R. R. Stebbing Collection. (7) i$, 2?? (all immature). Reg. no. 1936:11:2:3542-3581. South Georgia ; on Megaptera novaeangliae (Borowski) whale no. 355. Discovery Collection. (8) 6(3$, i$. Reg. no. 1936:11:2:3542-3581. South Georgia ; on Megaptera novaeangliae (Borowski), whale no. 373, genital region. Discovery Collection. (9) 2(&J, 2$$, 5 juvs. Reg. no. 1936:11:2:3542-3581. South Georgia ; on Megaptera novaeangliae (Borowski), whale no. 387. Discovery Collection. (10) 3(3$, 3??, 10 juvs. Reg. no. 1936:11:2:3542-3581. Deception Island, South Shetlands ; on Megaptera novaeangliae (Borowski). Discovery Collection. (n) 6$$, 4$?, 10 juvs. Reg. no. 1936:11:2:3542-3581. Saldanha Bay, South Africa, 29.9.1926 ; on Megaptera novaeangliae (Borowski), whale no. 1125. Discovery Collection. (12) ii(3$, i?. Reg. no. 1936:11:2:3542-3581. Durban, South Africa ; on Megaptera novaeangliae (Borowski), whale no. 03. Discovery Collection. (13) 2$$, 4??. Reg. no. 1936:11:2:3542-3581. Durban, South Africa, on Physeter catodon L., whale no. Dai. Discovery Collection. (14) 10$$, 7$$, 2 juvs. Reg. no. 1936:11:2:3542-3581. Saldanha Bay, South Africa, 6.8.1926 ; on Megaptera novaeangliae (Borowski), whale no. 918. Discovery Collection. (15) i6(3$, 9$$, 10 juvs. Reg. no. 1965:9:8:1-15. Paita, Peru, 05^5' S, 8iio' W, 19.10.1960, on Megaptera novaeangliae (Borowski), $ 13-3 m, whale no. PAi. Collected by Robert Clarke. (16) 2<&J. Reg. no. 1972:556:2. Varanger Fiord, Norway ; on Megaptera novae- angliae (Borowski). G. O. Sars Collection. (17) 50(3$, io$$ (immature). Reg. no. 1972:568:60. Off Russell Bay, Bay of Islands, New Zealand, August 1912 ; on Megaptera novaeangliae (Borowski). Terra Nova Collection. CATALOGUE OF WHALE-LICE 67 (18) 150(2(2??. Reg. no. 1972:569:150. Off Russell Bay, Bay of Islands, New Zealand, August 1912 ; on Megaptera novaeangliae (Borowski). Terra Nova Collection. (19) 14^, 6??. Reg. no. 1972:577:20. No locality data. No collection data. (20) 9(2(2, 6?? (immature), i juv. Reg. no. 1972:579:16. St. Vicente. Collected by R. T. Lowe. Cyamus catodontis Margolis, 1954 (1) 300(2(2?? (mature and immature). Reg. no. 1965:9:8:16-45. Paita, Peru 0505' S, 8iio' W, 30.9.1960 ; on Physeter catodon L., <$ 13-2 m, whale no- Pa870. Collected by R. Clarke. (2) 100(2(2??. Reg. no. 1965:9:8:16-45. Pisco, Peru, I347' S, 76i5' W, 17.10. 1960 ; on Physeter catodon L., <$ 13-6 m, whale no. Pi59- Collected by R. Clarke. (3) I 5^?? (mature and immature). Reg. no. 1965:9:8:16-45. Pisco, Peru, I347' S, 76i5' W, 17.1.1961 ; on Physeter catodon L., ^ 15-2 m, whale no. Pii6o. Collected by R. Clarke. (4) 150(2(2, 150$$. Reg. no. 1973:103:300. No locality data ; on Physeter catodon L., (2 54 ft, whale no. 20. Discovery Collection, 1951-52. Cyamus ceti (L., 1758) Lamark, 1801 (1) i(2, i$. Reg. no. 1911:11:8:23075-75. No locality data ; A. M. Norman Collection. (2) 3<2c?, 8??- Reg. no. 1928:12:1:2978-2984. Greenland ; on Balaena mysticetus L. T. R. R. Stebbing Collection. (3) 6^(2, 3??- Reg. no. 1956:10:10:340-343. Arctic Sea, 1893. University College, Dundee Collection (S.S. Eclipse). (4) 300(2(2, 200$$, 50?$ (immature), 50 juvs. Reg. no. 1972:556-600. North Europe ? No collection data. (5) 2(2c2, 4??. Reg. no. 1972:588:6. Murray Firth, Scotland. Leach Collection. (6) 6(2(2, *?, 6 juvs. Reg. no. 1972:559:13. No locality data. No collection data. (7) 9cft2> 10??. Reg. no. 1972:567:19. No locality data. No collection data. (8) 10(2(2, io$$, 20 juvs. Reg. no. 1972:578:40. Baffin Bay, 20.7.1894. University College, Dundee Collection (S.S. Eclipse}. (9) Reg. no. 6o4a, b, d, c, dry collection (Leach cab : 31). Britain. Leach Collection. Cyamus erraticus Roussel de Vauzeme, 1834 ( x ) 5c&2, 4?? (immature). Reg. no. 1912:10:2:1-7. 6 miles N.E. of Flugga, Shetland. Collected by Lovett. (2) 4(2(2, i$. Reg. no. 1928:12:1:2966-71. Durban, South Africa, 23.7.1908 ; on Eubalaena. T. R. R. Stebbing Collection. (3) 8(2(2, 10??. Reg. no. 1936:11:2:3532-3541. Saldanha Bay, South Africa, 23.7.1908 ; on Eubalaena glacialis australis Desmoulins, whale no. 1020. Discovery Collection. 68 R. J. LINCOLN & D. E. HURLEY (4) 2c&? 5??i i? (immature). Reg. no. 1972:560:8. Rockall, North Atlantic, 1.7.1914 ; on flipper of Eubalaena glacialis glacialis Miiller. No collection data. (5) I4cc?, 3$9. Reg. no. 1972:570:17. No locality data. Albany Museum, Grahamstown Collection. (6) !<. Reg. no. 1972:575:1. No locality data ; on Eubalaena glacialis glacialis Miiller. Collected by Professor Goldberg. Cyamus gracilis (Roussel de Vauzeme, 1834) (1) i<. Reg. no. 1907:12:2:218. Pacific Ocean. University College, Dundee Collection. (2) i$, i$ (immature). Reg. no. 1911:11:8:23069-70. No locality data. A. M. Norman Collection. (3) 8&J (immature), 4$$. Reg. no. 1928:12:1:2972-74. False Bay, Cape, South Africa. T. R. R. Stebbing Collection. (4) i$, 6 juvs. Reg. no. 1936:11:2:3582-3584. South Georgia ; on Eubalaena glacialis australis Desmoulins, whale no. 503. Discovery Collection. (5) Reg. no. 6o5a (dry collection), Leach cab : 31. British Sea. Leach Collection. Cyamus monodontis Liitken, 1873 (1) i specimen (slide collection) . No locality data ; on Monodon monoceros L. No collection data. (2) 3$$, 10 juvs. Reg. no. 1928:12:1:2975-77. No locality data ; on Monodon monoceros L. T. R. R. Stebbing Collection. (3) 2<<, 2$$. Reg. no. 1911:11:8:23071-74. No locality data ; on Monodon monoceros L. A. M. Norman Collection. (4) 5^, 5$$, 10 juvs. Reg. no. 1956:10:10:344-346. Davis Straits, Greenland, Canada ; on Monodon monoceros L. University College, Dundee Collection. (5) i$. Reg. no. 1972:561:1. No locality data. University College, Dundee Collection. (6) i$. Reg. no. 1972:562:1. No locality data ; on Monodon monoceros L. Collected by Nuttall. (7) ioo<^$$. Reg. no. 1972:563:100. Upernavik, Greenland, 1892. University College, Dundee Collection. Cyamus nodosus Liitken, 1860 (1) 4$$, 2$$ (immature), 14 juvs. Reg. no. 1907:12:2:212-217. Davis Straits, Greenland, Canada. University College, Dundee Collection. (2) 200^$$. Reg. no. 1879:26. Omenak, Greenland ; on Monodon monoceros L. Collected by E. Whymper. (3) 2(J(J, i?, i juv. Reg. no. 1911:11:8:23031-33. Omenak, Greenland. A. M. Norman Collection. (4) 2(, 4$?, 8$$ (immature). Reg. no. 1972:564:12. No locality data ; on Monodon monoceros L. University College, Dundee Collection. CATALOGUE OF WHALE-LICE 69 (5) i$. Reg. no. 1972:565:1. No locality data ; on Monodon monoceros L. Col- lected by Nuttall. (6) 5cJcJ, 3?$ (immature), 9 juvs. Reg. no. 1972:566:17. No locality data. University College, Dundee Collection. (7) 3 specimens (slide collection). No locality data ; on Monodon monoceros L. No collection data. Cyamus ovalis Roussel de Vauzeme, 1834 (1) 3c?c??$> attached to section of skin. Reg. no. 1911:10:16:1-10. South Georgia, 2.2.1911 ; on Eubalaena glacialis Muller. Collected by G. Cruick- shank. (2) I50&J??, attached to section of skin from lower maxilla. Reg. no. 1911:10:16: 11-20. South Georgia, 15.3.1911 ; on Eubalaena glacialis Muller. Collected by G. Cruikshank. (3) I0c, 20 juvs. Reg. no. 1911:11:8:23034-53. Iceland ; on Eubalaena glacialis glacialis Muller. A. M. Norman Collection. (4) 2<3cT, 3?$, 30 juvs. Reg. no. 1911:11:8:23054-23063. Pacific Ocean ; on Eubalaena glacialis japonica Lacepede. A. M. Norman Collection. (5) 1503$$?, attached to section of skin. Reg. no. 1912:6:15:1-20. South Georgia ; on Eubalaena glacialis australis Desmoulins. Collected by G. Cruikshank. (6) 6<^, 8$$, 30 juvs. Reg. no. 1912:10:2:8-27. 6 miles N.E. of Flugga, Shetland. Collected by Lovett. (7) 3 00 dc?$$> attached to three separate sections of skin. Reg. no. 1920:10:21:23- 42. Stromness Whaling Station, South Georgia ; on right whale (upper jaw). Collected by J. Rasmussen and Alex Lange. (8) 7dtf, 7?$, 6 juvs. Reg. no. 1936:11:2:3483-3494. South Georgia ; on Eubalaena glacialis australis Desmoulins, whale no. 503. Discovery Collection. (9) i$, 2$$, i juv. Reg. no. 1936:11:2:3483-3494. Saldanha Bay, South Africa, 26.8.1926 ; on Eubalaena glacialis australis Desmoulins, whale no. 1020. Discovery Collection. (10) 10^, 30$$, 100 juvs. Reg. no. 1972:571:140. Port Elizabeth. Collected by I. L. Drege. ( 11 ) 3<3<$> 20 juvs. Reg. no. 1972:572:23. Faeroerne Island, North Atlantic, 62 N, 07 W, June 1898 ; on Eubalaena glacialis glacialis Muller, $. No collection data. (12) 2<&, 2$$. Reg. no. 1972:573:4. No locality data ; on Eubalaena glacialis japonica Lacepede. Collected by W. N. Lockington. ( 1 3) 7c&> IO ?? 3 juvs. Reg. no. 1972:574:47. South Georgia ; from 'bonnet' of right whale. Collected by P. Stammwitz. (14) 3^, 9$9, 10 juvs. Reg. no. 1972:576:22. No locality data ; on Eubalaena glacialis glacialis Muller. Collected by Goldberg. (15) 20 specimens (dry collection). Port Jackson and South Africa. Leach Collection. yo R. J. LINCOLN & D. E. HURLEY Cyamus pacificus Liitken, 1873 ( J ) 5cTc?- Reg- n - I&77-3' No locality data ; on Megaptera novaeangliae (Borow- ski). Collected by W. N. Lockington. Cyamus scammoni Ball, 1872 ( J ) 3&? !? Reg- n - X 877:3. No locality data ; on Eschrichtius gibbosus (Erxleben). Collected by W. N. Lockington. (2) 30^$?. attached to section of skin. Reg. no. 1891:2:10:1-12. Wladiwostock, Victoria Bay, Sea of Japan. Collected by Fridolf Hook. Cyamus thomsoni Gosse, 1855 see Platycyamus ISOCYAMUS Gervais & Van Beneden, 1859 Isocyamus delphini (Guerin-Meneville, 1836) (1) 2<3c?. Reg. no. 1911:11:8:23067-68. No locality data ; on Globicephala melaena Traill (pilot whale). A. M. Norman Collection. (2) 4&?> 5??, 6 juvs. Reg. no. 1936:11:2:3585-3600. i445' N, i834'W, 27.10. 1925 ; on Steno bredanensis Lesson (rough-toothed dolphin), Dolphin A. Discovery Collection. (3) i(J, i$, ii juvs. Reg. no. 1936:11:2:3583-3600. 14^5' N, i834' W, 27.10. 1925 ; on Steno bredanensis Lesson, Dolphin B. Discovery Collection. (4) 9c?c??- Reg. no. 1972:547:9. Tonnybwlch Beach, Aberystwyth, Cardigan- shire ; on stranded porpoise. Collected by P. Miles. NEOCYAMUS Margolis, 1955 Neocyamus physeteris (Pouchet, 1888) (1) 4$$. Reg. no. 1965:9:8:46-65. Talcahuano, Chile, 3645' S, 73i2' W, 23.3.1961, on Physeter catodon L., $ 10-1 m, whale no. T6. Collected by R. Clarke. (2) isoni (Gosse, 1855) (i) i<$. Reg. no. 1907:12:2:219. No locality data. University College, Dundee Collection. CATALOGUE OF WHALE-LICE 71 (2) i(, i$. Reg. no. 1911:11:8:23077-78. No locality data ; on Hyperoodon ampullatus Forster. A. M. Norman Collection. (3) i$. Reg. no. 1956:10:10:347. No locality data. Collected by University College, Dundee. (4) Holotype. Ann. Mag. not. Hist. (2) 16 : 30-31. Reg. no. 1856:131 dry coll., Leach cab : 31. Portland Roads S. England ; taken from beneath the eye of stranded Bottlenose whale, Hyperoodon ampullatus Forster. Leach Collection. SCUTOCYAMUS Lincoln & Hurley, 1974 Scutocyamus parvus Lincoln & Hurley, 1974 (i) Holotype ?, io$$ ovig., 21$$ immature, 28$$, 37 juvs. Reg. no. 1973:105. Off Peterhead, Aberdeenshire, North Sea, 21.7.1933 ; on Lagenorhynchus albirostris Gray (white-beaked dolphin). Collected by Dr F. C. Fraser. Summary of hosts of whale-lice in the collections Host Mysticeti (whalebone whales) Gray whale, Eschrichtius gibbosus (Erxleben) Blue whale, Balaenoptera musculus (L.) Fin whale, Balaenoptera physalis (L.) Humpback whale, Megaptera novaeangliae (Borowski) North Pacific right whale, Eubalaena glacialis japonica Lacepede Southern right whale, Eubalaena glacialis australis Desmoulins North Atlantic right whale, Eubalaena glacialis glacialis Miiller Greenland right whale, Balaena mysticeti L. Odontoceti (toothed whales) Sperm whale, Physeter catodon L. Bottlenose whale, Hyperoodon ampullatus Forster Pilot whale, Globicephala melaena Traill Narwhal, Monodon monoceros L. Whale-lice Cyamus scammoni Dall Cyamus balaenopterae Barnard Cyamus balaenopterae Barnard Cyamus boopis Liitken Cyamus pacificus Lutken Cyamus ovalis R. de Vauzeme Cyamus erraticus R. de Vauzeme Cyamus gracilis (R. de Vauzeme) Cyamus ovalis R. de Vauzeme Cyamus erraticus R. de Vauzeme Cyamus ovalis R. de Vauzeme Cyamus ceti (L.) Cyamus boopis Lutken Cyamus catodontis Margolis Neocyamus physeteris (Pouchet) Platycyamus thompsoni (Gosse) Isocyamus delphini (Guerin-Meneville) Cyamus monodontis Lutken Cyamus nodosus L. Scutocyamus parvus Lincoln & Hurley White-beaked dolphin, Lagenorhynchus al- birostris Gray Rough-toothed dolphin, Steno bredanensis Isocyamus delphini (Guerin-Me'neville) Lesson REFERENCES HERSHKOVITZ, P. 1966. Catalog of living whales. Bull. U.S. natn. Mus. 246 : 1-259. LEUNG, Y.-K. 1967. An illustrated key to the species of whale-lice (Amphipoda, Cyamidae), ecto-parasites of Cetacea, with a guide to the literature. Crustaceana 12 (3) : 279-291. 72 R. J. LINCOLN & D. E. HURLEY LINCOLN, R. J. & HURLEY, D. E. 1974. Scutocyamus parvus, a new genus and species of whale- louse (Amphipoda : Cyamidae) ectoparasitic on the North Atlantic white-beaked dolphin. Bull. Br. Mus. nat. Hist. (Zool.) 26 : 59-64. Dr ROGER J. LINCOLN Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD Dr D. E. HURLEY NEW ZEALAND OCEANOGRAPHIC INSTITUTE P.O. Box 8009 WELLINGTON NEW ZEALAND A REVIEW OF LAEPHOTIS THOMAS, 1901 (CHIROPTERA : VESPERTILIONIDAE) By J. E. HILL INTRODUCTION IN recent years the African genus Laephotis Thomas, 1901 has been considered monotypic (e.g. Ellerman, Morrison-Scott & Hayman, 1953 : 78) with one subspecies, L. wintoni wintoni Thomas, 1901 in Kenya and a second, L. w. angolensis Monard, 1935 in Angola, Zambia, Botswana and the southern part of Zaire. However, Peterson (1971) while reporting a further example from Kenya reviewed much of the limited material of Laephotis available in collections and was led to suggest that wintoni and angolensis might represent distinct species, with the possibility that a third undescribed taxon might occur in Zambia. Setzer (1971) treated wintoni and angolensis as specifically distinct (as had Monard when describing the latter) and described two further species, namibensis from the Namib Desert, South West Africa, and botswanae from Botswana, Zambia and southern Zaire. Although this author referred to botswanae a number of the specimens from Zambia and southern Zaire hitherto allocated to angolensis, he did not have available all of the material so reported : in particular he did not examine the specimen (in the Harrison Zoological Museum, Sevenoaks, England) from Ndola, Zambia, which Peterson (1971 : 888) then thought could represent either the then unknown female of wintoni or possibly an unnamed taxon. More recently, Peterson (1973 : 602), in describing the first known female of wintoni has suggested that the specimen from Ndola may be more closely related to botswanae than to other specimens from Zambia or Zaire. Current accessions to the collections of the British Museum (Natural History) have included three female specimens of L. wintoni from Ethiopia, whence the genus has been hitherto unreported. These have led to a further examination of the seven examples of Laephotis already in the collections in London, together with six from Zaire in the Musee Royal de 1'Afrique Centrale, Tervuren, Belgium, and of two specimens in the Harrison Zoological Museum, one the apparently enigmatic example from Ndola. The four species recognized by Setzer (1971) are retained : wintoni and namibensis are apparently very similar but differ much more sharply from angolensis and botswanae than do these from each other. SYSTEMATIC SECTION LAEPHO TIS Thomas, 1901 Laephotis Thomas, 1901 : 460. TYPE SPECIES. Laephotis wintoni Thomas, 1901. Head low, flat-crowned ; muzzle broad, flat, the nostrils opening sublaterally ; ears moderate to large, when laid forward reaching to or beyond the tip of the Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 74 J- E. HILL muzzle, more or less triangular in outline ; anterior margin of the ear originating near the centre of the forehead, lacking any posteriorly directed basal lobe but with a distinct basal fold ; anterior margin convex proximally then less so or nearly straight ; tip rounded ; posterior margin of ear slightly emarginated just below tip, otherwise convex, with moderate rounded or semi-circular antitragal lobe, terminat- ing just behind the angle of the mouth. Tragus (Fig. i) as a rule rather wide, about one-third the length of the ear, its anterior margin slightly concave, tip bluntly pointed, anteriorly directed ; posterior margin of tragus convex or angular, a small triangular lobe at its base. Calcar extending along about two-thirds of the uropata- gial margin, with small, rounded post-calcarial lobe. Seven palate ridges in L. wintoni from Ethiopia, the first and second uninterrupted, the last reduced but also complete (Fig. 2) : however, Peterson (1973 : 602, fig. i) reported six ridges only in a subadult female of L. wintoni from Kenya. Skull with slightly flattened braincase ; slight lambdoid but no sagittal crest ; rostrum rather narrow, unexpanded, with little supraorbital inflation ; a shallow, median rostral depression ; zygomata slender ; narial emargination narrow, broadly U-shaped but slightly angular, extending posteriorly to or a little beyond a line joining the anterior margins of the anteorbital foramina ; pre-palatal emargination extending laterally a little beyond the inner faces of i 2 ~ 2 and posteriorly almost to a line joining the posterior faces of c 1 " 1 ; palate long, rather narrow, strongly domed ; maxillary toothrows usually almost parallel ; post-palatal extension moderate to long, its length from a line across the rear faces of m 3 ~ 3 to the tips of the pterygoid hamulars approaching or equal to the length of the palate from the same line to the back of the pre-palatal emargination (Table 2) ; short, broad post-palatal spine ; pterygoid hamulars strong, sharply deflected inwards ; bullae inflated, large. 21 I ^ Dentition i -, c -, pm -, m - = 32. Inner upper incisor (i 2 ) long, with secondary 3i 23 postero-internal cusp extending almost to its tip ; i 3 small, a little wider than long, its main cusp barely extending above the cingulum of i 2 , with small internal basal cusp, in toothrow, touching i 2 , separated from c 1 by a narrow space ; c 1 normal ; pm 4 a little wider than long, in contact with c 1 ; m 3 not reduced, with meta- cone and three commissures ; ij_ 3 tricuspid, imbricated, i 3 slightly the largest ; c t reduced, weak, little higher than pm 4 ; pm 2 reduced, about one-half the height and one-quarter or a little more the crown area of pm 4 , tightly compressed in tooth- row ; m 3 little reduced, the hypoconid and entoconid well developed. The genus occurs in Ethiopia and Kenya, in southern Zaire, Zambia, Botswana, South West Africa and in Angola. It appears to be related to Eptesicus. Laephotis wintoni Thomas, 1901 Laephotis wintoni Thomas, 1901 : 460. Kitui, Kenya, c. 3500 ft. Size large for the genus (length of forearm 36-41 mm, condylobasal length 15-2-15-8 mm) with very large ears (length from meatus 21 mm or more) ; tragus broad, at its widest at about one-third of its height from its base, its posterior margin strongly convex, sometimes slightly angular (Fig. la). Dorsal surface a shade of REVIEW OF LAEPHOTIS THOMAS, 1901 75 mid-brown, the pelage dark based with brown, slightly coppery tips ; ventral surface similar but paler, the hairs similarly dark based but with paler brown or, especially posteriorly, with buffy white tips. Skull elongate with long narrow rostrum ; palate long, the maxillary toothrows nearly parallel ; post-palatal extension very long, its length from a line across the rear faces of m 3 " 3 to the tips of the pterygoid hamulars equal or nearly equal to the length of the palate from the same line to the back of the pre-palatal emargination ; bony post-palate long, its length from a line across the rear faces of m 3 ~ 3 to the anterior edge of the mesopterygoid fossa exceed- ing the distance from the anterior edge of the mesopterygoid fossa to the tips of the pterygoid hamulars (Table 2). The female of wintoni has remained unknown until recently. Peterson (1971 : 888) speculated that a female specimen (1.2533) m the Harrison Zoological Museum, Sevenoaks, from Ndola, Zambia, considerably smaller than the male examples that he had examined, might represent it. Setzer (1971 : 264) noted that data presented by Peterson (p. 886) for specimens of Laephotis in the British Museum (Natural History) from Zaire reveal a slight sexual variation, females being on the average a little larger than males in external and cranial measurements. However, the skull had not then been extracted from the sole male specimen (B.M. 57.435, in alcohol) in this series (B.M. 57.435-438) which apparently represents two species (vide infra). The measurements given by Peterson show a female (B.M. 55.1135) from Zambia to be externally a little larger on the whole than a male (B.M. 55.1134) obtained at the same locality but so only in a limited number of cranial dimensions. Peterson (1973 : 601) has reported a subadult female of wintoni from Kenya which is comparable to male specimens in most dimensions. The three female specimens of wintoni now available from Ethiopia are consistently larger in most respects than two males from Kenya (Table i) or than the male and subadult female specimens reported from Kenya by Peterson (1971 : 885, 886 ; 1973 : 601). The number of specimens available is too small to establish any firm evidence of sexual variation in size, and the possibility remains that the population of wintoni in Ethiopia consists of larger individuals than does the population in Kenya. The species occurs (Fig. 3) in KENYA (Thomas, 1901 : 460 ; Harrison, 1961 : 292 ; Hayman & Hill, 1971 : 49 ; Peterson, 1971 : 885, 887, fig. i (head), pi. i (fig. 2) (skull) ; Setzer, 1971 : 262, fig. ic (tragus)) ; Peterson, 1973 : 601, fig. i (soft palate), and in ETHIOPIA (B.M. 72.4397-4399, from Koka, Shoa Province, 827' N, 39o6' E, at 1700 m). Laephotis namibensis Setzer, 1971 Laephotis namibensis Setzer, 1971 : 259, 263, fig. id (tragus). Kuiseb River, near Namib Desert Research Station, Gobabeb, South West Africa. No specimens are available for examination, the species being represented at present only by the female holotype and paratype in the collections of the United States National Museum of Natural History at the Smithsonian Institution, Washing- ton. From the description namibensis is characterized by its very large ears ; well- developed tragus and antitragus ; pale coloration ; large, relatively long and narrow 76 J. E. HILL skull ; relatively long, narrow palate ; and by great inflation of the bullae. It is evidently very similar to wintoni but has larger ears (length 24-25 mm) which are broader at the base, and a larger tragus. It is markedly paler in colour than wintoni, the dorsal surface pale drab, the ventral surface paler, the hairs tipped with white. Cranially, the braincase is more domed than in wintoni, the postorbital region more constricted, the zygomatic arches more arcuate in the vertical plane, the maxillary toothrows more nearly parallel and the bullae more inflated. Although the skull is rather longer than in male specimens of wintoni from Kenya, it is only marginally longer than in female specimens from Ethiopia (Table i). Laephotis angolensis Monard, 1935 Laephotis angolensis Monard, 1935 : 45- Tyihumbwe" (Chiumbwe River, a tributary of the Kasai, 15 km west of Dala), Angola. Smaller than wintoni or namibensis (length of forearm 32-35 mm, condylobasal length c. 12-9 mm) with smaller, narrower ears (length from meatus less than 16 mm) ; tragus (Fig. ib) smaller than in wintoni or namibensis (vide Setzer, 1971 : 263, fig. i), less broadened and rather more spatulate. Colour apparently similar to that of wintoni but the only available specimens are in alcohol and have been so for some years. According to Setzer (1971 : 260) namibensis is paler than angolensis. Skull very much smaller than in wintoni or namibensis (Table i), the bony part of the post- palatal extension much shorter than in wintoni, its length from a line across the rear faces of m 3 ~ 3 to the anterior edge of the mesopterygoid fossa less than the distance from the anterior edge of the mesopterygoid fossa to the tips of the pterygoid hamulars, not exceeding it as in that species (Table 2). A number of specimens have been allocated hitherto (Hayman & Hill, 1971 : 49) to angolensis. These include a further example from Angola, from a locality 35 miles east of Dande, in the collections of the American Museum of Natural History, New York (Hill & Carter, 1941 : 49) which is accepted as representative of angolensis by Setzer (1971 : 260 et seq.j. Specimens from Zambia, Zaire and Botswana were regarded previously as angolensis but Setzer (1971 : 260) took the sole example from Botswana as the holotype of a new species, botswanae, to which he referred such of the material from Zambia and Zaire as he was able to examine. Two (B.M. 55.1134- 1135) of the three specimens so far reported from Zambia were seen by Setzer : the third is the example (Harrison Zoological Museum 1.2533) discussed by Peterson (1971 : 885, 888 ; 1973 : 602) which apparently also represents botswanae (vide infra). The specimens (B.M. 57.436, 57.438) from Zaire seen by Setzer are from a series of ten collected at Mumene, 70 km east of Lumbumbashi ( = Elizabeth ville), Katanga, which, with a further three from the nearby locality of Musonge, 2 km to the west, were originally reported by Hayman (1957 : 43) . This author examined ten of these ; four (B.M. 57.436-438 from Mumene, B.M. 57.435 from Musonge) are in the collec- tions of the British Museum (Natural History) and a further six (M.R.A.C. 26.402- 406 from Mumene, M.R.A.C. 26.407 from Musonge) in the Musee Royale de 1'Afrique Centrale, Tervuren. Hayman, Misonne & Verheyen (1966 : 50) list nine specimens REVIEW OF LAEPHOTIS THOMAS, 1901 77 at Tervuren but no more than six can be found (Thys van den Audenaerde, in litt.), corresponding to the total examined in London by Hayman. Of these, eight must be referred to botswanae but two, B.M. 57.435 and B.M. 57.437 apparently represent not botswanae but angolensis as it is understood by Setzer (1971 : 260 et seq.). They agree closely with the descriptions of this species by Monard (1935 : 45) and Hill & Carter (1941 : 49) : their measurements (Table i) are similar to those of the holotype and to those of the second Angolan example as they are reported by Hill & Carter (p. 176) and Setzer (1971 : 261). The specimens differ from the others in the series from Zaire in smaller size, narrower tragus with generally a less angular posterior margin, and in slightly shorter, narrower rostrum and palate. The tragus of B.M. 57.435 is a little wider than that of B.M. 57.437 while among botswanae the tragus of M.R.A.C. 26.404 is rather narrow, with its posterior margin a little less angular than is usual in that series. Setzer (1971 : 262) notes that botswanae has a more massive rostrum than angolensis but the rostrum in B.M. 57.435 and B.M. 57.437 is no more than slightly shorter and not less massive when com- pared with the narrowest of botswanae. The rearward extension of the occipital crests, said by Setzer to be less in botswanae, does not differ consistently in the speci- mens examined. The toothrows of B.M. 57.435 are more divergent posteriorly than those of B.M. 57.437 to approach specimens referred to botswanae although, as Setzer notes of this species, its toothrows are generally more divergent posteriorly than in angolensis. The specimens available do not confirm the statement by Setzer that the bullae in botswanae are relatively as well as actually smaller than in angolensis : if B.M. 57.435 and B.M. 57.437 correctly represent angolensis then the bullae of bots- wanae are proportionately about the same in size and in actual terms a little larger. That angolensis and botswanae might occur together was suggested by Setzer (1971 : 262) : records (Fig. 3) of angolensis are thus restricted to ANGOLA (Monard, 1935 : 45 ; Hill & Carter, 1941 : 49, 176 ; Setzer, 1971 : 260, 263, fig. la (tragus)) and probably ZAIRE (Hayman, 1957 : 43 (in part) ; Hayman, Misonne & Verheyen, 1966 : 50 (in part) ; Peterson, 1971 : 885 (in part)). Laephotis botswanae Setzer, 1971 Laephotis botswanae Setzer, 1971 : 260, 263, fig. ib (tragus). 50 miles west, 12 miles south of Shakawe, Botswana. Size between angolensis and the large species wintoni and namibensis (length of forearm 34-38 mm, condylobasal length (13-5-14-3 mm) ; ears and tragus generally a little larger than in angolensis (length of ear from meatus more than 16 but less than 1 8 mm) but markedly smaller than in either of the large species. Tragus (Fig. ic) usually with a distinct angularity in its posterior margin at its widest point. Dorsal surface similar in colour to wintoni ; ventral surface paler than in that species, the hairs more liberally tipped with buffy white rather than pale brown. Rostrum relatively long, narrow ; maxillary toothrows slightly divergent posteriorly ; post- palatal region as in angolensis, the length of its bony part from a line across the rear faces of m 3 ~ 3 to the anterior edge of the mesopterygoid fossa less than the distance 7 8 J. E. HILL from the anterior edge of the mesopterygoid fossa to the tips of the pterygoid hamulars (Table 2). The specimen (Harrison Zoological Museum 1.2533) from Ndola, Zambia, which Peterson (1971 : 888) thought might represent the female of wintoni or possibly an 5 mm FIG. i. Tragus of (a) Laephotis wintoni ; (b) L. angolensis ; (c) L. botswanae. FIG. 2. Palate ridges of Laephotis wintoni. REVIEW OF LAEPHOTIS THOMAS, 1901 T 79 FIG. 3. Distribution of Laephotis wintoni ; A L. namibensis ; T L. angolensis L. botswanae. undescribed taxon and later (1973 : 602) closely related to botswanae proves in fact referable to this species. Although larger than specimens from Zaire or than those (B.M. 55.1134-1135) from Solwezi Boma, Zambia (i2io'S, 263o' E) its tragus (from the dry specimen) is angular posteriorly and the specimen agrees precisely in coloration with the other examples from Zambia, differing in this respect from the holotype of wintoni in its distinctly paler, more buffy white underparts, especially posteriorly. Specimens referable to this species have been recorded (Fig. 3) from BOTSWANA (Smithers, 1968 : 48, 49 (map), fig. a (head) (as angolensis) ; Setzer, 1971 : 260, 263, fig. ib (tragus) ; ZAMBIA (Ellerman, Morrison-Scott & Hayman, 1953 : 78 (as angolensis} ; Ansell, 1957 : 538 (as angolensis) ; Hayman, 1957 : 43 (as angolensis} ; Ansell, 1960 : 21 (as angolensis) ; Peterson, 1971 : 885, pi. i (fig. 2) (skulls, including specimen from Ndola) (as angolensis and Laephotis sp.) ; Setzer, 1971 : 260) ; ZAIRE (Hayman, 1957 : 43 (in part, as angolensis) ; Hayman, Misonne & Verheyen, 1966 : 50 (in part, as angolensis) ; Peterson, 1971 : 885 (in part, as angolensis) ; Setzer, 1971 : 260). 1 ! i rt rt .0 _U oj o3 ri rf PH &t OH ^ ^ Aweooq e c a e '3 13 13 t^ > rt C rf ^ .r-t .^H .^H nO 4)0) 5l)(U' 13 ' 13 ^- 1 *-HHHhHI-MWWhH C/) (/) - e iuo v"> 7*" T ?*" "9 !P Y"> ^< NN WNNNNNNNNNNN OOt^ MOONOiOO MONO* 10 o u"i *o 10 *o o ui oiQiduioo o o co i^ 1 jo q^Sugq 2 2 2 e ui-o 9 9 r 9 1 N N 9 9^9 ^M- inin^^inin 10^1-10 OOON ^-MION NON 0000 'IONONONON (ONON Cx ON -00 ON m ON NO roOt^^^^N .OOO rt- io!n inioio^w^inin in*n O O0t^ Tt-ONQOOOOO .Oro V roro T}-ro4j.Tt-rj-rj-ro 'fTj- ^ ^f ^ .(ONVOOOOO roiO 1 * ^ B JAL ob ob ob ob ob ob ob 9S'BDUl'Bjq 00 OO O 00 CO t^ 00 t^ ON ^^ tn ro c^> . | vo vo vo r^* in in vo 9S'BOurejq ^ ro n- 10 o o o vO 0000 OOO|Tf-rOMN OM^J- O' M OO O P q^PTAY ^ ^ ^ ^ ^ ^ ^ vo oo t^o r^t^c^c^ t>-t^t^. f^ w ro i CO oo o M - ^ " " VO .T M ro N t^ t^ in TAV Ol^moSXz 1 1 ON ob 00 ON ON ON ' ob 1 ob 11111 ''ob 4) ON M ^JOja^UI ^.S'69'T rororororororo roro iqOBI SSOJOB r>i -r 10 * ro ro roro roro^rorororo rororo O0t^ O N O>-IMON Oroin ,2 o> fe " ^ iqjo jo q^piAV 10 10 in in in ^'B^iqjoa^u'B in \o <* t^ vo ^M* *n ***> in if) *n ~*t~ ininin SSOJOB q^piA\ * * * -^- Tj- q^U9'T ro ro ro ro ro UIUBOOlApUOQ 2" IT 1? 1? I? qiU9I N N o 10 10 >n 10 10 10 roN I'rorororo rororj- G -a 5 jmilS JO OOOOMOrOrOM tOO t^ OOC^ in inrOOro rOO o 2 - g n 3 ^ 3u9j ^sa^Bajf) minovovoovo oo ro roro ^I'^t-Tf^J-Tf- Irt-io Jo fc i^ i^gui taoij _ * H U-N ^ ^_ OO> MiOinrocoOiMTt' ^ S , *8 S "i fe||6 ffi 1^9 jo q^Sugq ^ N N o3 N N 1 oiin f^vbooot^^-^ 111 OT fl 8 a U3 - O ^ _ jfr^-'ooinr^oonN MM [ UOI^BJ;ST9^I m^O-?O^ONON N N ,n2.ror^rororo ^-'t- "^vOvO^-Th-^- "ororo ONCOVO^J.J.^ i^ i_( i_, .J2 ^ s* s a si ^N in^i- f^r^c^t^io^ooo^-io^ ro roro rococororororocororororo *O *O O Of Ot- CH- CH- O O O Of Of *o Of CH- ia - * r-. o ^-- ^Q BM British Museum (Natur HZM Harrison Zoological Mu ROM Royal Ontario Museum USNM United States National AMNH American Museum of J 1UT? AP M.,c/So T?rolo /!Q 1* \trir REVIEW OF LAEPHOTIS THOMAS, 1901 81 TABLE 2 Palatal measurements (in mm) of Laephotis o fc _o I u 2 'be K wintoMZ BM 1.5.6.5* HZM 2.2030 BM 72.4397 BM 72.4398 BM 72.4399 (?) angolensis BM 57.435 BM 57.437 botswanae BM 57.436 BM 57.438 MRAC 26.402 MRAC 26.403 MRAC 26.404 MRAC 26.405 BM 55. 1134 BM 55. 1135 HZM 1.2533 $ ,-H , * .2 111 ^ O o3 > is en 1-1 fl o be tS (U *-i 43 "> S HH r fli be ^ ^ be ^ g S ft > S -^ J" ft 1 a 7-9 6-5 4-4 2-1 1-9 4-0 Kenya - - - - - - Kenya 8-1 6-8 4-4 2-4 1-8 4-2 Ethiopia 8-2 6-4 4'2 2-2 2-0 4-2 Ethiopia 8-5 6-8 4'3 2-5 1-9 4-4 Ethiopia 6-2 4-8 3-6 1-2 2-O 3-2 Zaire 6-1 4-8 3'5 i-3 1-9 3-2 Zaire _ 5'4 4-0 1*4 _ _ Zaire 6-7 5-2 3-8 1-4 2-0 3'4 Zaire 6-6 5'2 3'7 i '5 2-O 3'5 Zaire 6-6 5'2 3'7 I- 5 1-9 3'4 Zaire 6-4 5'2 3-8 1-4 1-9 3'3 Zaire 6-5 5-3 3-8 1-5 1-9 3'4 Zaire 6-5 5'2 3'7 I- 5 - - Zambia 6-4 5'2 3'7 I- 5 1-9 3'4 Zambia 7-1 5-6 4-1 I- 5 2-O 3'5 Zambia BM British Museum (Natural History), London. HZM Harrison Zoological Museum, Sevenoaks. MRAC Mus6e Royale de 1'Afrique Central, Tervuren. * Holotype. SUMMARY The majority of specimens of Laephotis hitherto reported in the literature are allocated to the appropriate one of the four species recognized in the most recent study of this genus, and these are briefly reviewed. Female specimens of L. wintoni are recorded from Ethiopia, whence until now the genus has been unreported. ACKNOWLEDGEMENTS My thanks are due to Dr M. J. Largen, of the Natural History Museum, Haile Selassie I University, Addis Ababa, who obtained the specimens from Ethiopia that 82 J. E. HILL led to this study and which have been donated to the British Museum (Natural History) : to Drs M. Poll and D. Thys van den Audenaerde of the Musee Royale de 1'Afrique Centrale, Tervuren, for the loan of the material of Laephotis in their care, and to Dr D. L. Harrison, who made available the specimens in his collections at Sevenoaks. REFERENCES ANSELL, W. F. H. 1957. Some mammals from Northern Rhodesia. Ann. Mag. nat. Hist. (12), 10 : 529-551. 3 tabs. 1960. Mammals of Northern Rhodesia. Lusaka: Government Printer. EIXERMAN, J. R., MORRISON-SCOTT, T. C. S. & HAYMAN, R. W. 1953- Southern African mammals 1758 to 1951 : a reclassification. London : British Museum (Natural History) . HARRISON, D. L. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. Jl E. Africa nat. Hist. Soc. 23 : 286-295. HAYMAN, R. W. 1957. Further notes on African bats. Rev. Zool. Bot. afr. 56 : 41-45. & HILL, J. EDWARDS. 1971. In: Meester, J. & Setzer, H. W. (Eds). The Mammals of Africa. An identification manual, i. Chiroptera. Washington: Smithsonian Institution. MISONNE, X. & VERHEYEN, W. 1966. The bats of the Congo and of Rwanda and Burundi. Annls Mus. r. Afr. cent. Ser. 8, Sci. Zool. No. 154, i-viii, 1-105, 2O P^ 8 - HILL, J. ERIC & CARTER, T. D. 1941. The mammals of Angola, Africa. Bull. Am. Mus. nat. Hist. 78 : 1-211, 36 figs., 17 pis. MONARD, A. 1935. Contribution a la mammalogie d' Angola et Prodrome d'une faune d' Angola. Archos Mus. Bocage, 6 : 1-314, 47 figs. PETERSON, R. L. 1971. Notes on the African long-eared bats of the genus Laephotis (family Vespertilionidae). Can. J. Zool. 49 : 885-888, i fig., i pi., i tab. 1973- The first known female of the African long-eared bat Laephotis wintoni (Vesperti- lionidae : Chiroptera). Can. J. Zool. 51 : 601-603, i fig. SETZER, H. W. 1971. New bats of the genus Laephotis from Africa (Mammalia : Chiroptera). Proc. biol. Soc. Wash. 84 : 259-264, i fig., i tab. SMITHERS, R. H. N. 1968. A checklist and atlas of the mammals of Botswana (Africa). ? Salisbury : Trustees of the National Museums of Rhodesia. THOMAS, O. On a new genus and species of Vespertilionine bat from East Africa. Ann. Mag. nat. Hist. (7), 7 : 460-462. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD THE GENUS FINMARCHINELLA SWAIN 1963 (CRUSTACEA : OSTRACOD A) AND ITS SPECIES By JOHN W. NEALE SYNOPSIS Six species of the genus Finmarchinella from the Northern Hemisphere are found to fall into two groups for one of which the new subgenus Barentsovia is instituted. The confusion surround- ing the four North Atlantic and Arctic species is analysed and the new species Finmarchinella (Barentsovia} curvicosta is described. INTRODUCTION IN 1963 Swain established the genus Finmarchinella with Cythereis finmarchica Sars 1866 as the type species. In 1957, however, Mandelstam had described the new species Nereina barentzovoensis from Russian Harbour, Novaya Zemlya, as the genotype of his new genus Nereina. In 1966 Ishizaki pointed out that Nereina was used for a gastropod described by Cristofori and Jan in 1832 and that Mandelstam's genus required a new name. The following year, Hazel (i967a, p. 18) examined the classification of Recent Hemicytheridae from off North-eastern North America and regarded Finmarchinella finmarchica and Nereina barentzovoensis as congeneric placing them both in Nereina and considering Finmarchinella to be a junior synonym. Subsequently Hazel (igGyb) noted that Nereina was preoccupied and since Fin- marchinella was available he validated Swain's genus which thus contained these two species and Cythereis angulata Sars 1866. In addition Nereina japonica Ishizaki 1966 from the Miocene and Pliocene of Japan and Finmarchinella uraniponnica Ishizaki 1969 from the Recent seas of western Honshu also belong here. These five species, together with a new Arctic species fall into two distinct groups. F. finmar- chica and F. uraniponnica are regarded as falling within the strict interpretation of Finmarchinella ; the remaining four species are separated and placed in the new subgenus Barentsovia based on F. (B.) barentzovoensis (Mandelstam). The principal aim of this paper, however, is to sort out the confusion that has arisen between the occidental species of the subgenus Barentsovia and place their taxonomy on a sound footing. This has been done on the basis of carapace characters since soft parts have not been available for study in most species. The uncontroversial Finmarchinella s.s. group is dealt with first. LOCATION OF MATERIAL Specimens preceded by 'BM' are in the collections of the British Museum (Natural History), those preceded by 'HU' are in the collections of Hull University. SYSTEMATIC DESCRIPTIONS Genus FINMARCHINELLA Swain 1963 TYPE SPECIES. Cythereis finmarchica Sars 1866. Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 3* 84 JOHN W. NEALE Subgenus FIN MARC BIN ELL A Swain 1963 Finmarchinella (F.) finmarchica (Sars 1866) (PI. i, figs. 6, 7 ; PL 2. figs, i, 5 & n) Cythereis finmarchica Sars, 1866 : 41. Hemicy there finmarchica, G. O. Sars, Sars, 1925 : 185-6, pi. LXXXV, fig. 3. DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Male left valve B.M. No. M aaSyA PI. i, fig. 6 0-688 0-331 Female left valve B.M. No. M 32876 PI. i, fig. 7 0-720 0-392 PI. 2, fig. ii Male carapace B.M. No. M 32876 PI. 2, fig. i 0-701 0-328 0-318 Female carapace B.M. No. M 32870 PI. 2, fig. 5 0-746 0-405 0-389 DISCUSSION. This species presents no difficulties in interpretation and only the original reference and the original author's later figure are given here in the synonymy. The characteristic features are the rather 'smoothed-out' appearance with well- developed reticulation and pitting but a lack of strong costation. In dorsal view the outline is smooth and more oval than in species of the subgenus Barentsovia (cf. PL 2, figs, i, 5 and PL 2, figs. 2-4). As in other species of the genus, sexual dimor- phism is very marked with the males much slimmer and lower in proportion to the length than the females (PL i, figs. 6, 7 ; PL 2, figs, i, 5). DISTRIBUTION. Since this species has not hitherto been confused with any other it is possible to use records in the literature to plot its distribution with some con- fidence (Fig. i). Sars originally described F. finmarchica from some empty shells from shell sand in Oxfjord, Finmark, and subsequently found it at Korshavn on the Norwegian coast further south. Besides Sars localities it has been found in Vadso and Lang Fjords (Brady & Norman, 1896, appendix). In the Norwegian and Barents Sea areas the author has found it in samples from Ernest Holt Station 2 (7029' N, I727' E) where it accounted for 2-18 per cent of the specimens, Station 6 (6954' N, I7oo' E, 1-62 per cent of the fauna), H.M.S. Vidal Station 29 (c. 6844' N, 4i23'5' E) and Station 46 on the Spitzbergen Shelf (75n-2' N, 22i4' E, 16-47 P er cent) . Widely distributed in high latitudes, it was recorded from three localities in the Davis Strait and has been found in material from the Hunde Islands (6852' N, 5307' W, 1-58 per cent), Holsteinsborg Harbour, Greenland (6655' N, 5325' W, 7-73 per cent), off Cape Flora, Franz Joseph Land (7957' N, 50oi' E, 1-25 per cent) and Russian Harbour, Novaya Zemlya (76i3' N, 624o' E, 0-97 per cent). Hazel (i967a) found it at Ungava Bay (6oo8' N, 6747' W) and gives a number of localities in the Gulf of Maine and on the Atlantic Shelf (Hazel, I967a, 1970). It was fairly widely recorded round Scotland and England by Brady (1868) and Brady and Nor- man (1889), many of whose records can be confirmed by reference to material in the Hancock Museum, Newcastle-upon-Tyne ; Ostenfeld and Wesenberg-Lund (1909) THE GENUS F IN M ARCH I N ELLA SWAIN Finmarchinella (F.) finmarchica (Sars 1866) f = fossil ZENITHAL EQUAL AREA FIG. i note it from the North Sea and the author has found it in the Celtic Sea in core sample 5652 (4938' N, 9i9' W) and from both top and bottom of the 88 cm core sample 5674 (495o-9' N, 9o8' W). Vos (1957) added the Chateau du Taureau (entrance to the River Morlaix Estuary). It has been widely encountered in the Bay of Biscay (Peypouquet, I97ib, gives 22 localities) but it is always rare and Peypouquet (i97ia) and Moyes & Peypouquet (1971) regard it as representing a palaeothanatocoenose of Holocene and Pleistocene age. This could explain the record from St Vincent, Cape Verd, by Brady & Norman (1889) but on balance one is inclined to agree with Hazel (i967a) who thought that it was most probably a mis-identification. It is well known as a fossil in the post-Tertiary deposits of Britain (Brady, Crosskey & Robert- son, 1874), Hazel (1968) has found it in the Pleistocene of a number of submarine canyons off the eastern seaboard of the United States and Lev (1969) has found it in his Normanicy there concinella community in the Kazantsevian deposits of the right bank of the River Yenesei and Cheshkaya Bay. Thus, F. finmarchica is essentially an Arctic and boreal-Arctic species whose southern limit appears to be about Ushant (Vos figures soft parts from the Chateau du Taureau locality). 86 JOHN W. NEALE Finmarchinella (Finmarchinella) uraniponnica Ishizaki 1969 Finmarchinella uraniponnica Ishizaki, 1969 : 217-218, pi. 26, figs. 12, 13 ; pi. 24, fig. 4 ; Ishizaki, 1971 : 83, pi. 3, fig. 3. This species is most easily differentiated from F. finmarchica by its more asym- metrical anterior and more sloping posterior margins in lateral view. The slight concavity in the anterior half of the ventral margin seems a little more accentuated also. Not so far recorded outside Honshu, it appears to be rather rare in Western Honshu where it was only found at one station in the Nakanoumi Estuary. Here it was accompanied by Cushmanidea subjaponica and Hermanites tosaensis which were also confined to this station, and seventeen other species which had a much wider distribution. In N.E. Honshu in Aomori Bay, however, Ishizaki (1971) has shown that it occurs much more commonly. Subgenus BARENTSOVIA subgen. nov. TYPE SPECIES. Finmarchinella barentzovoensis (Mandelstam, 1957). DIAGNOSIS. A subgenus of Finmarchinella which differs from Finmarchinella s.s. in the development of marked costation, particularly in the posterodorsal part of the valve and in the prominent development of the subcentral and eye tubercles. As shown below, the three occidental species of Barentsovia have been the subject of confusion amongst themselves and it is impossible to use the records in the litera- ture to establish their distribution unless there are accompanying illustrations which show the distinctive features. In consequence the distributions on the map (Fig. 2) use only the latter and material actually verified by the author. Finmarchinella (Barentsovia) barentzovoensis (Mandelstam, 1957) (PI. i, figs. 4, 5, 9 ; PI. 2, fig. 12) Cythere clathrata, Reuss var. nuda Brady, 1866 : 376-377, pi. 59, figs. 9, 10. Hemicythere angulata (Sars) Akatova, 1946 : 227, fig. 5. Nereina angulata (Sars) Hazel, 1967 (pars) : 19, pi. i, figs. 7, n. (non pi. i, figs. 8, 9, 10 = F. (B.) curvicosta.) Nereina barentzovoensis Mandelstam, 1957 : 180, pi. 3, figs. 7, 8. DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height (mm) (mm) Male left valve HU. 8o.R.23 PI. i, fig. 4 0-766 0-392 Female left valve HU. 8o.R.24 PI. i, fig. 5 0-831 0-457 Female left valve HU. 8o.R.25 PI. i, fig. 9 0-821 '447 PI. 2, fig. 12 DISCUSSION. This species has been confused with F. (B.) angulata (Sars). The earliest record of this species is that of Brady (1866) who figured it as a variety of THE GENUS FINMARCHINELLA SWAIN Finmarchinella (B.) angulata (Sars 1866) Finmarchinella (B.) barentzovoensis (Mande!stam1957) Finmarchinella (BJ curvicosta sp.nov. IHM. EQUAL AREA FIG. 2 Cythere clathrata Reuss which he called nuda, giving as localities the Hunde Islands and Norway. His figures show the characteristic ornamentation and this is con- firmed by a slide from the Hunde Islands in the Brady Collection in the Hancock Museum labelled 'Cythere angulata ? Sars = var : nuda Brady'. This contains five specimens, male and female right valves and two female left valves of F. (B.) barentzovoensis and a female right valve of F. (B.) angulata (Sars). Later Brady & Norman (1889) placed Cythere clathrata var. nuda in synonymy with Sars' species and the confusion has persisted. Akatova (1946) figured a specimen as Hemicythere angulata (G. O. Sars). More recently Hazel (i967a), commenting on the difficulty of interpreting Mandelstam's species from the published figures and lacking comparative material, figured two specimens of this species as Nereina angulata (Sars). The dif- ferences, however, are clear and well seen by comparing PI. i, figs. 4, 5 with PI. 2, fig. 8, and PI. 2, figs. 10, 12. The species is most easily recognized in lateral view by the strong rib which follows the anterior margin in its dorsal half and then curves in to the body of the valve at about mid-height. The posterodorsal quarter of the valve 88 JOHN W. NEALE surface is also particularly distinctive with two oblique ribs which run postero- dorsally from the subcentral tubercle and are linked by a very well-developed in- flation or tubercle at their posterodorsal termination. The lower of these two ribs is the stronger. DISTRIBUTION. Mandelstam's original material came from Russian Harbour, Novaya Zemlya, and material from this locality collected by Professor H. V. Howe in 1937 yielded 146 excellent specimens (46 per cent of them adults) of which three are figured here. Hazel's specimens came from Frobisher Bay (63io' N, 6745' W) and the Gulf of Maine (44o8' N, 68i3'W). Akatova figured a specimen from station 80 on the Novosiberian Shelf (j6$2' N, I3323' E). The present author has found it in British Museum (N.H.) samples collected by the 'Ernest Holt' at Stations i (7032' N, i823' E) and 6 (69^4' N, i7oo' E) and by H.M.S. Vidal at Station 46 (75n-2' N, 22i4' E) on the Spitzbergen Shelf, and in material from the Hunde Islands (6852' N, 53O7' W) in the Hancock Museum. The distribution of F. (B.) barentzovoensis is essentially Arctic (Fig. 2) , all the records (except for the somewhat anomalous specimen from the Gulf of Maine figured by Hazel) being from latitudes higher than 63 N and the maximum depth at which it has been recorded is 143 fathoms (Ernest Holt Station 6), the bulk of the occurrences being between 10-40 fathoms. It may thus be regarded as a relatively shallow-water Arctic form and it is possible that Hazel's Gulf of Maine right valve is a sub-Recent specimen. Finmarchinella (Barentsovia) angulata (Sars 1866) (PL i, fig. 8 ; PL 2, figs. 2,4, 6 & 10) Cythereis angulata G. O. Sars, 1866 : 46. Cythere angulata (Sars), Brady, 1868 : 409, pi. 26, figs. 39-42 ; Brady, Crosskey & Robertson, 1874 : 162-163, pi. 4, figs. 17-24, pi. 10, fig. 22 ; Brady & Norman, 1889 (pars) : 165 (non Cythere dathrata, var. nuda, Brady = F. (B.) barentzovoensis). Hemicythere angulata, G. O. Sars, Sars, 1925 : 187-188, pi. LXXXVI, fig. 2. 'Hemicy thermae' (genre ?) angulata (Sars), Wagner, 1957 : 61-62, pi. XXXVIII, figs, i, 2 ; Yassini, 1969 : 42-43, pi. XXXIX, fig. 17. Non Hemicythere angulata (G. O. Sars), Akatova, 1946 : 227, fig. 5 (= F. (B.) barentzovoensis). Non Finmarchinella angulata (Sars), Swain, 1963 : 813 (pi. 97, fig. 22 = juv. indet. ; pi. 99, fig. 9 & text-fig, nc. = F. (B.) curvicosta). Non Nereina angulata (Sars, 1865), Hazel, 1967 : 19 (pi. i, figs. 7, n = F. (B.) barentzovoensis ; pi. i, figs. 8, 9, 10 = F. (B.) curvicosta). DIMENSIONS OF FIGURED SPECIMENS. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Female left valve Oslo Museum F 1447.4 PL l > fig- 8 0-704 0-380 PI. 2, fig. 10 Female right valve Oslo Museum F 1447.6 PL 2, fig. 6 0-668 '357 Male carapace B.M. No. 1973.331 PL 2, fig. 2 0-701 0-367 o*3 2 5 Female carapace B.M. No. 1973.332 PI. 2, fig. 4 0-740 '4 I 5 '377 THE GENUS F IN M ARCHINELLA SWAIN 89 DISCUSSION. This species has been confused with both F. (B.) barentzovoensis and F. (B.) curvicosta in the past. Sars' illustration in the Crustacea of Norway (1925, pi. LXXXVI, fig. 2) shows a rib which slightly affects the lateral outline postero- dorsally. Sars (1925, p. 188) notes that he had taken this species 'in several places on the Norwegian coast from Christiana Fjord to Finmark, in the laminaria zone, though nowhere in any abundance'. The type material could not be traced but the Zoologisk Museum, Oslo, kindly searched its collections and provided a phial of spirit material labelled in Sars' own hand 'Hemicythere angulata G.O.S.' This yielded 13 specimens of F. (B.) angulata including four juveniles which showed that Sars' figure emphasized the essential features. Dorsally there is a rib which runs from approximately the mid-length of the valve towards the posterodorsal cardinal angle where it becomes accentuated, often affecting the outline posterodorsally in lateral view. The rib then turns downward at almost a right angle, or even more acutely, before disappearing at about mid-height (PI. i, fig. 8a ; PL 2, fig. 10). The posterior part of the dorsal margin is slightly concave in lateral view. F. (B.) angulata is most easily differentiated from F. (B.) barentzovoensis and F. (B.) curvicosta by the lack of any rib connecting the ventral end of this posterodorsal accentuation with the subcentral tubercle. Sars' material also shows the presence of a marginal rib anteriorly (PI. i, fig. 8a) which coincides with the outline at the anterior cardinal angle and ventrally, but lies some distance inside anteriorly where it follows the general curve of the outline but with the dorsal two-thirds forming a much flatter arc. This species also lacks the incurved part of the anterior rib at about mid-height which is characteristic of F. (B.) barentzovoensis. DISTRIBUTION. Sars found this species living in shallow water along the length of the Norwegian coast. It occurs at Russian Harbour, Novaya Zemlya, and Ernest Holt Station 6 in the Barents and Norwegian Sea areas and the author has also found juveniles in Cockburn Bank samples 5668 (495o' N, 9i8' W) and 5673 (495o' N, 909' W) although these Celtic Sea specimens are possibly not indigenous. Slides in the British Museum (N.H.) show that in the Greenland area it was found at Hol- steinsborg Harbour (where it occurred with almost equal numbers of F. (B.) curvicosta) and at Station 3 (693i' N, 56 01' W) of H.M.S. Valorous in 1875 ; in the Norwegian area at Hollingpollen, Drobak and Lungegaards-vandet, Bergen ; and round the Irish and Scottish coasts at Clew Bay, County Mayo, between the Cumbraes in the Firth of Clyde, off Tarbert, Loch Fyne and in the Minch. From the Brady Collection in the Hancock Museum, Newcastle-upon-Tyne, it has been possible to verify the pre- sence of this species in the Hunde Islands in the Greenland region ; in Stromness Bay, Scapa Flow, Rothesay Bay, Cumbrae, Rosneath, and Bo'ness in the Firth of Forth in the Scottish area and in Westport Bay in Ireland. From the same collection one can also confirm that it occurs fossil at Loch Gilp, Dalmuir, Colintraive, Cumbrae and in raised beaches at Oban and Tarbert. It also occurs fossil in the Quaternary of the Netherlands (Wagner, 1957) and Yassini (1969) found it at eight localities in the Bay of Biscay although he did not regard it as living in that area at the present day. Most records of this species are from less than 50 fathoms and only rarely has it been found at depths of more than 100 fathoms. Its known occurrence is shown in Fig. 2. go JOHN W. NEALE Finmarchinella (Barentsovia) curvicosta sp. nov. (PL i, figs. 1-3 ; PI. 2, figs. 3, 7-9, 13) Finmarchinella angulata (Sars) Swain, 1963 : 813-814, pi. 99, fig. 9 & text-fig, nc. non pi. 97, fig. 22 (= juv. indet.) Nereina angulata (Sars) Hazel, 1967 (pars) : 19, pi. i, figs. 8, 9, 10. non pi. i, figs. 7, n ( = F. (B.) barentzovoensis] . HOLOTYPE. A female carapace (mounted as separate valves) from Holsteinsborg Harbour, Greenland, B.M. No. 1973.310. PARATYPES. Nineteen specimens from Holsteinsborg Harbour, Greenland, including males, females and juveniles, B.M. Nos. 1973.311-329. OTHER MATERIAL. Fifty-two specimens in the collections of Hull University and Louisiana State University. DIMENSIONS OF FIGURED MATERIAL. Catalogue no. Figure Length Height Width (mm) (mm) (mm) Female left valve, holotype B.M. No. 1973.310 PI. i, fig. 3 0-808 0-451 PI. 2, fig. 13 Male left valve, paratype B.M. No. 1973.311 PI. i, fig. 2 0-743 0-383 Female right valve, paratype B.M. No. 1973.312 PI. i, fig. i 0-844 '45 PI. 2, fig. 8 Female left valve, paratype B.M. No. 1973.313 PI. 2, fig. 9 0-844 '454 Female carapace, paratype B.M. No. 1973.314 PI. 2, fig. 3 0-811 0-441 0-431 Female left valve HU. 8o.R.26 PI. 2, fig. 7 0-876 0-506 DIAGNOSIS. A species of Finmarchinella (Barentsovia} with gently convex postero- dorsal margin, well-developed costation and overall reticulation. An almost horizontal rib runs posteriorly from the subcentral tubercle to about five-sixths of the length where it joins a strong curved rib at an acute angle. The curved rib passes into a weaker oblique rib which runs back to the subcentral tubercle. The elongate loop formed by these costae usually contains two rows of polygonal fossae. Dorsal and ventral ribs complete the principal costation. DESCRIPTION. Valves in lateral view rounded-rectangular of typical Finmar- chinella shape. Posterodorsal margin gently convex, posterior margin concave. In dorsal view rounded subhexagonal. Eye tubercles and subcentral tubercle well- developed. Ornamentation of overall reticulation consisting of finely pitted poly- gonal fossae and costation. There are four principal costae, the most prominent of which is an almost horizontal rib which runs posteriorly from the subcentral tubercle. At about five-sixths the length this joins at an acute angle a short, curved, accen- tuated rib which runs towards the middle of the dorsal margin and generally forms the most prominent feature of the ornamentation. Dorsally, this strong, curved THE GENUS FIN M ARCHI N ELLA SWAIN 91 rib runs into a relatively weak oblique rib which runs anteroventrally back to the subcentral tubercle. This straight section of the curved rib is the most weakly developed of the principal costae. As a whole the costate pattern in this area forms an elongated loop with its apex at the subcentral tubercle. The loop normally contains two rows of polygonal fossae. A third principal rib starts just below the posterior part of the eye tubercle and runs obliquely in a posterodorsal direction to join the dorsal margin at about three-quarters of the length (PI. i, fig. 2a ; PI. 2, fig. 3). A fourth, slightly flexuous costa delimits the junction between the lateral and ventral surfaces and is very prominent at its posterior termination. Hinge antimerodont, anterior and posterior toothplates with 5-6 (usually 5) teeth and median locellate groove in the right valve. Left valve with complementary struc- tures. Normal pore canals large, simple and scattered. Marginal pore canals simple and straight ; approximately 36 anteriorly and 10 posteriorly. Anterior and posteroventral vestibules exceedingly narrow. Muscle scar pattern typical of the genus consisting of a vertical row of four adductor scars with three scars anteriorly. The long axes of the adductor scars lie obliquely to the length of the valve, declining anteriorly. The middle scars are very clearly divided. Sexual dimorphism marked, the males being lower in proportion to their length than the females. AFFINITIES AND DIFFERENCES. This species is most closely related to the two preceding species. It is most easily differentiated by the posterodorsal margin which is convex in lateral view compared with the concavity in this region seen in F. (B.) barentzovoensis (PI. i, figs. 4, 5) and F. (B.) angulata (PI. i, fig. 8a). Besides other differences in ornamentation F. (B.) curvicosta lacks the prominent incurving of the anterior rib of F. (B.) barentzovoensis and the curve of the posterodorsal rib is much flatter and lies well below the valve margin compared with F. (B.} angulata (cf . PL 2, figs. 13 and 10). DISTRIBUTION. F. (B.) curvicosta is essentially Arctic in distribution (Fig. 2). Slides in the British Museum (N.H.) show that it occurs in 10 fathoms at Holsteins- borg Harbour and 5-25 fathoms at Godhavn Harbour, Disco in the Greenland area and was dredged from 100 fathoms at Station 3 (693i' N, 56oi' W) by H.M.S. Valorous in 1875. The Brady Collection in the Hancock Museum, Newcastle-upon- Tyne, contains a slide of this species from the Hunde Islands (6852'N, 53O7' W), also in the Greenland area. It occurs at Kneeland Bay (625g' N, 6728' W) and Ungava Bay (6oo8' N, 6747' W) where it was recorded by Hazel (ig67a) as Nereina angulata (Sars) and Swain (1963) figured a female left valve from the Pleistocene Gubik Formation of Alaska, also as Finmarchinella angulata (Sars) . In the eastern hemisphere it was found quite commonly in 8 fathoms at Russian Harbour, Novaya Zemlya. Finmarchinella (Barentsovia) japonica (Ishizaki, 1966) Nereina japonica Ishizaki, 1966 : 143-144, pi. 19, figs. 1-4, text-fig, i, figs. 3, 4. DISCUSSION. This species shows the well-developed submedian and eye tubercles of Barentsovia. The costation is difficult to ascertain from the figure of the holotype 92 JOHN W. NEALE but in his description Ishizaki notes that the dorsal rib extends to the posterior end where it turns downward and disappears at mid-height. This, coupled with the general reticulation, is much more reminiscent of F. (B.) angulata (Sars) than of the other species of Barentsovia. It differs from F. angulata in the more prominent ventral rib. Ishizaki's species is exclusively fossil and has not so far been found outside the Miocene and Pliocene of Japan. CONCLUSIONS The genus Finmarchinella is essentially a cold water genus which includes non- costate (s.g. Finmarchinella} and costate (s.g. Barentsovia} groups. The confusion between species of the latter is easily resolved by reference to the shape and ornamen- tation of the valves. ACKNOWLEDGEMENTS It is a pleasure to thank the authorities of the British Museum (Natural History), London, the Hancock Museum, Newcastle-upon-Tyne, and the Zoologisk Museum, Oslo, who kindly arranged the loan of the material on which this work was based. REFERENCES AKATOVA, N. A. 1946. On the fauna of Ostracoda from the Novosiberian Shelf waters. In: Volume III. Results of the drifting expedition in the main northern seaways in the ice- breaker steamship 'Sedov' 1937-1940. Isdat. Glavsevmorputi, Moscow, Leningrad, 224-230, 8 figs. (In Russian.) BRADY, G. S. 1866. On New or imperfectly known Species of Marine Ostracoda. Trans. zool. Soc. London, 5 : 359-393, pis. 57-62. 1868. A Monograph of the Recent British Ostracoda. Trans. Linnean Soc. London, 26 : 353-495. pis. 23-41. CROSSKEY, H. W. & ROBERTSON, D. 1874. A Monograph of the post-Tertiary Entomo- straca of Scotland (including species from England and Ireland). Palaeont. Soc. Monogr. London, 1-232, pis. 1-16. & NORMAN, A. M. 1889. A Monograph of the Marine and Freshwater Ostracoda of the North Atlantic and of North-western Europe. Section I. Podocopa. Sci. Trans. Royal Dublin Soc. 4 (Series II) : 63-270, pis. 8-23. & NORMAN, A. M. 1896. A Monograph of the Marine and Freshwater Ostracoda of the North Atlantic and of North-western Europe. Part II. Sections II -IV. Myodocopa, Cladocopa, and Platycopa. Sci. Trans. Royal Dublin Soc. 5 (Series II) : 621-746, pis. 50-68. CRISTOFORI, J. DE & JAN, G. 1832. Conspectus methodicus testaceorum in collectione mea exstantium. Scientiae Naturalis Cultoribus : 8 pp. HAZEL, J. E. ig67a. Classification and Distribution of the Recent Hemicytheridae and Trachyleberididae (Ostracoda) off Northeastern North America. U.S. Geol. Surv. Prof. Paper 564 : 1-49, pis. i-n. ig67b. Corrections : Classification and Distribution of the Recent Hemicytheridae and Trachyleberididae (Ostracoda) off Northeastern North America. /. Paleont. 41 : 1284- 1285. THE GENUS FIN M ARCHIN ELLA SWAIN 93 HAZEL, J. E. 1968. Pleistocene ostracode zoogeography in Atlantic Coast submarine canyons. /. Paleont. 42 : 1264-1271, 3 text-figs. 1970. Atlantic Continental Shelf and Slope of the United States. Ostracode Zoogeography in the Southern Nova Scotian and Northern Virginian Faunal Provinces. U.S. Geol. Surv. Prof. Paper 529-E : i-v, Ei-E2i, 69 maps, n text-figs., 3 tables. ISHIZAKI, K. 1966. Miocene and Pliocene Ostracodes from the Sendai Area, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 37 : 131-163, pis. 16-19, I fig. 1969. Ostracodes from Shinjiko and Nakanoumi, Shimane Prefecture, Western Honshu, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 41 : 197-224, pis. 24-26, 16 figs., 2 tables. 1971. Ostracodes from Aomori Bay, Aomori Prefecture, Northeast Honshu, Japan. Sci. Rep. Tdhoku Univ., Ser. 2. Geology, 43 : 59-97, pis. 1-7, 8 figs., 4 tables. LEV, O. M. 1969. Analysis and Comparison of Ostracod Communities from Late Cenozoic Deposits in the Lower Reaches of the Yenesei, Ob', Pechora, Mezen and N. Dvina Rivers. In: Uchen. Zap. Paleont. Biostrat., nauch-issled. Inst. geol. Arktiki, 28 : 25-34, 5 P^ s - (I n Russian.) MANDELSTAM, M. I., SCHNEIDER, G. F., KUZNETSOVA, Z. V. & KATZ, F. I. 1957. New Genera in the Families Cypridae and Cytheridae. Ezheg. vses. paleont. obshch. 14 : 166-193, P^ 3 - I -IV. (In Russian.) MOVES, J. & PEYPOUQUET, J. P. 1971. Les Ostracodes indicateurs d'un paleorivage pleisto- cene en bordure du plateau continental du Golfe de Gascoyne. C. R. Somm. Stances Soc. Geol. France. Fasc. 44 : 219-220. OSTENFELD, C. H. & WESENBERG-LuND, C. 1909. Catalogue des especes de plantes et d'ani- maux observees dans le plankton recueilli pendant les expeditions periodiques depuis le mois d'Aout 1905 jusqu'au mois de Mai 1908. Publs Circonst. Cons. perm. int. Explor. Mer. No. 48. PEYPOUQUET, J. P. 1971 a. La distinction des biocenoses, thanatocenoses, paleothanato- cenoses : probleme fondamental sur une plateforme continentale. Bull. Inst. Geol. Aquitaine, 11 : 191-208, i map. i97ib. Inventaire de la microfaune d'Ostracodes de la region de Cap-Breton. Butt. Inst. Geol. Aquitaine, 11 : 209-217, i map, i table. SARS, G. O. 1866. Oversigt af Norges Marine Ostracoder, Norske Vidensk.-Akad. forhandlingar (1865), 1866, pp. 1-130. 1922-28. An Account of the Crustacea of Norway, 9 : 1-277, P^ 8 - I-H9- Bergen Museum. SWAIN, F. M. 1963. Pleistocene Ostracoda from the Gubik Formation, Arctic Coastal Plain, Alaska. /. Paleont. 37 : 798-834, pis. 95-99, 13 figs. Vos, A. P. C. DE. 1957. Liste annot^e des Ostracodes marins des environs de Roscoff. Archs Zool. exp. gen. 95 : 1-74, 29 pis., i table. WAGNER, C. W. 1957. ^ ur ^ es Ostracodes du Quaternaire recent des Pays-Bas et leur Utilisation dans V Etude geologique des Depots holocenes. Mouton & Co., The Hague, pp. 1-259, 50 pis., 21 figs. YASSINI, I. 1969. Ecologie des Associations d'Ostracodes du Bassin d'Arcachon et du Littoral Atlantique. Application a 1' Interpretation de quelques Populations du Tertiaire Aquitain. Bull. Inst Geol. Aquitaine, I-XXIV, 1-288, 39 pis. Dr JOHN W. NEALE Department of Geology THE UNIVERSITY HULL HU6 7RX 3t PLATE i Finmarchinella Magnification in all cases X 72 4 FIG. i. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female right valve, B.M. No 1973.312. a. Internal lateral view, b. Oblique dorsal internal view. Holsteinsborg Harbour, Greenland. FIG. 2. Finmarchinella (B.) curvicosta sp. nov. Paratype. Male left valve, B.M. No. 1973.311. a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 3. Finmarchinella (B.) curvicosta sp. nov. Holotype. Female left valve of carapace, B.M. No. 1973.310. a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 4. Finmarchinella (B.) barentzovoensis (Mandelstam) . Male left valve, HU. 8o.R.23. External lateral view. Russian Harbour, Novaya Zemlya. FIG. 5. Finmarchinella (B.) barentzovoensis (Mandelstam). Female left valve, HU. 8o.R.24. External lateral view. Russian Harbour, Novaya Zemlya. FIG. 6. Finmarchinella (F.) finmarchica (Sars). Male left valve, B.M. No. M. 3287 A. a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 7. Finmarchinella (F.) finmarchica (Sars). Female left valve, B.M. No. M.3287 B. a. External lateral view. b. Oblique anterior view. Holsteinsborg Harbour, Greenland. FIG. 8. Finmarchinella (B.) angulata (Sars). Female left valve, Oslo Museum F 1447.4. a. External lateral view. b. Oblique anterior view. Locality unknown. FIG. 9. Finmarchinella (B.) barentzovoensis (Mandelstam). Female left valve, HU. 8o.R.25. Oblique anterior view. Russian Harbour, Novaya Zemlya. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i PLATE 2 Finmarchinella FIG. i. Finmarchinella (F.) finmarchica (Sars). Male carapace, B.M. No. M. 3287 C. Dorsal view, x 74. Holsteinsborg Harbour, Greenland. FIG. 2. Finmarchinella (B.) angulata (Sars). Male carapace, B.M. No. 1973.331. Dorsal view x 74. Holsteinsborg Harbour, Greenland. FIG. 3. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female carapace, B.M. No. 1973.314. Dorsal view X73. Holsteinsborg Harbour, Greenland. FIG. 4. Finmarchinella (B.) angulata (Sars). Female carapace, B.M. No. 1973.332. Dorsal view x 72. Holsteinsborg Harbour, Greenland. FIG. 5. Finmarchinella (F.) finmarchica (Sars). Female carapace, B.M. No. M.3287 D. Dorsal view x 74. Holsteinsborg Harbour, Greenland. FIG. 6. Finmarchinella (B.) angulata (Sars). Female right valve, Oslo Museum No. F 1447.6. Internal view showing muscle scar pattern x 335. Locality unknown. FIG. 7. Finmarchinella (B.) curvicosta sp. nov. Female left valve, HU. 8o.R.26. External lateral view x 60. Russian Harbour, Novaya Zemlya. FIG. 8. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female right valve, B.M. No. 1973.312. a. Anterior part of hinge x 160. b. Posterior part of hinge x 160. Holsteinsborg Harbour, Greenland. FIG. 9. Finmarchinella (B.) curvicosta sp. nov. Paratype. Female left valve, B.M. No. 1973.313. Oblique ventral view x62. Holsteinsborg Harbour, Greenland. FIG. 10,. Finmarchinella (B.) angulata (Sars). Female left valve, Oslo Museum No. F 1447.4. Lateral view of posterodorsal part of valve x 160. Locality unknown. FIG. ii. Finmarchinella (F.) finmarchica (Sars). Female left valve, B.M. No. 3287 B. Lateral view of posterodorsal part of valve x 144. Holsteinsborg Harbour, Greenland. FIG. 12. Finmarchinella (B.) barentzovoensis (Mandelstam) . Female left valve, HU. 8o.R.25. Lateral view of posterodorsal part of valve x 144. Russian Harbour, Novaya Zemlya. FIG. 13. Finmarchinella (B.) curvicosta sp. nov. Holotype. Female left valve (of carapace), B.M. No. 1973.310. Lateral view of posterodorsal part of valve X 135. Holsteinsborg Harbour, Greenland. Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE 2 EUPLOTES RARISETA SP. N. (PROTOZOA: CILIATEA) A NEW SMALL MARINE HYPOTRICH By C. R. CURDS, B. JEAN WEST & JEAN E. DORAHY INTRODUCTION TUFFRAU (1960) listed 55 specimens of the genus Euplotes Ehrenberg, 1830 that had been described to that date, and in his extensive revision of the genus eliminated many of the species. In doing so, Tuffrau (1960) suggested that there were four characters of fundamental importance which should be used for taxonomic purposes at the species level. These characters were : the number of laterodorsal kinetics, the arrangement of the dorsal silver-line network, the number of frontoventral cirri and the shape of the non-dividing macronucleus. Borror (1968) added the appear- ance of the pellicular or cortical sculpturing that is often a feature of some Euplotes spp. as a further criterion. Using these characters, in addition to some of the more traditional features, Borror (1972) listed 43 species of Euplotes in his revision of the order Hypotrichida Stein, 1859. When both traditional and modern criteria are considered, the species of Euplotes described in the present paper does not conform to any of those listed by Borror (1972) nor Carter (1972). MATERIALS AND METHODS (a) Source and cultivation The hypotrich was originally contained in a sample of seawater collected from Aberystwyth, Wales. Some seawater was inoculated into a flask of Erdschreiber solution ('Medium i', Committee on Cultures, Society of Protozoologists, 1958) which was then incubated in the dark at room temperature (about 20 C) for several weeks. Under these conditions, Euplotes was the dominant ciliate which grew. Some of the cells so obtained were used by Miss Sheila Andrews to initiate a culture at the Cambridge Culture Centre of Algae and Protozoa (Euplotes sp. LB i624/2a). Later, a clonal culture was established (Euplotes sp. LB i624/2b) and most of the work described herein relates to that clone. The hypotrich was maintained in the dark at room temperature in conical flasks or test tubes containing Erdschreiber solution. Cultures were fed weekly with a few drops of a thick suspension of freeze-dried baker's yeast and subcultured at monthly intervals. The ciliate also grew well in 150 ml conical flasks containing 100 ml of seawater and 10-12 rice grains, but the maximum populations reached in these cultures were lower than those achieved by the former culture method. In order to establish the possible ecological range of this species, some experimental work was carried out with media consisting of various dilutions of Erdschreiber solution sup- plemented with baker's yeast and seawater supplemented with rice grains. In these Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 4 96 C. R. CURDS, B. J. WEST & J. E. DORAHY experiments, duplicate cultures in each medium were subcultured serially into flasks containing media of progressively lower salinity at 10 per cent (v/v) intervals. The inoculum size was always 4 ml per 100 ml of medium and a period of 1-5 weeks at room temperature was allowed between each transfer. In this way it was possible to establish the approximate lower salinity limit for this species. (b) Light microscopy Observations and measurements were made using both living and fixed material. Living organisms were slowed by immersion in methyl cellulose (5 per cent w/v in seawater), and osmium tetroxide vapour (2 per cent w/v) was found to be a suitable fixative for cells which were to be drawn or measured. Large numbers of Euplotes were fixed in bulk and their nuclei were stained using Dippell and Chao's modification of De Lamater's basic fuchsin stain (Sonneborn, 1950). The silver-line system was displayed using the 'wet' method modified by Chatton & Lwoff (1930) and Corliss (1953). (c) Scanning-electron microscopy The preparation of these marine organisms for the scanning-electron microscope proved to be rather more complex than the method described by Small and Marszalek (1969). It was found necessary to wash the cells 5 or 6 times prior to fixation in order to separate the cells from bacteria, yeast and debris. The first two washes were carried out by slow centrifugation (approx. 400 g) for a few seconds. In this case the majority of the Euplotes were retained in the supernatant. The cells were then repeatedly washed in membrane-filtered seawater and centrifuged at approxi- mately 1750 g. The cells in the final pellet were rapidly killed in osmium tetroxide vapour to prevent the cirri curling during fixation. Considerable effort was made to find a good method of fixation for these marine ciliates which was suitable for scan- ning-electron microscopy. The fixative suggested by Small & Marszalek (1969) was that used by Parducz (1967) and although this was acceptable in some respects, it had the disadvantage of disrupting the structure of the cirri and the adoral zone of membranelles (AZM). The fixative finally chosen, although it is still under review, consisted of equal parts of 2 per cent (w/v in distilled water) osmium tetroxide and a saturated aqueous solution of mercuric chloride. Sufficient sodium chloride was then added to give a final concentration of 3-3 per cent (w/v) ; this prevented the cells swelling during fixation. The three component parts were mixed immediately before use and then membrane filtered ; cells were fixed in the filtered solution over- night. Following fixation, the cells were washed repeatedly (at least 8 times) in membrane-filtered triple glass-distilled water and centrifuged at approximately 400 g between each wash. The washed cells were then frozen by being dropped onto pieces of aluminium floating on liquid nitrogen and were dried in a tissue dryer essentially as described by Small & Marszalek (1969). The pieces of aluminium covered with organisms were glued to stubs and coated with approximately 100 A of gold as described by Harris, Martin & Ogden (1972), specimens were examined with a Cambridge 'Stereoscan' Mk II scanning-electron microscope (Cambridge Instru- ments Ltd, Cambridge, England). EUPLOTES RARISETA SP. N. 97 Euplotes rariseta sp. n. Diagnosis Small (30-45 /xm long, 20-31 /mi wide), ovoid, marine hypotrich with 10 fronto- ventral, 5 transversal and 3 caudal cirri ; caudal cirrus below AZM stout. Ventral surface heavily sculptured with 6 posteriorly projecting ridges. Dorsal surface with 6 double-edged longitudinal ridges. Dorsal bristles sparse ; 6 kinetics with a maxi- mum of 6 bristles in the central kinetics. Dorsal silver-line system double of the patella type. Macronucleus S-shaped. Micronucleus small, situated anteriorly. Type slides showing silver-line systems, nuclear apparatus and whole mounts have been deposited in the slide collection of the Protozoa Section, BM (NH) : holotype Reg. No. 1972:11:1:11 ; paratype Reg. Nos. 1972:11:1:12-18. Body size and shape The size distribution data of Euplotes cells at the stationary and logarithmic phases of growth are given in Fig. i. It should be noted that Euplotes cells from the logarithmic phase are considerably larger (42 3 /mi long by 28 3 /mi wide) than are cells from the stationary phase (34-6 3 /mi long by 24-2 2-3 /mi wide). This is in agreement with the observations of Curds & Cockburn (1971) who noted that the size of the ciliate Tetrahymena pyriformis was related to its rate of growth. The data on size show that this species of Euplotes is amongst the smallest recorded for the genus. The dorsal surface is conspicuously sculptured with 6 double-edged longitudinal ridges (PL i, figs, a and c). To the left of each ridge is a parallel row of pits from which the short (2 /u.m long) dorsal cilia or bristles emerge. The ventral surface is also strongly sculptured (Fig. 2 ; PI. i, figs, a and c), particularly in the posterior region between the transverse cirri. There are 6 ventral ridges, 3 of which are prominent and consist of an oblique midventral ridge, and 2 ridges which run longitudinally flanking the 2 outermost transverse cirri. The other 3 ridges are less prominent and are more or less restricted to the area immediately between the transverse cirri. All 6 ventral ridges project slightly posteriorly between the trans- verse cirri and form fin-like structures (Fig. 2 ; Plate i, figs, b and c). The right buccal overture has a slight anterior evagination as in Euplotes alatus Kahl, 1932 (see Borror, 1968) but in the species under description it extends only just past the AZM. Ciliary organdies This Euplotes has 10 frontoventral cirri which are arranged as shown in PI. i, fig. b and in Figs. 2 and 3b. The arrangement closely resembles that of both Euplotes charon Ehrenberg, 1830 and Euplotes quinquecarinatus Gelei, 1950 (see Borror, 1968). There is the normal complement of 5 transverse cirri which arise between the ventral ridges and these, particularly cirri III i and IV i (following the numeration of Wallengren, 1900), are frequently seen to be frayed at the distal end (Fig. 2). The tendency for the transverse cirri to disrupt in this way is greatly increased during fixation. There are only 3 caudal cirri, and in this species the one 9 8 C. R. CURDS, B. J. WEST & J. E. DORAHY 99- 95- >80- E 5- 2'0 2'4 28 32 36 SIZE (microns) 40 44 48 FIG. i. Size distribution data of Euplotes rariseta. Curve A. Breadth of cells at stationary phase. Curve B. Breadth of cells during logarithmic growth. Curve C. Length of cells at stationary phase. Curve D. Length of cells during logarithmic growth. Stationary- phase cells were fixed in osmic acid vapour. Logarithmic-phase cells were measured alive immersed in methyl cellulose solution. 10/jm FIG. 2. Ventral aspect of Euplotes rariseta showing cirri and ventral ridges (camera lucida drawing). EUPLOTES RARISETA SP. N. 99 a b FIG. 3. Silver-line system of Euplotes rariseta, (a) dorsal surface, (b) ventral surface. below and nearest to the AZM is characteristically held stiffly out to the left side and is much stouter than the other two caudal cirri which arise beneath the central transverse cirri (Fig. 2). The presence of this large stiff caudal cirrus appears to be a feature unique to this species ; in the living cell, the cirrus seems to act as a rudder- like structure. The dorsal bristles are sparse in this species ; there are 6 rows and the mid-dorsal rows have only 6 bristles, in which respect this species resembles Euplotes bisulcatus Kahl, 1932 (see Borror, 1968). Nuclei The macronucleus of this species is an irregular S-shape (Fig. 4). In this respect it appears to be unique in the genus Euplotes. The posterior part of the macro- nucleus points anteriorly and ventrally towards the position of the cytostome. The micronucleus is situated close to the macronucleus on the left anterior edge of the body (Fig. 4). Silver-line system The geometry of the dorsal silver-line system is of the double or patella-type and closely resembles that of Euplotes raikovi Agamaliev, 1966 with a series of alternate longitudinal rows of narrow long polygons and wide short polygons (Fig. 3a). The ventral argyrome (Fig. 3b) consists of an irregular network of polygons in similar numbers to that of E. cristatus Kahl, 1932 (see Tuffrau, 1960). The dorsal silver-line network has been seen on some scanning-electron micrographs (PL i, fig. d). C. R. CURDS, B. J. WEST & J. E. DORAHY FIG. 4. Ventral view of the nuclei of Euplotes rariseta from a stained preparation. (M = macronucleus, m = micronucleus.) Ecological data Cells inoculated into Erdschreiber solution supplemented with yeast grew well in nine different dilutions of this medium from 100 down to 20 per cent (v/v). Cells transferred from 20 to 10 per cent Erdschreiber solution showed no growth during the 6 weeks following inoculation. Growth in seawater supplemented with rice grains was good in seven dilutions from 100 down to 40 per cent (v/v), but cells transferred from 40 to 30 per cent seawater showed good growth only after 3 weeks in culture. Subsequent transfer of cells from 30 to 20 per cent seawater led to poor growth after 5 weeks, and no growth was observed in 10 per cent seawater. Contractile vacuole activity was not observed in organisms from any of the tested dilutions of seawater or Erdschreiber solution. This is in agreement with the findings of Yocum (1934), who found that the activity of the contractile vacuole in Euplotes patella Ehrenberg, 1831 diminished and became imperceptible at concentrations of seawater above 10 per cent. Yocum (1934) also found that E. patella would survive well and divide in 66 per cent seawater. In view of the finding that the species of Eiiplotes described in the present paper will survive and divide in dilutions of seawater or Erdschreiber solution between 20 and 100 per cent, it seems likely that, in nature, this organism could be found in both marine and brackish environments. However, for the lower dilutions tested, organisms were not as abundant in seawater medium as they were in Erdschreiber medium, and it is possible that in these experiments the amounts of substances other than sodium chloride were limiting growth. DISCUSSION AND CONCLUSIONS In the past, some authors have noted that the number of caudal cirri is not as constant as the others, for example, Euplotes mutabilis Tuffrau, 1960 has 4-5 caudals, E. raikovi has 2-3 caudals and there are other similar examples. In view of these observations perhaps the use of the number of caudal cirri as a character should be regarded with caution. Although E. octocirratus Agamaliev, 1967 is the only other species of Euplotes which has precisely the same numbers of cirri on the ventral surface, that is 10 frontoventrals, 5 transversals and 3 caudals, one should EUPLOTES RARISETA SP. N. 101 consider those species with a 10 : 5 : 4 cirri complement. These include the follow- ing : E. alatus, E. balteatus Kahl, 1932, E. crenosus Tuffrau, 1960, E. cristatus, E. harpa Stein, 1859, E. indentatus Carter, 1972, E. inkystans Chatton in Tuffrau, 1960, E. minuta Yocum, 1930, E. magnicirratus Carter, 1972, E. quinquecarinatus , E. roscoffensis Dragesco, 1966, E. trisulcatus Kahl, 1932, E. tuffraui Berger, 1965 and E. vannus Minkjewicz, 1901. In all these species the shapes of the macronuclei are quite unlike that of the species described in this paper and furthermore the silver- line systems of these species are different. It is unfortunate that the dorsal and ventral silver-line systems have not been described for all the known species of Euplotes ; however, of those listed above none have 6 dorsal kinetics as found in E. rariseta ; only E. octocirratus and E. trisulcatus have 7 kinetics, the remainder have 8 or more. There are four other species with 6 dorsal kinetics and these are E. raikovi, E. strelkovi Agamaliev, 1967, E. tegulatus Tuffrau, 1960 and E. balticus Kahl, 1932. None of these four species have the cor- rect numbers of cirri and furthermore all possess far too many dorsal kinetosomes. E. bisulcatus is the only other species with a maximum of 6 dorsal kinetosomes per kinety but there are 8 dorsal kinetics in that species. Euplotes rariseta differs from all previously described species of Euplotes in pos- sessing an S-shaped macronucleus and a very stout caudal cirrus. In addition, although the remaining characters have been observed amongst other species of Euplotes, it is evident from the information given above that none of these species has the same combination of characters. For these reasons, it is considered that the species of Euplotes described in this paper is sufficiently distinct from all others to be designated as a separate species, and because of the paucity of dorsal bristles has been named Euplotes rariseta. Following the revised classification of the Com- mittee on Taxonomic Problems of the Society of Protozoology (Honigberg et al. 1964) Euplotes rariseta is placed into class Ciliatea Perty, 1852, order Hypotrichida Stein, 1859, family Euplotidae Ehrenberg, 1838. ACKNOWLEDGEMENTS The authors wish to thank Mr R. Harris, Department of Zoology, British Museum (Natural History) , for his considerable aid and advice on the treatment of specimens for scanning-electron microscopy. We are indebted to Dr R. H. Hedley and Mr C. G. Ogden, British Museum (Natural History), and to Mr E. A. George and Dr F. C. Page, Culture Centre for Algae and Protozoa, who read the original manu- script and made several constructive suggestions. Part of this work was carried out while B. J. W. was employed by the Natural Environment Research Council, Culture Centre for Algae and Protozoa, Cambridge. REFERENCES BORROR, A. C. 1968. Systematics of Euplotes (Ciliophora, Hypotrichida) ; towards union of the old and the new. /. Protozool. 15 (4) : 802-808. 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Protozool. 19 (i) : 1-23. 102 C. R. CURDS, B. J. WEST & J. E. DORAHY CARTER, H. P. 1972. Infraciliature of eleven species of the genus Euplotes. Trans. Amer. Microsc. Soc. 91 (4) : 466-492. CHATTON, E. & LWOFF, A. 1930. Impregnation, par diffusion argentique, de 1'infraciliature des cilies marins et d'eau douce, apres fixation cytologique et sans dessication. C. r. Seanc. Soc. Biol. 104 : 834-836. COMMITTEE ON CULTURES, SOCIETY OF PROTOZOOLOGISTS. 1958. A Catalogue of Laboratory Strains of Free-living and Parasitic Protozoa. /. Protozool. 5 (i) : 1-38. CORLISS, J. O. 1953. Silver impregnation of ciliated protozoa by the Chatton-Lwoff technic. Stain Tech. 28 : 97-100. CURDS, C. R. & COCKBURN, A. 1971 . Continuous monoxenic culture of Tetrahymena pyriformis. J. gen. Microbiol. 66 : 95-108. HARRIS, R. H., MARTIN, B. S. & OGDEN, C. G. 1972. Notes on the preparation of natural history specimens for scanning electron microscopy. Bull. Br. Mus. nat. Hist. (Zool). 24 (3) : 223-228. HONIGBERG, B. M., BALAMUTH, W., BOVEE, E. C., CORLISS, J. O., GOJDICS, M., HALL, R. P., KUDO, R. R., LEVINE, N. D., LOEBLICH, A. R., WEISER, J. & WENRICH, D. H. 1964. A revised classification of the Phylum Protozoa. /. Protozool. 11 pp. 7-20. PARDUCZ, B. 1967. Ciliary movement and co-ordination in ciliates. Int. Rev. Cytol. 21 : 91- 128. SMALL, E. B. & MARSZALEK, D. S. 1969. Scanning electron microscopy of fixed, frozen and dried protozoa. Science, N.Y. 163 : 1064-1065. SONNEBORN, T. M. 1950. Methods in general biology and genetics of Paramecium aurelia. J. exp. Zool. 113 : 84-147. TUFFRAU, M. 1960. Revision du genre Euplotes, fondle sur la comparaison des structures superficielles. Hydrobiologia, 15 : 1-77. WALLENGREN, H. 1900. Zur Kenntnis der vergleichenden Morphologic der Hypotrichen. Bih. K. svenska Vetensk A had. Handl. 26 : 1-31. YOCUM, H. B. 1934. Observations on the experimental adaptation of certain fresh-water ciliates to seawater. Biol. Bull. mar. biol. Lab. Woods Hole, 67 : 273-276. Dr COLIN R. CURDS Miss B. JEAN WEST, B.Sc.* Mrs JEAN E. DORAHY, M.Sc. CULTURE CENTRE OF ALGAE AND PROTOZOA Department of Zoology STOREY'S WAY BRITISH MUSEUM (NATURAL HISTORY) CAMBRIDGE CROMWELL ROAD LONDON SW7 5BD * Present address. Microbiology Department, School of Biological Sciences, University of Bath, England. 4t PLATE i Scanning micrographs of Euplotes rariseta a. Dorsal view showing ridges and rows of dorsal bristles ( x 2-ik) b. Ventral view showing the arrangement of cirri and ventral ridges ( x i-8k) c. Posterolateral aspect showing double-edged dorsal ridges ( x 2-yk) d. Dorsal surface showing argyrome ( x 6-ik) Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i OBSERVATIONS ON CLONAL CULTURES OF EUGLYPHA ACANTHOPHORA AND EUGLYPHA STRIGOSA (TESTACEA : PROTOZOA) By R. H. HEDLEY, C. G. OGDEN & JUNE I. KRAFFT INTRODUCTION MANY testate amoebae are known only from their original descriptions, based on observations made at the magnification limits of the optical microscope. The introduction of scanning-electron microscopy for surface ultrastructure examination now allows more accurate descriptions to be made of these animals. The present account describes the shell morphology and biology of two species of Euglypha Dujardin, 1841, namely Euglypha acanthophora (Ehrenberg, 1841) and Euglypha strigosa (Ehrenberg, 1871), both of which have been recently isolated from samples of sewage sludge and moss respectively, collected in southern England. This is the third paper in a series dealing with testate amoebae. The previous papers (Hedley and Ogden, 1973, 1974) are devoted to the biology and fine structure of Euglypha rotunda and Trinema lineare respectively. Fine structure observations on the cytoplasm of E. acanthophora and E. strigosa will be the subject of a separate publication. The genus Euglypha belongs to the family Euglyphidae - the classification adopted here is that proposed by Loeblich and Tappan (1961) : Class RHIZOPODEA Von Siebold, 1845 Subclass FILOSIA Leidy, 1879 Order GROMIDA Claparede and Lachmann, 1859 Superfamily EUGLYPHACEA Loeblich and Tappan, 1961 Family EUGLYPHIDAE Wallich, 1864 test hyaline, symmetrical, elongate, composed of rounded siliceous scales, aperture rounded or elongate ; one nucleus. Previous work - biology Euglypha acanthophora. Most of the early biological observations of Euglypha were made on this species. These reports (Leidy, 1877 ; Gruber, 1881 ; Blochmann, 1887 ; Schewiakoff, 1888 ; Carter, 1889 ; Penard, 1890 ; Reukauf, 1912 ; Popoff, 1912) dealt mainly with nuclear division and reproduction by binary fission. Carter (1865, 1889), Reukauf (1912) and Popoff (1912) also described another process of division in which many individuals were produced at one time, but all the stages of this division were not seen. A report by Jones (1929), made from observations on an Bull. BY. Mus. nat. Hist. (Zool.) 27, 2 5* 104 R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT old protozoan culture, described a flagellate stage for E. acanthophora. This has not been confirmed and it is possible that the cultures were contaminated. Encystation was observed by Penard (1890, 1902), Leidy (1879), Popoff (1912) and Reukauf (1912). Leidy (1879) described the cyst as being enclosed in two distinct envelopes, and Popoff (1912) illustrated the various stages in the process of encystation. When two individuals were observed joining to form a single large specimen and immediately formed a cyst, Reukauf (1912) suggested that this was a copulation cyst which at excystation produced many new individuals. In studies on the distribution of testaceans in relation to their habitats, Heal (1961) found E. acanthophora in small numbers associated with fen areas, whereas de Graaf (1956) considered this species to have no particular associations. Euglypha strigosa. This species has been recorded from both wet and dry habitats, de Graaf (1956) noted that in a Netherland fen it preferred drier Sphagnum, whilst Heal (1961) described it as a eury topic species occurring in large numbers in bog hummocks in Northern England and also noted (Heal, 1962) that it appeared to be restricted to the lower levels of the Sphagnum. Differences in shell size and shape of specimens of E. strigosa were described by Chardez and Leclercq (1963), who suggested that these differences could be related to the type of habitats in which they occur. Previous work - taxonomy In a review of the British Freshwater Rhizopoda fauna Cash et al. (1915) summarize previous taxonomic reports of both Euglypha acanthophora and E. strigosa. The present account excludes these references and is restricted to subsequent descrip- tions. Euglypha acanthophora (Ehrenberg, 1841). This species was originally described as E. alveolata but was made a synonym of E. acanthophora by Cash et al. (1915) who gave a detailed account of their reasons, and this has since been accepted by most authors. They (Cash et al. 1915) redescribed E. acanthophora and also two varieties which differed mainly in size, and the shape and number of the elongated shell- plates. Further redescriptions and sizes were given by Hoogenraad and de Groot (1940) and Decloitre (1962, 1965), the latter author also described (Decloitre, 1962) seven varieties of E. acanthophora. Euglypha strigosa (Ehrenberg, 1872). Cash et al. (1915) described this species as 'the most generally distributed and numerous of the Euglyphae ', easily distinguished from E. ciliata and E. compressa by its thickened apertural-plates and circular aper- ture. In a review of the genus Euglypha, Decloitre (1962) redescribed E. strigosa and three varieties. Additional sizes and descriptions were given by Bartos (1963) and Decloitre (1964). The surface ultrastructure of the shell of E. strigosa was described by Cambar et al. (1964), from metal and carbon-shadowed preparations. MATERIALS AND METHODS Euglypha acanthophora was isolated from a sample of sewage sludge from Maple Lodge Works of the Colne Valley Sewage Board, Hertfordshire, in December, 1972. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 105 Euglypha strigosa was isolated from a sample of sphagnum moss collected in the New Forest, near Christchurch, Hampshire, in April, 1970. Both species were obtained from crude cultures made from small portions of each type of material placed in a shallow layer of culture liquid and kept in the laboratory at room temperature, 18-20 C. Agnotobiotic cultures were kept in small plastic containers, on a thin layer of agar agar (i per cent in distilled water) with a sterilized wheat grain added prior to setting and covered with a shallow layer of culture liquid. The culture liquid was a 5 per cent (w/v) solution of soil extract, plus 100 mg/1" 1 sodium nitrate and 15 mg/l" 1 sodium dihydrogen orthophosphate, in distilled water. E. strigosa has more recently been kept on 1-5 per cent agar in amoeba saline, with a sterilized rice grain added and covered with a shallow layer of Prescott and James solution (Prescott & James, 1955). Clonal cultures were established by isolating single active animals. One clone has been used subsequently to produce the working cultures of each species. These clones have been deposited, E. strigosa (No. 1520/2), E. acanthophora (No. 1520/3), and maintained at the Culture Centre of Algae and Protozoa, The Natural Environ- ment Research Council, Cambridge, England. Subcultures made at 4-5 week intervals are adequate to maintain active animals that readily feed and reproduce. Optical microscopy. Specimens were examined by both phase-contrast and bright- field illumination, either live or after fixation with Schaudinn's fluid. Smears of fixed animals were stained with either borax carmine or iron haematoxylin. Scanning-electron microscopy. Live specimens were either cleaned by transferring them from the culture vessels through several changes of triple-distilled water, or fixed in glutaraldehyde and then thoroughly washed. They were then manipulated using a single-hair brush onto a small cover-slip, previously cleaned with acetone and lint-free tissue, to which they adhere when dry. To obtain single siliceous shell- plates, an individual clean specimen was placed on a small segment of cover-slip and covered with a small drop of concentrated sulphuric acid. This was heated gently to evaporate the acid and liberate the plates from the organic cement. These prepared cover-slips were secured to 'Stereoscan' stubs with an electrically conductive paint, 'Silver Dag', prior to being coated evenly with 10-15 nm f gld using the method recommended by Harris et al. (1972). The stubs were examined on either the Cam- bridge Stereoscan Mk II or Stereoscan 600, operating at 15 or 20 kV and the results were recorded on Ilford HP4 film. DISTRIBUTION E. acanthophora is usually found in damp mosses, submerged vegetation and standing water, whilst E. strigosa is found in similar habitats and additionally in drier mosses and soil. A full list of locality records and references was given by Decloitre (1962). Since this review several authors, namely, Bartos (1963), Bonnet (1966), Decloitre (1964, 1965) and Stepanek (1963), have recorded these species from other localities and these are included here. io6 R. H. MEDLEY, C. G. OGDEN & J. I. KRAFFT The following lists illustrate the geographically widespread distribution of both species : Euglypha acanthophora EUROPE NORTH AMERICA SOUTH AMERICA : AFRICA : ASIA : AUSTRALASIA : England, Wales, Scotland, Ireland, France, Holland, Belgium, Germany, Spain, Switzerland, Hungary, Czechoslovakia, Luxembourg, Iceland and Lapland. Greenland, Canada and the United States of America. Venezuela and Colombia. Congo, Cameroons, French West Africa, South Africa and Madagascar. China, Japan, Java and Sumatra. Australia and New Zealand. Euglypha strigosa EUROPE : NORTH AMERICA SOUTH AMERICA ; AFRICA : ASIA : AUSTRALASIA : ANTARCTICA : England, Wales, Scotland, Ireland, France, Belgium, Holland, Spain, Germany, Switzerland, Italy, Hungary, Czechoslovakia, Bulgaria, Rumania, Spitzbergen, Finland and Iceland. Greenland, Canada and the United States of America. Central America, Colombia, Venezuela and Chile. South Africa, French West Africa, Madagascar and the Canary Islands. Krakatoa Island, Borneo and Java. Australia, Tasmania and New Zealand. Adelaide Island. MORPHOLOGY AND VARIATION Euglypha acanthophora. The shell has a mean length of 70-8 /u,m 9-6 /mi and a mean breadth of 39-0 /zm 4-4 ju,m ; it is elliptical in shape through the major axis and circular through the minor axis (PL i, fig. A), and is made up of three different types of siliceous plates- apertural-plates (PL i, fig. E), shell-plates (PL i, fig. B) and elongated shell-plates (PL i, fig. B). The aperture is terminal, circular and has a mean diameter of 18-4 /u,m 4-9 /u,m, it is surrounded by between 10 and 13 evenly spaced apertural-plates (PL i, fig. D). Each apertural-plate is roughly circular in shape and varies in size from 8-1 /u,m to 12-5 /u,m in length and 7-2 /u,m to 11-4 ju,m in width. The dentate edge of the apertural-plate is slightly thickened and carries a median tooth with either four or five smaller lateral teeth on each side (PL i, figs. C and E). The apertural-plates are found not only bordering the aper- ture, but are also seen in the second and third row inside the aperture (PL i, fig. E). There are approximately 200 shell-plates which range in size from 10-9 /urn to 12 ju.ni in length and 7-9 /u.m to 9-1 /mi in width. They are roughly circular in shape with the anterior edge usually having three small pointed projections (Fig. la), and are arranged regularly in alternate longitudinal rows (PL i. fig. A). In the posterior CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 107 a b c FIG. i. Diagram to show the comparative shape of shell-plates (a) E. acanthophora from anterior region, (b) E. acanthophora from posterior region, (c) E. strigosa. region of the shell there are normally between one to six elongated shell-plates which have one edge tapering to a fine point (PL i, fig. B). These plates are approximately twice as long as a normal shell-plate, nevertheless, the projections break easily so that they are often seen to be short and end bluntly. The pointed edge of these elongated shell-plates is usually directed posteriorly so that they are seen to project from the outline of the shell, but on occasions they follow the curvature of the shell and are difficult to see by light microscopy. Abnormally shaped shells occur occasionally, although the incidence of such forms appears to be less than three per cent. They are usually specimens with either deformed anterior regions or small specimens, about half or two-thirds normal length, with wide apertures. The first type is the most frequent abnormal shell form, having a normal arrangement of shell-plates in the posterior region but anterior to the middle region the shell-plates are compacted together haphazardly, so that the shell only attains half the normal length. These deformed shells usually collapse when air- dried whilst normal shells retain their shape. The animals inside these deformed shells appear to possess the normal cytoplasmic organelles. Euglypha strigosa. The shell has a mean length of 81-3 /mi 3-8 /mi and a mean breadth of 42-2 /mi 6 /mi ; it is elliptical in shape through both the major and minor axes (PI. 2, fig. A). Two distinct types of siliceous plates make up the shell, apertural-plates and shell-plates, and in addition there are numerous elongated siliceous spines (PI. 2, fig. A). The aperture is terminal and circular, with a mean diameter of 14-9 /mi + 0-8 /mi ; it is surrounded by between 10 and 13 evenly spaced, apertural-plates - the usual number of apertural-plates is n (PI. 2, fig. E). Each apertural-plate is roughly circular in shape and varies in size from 9-2 /mi to 10-8 /mi in length and 7-8 /mi to 8-5 /mi in width. One margin is distinctly thicked and carries a large median tooth with either three or four smaller teeth on each side (PI. 2, fig. C). The shell-plates are roughly oval in shape (Fig. ic), with a size range of from 8-0 /mi to 11-3 /mi in length and 4-1 /mi to 6-4 /mi in width, and they are arranged in regularly spaced, alternate longitudinal rows. The thin, elongated spines vary in shape and size from small pear-shaped spines approximately 2 /mi long to thin io8 R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT spines 23 p,m long, and they project from any part of the shell surface (PL I, fig. A). They are found at the junction of shell-plates, where they are held in position by a similar organic cement material to that which holds the shell-plates together in E. rotunda (Hedley and Ogden, 1973). This has been confirmed from the present authors unpublished observations of the fine structure of E. strigosa. The spines are single structures, although two or more are often seen projecting from the same junction as though they were connected. They usually project from the surface at right-angles, but they may lie close to the surface and point in any direction. There appears to be no regular arrangement in the distribution of the spines, although usually only the smaller spines are seen at the junction of the apertural-plate and shell-plate region, and the largest spines seem to lie along the lateral margins. The spines appear to be removed easily by abrasive action (PI. 2, fig. B). No abnormal forms of this species have been seen in our cultures. REPRODUCTION Euglypha acanthophora. Growth curves produced from observations made on three replicate cultures gave an estimated doubling time of between 2-3 and 2-8 days. Cytoplasmic division proceeds in a similar sequence to that previously described for E. rotunda by Hedley and Ogden (1973) . The approximate time taken to produce the daughter shell is 25 minutes, whereas the time taken for the completion of binary fission is two hours. The animals are usually active after division with strong pseudo- podial movement (Fig. 2) . Eight specimens with abnormal apertures were isolated and after six days had divided to produce 27 normal individuals. Observations on individual, dividing abnormal forms, such as those with a wide aperture and others being half normal size, show that they often produced smaller individuals with apparently the normal shell shape. One abnormal animal at division produced an almost normal shell, a small shell and some loose shell-plates (Fig. 3). Cytoplasm was present in the larger daughter shell, and the group did not separate after the normal time taken for division, but moved after this time as a group. During division in a normal individual, at the time when all the reserve shell-plates had passed into the pseudopodial trunk, a small circle of shell-plates became detached from the posterior end of the daughter shell. Binary fission proceeded normally and the two individuals separated, leaving behind the circle of shell-plates. Euglypha strigosa. An estimated doubling time of between 3-9 and 4-3 days was obtained from growth curves produced from observations on three replicate cultures. Division is preceded by the retraction of the filose pseudopodia, followed by the extrusion of a large pseudopodial trunk and shortly after by the apertural-plates (PI. 2, fig. D). The shell-plates are then arranged progressively until the daughter- shell is almost complete, at which point the spines are pushed out between the shell- plates with considerable movement. This continues for a short time after the shells are equal in size, but then the movement of both shell-plates and spines ceases abruptly. The time taken to produce a daughter-shell is approximately 50 minutes, with complete binary fission taking about three hours. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) 109 / 10 FIG. 2. Drawings of a moving specimen of E. acanthophora observed over a period of 30 minutes, to show the diversity of pseudopodial movement ; the arrow denotes the direction of movement. It was noted that in preparations of dividing specimens for scanning electron microscopy, the daughter-shell consistently collapsed. This is possibly due to an incomplete stabilization of the organic cement lining. no R. H. HEDLEY, C. G. OGDEN & J. I. KRAFFT FIG. 3. Diagram representing the division products of an abnormal form ; A - abnormal form, B - apparently normal daughter and C - a small form without cytoplasm. ACKNOWLEDGEMENT We would like to acknowledge the technical assistance of Miss E. M. Sides, who was responsible for collecting and isolating E. strigosa. REFERENCES EARTHS, E. 1963. Rhizopoden einiger Moosproben aus Java. Ada Universitatis Carolin. : 119-190. BLOCHMANN, F. 1887. Zur Kenntnis der Fortpflanzung von Euglypha alveolata Duj . Morph. Jb. 13:173-183. BONNET, L. 1966. Le peuplement thecamoebien de quelques sols du Chili (I). Protistologica 2 : 113-139. CAMBAR, R., LE BLANC, M., MERCIER, M. & THOMAS, R. 1964. Etude en microscopie electroni- que des microstructures superficielles de Protistes fossiles ou vivants. C. r. Acad. Sci. (Paris), 258 : 3554-3555- CARTER, H. J. 1865. On the fresh- and salt-water Rhizopoda of England and India. Ann. Mag. nat. Hist. (3) 15 : 277-293. 1889. Ramulina parasitica, a new species of fossil foraminifera infesting Orbitolites mantelli, var. theobaldi, with comparative observations on the process of reproduction in the Myce- tozoa, freshwater rhizopoda and foraminifera. Ann. Mag. nat. Hist. (6) 41 : 94-101. CASH, J., WAILES, G. H. & HOPKINSON, J. 1915. The British Freshwater Rhizopoda and Heliozoa. Vol. 3, Rhizopoda, part 3. The Ray Society London. 156 pp., 24 pis. CHARDEZ, D. & LECLERCQ, J. 1963. Variabilite des populations d' Euglypha strigosa (Ehren- berg) Leidy en fonction de 1'habitat (Rhizopoda testacea). Bull. Inst. agron. Stns Rech. Gembloux, 31 : 21-27. CLONAL CULTURES OF EUGLYPHA (PROTOZOA) in DECLOITRE, L. 1962. Le genre Euglypha Dujardin. Arch. Protistenk. 106 : 51-100. 1964. Thecamoebiens de la Xlleme Expedition Antarctique Fran9aise. Territoire des Terres Australes et Antarctique Francaises. Expeditions Polaires Fran9aises (Missions Paul-Emile Victor). Publ. No. 259 : 1-47. 1965. Amoebida testacea (Rhizopoda). Zoology Iceland, 2 (i) : 1-58. GRAAF, FR. de 1956. Studies on Rotatoria and Rhizopoda from the Netherlands. I. Rotatoria and Rhizopoda from the 'Grote Huisven'. Biol. Jaarb. 23 : 145-217. GRUBER, A. 1881. Der Theilungsvorgang bei Euglypha alveolata. Z. wiss. Zool. 35 : 431-439. HARRIS, R. H., MARTIN, B. S. & OGDEN, C. G. 1972. Notes on the preparation of natural history specimens for scanning electron microscopy. Bull. Br. Mus. nat. Hist. (Zool.) 24 : 223-228. HEAL, O. W. 1961. The distribution of testate amoebae (Rhizopoda : Testacea) in some fens and bogs in Northern England. /. Linn. Soc. (Zool.) 44 : 369-382. 1962. The abundance and micro-distribution of testate amoebae (Rhizopoda : Testacea) in Sphagnum. Oikos, 13 : 35-47. HEDLEY, R. H. & OGDEN, C. G. 1973. Biology and fine structure of Euglypha rotunda (Testacea : Protozoa). Bull. Br. Mus. nat. Hist. (Zool.) 25 : 119-137. 1974. Observations on Trinema Hneare Penard (Testacea : Protozoa). Bull. Br. Mus. nat. Hist. (Zool.) 26 : 185-199. HOOGENRAAD, H. R. & DE GROOT, A. A. 1940. Fauna van Nederland. Vol. 9, Zoetwater- rhizopoden en Heliozoen. Leiden. 302 pp. JONES, P. M. 1929. Indications of a flagellate stage in Euglypha alveolata. Am. Nat. 63 : 565- 570. LEIDY, J. 1877. The birth of a Rhizopod. Proc. A cad. nat. Sci. Philad. 29 : 261-265. 1879. Freshwater Rhizopods of North America. In: Vol. 12, United States Geological Survey of the Territories. Washington. 324 pp. LOEBLICH, A. R. & TAPPAN, H. 1961. Suprageneric classification of the Rhizopodea. /. Paleontology, 35 : 245-330. PENARD, E. 1890. Etudes sur les Rhizopodes d'eau douce. Mem. Soc. Phys. Hist. nat. Geneve, 31 : 1-230. 1902. Faune Rhizopodique du Bassin du Leman. Geneva. 700 pp. POPOFF, M. 1912. Uber die geschlechtliche Fortpflanzung von Euglypha alveolata Duj. Arch. Protistenk. 25 : 8-26. PRESCOTT, D. M. & JAMES, T. W. 1955. Culturing of Amoeba proteus on Tetrahymena. Exp. Cell. Res. 8 : 256-258. REUKAUF, A. 1912. Zur Encystierung von Euglypha alveolata. Zool. Anz. 39 : 372-375. SCHEWIAKOFF, W. 1 888. Uber die karyokinetische Kerntheilung der Euglypha alveolata. Morph. Jb. 13 : 193-258. SxfipANEK, M. 1963. Die Rhizopoden aus Katanga ( Kongo- Afrika). Annls Mus. r. Afr. cent. 117 : 9-91. Dr R. H. HEDLEY C. G. OGDEN JUNE I. KRAFFT BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD 5*t PLATE i A. Lateral view of E. acanthophora showing the arrangement of the shell-plates. x 1500 B. Preparation of shell-plates to illustrate two elongated shell-plates. X 3000 C. Lateral view of single apertural-plate with characteristic teeth. X 7600 D. Aperture of E. acanthophora showing the arrangement of the apertural-plates. x 2900 E. Ventral view of apertural-plate preparation, to show thickened dentate edge and the arrangement of three rows of aperture-plates. X 3500 Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE i PLATE 2 A. Lateral view of E. strigosa to show the distribution of siliceous spines. x 1200 B. Area of shell surface illustrating the holes left by removal of spines. x 5000 C. Lateral view of a single apertural-plate. X 4700 D. Apertural view of the first stage of division ; note the large pseudopodial trunk (arrowed) with some daughter-shell apertural-plates arranged on it. X 2700 E. Aperture of E. strigosa with eleven evenly spaced apertural-plates. x 3000 Bull. Brit. Mus. nat. Hist. (Zool.) 27, 2 PLATE 2 D DESCRIPTIONS OF THREE SPECIES OF EUPLOTES (PROTOZOA : CILIATEA) By C. R. CURDS INTRODUCTION IN HIS revision of the order Hypotrichida Stein, 1859, Borror (1972) listed 43 species of the genus Euplotes Ehrenberg, 1830 and in the same year Carter (1972) added a further four new species. The latter author suggested the following characters constitute a reliable basis for separating species within the genus ; the pattern of the dorsal interkinetal silver-line network or argyrome, the number of dorsolateral kinetics, the shape of the adoral zone of membranelles and the number of membran- elles therein, the number and arrangement of the ventral cirri and the shape of the non-dividing macronucleus. Most of these characters were initially introduced by Tuffrau (1960) while Borror (1968) added the appearance of the cortical sculpturing that is sometimes a feature of the dorsal and ventral surfaces of Euplotes spp. Three species of Euplotes are described in the present paper. The first two are freshwater forms that were isolated from samples of activated sludge and these were subsequently identified as E. moebiusi forma quadricirratus and E. affinis forma tricirratus respectively. The third is a euryhaline species from Austria which does not conform to any of those species described in the literature when both traditional and modern criteria are taken into consideration. Euplotes moebiusi Kahl, 1932 and the variety with four caudal cirri, E. moebiusi forma quadricirratus Kahl, 1932, have not been described in the literature since their first brief descriptions and illustrations by Kahl (1932). However, this species has regularly been observed in activated-sludge plants treating sewage and industrial waste waters over many years (Curds & Cockburn, 1970 ; Ministry of Technology 1968). Photographs of E. moebiusi were published by Klein (1958) in order to demonstrate the 'dry' silver method but these were not sufficiently comprehensive for taxonomic purposes. Euplotes affinis Dujardin, 1841 and its variety with three caudal cirri E. affinis forma tricirratus Kahl, 1932 are also examples that have not been described since their originals and yet have been seen regularly in aerobic waste-treatment processes (Curds & Cockburn, 1970) and in other organically polluted situations (Bick, 1972). Tuffrau (1960) thought it likely that both E. moebuisi and E. affinis were synonyms for the species E. charon Ehrenberg, 1830. MATERIALS AND METHODS (a) Source and cultivation A clonal culture of Euplotes moebiusi was isolated direct from an activated-sludge sample obtained from Maple Lodge Sewage Treatment Works, Rickmansworth, Hertfordshire. This species was maintained in freshwater Erdschreiber solution Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 6* n 4 C. R. CURDS ('Medium i', Committee on Cultures, Society of Protozoologists, 1958), either in test-tubes or in Petri dishes. The largest populations were obtained in Petri dishes containing a thin layer of Musgrave and Clegg's agar (2-5 per cent agar, 0-5 per cent sodium chloride and 0-5 per cent Liebig's beef extract in distilled water) which was streaked with the bacterium Klebsiella aerogenes (National Collection of Industrial Bacteria, NCIB 8017) as a food supply and then flooded with Erdschreiber solution. Subcultures were transferred at monthly intervals. Euplotes affinis was collected by Mr A. Cockburn from a sample of activated sludge taken from an experimental small-scale pilot plant operated at the Water Pollution Research Laboratory, Stevenage, Hertfordshire. It was sent to the British Museum (Natural History) as a clonal culture and was maintained in a similar manner to E. moebiusi. The third hypotrich, a small euryhaline Euplotes sp., was originally collected by Professor E. Tschermak from a freshwater source in Schlospark Schonbrunner in Vienna which is the locus classicus for the alga Ruttnera spectabilis Geitler, 1942 (see Geitler, 1942, 1943). Samples of this alga were sent to Dr Mary Parke at the Marine Biological Station at Plymouth where the hypotrich was first noticed and cultured. Cultures of the ciliate were subsequently deposited with the NERC Culture Collection of Algae and Protozoa where it was cultivated in saltwater Erdechreiber solution. Later, a clonal culture was established in freshwater and marine media at the British Museum (Natural History) and the descriptions herein relate to organisms from that clone. This small Euplotes sp. could be maintained equally well in test-tubes or plastic Petri dishes containing either fresh or sea water Erdschreiber solutions. Cul- tures were kept in the dark at room temperature, and were fed at weekly intervals with a few drops of a thick suspension of baker's yeast. Cultures were transferred at monthly intervals. (b) Microscopy Light microscopy and the methods used for observations and measurements were similar to those described by Curds, West & Dorahy (1974). Silver-line systems were displayed using the 'wet' method of Corliss (1953) in the cases of Euplotes affinis and the small euryhaline species. The 'dry' method of Klein (1958) was used to show the silver-line system of E. moebiusi. The latter method proved to be far more reliable and quicker than the conventional 'wet' method. As Klein (1958) pointed out, the success of the 'dry' method depends on the cell drying and dying more or less simultaneously and this was achieved by removing excess moisture with the aid of screws of paper tissue and by flicking single cells out from drops of liquid onto dry parts of the slide by means of an eyelash mounted in a glass rod. Nuclei were stained using Dippell and Chao's modification of De Lamater's basic fuchsin method described by Sonneborn (1950). The nuclei of E. moebiusi and E. affinis were stained after fixation on the slide by air-drying with equal success as the con- ventional method of chemical fixation. The techniques used for scanning-electron microscopy of the small euryhaline Euplotes sp. were similar to those previously described by Curds et al. (1974) with the exception of the fixation methods. Here the hypotrichs were not killed in osmium THREE SPECIES OF EUPLOTES 115 tetroxide vapour, and fixation was best using the osmium-meruric chloride fixative (Parducz, 1967) which was recommended by Small & Maraszalek (1969). A com- parison of the results obtained by the fixation methods of Curds et al. (1974) and those of Small & Marszalek (1969) for this species is demonstrated in PI. i. Plate la shows a cell fixed in Parducz's solution following the recommendations of Small & Mars- zalek (1969) where PI. ib shows a similar cell that had been killed in osmium vapour fixed in a solution containing equal parts of 2 per cent (w/v) osmium tetroxide and saturated mercuric chloride solutions. It is apparent that the Parducz's fixative was far better for this species than were the methods of Curds et al. (1974), whereas the reverse was true for the species Euplotes rariseta Curds, West & Dorahy, 1974. These results suggest therefore that the choice of fixative may vary with the species under consideration. RESULTS Euplotes moebiusi Kahl, 1932 DIAGNOSIS. Medium (60 /-mi long, 40 /Am wide), ovoid freshwater hypotrich with 10 fronto ventral, 5 transverse and 4 caudal cirri. Ventral surface heavily sculptured with 7 ridges, dorsal surface with 5 longitudinal ridges. Adoral zone with 35-40 membranelles which extend two-thirds the length of the cell. Dorsal silver-line system with 5 longitudinal rows of narrow polygons interspersed with an irregular network of larger polygons ; 7 dorsolateral kinetics bearing a maximum of n dorsal cilia. Macronucleus 3-shaped, micronucleus small, situated anteriorly. Slides showing silver-line systems, ventral ridges and nuclei have been deposited in the slide collection of the B.M. (N.H.), Reg. Nos. 1973:4:14:1-5. DETAILED DESCRIPTION. It can be seen from the size distribution data given on Fig. i that Euplotes moebiusi is a medium-sized species and is 62-25 6'6 /u,m long and 39-45 5-87 /u,m wide. The outline shape of the body is oval and there is a definite notch at the anterior end of the body where the adoral zone of membranelles (AZM) begins. The ventral surface is heavily sculptured with 7 ridges. (Fig. 2). One flattened ridge runs along the edge of the peristome and terminates posteriorly in a sharp point. One short ridge is restricted to the anterior half of the body and lies between the front-ventral cirri separating cirrus streak I and II from streak III (using the method of cirrus numeration of Wallengren, 1900). Three short ridges are confined to the posterior and lie between the transverse cirri. One ridge stretches the entire length of the body beginning at the anterior notch (between streaks III and IV) and terminating between the transverse cirri III i and IV i. One medium length ridge is restricted to the central portion of the body and separates cirri V 3 and VI 2 from cirrus V 2. The positions and shapes of these ridges are similar to those figured by Kahl (1932). There are 5 longitudinal ridges on the dorsal surface. Euplotes moebiusi has 10 frontoventral cirri which are distributed as shown in Figs . 2 and 3b . The arrangement resembles that of E . char on . There are 5 transverse and 4 caudal cirri. No specimens were observed with 3 caudal cirri as was shown in the original descriptions by Kahl (1932). The AZM is composed of 35-40 membran- elles and its extends two-thirds of the way down the body (Fig. 2). 6** n6 C. R. CURDS I 99- S 98 ~ 3. 95- ^-290- CO 80- > >- CO -4. = ;-= 6oH 01 La <-> 03 i: -a 40- 20- 20 30 40 50 SIZE (microns) 60 70 FIG. I. Size distribution data of Euplotes moebiusi (broken lines) and Euplotes affinis (continuous lines) . Triangles denote width of cells, circles denote length. FIG. 2. Ventral aspect of Euplotes moebiusi showing cirri and ridges (scale represents 10 pim). THREE SPECIES OF EUPLOTES 117 The geometry of the dorsal silver-line system of Euplotes moebiusi differs from any of those published. It consists (Fig. 3a) of 5 ladder-like longitudinal rows of narrow polygons with the pits of the dorsal cilia or bristles positioned on the right. In other Euplotes spp. with a double patella-type of dorsal argyrome the dorsal pits are situated on the left of the rows of smaller polygons. In addition, an irregular network of polygons, resembling that of E. mutabilis Tuffrau, 1960, is sandwiched between the ladder-like rows. There are 7 dorsolateral kinetics with the central kinetics bearing a maximum of n dorsal bristles. The ventral silver-line network (Fig. 3b) is a conventional series of irregular polygons whose general pattern closely resembles that of E. patella Muller, 1773 (see Tuffrau, 1960). a b FIG. 3. Silver-line system of Euplotes moebiusi, (a) dorsal surface, (b) ventral surface. The macronucleus of Euplotes moebiusi is an irregular 3 -shape (Fig. 4) resembling that of E. plumipes Stokes, 1884 (see Tuffrau, 1960), except that the posterior tail is shorter than in that species. The micronucleus is small and lies very close to, and sometimes overlaps, the left anterior edge of the macronucleus. Euplotes affinis Dujardin, 1842 Small (38 /um long, 26 /*m wide), ovoid freshwater hypotrich with DIAGNOSIS. 9 fronto ventral, 5 transverse and 3 caudal cirri. Ventral surface sculptured with 3 prominent ridges and dorsal surface with 5 longitudinal ridges. AZM with 18-20 n8 C. R. CURDS FlG. 4. M Ventral view of the nuclei of Euplotes moebiusi from a stained preparation (M = macronucleus, m = micronucleus) . membranelles which extends two-thirds the length of the cell. There is a small undulating membrane. Dorsal silver-line system of the double eurystomus type with 7 dorsolateral kinetics and a maximum of 9 dorsal cilia in the central kinetics. The macronucleus is 3 -shaped and there is a small anterior micronucleus. Slides showing silver-line systems and nuclei have been deposited in the slide collection of the B.M. (N.H.), Reg. Nos. 1973:9:26:1-10. DETAILED DESCRIPTION. Euplotes affinis is one of the smaller freshwater species whose dimensions are 38-4 4-3 /^m long and 25-8 4-0 /mi wide. The size distribu- tion data of this species are compared with those of E. moebiusi in Fig. i. The outline shape of E. affinis resembles that of E. moebiusi and there is a marked notch at the anterior of the cell which denotes the origin of the AZM. The ventral surface is heavily sculptured by 3 longitudinal ridges (Fig. 5) which travel almost the entire length of the cell. The outer pair of ventral ridges flank the transverse cirri at the posterior end of the body and terminate anteriorly between cirrus streaks I and II and between streaks V and VI. There are 3 other minor ridges that are restricted to the posterior end of the cell and these lie between the transverse cirri. The positions and shapes of the complete ventral ridging conforms closely with those figured by Kahl (1932). The dorsal surface is also heavily sculptured with 5 longi- tudinal ridges. Euplotes affinis has 9 frontoventral cirri whose distribution is shown in Figs. 5 and 6b. There are 5 transverse cirri and 3 caudal cirri ; one of the caudals is larger than the others and is held out stiffly to the right in a manner similar to that of E. rariseta (see Curds et al. 1974). No specimens have been observed with 4 caudal THREE SPECIES OF EUPLOTES 119 cirri as was shown in the original descriptions of E. affinis by Dujardin (1841), although Kahl (1932) described the variety E. affinis forma tricirratus which had 3 caudal cirri. The AZM of E. affinis extends two-thirds the length of the body and is composed of 18-20 membranelles which is approximately half the number found in E. moebiusi. m FIG. 5. Ventral aspect of Euplotes affinis showing cirri, ridges and nuclei (scale represents 10 (zm, M = macronucleus, m = micronucleus). The dorsal and ventral silver-line systems are shown in Fig. 6. The geometry of the dorsal argyrome (Fig. 6a) is of the double type resembling that of Euplotes eurystomus Wrzesniowski, 1870. There are 7 dorsolateral kinetics in E. affinis and the central kinetics bear a maximum of 9 dorsal cilia. The ventral silver-line system is of a common type consisting of a series of relatively few polygons. The macronucleus is 3-shaped and resembles those of Euplotes harpa Stein, 1859 and E. plumipes (see Tuffrau, 1960). The micronucleus is small and is situated at the anterior edge of the macronucleus. Euplotes parkei sp. n. DIAGNOSIS. Small (41 /zm long, 30 /mi wide) euryhaline species ; broadly oval in outline. Dorsal surface with 6 low longitudinal ridges and ventral surface with 7 minor ridges. AZM approximately two-thirds body length, composed of 18 mem- branelles. A deep pocket near the cytostome bears an undulating membrane. Usually 8, but rarely 9, fronto ventral cirri ; 5 transverse and 4 caudal cirri. There are 8 dorsolateral kinetics with a maximum of n dorsal cilia in the central kinetics. C. R. CURDS Dorsal silver-line system of the double eurystomus-type. Macronucleus C-shaped with anteriorly situated micronucleus. Type slides showing silver-line systems and nuclei have been deposited in the slide collection of the B.M. (N.H.). Holotype Reg. No. 1973:4:2:1, and paratype Reg. Nos. 1973:4:2:2-5. a b FIG. 6. Silver-line system of Euplotes affinis, (a) dorsal surface, (b) ventral surface. DETAILED DESCRIPTION. This is a small euryhaline species (41-215-6 /urn long, 30-615-3 /u,m wide) whose size distribution data are given in Fig. 7. It is broadly oval in outline shape with the dorsal surface sculptured with 6 relatively low longi- tudinal ridges. To the left of each ridge is a parallel row of pits from which the short (2 /urn) dorsal bristles emerge (PI. i, figs, c & d). The ventral surface also bears 7 longitudinal ridges (PI. i, fig. a) but these are not as prominent as in Euplotes moebiusi. One ventral ridge travels the complete length of the body along the extreme edge of the peristome, while the other 6 are relatively short and are confined to the posterior half of the cell. The transverse cirri arise from between these 6 minor ridges. The AZM extends two-thirds the length of the body and is composed of 18 membranelles. There is an undulating membrane situated in a deep pocket on the right of the peristome in the proximity of the cytostome. This species of Euplotes usually bears 8 frontoventral cirri which are arranged as shown in Figs. 8, gb and PI. i, fig. a ; however, a gth frontoventral cirrus is occasion- ally present within the same clone and this lies in a position V 2 (PI. i, fig. b). The 9th frontoventral cirrus has been found only in animals cultured in freshwater Erdschreiber solution even though the marine cultures have been searched thoroughly THREE SPECIES OF EUPLOTES 121 = . 99- CO 98- o. o 95- ^ QJ "s 90n o CO as 80- CD +- t 60-| cB -g 4 H Q_ 2 Q_ a> 03 20- 10- 5- 2- 1- 20 30 40 SIZE (microns) 50 60 FIG. 7. Size distribution data of Euplotes parkei. Triangles denote width and circles length of the cells. for this variant. It is not yet known if the salinity of the culture plays any part in promoting this type of intraspecific polymorphism. The 5 long transverse and 4 caudal cirri were constant in their numbers. There are 8 dorsolateral kinetics in this species and the central kineties bear a maximum of n dorsal cilia. The dorsal and ventral silver-line systems are shown in Fig. 9. The disposition of the dorsal argyrome (Fig. ga) is of the double type resembling that of Euplotes eurystomus. The ventral silver-line system consists of a series of few but large polygons and in this respect resembles that of E. cristatus Kahl, 1932 (see Tuffrau, 1960). The dorsal argyrome can be seen on some scanning- electron micrographs (PI. i, figs, c & d) as a series of tiny specks. The macronucleus (Fig. 8) is C-shaped and the micronucleus is situated close to the left anterior edge of the macronucleus. DISCUSSION AND CONCLUSIONS Euplotes moebiusi is one of the nine species of the genus whose silver-line system had not been fully described, and its identity relied solely upon the brief description of Kahl (1932). It is evident from the description of the silver-line system given in 122 C. R. CURDS FIG. 8. Ventral aspect of Euplotes parkei showing cirri, ridges and nuclei (scale represents 10 pun, M = macronucleus, m = micronucleus) . a b FIG. 9. Silver-line system of Euplotes parkei, (a) dorsal surface, (b) ventral surface. THREE SPECIES OF EUPLOTES 123 the present paper that E. moebiusi is a distinct species and is not a synonym of E. charon as was suggested by Tuffrau (1960). Before the completion of the present studies it was the opinion of the author that Euplotes moebiusi was likely to be a synonym of E. affinis because of the similarities between the two species which may be listed thus ; they both have the same general shape and size with a definite anterior notch ; both have ventral ridges of similar appearance ; both E. moebiusi and E. affinis are reported to have 4 and 3 caudal cirri variants called E. moebiusi forma quadricirratus and E. affinis forma tricirratus respectively ; both have 3 -shaped macronuclei and both may be found in organically polluted freshwater habitats. This meant that there was only one known character left to separate the two species, namely the presence or absence of frontoventral cirrus V 2. In view of the findings presented in this paper concerning the intra- specific polymorphism of E. parkei due to the variability of cirrus V 2 it is evident that the presence or absence of this cirrus is not as reliable a character as was origin- ally believed. The results presented here however have clarified the situation considerably and E. affinis can now be distinguished from E. moebiusi by the 5 characters listed in Table i. The most reliable character is the geometry of the dorsal silver-line system which is completely different in the two species (compare Figs. 3b & 6b). In the author's opinion E, affinis should not be regarded as a synonym of E. charon as was suggested by Tuffrau (1960), and there are suffi- cient reliable characters to regard E. affinis as a species distinct from all others. TABLE i List of characters which serve to distinguish between Euplotes moebiusi and Euplotes affinis Character Number of frontoventral cirri Number of membranelles in AZM Maximum number of dorsal cilia in mid- dorsolateral kinetics Dorsal argyrome Ventral argyrome E. moebiusi 10 35-40 ii Complex E. affinis 9 18-20 Double eurystomus Many small polygons Few large polygons Agamaliev (1967) reported intraspecific polymorphism in the number of fronto- ventral cirri in his description of Euplotes raikovi Agamaliev, 1966. In the Caspian Sea strain of E. raikovi, Agamaliev (1967) noted that there were 7 or 8 frontoventral cirri and indicated that cirrus V 2 was that which did not develop in some specimens. However in a recent paper, Washburn & Borror (1972) described a strain of E. raikovi, isolated from a sand sample taken from the New Hampshire coast of the U.S.A., in which they could not find an 8th cirrus (cirrus V 2) although they did observe a completely barren plaque in each case. Negative evidence such as this can never be conclusive and one must accept that Agamaliev's strain did exhibit polymorphism of cirrus V 2 as he claimed, particularly in the light of the photo- graphic evidence presented in this paper where there can be no doubt that E. parkei may have 8 or 9 completely normal frontoventral cirri. More work must be carried 124 C - R - CURDS out on the morphogenesis of E. parkei particularly on the fate of cirrus V 2, but the evidence so far obtained indicates that there is not even a barren plaque in the case of E. parkei specimens with 8 frontoventral cirri. Furthermore, more work is needed to test adequately whether or not intraspecific polymorphism such as this can be induced by adjusting the salinity of the culture medium or if the results so far obtained can be attributed to pure chance. It is possible that one of the reasons why Euplotes parkei has remained unnoticed until now is because none of its characters, in isolation, will distinguish this species from all others. A combination of characters is required to do so, but there can be little doubt that this small euryhaline Euplotes is a separate and distinct species. The following species have the combination of characters - a double dorsal argyrome with 8 or 9 frontoventral, 5 transverse and 4 caudal cirri - E. aediculatus Pierson, 1943 ; E. amieti Dragesco, 1970 ; E. aspheronicus Agamaliev, 1966 ; E. diadaleos Diller & Kounaris, 1966 ; E. eurystomus ; E. octocarinatus Carter, 1972 ; E. patella ; E. patella forma latus Agamaliev, 1967 ; E. plumipes ; E. tegulatus Tuffrau, 1960 ; E. tuffraui Berger, 1965 ; E. variabilis Stokes, 1887 (see Carter, 1972) ; and E. zenkewitchi Burkovsky, 1970. However only four of these, E. aediculatus, E. eurystomus, E. variabilis and E. octocarinatus, have 8 dorsolateral kinetics. Since E. octocarinatus has a patella-like double dorsal argyrome there are only three remain- ing species with which to compare and contrast E. parkei. All three of these Euplotes spp. are large (over 100 jum long) and have at least 40 membranelles in the AZM, whereas E. parkei is small (under 50 /um long) and has less than 20 mem- branelles. Furthermore, the shapes of the macronuclei differ and all three species have many more dorsal cilia than E. parkei. It is evident therefore that Euplotes parkei differs from all previously described species of Euplotes and the differences are considered to be sufficiently distinct for this organism to be designated as a separate species. It is named Euplotes parkei after Dr Mary Parke of the Marine Biological Station Plymouth who first isolated and cultivated this hypotrich. Following the revised classification of the Committee on Taxonomic Problems of the Society of Protozoology (Honigberg et al., 1964), Euplotes parkei is placed into class Ciliatea Perty, 1852, order Hypotrichida Stein, 1859, family Euplotidae Ehrenberg, 1838. ACKNOWLEDGEMENTS The author is indebted to Mrs J. E. Dorahy, Department of Zoology, British Museum (Natural History), for her technical assistance, to Miss B. J. West, NERC Culture Centre for Algae and Protozoa, who made the original silver preparations of Euplotes parkei, to Mr A. Cockburn, Water Pollution Research Laboratory of the Department of the Environment, who isolated E. affinis, to Miss M. Yeoh, Maple Lodge Sewage Treatment Works, Rickmansworth, who supplied samples of activated sludge containing E. moebiusi and to Mr R. Harris, Department of Zoology, British Museum (Natural History), for his aid and advice on the preparation of ciliates for scanning-electron microscopy. THREE SPECIES OF EUPLOTES 125 REFERENCES AGAMALIEV, F. G. 1967. Faune cilies mesopsammiques de la cote ouest de la Met Caspienne. Cahiers de Biol Mar. 8 : 359-402. BICK, H. 1972. Ciliated Protozoa. An illustrated guide to the species used as biological indicators in freshwater biology. World Health Organization, Geneva. BORROR, A. C. 1968. Systematics of Euplotes (Ciliophora, Hypotrichida) ; towards union of the old and the new. /. Protozool. 15 (4) : 802-808. - 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Protozool. 19 (i) : 1-23. CARTER, H. P. 1972. Infraciliature of eleven species of the genus Euplotes. Trans. Amer. Microsc. Soc. 91 (4) : 466-492. COMMITTEE ON CULTURES, SOCIETY OF PROTOZOOLOGISTS. 1958. A Catalogue of Laboratory Strains of Free-living and Parasitic Protozoa. /. Protozool. 5 (i) : 1-38. CORLISS, J. O. 1953. Silver impregnation of ciliated protozoa by the Chatton-Lwoff technic. Stain Tech. 28 : 97-100. CURDS, C. R. & COCKBURN, A. 1970. Protozoa in biological sewage-treatment processes - I. A survey of the protozoan fauna of British percolating niters and activated-sludge plants. Wat. Res. 4 : 225-236. WEST, B. J. & DORAHY, J. E. 1974. Euplotes rariseta sp. n. (Protozoa, Ciliatea), a new small marine hypotrich. Bull. Br. Mus. nat. Hist. (Zool), 27 (2) : 95-102. DUJARDIN, F. 1841. Histoire Naturelle des Zoophytes. Infusoires. Paris. 678 pp. GEITLER, L. 1942. Neue luftlebige Algen aus Wien. Ost. hot. Z. 41 (i) : 49-51. 1943- Eine neue atmophytische Chrysophycee, Ruttnera spectabilis, nov. gen., nov. spec. Int. Revue ges. Hydrobiol. 43 : 100-109. HONIGBERG, B. M., BURBANCK, W., BOVEE, E. C., CORLISS, J. O., GOJDICS, M., HALL, R. P., KUDO, R. R., LEVINE, N. D., LOEBLICH, A. R., WESIER, J. & WENRICH, D. H. 1964. A revised classification of the Phylum Protozoa. /. Protozool. 11 : 7-20. KAHL, A. 1932. Urtiere oder Protozoa. I : Wimpertiere oder Ciliata (Infusoria), eine Bearbeitung der freilebenden und ectocommensalen Infusorien der Erde, unter Ausschluss der marinen Tintinnidae. Un: Dahl, F. Die Tierwelt Deutschlands, Teil 25, pp. 399-650. G. Fischer, Jena. KLEIN, B. M. 1958. The 'dry' silver method and its proper use. /. Protozool. 5 (2) : 99-103. MINISTRY OF TECHNOLOGY. 1968. Protozoa in sewage-treatment processes. In: Notes on Water Pollution, No. 43. H.M.S.O., London. PARDUCZ, B. 1967. Ciliary movement and co-ordination in ciliates. Int. Rev. Cytol. 21 : 91- 128. SMALL, E. B. & MARSZALEK, D. S. 1969. Scanning electron microscopy of fixed frozen and dried Protozoa. Science, N.Y. 163 : 1064-1065. SONNEBORN, T. M. I95O. Methods in general biology and genetics of Paramecium aurelia. J. exp. Zool. 113 : 84-147. TUFFRAU, M. 1960. Revision du genre Euplotes, fondee sur la comparaison des structures superficielles. Hydrobiologia, 15 : 1-77. WALLENGREN, H. 1900. Zur Kenntnis der vergleichenden Morphologie der Hypotrichen. Bih. K. svenska Vetensk Akad. Handl. 26 : 1-31. WASHBURN, E. S. & BORROR, A. C. 1972. Euplotes raikovi Agamaliev, 1966 (Ciliophora, Hypotrichida) from New Hampshire : Description and morphogenesis. /. Protozool. 19 (4) : 604-608. Dr C. R. CURDS Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD 6*f PLATE i Scanning-electron micrographs of Euplotes parkei a. Ventral view showing ridges and cirri. Cell fixed in Parducz's (1967) fixative. b. Ventral view of a 9 frontoventral cirri variant. Cell killed and fixed following the methods of Curds et al. (1974). c. Dorsal view showing the outline of the dorsal argyrome as specks. d. Dorsal view showing ridges, dorsal cilia and outline of argyrome. Bull. Brit. Mus. nat. Hist. (Zool). 27, 2 PLATE i NEW RECORDS OF BATS FROM SOUTH- EASTERN ASIA, WITH TAXONOMIC NOTES By J. E. HILL INTRODUCTION BATS from Malaya, Java and Sulawesi (Celebes) identified in recent years at the British Museum (Natural History) have included specimens representing a number of poorly known species and thereby of taxonomic interest and importance, or which provide further distributional records. The majority of the Malayan specimens have come to London through the agency of Lord Medway and were collected by him or by Mr G. C. Yong. A further interesting specimen from Malaya has been provided by Dr D. R. Wells of the School of Biological Sciences, the University of Malaya. The Indonesian specimens are from a collection submitted for identifica- tion by Captain P. F. D. Van Peenen, M.C., U.S.N., Officer in Charge, U.S. Naval Medical Research Unit No. 2, Djakarta Detachment. My thanks are due also to Dr G. G. Musser and Dr Karl F. Koopman of the American Museum of Natural History, New York, who waived a prior claim to the Sulawesian specimens, and to Dr H. W. Setzer of the United States National Museum of Natural History, the Smithsonian Institution, Washington, who arranged the loan of one of the specimens discussed. Measurements are in millimetres : unless otherwise indicated, the specimens have been donated to the collections of the British Museum (Natural History). SYSTEMATIC SECTION ^ Chironax (?) melanocephalus (Temminck, 1825) Sulawesi : Soroako, south Sulawesi. $ (young adult) B.M. 73.1802. Chironax has been unreported hitherto from Sulawesi. This young adult specimen differs in a number of features from Malayan material referred to C. melanocephalus and apparently also in some ways from Javan specimens, of which none is available for comparison. Consequently, it is referred to C. melanocephalus with considerable hesitation. This Sulawesian specimen lacks the distinctive blackish cap usually characteristic of Chironax although the nape and crown are darker brown than the back which is warm brown, tinged with grey over the shoulders. The throat and the sides of the neck are creamy white, the belly yellowish white and the flanks brown, this colour extending across the hinder part of the ventrum. In colour the specimen agrees quite well with the original description by Temminck (1825 : 190) of specimens from Bull. Br. Mus. nat. Hist. (Zool.) 27, 2 7 128 J. E. HILL Java : it differs from mainland specimens in its generally paler dorsal surface and also lacks any rufous or orange at the sides of the neck, a feature sometimes found in mainland examples. There are some differences in wing structure when a comparison is made with mainland and Javan specimens. In particular, the fifth metacarpal is relatively a little longer, and the second phalanges of the third, fourth and fifth digits are rela- tively a little shorter, those of the fourth and fifth digits not exceeding in length the first phalanges of their respective digits as they do in continental and Javan examples. The relevant wing structures are summarized in Table i and in these respects it can be seen that the Sulawesian specimen approaches the closely related genus Balionycteris. TABLE i Wing indices of Chironax and Balionycteris Length of forearm = 1000 Chironax Balionycteris Malaya Java* Sulawesi Third digit Metacarpal 677-711 640 635 705 710-780 First phalange 474~533 4 8 9 4 88 57 474~54 8 Second phalange 584-657 640 623 568 559-653 Fourth digit Metacarpal 617-679 617 578 641 693-739 First phalange . 368-415 373 378 373 374-43 Second phalange 396-444 418 422 360 352-415 Fifth digit Metacarpal 659-704 640 622 708 712-764 First phalange 3i 8 ~36o 344 322 319 330-364 Second phalange 339~3 8 4 34 8 344 3*7 3^-354 * 'Co-types' of C. melanocephalus, from Andersen (1912 : 676). Cranially the specimen from Sulawesi is a little smaller than the mainland examples but its cranial dimensions fall within the range of variation reported for the Javan 'cotypes' by Andersen (1912 : 678). The supraorbital region is a little more swollen medianly than in continental specimens and the postorbital processes are more massively developed. As might be expected in a young adult, the premaxillae are not solidly fused anteriorly. The post-canine teeth are generally rather smaller than are those of specimens from the mainland, with pm 4 more rounded, less rectangular in outline, while pm 3 lacks completely the antero-external cusp customary in C. melanocephalus. This cusp, however, is very small in some Malayan specimens. Chironax melanocephalus has been reported from Java, Sumatra (Chasen, 1940 : 28, 30), Nias Island (Thomas, 1923 : 252), Malaya (Chasen, loc. cit. ; Hill, 1961 : 640) and from southern Thailand (Hill and Thonglongya, 1972 : 181), who also reported further Malayan specimens. The presence of Chironax in Sulawesi is not unexpected BATS FROM SOUTHEASTERN ASIA 129 but the exact taxonomic status of the Sulawesian population must remain to some extent uncertain until further specimens from Sulawesi and Java are available. Measurements of the specimen from Sulawesi : length of forearm 44-2 ; length of third metacarpal 31-1 ; length of its first phalange 22-4 ; length of its second phalange 25-1 ; length of fourth metacarpal 28-3 ; length of its first phalange 16-5 ; length of its second phalange 15-9 ; length of fifth metacarpal 31-3 ; length of its first phalange 14-1 : length of its second phalange 14-0 ; greatest length of skull 22-2 ; condylobasal length 21-4 ; condylocanine length 20-8 ; palatal length n-i ; palation to incisive foramina 9-3 ; palation to basion 8-2 ; length orbit-nares 5-2 ; width of braincase 9-8 ; mastoid width 10-0 ; zygomatic width 14-3 ; n^-m 1 (crowns) 6-3 ; p 4 -p 4 (crowns) 6-3 ; lachrymal width 6-1 ; c 1 -^ 1 (cingula) 4-3, (alveoli) 4-0 ; postorbital width 5-5 ; interorbital width 4-9 ; width of mesopterygoid fossa 2-9 ; width between p 4 -p 4 3-8 ; width between bases of canines 1-9 ; orbital dia- meter 5-6 ; length of mandible from condyle 16-0 ; length of complete mandible from both condyles 15-2 ; coronoid height 7-3 ; c-m 1 (crowns) 6-9 ; c-m 2 (crowns) 7-5 (in order of Andersen, 1912 : 678, with interpolations). Measurements of teeth : length x width of pm 3 1-7x1-1 ; of pm 4 1-7x1-2 ; of m 1 1-3x0-9: of pnij 0-7 x 0-7 ; of pm 3 1-6x1-1 ; of pm 4 1-7 x 1-2 ; of nij 1-4x0-9; of m 2 0-8x0-6 (notation of teeth based on Andersen, 1912 : 680). It may be noted that by error the illustration of Chironax in the editions and print- ings of Walker (1964, 1965, 1968) is not of C. melanocephalus but of a young Rousettus. ~ Rhinolophus pusillus Temminck, 1834 Malaya : Pasang Kamunting, Kg. Ginting, Penang. $ B.M. 73.608. Andersen (1905 : 121) summarized the many different forms of Rhinolophus before then confused under Rhinolophus minor Horsfield, 1824, envisaging a lepidus group composed of lepidus, minor and subbadius subgroupings. Later, the same author (1918 : 376) in a paper issued on his behalf by Oldfield Thomas renamed this the pusillus group and added a number of briefly diagnosed new forms. Tate and Archbold (1939 : 3) listed the names allocated to the group, retaining the sub- divisions proposed by Andersen. These authors included R. minor and R. pusillus as distinct species but Andersen (1905 : 126) considered pusillus a synonym of minor : however, Rhinolophus minor Horsfield, 1824 is preoccupied (Chasen, 1940 : 38 ; Ellerman & Morrison-Scott, 1951 : 116). The species has not hitherto been recorded from the Malay Peninsula : this speci- men from Penang agrees closely with a small series from Java and with one specimen from Madura identified by Andersen as R. pusillus, the species that he called R. minor in his early investigation (1905 : 121) of the group. There is close agreement also between the Penang specimen and a series in the collections of the British Museum (Natural History) from the island of Tioman, off the east coast of Malaya. Rhinolophus pusillus is small, easily recognized by its upright triangular connecting process and small skull (Table 2). 7* 1 3 o J. E. HILL it 41 fl rt ti C C too?* E2 o3 rt rt CH f* 3 g a g 2^ >>>>csS5ogg M ;- ;- ^ g ^ H H H E ui-o p I o O* N c^ O co ^* io vb vb vb vo o 9}.9jduioo jo l-t *$ ON M b I b ON b IO ON O O O ON ON O 8 UI-O vO CO OO vp CO >O IO IO if) if) 8 UI- 8 UI T o- t o OO OO CO CO I CO CO CO ON CO 00 ro ro ro ro -s; J "o 8 ft! 9S'BOUIBjq jo M op op ON 00 t^ vb vb vb I vb CO 00 IO M N p N c^ I c^ f^ cb ob ob cb HO O t*^ vO t^ 00 I CO CO CO CO | CO jo i i -) CO CO vp M H M jo cocococo rorororororo X9g CH-CH-CH-Of'ri^CH-CH-CH- I *O H . H . M . H O M M M ON 00 00 MVOOvO M 65 mm S.L. Length of lower jaw 40-8-51-0, M = 45-6 (37-6-46-2, M = 42-0) per cent of head, 1-6-2-0 (modal range 1-8-2-0) times longer than broad in specimens of all sizes. Length of upper jaw 31-8-44-0, M = 38-4 (30-6-37-6, M = 33-3) per cent of head. Mouth slightly oblique, the posterior tip of the maxilla reaching a vertical through the anterior margin of the orbit, or to a point slightly posterior to that line. Gill rakers. Of variable form, but usually with the upper 2 or 3 rakers on the first gill arch flattened and branched, those on the middle section of the arch relatively slender, and the lowermost I or 2 rakers reduced ; no pseudorakers are developed 156 P. H. GREENWOOD FIG. 8. Haplochromis macconneli. Dentition. A : Outer teeth from the premaxilla. B : Outer teeth from the dentary. In both, the teeth are from an anterolateral position in the jaw and are viewed labially. (see p. 143). There are 7 (fi), 8 (f8) or 9 (121) rakers on the lower limb of the first gill arch. Scales. Ctenoid, those on the chest very small but grading in size with those on the subpectoral region (that is, the demarcation between small and large scales in this region is less abrupt than in H. rudolfianus ; see p. 154). Five and a half to 7 (mode 6) scales between the dorsal fin origin and the lateral line, 7-9 (modes 8 and 9), rarely 5, between the pectoral and pelvic fin bases. Cheek with 3 or 4 rows of imbricating scales. In most specimens there is a small naked area (about the area of one scale) in front of the first dorsal fin spine. Fins. Dorsal with 22 (f6), 23 (fi6) or 24 (f8) rays, comprising 13 (12), 14 (fi6), 15 (fn) or 16 (fi) spinous and 8 (fn), 9 (fi8) or 10 (fi) branched rays. Anal with 3 spines and 7 (f2), 8 (25) or 9 (13) branched rays. First ray of the pelvic fin slightly produced, more so in larger fishes. Caudal truncate, scaled on its proximal quarter (mode) to third. Pectoral fin 25-8-34-5, M = 28-6 (23-7-28-3, M = 26-1) per cent of standard length, and 73-0-92-5, M = 81-1 (63-5-77-0, M = 72-3) per cent of head. Teeth. In fishes more than 50 mm S.L. the outer row of premaxillary teeth is composed of unequally bicuspid teeth anteriorly, but of tricuspids laterally and posteriorly (Text-fig. 8) ; the median cusp of the tricuspids, and the major cusp of the bicuspids is slightly incurved. Many specimens have some tricuspids intercalated amongst the biscuspids anteriorly, and in a few fishes almost the entire outer row is composed of tricuspid teeth. When there is a mixture of bi- and tricuspids the latter predominate. At all positions in the tooth row the bi- and tricuspids are of equal size. Tooth morphology and arrangement in the lower jaw are similar to those in the upper, although more individuals have only tricuspid teeth present ; a predominantly bicuspid outer row is rarely encountered. Fishes in the size range 20-40 mm S.L. have mainly bicuspid teeth in the upper jaw, with those tricuspids present restricted to a posterolateral position in the row. LAKE RUDOLF HAPLOCHROMIS SPECIES 157 B FIG. g. Haplochromis macconneli. A : Neurocranium in right lateral view. B : Neuro- cranium, dorsal surface. C : Bones of the infraorbital series, right side. D : Right lower jaw, viewed from a slightly ventrolateral position. Bicuspid teeth also predominate in the lower jaw, but some unicuspid (and slender) teeth may occur posterolaterally. Tooth form is like that of the larger fishes. There are 41-64 (M = 52) outer premaxillary teeth in fishes 50-77 mm S.L., and 24-42 in specimens 22-39 mm long. Irrespective of a fish's size, the inner teeth in both jaws are all tricuspid and small, and are usually arranged in a single series but double rows are encountered occasion- ally. OSTEOLOGY. The characteristic hypertrophy in the cephalic laterosensory canal system has been commented upon above (see also Text-figs. 7 and 9). Canal bones in the pectoral skeleton are also affected, and the otic region of the skull is noticeably inflated (see Text-fig. 9). The neurocranium (Text-fig. 9) departs from the generalized Haplochromis type (as seen in H. mdolfianus, Text-fig. 3) and clearly approaches that found in H. saxicola and allied species in the Lake Victoria species flock (Greenwood, 1974). In other words, the preotic region of the skull is slightly more elongate than in the generalized type, and associated with this and the shallower braincase, the preotic skull roof is straighter and slopes upwards at a smaller angle. The supraoccipital 158 P. H. GREENWOOD crest is lower than in most generalized types of skull, and the otic region is narrower. Although all the canal bones of the skull (and also those bones encasing the inner ear) are noticeably inflated, the dermopterotics show the greatest hypertrophy of all (see Text-fig. gA and B). The lower pharyngeal bone (Text-fig. 10) is triangular in outline (length and breadth of the dentigerous surface almost equal), is relatively slender, and has an anterior blade that is neither noticeably long nor noticeably short. The teeth on this bone are rather sparsely arranged in 16-20 irregular rows. Without exception, the teeth are weakly bicuspid, with a low, blunt or even barely visible anterior cusp and a crown that slopes gently into the sharper and larger posterior cusp ; all are fine and compressed but those in the two median rows are somewhat coarser. FIG. 10. Haplochromis macconneli. Lower pharyngeal bone in occlusal view. Vertebrae and the caudal fin skeleton. Vertebral counts for the 24 specimens radiographed are : 27 (fi), 28 (f8), 29 (fi4) or 30 (fi), comprising 12 (fi6) or 13 (f8) abdominal and 15 (fi), 16 (fi5) or 17 (f8) caudal centra. The fused first ural and first preural centra are excluded from these counts. The caudal fin skeleton is unusual in showing a high degree of fusion between various hypurals (or if not fusion, such extremely close apposition as to be in- distinguishable from fusion on radiographed specimens). The extent of fusion or apposition is generally complete, but in a few fishes there are short unfused sections between the otherwise conjoined elements. About 77 per cent of the 26 specimens radiographed showed some degree of fusion between various hypurals. Only 6 specimens (ca 23 per cent of the sample) had all 5 hypurals completely free from each other. Most frequently (i.e. in 20 fishes) hypurals I and 2 are fused. In the upper part of the skeleton the commonest pattern of fusion is to have hypurals 3 and 4 fused, but hypural 5 free (n specimens) ; only 4 specimens had all 3 upper hypurals fused, and 5 fishes showed no fusion between any of the 3 upper hypurals. In H. rudolfianus, it will be recalled, only 2 out of 30 specimens (i.e. about 7 per cent) had any fused hypurals ; in both these fishes fusion was between hypurals i and 2 (see p. 146). LAKE RUDOLF HAPLOCHROMIS SPECIES 159 For comparison with the situation in Lake Rudolf, the caudal skeleton in several endemic Haplochromis species from other lakes was examined. Of 174 specimens (representing 12 species) from Lake George, Uganda, about 8 per cent showed some (but never complete) fusion between elements in both the upper and the lower parts of the skeleton. The frequency of fusion seems equally distributed amongst the species examined. In Lake Victoria, too, hypural fusion is relatively rare. One hundred and seventy-eight specimens representing 22 species (with a modal sample size of 10 specimens per species) were examined. Of these, 20 specimens (i.e. about 13 per cent) had fused hypurals. The commonest pattern here is of fusion between hy- purals 3 and 4, less frequently it occurs between hypurals i and 2, and only once was it recorded, with certainty, between hypurals i and 2, and 3 and 4 in the same individual (although two other individuals may show this pattern). The Lake Victoria species examined are from several phyletic lineages within the Haplochromis flock of that lake and cover a wide range of body forms. Far fewer specimens are available of the four endemic Haplochromis species from Lake Albert. Of these, H . bullatus Trewavas has 3 out of 18 fishes (i.e. about 17 per cent) with hypurals i and 2 fused (or very closely apposed), while H. avium Regan (9 specimens) and H. albertianus Regan (15 specimens) have none. Both the latter species, however, have some individuals in which the hypurals are closely apposed. Thus, even allowing for the small sample sizes involved in this survey, it does seem that the frequency of hypural fusion in H, macconneli is exceptionally high. The significance of this phenomenon remains unexplained. VISCERA. Because of poor preservation it is impossible to measure precisely the length of the gut in H. macconneli ; I would estimate the intestine to be about i^ times longer than the standard length. In strong contrast to H. rudolfianus there is no trace of melanin in the peritoneal tissue and neither are the gonadial walls pig- mented (see p. 146 above). COLORATION. In life, adult females are a pale greenish-fawn with traces of greenish iridescence on the flanks. All fins are colourless except for three conspicuous and bright yellow spots on the anal. Live colours for males have not been recorded. Preserved colours. I suspect that the material I have examined is somewhat bleached and thus the coloration is probably lighter than it might otherwise have been. There is apparently little sexual dichromatism. In both sexes the ground colour of the body and head is a pale yellowish-fawn with no tonal variation between dorsum and venter. All fins are hyaline with, in both sexes, dark maculations on the proxi- mal third to half of the caudal fin. The males examined have the dorsal fin somewhat darker than that of the females, and there is a fairly dense aggregation of melano- phores on the anterior third of the pelvic fins. (These specimens may, however, be less bleached than are the females.) ECOLOGY. Judging from Mr Hopson's records, H. macconneli is confined to water more than 20 m deep, and is probably most abundant in deeper water (i.e. at depths 160 P. H. GREENWOOD of 50-70 m). Some of the smallest specimens collected (15-20 mm S.L.) were caught at a depth of 75 m, and none, of any size, has been recorded from littoral habitats. The pale coloration of this species and the absence of melanic pigments in the peritoneum and gonads may well be correlated with this deep-water distribu- tion. No data are available yet on the feeding habits of the species. Breeding biology. The marked sexual dimorphism in adult size has been noted already (p. 154). Despite a very thorough search, Mr and Mrs Hopson were at first unable to find any males amongst the several hundreds of H. macconneli they ex- amined. Eventually two males, 31-0 and 35-5 mm S.L., were identified in a catch made some 2-4 km east of North Island at a depth of 30 m. Both these fishes appear to be sexually adult ; one is probably in an advanced stage of ripening, the other at a slightly earlier stage of development. Females are certainly adult at a length of ca 50 mm and it seems likely that they may mature at a smaller size (Hopson in litt.}. Be that as it may, females seem to grow to a much greater length than do males. In a sample of 21 adult females examined, only 2 have both ovaries equally developed. Eleven fishes have the right ovary much larger than the left one (at all stages of oogenesis), and 8 have only the right ovary developed. Unlike H. rudolfi- anus, there is no trace of melanization in the ovarian wall of H. macconneli. No details are available on the breeding habits or seasons of the species. DIAGNOSIS AND AFFINITIES. Haplochromis macconneli is readily distinguished from the other Lake Rudolf species by the hypertrophy of its cephalic laterosensory canal system, its dentition (see p. 156), and in specimens > 35 mrn S.L., from H. rudolfianus by several morphometric characters (deeper cheek and preorbital, longer snout, smaller eye and longer jaws). Indeed, H. macconneli is so distinct morphologically from H. rudolfianus and H. turkanae (and the putative fourth species) that it is difficult to establish its phyletic affinities with these other taxa. In addition to the characters listed above, H. macconneli also differs from H. rudolfianus (and from the undescribed species too) in the shape of its neurocranium, which is of a more specialized type (see above, p. 157 and Greenwood, 1974). Almost certainly the hypertrophy of the cephalic lateral line canals is an adaptation associated with the deep-water habitat of H. macconneli, and strongly suggests that the species evolved within the lake. No fluviatile Haplochromis exhibits this specialization. It is interesting to note that the cephalic lateral line canal hypertrophy seen in H. macconneli is much greater than that found in any known Haplochromis species living at comparable depths in Lake Victoria (Greenwood & Gee, 1969). Amongst Haplochromis-gToup species an equivalent hypertrophy is seen only in H. bullatus of Lake Albert, and in species of the endemic Lake Malawi genus Trematocranus (Trewavas, 1935). Species of another Malawi endemic, Aulonocara, show greater development in certain parts of the system (the infraorbital series for example), but otherwise exhibit a level of hypertrophy comparable with that of H. macconneli. Trematocranus and Aulonocara are manifestly more closely related to each other and to other taxa from Lake Malawi (Trewavas, 1935 ; personal observations) than LAKE RUDOLF HAPLOCHROMIS SPECIES 161 to any species occurring outside the lake. Neither need be considered further in the possible phylogeny of H. macconneli. On purely morphological grounds H. bullatus of Lake Albert could be considered the nearest living relative of H. macconneli. Both species share the specialization of enlarged laterosensory canals on the head, and both share (with other species from Lakes Albert and Rudolf) certain peculiarities in the predorsal and thoracic squama- tion patterns (see above, p. 144). The latter character is, however, difficult to assess with respect to its being a primitive or a derived one. At present all that can be said is that the pattern is not encountered amongst the Haplochromis species of Lake Victoria nor is it seen in the fluviatile Haplochromis of Kenya, Uganda and Tanzania. It is rarely encountered in the Haplochromis species flock of Lakes Edward and George (where it is known from two species, H. pharyngalis Poll and H. petronius Greenwood ; see Greenwood, 1973), but it does seem to characterize the Haplochromis of the River Nile, Lake Albert, Lake Rudolf and the River Zaire (personal observations ; also Greenwood, 1971). Haplochromis macconneli differs from H . bullatus in the shape of its neurocranium (see p. 157 above) which is like that found in the moderately specialized insectivore- piscivore radiation in Lake Victoria (see Greenwood, 1974, pp. 80-93). This difference would not, of course, debar H. bullatus from consideration as the living plesiomorph sister species of H. macconneli. But the fact that the Lake Rudolf and Lake Albert basins have never been interconnected (and if there had been some riverine connection, the probability that any presumed common ancestor of the two species would itself have been abyssal in habits) seems to rule out any such close phyletic relationship. Interspecific similarities in laterosensory canal hypertrophy are thus to be interpreted as the product of parallel evolution. The shared peculiari- ties in squamation patterns may well reflect a common ancestry but this is likely to be a relatively distant one (see below, p. 162). It seems probable, therefore, that the relationships of H. macconneli should be sought amongst the species of Lake Rudolf. Two interpretations seem possible. First, H. macconneli may be an immediate derivative of an H. rudolfianus-like ancestor (i.e. H. rudolfianus and H. macconneli may be true sister species). Alter- natively, H. macconneli might be the apomorph survivor of another lineage the relatively plesiomorph, that is H. rudolfianus-like members of which have become extinct (unlike the situation in Lake Victoria, for example, where it is possible to follow, from species still extant, the specialization of a lineage ; see Greenwood, 1974). DISCUSSION Even with the discovery of two and possibly three new Haplochromis species, the total fish fauna of Lake Rudolf still stands low on the scale of endemicity in African lakes. Its Haplochromis species flock also shows a low level of adaptive radiation, probably lower than that of Lake Albert (see Trewavas, 1938 ; Greenwood, 1971) where an anatomically specialized mollusc-eater, a specialized grazer on epiphytes and a species adapted for life in deep water have evolved. 162 P. H. GREENWOOD This comparison must, however, be interpreted with care. For one thing, H. mahagiensis David & Poll (the mollusc-crushing species) of Lake Albert may well belong to a different lineage from that of the other species in the lake. It could be the local representative of a fluviatile mollusc-crushing species represented elsewhere by H. straeleni Poll and H. vanderhorsti Greenwood (see Greenwood, 1954 & iQSQa, for discussion). Furthermore, a temporal element is probably involved. There is little evidence of Lake Albert having dried out at any time in its history, but Lake Rudolf probably was severely reduced, or even completely desiccated, during the middle part of the Pleistocene (see Fryer & lies, 1972, for review). Refilling of the Rudolf basin appears to have been through what is now the River Sobat at some time within the later Pleistocene. Subsequently the connection was broken and has never been re-established. Such an historical background has two consequences, namely that Lake Rudolf is to be considered a relatively young lake, and that its colonizers (or, perhaps more accurately, its recolonizers) were species of Nilotic origin. The relative youth of Lake Rudolf, coupled with the nature of its basin may, as Fryer & lies (1972) suggest, account for the paucity of endemic species and, I would also suggest, for the muted adaptive radiation seen amongst the three or four Haplochromis species that evolved there. With two exceptions, all the Haplochromis species of Lake Albert are apparently confined to the basin of that lake. The two more widely distributed species, H. wingatii (Blgr.) and H. loati Greenwood, both have dental specializations that are not shared with any Lake Rudolf taxa (Greenwood, 1971). Thus, it seems unlikely that an extant Albertine Haplochromis species was the original recolonizer of Lake Rudolf in later Pleistocene times. Nor does it seem probable that the recolonizer closely resembled any species from the Nile (amongst which must be considered H, wingatii and H. loati or their ancestors). In this situation the only conclusion that can be drawn is that some fluviatile species, now extinct, provided the founder population for the Lake Rudolf microflock. It is, of course, possible that the present-day Rudolf species (or some part of them) are descendants from the relicts of a previous flock, possibly a more complex one, that inhabited the early Pleistocene lake and which survived the subsequent period of desiccation. The neurocranial differences between H. rudolftanus and H. maccon- neli might be explained in this way. If this was the history of the present-day species then it follows that their ancestors were derived not from the Nile (which did not then exist in its present form ; see Berry & Whiteman, 1968), but from a river that originated in the eastern highlands and emptied into the developing Nile system in the region of the present River Sobat. Shared peculiarities in the squamation pattern of the thoracic and predorsal regions hint at a common ancestry for the Lake Rudolf and Lake Albert Haplochromis species (see above, p. 161). This character complex is not found in the H. bloyeti-like species group that is widespread in the rivers of Uganda, Kenya and Tanzania. Members of this species complex are thought to be close relatives of the ancestral species which gave rise to the sister species flocks in Lake Victoria and Lakes Edward and George (see Greenwood, 1973 & 1974). Fishes in these flocks, with one exception in Lake George and one in Lake Edward, all lack the Albert-Rudolf scale LAKE RUDOLF HAPLOCHROMIS SPECIES 163 pattern. The exceptional species, H. petronius and H. pharyngalis, resemble the Nilotic H. wingatii in several features as well as the one of scale pattern, and they may represent an exotic element amongst the otherwise H. bloyeti-like. derivatives inhabiting these lakes (Greenwood, 1974). The possible phyletic relationship of Haplochromis species from Lakes Rudolf and Albert raises some interesting points of zoogeography. The lakes are several hundred kilometres apart and any form of past interconnection they may have had would have been of an indirect nature (see above, p. 162). In contrast, Lake Albert is close to Lake Edward and the lakes are in direct contact through the River Semliki. Yet, their faunas, both cichlid and non-cichlid, are quite distinct (Greenwood, 1959^. The present barrier to faunal interchange, principally the Semliki rapids, is clearly an effective one. I have argued elsewhere for a close phyletic relationship between the Haplochromis species of Lakes Edward and Victoria and for their derivation in parallel from a common ancestor that once inhabited the westward flowing rivers of eastern Africa during the Pleistocene (Greenwood, 1973). It seems now that perhaps this concept should be qualified by postulating an ancestral species that lived in some but not all of those rivers. The reasoning behind this qualification is, of course, the presence of species in Lake Albert that would seem to be derived from a different lineage, a lineage that also gave rise to the species of the Nile and, possibly, Lake Rudolf as well (see above, p. 161). Furthermore, it is possible that the Haplochromis species of the River Zaire may share ancestry with these species (p. 161). The evidence upon which these postulated phylogenies are based, a shared scale pattern, is admittedly tenuous, particularly since it is not yet possible to determine which of the two types is to be considered the primitive condition. Further research is planned to investigate the phylogeny of Haplochromis-group species and I would certainly not consider the ideas expressed here as more than a working hypothesis. The presence of two species with Albert-Rudolf scale patterns (H. petronius and H. pharyngalis) in Lakes George and Edward demands explanation. Three pos- sible explanations can be considered. First, the ancestor of these species made its way into the Lake Edward basin in fairly recent times and via the River Semliki. The likelihood of this, however, is reduced by the fact that no reciprocal exchange of Haplochromis seems to have taken place (although the two lakes share another cichlid Sar other odon leucostictus [Trewavas]). Second, the prerift rivers each may have contained species of both squamation types. The absence of species with the Albert-Rudolf pattern from most rivers in eastern Africa (and probably from Lake Victoria as well) would seem unlikely if both types had been represented there previously. The third possibility is one based on the assumption of there having been a single basin in which the proto-Lakes Albert and Edward developed, probably as a series of partly interconnected small and swampy lakes. The northern region of this basin (the future Lake Albert) could have been fed by rivers in which the Haplochromis species had an Albert-Rudolf facies, while the southern end (future Lakes Edward and George) was fed by rivers with Haplochromis of a Victoria-Edward facies. The next assumption would be that only a limited exchange of species took place between the two regions before their continuity was broken. Victoria-type species, if any penetrated to the north, were, presumably, unsuccessful in that 164 P. H. GREENWOOD environment or in competition with the Albert-Rudolf types. Such a general contact between the early lakes may also account for the similarity in their Pleisto- cene fish faunas (see Greenwood, I95gb ; also new and unpublished observations), although one can equally argue that a fauna of this type was widespread in the prerift river systems. Whatever the explanation, it does seem that both Lake Albert and Lake Rudolf have, since their inceptions, been relatively isolated from each other and from other water bodies in eastern Africa. ACKNOWLEDGEMENTS I am particularly grateful to Mr and Mrs A. J. Hopson for, in the first place, sending me the specimens on which this paper is based, and then for the great pains they took to obtain extra information and specimens. To my colleague Mr Gordon Howes goes my gratitude for all the help he has given in the preparation of the paper, for his draughtsmanship displayed in Text-figs. 5, 7 and 8, and for his skill in radio- graphing several hundred specimens. Finally, I thank my colleague Dr Keith Banister whose visit to Lake Rudolf (made since this paper went to press) led to his providing me with valuable extra information on Crater Lake species, and on the scarcity of H. macconneli males ; despite diligent searches he found none of the latter. REFERENCES BEADLE, L. C. 1932. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. 4. The waters of some East African lakes in relation to their fauna and flora. /. Linn. Soc. (Zool.}, 38 : 157-211. BERRY, L. & WHITEMAN, A. J. 1968. The Nile in the Sudan. Geogrl. J. 134 : 1-37. FRYER, G. & ILES, T. D. 1972. The Cichlid Fishes of the Great Lakes of Africa. Their Biology and Evolution. Oliver and Boyd. Edinburgh. GREENWOOD, P. H. 1954. O n two species of cichlid fishes from the Malagarazi River (Tangan- yika), with notes on the pharyngeal apophysis in species of the Haplochromis group. Ann. Mag. nat. Hist. (12), 7 : 401-414. I959a. The monotypic genera of cichlid fishes in Lake Victoria, Part II. Bull. Br. Mus. nat. Hist. (Zool.), 5 : 163-177. !959b. Quaternary fish fossils. Explor. Pare. natn. Albert Miss. J. de Heinzelin de Braucourt, 4 : 1-80. 1965. On the cichlid fishes of Lake Nabugabo, Uganda. Bull. Br. Mus. nat. Hist. (Zool.), 12 : 3I3-357- 1971. On the cichlid fish Haplochromis wingatii (Blgr.) and a new species from the Nile and Lake Albert. Revue Zool. Bot. afr. 84 : 344-365. 1973- A revision of the Haplochromis and related species (Pisces : Cichlidae) from Lake George, Uganda. Bull. Br. Mus. nat. Hist. (Zool.), 25 : 139-242. 1974- The cichlid fishes of Lake Victoria, east Africa : the biology and evolution of a species flock. Bull. Br. Mus. nat. Hist. (Zool.), Suppl. No. 6 : 1-134. & GEE, J. M. 1969. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), Part VII. Bull. Br. Mus. nat. Hist. (Zool.), 18 : 1-65. TREWAVAS, E. 1933. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. ii. The cichlid fishes. /. Linn. Soc. (Zool.}, 38 : 309-341. LAKE RUDOLF HAPLOCHROMIS SPECIES 165 TREWAVAS, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10), 16:65-118. 1938. Lake Albert fishes of the genus Haplochromis. Ann. Mag. nat. Hist, (n), 1 : 435- 449- 1973- On the cichlid fishes of the genus Pelmatochromis with proposal of a new genus for P. congicus ; on the relationship between Pelmatochromis and Tilapia and the recognition of Sarotherodon as a distinct genus. Bull. Br. Mus. nat. Hist. (Zool.), 25 : 1-26. WORTHINGTON, E. B. 1932. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. I. General introduction and station list. /. Linn. Soc. (Zool.), 38 : 99-119. & RICARDO, C. K. 1936. Scientific results of the Cambridge expedition to the East African lakes, 1930-1. 15. The fish of Lake Rudolf and Lake Baringo. /. Linn. Soc. (Zool.), 39 : 353-389. P. H. GREENWOOD, D.Sc. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SWy 5BD A LIST OF 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. I974- 375- Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol B$4 sNU /v A REVIEW OF SCOTOECUS THOMAS, 1901 (CHIROPTERA : VESPERTILIONIDAE) J. E. HILL BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 4 LONDON: 1974 /v OCTI974 A REVIEW OF SCOTOECUS THOMAS, (CHIROPTERA : VESPERTILIONIDAE) BY JOHN EDWARDS HILL Pp. 167-188 ; 4 Text- figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 4 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 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. 27, 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.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 20 September 1974 Price 1.05 A REVIEW OF SCOTOECUS THOMAS, 1901 (CHIROPTERA : VESPERTILIONIDAE) By JOHN E. HILL INTRODUCTION CURRENT classification of the genus Scotoecus rests basically on Ellerman, Morrison- Scott & Hayman (1953 : 83) who recognized two species in southern Africa, one (albofuscus) with pale wings, the other (hirundo) with dark wings. Hayman (1963 : 104) expanded this opinion in a brief review, and allocated the named forms then in the genus to one or other of these as subspecies. Rosevear (1965 : 300) retained faldbae from northern Nigeria as a distinct species, but Hayman & Hill (1971 : 36) recognized only albofuscus and hirundo as valid species. However, specimens examined recently at the British Museum (Natural History) have suggested that the dark-winged group is composed of two species, one smaller, for which hirundo is available, the other a little larger in size, for which the earliest name is hindei. Hitherto, Scotoecus has been considered exclusively African, but a study of the Indian species first described as Scotophilus pallidus by Dobson (1876 : 186) and nowadays generally referred to Nycticeius shows it to be more closely allied to Scotoecus, to which it is transferred. Measurements throughout are in millimetres : minimum and maximum values are given for series, with (in parentheses) the number of specimens measured. SYSTEMATIC SECTION SCOTOECUS Thomas, 1901 Scotoecus Thomas, 1901 : 263. Type species Scotophilus albofuscus Thomas, 1890. DIAGNOSIS. Similar to Scotozous and Nycticeius but with spatulate or nearly spatulate tragus (Fig. i) ; braincase elevated frontally to give a slightly convex frontal profile ; rostrum greatly widened at the lachrymals ; narial and pre- palatal emarginations exceptionally deep ; zygomata tenuous, usually lost in pre- paration ; and anterior face of the upper canine flattened and grooved. DESCRIPTION. Small (length of forearm 28-38) ; muzzle broad, flattened, anteriorly sparsely haired, nares circular, opening obliquely ; ear rounded with rounded tip, its anterior margin lacking any basal lobe ; tragus spatulate or nearly so with small, more or less triangular posterior basal lobe ; calcar extending along two thirds of the uropatagial margin with small post-calcarial lobe ; penis sometimes exceptionally long. Skull low, braincase broad, with slight occipital helmet and low lambdoid and sagittal crests ; skull slightly elevated frontally to produce a faintly convex profile ; interorbital region wide, rostrum broad, much expanded at lachrymals ; anteorbital foramen moderate or large, closed by strong, nearly vertical bar ; small supra- orbital tubercles often present ; narial emargination wide, U-shaped or slightly i 7 o J. E. HILL V-shaped, the apex rounded, deep, extending posteriorly at least to a line joining the anterior margins of the anteorbital foramina, sometimes to a line joining the anterior margins of the orbits ; pre-palatal emargination wide, extending posteriorly to a point lying between a line joining the anterior faces of the large upper premolars (pm 4 - 4 ) and a line joining the anterior faces of the first upper molars (m 1 - 1 ) ; zygo- mata tenuous, in African species often lost in preparation ; maxillary toothrows not markedly convergent anteriorly ; a narrow post-palatal spine and shallow basioccipital depressions. 1123 1113 Dental formula i-, c-, pm -, m - = 32 or i -, c-, pm -, m - = 30. Massive, 3i 23 3123 unicuspid inner upper incisor (i 2 ), in contact with the canine or nearly so ; anterior face of upper canine flattened, grooved, sometimes prominently so ; anterior upper premolar (pm 2 ) minute or absent, when present compressed into a lingual recess between canine and the second upper premolar (pm 4 ) which are otherwise in contact ; pm 4 with antero-medial cusp or protocone low or absent ; upper molars (m 1 ' 3 ) with protocones prominently developed ; the lingual shelves widely separated, m 3 not reduced, its crown area exceeding one half the crown area of m 1 or m 2 , with promi- nent metacone, metastyle and three commissures ; lower incisors (ij.g) imbricated ; anterior lower premolar in the Indian species reduced to one third to one half the crown area of the second lower premolar (pm 4 ) which is not compressed in the toothrow ; in the African species pm 2 a little less in crown area than pm 4 , the latter slightly shortened antero-posteriorly, compressed between pm 2 and the first lower molar (m^ ; third lower molar (m 3 ) not reduced or only very slightly reduced, its posterior triangle not at all or only a little smaller than the anterior triangle, hypo- conid and entoconid present, the latter slightly the lesser. LITERATURE. Trouessart, 1904 : 85 (spelt Scotaecus) ; Miller, 1907 : 217 (generic diagnosis) ; Wettstein, 1918 : 654 (review, key) ; Simpson, 1945 : 59 (incorporated into Nycticeius) ; Ellerman, Morrison-Scott & Hayman, 1953 : 83 (status, partial key, a subgenus of Nycticeius) ; Hayman, 1963 : 104 (review, tooth formula, variability of pm 2 , a subgenus of Nycticeius) ; Koopman, 1965 : 17 (status, generically valid) ; Rosevear, 1965 : 289, 297 (status, description, partial key, generically valid) ; Ansell 1967 : 21 (as a genus) ; Hayman & Hill, 1971 : 36 (key, notes, distribution, subgenus of Nycticeius) ; Koopman, in litt., in Hayman & Hill, 1971 : 36 (a subgenus of Nycticeius}. DISTRIBUTION AND SPECIES. Indian and African, in India a semi-desert species inhabiting tropical thorn forest, in Africa occurring chiefly in open woodland in three species with few records from the high forest. The Indian species (first de- scribed as Scotophilus pattidus by Dobson, 1876 : 186) was for many years referred to Scoteinus (see below) but is here allocated to Scotoecus on account of its aural and rostral features, together with its grooved upper canines. Its features are less extreme than are those of the African species of Scotoecus and apparently it forms a link between these and Nycticeius, with which in recent years it has been associated The African members of the genus may be divided into two sections, the one pale- winged, with brownish ventral surface, the other dark-winged, the ventral surface REVIEW OF SCOTOECUS THOMAS, 1901 171 pale, usually some shade of greyish. These correspond to the two species currently recognized in Africa by most modern authors. The examination of Ethiopian speci- mens and of specimens from Cameroon and Uganda in the course of preparing this study has suggested the further division of the dark-winged section into two species, one smaller, the other larger ; however, the genus is still known from relatively few specimens and the classification presented here is to that extent provisional. The species may be keyed : 1 Tragus long, relatively narrow ; zygomata not cartilaginous ; pm 2 reduced, its crown area one third to one half that of pm 4 ; this tooth not compressed in toothrow ...... pallidus (p. 172) Tragus short, relatively wide ; zygomata cartilaginous ; pm 2 unreduced, its crown area subequal to that of pm 4 ; this tooth compressed in toothrow ........... 2 2 Tragus more or less parallel-margined, the upper part of its anterior margin not prolonged anteriorly ; outer part of wing membrane pale ; ventral surface of body brown ; braincase slightly flattened albofuscus (p. 174) Tragus slightly hatchet-shaped, the upper part of its anterior margin projected forward ; outer part of wing membrane dark ; ventral surface of body greyish, at most tinged with brown ; braincase slightly elevated ........... 3 3 Smaller, length of forearm generally less than 33-0, condylobasal length 13-2 or less, c-m 3 5-1 or less . . . . . . hirundo (p. 177) - Larger, length of forearm generally exceeding 32-0, condylobasal length 13-5 or more, c-m 3 5-2 or more ..... hindei (p. 179) HISTORY. First separated from Scotophilus at the generic level by Thomas (1901 : 263), Scotoecus was considered generically valid by Miller (1907 : 217), who allied it with Nycticeius and Scoteinus. Allen (1939 : 98) listed Scotoecus as a distinct genus for Africa but since that date opinions of its status have diverged. Simpson (1945 : 59) incorporated it into Nycticeius, a lead followed by Ellerman, Morrison- Scott & Hayman (1963 : 83) who treated Scotoecus as a subgenus of Nycticeius with the comment that the characters given by Thomas and Miller seemed more subgeneric than generic. Hayman (1963 : 104) and Hayman & Hill (1971 : 36) adopted this view but Rosevear (1965 : 289) noted that Scotoecus is readily distinguished from Nycticeius and considered that it merited generic separation, a view followed by Ansell (1967 : 21) and shared by Koopman (1965 : 17) who did not consider Scotoecus congeneric with Nycticeius but later (in litt., in Hayman & Hill, 1971 : 36) revised this opinion to accord Scotoecus subgeneric rank only. REMARKS. There seem no good grounds for considering Scotoecus congeneric with Nycticeius : the form of the tragus, the elevated cranium and widened rostrum with deep anterior emarginations, and the grooved upper canines of Scotoecus separate it sharply from this genus. The convexity of the frontal area in Scotoecus, its broad rostrum with deep emarginations and its weak zygomata present a parallel with Nyctalus, in which, however, no modification of the upper incisors has occurred. 172 J. E. HILL Scotoecus pallidus (Dobson, 1876) ? Vespertilio noctulinus Geoffrey, 1831 : 92, pi. 3. Bengal, India. Scotophilus pallidus Dobson, 1876 : 186, Appendix D, 208. Mian Mir, Punjab, northwestern India. DIAGNOSIS. Differing from the African S. albofuscus, S. hirundo and S. hindei in thicker, more fleshy ears, more swollen at the antitragus ; tragus (Fig. i) longer and relatively narrower, its tip more expanded ; narial emargination faintly V-shaped, its lateral margins slightly convergent posteriorly, the apex rounded ; pre-palatal emargination narrower ; anteorbital foramen smaller ; zygomata tenuous but not cartilaginous ; anterior lower premolar (pm 2 ) reduced, its crown area one third to one half that of the second lower premolar (pm 4 ) which is not compressed in the toothrow ; third lower molar (m 3 ) slightly reduced, the posterior triangle a little narrower than the anterior triangle. Differs further from S. hirundo and S. hindei in less inflated and less elevated braincase, in this respect resembling 5. albofuscus. DESCRIPTION. Large for the genus (length of forearm 34-38) ; lips fleshy ; ear moderate, reaching about halfway to tip of muzzle, with fleshy membrane and anti- tragal thickening ; anterior margin of tragus straight, tip rounded, anteriorly directed, posterior margin convex, penis not exceptionally developed. Dorsal surface pale brown, tinged with fawn, ventral surface paler, greyish white. Skull relatively massive, with (especially in older individuals) low occipital helmet and posterior sagittal and lambdoid crests ; slight supraorbital ridges with incipient supraorbital tubercles ; anterior margin of orbit nearly vertical, the supraorbital and lachrymal regions swollen ; anteorbital foramen moderate, closed by narrow bar ; narial emargination wide, faintly V-shaped, slightly narrowed posteriorly, its lateral margins slightly convergent, extending posteriorly a little more than halfway to the anterior orbital margin ; pre-palatal emargination wide, U-shaped, extending pos- teriorly to a line joining the anterior faces of the second upper premolars (pm 4 " 4 ) ; zygomata narrow, tenuous but not cartilaginous ; post-palatal extension narrow, with small, delicate post-palatal spine. ii 13 Dental formula i -, c -, pm -, m - = 30. Upper canine with flattened, faintly grooved anterior face ; small upper premolar (pm 2 ) absent ; large upper premolar (pm 4 ) compressed between the canine and the first upper molar (m 1 ), with small antero-medial cusp or protocone ; m 1 ' 3 with strong protocones, their lingual shelves widely separated ; anterior lower premolar (pm 2 ) much reduced, one third to one half the crown area of the second lower premolar (pm 4 ) and less than one half its height, compressed between the canine and pm 4 ; third lower molar (m 3 ) very slightly reduced, posterior triangle a little narrower than the anterior triangle, hypoconid and entoconid slightly smaller than protoconid, paraconid and metaconid. MEASUREMENTS. Length of forearm (14) 34-1-38-4 ; condylobasal length (8) I4'i-i4'9 ; condylocanine length (8) 13-9-14-7 ; least interorbital width (10) 4-0-4-4 ; zygomatic width (i) 10-5 ; width of braincase (9) 7-3-8-1 ; mastoid width (7) 8-9-9-7 ; ^-c 1 (9) 4-9-5-5 ; m 3 -m 3 (9) 6-5-6-9 ; c-m 3 (u) 5-2-5-7 ; c-m 3 (n) 5-8-6-2. REVIEW OF SCOTOECUS THOMAS, 1901 173 LITERATURE. Dobson, 1876 : 186 (description), 208 (holotype listed) ; 1878 : 257, 258 (in Scoteinus, a subgenus of Scotophilus), 264 (description repeated), pi. xv, fig. 3 (ear) ; Blanford, 1891 : 222 (further description) ; Trouessart, 1897 : 119, 1904 : 85 (in Scoteinus, a subgenus of Scotophilus) ; Siddiqi, 1961 : 125 (Pakistan localities reviewed) ; Sinha & Chakraborty, 1971 : 53 (cranial features, measure- ments), figs, i A, 2 A, 3 A (skull). DISTRIBUTION. WEST PAKISTAN (Sind : Dobson, 1877 : 310 ; Anderson, 1881 : 137 ; Wroughton, 1916 : 752 ; Siddiqi, 1961 : 125 ; Sinha & Chakraborty, 1971 : 54). INDIA (Punjab : Dobson, 1876 : 186, 208, 1877 : 310, 1878 : 264 ; Anderson, 1881 : 137 ; Allen, 1908 : 48. Uttar Pradesh ; Bihar : Khajuria, 1951 : 120 ; Sinha & Chakraborty, 1971 : 54 ; specimens collected at Bahgownie, Darbhanga District, Bihar by N. A. Baptista in the collections of the British Museum (Natural History)). The record from Massowa, Eritrea, Ethiopia by Thomas & Doria (1886 : 206) in fact refers to Nycticeius schlieffenii, these authors having been misled by a com- parison with a specimen of schlieffenii with an erroneous locality, identified by Dobson as pallidus (Thomas, 1890 : 86). HISTORY. A rarely collected species to which there are relatively few references. Described as a Scotophilus, the species was placed by Dobson (1878 : 257, 258) and by Trouessart (1897 : 119, 1904 : 85) in the subgenus Scoteinus, subsequently elevated to generic rank by Miller (1907 : 217) but diagnosed by this author without reference either to its type species (designated by Miller, p. 217) Scotophilus emarginatus Dobson, 1871 or to Scotophilus pallidus Dobson, 1876. The type species has been shown by Sinha & Chakraborty (1971 : 53) to be congeneric with the species known for many years as Scotomanes ornatus (Blyth, 1851)* and pallidus does not in any event conform to the diagnosis of Scoteinus by Miller : the lachrymal region is widened as in the African species of Scotoecus and the third molars (m f) are almost un- reduced, the narrowing of the rostrum and reduction of m f being the chief diag- nostic features of Scoteinus as understood by Miller, who had seen only the Australian species balstoni and greyii and the African schlieffenii, all nowadays usually referred to Nycticeius. REMARKS. As might be expected from the disjunct distribution, the Indian 5. pallidus differs more widely from its African congeners than these do from each other. In particular, a number of its features are less extreme than in the African species and its dentition approaches more nearly to the type exemplified by Nycticeius. There seem no good grounds for referring pallidus to this genus, however, since in Nycticeius the rostrum is not broadened and the upper canine is rounded anteriorly, not at all flattened or grooved : the same is true of the Australian species referred hitherto to Scoteinus and more recently to Nycticeius. In these, additionally, a greater measure of reduction of the third molars (m f ) has occurred. * The generic synonymy was not discussed by Sinha and Chakraborty. Scoteinus and Scotomanes were proposed simultaneously by Dobson (1875: 371) Scoteinus having line priority. Article 24 of the International Code of Zoological Nomenclature provides that in such a case the relative priority of the names is determined by the action of the first reviser. Sinha and Chakraborty may be considered to be first revisers and continue to use Scotomanes. In view of the varied uses of Scoteinus this action is in accord with Recommendation 2^A of the International Code, which suggests that the name that ensures stability and universality of nomenclature be selected. 174 J- E - HILL Vespertilio noctulinus I. Geoffrey, 1831. Tate (1942 : 282) and Ellerman & Morrison-Scott (1951 : 177) suggested that this name should be associated with pallidus rather than considered a synonym of Scotophilus temminckii ( = S. kuhlii) as it had been hitherto. It has a long history in the early literature : the description is repeated by Temminck (1840 : 211), who, in a supplement to the same work, subsequently provided (p. 266) a further description with details of the dentition, referring specimens from Singapore in the collections of the Rijksmuseum van Natuurlijke Historic, Leiden, to noctulinus. Cantor (1846 : 185), in a catalogue of the mammals of the Malay Peninsula referred the name to the synonymy of Scoto- philus temminckii (= 5. kuhlii) with the comment that Nycticeius noctulinus, Temminck referred to the young of the species, a view supported by Jentink (1888 : 183) who listed a young example of 'Vespertilio noctulinus Is. Geoffroy' as Scotophilus temminckii. Wagner (1840 : 543, 1855 : 765) also provided further descriptions : Fitzinger (1870 : 367) gave a detailed synonymy to that date, with another descrip- tion (as Nycticejus noctulinus) but Dobson (1876 : 120, 1878 : 258) referred the name to the synonymy of Scotophilus temminckii. Here it has since remained (Blanford, 1891 : 320, although this author evidently had some reservations, placing a query after the allocation ; Trouessart, 1897 : 118) until recent years. It is of interest to note that Oldfield Thomas has made a marginal note in his copy of Dobson's Cata- logue (1878) which reads 'Size of a pipistrelle! F.a. 36 (Not a Scotophilus} !' -the grounds on which Tate suggested association with pallidus. Scotoecus albofuscus (Thomas, 1890) DIAGNOSIS. Similar to S. pallidus but smaller ; ears not especially fleshy ; tragus (Fig. i) smaller, spatulate, not prolonged anteriorly ; distal part of wing whitish ; supraorbital region widely expanded ; lateral margins of narial emargina- tion not convergent posteriorly ; anteorbital foramen large ; zygomata cartila- ginous ; anterior lower premolar (pm 2 ) unreduced and third lower molar (m 3 ) barely reduced. DESCRIPTION. Small (length of forearm 28-5-31-0) ; ear membrane and anti- tragus relatively thin ; tragus small, spatulate, upper part of its anterior margin not prolonged anteriorly. Dorsal surface brown, ventral surface similar but slightly paler ; wing blackish brown from body to a line joining elbow to knee but whitish from forearm, the forearm and digits dark ; interfemoral membrane dark brown. Skull with broad, low, flattened braincase ; supraorbital region widely expanded ; narial emargination U-shaped, its lateral margins not convergent posteriorly ; anteorbital foramen large, closed by a strong, nearly vertical bar ; zygomata car- tilaginous, often lost in preparation. ii 13 Dental formula i -, c -, pm -, m - = 30. Upper canine with flattened, strongly 312 3 grooved anterior face ; small upper premolar (pm 2 ) absent in all eight specimens examined and in one (No. 22412, Musee Royale d'Afrique Centrale, Tervuren) seen by Dr F. de Vree (in litt.) ; large upper premolar (pm 4 ) with antero-medial cusp or REVIEW OF SCOTOECUS THOMAS, 1901 175 a i L mm i i i. j FIG. i. Tragus of (a) Scotoecus pallidus. (b) Scotoecus albofuscus. (c) Scotoecus hirundo. (d) Scotoecus hindei. protocone low or absent ; upper molars (m 1 ' 3 ) with strongly developed protocones ; anterior lower premolar (pm 2 ) unreduced, its crown area almost equal to that of the second lower premolar (pm 4 ) which is compressed in the toothrow and is slightly flattened antero-posteriorly ; third lower molar (m s ) barely reduced, the width of its posterior triangle nearly equal to that of the anterior triangle. DISTRIBUTION AND SUBSPECIES. Known from a limited number of specimens from localities in West Africa, Zaire, Tanzania and Malawi. Following Hayman (1963 : 105), Hayman & Hill (1971 : 37) recognized three subspecies, 5. a. albofuscus in West Africa, 5. a. cinnamomeus Wettstein, 1916 in the Sudan and S. a. woodi Thomas, 1917 in southern and eastern Zaire, Tanzania and Malawi. However, cinnamomeus is based upon an example of Nycticeius schlieffenii*. Scotoecus albofuscus albofuscus (Thomas, 1890) Scotophilus albofuscus Thomas, 1890 : 84. Bathurst, Gambia. DIAGNOSIS. Upper canine with rounded base, its antero-internal margin not sharply angular, postero-internal margin rounded, lingual shelf narrow. MEASUREMENTS. Length of forearm (4) 29-8-31-0 ; condylobasal length (3) 12-7-13-5 ; condylocanine length (3) 12-8-13-3 ; least interorbital width (3) 4-1-4-4 ; zygomatic width ; width of braincase (4) 7-3-7-7 ; mastoid width (3) 8-2-9-0 ; ci-c 1 (3) 4-4-5-1 ; m 3 -m 3 (3) 6-6-6-7 ; c-m 3 (4) 4'7~5' '> c-m 3 (2) 5-1-5-3. * The status of cinnamomeus Wettstein, 1916 from Kordofan has been reviewed in some detail by Kock (1969: 188). The preliminary description by Wettstein (1916: 191) was succeeded by a more detailed study (1918: 653) with illustrations (p. 653, fig. 12) of the ear and tragus and more particularly (pi. 4, n g s - 5' 5 a ~b) of the skull and dentition. As Kock points out, these ally cinnamomeus undoubtedly with Nycticeius schlieffenii. This opinion is supported by Koopman (in litt., in Hayman & Hill, 1971: 37) who has examined the holotype in Vienna. 176 J. E. HILL FIG. 2. Distribution of Scotoecus albofuscus. LITERATURE. Rosevear, 1965 : 298, 299 (description), fig. 78 (skull), 305, fig. 8oa (tragus). DISTRIBUTION. SENEGAL (Badi : Thomas, 1911 : 119, as a location in Gambia) ; GAMBIA ; SIERRA LEONE (both Rosevear, 1965 : 299, from specimens in British Museum (Natural History)); IVORY COAST (Ayeremou (= Ahieremou, 6i4' N., 456' W. ?), specimens in the Museum d'Histoire Naturelle, Geneva (F. de Vree, in litt.)) ; NIGERIA (Dollman, 1908 : 546). Scotoecus albofuscus woodi Thomas, 1917 Scotoecus woodi Thomas, 1917 : 280. Chiromo, southern Malawi, 200 feet. DIAGNOSIS. Upper canine with angular base, its antero-lateral margin forming a right angle, postero-internal margin angular and projecting, the lingual shelf wide. MEASUREMENTS. Length of forearm (5) 28-5-30-5 ; condylobasal length (4) 12-9-13-7 ; condylocanine length (4) 12-9-13-7 ; least interorbital width (4) 4-1-4-3 ; zygomatic width ; width of braincase (4) 7-2-7-6 ; mastoid width (3) 8-8-9-0 ; REVIEW OF SCOTOECUS THOMAS, 1901 177 c 1 -^ (4) 4-8-5-1 ; m 3 -m 3 (4) 6-3-6-8 ; c-m 3 (4) 4-8-5-1 ; c-m 3 (4) 5-1-5-5 (speci- mens all from Chiromo, Malawi). Dr F. de Vree (in litt.) has measured a specimen (No. 22412) in the Musee Royale d'Afrique Centrale, Tervuren, from Baudouinville (07O3' S., 2g42' E.), Zaire : length of forearm 29-4 ; condylobasal length 12-9 ; least interorbital width 4-6 ; zygomatic width ; width of braincase 7-3 ; mastoid width c. 8-4 ; m 3 -m 3 6-7 ; c-m 3 5-2 ; c-m 3 (alveoli) 4-8 ; c-m 3 (alveoli) 5-5. LITERATURE. Harrison, 1958 : 98, fig. i (whole animal), fig. 2 (head). DISTRIBUTION. ZAIRE (Hayman, 1957 : 44 ; Hayman, Misonne & Verheyen, 1966 : 56) ; TANZANIA (Harrison, 1958 : 96 ; Vesey-Fitzgerald, 1964 : 64) ; MALAWI ; probably in ZAMBIA (Ansell, 1960, Appendix A, p. no, Appendix B, p. 117). REMARKS. According to Thomas (1917 : 280), woodi of southern Africa can be distinguished from the West African albofuscus by its smaller size, proportionately broader skull and by the different shape of the base of the upper canine. The small series of albofuscus and woodi available in the collections of the British Museum (Natural History) do not support the statement that woodi is significantly smaller than albofuscus and it appears to differ in size from albofuscus only in a generally very slightly wider rostrum, equalled in width by one of albofuscus. Scotoecus hirundo (de Winton, 1899) Scotophilus hirundo de Winton, 1899 : 355. Gambaja, Ghana, 1300 feet. DIAGNOSIS. Similar in most respects to 5. albofuscus but anterior margin of tragus (Fig. i) projected anteriorly in its distal part to give the tragus a slightly hatchet-shaped outline, its upper margin nearly horizontal, the junction of the upper and posterior margins angular, slightly obtuse ; wing membranes wholly dark ; ventral surface of body some shade of grey ; braincase slightly inflated and rounded dorsally, less flattened ; pm 2 usually but not always present, the dental formula .11 23 .11 13 i -, c -, pm -, m - = 32 or i -, c -, pm -, m - = 30. 3123 ii 23 VARIABILITY OF PM 2 . The presence of a minute anterior upper premolar (pm 2 ) sandwiched tightly between the canine and pm 4 is variable in 5. hirundo, pm 2 being totally absent from three of the nineteen specimens examined. The tooth is present in both sides of the jaw in a specimen (in the Museum National d'Histoire Naturelle, Paris) from Saboya, Senegal. Dr F. de Vree (in litt.) has examined further specimens (in the Museum d'Histoire Naturelle, Geneva) from the Bandia Forest, Senegal : two of these possessed the small pm 2 but so far as could be seen, this was lacking from a third. It is present in both sides of the jaw in one from Sierra Leone but totally absent from another (Hayman, 1963 : 105). There is no trace of pm 2 in the holotype from Ghana : it is present on both sides of the jaw in one specimen from the Cameroon but in another present in the left hand side only. Of ten examples from Ethiopia, eight have the tooth on both sides of the jaw, in one it is present in the right hand side only and in one it is totally absent. Three specimens from Uganda have pm 2 present on both sides of the jaw. 178 J. E. HILL MEASUREMENTS. Length of forearm (18) 29-7-32-8 ; greatest length of skull (15) 12-6-13-7 ; condylobasal length (16) 12-1-13-0 ; condylocanine length (16) 12-1-13-1 ; width of rostrum (18) 5-9-6-5 ; width across anteorbital foramina (18) 4-6-5-2 ; least interorbital width (18) 4-2-4-6 ; zygomatic width (i) 10-3 ; width of braincase (16) 7-0-7-7 ; mastoid width (13) 8-0-8-8 ; c 1 -c 1 (18) 4-3-5-0 ; m 3 -m 3 (18) 5-9-6-7 ; c-m 3 (18) 4-7-5-1 ; length of complete mandible (15) 8-9-9-9 > c-m 3 (18) 4-9-5-4. Dr F. de Vree (in litt.) has measured three specimens (see above) (one skull only prepared) from the Bandia Forest, Senegal : length of forearm 31-0-32-3 ; greatest length of skull 13-9 ; condylobasal length 12-9 ; least interorbital width 4-7 ; width of braincase 7-8 ; mastoid width 8-9 ; m 3 -m 3 6-5 ; c-m 3 5-3 ; c-m 3 (alveoli) 4-9 ; length of mandible 10-1, c-m 3 (alveoli) 5-3. LITERATURE. Rosevear, 1965 : 300 (further description), 305, fig. 8oc (tragus). DISTRIBUTION. SENEGAL (Saboya, specimen in Museum National d'Histoire Naturelle, Paris ; Bandia Forest, specimens in the Museum d'Histoire Naturelle, Geneva (F. de Vree, in litt.)) ; Sierra Leone (Hayman, 1963 : 105) ; GHANA ; CAMEROON (Yagoua, specimens obtained by F. de Vree and W. Verheyen) ; ETHIO- PIA (Gambela, 515 and 1768 metres, 8i5' N., 3435' E., and Didessa River, Wollega Province, 1190 metres, 9O2' N., 36O9' E., specimens in British Museum (Natural History)); UGANDA (Budongo, i39' N., 3i35' E., specimens in British Museum (Natural History)). REMARKS. In recent years (following Hayman, 1963 : 104) the named forms in the dark-winged section of Scotoecus have been considered subspecies of hirundo, the earliest name. However, measurements of specimens in the collections of the British Museum (Natural History) and of specimens examined in 1971 for Dr F. de Vree, Antwerp, suggest that two species are involved, basically one of smaller size and northern distribution for which the prior name is hirundo, and a second species of larger size and predominantly more southern distribution for which the earliest name is hindei. A specimen (B.M. 72.4421) from the Didessa River, Wollega Province, Ethiopia, is in close agreement with hirundo from West Africa : another (B.M. 72.4423) from the same locality is clearly referable to hindei. These are supported by a series of specimens (B.M. 70.2263-2270) and a further example (B.M. 72.4420), all from Gambela, western Ethiopia, which agree with hirundo : others (B.M. 70.2262, 72.4422) from Bulcha, Lake Margherita, some 200 miles to the southeast of Gambela, are referable to hindei. Scotoecus has apparently not been collected hitherto in Ethiopia. The two also occur almost sympatrically in Uganda : specimens (B.M. 74.1-5) (including pregnant or lactating females) obtained at Budongo (i39' N., 3i35' E.) by J. F. Kingdon and clearly referable to hirundo contrast sharply in size with others (B.M. 63.1151, 65.3435) referable to hindei from West Madi (33o' N., 3i35' E.) and Nabumali, South Bugisu (o59' N., 34i2' E.). Furthermore, two specimens obtained at Yagoua, northern Cameroon, by F. de Vree and W. Verheyen agree very closely with hirundo and were collected concurrently at that locality with two further larger specimens referable with little doubt to falabae, here considered to be a subspecies of hindei. No Scotoecus has been reported hitherto from the REVIEW OF SCOTOECUS THOMAS, 1901 179 Cameroon. There is evidence to suggest, therefore, that in Ethiopia a small species (hirundo) occurs with or in close proximity to a larger species (hindei) : both occur in nearby localities in Uganda and are again sympatric in the Cameroon. Measure- ments of hirundo and of hindei from the areas of sympatry or near sympatry are contrasted in Table i. FIG. 3. Distribution of Scotoecus hirundo. Scotoecus hindei Thomas, 1901 DIAGNOSIS. Similar to S. hirundo but larger, especially cranially, with the supra- orbital and anteorbital regions wider and more greatly expanded. VARIABILITY OF PM 2 . The minute anterior upper premolar (pm 2 ) is generally present in 5. hindei : its presence in twenty-six examples in the collection of the British Museum (Natural History) is variable only in those from Ethiopia and Kenya. The tooth is present on both sides of the jaw in two specimens from Ethiopia, but in a third example present only in the left-hand side of the jaw. Among Kenyan specimens, the tooth is present on both sides in eight, present in the right-hand side but absent from the left in another, and totally absent from two further examples, one the holotype of S. hindei hindei. Hollister (1918 : 94) notes that pm 2 is com- pletely absent from one of five Kenyan specimens in the Museum of Natural History, i8o J. E. HILL FIG. 4. Distribution of Scotoecus hindei. Records referred to 5. h. albigula are indicated by unshaded circles O- the Smithsonian Institution, Washington, but that an alveolus can be seen on one side. The tooth is present in a further specimen reported (as albigula) by this author from Kiriba, Sudan (see below). Furthermore, Dr F. de Vree notes (in litt.) that pm 2 is absent from a specimen (M.R.A.C. 22192) in the Musee Royale d'Afrique Centrale, Tervuren, from Kapalowe, Katanga Zaire. Allen (1911 : 330) describes pm 2 in detail. DISTRIBUTION. Nigeria, Cameroon and Sudan to Ethiopia and Somalia ; Kenya, Uganda and Tanzania to Zambia ; Zaire ; Angola. Scotoecus hindei hindei Thomas, 1901 Scotoecus hindei Thomas, 1901 : 264. Kitui, Kenya, 1150 metres (in original description as 3500 feet). (?) Scotoecus artinii de Beaux, 1923 : 98. Archer's Post, northern Guaso Nyiro, Kenya. MEASUREMENTS. Length of forearm (15) 32-0-37-2 ; greatest length of skull (14) 13-8-15-0 : condylobasal length (15) 13-4-14-4 ; condylocanine length (15) 13-3- 14-4 ; width of rostrum (14) 6-6-7-8 ; width across anteorbital foramina (14) REVIEW OF SCOTOECUS THOMAS, 1901 181 5-1-6-0 ; least interorbital width (15) 4-4-4-8 ; zygomatic width (i) 10-7 ; width of braincase (15) 7-3-8-0 ; mastoid width (15) 8-8-9-7 '< ^-c 1 (14) 5-0-5-6 ; m 3 -m 3 (14) 6-6-7-4 ; c-m 3 (16) 5-3-5-7 ; length of complete mandible (13) 10-0-11-2 ; c-m 3 (16) 5-5-6-4. Dr F. de Vree (in litt.) has measured a specimen (see above) from Kapalowe, Katanga, Zaire : length of forearm 33-2 ; greatest length of skull 14-7 ; condylobasal length 13-8 ; least interorbital width 4-5 ; zygomatic width 10-5 ; width of braincase 7-6 ; mastoid width 9-1 ; m 3 -m 3 7-0 ; c-m 3 5-4 ; c-m 3 (alveoli) 4-9 ; length of mandible 10-1 ; c-m 3 (alveoli) 5-3. DISTRIBUTION. SUDAN (G. M. Allen, 1914 : 349, as Scoteinus schlieffenii, identified as Scotoecus by Koopman, 1965 : 16 ; Hollister, 1918 : 94, from Kiriba, 'Uganda', a locality in southern Sudan according to Kock, 1969 : 192, as albigula ; Kock, 1969 : 192) ; ETHIOPIA (from Bulcha, Lake Margherita, 1800 metres, 6n' N., 38io' E., and from Didessa River, Wollega Province, 1190 metres, 902' N., 3609' E., speci- mens in British Museum (Natural History)) ; SOMALIA (de Beaux, 1924 : 155, as artinii ; Funaioli, 1971 : 29, 64, as hirundo) ; ZAIRE (J. A. Allen, 1916 : 447, as Scoteinus schlieffenii, identified as Scotoecus hirundo hindei by Koopman, 1965 : 16 ; Hayman, 1954 : 291 ; Leleup, 1956 : 77 ; Anciaux de Faveaux, 1958 : 271 ; Hay- man, Misonne & Verheyen, 1966 : 56) ; UGANDA (Hayman & Hill, 1971 ; 36, as albigula) ; KENYA (G. M. Allen, 1911 : 330 ; Dollman, 1914 : 309, in part Nos. 148, 149, 150, as Scoteinus schlieffenii, re-identified in the present study ; Hollister, 1918 : 94 ; Harrison, 1961 : 293) ; TANZANIA (a manuscript note by R. W. Hayman in a British Museum (Natural History) copy of Hayman & Swynnerton, 1951 (p. 294), records a specimen from Lake Mangona, but this example not seen ; Dodoma, specimen in British Museum (Natural History)). Scotoecus artinii de Beaux, 1923. The precise status of artinii is far from clear. Koopman (1965 : 17) is inclined to regard artinii, hindei and a third more southerly form, albigula, as synonymous and notes that a specimen reported from Niangara, eastern Zaire, by J. A. Allen (1916 : 447) as Scoteinus schlieffenii is in fact a Scotoecus, referring it to S. hirundo hindei (= S. hindei hindei} with the remark that its best agreement seems to be with artinii. Hayman & Hill (1971 : 36) listed artinii as a subspecies of hirundo (S. hindei as here understood), considering (p. 37) on the basis of specimens from the Cherangani Hills, western Kenya, that possibly it might be a small form distinct from both hindei and albigula. Three specimens attributed to artinii by R. W. Hayman were collected in 1966 by Mr A. N. Start at the Wei Wei River, Sigor, northeastern Cherangani Hills, Kenya, at about 3000 feet, the same collector subsequently obtaining in 1967 further examples (now B.M. 68.481-482) at the same locality. Mr Start also obtained three specimens attributed to albigula by R. W. Hayman from the Cherangani Hills at 6000 feet in 1965. Of these eight it has been possible to examine only B.M. 68.481-482, but the collections of the British Museum (Natural History) include three further examples (B.M. 14.7.31.15-17) of 5. hindei from the Wei Wei River, formerly identified as Scoteinus schlieffenii (Nos. 148, 149, 150 of Dollman, 1914 : 309). Male specimens from the Wei Wei River agree quite closely with the male holotype of 5. hindei hindei and with male specimens referred to this subspecies from the 182 J. E. HILL Lorian Swamp and the Guaso Nyiro, differing only in their slightly more slender canines. In this respect they resemble the male examples B.M. 63.1151 from West Madi, Uganda, and B.M. 65.3435 from Nabumali, South Bugisu, Uganda. How- ever, a single male specimen collected 30 miles northwest of Baringo, Kenya, bridges this difference. Specimens from the Wei Wei River and from Uganda are generally a little smaller than specimens from more easterly locations and are similar in size to the specimen from Zaire measured by J. A. Allen (1916 : 447) (as Scoteinus schlieffenii}. On the whole they are a little larger than specimens of S. hindei from the northern Cameroon. De Beaux gives few diagnostic characters for artinii but his measurements indicate a short skull with the palatal width and toothrow dimensions of 5. hindei : Hollister (1918 : 94) reports hindei from Archer's Post, Kenya, the type locality of artinii. For the present artinii is considered a provisional synonym of S. h. hindei but there remains the possibility that a subspecies with slender canines is to be found in western Kenya, Uganda and eastern Zaire and it is to this that perhaps artinii refers. If artinii can be distinguished then three subspecies of hindei occur in Kenya, namely S. h. hindei to the north, east and south, S. h. artinii to the west and northwest and 5. h. albigula to the southwest. It is possible also that the rather larger albigula may be found to be specifically valid, but direct sympatry with hindei (including artinii) has yet to be demonstrated. The reported occurrences of the three forms in East Africa display a degree of overlap. Scotoecus hindei albigula Thomas, 1909 Scotoecus albigula Thomas, 1909 : 544. Kirui, Mount Elgon, Kenya, 6000 feet. DIAGNOSTIC. Larger than S. h. hindei, with larger, more massive teeth ; canines longer and heavier. MEASUREMENTS. Length of forearm (3) 35-0-38-5 ; greatest length of skull (3) 15-3-15-5 ; condylobasal length (3) 14-7-14-8 ; condylocanine length (3) 14-8-15-0 ; width of rostrum (3) 7-2-7-7 ; width across anteorbital foramina (3) 5-7-6-0 ; least interorbital width (3) 4-5-4-9 ; zygomatic width ; width of braincase (3) 8-0-8-1 ; mastoid width (3) 9-6-9-9 ; c l -c l 5-3-5-8 ; m s -m 3 (3) 7-5-8-0 ; c-m 3 (5) 5-9-6-0 ; length of complete mandible (3) 11-2-11-6 ; c-m 3 6-5-6-6. LITERATURE. Monard, 1935 : 52 (notes, description repeated, in French) ; Hill & Carter, 1941 : 52 (notes, descriptive data), 177 (measurements of Angolan specimen, from Monard). DISTRIBUTION. KENYA (three examples collected in 1965 by A. N. Start in the Cherangani Hills, east of Mount Elgon, northeast of Kitale, Kenya, at 6000 feet, examined by R. W. Hayman : see Hayman & Hill, 1971 : 36, 37) ; ANGOLA (Monard, *935 : 5 2 I Hill & Carter, 1941 : 52, 177 ; Hayman, 1963 : 104) ; ZAMBIA (Wrough- ton, 1907 : 4 ; Lancaster, 1953 : 18, as Scotoecus woodi, identified as Nycticeius hirundo hindei (= Scotoecus hindei hindei) by Ansell, 1960 : 23, Appendix A, p. no, Appendix B, p. 117 ; Ansell, 1967 : 21, as 5. hirundo hindei (= S. hindei hindei). Hollister (1918 : 94) recorded albigula from Kiriba, Uganda, but Kock (1969 : 192) REVIEW OF SCOTOECUS THOMAS, 1901 183 considers this to represent S. hirundo hindei (= S. hindei hindei) and notes that Kiriba is in the southern Sudan, 10 miles south of Gondokoro. REMARKS. The dimensions of B.M. 7.1.11.6 from Petauke, Zambia (Wroughton 1907 : 4), are similar to those of S. h. albigula, and this specimen has the generally heavier canines and more massive cheekteeth associated with this form, to which it is referred. Specimens in the Kaffrarian Museum (K.M. 1982, 1983) reported as Scotoeciis woodi from the Fort Jameson District by Lancaster (1953 : 18) are identified as 5. hirundo hindei (= S. hindei hindei) by Ansell (see above). These may in fact also be referable to albigula, and provisionally are listed as such here, as is a specimen reported from Mfuwe Camp, Zambia by Ansell, 1967 : 21 as S. hirundo hindei (= S. hindei hindei} but which is larger than this subspecies. It is possible that further specimens may show albigula to be specifically distinct. It occurs as far north in Kenya as Mount Elgon and the Cherangani Hills, while S. h hindei has been obtained from more southerly localities in Katanga, Zaire and in Tanzania. However, in Kenya albigula has been reported only from high elevations, 5. h. hindei only from lower altitudes, and direct sympatry has yet to be shown to occur. Scotoecus hindei falabae Thomas, 1915 Scotoecus falabae Thomas, 1915 : 447. Kabwir, northern Nigeria, 2500 feet. DIAGNOSIS. Similar in most respects to 5. h. hindei but slightly smaller and dorsally a little paler. MEASUREMENTS. Length of forearm (8) 32-0-35-2 ; greatest length of skull (7) 13-8-14-4 ; condylobasal length (6) 13-2-13-5 ; condylocanine length (6) 13-5-13-7 ; width of rostrum (7) 6-5-7-0 ; width across anteorbital foramina (7) 5-0-5-3 ; least interorbital width (7) 4-3-4-6 ; zygomatic width ; width of braincase (6) 7-3-7-9 ; mastoid width (6) 8-4-9-0 ; c^c 1 (7) 4-7-5-0 ; m 3 -m 3 (7) 6-6-6-9 '> c-m 3 (8) 5-2-5-5 ; length of complete mandible (5) 9-8-10-0 ; c-m 3 (7) 5-6-5-8. LITERATURE. Rosevear, 1965 : 300 (further description, as a species), 305, fig. Sob (tragus). DISTRIBUTION. NIGERIA (Kabwir ; Yaba ; Jos, specimens in British Museum (Natural History)) ; CAMEROON (Yagoua ; Mokolo (Mayo Louti), specimens ob- tained by F. de Vree and W. Verheyen). REMARKS. Specimens from the northern Cameroon are very similar in size to 5. h. falabae from Nigeria but on the whole have slightly more massive canines and, to a lesser extent, more massive cheekteeth. In this respect they approach 5. h. hindei. There is apparently no sexual dimorphism in size either in 5. hirundo or in 5. hindei : however, female specimens in both have slightly smaller, more slender canines than do male examples. The canines of specimens attributed to 5. hirundo are generally less massive than are those of specimens referred to S. hindei, but in the case of S. hirundo and 5. hindei falabae the canines of males of hirundo are almost exactly similar in size to those of females of falabae. 184 J. E. HILL The only localities at which specimens attributed to S. hirundo have been found so far to occur sympatrically with others referred to 5. hindei are Yagoua, northern Cameroon (Scotoecus having been hitherto unreported from the Cameroon), and Didessa River, Wollega Province, Ethiopia, with near sympatry at other Ethiopian localities and in Uganda. From Yagoua, F. de Vree and W. Verheyen obtained four examples of Scotoecus, two males and two females. The two male specimens are quite clearly referable to S. hindei falabae with which they are in close agreement. The two female specimens have considerably shorter skulls and agree closely with females of 5. hirundo. F. de Vree and W. Verheyen also obtained a male and female from Mokolo (Mayo Louti) but these are of similar size, agreeing in this respect with falabae to which they are referred. The females from Yagoua are smaller than the female of falabae from Mokolo and consequently are referred to 5. hirundo. Measure- ments of these specimens, and of others from Ethiopia and Uganda, appear in Table i. TABLE Measurements (in millimetres) of specimens from areas o Registration No. Sex Length of forearm 5. hirundo B.M. 72.4421 9 3i'7 B.M. 70.2267 6" 32-0 B.M. 70.2269 6* 31-1 B.M. 70.2263 1 70 2266 1 29-7 B.M. 70.2268 r B.M. 70.2270 J (6) B.M. 72.4420* $ 31-8 B.M. 74.4 3 32-0 B.M. 74.1 9 32-5 B.M. 74.2 9 30-4 B.M. 74.3 9 30-8 B.M. 74.5* $ 30-1 1.430 9 3i-5 I-43I ? 32-8 S. hindei hindei B.M. 72.4423 6* 34-1 B.M. 70.2262 6* 37'2 B.M. 72.4422 6* 33'4 B.M. 63.1151 6* B.M. 65.3435 $ 32-7 S. hindei falabae I-4I5 6* 32-9 1.429 6* 32-6 1.667 6* 32-0 Greatest length Condylo- Condylo- of basal canine skull length length 1.682 33-5 12-6 12-8 13-4 (5) 13-2 13-2 13-5 14-2 14-8 14-4 14-2 13-8 13-9 14-0 13-9 12-4 12-7 I2-I 12-5 (5) 12-4 12-7 12-9 12-5 12-4 12-7 13-6 14-2 13-6 13-9 13-7 13-4 I3-3 13-2 13-2 12-3 12-6 I2-I 12-5 (5) 12-4 12-8 13-0 12-6 12-5 12-9 13-6 14-2 13-5 13-8 13-9 13-5 13-5 13-5 13-6 Rostral width 6-1 6-3 6-5 6-1 -6-4 (6) 6-0 6-5 6-4 6-2 6-2 6-0 7-1 6-7 6-6 6-7 6-5 6-8 7-0 6-5 Width across Least anteorbital interorbit foramina width 4-6 4-9 5'2 4'7 -5-0 (6) 4-6 5'2 5' 4-6 4-9 4-8 5'4 5'3 5'3 5'3 5-2 4-2 4'4 4-2 4'2 4-5 (6) 4-2 4'5 4'7 4'4 4'3 4'3 4-8 4-6 4-6 4-6 4'7 4'4 4'3 4-5 4'4 Young adult REVIEW OF SCOTOECUS THOMAS, 1901 185 ACKNOWLEDGEMENTS My thanks are due to Dr M. J. Largen of the Haile Selassie I University, Addis Ababa, Ethiopia, whose specimens, donated to the British Museum (Natural History), first suggested this study. Dr F. de Vree of the Laboratorium voor Algemene Dierkunde, Rijksuniversitair Centrum, Antwerp, Belgium, has generously allowed me to make use of his important specimens (collected with Dr W. Verheyen) from the northern Cameroon, and, furthermore, has been especially helpful in pro- viding measurements and data from specimens in other European collections, drawn from his own records. I am indebted also to Dr V. Aellen of the Museum d'Histoire Naturelle, Geneva, Switzerland, who has given permission for the publication of data taken from specimens in the collections in his care, and to Mr J. F. Kingdon, whose specimens from Uganda proved invaluable to the course of the work. mpatry or near sympatry between 5. hirundo and S. hindei Width gomatic of Mastoid width braincase width m 3 -m 3 c-m 3 Length of complete mandible c-m 3 Location 7-1 7-5 7-2 7'3 7-5 (4) 7-0 7'5 7-6 7-1 7'3 7'3 8-2 8-3 8-2 -8-3 (5) 7'7 8-3 8-3 8-2 4'3 4'7 4-8 4-6 4'7 (6) 4'3 4-8 4'7 4-6 4'5 4'5 6-0 6-6 6-6 6-3 6-5 (6) 5'9 6-3 6-6 6-3 6-6 6-7 4'7 4-9 4'7 4-8 (6) 4'7 4-8 4'9 9'4 8-9 .9-9 (6) 9-1 9'9 9-6 9'4 9-0 9-6 4'9 5'3 5'3 4-9 5-2 (6) 5-i 5'5 5'3 5'4 Didessa, Ethiopia Gambela, Ethiopia Gambela, Ethiopia Gambela, Ethiopia Gambela, Ethiopia Budongo, Uganda Budongo, Uganda Budongo, Uganda Budongo, Uganda Budongo, Uganda Yagoua, Cameroon Yagoua, Cameroon 7-5 8-9 5-1 6-9 5-3 io-o 5-6 Didessa, Ethiopia 7-8 9-4 5-2 7-0 5-7 10-6 6-1 Bulcha, Ethiopia 7-9 8-8 5-4 5-8 Bulcha, Ethiopia 7-3 8-8 5-3 6-9 5-3 10-3 5-7 West Madi, Uganda 8-0 9-2 5-2 6-5 5-4 10-6 5-8 Nabumali, Uganda 7-5 8-4 4-8 6-7 5-5 io-o 5-8 Yagoua, Cameroon 7-3 8-6 5-0 6-9 5-3 io-o 5-8 Yagoua, Cameroon 7-4 8-6 4-9 6-7 5-2 9-8 5-6 Mokolo (Mayo Louti) , Cameroon 7-4 8-9 4-7 6-8 5-3 io-o 5-7 Mokolo (Mayo Louti), Cameroon 186 J. E. HILL SUMMARY The vespertilionid genus Scotoecus is reviewed in detail, and is considered to include four species, one, pallidus, from India, having been referred at one time to Scoteinus and more recently to Nycticeius. The three remaining, albofuscus, hirundo and hindei are African : evidence is brought forward to support the view that hirundo and hindei are specifically distinct, not conspecific as thought by the majority of modern authors. Both are reported for the first time from the Cameroon and from Ethiopia. REFERENCES ALLEN, G. M. 1908. Notes on Chiroptera. Bull. Mus. comp. Zool. Harv. 52 : 25-62, i pi. 1911. Bats from British East Africa. Bull. Mus. comp. Zool. Harv. 54 : 321-331. 1914. Mammals from the Blue Nile Valley. Butt. Mus. comp. Zool. Harv. 58 : 305-355, i fig. 1939- Checklist of African mammals. Bull. Mus. comp. Zool. Harv. 83 : 1-763. ALLEN, J. A. 1917. In ALLEN, J. A., LANG, H. & CHAPIN, J. P. The American Museum Congo Expedition collection of bats. Part I. Systematic List. Butt. Am. Mus. nat. Hist. 37 : 405-478, 15 figs., i pi. ANCIAUX DE FAVEAUX, F. M. 1958. Speologica africana. Chiropteres des grottes du Haut-Katanga (Congo Beige). Bull. Inst. fr. Afr. noire Ser. A, 20 : 263-275. ANDERSON, J. 1881. Catalogue of Mammalia in the Indian Museum, Calcutta. Part I. Primates, Prosimiae, Chiroptera, and Insectivora. Calcutta. ANSELL, W. F. H. 1960. Mammals of Northern Rhodesia. Lusaka. 1967. Additional records of Zambian Chiroptera. Arnoldia, (Rhodesia), 2 (38) : 1-29. BLANFORD, W. T. 1891. The Fauna of British India, including Ceylon and Burma. Mammalia. Part II. London. CANTOR, T. 1846. Catalogue of the Mammalia inhabiting the Malayan Peninsula and islands. /. Asiat. Soc. Bengal, 15 : 171-203. DE BEAUX, O. 1924. Mammiferi della Somalia Italiana. Elenco delle specie e sottospecie finora riconsciute presenti, con annotazioni di sistematica e zoo-geografia. Atti Soc. ligust. Sci. nat. geogr. N.S. 3 : 149-168. DE WINTON, W. E. 1899. On mammals collected by Lieut. -Colonel W. Giffard in the Northern Territory of the Gold Coast. Ann. Mag. nat. Hist. (7), 4 : 353-359- DOBSON, G. E. 1876. Monograph of the Asiatic Chiroptera and Catalogue of the species of bats in the collection of the Indian Museum, Calcutta. London. 1877. Notes on a collection of Chiroptera from India and Burma, with descriptions of new species. /. Asiat. Soc. Beng. 46 : 310-313. 1878. Catalogue of the Chiroptera in the Collection of the British Museum. London. DOLLMAN, G. 1908. On a collection of bats from Yola, northern Nigeria, collected by Mr. G. M. Webster. Ann. Mag. nat. Hist. (8), 2 : 545-547. 1914. Notes on a collection of East African mammals presented to the British Museum by Mr. G. P. Cosens. Proc. zool. Soc. Lond. 307-318. ELLERMAN, J. R. & MORRISON-SCOTT, T. C. S. 1951. Checklist of Palaearctic and Indian Mammals, 1758-1946. London. - & HAYMAN, R. W. 1953. Southern African Mammals, 1758-1951 : a reclassifica- tion. London. FITZINGER, L. J. 1870. Kritische Durchsicht der Ordnung der Flatterthiere oder Handfliigler (Chiroptera). Familie der Fledermause ( Vespertiliones) . V. Abtheilung. Sber. Akad. Wiss. Wien. 62, (i) : 353-448. FUNAIOLI, U. 1971. Guida breve dei Mammiferi della Somalia. Oltremare. GEOFFROY, M. I. 1831. In BELANGER, M. C. Voyage aux Indes-Orientales, etc. Zool. Paris. REVIEW OF SCOTOECUS THOMAS, 1901 187 HARRISON, D. L. 1958. Two bats (Microchiropetra) new to the fauna of Tanganyika Territory. Durban Mus. Novit. 5 : 95-98, 2 figs. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. // E. Africa nat. Hist. Soc. 23 : 286-295. HAYMAN, R. W. 1954. Notes on some African bats, mainly from the Belgian Congo. Rev. Zool. Bot. afr. 50 : 277-295. 1957- Further notes on African bats. Rev. Zool. Bot. afr. 56 : 41-45. 1963. Mammals from Angola, mainly from the Lunda District. Publcoes cult. Co. Diam. Angola, No. 66 : 81-140, 15 figs. & HILL, J. EDWARDS 1971. In MEESTER, J. & SETZER, H. W. The Mammals of Africa. An Identification Manual. Part 2. Chiroptera. Washington. MISONNE, X. & VERHEYEN, W. 1966. The bats of the Congo and of Rwanda and Burundi. Annls Mus. r. Afr. cent. ser. in 8, Sci. zool. No. 154 : i-viii, 1-105, 2O P* s - HILL, J. ERIC & CARTER, T. D. 1941. The mammals of Angola, Africa. Bull. Am. Mus. nat. Hist. 78 : 1-211, 34 figs., 17 pis. HOLLISTER, N. 1918. East African mammals in the United States National Museum. Part i. Insectivora, Chiroptera and Carnivora. Bull. U.S. natn. Mus. No. 99 : 1-194, 55 P^ s - JENTINK, F. A. 1887. Museum d'Histoire Naturelle de Pays-Bas etc. Tome XII. Cata- logue Systdmatique de Mammiferes. (Rongeurs, Insectivores, Cheiropteres, Edentes et Marsupiaux.) Leiden. KHAJURIA, H. 1951. Taxonomic studies on some Indian Chiroptera. Rec. Indian Mus. 50 : 113-128. KOCK, D. 1969. Die Fledermaus-Fauna des Sudan (Mammalia, Chiroptera). Abh. sencken. naturforsch. Ges. 521 : 1-238, 20 figs. KOOPMAN, K. F. 1965. Status of forms described or recorded by J. A. Allen in 'The American Museum Congo Expedition Collection of Bats'. Am. Mus. Novit. No. 2219 : 1-34. LANCASTER, D. G. 1953. A check list of the mammals of Northern Rhodesia. Lusaka. LELEUP, N. 1956. La faune cavernicole du Congo Beige et considerations sur les Coleopteres reliques d'Afrique intertropicale. Annls Mus. r. Congo Beige, ser. in 8, Sci. zool. No. 46 : 1-170, 118 figs., 5 pis. MILLER, G. S. 1907. The families and genera of bats. Bull. U.S. natn. Mus. 57 : i-xvii, 1-282, 49 figs., 14 pis. MONARD, A. 1936. Contribution a la mammalogie d'Angola et Prodrome d'une faune d'Angola. Arches Mus. Bocage, 6 : 1-309. ROSEVEAR, D. R. 1965. The Bats of West Africa. London. SIDDIQI, M. S. 1961. Checklist of mammals of Pakistan with particular reference to the mammalian collection in the British Museum (Natural History), London. Biologia, Lahore 7 : 93-225. SIMPSON, G. G. 1945. The principles of classification and a classification of mammals. Bull. Am. Mus. nat. Hist. 85 : i-xvi, 1-349. SINHA, Y. P. & CHAKRABORTY, S. 1971. Taxonomic status of the vespertilionid bat, Nycti- cejus emarginatus Dobson. Proc. zool. Soc. Calcutta 24 : 53-59, 3 figs., i tab. SWYNNERTON, G. H. & HAYMAN, R. W. 195 1. A checklist of the land mammals of the Tanganyika Territory and the Zanzibar Protectorate. // E. Africa nat. Hist. Soc. 20 : 274- 392. TATE, G. H. H. 1942. Results of the Archbold Expeditions. No. 47. Review of the Ves- pertilionine bats, with special attention to genera and species of the Archbold Collections. Bull. Am. Mus. nat. Hist. 80 : 221-297, 5 n s - TEMMINCK, C. J. 1840. Monographies de Mammalogie. Paris. THOMAS, O. 1890. Description of a new Scotophilus from the Gambia, with remarks on some of the allied species. Annali Mus. civ. Stor. nat. Giacomo Doria (2), 9 : 84-88. 1901 . A new Scotophiline bat from British East Africa, with the description of a new genus of the group. Ann. Mag. nat. Hist. (7), 7 : 263-265. 1909. New African mammals. Ann. Mag. nat. Hist. (8), 4 : 512-549. i88 J. E. HILL THOMAS, O. 1911. List of small mammals obtained by Mr. G. Fenwick Owen on the Upper Gambia and in Fouta Djallon. Ann. Mag. nat. Hist. (8), 8 : 117-124. 1915. A new bat from Northern Nigeria. Ann. Mag. nat. Hist. (8), 16 : 447-448. 1917. A new bat of the genus Scotoecus. Ann. Mag. nat. Hist. (8), 19 : 280-281. & DORIA, G. 1886. Note intorno al alcuni Chirotteri appartenenti al Museo Civico di Geneva e descrizione di due nuove specie del genere Phyllorhina. Annali Mus. civ. Stor. nat. Giacomo Doria (2), 4 : 201-207. TROUESSART, E.-L. 1897. Catalogus mammalium tarn viventium quam fossilium. Tomus I. Primates, Prosimiae, Chiroptera, Insectivora, Carnivora, Rodentia, Pinnipedia. Berlin. 1904. Catalogus mammalium tarn viventium quam fossilium. Quinquennale supple- mentum. Berlin. VESEY-FITZGERALD, D. F. 1964. Mammals of the Rukwa Valley. Tanganyika Notes Rec. No. 62 : 61-72. WAGNER, J. A. 1840. Die Sdugethiere in Abbildungen nach der Natur mit Beschreibung von D. Johann Christian Daniel von Schreber. Supplementband i . Leipzig. 1855. Die Sdugethiere in Abbildungen nach der Natur mit Beschreibung von D. Johann Christian Daniel von Schreber. Supplementband 5. Leipzig. WETTSTEIN, O. v. 1916. Neue Affen und Fledermause aus Nordost-Afrika. Anz. Akad.Wiss. Wien. 53 : 189-192. 1918. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Kaiserliche Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommen zoologischen Expedition nach dem Anglo- Agyptischen Sudan (Kordofan) 1914. II. Bearbeitung der auf der Expedition gesammelten Vogel und Saugethiere. Denkschr. Akad. Wiss. Wien. (1917), 1918, 94 : 555-693, 13 figs., 4 tabs., map. WROUGHTON, R. C. 1907. On a collection of mammals made by Mr. S. A. Neave in Rhodesia, north of the Zambesi, with field notes by the collector. Manchester Memoirs, 51 (5) : 1-39. 1916. Bombay Natural History Society's Mammal Survey of India, Burma and Ceylon. Report No. 24. Sind. /. Bombay nat. Hist. Soc. 24 : 749-758. J. E. HILL Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON SW7 5BD A LIST OF 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 sNU :; CATALOGUE OF THE TYPES TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) I. SUPERFAMILY PSEUDOTRACHEATA R. J. LINCOLN AND J. P. ELLIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 5 LONDON: 1974 CATALOGUE OF THE TYPES OF TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) I. SUPERFAMILY PSEUDOTRACHEATA BY ROGER ]. LINCOLN AND JOAN P. ELLIS Pp. 189-246 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 27 No. 5 LONDON: 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, issued in Jive series corresponding to the 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. 27, No. 5, 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.) Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 23 December, 1974 Price 3.10 CATALOGUE OF THE TYPES OF TERRESTRIAL ISOPODS (ONISCOIDEA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) I. SUPERFAMILY PSEUDOTRACHEATA By ROGER J. LINCOLN AND JOAN P. ELLIS THE superfamily Pseudotracheata comprises the following ten families of terrestrial isopods : Cylisticidae, Porcellionidae, Balloniscidae, Trachelipidae, Atlantidiidae, Armadillidiidae, Eubelidae, Sphaeroniscidae, Actoeciidae, Armadillidae, encompas- sing all those groups of woodlice which possess the characteristic pseudotracheal respiratory apparatus (Vandel, 1962, 1963, 1968, 1970). The collections of the British Museum (Natural History) are very rich in type material of this group of woodlice, acquired in large part from the extensive collec- tions of Gustave Budde-Lund, A. M. Norman and Karl W. Verhoef, and to a lesser extent from C. L. Koch, Adrien Dollfus, J. Omer-Cooper, K. H. Barnard, W. E. Collinge and others. This important collection of types has now been indexed and catalogued for the first time, and it is hoped at a future date to provide a similar catalogue listing the rest of the holdings of woodlice types. The catalogue contains in all some 672 separate items relating to 522 species in 77 genera. The genera are listed alphabetically within families, and the species are similarly listed within the appropriate genus. All available information, including the reference to the original description, number and sex of specimens, type status, locality, date, collector, method of acquisition and British Museum (Natural History) registration number, is given with each item. Unless otherwise stated the material is preserved in spirit. Where the author's name appears in brackets it indicates that the entry is made in a combination other than the original combination, in which case the latter is given in square brackets thus : albicornis (Dollfus) (iSg6b : 3) [Lucasius albicornis] Where a particular species is now considered a junior synonym of another species the latter is also indicated in brackets after the entry : sphinx Verhoeff (i93ia : 533) [Transferred to Porcellio lamettatus Uljanin] A full index of species, genera and families is provided at the end of the catalogue. The collection also contains a large number of specimens labelled as types, mostly from the Budde-Lund collection, for which no reference can be found in published literature. It seems probable 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 167 items is given in Appendix i, and a list of the manuscript names has been deposited in the library of the British Museum (Natural History). I9 2 R. J. LINCOLN AND J. P. ELLIS Family GYLISTICIDAE CYLISTICUS Schnitzler annulicornis Verhoeff (igoSb : 184) SYNTYPES: three females. Reg. no. 1907:11:4:36-38. Toscana region, Italy. Purchased from K. W. Verhoeff. biellensis Verhoeff (i93oa : 176) SYNTYPES : two females. Reg. no. 1930:5:26:54-55. Piemonte, Italy. Purchased from K. W. Verhoeff. SYNTYPES: two males; six females. Reg. no. 1937:7:6:109-113. Piemonte, Italy. Purchased from K. W. Verhoeff. caucasius Verhoeff (1917 : 157) SYNTYPES: male. Reg. no. 1921:6:10:46. Gagri, Caucasus Mts. Collected by N. Lignau. Purchased from K. W. Verhoeff. dentifrons Budde-Lund (1885 : 81) HOLOTYPE : fragments. Reg. no. 1921:10:18:4212. Astrakhan, USSR. Budde-Lund Collection. esterelanus Verhoeff (1917 : 155) SYNTYPE : female. Reg. no. 1921:6:10:45. Esterel Mts, France. Purchased from K. W. Verhoeff. gracilipennis Budde-Lund (1885 : 79, 229) SYNTYPE: male. Reg. no. 1911:11:8:10063. Rome, Italy. Collected by Bergs0e. Norman Collection (ex Copenhagen Museum). SYNTYPE : male. Reg. no. 1921:10:18:4213. (Redetermined as Cylisticus esterelanus Verhoeff.) Pierrefeu, France. Collected by E. Simon. Budde-Lund Collection. SYNTYPES : three males ; three females. Reg. no. 1921:10:18:2414-4218. Rome, Italy. Collected by Bergs0e. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1956:10:10:201-202. Rome, Italy. Collected by Bergs0e. Presented by University College, Dundee. nasatus Verhoeff I93ia : 527 SYNTYPE : one juvenile. Reg. no. 1931:4:27:42. Villa Napoleon, Elba I. Collected by K. Strasser. Purchased from K. W. Verhoeff. SYNTYPE : female. Reg. no. 1930:5:26:52. Ormea, Liguria, Italy. Purchased from K. W. Verhoeff. pallidus Verhoeff (19280 : 159) SYNTYPE : female. Reg. no. 1930:5:26:53. Liguria, Italy. Purchased from K. W. Verhoeff. plurnbeus Verhoeff (igoib : 73) SYNTYPE : female. Reg. no. 1901:9:19:41. Lake Garda, Italy, Purchased from K. W. Verhoeff. plurnbeus bergomatius Verhoeff (19280 : 162) SYNTYPES: one male ; one female. Reg. no. 1928:7:4:107-108. Bergamo, Italy. Pur- chased from K. W. Verhoeff. plurnbeus umbricus Verhoeff (19280 : 163) SYNTYPES: one male; one female. Reg. no. 1928:7:4:105-106. Umbria, Italy. Pur- chased from K. W. Verhoeff. pontremolensis Verhoeff (i936a : 112) SYNTYPES: one male; three females. Reg. no. 1937:7:6:114-117. Spezia, N. Italy. Purchased from K. W. Verhoeff. suberorum Verhoeff (193 la : 526) SYNTYPE : juvenile. Reg. no. 1931:4:27:41. Monte Massoncello, Italy. Purchased from K. W. Verhoeff. transsilvaticus Verhoeff (igoSb : 185) SYNTYPE: female. Reg. no. 1907:11:4:39. Siebenbiirgens. Purchased from K. W. Verhoeff . TYPES OF TERRESTRIAL ISOPODS 193 Family PORCELLIONIDAE ACAEROPLASTES Verhoeff argentarius (Verhoeff) (i93ia : 541) [Metoponorthus (A.) argentarius] SYNTYPE : female. Reg. no. 1931:4:27:62. Monte Argentario, Italy. Purchased from K. W. Verhoeff. melanurus (Budde-Lund) (1885 : 181) [Metoponorthus melanurus] SYNTYPES : two females. Reg. no. 1921:10:18:5273-5274. Porto Calvien, Corsica. Collected by E. Simon. Budde-Lund Collection. SYNTYPES: two males; one female. Reg. no. 1956:10:10:145-146. Bona, Algeria. Collected by Meinert. Presented by University College, Dundee. AGABIFORMIUS Verhoeff hirtus (Aubert & Dollfus) (1890 : 8) [Lucasius hirtus] SYNTYPE : male. Reg. no. 1911:11:8:10532. Marseille, France. 'In soil at the foot of a wall.' Collected by A. Dollfus. Norman Collection. SYNTYPE : female. Reg. no. 1921:10:18:4023. Marseille, France. Budde-Lund Collec- tion. minutus (Budde-Lund) (igoga : 8) [Angara minuta] SYNTYPES : three females. Reg. no. 1921:10:18:4028-4030. Porto Tunis, Tunisia. Col- lected by F. Silvestri. Budde-Lund Collection. SYNTYPES: one male; one female. Reg. no. 1921:10:18:4031-4032. Sidi Bel Akaren. Collected by F. Silvestri. Budde-Lund Collection. obtusus (Budde-Lund) (igoga : 8) [Angara obtusa] SYNTYPES: one male; four females. Reg. no. 1921:10:18:4061-4065. Cairo, UAR. 18.5.1901. Collected by L. A. Jagerskjold. Budde-Lund Collection. rufobrunneus Omer-Cooper (1923 : 401) SYNTYPE: fragments. Reg. no. 1922:5:18:6. Amara, Iraq. 13.2.1918. Collected by W. E. Evans. Presented by J. Omer-Cooper. SYNTYPES : micropreparations of appendages : six males, one female. Reg. no. 1922:5: 18:51-57. Amara, Iraq. 13.2.1918. Collected by W. E. Evans. Presented by J. Omer- Cooper. CAEROPLASTES Verhoeff porphyrivagus (Verhoeff) (1918 : 134) [Metoponorthus (C.) porphyrivagus] SYNTYPE : female. Reg. no. 1921:6:10:31. French Riviera. Purchased from K. W. Verhoeff. DESERTONISCUS Verhoeff subterraneus Verhoeff (igsob : 120) SYNTYPE : female. Reg. no. 1930:5:26:73. Turkestan, USSR. Purchased from K. W. Verhoeff. ENNURENSIS Collinge hispidus Collinge (19153, : 143) PARATYPE : female. Reg. no. 1919:4:26:519. Ennur, near Madras, India. 'Under stones on sand 19.10.1913.' Collected by N. Annandale. Collinge Collection. 194 R - J- LINCOLN AND J. P. ELLIS GERUFA Budde-Lund hirticornis Budde-Lund (igogb : 59) SYNTYPES : two females. Reg. no. 1921:10:18:1468-1469. Cape Flats, South Africa. Collected by L. Schultze. Budde-Lund Collection. macrops Barnard (1932 : 276) SYNTYPES : four males ; nineteen females. Reg. no. 1933:1:25:208-215. Oudebosch, River Zonder End Mts, Cape Province, South Africa. Collected by K. H. Barnard. Pre- sented by the South African Museum. marmorata Barnard (1932 : 277) SYNTYPES : four males ; thirty-six females. Reg. no. 1933:1:25:216-227. George, Cape Province, South Africa. Collected by K. H. Barnard. Presented by the South African Museum. montana Barnard (1932 : 275) SYNTYPES : five females. Reg. no. 1933:1:25:204-207. Riversdale Mts, Cape Province, South Africa. Collected by K. H. Barnard. Presented by the South African Museum. HALOPORCELLIO Verhoeff penicilliger Verhoeff (1917 : 170) HOLOTYPE : male. Reg. no. 1921:6:10:78. Yafo, Israel. Collected by Aharoni. Pur- chased from K. W. Verhoeff. sphinx Verhoeff (i93ia : 533) [Transferred to Porcellio lamellatus Uljanin] SYNTYPE : female. Reg. no. 1931:4:27:61. Populonia, Massoncella, Italy. Purchased from K. W. Verhoeff. HEMILEPISTUS Budde-Lund bodenheimeri Verhoeff (i93ib : 40) [Transferred to Hemilepistus reaumuri (Audouin & Savigny)} SYNTYPE : female. Reg. no. 1931:4:27:68. Palestine. Collected by Bodenheimer. Purchased from K. W. Verhoeff. SYNTYPES: one male; three females. Reg. no. 1938:7:7:41-44. Palestine. Collected by Bodenheimer. Purchased from K. W. Verhoeff. cristatus Budde-Lund (1885 : 153) SYNTYPES: one male ; three females. Reg. no. 1911:11:8:10443-10446. Serdscen, Iran. Norman Collection. SYNTYPES : one male ; one female. Reg. no. 1956:10:10:156-157. Serdscen, Iran. Presented by University College, Dundee. elongatus Budde-Lund (1885 : 160) HOLOTYPE : female. Reg. no. 1921:10:18:4103. 'Taschburun in Transcaucasius.' June 1879. Collected by A. Brandt. Budde-Lund Collection. nodosus Budde-Lund (1885 : 159) SYNTYPES: one male; one female. Reg. no. 1921:10:18:4115-4116. 'Tschinas, Rossia Asiatica.' 1878. Collected by V. Russow. Budde-Lund Collection. palaestinus Verhoeff (i93ib : 38) SYNTYPES : two males. Reg. no. 1931:4:27:65-67. Palestine. Purchased from K. W. Verhoeff. SYNTYPES: three males ; three females. Reg. no. 1938:7:7:35-40. Palestine. Purchased from K. W. Verhoeff. schirasi Lincoln (1970 : 127) PARATYPE : male. Reg. no. 1970:199:1. Shiraz, Iran. Collected by Kollar. Budde- Lund Collection. zachvatkini Verhoeff (i93ob : 122) [Transferred to H. nodosus Budde-Lund] SYNTYPE : female. Reg. no. 1930:5:26:74. Turkestan, USSR. Purchased from K. W. Verhoeff. TYPES OF TERRESTRIAL ISOPODS 195 HEMIPORCELLIO Collinge carinatus Collinge (igisa : 145) PARATYPE : female. Reg. no. 1919:4:26:458. Lake Chilka, Rambha, Ganjam District, Madras, India. 'At edge of Lake, 1913.' Collinge Collection. imtnsi (Collinge) (i9i4a : 207) \Povcellio immsi] HOLOTYPE : female. Reg. no. 1914:8:26:1. Allahabad, India. 16.9.1907. Collected and presented by A. D. Imms. PARATYPES : one male ; one juvenile female. Reg. no. 1919:4:26:456-457. Allahabad, India. 16.9.1907. Collected by A. D. Imms. Collinge Collection. INCHANGA Barnard natalensis Barnard (1932 : 278) SYNTYPES : five males ; eleven females. Reg. no. 1933:1:25:228-233. Inchanga, Natal, South Africa. 1917. Collected by K. H. Barnard. Presented by the South African Museum. LEPTOTRICHUS Budde-Lund politus Omer-Cooper (1923 : 402) SYNTYPES: three males; one female. Reg. no. 1922:5:18:7-10. Amara, Iraq. 1918. Collected by R. G. Tame. Presented by J. Omer-Cooper. SYNTYPES : micropreparations of appendages : three males ; six females. Reg. no. 1922:5:18:35-43. Amara, Iraq. 1918. Collected by R. G. Tame. Presented by J. Omer-Cooper. LUCASIUS Kinahan pallidus (Budde-Lund) (1885 : 134) [Porcellio pallidus] SYNTYPE : male. Reg. no. 1921:10:18:5182. Algier. Budde-Lund Collection. SYNTYPES : three males. Reg. no. 1921:10:18:5183-5185. Tlemcen, Algeria. Collected by E. Simon. Budde-Lund Collection. SYNTYPE : female. Reg. no. 1921:10:18:5186. Mid-France. Collected by E. Simon. Budde-Lund Collection. MAHEHIA Budde-Lund bicornis Budde-Lund (1913 : 376) SYNTYPES : one male ; two females. Reg. no. 1913:1:8:64-66. Silhouette I., Seychelles, Indian Ocean. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. SYNTYPES : Seven males ; seven females. Reg. no. 1921:10:18:997-1008. Silhouette I., Seychelles, Indian Ocean. Budde-Lund Collection. laticauda Budde-Lund (1913 : 376) SYNTYPES : six males ; one female. Reg. no. 1913:1:8:58-63. Cascade, Mt Alphonse, Mahe I., Seychelles, Indian Ocean. 1800 ft, 'in base of Pandanus leaf. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. tnaculata Budde-Lund (1913 : 375) SYNTYPE : female. Reg. no. 1913:1:8:50. Mt Alphonse, Mahe I., Seychelles, Indian Ocean. 3.12.1905. Collected by the 'Sealark' Expedition. Presented by J. S. Gardiner. SYNTYPE : female. Reg. no. 1921:10:18:1021. Top of Mt Morne, Mahe I., Seychelles, Indian Ocean. 2700 ft. Budde-Lund Collection. SYNTYPE : few fragments only. Reg. no. 1921:10:18:1022. Seychelles, Indian Ocean. Budde-Lund Collection. I 9 6 R. J. LINCOLN AND J. P. ELLIS METOPONORTHUS Budde-Lund approximates Budde-Lund (1885 : 185) SYNTYPE : male. Reg. no. 1921:10:18:5230. Sebastopol. Budde-Lund Collection. benaci L. Koch (1901 : 56) SYNTYPES : one male ; three females. Reg. no. 1925:7:22:697-700. Torbole, Italy. Koch Collection. cilicius Verhoeff (1918 : 138) SYNTYPE : female. Reg. no. 1921:6:10:35. Toros Gari Mts, Cilicia, Turkey. Purchased from K. W. Verhoeff. coxalis Budde-Lund (1885 : 175) SYNTYPE: female. Reg. no. 1911:11:8:10520. 'Ouled Anteurs', Algeria. Collected by E. Simon. Norman Collection. SYNTYPES: one male; two females. Reg. no. 1921:10:18:5235-5237. 'Rochers des Ouled Anteurs', Algeria. Collected by E. Simon. Budde-Lund Collection. frontosus Budde-Lund (1885 : 183) SYNTYPE : male. Reg. no. 1911:11:8:10525. Biskra, Algeria. Collected by Meinert. Norman Collection (ex E. Simon collection). SYNTYPES: one male; one female. Reg. no. 1921:10:18:5239-5240. Collected by Meinert. Budde-Lund Collection. fuscomarmoratus Budde-Lund (1885 : 189) SYNTYPE: female. Reg. no. 1911:11:8:10521. Oran, Algeria. Collected by E. Simon. Norman Collection. SYNTYPES: three females. Reg. no. 1921:10:18:5241-5243. Oran, Algeria. Collected by E. Simon. Budde-Lund Collection. gravei Verhoeff (1918 : 135) SYNTYPE : male. Reg. no. 1921:6:10:36. Sicily. Purchased from K. W. Verhoeff. hispida (Miers) (i877b : 676) {Porcellio (Porcellionides) hispida] SYNTYPES : one male ; one female. Reg. no. 1879 : 21. Mongolia. Presented by Prof. Wrzesniowsky. lacteolus Budde-Lund (1885 : 186) SYNTYPE: female. Reg. no. 1911:11:8:10492. 'Bou-Saada', Algeria. Collected by E. Simon. Norman Collection. SYNTYPES : one male ; one female. Reg. no. 1921:10:18:4400. 'Bou-Saada', Algeria. Collected by E. Simon. Budde-Lund Collection. littoralis Budde-Lund (1885 : 179) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:5247-5248. 'Kertsch', Crimea. Collected by V. N. Uljanin. Budde-Lund Collection. SYNTYPES: two males; one female. Reg. no. 1921:10:18:5249-5251. (Redetermined Protracheoniscus occidentalis Vandel.) Montpellier, France. Collected by E. Simon. Budde- Lund Collection. SYNTYPES: one male; one female. Reg. no. 1921:10:18:5252-5253. Crimea, Ukraine, USSR. Collected by V. N. Uljanin. Budde-Lund Collection. myrmicidarum Verhoeff (1918 : 132) [Transferred to M. (Myrmeconiscus) myrmecophilus myrmecophilus (Stein)] SYNTYPES: one male ; one female. Reg. no. 1921:6:10:37-38. Sicily. Purchased from K. W. Verhoeff. nigrobrunneus Budde-Lund (1896 : 47) SYNTYPE : head only (both antennae and mouthparts from right-hand side missing). Reg. no. 1921:10:18:5328. Kappari, Greece. 14.6.1887. Collected by Oertzen. Budde- Lund Collection. philoscoides Budde-Lund (1885 : 175) SYNTYPES: three females. Reg. no. 1911:11:8:10506-10508. Algeria. Collected by E. Simon. A. M. Norman Collection (ex Dollfus collection). TYPES OF TERRESTRIAL ISOPODS 197 SYNTYPES : six females. Reg. no. 1921:10:18:5329-5334. Oned-Zitoun, Algeria. May 1883. Collected by E. Simon. Budde-Lund Collection. SYNTYPES: two males ; one female. Reg. no. 1928:12:1:1518-1520. Algeria. Collected by E. Simon. Stebbing Collection (ex Dollfus collection). SYNTYPES : two females. Reg. no. 1956:10:10:141-142. Oned-Zitoun, Algeria. Col- lected by E. Simon. Presented by University College, Dundee. pruinosus anconanus Verhoeff (ig28c : 144) SYNTYPE : male. Reg. no. 1928:7:4:89. Ancona, Italy. Purchased from K. W. Verhoeff. SYNTYPES : two males ; one female. Reg. no. 1973:152:3. Mid-Italy. Larwood Collec- tion (ex Verhoeff collection). pruinosus meleagris Budde-Lund (1885 : 168) [Metoponorthus meleagris] SYNTYPES: two males ; seven females. Reg. no. 1921:10:18:5279-5287. France ('Gallia Meridionalis'). Collected by E. Simon. Budde-Lund Collection. sabuleti Budde-Lund (1885 : 186) SYNTYPES : one male ; one female. Reg. no. 1921:10:18:3920-3921. Biskra, Algeria. Budde-Lund Collection. SYNTYPES: two females. Reg. no. 1921:10:18:3922-3923. Algeria. Budde-Lund Collec- tion. SYNTYPE : male. Reg. no. 1921:10:18:4735. Biskra Mt, Algeria. Collected by Tat- schanowski. Budde-Lund Collection. sexfasciatus Budde-Lund (1885 : 167) SYNTYPES : five females. Reg. no. 1921:10:18:5306-5310. Algiers. Collected by E. Simon. Budde-Lund Collection. sexfasciatus asifensis Verhoeff (i938b : 63) [Metoponorthus asifensis] SYNTYPE : female. Reg. no. 1938:7:7:45. Atlas, Morocco. Purchased from K. W. Verhoeff. virgatus Budde-Lund (1885 : 182) SYNTYPES: two males; nine females. Reg. no. 1921:10:18:5514-5525. Florida, N. America. Collected by Leuckart. Budde-Lund Collection. viridis Budde-Lund (1885 : 179) SYNTYPES : three males. Reg. no. 1921:10:18:5526-5527. Bona, Algeria. Collected by E. Meinert. Budde-Lund Collection. MICA Budde-Lund tardus (Budde-Lund) (1885 : 305) [Porcellio tardus] SYNTYPE: female. Reg. no. 1911:11:8:10533. Tlemcen, Algeria. Collected by E. Simon. Norman Collection. PORCELLIO Latreille acustiserra Barnard (1940 : 358) SYNTYPES: thirteen males; twenty females. Reg. no. 1949:2:2:138-162. Lakes of Addas, shore of Hora Harsadi, Ethiopia, 7000 ft. 3.12.1936. Collected and presented by J. Omer-Cooper. aegaeus Verhoeff (igo7a : 257) SYNTYPE : female. Reg. no. 1907:11:4:44. Naxos I., Greece. Purchased from K. W. Verhoeff. albicornis (Dollfus) (i8g6b : 3) [Lucasius albicornis] SYNTYPE: male. Reg. no. 1911:11:8:10529. Ficuzza, Palermo, Sicily. 900 m. Norman Collection (ex Dollfus collection). SYNTYPES : not separated from other (non-type) specimens. Reg. no. 1921:10:18:5163- 5172 (part). Ficuzza, Palermo, Italy. Budde-Lund Collection (ex Dollfus collection). 198 R. J. LINCOLN AND J. P. ELLIS albinus Budde-Lund (1885 : 142) SYNTYPE : male. Reg. no. 1921:10:18:5163. Onargla, Sahara, Algeria. Collected by E. Simon. Budde-Lund Collection. albomarginatus Vogl (1875 : 516) SYNTYPES : four males. Reg. no. 1925:7:22:449-451. Syra I., Cyclades, Greece. Col- lected by Ebner. Koch Collection. (illuaudi Dollfus (1893 : 52) SYNTYPES : two males. Reg. no. 1921:10:18:4249-4250. Canary Is. Collected by C. Alluaud. Budde-Lund Collection. atnoenus Dollfus (i829b : 178) [Transferred to Porcellio violaceus Budde-Lund] SYNTYPES: two males; one female. Reg. no. 1911:11:8:10396-10398. Cuenca, Spain. Norman Collection. angustulus Budde-Lund (1885 : 146) SYNTYPE: female. Reg. no. 1911:11:8:10429. Bou-Saada, Algeria. Collected by E Simon. Norman Collection (ex Dollfus collection). SYNTYPES : one male ; four females. Reg. no. 1921:10:18:3792-3796. Algeria. Col- lected by E. Simon. Budde-Lund Collection. ater Budde-Lund (1896 : 45) SYNTYPES: one male ; one female. Reg. no. 1921:10:18:4284-4285. Rothenthurm Pass, Transylvania, Rumania. Budde-Lund Collection. auritus Budde-Lund (1885 : 126) SYNTYPE : female. Reg. no. 1921:10:18:4978. Seville, Spain. Collected by Fr. Meinert. Budde-Lund Collection. barroisi Dollfus (i8g2c : 127) SYNTYPE : female. Reg. no. 1911:11:8:10400. 'Ouadys de la Mer Morte.' Collected by T. Barrois. Norman Collection (ex Dollfus collection) batesoni Collinge (i9i5b : 461) PARATYPE : male. Reg. no. 1919:4:26:540. S. Spain, 1894. Collected by W. Bateson. Collinge Collection. bistriatus Budde-Lund (1885 : 88) SYNTYPE : male. Reg. no. 1921:10:18:4288. Istanbul, Turkey. Collected by Jelski. Budde-Lund Collection. blattarius Budde-Lund (1885 : 131) SYNTYPE : male. Reg. no. 1921:10:18:4286. Algeria. Budde-Lund Collection. brevipennis Budde-Lund (1885 : 97) SYNTYPES : one male ; three females. Reg. no. 1921:10:18:3798-3801. Algeria. Col- lected by E. Simon. Budde-Lund Collection. btuldelundi Simon (1885 : 10) SYNTYPE : female. Reg. no. 1911:11:8:10428. Tunisia. Norman Collection (ex Dollfus collection). SYNTYPES: two males ; one female. Reg.no. 1921:10:19:4979-4981. Tunisia. Budde- Lund Collection (ex Simon collection). calrnani Omer-Cooper (1923 : 399) SYNTYPES: two females. Reg. no. 1922:5:18:4-5. Baku, Azerbaydzhan, USSR. Col- lected by P. A. Buxton. Presented by J. Omer-Cooper. canariensis Dollfus (1893 : 5) SYNTYPE: male. Reg. no. 1911:11:8:10409. Canary Is. Collected by C. Alluaud. Normal Collection (ex Dollfus collection). SYNTYPES : two males ; one female ; one juvenile. Reg. no. 1921:10:18:4289-4291. Canary Is. Collected by C. Alluaud. Budde-Lund Collection. carthaginensis Silvestri (1897 : 415) [Transferred to Porcellio olivieri Audouin] SYNTYPES: four males; seven females. Reg. no. 1921:10:18:3802-3811. Carthage (ruins), Tunisia. Budde-Lund Collection. TYPES OF TERRESTRIAL ISOPODS 199 cayennensis Miers (iSyyb : 667) [Transferred to Porcellio scaber Latreille] SYNTYPES : two females. Reg. no. 1941:6:27:15. Cayenne, Guiana. Presented by Prof. Wrzesniowsky. cognatus L. Koch (1901 : 52) SYNTYPE : female. Reg. no. 1925:7:22:453-454 (part). Meran Molde. Koch Collection. SYNTYPE : female. Reg. no. 1925:7:22:453-454 (part). Ratzes. Koch Collection. contractus Dollfus (i8g2c : 8) SYNTYPE: male. Reg. no. 1892:12:6:1. Syria. Presented by W. D'Arcy Thompson. SYNTYPES: one male ; one female. Reg.no. 1911:11:8:10384-10385. Syria. Collected by L. Bleuse. Norman Collection. SYNTYPES: two males. Reg. no. 1928:12:1:1464-1465. Saida, Syria. Collected by L. Bleuse. Stebbing Collection. cruentatus L. Koch (1901 : 37) SYNTYPES : one male ; one female. Reg. no. 1925:7:22:479-480. Sugenheim, W. Ger- many. Koch Collection. debueni Dollfus (i892b : 176) SYNTYPES: three females. Reg. no. 1911:11:8:10393-10395. Villa Rutis, Coruna, Spain. Collected by Bolivar. Norman Collection. SYNTYPE : female. Reg. no. 1921:10:18:4360. Coruna, Spain. Collected by Bolivar. Budde-Lund Collection. dispar Verhoeff (190 id : 407) SYNTYPES: one male; two females. Reg. no. 1901:9:19:86-88. Coimbra, Portugal. Collected by Moller. Purchased from K. W. Verhoeff. SYNTYPE: female. Reg. no. 1907:11:4:72. Coimbra, Portugal. Collected by Moller. Purchased from K. W. Verhoeff. evansi Omer-Cooper (1923 : 398) HOLOTYPE : male. Paratypes : one male; one female. Reg. no. 1922: 5:18:1-3. Amara, Mesopotamia, Iraq. Collected by C. L. Boulenger. Presented by J. Omer-Cooper. PARATYPES : micropreparations of appendages : two males ; five females. Reg. no. 1922:5:18:44-50. Amara, Iraq. Collected by C. L. Boulenger. Presented by J. Omer- Cooper. ficorutn Verhoeff (i93ia : 532) SYNTYPE : male. Reg. no. 1931:4:27:57. Monte Massoncello, Italy. Purchased from K. W. Verhoeff. ficulneus Budde-Lund (1885 : 98) SYNTYPES : two males ; two females. Reg. no. 1921:10:18:4995-4998. Syria. Collected by E. Simon. Budde-Lund Collection. SYNTYPES : two females. Reg. no. 1956:10:10:228-229. Syria. Collected by E. Simon. Presented by University College, Dundee. flavocinctus Budde-Lund (1885 : 109) SYNTYPES : two females. Reg. no. 1956:10:10:226-227. Spain. Collected by L. Lund and Fr. Meinert. Presented by University College, Dundee. flavovittata Miers (i877b : 669) [Transferred to Metoponorthus pruinosus (Brandt)] SYNTYPES : four females. Reg. no. 1879:21. Cayenne, Guiana. Presented by Prof. Wrzesniowsky. gallicus Dollfus (1903 : 63) SYNTYPES : two females. Reg. no. 1907:11:4:73-74. Pyr6n6es. Purchased from K. W. Verhoeff. gemtnulatus Dana (1853 : 725) [Transferred to Porcellio scaber Latreille] SYNTYPES : five males ; one female. Reg. no. 1877:3. San Francisco, N. America. Presented by W. N. Lockington. graniger Miers (1876 : 226) HOLOTYPE : female. Reg. no. 1973:512:1 (original registration no. 1854:4). New Zealand. Presented by Col. Bolton. R. J. LINCOLN AND J. P. ELLIS Reg. no. 1921:10:18:3812. Spain. Collected by Reg. no. 1921:10:18:4425- Spain. Collected by Fr. Bona, Algeria. Collected by Fr. Meinert. Reg. no. 1921:10:18:4632-4633. 'Ordus.' Pre- Pre- granuliferus Budde-Lund (1885 : 128) SYNTYPES : one female ; one juvenile. L. Lund. Budde-Lund Collection. imbutus Budde-Lund (1885 : 145) SYNTYPES : not separated from other (non-type) specimens. 4436 (part). Sicily. Budde-Lund Collection. imbutus pellegrinensis Verhoeff (igoSa : 366) SYNTYPE : male. Reg. no. 1921:6:10:32. Sicily. Purchased from K. W. Verhoeff. incanus Budde-Lund (1885 : 102) SYNTYPES : two females. Reg. no. 1921:10:18:4437-4438. Meinert. Budde-Lund Collection. intercalarius Budde-Lund (1885 : 104) SYNTYPE : female. Reg. no. 1921:10:18:4439. Budde-Lund Collection. interpolator Budde-Lund (1885 : 93) SYNTYPES : specimens coated with mould. Budde-Lund Collection. jelskii Miers (iSyyb : 668) [Transferred to Metoponorthus pruinosus Brandt] SYNTYPES: one male; two females. Reg. no. 1879:21. Maraynioe (?), Guiana, sented by Prof. Wrzesniowsky. SYNTYPES: three males; three females. Reg. no. 1879:21. Pumamarca, Peru, sented by Prof. Wrzesniowsky. SYNTYPES : nine males ; ten females. Reg. no. 1887:20. Caracas, Guiana. Presented by Dr Ernst. latissimus Budde-Lund (1885 : 95) [Transferred to Porcellio spatulatus (Costa)] SYNTYPES: three males; five females. Reg. no. 1921:10:18:4988-4992. Corsica (collected by E. Simon) and Sardinia (collected by Keitel). Budde-Lund Collection. lepineyi Verhoeff (i937b : 305) SYNTYPES: one male; three females. Reg. no. 1938:7:7:7-10. Haute Atlas, Morocco. Collected by J. de Le'piney. Purchased from K. W. Verhoeff. longicauda Budde-Lund (1885 : 112) SYNTYPE : male. Reg. no. 1911:11:8:10358. Algeria. Collected by Fr. Meinert. Norman Collection. SYNTYPES: two females. Reg. no. 1911:11:8:10359-10360. Algeria. Collected by E. Simon. Norman Collection. SYNTYPES : one male ; five females. Lund Collection. SYNTYPES : one male ; one female, by University College, Dundee. longipennis Budde-Lund (1885 : 91) SYNTYPES : two females. Reg. no. 1921:10:18:4475-4476. Collected by Nordmann. Budde-Lund Collection. maculipes Budde-Lund (1885 : 105) SYNTYPES: two males; two females. Reg. no. 1921:10:18:4488-4491. Collected by Sorensen. Budde-Lund Collection