PROCEEDINGS OF THE California Academy of Sciences Volume 43 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1982-1984 PUBLICATIONS COMMITTEE Sheridan Warrick, Editor Frank Almeda Daphne G. Fautin Tomio Iwamoto Frank H. Talbot (US ISSN 0068-547X) The California Academy of Sciences Golden Gate Park San Francisco, California 94 1 1 8 PRINTED IN THE UNITED STATES OF AMERICA BY ALLEN PRESS, INC., LAWRENCE, KANSAS INDEX TO VOLUME 43 (Compiled by Lillian J. Dempster) New names in boldface type Acanthocybium solanderi 30 1 Acarina 14 Acestrorhynchus 206 Acridium coloratum 49-50 xanthopterum 44, 5 1 Ahlia 60 egmontis 64 Albula chinensis 210 lAlbula chinensis 210 Alloeocarpa 239 sp. 239 Allopora 245 Alopias superciliosus 89 vulpinus 89 Alopiidae 94-96 Alytes obstetricans 217 Amazonsprattus 3 1 7-32 1 scintilla 317-321 Ambassis 314 ^«c/20v/e//a317, 321 Andricus kollari 132 Anthias macrophthalmus 302 ^;7/m3' 126 figulus 126 Aplidium californicum 244 Aplochitonidae 203, 215 Arctocephalus doriferus 229-230 Argentinoidei 20 1 Ascidiacea 239 Atherinops 314 Atractoscion nobilis 232 Aulopodidae 201 Balanus glandula 244 nubilis 244-245 Balitora 68 Baltimora recta 49 5an7/M5 151, 156-157 Bdellostoma cirrhatum 264 dombeyi 255 Forsteri 264 heptatrema 250, 264 Benthenchelys 60 cartieri 64 Berycidae 3 1 1 5^/7x311 5/aA^a 269-270, 281-282 elliptica 270-27 1 tuberculata 270, 281-282 Blakeae 269, 282 Bosminiopsis deitersi 320 Brycon 142 Calophysoides 275 Calyptrella 275 Carabidae 159 Carchariniformes 93 Carcharodon carcharias 89, 107, 221-238 Carinotetraodon 15 lorentziana 49 parqui 49 strigillatum 49 Cetengraulis juruensis 317 Cetorhinidae 94-96 Cetorhinus 95-96, 109-110 maximus 96, 232 Chaenogobius sp. 181 Chaetodon 3 1 1 striatus 3 1 1 substriatus 3 1 1 Characoidei 201 C/ze/a 144 Chelonodon 2 Chlorophthalmidae 201 Chonerhinos 1-16 africanus 1, 4, 7-10 amabilis 1, 3-7, 11-15 modestus 1-15 naritus 1-2, 5 nefastus 1, 3-5, 9-11, 13-16 remotus 1, 3-5, 9, 11-15 silus 1,3-5, 9, 11-16 Chonerhinus 2 africanus 7 modestus 5 naritus 5 Chromacrini 43 Chromacris 43-58 color ata 44-51, 54 colorata group 45, 47, 51 colorata- miles group 50 /c?erM5 44, 46-48, 50, 54-56 m//e5 44-48, 51, 55 minuta 43-47, 5 1 nuptialis 44-48, 53 [323] 324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43 peruviana 44, 46-47, 49-50, 54, 56 psittacus 46-48, 50, 54 psittacus pacificus 43-44, 47, 54-55 psittacus psittacus AA-A5, 54-55 speciosa 43-53 trogon 46-50, 54-55 trogon group 45, 47, 50-51, 54 trogon intermedia 43-45, 54 trogon trogon 44-45, 53-54 Chthamalus dalli 244 aadocera317, 319 Clavilithes macrospira 314 Clidemia 269, 27 1,274 petiolaris 275 saltuensis 27 A tetrapetala 269, 271-274 trichosantha 269, 273-275 Clupeidae218, 317, 321 Cnemidocarpa 247 Coleoptera 159 Colomesus 15 Compositae 49 Cookeolus 302, 304-305, 308, 312, 314 Zjoo;75 301-309, 312,314 Cynoscion nobilis 232 Cyprinidae 67, 142-143, 148, 151-152, 155-157 Cyprinus balitora 67, 74 sucatio 67, 69 Dallia pectoralis 199-200 Dalophis riipelliae 20-2 1 Danio 151 Dasyatidae 283-300 Dasyatis 284-289, 292, 295, 298 americana 285 centroura 285-286, 295, 299 garouaensis 283-284, 286, 289-290, 292-295, 298-300 margarita 283-284, 286-290, 292-295, 298-299 margaritella 283-284, 286-295, 298-299 marmorata 285 pastinaca 285-286 rudis 285 sp. 287, 292 ujo 286 ukpam 283-289, 294-300 violacea 285 Dendrodoa 239, 246-247 abbotti 239-248 carnea 239, 245-247 grossularia 245-247 grossularia-carnea-A\i\ioXt\ series 247 (Styelopsis) 239, 246-247 (Styelopsis) abbotti 239-248 uniplicata 246-247 Diplomystes 154-155 Diptera317 Drimys 277 Eleotrididae 32 1 Elopomorpha 2 1 8 Elops 194, 217-218 hawaiiensis 200, 2 1 8 Engraulididae 317, 321 Enhydra lutris 229, 231 Eperlanus 204 chinensis 204-205 Epiactis prolifera 244 Epigonus 314 Epinephelus spp. 30 1 Eptatretus 249-267 burgeri 249 carlhubbsi 249-267 cirrhatus 249-250, 254-261, 264-265 deani 250 heptatrema 264 hexatrema 249 laurahubbsi 249-267 polytrema 262 springeri 254 5?0Mm 250, 255, 262 strahani 249-267 undescribed 262 Eremopodes sp. 4 1 Esomus 148 Eugomphodus 94 tricuspidatus 94 Eurystomella bilabiata 244 Exoglossum 143 Fm^m niphobles 14 Galaxias 194 Galaxiidae 203, 214-216 Galaxioidea 203 Galaxioidei 215 Galeocerdo cuvier 89, 229 Galeomorphii 93 Galeorhinus zyopterus 232 Garra 75 Gastromyzon 75 Geochelone 111-121 abingdonii 1 12-1 13, 1 15, 1 19 decA:/ 112-113, 115, 119 chathamensis 112-113, 115, 119 chilensis 1 1 9 fifarwm/ 1 12-1 13, 1 15, 1 19 elephantopus 112-113 ephippium 111-113, 115-120 galapagoensis 1 1 3 ^m/?m 111-113, 115-120 hesterna 1 1 9 hoodensis 112-115, 119 macrophyes 1 1 3 INDEX microphyes 112-115, 119 nigra 1 1 3 nigrita 111-120 phantastica 112-115, 119 porteri 113-114 sp. 114 vandenburghi 112-115, 119 vicina 112-116, 119 vosmaeri 1 1 1 wallacei 112-114, 119 Ginglymostoma 109 Glaucosoma 3 1 4 Glenoglossa 59-66 wassi 59-66 Graffenrieda 269, 275 bella 269, 275-277 galeottii 275 gracilis 275 micrantha 275, 277 Gryllus locusta miles 5 1 speciosus 47, 51 Gymnomuraena marmorata 22 Gymnothorax 18, 20 enigmaticus 17-24 leucacme 17, 22 petelli 17,20,22 "petelir form 20-21 reticularis 20-2 1 rueppelliae 17, 19-22 rupelli 20 ruppeli 17, 20 ''ruppelir 21 5/g«//er 17, 21-22 undulatus 22 waialuae 17, 22 Hemisalanginae 206, 210 Hemisalanx 206, 210 prognathus 185, 206, 208, 210 Hemitrygon 2S1 ukpam 295 Hepsetus 194, 206 Heptatrema cirrata 264 Heptatretus banksii 250, 264 Heterodontiformes 93 Heteropriacanthus 301-315 cruentatus 301, 304-305, 307, 309-311 Hexanchus griseus 89 Himantura 284 5/^«(/e/- 290, 292, 294, 298 Homaloptera 75 *a«fon 250, 264 cirrhata 264 Hoplias 206 Hylaeus pec t oralis 1 36 Hymenoptera 27, 123 Hypolophus 284 Hypomesus 194 o//fifw5 2 1 5 ///5/za317 /^wr/z^ 110 brasiliensis 110 I sums spp. 233 Jenkinsia 3 1 7 Lactarius 314 Lamnidae 94-96 Lamniformes 89, 93 Leguminosae 49 Lepidogalaxias 194-195, 214-217 salamandroides 2 1 5-2 1 6 Leptasterias pusilla 244 Leucosoma 206, 210 chinensis 205, 210 reeve5/ 206, 210 Leuresthes 314 Lipara lucens 1 36 Lovettia 2 1 5 Luciobrama 152-153 Luciosoma 143 Lycium 49 cestroides 49 Lycodontis 20 Macrochirichthys 144, 151 Macrocystis pyrifera 232 Mallotus 215 Medicago saliva 49 Megachasma 87-1 10 pelagios87-110 Megachasmidae 87-1 10 Melastomataceae 269-282 Metasalanx 206 coreanus 206, 210 M/coma 269, 275, 277, 279 arboricola 269, 277-279, 281 cordata 279 mollicula 279 Miconieae 271, 277 Microdonophis fowleri 23 Microphilypnus 321 Mirounga angustirostris 229-230 Mitsukurina 105, 107 owstoni 94 Mitsukurinidae 94, 96 Mixodigma leptaleum 1 10 Mixodigmatidae 1 1 Monotreta 2, 1 5 interrupt a 17, 22 umbrofasciata 17, 21 xanthopterus 22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43 Muraenichthys 60, 62 Muraenidae 18 Mycteroperca spp. 301 Myctophidae 200 Myliobatis californica 230-232 Myrophinae 59-60 Myrophis 60 frio 65 Myxine 251-252, 254 glutinosa 255 Nebria 159-177 acuta acuta 167 acuta sonorae 159, 167 altisierrae 159-161, 169, 171, 176 arkansana 167 arkansana arkansana 167 arkansana edwardsi 167 arkansana fragilis 159, 167 arkansana uinta 159, 167 calva 159, 164, 170-171, 173, 177 campbelli 159, 161, 169, 171, 176 crassicornis 166-167 crassicornis intermedia 159, 166 fragilis 159, 167 fragilis fragilis 1 6 7 fragilis teewinot 159, 167 g^Wm albimontis 159, 163, 169, 171, 173, 177 haida 159, 162, 169, 176 intermedia 159, 166-167 jeffreyi 159, 162, 169, 176 labontei 159, 163-164, 170, 177 louiseae 159, 162-163, 169, 176 piute 167-168 piute piute 159, 167 piute se\ier\ 159, 164-165, 168, 170, 172, 174, 177 piute utahensis 159, 168 sierrablancae 159, 164, 170-171, 174, 177 sonorae 159, 167 steensensis 159, 165, 170, 172, 175, 177 trifaria 167-168 trifaria catenata 1 64 trifaria coloradensis 165 m/ana pasquineli 159, 165-166, 170, 172, 175, 177 trifaria piute 159, 167-168 trifaria tetonensis 159, 167 trifaria trifaria 159, 164-165, 167 trifaria utahensis 159, 168 utahensis 168 virescens 160-161 wallowae 159, 161-162, 169, 171, 173, 176 Nebriini 159-177 Neenchelys 59-60, 62-64 buitendijki 64-65 daedalus 59-66 microtretus 62, 64-65 parvipectoralis 64 spp. 64 Nemamyxine 251-254 Neomysis sp. 181 Neomyxine 251, 254 NeosalanxXn, 184, 186, 198,201-205,210-212,214, 219 andersoni\19, 181-182, 184, 186,209,211-212, 219 brevirostris 1 79, 1 8 1 , 1 86-1 87, 209, 2 1 1-2 1 2, 2 1 9 hubbsi 212 jordani \79-lSl, 183-186, 190-192, 195, 197, 199, 201-202, 204, 209-212, 219 regani 2 1 2 reganius 179, 181, 186, 202, 212 tangkahkeii taihuensis 212, 219 Neoscopelidae 200 Neomacheilus 72 Notomyxine 254 Notoplana acticola 244 Novumbra hubbsi 217 Odontaspididae 94, 96 Odontaspis 94, 96 Ophichthidae 23, 59-60 Ophichthinae 60 Ophichthus erabo 17, 23-24 garretti 23 retifer 17, 23 Ophiodon elongatus 231-232 Opsariichthys 154-155 Orectolobiformes 93 Orthoptera 43 Osmeridae 190, 199-201, 203, 215-216 Osmeroidea 179, 203 Ostariophysi 180-181,317 Osteochilus 154 Otolithus (Pagellusl) gregarius 3 1 3 (Sparidarum) gregarius 3 1 3 (Sparidarum) rutoti 3 1 3 Oxygaster 144 Pamphorichthys 321 Pangasius 13 polyuranodon 13-14 Paramyxine 251, 253, 255 Paraprotosalanx 204-205, 211 andersoni 205 Parasalanx 202, 206-207, 210 acuticeps 206, 208 angusticeps 207-208, 210 annitae 206-208 ariakensis 210 cantonensis 207-208, 210 gracillimus 206-208, 210 INDEX 327 longianalis 206-208 (Salanx) 202 IParasalanx gracillimus 207 Parexoglossum 143 PellonaZll Petrolisthes 244 Petromyzon cirrhatus 264 Phoca vitulina 236 Pisces 203, 301 Platycara 68 Plecoglossidae 200, 203, 215 Plecoglossus 203, 215 Plecoptera 14 Plectognathi 14 Podocarpus 277 Poecilia 321 Poeciliidae 321 Potamotrygon 283 garouaensis 292 Potamotrygonidae 283 Priacanthidae 301-315 Priacanthus 301-302, 310, 312-313 alalaua 301-303, 305, 307-310 arenatus 301, 310 cruentatus 301-302, 304, 309-310 hamrur 301, 310 macracanthus 301, 310 meefc/301, 310 tayenus 30\, 310 Prionace glauca 228 Pristigaster cayana 3 1 7 Pristigenys 301, 304, 311-314 bella30\, 310, 314 caduca 3\0, 313 dentifer 301, 310, 314 macrophthalmus 3 1 1 rM?o?/310, 312-313 spectabilis 310, 314 spp. 314 substriata 301, 304, 310-312, 314 substriatus 3 1 1 Protosalanginae 179, 181, 185, 189-190, 192, 194, 198, 202,204,206,211 Protosalanx 179, 182-185, 189, 192, 194, 198-199, 202,204-205, 211,214,216,218 andersoni 205, 209, 211 brevirostralis 2 1 2 brevirostris 209, 211-212 chinensis 179-181, 183-185, 187, 192-193, 196, 199, 201-202, 205, 208, 210-211, 214 hyalocranius 205, 211-212 tangkahkeii 2 1 1 IProtosalanx tangkahkeii 212 Prototroctes 203 Prototroctidae 20 1 Pseudocarcharias 105 kamoharai 94 Pseudocarchariidae 94, 96 Pseudomyrophis 59-60, 64-65 atlanticus 65 micropinna 65 nimius 65 spp. 64-65 Pseudopriacanthidae 3 1 3 Pseudopriacanthus 302, 304, 311-313 alius 3\2 niphonius 3 1 2-3 1 3 serrula 301-305, 307, 309, 312 Psilorhynchidae 67 Psilorhynchus 67-70, 73, 75 balitora 67-76 balitora-\\kt species 68 gracilis 67-76 homaloptera 67-69, 72, 75 homaloptera rowleyi 67 pseudecheneis 67-69, 72 sucatio 67-70, 72-75 sucatio var. damodarai 13-1 A variegatus 68, 74 Quercus spp. 277 Raja pastinaca 286 Rasbora 151 Reganisalanx 206 normani 206-20S, 210 "^Reganisalanx brachyrostralis 210 Retropinnidae 201, 203, 215 Rhiniodon 109-110 rypM5 89, 96 Rhinosardinia amazonica 3 1 7 /cren^ 55 latipennis 44, 53 m/te 52 miles Var. C 52 nuptialis 53 opulenta 44, 55-56 ;7ed/e5 44, 49-50 peruviana 56 speciosa 52 5/0/// 44, 51 trogon 53 Romalea psittacus 54 Romaleidae 43 Romaleinae 43, 47 i?M/?M5 136 Sagraea 21 A Salangichthyinae 179, 181-182, 186, 189-190, 192, 194-195, 198,202,204, 211 Salangichthys 184, 186, 202-203, 211-213 ishikawae 179-181, 185-186, 199, 204, 209, 213 kishinouyei 2 1 3 microdon 179, 181-187, 201-202, 204, 209, 212- 215,218 328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43 Salangidae 179, 181-183, 185, 194-195, 198-206,210- 211,213-218 Salanginae 179, 181-182, 185, 189-190, 192-195, 198, 204-206,210,218 Salangoidea 179-200 Salanx 179, 184, 201-204, 206-207, 210-211 acuticeps 206 argentea 2 1 1 ariakensis 181, 185-186, 206-208, 210, 213 brachyrostralis 210 chinensis 181, 187, 204-205, 210, 216 cuvieri 180-182, 186, 188, 192, 194, 197, 199, 202, 206-208, 210 {Hemisalanx) 186, 210 {Hemisalanx) prognathus 179, 210 hyalocranius 204-205, 208, 212 (Leucosoma) 186, 198, 210 (Leucosoma) reevesi 179, 198 (Leucosoma) reevesii 210 prognathus 181, 185-186, 189, 195, 208, 210 reevesi 179, 181, 186, 208, 210 reevesii 2 1 (Salanx) 186,202,207,210 (Salanx) ariakensis 179, 206 (Salanx) cuvieri 179, 207 ?Salanx argentea 212 brachyrostralis 210 cuvieri 206 Salmacina 244 Salmo 194 Salmonidae 200-201, 203, 214 Salmoniformes 179, 181, 195, 199-200, 214-215, 218 Salmonoidea 179, 203 Salmonoidei 203 Salmostoma 144, 151 Sardina pilchardus 200 Sardinops sagax 232 Schismorhynchus 60, 62 Schultzidia 60 Sciaena hamrur 302 Scorpaenichthys marmoratus 231-232, 234 Securicula 151 Seriola lalandi 30 1 Sideria chlevastes 1 7, 22 Siluriformes 20 1 Solanaceae 49 Solanum 49 argentinum 49 elaeagnifolium 49 verbascifolium 49 Somniosus microcephalus 89, 105 pacificus S9, 105 (Somniosus) 105 Sparidarum rutoti 3 1 2 Sphecidae27, 123 Sphex figulus 125-126 fuliginosa 126 fuliginosus 123, 126-127 Sphyrna mokarran 89 Spirinchus 194 Spirobis 244 Squalus 228 aca«?/z/a5 231-232 Stereolepis gigas 30 1 5/ye/a 246-247 uniplicata 247 Styelopsis 246 Sundasalangidae 179, 181-182, 184-185, 189-190, 192, 194-195, 199, 201-202, 204, 211, 213, 218 Sundasalanx 179, 181, 185, 192-193, 198, 201, 204, 213-214,218 microps 170-181, 186, 191-192, 196, 198, 200- 202,209,213-214 praecox 179, 181, 186, 193, 196, 209, 213-214 Synodontidae 201 Tachysphex ll-'M acanthophorus 27, 29, 39-41 alayoi 29 apricus27, 29-31, 34 arizonac 27, 29-30, 32, 35 arinatus27, 29,40-41 ashmeadii 3 1 belfragei 36, 41 bohartorum 27, 29-31 brevicornis 29 brulli group 28-29, 36, 39-40 crenulatus 35-36 fulvit arsis 29 glabrior 31,36 idiotrichus 27, 29, 31, 34, 36 irregularis 27, 29, 31-32, 34, 36 julliani group 28-29 krombeini 34 krombeiniellus 27, 29, 41 lamellatus 27, 29-30, 32, 35-37 maurus 4 1 inenkei27, 29,41-42 mirandus 27, 29, 32-34 mundus 29, 36, 40-41 musciventris 27, 29, 33-35 occidentalis 27, 29, 34-35 papago 27, 29, 35 pechumani 30-3 1 pompiliformis group 28-29, 31, 36 psilocerus 30, 35 semirufus 33-34 Solaris 27, 29, 35 sonorensis 32, 37 spatulifer 27, 29, 35-36 spinulosus 40 tarsatus 34 terminal us group 28-29 texanus 30 undescribed species 33 verticalis 27, 29, 31, 36-37 yolo 27, 29, 37-39 yuma 27, 29, 39 Taeniura 284 grabata 285 Tetraodon 2, 1 5 modest us 2 Tetraodon {Arothron) modestus 7-8 modestus 2 naritus 2 Tetraodontidae 1, 14-15 Tetraodontiformes 14 Thryssocypris 141-158 smaragdinus 141-158 tonlesapensis 141-158 Thunniis albacares 30 1 obesus 301 Thysanopoda pectinata 108 Topobea 269-270, 281-282 brenesii IIQ calophylla 269, 280-282 durandiana 282 elliptica 269-270, 281 pittieri 2Sl Trichoptera 14 Trygon 287, 295 margarita 287, 290 ukpam 295 Trypanorhynchida 110 Trypoxylon 125-126 apicale 126-127 apicalis 126 fieuzeti 127, 131 figulus 123-140 figulus barbarum 123, 127, 131 figulus koma 123, 132 figulus major 126 figulus forma major 1 26 figulus var. mayor 136 figulus -var. majus 126 figulus media 1 36 figulus forma me^/a 136 figulus var. me^/a 136 figulus medium 132, 136 figulus var. medium 136 figulus minor 132 figulus forma m/>?or 132 figulus var. minor 132 figulus minus 132, 136 figulus var. m/«u5 132 figulus minus var. rwZ)/ 1 36 figulus yezo 123, 127 ma/M5 123, 127 medium 123-140 w/'nM5 123-140 rwi/ 123, 139 Umbridae217 Urogymnus 283-284, 286-287, 295 africanus 285 asperrimus 285 Urolophoides 284 Uropterygius 22 alboguttatus 17, 22 kamar 22 marmoratus 17, 22-23 xanthopterus 17, 22 Verbesina encelioides 49 Xenopterus 1-3, 9 bellengeri 2 naritus 2-3 Xestotrachelus 43-58 hasemani 44, 56-57 robustus 44-46, 48, 50, 55-57 Zalophus californianus 77-85, 229-230 Zoniopoda robusta 56 PROCEEDINGS, OCCASIONAL PAPERS, AND MEMOIRS of the CALIFORNIA ACADEMY OF SCIENCES INSTRUCTIONS TO AUTHORS Publications The California Academy of Sciences publishes at irregular intervals three series containing re- sults of original scientific research. 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Leaf-cutter ants of Oregon. Amer. Bull. Entomol. 43:1-8. Smith, A. T., and L. O. Lee. 1969. Social or- ganization of hymenopterans. Ecology 1 14:3 2- 49. Edited books: White, L. M. Q., and J. T. Brown, eds. 1974. The biota of Gondwanaland. Harper and Row, Chicago, Illinois. 640 pp. Articles/chapters in edited books: Johnson, L. B., D. F. Black, and R. M. Hobbs. 1956. The acacias. Pp. 1093-1235 in Aus- tralian flora, I. Q. Doyle and M. Henry, eds. Kangaroo Press, Sydney, New South Wales, Australia. Dissertation or thesis: Green, J. P. 1960. Osmoregulation in sipun- culids. Ph.D. Dissertation, University of Wy- oming, Laramie, Wyoming. 122 pp. For further guidance, consult the Editor, Sci- entific Publications, California Academy of Sci- ences. CONTENTS OF VOLUME 43 Pages No. 1. Roberts, Tyson R. The Southeast Asian freshwater pufferfish genus Chone- rhinos (Tetraodontidae), with descriptions of new species. PubUshed June 15, 1982 1-16 No. 2. McCosKER, John E., and John E. Randall. Synonymies of Indian Ocean eels, with the description of Gymnothorax enigmaticus, a moray previously known as G. Ruppeli. Published June 15, 1982 17-24 No. 3. PuLAwsKi, WojciECH J. Ncw species of North American Tachysphex wasps (Hymenoptera, Sphecidae). Published June 15, 1982 27-42 No. 4. Roberts, H. Radclyffe, and Carlos S. Carbonell. A revision of the grass- hopper genera Chromachs and Xestotrachelus (prihoptera, Romaleidae, Roma- leinae). Published November 4, 1982 43-58 No. 5. McCosKER, John E. A new genus and two new species of remarkable Pacific worm eels (Ophichthidae, subfamily Myrophinae). Published November 4, 1982 59-66 No. 6. Rainboth, Walter J. Psilorhynchus gracilis, a new cyprinoid fish from the Gangetic lowlands. Published July 6, 1983 67-76 No . 7 . Le Boeuf, Burney J . , David Aurioles, Richard Condit, Claudio Fox, Robert GisiNER, Rigoberto Romero, and Francisco Sinsel. Size and distribution of the California sea lion population in Mexico. Published July 6, 1983 77-85 No. 8. Taylor, Leighton R., L. J. V. Compagno, and Paul J. Struhsaker. Megamouth— a new species, genus, and family of lamnoid shark {Megachasma pelagios, family Megachasmidae) from the Hawaiian Islands. Published July 6, 1983 87-110 No. 9. Crumly, Charles R. The cranial morphometry of Galapagos tortoises. Pub- lished January 17,1 984 111-121 No. 10. PuLAwsKi, WojciECH J. The status of Trypoxylon figulus (Linnaeus, 1758), medium De Beaumont, 1945, and minus De Beaumont, 1945 (Hymenoptera: Sphecidae). Published January 17, 1984 123-140 No. 11. Roberts, Tyson R., and Maurice Kottelat. Description and osteology of Thryssocypris, a new genus of anchovylike cyprinid fishes, based on two new species from Southeast Asia. Published January 17, 1984 141-158 No. 12. Kavanaugh, David H. Studies on Nebriini (Coleoptera: Carabidae), V. New Nearctic Nebria taxa and changes in nomenclature. Published July 12, 1984... 159-177 No. 13. Roberts, Tyson R. Skeletal anatomy and classification of the neotenic Asian salmoniform superfamily Salangoidea (icefishes or noodlefishes). Published July 12, 1984 179-220 No. 1 4. Tricas, Timothy C, and John E. McCosker. Predatory behavior of the white shark {Carcharodon carcharias), with notes on its biology. Published July 12, 1984 221-238 No. 1 5. Newberry, Andrew Todd. Dendrodoa (Styelopsis) abbotti, sp. no v. (Styelidae, Ascidiacea) from the Pacific Coast of the United States, and its impact on some gonadal criteria of its genus and subgenus. Published September 19, 1984 239-248 [iii] Pages No. 16. McMillan, Charmion B., and Robert L. Wisner. Three new species of seven- gilled hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean. Published De- cember 11, 1984 249-267 No. 17. Almeda, Frank. New and noteworthy additions to the Melastomataceae of Panama. Published December 11, 1984 269-282 No. 18. CoMPAGNO, Leonard J. V., and Tyson R. Roberts. Marine and freshwater stingrays (Dasyatidae) of West Africa, with description of a new species. Pub- lished December 11,1984 283-30a No. 19. Fitch, John E., and Stephen J. Crooke. Revision of Eastern Pacific catalufas (Pisces: Priacanthidae) with description of a new genus and discussion of the fossil record. Published December 11, 1 984 30 1-3 1 5 No. 20. Roberts, Tyson R. Amazonsprattus scintilla, new genus and species from the Rio Negro, Brazil, the smallest known clupeomorph fish. Published December 1 1 , 1 984 3 1 7-32 1 Index to Volume 43 323-329 Instructions to Authors 331-332 [iv] PROCEEDINGS OF THE CALIFORNIA ACADEMY jOF SGIBNjQES Vol. 43, No. 1, pp. 1-16, 10 figs. lui- 'J 198;* PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2 Figure 7. Adult Ophichthus erabo, from Jordan and Snyder (1901). curators and staffs of many museums for allow- ing us to examine specimens under their care. Randall's collections were made possible in part by grants from the National Geographic Society and the American Philosophical Society. A por- tion of McCosker's work was supported by funds from the Charline Breeden Foundation. Literature Cited Bleeker, p. 1859. Over eenige vischsoorten van de Zuid- kustwateren van Java. Nat. Tijdschr. Neder. -Indie 19:329- 352. . 1864. Atlas ichthyologique des Indes Orientales Neer- landaises. Vol. 4. Amsterdam. 132 p. Bliss, R. 1883. Descriptions of new species of Mauritian fish- es. Trans. Soc. Roy. Arts Sci., Maurice 13:45-63. Bloch, M. E. 1795. Naturgeschichte der Auslandischen Fische. Vol. 9. Chave, E. H., andH. A. Randall. 1971. Feeding behavior of the moray eel, Gymnothorax pictus. Copeia 1971 (3):570-574. Fowler, H. W. 1935. South African fishes received from Mr. H. W. Bell-Marley in 1935. Proc. Acad. Nat. Sci., Phila- delphia 87:361-408. . 1956. Fishes of the Red Sea and southern Arabia. 1 . Branchiostomida to Polynemidae. Weizmann Sci. Press, Jerusalem. 240 p. GosLiNE, W. A. 1958. Central Pacific eels of the genus Urop- terygius, with the descriptions of two new species. Pac. Sci. 12(3):22 1-228. GiJNTHER, A. 1910. Andrew Garretfs Fische der Siidsee, . . . Heft IX. J. Mus. Godeffroy, Hamburg 17:389-515. HoBSON, E. S. 1974. Feeding relationships of teleostean fish- es on coral reefs in Kona, Hawaii. Fish Bull.. U.S. 72(4):91 5-1031. Jenkins, O. P. 1904. Report on collections of fishes made in the Hawaiian Islands, with descriptions of new species. U.S. Bur. Fish. Fish. Bull. 22:417-511. Jordan, D. S., and C. H. Gilbert. 1883. Description of a new muraenoid eel from the Galapagos Islands. Proc. U.S. Natl. Mus. 6:208-210. , and J. O. Snyder. 1901. A review of the apodal fishes or eels of Japan, with descriptions of 19 new species. Proc. U.S. Natl. Mus. 23:837-890. Kaup, J. 1856. Uebersicht der Aale. Arch. Naturges. 22(l):41-77. Klunzinger, C. B. 1871. Synopsis der Fische des Rothen Meeres. II. Theil. Verh. Zool.-Bot. Ges. Wien 21:441-668. McClelland, J. 1845. Apodalfishesof Bengal. J. Nat. Hist. Calcutta 5:150-226. McCosKER, J. E. 1979. The snake eels (Pisces, Ophichthidae) of the Hawaiian Islands, with the descriptions of two new species. Proc. Calif. Acad. Sci., ser. 4, 42(2):57-67. , and J. E. Randall. 1977. Three new species of Indo-Pacific moray eels (Pisces: Muraenidae). Proc. Calif. Acad. Sci., ser. 4. 41(3): 161-168. , AND R. H. Rosenblatt. 1975. The moray eels (Pisces: Muraenidae) of the Galapagos Islands, with new records and synonymies of extralimital species. Proc. Calif. Acad. Sci., ser. 4, 40(13):417-427. Randall, J. E. 1973. Tahitian fish names and a preliminary checklist of the fishes of the Society Islands. Occ. Pap. Bernice P. Bishop Mus. 24(1 1): 167-2 14. RiJppELL, W. P. E. S. 1852. Verzeichniss der in dem Museum der Senckenbergischen . . . Fische und deren Skelette. Frankfurt-a-M. ScHULTZ, L. P., AND COLLABORATORS. 1953. Fishes of the Marshall and Marianas islands. Families from Asymme- trontidae through Siganidae. U.S. Natl. Mus. Bull. 202, 1. 685 p. Smith, J. L. B. 1962. The moray eels of the western Indian Ocean and the Red Sea. Ichthyol. Bull. Rhodes Univ. 23:421-444. Snyder, J. O. 1904. A catalogue of the shore fishes collected by the steamer "Albatross" about the Hawaiian Islands in 1902. U.S. Bur. Fish. Fish. Bull. 22:513-538. Weber, M., and L. F. de Beaufort. 1916. The fishes of the I ndo- Australian Archipelago. Vol. 3. Leiden. 455 p. PROCEEDINGS- CALIFORNIA ACADEMY OF SCIENCES 1986 Vol. 43, No. 3, pp. 27-42 Uu J June 15, 1982 NEW SPECIES OF NORTH AMERICAN TACHYSPHEX WASPS (HYMENOPTERA, SPHECIDAE) By Wojciech J. Pulawskl California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: The following new North American species of Tachysphex are described (type-localities are given in parentheses): acanthophorus (Arizona: Willcox), apricus (California: Borrego Valley), arizonac (Arizona: 2 mi. NE Portal), armatus (Nevada: Sandy), bohartorum (California: Boca), idiotrichus (Arizona: 5 mi. W Portal), irregularis (California: Hallelujah ivinc^on), krombeiniellus (Florida: Levy County), lamellatus (Mexico: Sonora: Alamos), menkei (California: Borrego Valley), mirandus (California: Palm Springs), musciventris (California: Borrego), occidentalis (California: 12 mi. E Lone Pine), papago (Arizona: Nogales), Solaris (California: Borrego Valley), spatulifer (California: Arroyo Seco Camp), verricalis (California: 9 mi. W Beaumont), yuma (Mexico: Baja California: La Paz), and yolo (California: Davis). Introduction For several years, I have been working on a monographic revision of North American Tachysphex. Because of the size of this under- taking, it will be some time before it is finished. Therefore, I am describing some of the new species now so their names will be available to those persons working on Tachysphex behavior. Furthermore, many hundreds of paratypes have been deposited in 34 collections in the USA and abroad, and it is desirable to validate these manuscript names now to avoid their possible use as nomina nuda in the works of others. The descriptions given below are restricted to those features which enable unambiguous recognition of each species. More complete characteriza- tions will be given when my revision is pub- lished. The terminology used below is based mainly on Bohart and Menke (1976). A few terms which need clarifications are the following: clypeus: the clypeus has a midsection and two lateral sections; the midsection usually has a densely punctate, setose basomedian area, a sparsely punctate shiny bevel, and a marginal lip. scutum: this term is used here for brevity's sake instead of mesoscutum. tergum, sternum: short terms for gastral tergum, gastral sternum. Many collectors are cited numerous times in the lists of material examined. Their names have been abbreviated to initials, as follows: ASM, A. S. Menke: BV, B. Villegas: DRM. D. R. Miller: EEC. E. E. Grissell: EIS, E. I. Schlin- ger: GEB, G. E. Bohart: GDB, G. D. Butler; FDP, F. D. Parker; FGW, F. G. Werner: FXW, F. X. Williams: HKC, H. K. Court; JCH, J. C. Hall; JAP, J. A. Powell; JMD, J. M. Davidson; JWMS, J. W. MacSwain; LAS, L. A. Stange; MAC, M. A. Cazier; MEI, M. E. Irwin: MSW, [27] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 M. S. Wasbauer; PDH, P. D. Hurd; PMM, P. M. Marsh; PFT. P. F. Torchio; RCB, R. C. Bechtel; RMB, R. M. Bohart; ROS, R. O. Schuster; RRD. R. R. Dreisbach; TG, Terry Griswold; WJP, W. J. Pulawski. In the geographic names below, the following words have been abbreviated: County, Co.; Creek, Cr.; Highway, Hwy.; Island, I.; miles, mi.; Mountain(s), Mt(s).; River, R.; Station, Sta. The name Lower California has been used for the peninsula rather than Baja California, be- cause the latter may refer either to the peninsula or to a state in Mexico. Altitudes and distances are given as they appear on the original labels — in feet and miles. Multiplying the distances in miles by 1.609 and the elevations in feet by 0.3048 will convert them into kilometers and meters, respectively. Sources of Material and Acknowledgments The specimens described in this paper came from institutional and private collections listed below. The initials preceding the names are the abbreviations by which institutions or private collections are referred to in the text. AMNH: American Museum of Natural History, New York. New York (M. Favreau) ANSP: Academy of Natural Sciences of Philadelphia, Phila- delphia, Pennsylvania (D. C. Rentz) ASU: Arizona State University, Department of Zoology, Tempe, Arizona (F. F. Hasbrouck) BMNH: British Museum (Natural History), London, England (C. R. Vardy) CAS: California Academy of Sciences, San Francisco, Cali- fornia (P. H. Arnaud. Jr., T. J. Zavortink, W. J. Pulawski) CIS: California Insect Survey, Division of Entomology, Uni- versity of California, Berkeley, California (H. Daly) CNC: Canadian National Collections of Insects, Arachnids and Nematodes, Biosystematics Research Institute, Otta- wa, Ontario (J. R. Barron) CSDA: California Department of Food and Agriculture. Sac- ramento, California (M. S. Wasbauer) CSU: Colorado State University, Department of Zoology and Entomology, Fort Collins, Colorado (H. E. Evans) CU: Cornell University, Department of Entomology and Lim- nology, Ithaca, New York (L. L. Pechuman) FSCA: Florida State Collections of Arthropods, Gainesville, Florida (E. E. Grissell) HKT: H. K. Townes, American Entomological Institute, Ann Arbor, Michigan INHS: Illinois State Natural History Survey, Urbana, Illinois (W. E. LaBerge) KU: University of Kansas, Snow Entomological Museum. Lawrence, Kansas (G. W. Byers) KVK: K. V. Krombein, Arlington, Virginia (private collec- tion), now in USNM LACM: Natural History Museum of Los Angeles County, Los Angeles, California (R. R. Snelling) MCZ: Museum of Comparative Zoology at Harvard Univer- sity. Cambridge, Massachusetts (J. Lawrence, J. C. Scott, M. K. Thayer) MPM: Milwaukee Public Museum, Milwaukee, Wisconsin (J. K. Lawton) NYSU: New York State University. College of Environmen- tal Sciences and Forestry, Department of Environmental and Forest Biology, Syracuse, New York (F. E. Kurczews- ki) OSDA: State of Oregon Department of Agriculture, Salem, Oregon (R. L. Westcott) OSU: Oregon State University. Department of Entomology, Corvallis. Oregon (P. Oman. G. R. Ferguson) TG: Terry Griswold, % Bee Biology & Systematics Labora- tory, Utah State University, Logan, Utah (private collec- tion) UAE: University of Alberta, Department of Zoology, Ed- monton, Alberta (A. L. Steiner) UAT: University of Arizona, Department of Entomology, Tucson, Arizona (F. G. Werner) UCD: University of California, Davis, Department of Ento- mology, Davis, California (R. M. Bohart, R. O. Schuster) UCR: University of California, Riverside, Department of Bi- ological Control, Riverside, California (S. Frommer) UFG: University of Florida, Department of Entomology and Nematology, Gainesville, Florida (B. Saffer) UGA: University of Georgia. Department of Entomology, Athens, Georgia (R. W. Matthews. C. L. Smith) UIM: University of Idaho, Department of Entomology, Mos- cow, Idaho (W. F. Barr) UMSP: University of Minnesota. Department of Entomology and Zoology, St. Paul, Minnesota (P. J. Clausen) USNM: United States National Museum of Natural History (Smithsonian Institution), Washington, D.C. (A. S. Menke, K. V. Krombein) USU: Utah State University, Department of Zoology, Logan. Utah (G. E. Bohart, F. D. Parker, Terry Griswold) WJP: Wojciech J. Pulawski, % California Academy of Sci- ences, San Francisco, California (private collection) WSU: Washington State University, Department of Entomol- ogy, Pullman, Washington (M. T. James, R. Zack) I express my sincere thanks to the curators and other persons who kindly submitted speci- mens for study. I feel especially indebted to R. M. Bohart, A. S. Menke, K. V. Krombein, and F. F. Kurczewski who helped me in many ways. Species Groups Sixteen species groups are recognized in Tachysphex (sec Pu\awski 1971, 1974, 1977), but only four of them are represented in North America. They are: the pompiliformis , tenni- natiis, brullii, and juUiani groups. The species described in this paper belong to the pompili- formis and bntUii groups which are defined as follows: The pompiliformis group lacks peculiarities which characterize other groups and thus pos- sibly is a heterogenous assemblage of conve- nience. The propodeal hindface in this group is inclined, the female pygidial plate is not broad- NEW SPECIES OF NORTH AMERICAN TACHYSPHEX ened and without peculiar microsculpture. the preapical bristfes on the female gastral segments are not thickened, and the male sterna are pru- inose (except in niirandus). By comparison, in thejiilliani group the propodeal hindface is ver- tical or nearly so. male sterna are glabrous or sparsely pruinose. and in the females of most species the preapical bristles of gastral segments IV and V are thickened, and the pygidial plate is broadened or has a peculiar microsculpture. The vertex is simple in the pompiliformis group, while in the terminatus group a swelling is pres- ent behind each hindocellus. Unlike the bruUii group, the apical female tarsomeres are simple (see that group for details). The pompilifonnis group is cosmopolitan. Its species prey upon acridid nymphs, but the Palearctic species ////- vitarsis collects tettigonids. The following new species are members of the pompilifonnis group: apricus, arizonac, bohartoruin, idiotri- chiis, irregularis, lamellatus, mirandus, musci- ventris, occidentalis, papago, Solaris, spatuli- fer. verticalis, yolo, and yuma. The brullii group is characterized by the pe- culiar apical female tarsomeres: dorsum convex, apicoventral margin produced into a lobe or at least convex, and vertex variously modified (covered with erect setae except glabrous ba- sally, or angulale basally in lateral view, or densely spinose). In other groups the dorsum is scarcely convex, the apicoventral margin is straight or nearly so, and the venter is evenly covered with setae which are usually inclined (but erect in verticalis), and it may have a few spines in some species. Furthermore, the pro- podeal dorsum setae are erect or inclined back- wards in most species of the brullii group, but only laterally so in acanthophorus, alayoi, ar- matus, many individuals of mundus, and some Australian species. Setae are inclined obliquely cephalad in the Australian species brevicornis and in most species of other groups. The brullii group is widespread throughout all zoogeo- graphic regions. Some species prey upon tetti- gonids, while others are blattid collectors. The following new species are members of this group: acanthophorus, armatus, krombeiniel- lus, and menkei. Species of the Pompiliformis Group Tachysphex apricus sp.n. Etymology. — The specific name apricus is a Latin word meaning exposed to the sun. Diagnosis. — Tachysphex apricus differs from other species of the pompiliformis group by the setal pattern of its propodeal dorsum: median setae are inclined cephalad, but the lateral setae are directed obliquely backwards and join api- comesally. Some species of the brullii group (e.g., acanthophorus) have an identical pattern, but the unspecialized apical female tarsomere of apricus is distinctive. The male of apricus can be distinguished by the compressed femoral notch whose glabrous bottom forms an obtuse, longitudinal crest. T. idiotrichus has a similar crest, but unlike that species the body vestiture is short in apricus. Unlike most species of the pompiliformis group, the propodeal side of apri- cus is alutaceous, shiny, impunctate or minutely punctate. Geographic Distribution. — Xeric areas between southern Texas, southern Nevada, and southern California, and also Lower California. Material Examined. — Holotype: 6 . California, San Diego Co., Borrego Valley, 3 May 1956, P. D. Hurd (UCD). Paratypes: 38 9 , 60 eth:j ■ McNary H. E. Evans (19, CSU). Presidio: 3 ml e PreVldio. H. E. Evans (I 9, MCZ). • Literature Cited BoHART, R. M., AND A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. University of California Press, Berkeley, Los Angeles, London. 1 color pi., ix + 695 p. PuLAWSKi,. W.*J«. 1971. Les Tachysphex Kohl (Hym., Spheci- da^ d'ela region palearctique occidentale et centrale. Pari- stwowe Wydawnictwo Naukowe, Wroclaw. 464 p. . 1974. A revision of the Neotropical Tachysphex (Hym., Sphecidae). Polskie Pismo Ent. 44:3-80. . 1977. A synopsis of Tachysphex Kohl (Hym., Spheci- dae) of Australia and Oceania. Polskie Pismo Ent. 47:203- 332. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 43, No. 4, pp. 43-58, 27 figs. * s Novetnber 4, 1982 UBRAHY "4ory I •^OV 9, 2 198? A REVISION OF THE GRASSHOPPER GENERA CHROMACRIS AND XESTOTRACHELU^^^-^ — (ORTHOPTERA, ROMALEIDAE, ROMALEINAE) By H. Radclyffe Roberts Department of Entomology, Academy of Natural Sciences, 19th and the Parkway, Philadelphia, Pennsylvania 19103 and Carlos S. Carbonell* Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boavista, 20942 Rio de Janeiro, RJ, Brazil Abstract: Illustrations, diagnoses, and distributional records are given for 14 species and subspecies of the Neotropical grasshopper genera Chromacris and Xestotrachelus, of which 1 species, Chromacris minuta, and 2 subspecies, C. trogon intermedia and C. psittacus pacificus, are described as new, and 4 names are newly synonymized. Known information concerning food plants, oviposition, habitat, and predator defense for Chro- macris speciosa is briefly reviewed. (Orthoptera, Romaleidae, Romaleinae, grasshoppers. Neotropical, new taxa, taxonomic revision) Introduction treatment of the genus, a revisionary study ap- The grasshopper species of the genus Chro- peared to be needed for this common and wide- macris are of a striking color, usually a glossy spread group of grasshoppers. The monotypic green with yellow markings, and red or yellow genus Xestotrachelus of southern South Amer- wings. They occur in the humid areas of the ica is included in this study because of its sim- American tropics from Mexico to Argentina. As ilarity to Chromacris and because it is the only a general rule, but one species occurs at any one closely related genus. locality. Adults are usually seasonal in their ap- The subfamily Romaleinae currently includes pearance, so that some months of the year a about 48 genera. Rehn and Grant (1959) erected species may appear to be absent from a local 16 tribes in this subfamily and proposed the tribe fauna. Because most of the taxa of the genus Chromacrini for the genera Chromacris and have been poorly defined or understood, the ap- Xestotrachelus. Because 10 of their tribes in- plication of a number of their names has been elude but one or two genera each, and they give uncertain, and there has been no comprehensive no distinguishing characters for these tribes, it does not appear useful to recognize the tribe * Bolsista. Conselho Nacional de Desenvolvimento Cienti- Chromacrini and Other SUCh tribes of the fico e Tecnologico, Brazil. subfamily. [43] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 FiGUF Chromacris speciosa Chromacris speciosa, Casupa, Florida, Uruguay. The genus Chromacris includes at least eight species, two of which are polytypic, each having two subspecies, and two other species that we tentatively recognize pending further informa- tion. One species, C. minuta, and two subspe- cies, C. trogon intermedia and C. psittaciis pa- cificus, are described as new. There are five junior synonyms, three of which we newly syn- onymize. Specimens belonging to various collections are indicated by the following abbreviations: ANSP, Academy of Natural Sciences of Phila- delphia; CACS, Dr. C. A. Campos Seabra Col- lection; CSC, Carlos S. Carbonell Collection, Montevideo, Uruguay; CHFR, C. H. F. Rowell Collection, Zoologisches Institut der Universi- tat, Basel, Switzerland; FCZ, F. Carrasco Col- lection, Cuzco, Peru; MNHN, Museum Nation- al d'Histoire Naturelle, Paris, France; MZSP, Museu de Zoologia, Universidade Sao Paulo, Brazil; UMMZ, University of Michigan Mu- seum of Zoology, Ann Arbor, Michigan, U.S.A. Acknowledgments We are grateful to our many friends for their help and advice. We especially thank the follow- ing persons and their respective institutions for the loan of specimens. Dr. Irving J. Cantrall, UMMZ; Drs. F. Carrasco and J. A. Escalante, University of Cuzco, Peru; Dr. Marius Des- camps and Christiane Amedegnato, MNHN; Dr. Kurt K. Giinther, Museum fiir Naturkunde, Humboldt Universitat, Berlin, DDR; and the late Dr. H. Reichardt, MZSP. Species List In the following list of taxa we have attempted to place similar or related taxa as close to one another as practical. The number assigned to each taxon matches the number in the text. Ju- nior synonyms are given (in italics) below each numbered taxon. 1. Chromacris colorata (Serville) Rhomalea pedes Pictet and Saussure 2. Chromacris minuta n.sp. 3. Chromacris miles (Drury) 4. Chromacris speciosa (Thunberg) Acridium xanthopterum Hahn Rhomalea stolli Pictet and Saussure 5. Chromacris nuptialis (Gerstaecker) Rhomalea latipennis Pictet and Saussure 6. Chromacris trogon trogon (Gerstaecker) 7. Chromacris trogon intermedia n.subsp. 8. Chromacris psittacus psittacus (Gerstaecker) 9. Chromacris psittacus pacificus n.subsp. 10. Chromacris icterus (Pictet and Saussure) Rhomalea opulenta Gerstaecker 1 1. Chromacris peruviana (Pictet and Saussure) 12. Xestotrachelus robustus (Bruner) Xestolrachelus hasemani Bruner ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS Xestotrachelus robustus Figure 2. Xestotrachelus robustus. Chapada dos Guimaraes. Mato Grosso, Brazil. Key to Species of Chromacris 1. Exterior of small aedeagus membra- nous (Figs. 19-20). First two plaits or anterior fold of wings entirely black to base (Figs. 3-6). Colorata group _^^ 2 1 ' . Exterior of aedeagus sclerotized form- ing a short collarlike structure around the phallotreme opening (Figs. 22-25). First two plaits or anterior fold of wings yellow on basal half (Figs. 7-9), or some black on basal half of first plait (Fig. 10). Trogon group 6 2( 1). Yellow on part of antennae. Stridulat- ing structures weakly developed (Fig. 15) 3 2'. Antennae entirely black. Stridulating structures well developed (Figs. 16- 17) 4 3(2). Basal portion of antennae yellow (Mexico to Costa Rica) .__. C. colorata y . Distal portion of antennae yellow (Acapulco, Mexico) C. miniita 4(1'). Membrane of tegmina dark brown to black with strongly contrasting straw- colored veins (inland south central Brazil) C. nuptialis 4'. Not as above 5 5(4'). Posterior yellow margin of pronotum interrupted by black or green at angle between disc and lateral lobes. Rim of coxal articulation on mesathorax and metathorax entirely or partially yellow (South America) C. speciosa 5'. Posterior yellow margin of pronotum not interrupted at angle between disc and lateral lobes. Rim of coxal artic- ulation on mesathorax and metathorax entirely green (southeastern Mexico) C. miles 6(1'). Antennae entirely black 7 6'. Antennae with yellow tips 10 7(6). No bands on hind tibiae 8 7'. Yellow bands on hind tibiae 9 8(7). Yellow bands lacking on all legs (Cos- ta Rica) C. trogon trogon 8'. Yellow bands on hind femora (Guate- mala, Belize, and Honduras) C trogon intermedia 9(7'). Proximal yellow band on hind femora interrupted on outer ventral portion (Costa Rica to northern Colombia and Venezuela) C. psittacus psittacus 9'. Proximal yellow band on hind femora PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 Figures 3-11. Male hind wings of seven of the eight species of Chromacris (3-10) C. nuptialis being omitted because of similarity to C. speciosa (6), and Xestotrachelus robustus (1 1), ail at same scale and from the following localities: (3) colorata Medellin de Bravo, Veracruz, Mexico; (4) minuta Acapulco, Guerrero, Mexico; (5) miles Boloyuc, Quintana Roo, Mexico; (6) speciosa Aratinga, Rio Grande do Sul, Brazil; (7) trogon San Lorenzo, Alajuela, Costa Rica; (S) psillacus Cabima, Panama; (9) icterus Tabatinga, Amazonas, Brazil; (\0) peruviana Satipo, Junin, Peru; (I l)X. robustus Cerro Cora, Amambay, Paraguay. ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS entire (western Colombia and western Ecuador) C. psittacus pacificus 10(6'). Hind tibiae with single, distal, yellow band (southeastern Colombia to north- eastern Peru) C. icterus 10. Hind tibiae with two yellow bands (Peru) C peruviana Chromacris Walker Chromacris Walker, 1870:643. [Type-species Gryllus spe- ciosus Thunberg, 1824, by subsequent designation of Rehn 1904:532.] Diagnosis. — Neither pronotal crest nor pro- truding fastigium present; fully alate (Fig. 1). Medium size, body length of females 33 mm (C. minuta) to 55 mm (C. icterus and C. peruviana). Glossy olive-green to dark green with contrast- ing yellow or yellow tinged with red markings. These markings may be much reduced as in Pe- ruvian specimens of C. speciosa. Hind wings various shades of red, orange, or yellow with contrasting black pattern (Figs. 3-11) character- istic of this genus and Xestotrachelus, distin- guishing them from all other genera of subfam- ily. Genitalia (Figs. 19-26) rather uniform throughout the genus. Noteworthy are weakly developed aedeagal valves. Classification. — Two groups of Chroma- cris can be recognized. One group including tro- gon, psittacus, icterus, and peruviana (trogon group) have the proximal half of the first two plaits of the hind wings without black on the anterior portion of first two plaits (Fig. 10). In contrast, first two plaits of group containing co- lorata, minuta, miles, speciosa, and nuptialis (colorata group) are entirely black (Figs. 3-6). Aedeagal valves of trogon group are externally sclerotized (Fig. 22-25) whereas those of the co- lorata group are small membranous lobes (Figs. 19, 20). Trogon group members common- ly have yellow wings, occasionally orange, and rarely red. Wings of colorata group mem- bers are commonly red, orange in some geo- graphical areas, and frequently yellow in part of Atlantic coastal area of Brazil. In the trogon group, prosternal tubercle small and pointed with concave sides as seen in profile. In con- trast, C. miles, C. speciosa, and C. nuptialis have a large, bluntly pointed tubercle that is variable in form, even intra-specifically. On the other hand, tubercle of C colorata and C. mi- nuta more nearly resembles that of the trogon group in its small size, may be pointed, but sides in profile are rarely concave. C. miles and its two closely related species, C. speciosa and C nuptialis, have a well-developed stridulatory ap- paratus, as in many other species of the Ro- maleinae. The serrate cross veins (SV) between the first (lA) and second (2 A) anal veins of the second plait (Figs. 16, 17) contact the raised scraper veins on underside of tegmen when wings are folded. Tympanate or fenestrate areas (TA) on either side of these cross veins are well developed as resonators. This stridulatory ap- paratus is more weakly developed in other species of Chromacris and, in many cases, may not be functional. For example, compare the wing of C icterus (Fig. 15) with those in Figures 16 and 17. In summary, the trogon group is dis- tinguished from the colorata group by the ae- deagal valves and black pattern of hind wings, and the colorata group is divided into two subgroups by shape of prosternal tubercle and stridulatory apparatus. Recognition of species in the genus Chroma- cris has been difficult because of the lack of morphological characters and dependence on color and color pattern. The trogon group of four species, including two subspecies, can be rea- sonably well defined by a combination of differ- ent color-pattern characters (Table 2). The illus- trations of the aedeagus of this group (Figs. 22- 25) may suggest species differences, but individ- ual variation is such that clear distinctions be- tween species are not evident. The wide-ranging C. speciosa of South America has been most puzzling because of the great amount of geo- graphical variation of color and color pattern and variation within a local population. It has been difficult to decide whether we are dealing with species, subspecies, or just color forms. As an example of color forms, about half of the 21 specimens of C. speciosa recorded from Flores- ta de Tijuca near Rio de Janeiro, Brazil, have yellow wings and the other half orange wings, with no intermediates. We have concluded that it is most practical to treat C speciosa, at least for the present, as but one highly variable species. We need more information on C miles and C. nuptialis to understand their status as species and their relationship to C. speciosa. General Observations. — Most of the avail- able data for the genus refer to C. speciosa. The biology of this species was studied in Tucuman, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 C. speciosa b ^2Aa 18 -1 'I'W^ 'M 'M- '1 T TJ TA- N?-*^r^ SV^ ^fiwT L L aT^ TA- S ^^^ ■A>. r o b u s t u s / //O^ Qui -1A 2A 2Aa 2Aa Figures 1 2-14. Male hind wings of three species of Chromacris showing relative position of radius (R) median (M) fork to the second anal accessory (2Aa) fork. The 2Aa fork is more distant from the base of wing than the R-M fork in C. speciosa (Fig. 14) and its related species, C. miles and C. nuptialis. whereas in C. trogon and C. psittacus (Figs. 12 and 13) and all other species of the genus, the 2Aa fork is equidistant or closer to base of wing than the R-M fork. The If is the first and the 2f is the second fold line of wing. Figures 15-18. Details of stridulatory area of male hind wings of three species of Chromacris (Figs. 15-17) and Xesto- trachelus robustus (Fig. 18). Note well-developed tympanate areas (TA) in Figs. 16-18. Terminology of wing venation follows Ragge (1955). SV, serrate veinlets or cross veins; TA, tympanate or fenestrate areas; R, radial vein; Rs, radial sector; Cul, first cubital vein; lA, first anal vein; 2A, second anal vein. Argentina, by Barrera and Turk ( 1977). Data on the biology of C. colorata have been reported from Monterrey, Mexico, by Pretto-Malca (1968), at the other extreme for the distribution of the genus. Some data on the food of other species exist, mainly in papers of applied ento- mology. Some aspects of the general biology of the species of the genus, such as their general ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS preference for solanaceous and composite plants and the gregariousness of their juveniles, are generally known by all entomologists who have collected these insects in the field. OVIPOSITION AND DEVELOPMENT. — EggS of C. speciosa are laid in the soil, the top of the egg-pod 10 to 20 mm under the surface. Eggs are cemented together in the pods, but not embedded in the frothy secretion which covers the egg-pods of other acridoids. This secretion just forms the upper half of the pod, while the egg-mass is bare and usually 15 mm long, 8.5 mm wide. The number of eggs in each pod varies between 61 and 70, with a mean of 66 eggs (Bar- rera and Turk 1977). This species grows from hatching to imago, under the Tucuman climate, in 30 to 60 days, passing through five instars in the male and six instars in the female. Nymphs are black with red and some white markings. Sexual maturity is attained 10 days after the last molt. Adults mate repeatedly and females lay at least two pods. The insects usually disperse af- ter reaching the imaginal stage (Turk and Bar- rera 1976). Pretto-Malca (1968) stated that egg- pods of C. colorata contain an average of 35 eggs and that the insect reaches the imaginal stage through six nymphal instars, and sexual maturity about 18 days later. Nymphs of this species are highly gregarious. Food Plants. — Chromacris speciosa prefers solanaceous plants. Turk and Barrera (1976) re- ported its feeding on Cestrum parqui, C. stri- gillatum, C. lorentziana, Lycium cestroides. Solarium argentinum and S. verbascifolium (So- lanaceae), and also on Verbesina encelioides (Compositae) and alfalfa {Medicago sativa, Le- guminosae). They bred to maturity nymphs of this species found on alfalfa, using only this plant for food, and development was normal. They reported that in laboratory breedings nymphs which were first fed on Lycium ces- troides readily changed to other species of Lyci- um, but would starve to death rather than accept Solatium or genera of other plants. Conversely, nymphs started on Solanum would not accept Lycium. Thus, food plants, at the generic level, are determined by the first food of the newly hatched nymphs. The same feeding experiments demonstrated that this species would not eat species of the grass family (Turk and Barrera 1976). Carrasco (1962), however, reported C. speciosa and another unidentified species (evi- dently C. peruviana according to Carrasco' s de- scription) feeding on rice plants and doing con- siderable damage to this crop in Peru. Guagliumi (1973) mentioned C. speciosa as feeding on sug- ar cane in northeastern Brazil. Astacio-Cabrera (1975) reported C. colorata in Nicaragua on the composite Baltimora recta, and Pretto-Malca (1968) has stated that this species in Mexico usually feeds, and has been bred on, Solanum elaeagnifolium. Rowell (1978) reported the So- lanaceae as the preferred food of C. trogon in Costa Rica. Behavior. — The gregarious stages of Chro- macris, which are usually seen forming large groups on the tops of their food plants, together with their bright and contrasting coloration sug- gest that they are unpalatable or poisonous to predators and that their coloration is premoni- tory. One of us (H.R.R.) recently observed 20 to 30 conspicuous nymphs on top of a tussock of grass two or three meters from a small sola- naceous shrub stripped of its leaves, which sug- gests that the gregarious behavior is a part of the premonitory defense. After reaching the imaginal stage, these insects tend to disperse. Adults are very visible during flight, but once they alight on vegetation, they seem to disap- pear after the display of their colorful wings sud- denly ceases. Habitat. — The species of this genus are usu- ally found at forest edges and clearings, road- sides, edges of cultivated fields, and nearly all places where herbaceous solanaceous and com- posite plants occur. They seem to avoid heavy forest and prairie habitats. 1. Chromacris colorata (Serville) Acridium coloratum Serville, 1839:674. [Holotype, MNHN, no longer extant, said to come from South Carolina, USA, obviously in error. Mention of the 8-10 basal segments of the antennae as yellow and others black clearly indicates that it is the Mexican species.] Rhomalea pedes Saussure, 1859:392. [Lectotype, d. Geneva Museum; Mexico. So labelled by C.S.C. and here desig- nated.] Diagnosis. — Eight to 12 proximal segments of antennae yellow, distal segments black — a unique color pattern for the genus. Posterior margin of the pronotum yellow, but no yellow patches on midportion of lateral lobes, as is usu- al in C. speciosa. Three yellow bands on hind femur and only one distal band on hind tibia. Middle leg with one band on tibia, femur, and PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 22 C. trogon 23 C. psittacus 24 C. icterus 25 C. peruviana Figures 19-25. Dorsal and lateral views of aedeagi. Figs. 19 and 20 are examples of colorata-miles group. Figs. 22-25 are examples of trogon group. Fig. 21. Xestotrachelus rebus tus from Cerro Cora, Amambay, Paraguay. Fig. 19. Chromacris colorata from Medellin de Bravo, Veracruz, Mexico. Fig. 20. C. speciosa from Resistencia, Chaco, Argentina. Fig. 22. C. trogon from San Lorenzo, Aulajuela, Costa Rica. Fig. 23. C. psittacus from Las Pavas, Santa Marta Mts., Colombia. Fig. 24. C. icterus from Villavicencio, Colombia. Fig. 25. C. peruviana from Aucayacu, Huanuco, Peru. trochanter. Fore legs with one band on tibia. Hind wings rich cherry-red, with black pattern (Fig. 3) similar to that in C. speciosa. Prosternal tubercle tapers to a narrow point, much as in C. psittacus and other yellow- winged species. The small, short aedeagus formed by a pair of papil- lose, flattened lobes (Fig. 19) slightly sclerotized internally. Aedeagus similar to others of the co- lorata group. Distribution. — Tropical humid areas of Mexico south to Costa Rica. Of common occur- rence July to September. Specimens. — Mexico. States of Nuevo Leon, Tamaulipas, San Luis Potosi, Veracruz, Oaxaca, Guerrero, Morelos, Ja- lisco, Nayarit, Sinaloa, and Yucatan. Guatamala. El Peten: 3 mi [4.8 km] S Tikal, 16 Aug. 1974 (J. C. Lee), 6d, 39. 22 mi. [35 km] nw Poptun, 15 Jul. 1974 (J. C. Lee), Id, 39. BeUze. Rio Grande, Aug. 1931 (J. J. White), 39. Nicaragua. Managua: Sep. 1955 (E. Morales-Agacino), 2d. 19, CSC. Costa Rica. Guanacaste: Caiias, Sep. 1965 (C. H. F. Row- ell), Id, 19, CHFR. Comments. — Specimens from Volcan Colima (Jalisco, Mexico) and Guanacaste (Costa Rica) agree with Serville's description of Acridium coloratiim in lacking any banding on fore and middle legs and on hind tibia, and much reduced banding on hind femora. The type-specimen of Rhomalea pedes has fully banded legs, as have most Mexican specimens. Costa Rican speci- mens show small yellow marks on the prozonal part of the lateral lobes of the pronotum, as is ROBERTS AND CARBONELL; REVISION OF CHROMACRIS AND XESTOTRACHELUS also true of some specimens of C. speciosa. These Costa Rican specimens also have 12 seg- ments of the antennal flagellum yellow, the fol- lowing 4 segments part black and part yellow, and only the tip of the antennae entirely black. The Costa Rican specimen has no yellow on posterior margin of pronotum. 2. Chromacris minuta n.sp. Diagnosis. — Six to eight apical antennal seg- ments yellow, other segments black. Hind wings orange-red rather than cherry-red as in C co- lorata. Body color pale olive-green with greatly reduced yellow markings. No yellow bands on legs except weakly developed proximal and me- dian bands on hind femur. Prosternal tubercle short and conical. End of aedeagus much as in C colorata. Body size small for the genus, males 26-31 mm, females 35-39 mm from fas- tigium to end of wings. Specimens. — Holotype: 6, ANSP, Acapulco, Guerrero, Mexico, 13 Aug. 1935 (H. R. Roberts, E. R. Helwig). Paratypes: Mexico. Guerrero: 5 mi. [8 km] n Acapulco, 15 Sep. 1940 (C. Bolivar, H. R. Roberts), 16, 19. Same data as for holotype, 6(J , 8 9 , 3 juv. Comments. — This species is most similar to C. colorata. It should be looked for elsewhere along the Pacific coast of Mexico. A female of C. colorata from between Tierra Colorada and Ri'o Papagayo, about 40 km north of Acapulco, shows some reduction of yellow banding of the legs as occurs in C. minuta, but in other respects is typical of C. colorata. 3. Chromacris miles (Drury) Gryllus locusta miles Drury, 1773:79, pi. 42, fig. 2. [Holo- type, 9 , not found in British Museum or Oxford collections, but excellent figure should suffice; "Bay of Honduras."] Diagnosis. — Red wing pattern (Fig. 5) similar to that in C colorata and others of the colorata group. Resembles C colorata in lacking yellow in middle area of lateral lobe of pronotum. This condition occurs rarely in C speciosa. Similar to C speciosa in having entirely black antennae and two yellow bands rather than one on hind tibiae, but differs in having yellow on hind mar- gin of pronotum interrupted by black or green only on midline, whereas in C. speciosa it is interrupted on midline and both sides at the an- gles that limit disc from lateral lobes of meta- zona. Also, in C. miles rim of coxal articulation on mesathorax and metathorax green, whereas it is entirely or partially yellow in C. speciosa. Specimens.— Mexico. Quintana Roo: 5 mi. [8 km] se Po- lyuc (Boloyuc), 28 Jul. 1%0 (P. M. Litchfield), M, UMMZ. Veracruz: Laguna Verde, Aug. 1974 (M. Descamps), a small series of males and females, MNHN. Comments.— It has been thought that Dru- ry's name should be applied to Thunberg's South American species, C. speciosa. Finding specimens from Veracruz and the peninsula of Yucatan which closely match Drury^s figure supports the existence of a distinct species in the Bay of Honduras region. Based on the black pattern of the hind wings, this species belongs to the colorata group and is closest to C spe- ciosa in the strong development of the stridu- latory areas on the hind wing. Its color pattern is also more similar. C. speciosa, however, does not occur north of Colombia, and C. miles oc- curs within the range of C. colorata. More in- formation on the distribution of the genus in this region is needed to clarify our understanding of Drury's species. 4. Chromacris speciosa (Thunberg) Gryllus speciosus Thunberg, 1824:404. [Lectotype, 9 , so la- belled by C.S.C. and here designated; two male syntypes also examined; Uppsala Museum; Brazil]. Acridium xanthopterum Hahn, 1835, table A, fig. 2. [Holo- type unknown; Brazil. Hahn attributes the name to Perty in "Ins. nov bras.,"' but no such reference has been found. Black pattern of hind wing shown in illustration identifies it as the yellow-winged form of this species. New synonym.] Rhomalea stolli Pictet and Saussure, 1887:351. [Lecto- type, 3 , so labelled by C.S.C. and here designated; Geneva Museum; Bahia, Brazil. Synonym by Kirby 1910:373.] Diagnosis. — Varies geographically and local- ly. Red winged over most of its range, but in lowland coastal area from Bahia, Brazil, to Uru- guay, wings are frequently orange or yellow, and tegmina tend to be green rather than green tinged with red. Yellow-winged individuals readily distinguished from the typically yellow- winged species of the trogon group by entirely black basal half of the first two plaits (anterior or first paired fold of wing) (Fig. 6). In Paraguay, Argentina, and Uruguay wings tend to be or- ange-red, and yellow markings are tinged with red. In Bolivia, Peru, and Ecuador reduction and variation in leg banding evident. Elsewhere, hind femur almost always has three pale bands and the hind tibia two pale bands. Specimens from Ecuador and Peru lack hind tibial bands. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. Table 1. Variation in Banding of Hind Femur of Chromacris speciosa (22 specimens from Ecuador and Peru, 60 specimens from Santa Cruz, Bolivia). Banding of hind femur Ecuador, Peru Bolivia Strong Weak Absent Strong Weak Absent Proximal 17 5 40 20 Median — 10 12 5 37 18 Distal — 1 21 27 3 30 and hind femur usually has a strong proximal band, median band may be weak or absent, and distal band nearly always absent. Some 60 spec- imens from city of Santa Cruz region of eastern BoHvia show great variation in presence or ab- sence of various bands. Hind tibia may have a distal band or none. Hind femur usually has strong proximal band; it is never absent; median band usually weakly developed; distal band may be strongly developed (Table 1). Also, in the Andean region north to Ecuador, body color and tegmina darker. Specimens. — Unless otherwise noted all specimens have red hind wings, three yellow bands on hind femur, two yellow bands on hind tibia, and tegmen tinged with red. Colombia. Magdalena: Aracataca, 4-10 Aug. 1920 (Rehn, Hebard), 8^ f^San Salvador Fernandina -J ^\ < T3 -O ii ii £ Q c o S 3 0- Cu OS c/5 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43. No. 9 Figure 2. Measurements taken rrom Galapagos tortoise skulls (see Table 2 for explanations of abbreviations). G. hoodensis: CAS 8121, 8122. G. microphves: CAS 8158. G. nigrita: CAS 8381, 8289, 8286, 8385; MVZ 67613-67615, 59528, 67624-67629, 67631- 67633; USNM 104330-104331. G. phantastica: CAS 8101. G. vandenburghi: CAS 8141. G.v/cma: CAS 8 179, 8 193, 8 177; USNM 129247. G. wallacei (probably an invalid form, fide MacFarland et al. 1974a): CAS 8134. Geochelone sp. (but definitely Galapagos tortoise species): AMNH 7288, 42961, 63415, 36420, 36568-36570, 63416; CAS 8298, 8404, 841 1, 8409, 8402, 8377, 8407, 8410, 8403, 8414, 8397, 84 1 2, 8272; Calif. State Univ., Fullerton Coll. 3 uncat.; FMNH 13523, 1 uncat.; LACM (Vert. Paleo.) pr 63, pr 58, pr 64; MCZ 46606, 11070, 11069, 32098, 1905, 4668; MVZ 80075; SDSNH 56605, 55458; USNM 65896. 102904, 129393, 15192, 29338. 29305, 29254, 29252, 15190, 15193, 29256. Means and standard deviations were calculat- ed for each of the 16 measurements and corre- lation coefficients were also calculated. At the recommendation of Fritts (pers. comm.), I followed the last thorough taxonomic review (Van Denburgh 1 9 14) in which the different forms were given species-level designations. The species names used by Van Denburgh (1914) are fol- lowed with one exception; G. porteri is consid- ered a junior synonym of G. nigrita {fide Fritts in press). Statistical comparisons between island Table 2. Skull Measurements Recorded for Galap- agos Tortoises. (All measurements taken with dial calipers and recorded to nearest 0.01 mm.) I 'ariable— Description B— Basicranial length WAT — Width of skull at anterior tympanic opening WO- Width between orbits HN — Height of external narial opening WN — Width of external narial opening LB— Length of basisphenoid WB- Width of basisphenoid WZ— Width of quadratojugal WP- Width of postorbital WS-Width ofjugal DPV — Distance (greatest) from prepalatine foramina (or fo- ramen, if only one present) to vomer LP— Length of prootic WFS — Width of prootic at stapedial foramen PW — Width of pterygoid waist APW — Width of anterior prcmaxillac PC— Length of sagittal contact of prefrontals CRUMLV: TORTOISE SKULLS Table 3. Means and Standard Deviations for 16 Variables in Five Galapagos Tortoise Species. Measurements are illustrated in Figure 2 and abbreviations are listed in Table 2. Most sample sizes are small; all measurements are in millimeters. G. ephippium G. gunihen G. nigra a G. V cina G. chathamensis (N = = 9) (N = 15) (N = 18) (N = 4) (N = = 6) Variable X SD X SD X SD X SD X SD B 96.7 11.4 128.0 21.4 121.5 39.2 109.0 49.2 98.1 27.3 WAT 73.9 9.2 106.6 19.9 98.4 31.8 86 38.9 80.4 25.0 WO 25.1 3.1 35.4 7.3 37.0 13.3 28 4 12.6 28.4 7.7 HN 12.5 2.0 18.6 3.4 18.6 6.0 16 1 7.3 13.9 4.1 WN 17.0 2.2 25.1 4.4 23.1 7.7 21 3 9.2 18.5 4.7 LB 13.3 3.2 18.7 4.2 14.7 4.3 18 1 8.9 14.7 5.6 WB 14.6 2.2 19.1 4.3 17.1 4.6 15 8 7.2 13.8 , 3.4 WZ 9.3 3.6 14.3 4.4 13.3 4.7 12 6 6.8 10.1 3.5 WP 7.0 2 T 9.5 2.5 9.0 3.5 8 8 4.5 7.3 3.0 WS 7.3 2.3 12.0 3.0 9.5 4.0 9 6 5.1 7.9 3.1 DPV 3.2 0.8 4.2 0.8 4.2 1.5 3 7 2.4 3.1 1.0 LP 14.1 2.1 21.0 5.6 18.1 6.5 14 8 6.6 15.2 3.7 WFS 10.0 2.0 16.0 6.1 12.8 6.1 8 9 6.3 12.5 6.2 PW 19.2 1.8 25.9 5.1 26.1 8.4 21 9 8.6 19.0 4.0 APW 10.5 1.8 15.2 2.4 14.1 4.9 11 8 8.0 10.5 3.0 PC 8.6 1.5 10.6 4.0 13.3 5.4 8 5 3.9 8.2 3.9 populations were hampered by incomplete lo- cality data; 50 of 1 1 6 specimens (43%) examined possessed doubtful or unknown locality data. The specimens without locality data were readily identified as Galapagos tortoises, but could not be identified to species without locality data. These specimens were used in the computation of correlation coefficients and in factor analysis, but could not be used in other statistical proce- dures. To facilitate my analyses, populations were combined based on the shell types advocated by Van Denburgh( 1914) and Yr'ms {in press). Thus, the saddlebacked forms {G. abingdonii [N = 2]. G. phantastica [N = 1]. G. becki [N = 1], G. hoodensis [N = 2]. and G. ephippium [N = 9]) were combined, yielding a sample of 1 5 individ- uals. The non-saddlebacked forms (intermediate and domed shells of Van Denburgh 1914) were also combined, forming a larger sample of 48 individuals (G. chathamensis [N = 6]. G. dar- wini [N = 2]. G. guntheri [N = 15]. G. micro- phyes [N = 1]. G. nigrita [N = 18]. G. vicina [N = 4], and G. vandenburghi [N = 1]). These larger samples were then compared to determine whether cranial variation mirrored the already well known shell variation. Comparisons were also made among G. ephippium. G. guntheri. and G. nigrita to determine whether noncombined and combined samples contained the same mag- nitude of variation. The Statistical Package for the Social Sciences (SPSS) was used on the WYLBUR facility at the Campus Computer Information Service (CCIS) at Rutgers— The State University for initial data examination. Final statistical analyses were ac- complished using SPSS programs available Table 4. Means and Standard Deviations for Saddle- backed Tortoises Represented by Specimens of Five Species and Non-saddlebacked Tortoises Represented b\ Speci- mens OF Seven Species. Measurements are illustrated in Figure 2 and abbreviations are listed in Table 2; all measurements are in millimeters. Saddlebacked Nonsaddlebacked (N = = 15) (N = = 48) Variables X SD X SD B 98.9 15.9 116.5 37.1 WAT 75.8 13.0 96.0 30.4 wo 26.4 4.6 32.9 11.2 HN 13.2 2.9 17.1 5.6 WN 17.5 2.4 22.6 7.2 LB 13.8 3.2 16.0 5.7 WB 14.5 2.0 17.1 5.5 wz 9.5 3.4 13.1 5.3 WP 7.1 2.2 8.8 3.5 WS 7.3 2.3 9.9 3.9 DPV 2.9 1.2 3.9 1.4 LP 14.4 2.7 18.4 6.2 WFS 10.1 1.8 13.7 6.4 PW 20.1 2.9 24.6 7.8 APW 10.5 1.7 13.2 4.5 PC 7.9 1.4 10.2 5.2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43, No. 9 Table 5. Correlation Coefficients Between All the Skull Measurements Illustrated in Figure 2 and Abbreviated IN Table 2. All specimens measured are combined into a single sample. Nevertheless, all coefficients are significant to at least the P = 0.05 level. B WAT WO HN WN LB WB WZ WP WS DPV LP WFS PW APW WAT WO HN WN LB WB WZ WP WS DPV LP WFS PW APW PC 98 94 94 96 94 93 97 96 93 95 88 86 79 84 88 92 92 87 88 92 87 84 86 86 81 87 78 84 87 86 88 83 87 79 80 88 91 92 87 88 91 86 89 84 84 69 69 69 64 72 60 68 64 66 95 94 91 91 94 85 91 86 84 90 89 87 86 89 84 85 78 77 92 91 90 90 91 78 88 85 83 93 93 89 91 94 84 89 82 83 60 56 55 63 60 38 49 43 48 91 .63 87 .55 .91 83 .69 .90 .82 89 .68 .86 .82 .85 48 .49 .55 .50 .54 through the Office of Computer Services (OCS) at the Smithsonian Institution. Simple descrip- tive statistics, Hnear regression, factor analysis, and stepwise discriminant analyses were used to summarize observed cranial variation. Results Geochelone ephippiimi appears to have the smallest skull and G. gunthen the largest skull of Galapagos tortoises (Table 3), but when max- imum basicranial lengths (mean plus two stan- dard deviations) are compared, G. nignta ap- pears to possess the largest skull (B^^a,, = 171 mm for G. guntheri, 200 mm for G. nigrita). The efficacy of this procedure is in some doubt since the Bn^a^ for G. vicina exceeds that of G. nigrita, even though no skull of the former is anywhere near as large as the latter. This may be the prod- uct of a small sample size for G. vicina, repre- sented by only four specimens. The largest skulls in these samples are G. ephippiimi. 114.0 mm; G. guntheri, 157.7 mm; G. nigrita, 157.6 mm; and G. vicina, 142.7 mm. The G. nigrita sample includes the two smallest tortoises measured, which depresses the mean basicranial length and elevates the standard deviation. Combined samples clearly show a size differ- ential between saddlebacked and domed tortois- es; saddlebacked tortoises have smaller skulls. This is supported by all 16 variables (see Table 4). All correlation coefficients were significant to at least the P = 0.05 level (Table 5). Some vari- ables, however, did not correlate as highly with other variables. Examples include PC, DPV, and LB. Because intervariable correlation was so high, linear regression showed slight, if any, tendency toward curvilinearity. The intercepts for saddle- backed forms were lower than the intercepts for non-saddlebacked forms, reflecting the differ- ence in size between the two groups. Slopes, how- ever, were practically identical. As an example, linear equations relating WO to LB for saddle- backed and non-saddlebacked tortoises have slopes of 1.38 and 1.37, respectively, whereas intercepts are 7.92 and 1 1.74, respectively (r = 0.75 for saddlebacks and 0.69 for nonsaddle- backs, P < 0.005 for both). Factor analysis yielded three factors, the first accounted for almost 95% of the data variance (see Table 6). Before rotation all 16 variables correlated most highly with this first factor. Ro- tation simplifies vectors derived by the analysis procedure and is necessary because factor anal- ysis problems have more than one solution. There are two general rotation techniques: orthogonal and oblique. Orthogonal rotation solutions de- rive vectors along axes of data variation that are perpendicular to one another and thus uncorre- lated. Oblique techniques, on the other hand, do not require that vectors be orthogonal, so vectors can be correlated. Even after varimax rotation, an orthogonal technique that simplifies the col- umns of a factor matrix by maximizing factor- variable loadings, 12 of the 1 6 variables correlate most highly with factor one. Varimax rotation CRUMLV: TORTOISE SKULLS 117 oo ® Factor 3 Figure 3. A plot of factor scores for factors two and three. Geochelone nigrila (solid circles). G. gunthen (cross-hatched circles) and G. ephippiuin (open circles). When factor scores for all tortoises are plotted there is a prominent trend from the lower-left to upper-right quadrant. Although this general trend for all tortoises is suggestive of a positive trend toward increased snout elongation with increased robustness (as illustrated by G. nignta), the points for G. ephippium and G. gunthen show a negative relationship between robustness and snout elongation. was chosen because it maximizes the variation accounted for by the factor vectors without all the variables loading highly on the same factor, as occurs in quartimax rotation. Identifying vectors of data variation is spec- ulative: but it seems likely that factor one sum- marizes variation in size. Thus, 95% of the vari- ation in Galapagos tortoise skulls may be the result of variation in size. The other two factors are more difficult to interpret, partly because so little variation (only 5%) is summarized by these factors. Factor two summarizes variation in cra- Table 6. Statistics Produced by Factor Analysis Using Varimax Rotation. All specimens were included in this anal- ysis. Abbreviations used in the summarized factor matrix are listed in Table 2. Eigenvalues are measures of the relative importance of the factors. Factor 1 2 3 Eigenvalue 13.19 0.47 0.28 % Variation 94.6 3.4 2.0 Summarization of WFS 0.82 WP 0.63 PC 0.75 Factor Matrix LB 0.80 DPV 0.60 HN 0.51 LP 0.78 PW 0.53 WAT 0.76 WO 0.53 WS 0.76 WZ 0.72 B 0.76 WB 0.75 WN 0.73 HN 0.72 nial width and the width of skull arches, em- phasizing WO, WP, WZ, PW. and DPV. There- fore, factor two could be identified as some measure of robustness. Factor three, emphasiz- ing PC and HN, suggests there is variation in the anterior part of the skull. A high factor three score results from an increase in PC and HN. This results from elongating the anteromedial portion of the triturating surface, which concom- itantly yields a longer skull. A bivariate plot of the second and third factor scores for G. nigrita, G. guntheri, and G. ephip- pium (Fig. 3) indicates that as skulls become more robust, the anterior nasal part of the skull elon- gates; as robustness increases the skull becomes relatively longer. However, examining the indi- vidual points for G. guntheri and G. ephippium suggests just the opposite; as robustness increases elongation decreases. This negative relationship seems more pronounced in G. gunthen. Three separate discriminant function analyses were done: one for G. nigrita, G. ephippium. and G. guntheri: one for the combined samples: and one comparing small samples to larger samples. In the first analysis, the three forms were distin- guished by two factors (Table 7). Factor one sum- marized variation in 14 of the 16 variables but accounted for only 54.9% of the data variance. A high canonical correlation coefficient and a low Wilks's lambda indicate that this factor is good 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9 illl " 2 -6 ■4 2 2ee 4^ E g ,',' 2 e e Figure 4. A plot of the discriminant scores derived from an analysis that included Geochelone guntheri, G. nignta. and G. ephippium. Discriminant factor one is the horizontal axis and discriminant factor two is the vertical axis. Statistical sep- aration of these three populations is marked. High positive scores along the horizontal axis indicate small size, whereas high positive scores along the vertical axis indicate a poorly exposed basisphenoid. Upper case letters indicate group cen- teroids. at distinguishing groups. Geochelone ephippium is separated from the other two forms by this factor. The discriminating variables are nega- tively correlated to factor one, and the species with a small skull is differentiated from the two species with large skulls, suggesting that factor Table 7. Statistics Produced by a Discriminant Analysis of G. ephippium, G. guntheri and G. nigrita. Ab- breviations are listed in Table 2. Eigenvalues are measures of the relative importance of the factors; high canonical corre- lation coefficients (near I) and low Wilks's lambdas (near 0) indicate that factors are good discriminators. Discriminant function 1 2 Eigenvalue 7.06 5.82 % Variation 54.8 45.2 Canonical correlation 0.94 0.92 Wilks's lambda 0.02 (7^= 0.003) 0.15(^ = 0.02) Pooled within groups HN -0.22 LB -0.21 correlations bet. LP -0.20 WB -0.17 canonical WN -0.19 discr. fncts. & WAT -0.1 9 discr. variables APW-0.18 WO -0.18 Groups delineated G. ephippium G. nigrita from others from G. guntheri Figure 5. A histogram illustrating the results of a discrim- inant analysis of the saddlebacked and non-saddlebacked forms. The saddlebacked species are in the upper histogram, the non- saddlebacked species are in the lower histogram. The arrows indicate the median in each class. The discriminant scores (high positive scores indicate large size) are on the lower axis and the number of individuals are represented by left-hand axis. Although the saddlebacked and non-saddlebacked forms are clearly different sizes, there is significant overlap. one is an inverse measure of size. Factor two, which also has a high canonical correlation coef- ficient and a low Wilks's lambda, distinguishes G. nigrita from G. guntheri and accounts for the remaining variation in the data. Two variables are highly correlated with this second factor, LB and WB. Geochelone nigrita has high positive values for discriminating factor two, indicating that the basisphenoid is poorly exposed. Figure 4 graphically illustrates the completeness of sep- aration. Standardized canonical discriminant function coefficients are available upon request. These coefficients can be used to calculate discriminant scores for individual specimens whose identity is unknown; but choices are restricted to the pop- ulations originally compared (in this case G. ephippium, G. guntheri, or G. nigrita). The second discriminant analysis applied to the combined samples. Because only two groups were analyzed, a single discriminating factor was computed. The Wilks's lambda was not low, sug- gesting that the two groups cannot be easily dis- tinguished. The size differential between saddle- backed and non-saddlebacked tortoises is readily apparent (Fig. 5). Standardized canonical dis- criminant function coefficients arc available upon request. The third discriminant analysis compared CRUMLV: TORTOISE SKULLS Table 8. Classification Results oe a Discriminant Analysis Classification Procedure. Individual specimens were classified to one of three species: G. ephippiuin (a saddlebacked species). G. guntheri (an intermediate form) or G. nignta (a domed form). Asterisk indicates invalid taxon {fide MacFarland et al. 1974a). Shell type Trivial name Sex Mus. no. Classified as domed vandenburghi ? CAS 8141 ephippium intermediate chathamensis ? CAS 8133 ephippium intermediate chathamensis 7 CAS 8131 ephippium intermediate chathamensis 9 USNM 29255 ephippium intermediate chathamensis male CAS 8127 ephippium intermediate chathamensis ? CAS 8130 ephippium intermediate chathamensis ? CAS 8128 ephippium intermediate darwini female CAS 8106 ephippium intermediate darwini male CAS 8108 guntheri intermediate microphyes male CAS 8158 guntheri intermediate vicina male CAS 8179 ephippium intermediate vicina female CAS 8193 ephippium intermediate vicina 7 USNM 129247 ephippium intermediate vicina male CAS 8177 guntheri intermediate wallacei* male CAS 8 134 guntheri saddlebacked abingdonii •;' USNM 29269 guntheri saddlebacked ahingdonii male CAS 81 12 guntheri saddlebacked becki female CAS 8120 ephippium saddlebacked hoodensis male CAS 8121 ephippium saddlebacked hoodensis female CAS 8122 ephippium saddlebacked phantastica male CAS 8101 guntheri small samples of tortoise species to large sam- ples. Small samples were classified by the dis- criminant fiinction classification procedure to one of three species {G. guntheri, G. ephippium. G. nigrita). The results of this procedure are sum- marized in Table 8. Some species with inter- mediate shell types {fide VanDenburgh 1914) were classified as saddlebacked species (e.g.. G. chath- amensis was classified as G. ephippium). whereas other species with intermediate shell types were classified as G. guntheri. an intermediate form. No species was classified as a domed form. Skull variation did not parallel shell variation in any meaningful way. Discussion Small sample sizes and the paucity of accurate locality data limit the utility of this study. There- fore, samples were combined. (Thorpe. 1976. discusses the ramifications of such procedures.) Because most of the specimens in the United States were examined, this limitation cannot be overcome without costly and time-consuming removal of skulls from skins and stuffed speci- mens of known provenance. The choice of a putative ancestral morphotype makes an enormous difference in how one in- terprets evolutionary processes, patterns, and mechanisms. The size of the ancestral Galapagos tortoise is not known. Auffenberg (1971) be- lieved that the fossil Geochelone hesterna was a likely ancestral candidate for Galapagos tortoises as well as Geochelone chilensis from Argentina. The skull of G. hesterna is very much like a Galapagos tortoise skull. Although it is not as large as that of the largest of Galapagos domed tortoises, it is larger than that of the small sad- dlebacked tortoises. Thus, I favor an interme- diate-sized ancestor for Galapagos tortoises, per- haps something smaller than G. guntheri. If so, then G. nigrita is the result of continued gigan- tism and G. ephippium is the result of dwarfism. Why is there such flimsy coincidence between shell variation and cranial variation in Galapa- gos tortoises? Zangerl and Johnson (1957) and Zangerl (1969) have intimated that much of the shell variation observed in most species has little effect on an individuafs survival or fitness. Fritts {in press) has shown the contrary for Galapagos tortoises. But this selection on shell morphology does not seem to apply to skull morphology. What other selective factors could be molding skull morphology? I tend to agree with Bramble (1971), who felt that biomechanical constraints on chewing are the primary sources of selection upon turtle skulls. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43. No. 9 How Strong are these selective forces? Selective factors imposed by diet are known to be espe- cially strong in other animals from Galapagos. For example, Boag and Grant (1981) discovered that finches in Galapagos experienced intense se- lection upon beak size and shape as the result of a one-year drought. Because of the long life of tortoises and their ability to survive long periods without food or water, however, short-term en- vironmental changes such as those described by Boag and Grant are unlikely to affect tortoises as severely. Another source of selective pressure is possi- ble. During intraspecific agonistic encounters (Fritts, pers. comm.), the victor is the individual capable of raising its head the highest. Could apparent head width also affect the outcome of these battles? Interestingly, the relative head width of Geochelone guntheh and G. ephippium increases with size. These tortoises inhabit low dry islands (or parts of islands) where carrying capacities of the habitat may be lower and in- traspecific competition therefore higher. In con- trast, relative head width in G. nigrita decreases with size. This tortoise lives on a higher moist island where carrying capacities may be higher and intraspecific competition may not be as in- tense. Also, this apparent decrease in relative width actually accompanies an increase in the length of the masticatory surface area, perhaps allowing more efficient mastication. Acknowledgments I thank L. Barnes (LACM), R. Crombie (USNM), R. Drewes (CAS), G. Foley (AMNH). T. Fritts (SDSNH), A. Leviton (CAS), H. Marx (FMNH), W. Presch, A. Resetar (FMNH), J. Ro- sado (MCZ), H. Voris (FMNH), D. Wake (MVZ), E. Williams (MCZ), G. Zug (USNM), and R. Zweifel (AMNH) for making specimens avail- able. I also thank R. Crombie, C. Ernst, T. Fritts, S. McDowell, E. Meyer, K. Miyata, D. Stead- man, and G. Zug for reading parts or all of the manuscript and providing helpful suggestions. Funding, for which I am most thankful, came from a Sigma Xi Grant-In-Aid of Research, the Alvarado Community Hospital Research Foun- dation, the Vertebrate Zoology Reserve Fund of San Diego State University, and the Theodore Roosevelt Memorial Fund of the American Mu- seum of Natural History. Data analysis, begun at Rutgers— The State University with Depart- ment of Zoology and Physiology support, was completed with the assistance of the Office of Computer Services during my tenure as a Smith- sonian Predoctoral Fellow at the National Mu- seum of Natural History. Literature Cited Arnold. E. N. 1 979. Indian Ocean giant tortoises: their sys- tematics and island adaptations. Phil. Trans. R. Soc. Lend. (8)286:127-45. AuFFENBERG. W. 1971. A ncw fossil tortoise. with remarks on the origin of South American testudinines. Copeia 1 97 1: 106-17. Boag, P. T., AND P. R. Grant. 1981. Intense natural selec- tion in a population of Darwin's finches (Geospizinae) in the Galapagos. Science 214:82-85. Bramble. D. M. 1971. Functional morphology, evolution, and paleoecology of gopher tortoises. Ph.D. thesis, Univ. Calif, Berkeley. 341 p. Crumly, C. R. 1980. The cranial osteology and evolution of the tortoise genus Geochelone (Testudines, Testudinidae). M.S. thesis, San Diego State Univ. 338 p. . 1982. A cladistic analysis ofGeo(r/7e/o«f using cranial osteology. J. Herpetol. 16(3):2I5-34. Duellman, W. E., T. Fritts, and A. Leviton. 1978. Mu- seum acronyms. Herp. Rev. 9(I):5-9. Fritts, T. H. [in press]. Morphometries of Galapagos tor- toises: evolutionary implications. Garman, S. 1917. The Galapagos tortoises. Mem. Mus. Comp. Zool. 30:262-96. Gunther. A. 1875. Description of the living and extinct races of gigantic tortoises. Parts I and II: The tortoises of the Galapagos Islands. Phil. Trans. R. Soc. Lond. (B) 165: 251-84. . 1877. The gigantic land tortoises (living and extinct) in the collection of the British Museum. British Museum, London. 96 p. MacFarland, C. G., J. Villa, and B. Toro. 1974a. The Galapagos giant tortoises (Geochelone elephantopus). Part I: The status of the surviving populations. Biol. Conserv. 6: 118-33. . 1974b. The Galapagos giant tortoises (Geochelone elephantopus). Part II: Conservation methods. Biol. Con- serv. 6:198-212. Marlow. R.. and J. L. Patton. 1981. Biochemical rela- tionships of the Galapagos tortoises (Geochelone elephan- topus). J. Zool. London 195:413-22. Pritchard, p. C. H. 1979. Encyclopedia of turtles. T. F. H. Publishers. 895 p. Rothschild, L. 1901. A new land tortoise from the Gala- pagos Islands. Nov. Zool. 8:372. . 1902. Description of a new species of gigantic land turtle from the Galapagos Islands. Nov. Zool. 9:619. . 1903. Description ofa new species of gigantic tortoise from Indefatigable Island. Nov. Zool. 10:1 19. . 1915. The gigantic land tortoises of the Galapagos Islands in the Tring Museum. Nov. Zool. 22:403-17. Thorpe, R. S. 1976. Biometrical analysis of geographic vari- ation and racial affinities. Biol. Rev. 51:407-52. Van Denburgh, J. 1907. Expeditionof the California Acad- emy of Sciences to the Galapagos Islands. 1905-1906. Part CRUMLY: TORTOISE SKULLS j -, , L Preliminary descriptions offour new races of gigantic land . 1977. Liste der rezenten Amphibien un Reptilien tortoises from the Galapagos Islands. Proc. Calif Acad. Sci. Testudines. Crocodylia, Rhyncocephalia. Das Tierrich 100- 1:1-16. 1-174. . 1914. Expedition ofthe California Academy of Sc- Zangerl, R. 1969. The turtle shell. Pages 31 1-^19 //; Cans ences to the Galapagos Islands. 1905-1906. Part X: The C, A. d'A. Bellaris, and T. S. Parsons. Biologv ofthe Rep- gigantic land tortoises ofthe Galapagos Archipelago. Proc. tilia, vol. 1, Academic Press New York Cahf Acad. Sci.. ser. 4, 2:203-374. . ^^d R. G. Johnson." 1957. The nature of shield Wermuth. H.. and R. Mertens. 1961. Schildkroten, kro- abnormalities in the turtle shell. Fieldiana: Geo!. 10:341- kodile, und bruckenechsen. Gustav Fischer Verlag, Jena. 62 422 p. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco. California 941 18 PROCEEDINGS OF THE CALIFORNIA ACAaEM^g9J6^(:iENCES LIBRARY ' Vol. 43, No. 10, pp. 123-140. 8 figs., 1 table J*N 271984 January 17, 1984 THE STATUS OF T /? W^^J^SrmMmim. US (LINNAEUS, 1758), MEDIUM DE BEAUMONT, 1945, AND MINUS DE BEAUMONT, 1945 (HYMENOPTERA: SPHECIDAE) By Wojciech J. Pulawski California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Abstract: Trypoxylon figulus (Linnaeus, 1758), medium de Beaumont, 1945, and minus de Beaumont, 1945, currently confused under the name figulus, are separated on the basis of newly discovered characters. T. majus Kohl, 1883, figulus barbarum de Beaumont, 1957, and figulus yezo Tsuneki, 1956, are newly synonymized with^^«/«i, anA figulus koma Tsuneki, 1956 is newly synonymized with minus. Neotypes are designated for Sphex fuliginosus Scopoli, 1763, and Trypoxylon majus Kohl, 1883, both synonyms o{ figulus, and a lectotype is designated for Trypoxylon rubi Wolf, 1959, a synonym of medium. Introduction De Beaumont (1945) was first to observe that Trypoxylon figulus of European authors actually consisted of three phena. Their status has been controversial over the years. De Beaumont ( 1 945, 1964a) and Richards (1980) called them vari- eties, Bluthgen (1951) gave them species rank, and Wolf (1959) and Bohart and Menke (1976) treated them as subspecies. The last interpreta- tion is untenable, since the three phena are large- ly sympatric. Tsuneki (1981) regarded medium as a good species, characterized by both external and genitalic characters, and considered minus as a simple form of figulus. According to Valkeila (1961). specimens reared from one nest mostly are one phenon, but he reported that two phena (e.g., majus and minus) are found in some nests. He concluded that all three are individual variants of one species. Un- fortunately, Valkeila's data cannot be verified. I have examined all of his specimens, which are presently kept at Helsinki University. Some specimens have identification labels by de Beau- mont, but not a single label refers to nests or cells from which specimens were reared. Possibly Valkeila misidentified some specimens, but this cannot be determined because his identification labels give the name figulus only, without ref- erence to form or varietal names. Another pos- sible explanation is that offspring of two nests were accidentally confused. A thorough examination of the three phena, based on more than 3800 specimens from many countries, convinced me that actually they are good species. My opinion is based on the follow- ing evidence: 1. Morphology. Although some males of fi- gulus and minus cannot be distinguished with certainty, females are separated by structural gaps and do not intergrade; also the male of medium is easily recognized by its peculiar gonoforceps. Some previously unnoticed characters (antero- [123] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43. No. 10 ventral mesothoracic process, length of thoracic vestiture, female hindcoxal pit) are especially helpful in recognizing the three species. 2. Rearing. Of 91 specimens reared from trap nests in Liege, Belgium, by A. Jacob-Remacle, and examined by me, all are minus (7 9,5 5 reared in 1976, 44 2, 35 S reared in 1981). This resuU contradicts Valkeila's conclusions: If the three phena really are variants of one species, then one would expect some variation of phena in the Liege trap nest material. 3. Geographic distribution. All three species are largely sympatric, but only figidus and me- dium have been found in Great Britain, only figulus and mimis in the Iberian Peninsula, and only figulus in North Africa and North America. If the three phena were just individual forms of one species, some variation should have oc- curred in North American populations, and all three phena should have been observed in Great Britain (all three do occur in the Netherlands, where climatic and ecological conditions are practically identical to those in England). 4. Habitat preference. Of 429 specimens col- lected by J. Leclercq in waterbowl traps in Liege, Belgium, in 1980 and 1981 (and examined by me), 4 are medium and the remainder are minus (202 2, 223 S). Not a single figidus was trapped, in spite of the fact that this species is common in Belgium. I fully agree with Tsuneki (1981) that some of the previously used characters are not reliable. For example, the mesopleural punctures of ^7- gulus are dense and well defined according to de Beaumont (1945, 1964a), but in the smallest males they actually are as sparse and minute as in most minus. Such characters have not been used here. Proper mounting is critical for studying the species considered. For example, the anteroven- tral mesothoracic region must not be damaged by the pin. When pinning the specimens, one should insert the pin so that it passes through the membrane between the mesothorax and fore- coxae (which then extend laterad). In this posi- tion, the anteroventral mesothoracic region is easily visible. Unfortunately, many European collectors mount their specimens on cardboard rectangles with glue or minutiae (venter down rather than on a side), and such specimens must be relaxed and remounted before examination. In the text below the locality records are ar- ranged according to current administrative di- visions for each country except Sweden and Fin- land, for which biogeographic provinces have been used. Localities given on specimen labels but not found on available maps or in gazeteers have not been considered. An exclamation mark preceding the word Ho- lotype or Neotype in the bibliographic citations indicates that the type has been examined. Sources of Material The specimens examined came from institu- tional and private collections listed below. The acronyms preceding the names are the abbrevi- ations by which these collections are referred to in the text. AKM: Aimo K. Merisuo, Turku, Finland AWE: Father Andreas W. Ebmer, Linz, Austria BB: Padre Bruno Bonelli, Cavalese, Italy BMNH: British Museum (Natural History), London, England (Mr. C. R. Vardy) CAS: California Academy of Sciences, San Francisco, Cali- fornia (W. J. Pulawski) CNC: Canada National Collection of Insects, Arachnids and Nematods, Biosystematics Research Institute, Ottawa, On- tario CU: Cornell University, Department of Entomology and Lim- nology, Ithaca, New York (Dr. L. L. Pechuman) DBB: Major Donald B. Baker, Ewell. Surrey, England DEI: Institut flir Pflanzenschutzforschung der Akademie der Landwirtschaftswissenschaften der DDR, Zweigstelle Ebers- walde, Abteilung Taxonomie der Insekten (formerly Deutsches Entomologisches Institut), Eberswalde-Finow (Dr. J. Oehlke) FIS: ForschungsinstitutSenckenberg, Frankfurt am Main, Fed- eral Republic of Germany (Dr. J. -P. Kopelke) FJS: Seflor Francisco J. Sudrez, Almeria, Spain FSAG: Faculte de Sciences Agronomiques, Gembloux, Bel- gium (Dr. J. Leclercq) GP: Signor Guido Pagliano, Turin, Italy GVR: Mr. Gerard van Rossem, Wageningen, The Netherlands HD: Dr. Holger Dathe, Forschungsstelle fiir Wirbeltierfor- schung, Berlin, German Democratic Republic HW: Herr Heinrich Wolf Plettenberg, Federal Republic of Germany HY: Helsingin Yliopisto (=University of Helsinki), Depart- ment of Agricultural and Forest Zoology, Finland, including E. Valkeila collection (Dr. Martti Koponen) lEE: Instituto Espai^ol de Entomologia, Madrid, Spain (Dr. E. Mingo Perez) JG: Dr. Joseph Gusenleitner, Linz, Austria KMG: Mr. Kenneth M. Guichard, % British Museum (Natural History), London, England KS: Professor Dr. Konrad Schmidt, Zoologisches Institut der Universitat, Karlsruhe, Federal Republic of Germany KT: Professor Katsui Tsuneki, Mishima, Japan LEM: Lyman Entomological Museum & Research Laboratory, Ste. Anne de Bellevue. Quebec. Canada (Dr. A. Finnamore) MGA: Muzeul de Istorie Naturala Grigore Antipa, Bucharest, Rumania (Mrs. X. Scobiola Palade) PULAWSKI: THE STATUS OF TRYPOXYLOS FIGVLVS, MEDIUM. AND MINI'S Table 1. Ancestral and Derived Character States of Three Species in the Genus Trypoxylos Character Ancestral Derived 1. Thoracic pilosity 2. Anteroventral mesothoracic process 3. Free margin of female clypeus 4. Female hindcoxal pit 5. Sete of hindcoxal pit 6. Male apical flagellomere 7. Gonoforceps process shorter (as in medium) absent straight or sinuate circular evenly distributed shorter (as in medium) absent longer (as in figulus) present concave oblong channel-like structure longer (as in figulus) present MHNG: Museum d'Histoire Naturelle de Geneve. Switzerland (Dr. CI. Besuchet) MSNM: Museo Civico di Storia Naturale, Milano, Italy (Dr. C. Leonardi) MCZ: Museum of Comparative Zoology, Harvard University. Cambridge, Massachusetts NHMB: Naturhistorisches Museum Bern. Bern. Switzerland (Dr. H. D. Volkart) NHMV: Naturhistorisches Museum. Vienna. Austria (Dr. M. Fischer) NRS: Naturhistoriska Riksmuseet, Stockholm V. Sweden (Mr. S. Erlandsson) RMNH: Rijksmuseum van Natuurlijke Historic. Leiden. The Netherlands, including collections of J. P. van Liih and P. M. F. Verhoefr(Dr. K. van Achterberg) SFG: Dr. Severiano Fernandez Gayubo. Departamento de Zoologia. Universidad de Salamanca. Spain SMT: Staatliches Museum fur Tierkunde. Dresden, German Democratic Republic (Dr. Regine Eck) TMB: Termeszettudomanyi Muzeum. Budapest. Hungary (Dr. J. Papp) TN: Mr. Toshiaku Nambu, Yoni-machi, Saitama Prefecture, Japan USNM: United States National Museum (Smithsonian Insti- tution), Washington, D.C. VH: Dr. Volk Haeseler, Universitat Oldenburg. Oldenburg. Federal Republic of Germany VLK: Dr. Vladimir L. Kazenas. Zoological Institute. Kazakh Academy of Sciences. .\lma Ata. USSR WJP: Wojciech J. Pulawski. San Francisco. California WSU: Washington State University. Department of Ento- mology, Pullman, Washington ZMB: Museum flir Naturkunde an der Humboldt Universitat zu Berlin, German Democratic Republic (Dr. F. Koch) ZMH: Zoologisches Institut und Zoologisches Museum der Universitat Hamburg. Federal Republic of Germany (Dr. R. Abraham) ZMK: Zoological Museum. Copenhagen, Denmark (Dr. O. Lomholdt) ZMMU: Zoological Museum. Moscow State University. Mos- cow. USSR (Dr. L. V. Zimina, via Dr. A. P. Rasnitsyn) ZMUB: Zoological Museum, University of Bergen, Norway (Dr. Lita Greve Jensen) ZSM: Zoologische Staatssammlung Miinchen. Federal Repub- lic of Germany (Dr. E. Diller) Acknowledgments Study of the Linnean type of Sphex figulus at the BurUngton House, London, was possible ow- ing to the kind assistance of Michael C. Day of the British Museum (Natural History). North American specimens of figulus belonging to var- ious U.S. and Canadian institutions listed above (except for LEM) were kindly forwarded by Rolin C. Coville, University of California. Berkeley, who had them on loan, Arnold S. Menke and Eric E. Grissell critically reviewed the manu- script and made many valuable suggestions. Da- vid H. Kavanaugh commented on the phyloge- netic trees. Mary Ann Tenorio drew the phylogenetic schemes and distributional maps, and Donald J. Becker took the photographs with a Hitachi S-520 scanning electron microscope. Phylogenetic Relationships Reconstructing phyletic relationships between three isolated species of a large genus like Try- poxylon is precarious, because polarities of mor- phological transformations can easily be misin- terpreted. With this restriction in mind, I nevertheless think it worthwhile to analyze the relationships between figulus. medium, and mi- nus. Their ancestral and derived character states, based on outgroup comparisons, are shown in Table 1. Based on the above table, the three possible phylogenetic trees (only dichotomic trees are considered) are as shown in Fig. 1 . Tree B is the most probable, since no single derived character state is shared by any two of the three species in the schemes A and C. Furthermore, trees A and C imply a parallel development of the elongate male flagellomere XI in figulus and minus, an unlikely event. Key to the Species 1 . Female: clypeal free margin evenly concave between orbit and median projection (Fig. 7A); mesopleural setae around scrobe shorter than midocellar diameter; hindcox- 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43. No. 10 Figure 1 . Hypothetical phylogenetic trees of Trypoxylonfigidits (fig), medium (med). et minus (min). Open circles: generalized character states. Solid circles: derived character states. Numbers refer to characters listed on p. 125. al pit nearly circular, its setae forming pe- culiar, channel-like structure (Fig. 7B-D). Male: maximum length of flagellomere X 0.75-0.9 times its width; maximum length of flagellomere XI generally 2.0-2.2 times its basal diameter (occasionally 2.4 times); gonoforceps with externoventral expansion at about midlength (Fig. 7E, F) medium de Beaumont Female: clypeal free margin (Fig. 2A, 5A) sinuate to nearly straight, except concave in occasional western European figulus in which mesopleural setae around scrobe are generally longer than midocellar diameter, and hindcoxal pit is generally oblong; setae of hindcoxal pit not forming channel-like structure. Male: maximum length of fla- gellomere X 0.5-0.8 its width; maximum length of flagellomere XI 2.2-3.6 times its basal diameter; gonoforceps not expanded exteroventrally (Fig. 2E, F) 2 Mesothorax without antero ventral process'; mesopleural setae around scrobe in most specimens longer than midocellar diameter (shorter in some individuals). Female: clyp- eal free margin sinuate between lobe and orbit (Fig. 2A), hindcoxal pit oblong (Fig. ' In occasional males the anteroventral mesothoracic process is absent and the mesopleural setae are shorter than the mid- ocellar diameter. Such specimens can be either figulus with unusually short mesopleural setae, or mmus without meso- thoracic process. I cannot find characters for distinguishing them. 2B-D) or (some specimens) circular, eva- nescent in Japanese specimens figulus (Linnaeus) - Mesothorax with anteroventral process (Fig. 5B, C) in more than 95% of specimens; mesopleural setae around scrobe shorter than midocellar diameter. Female: clypeal free margin almost straight between lobe and orbit (Fig. 5A), hindcoxal pit circular (Fig. 5D-F) mimts (de Beaumont) Trypoxylon figulus (Linnaeus) Sp hex figulus Linnaeus, 1758:570. ! Holotype: 9. Sweden, Uppsala (Linnean Society, London). — Day, 1979:62. — In Trypoxylon: Latreille, 1802:79: Tsunekj, 1981:15 (rede- scription, geographic variation). — In Apius: Jurine, 1807: 142. Sphex fullginosus Scopoli, 1763:292 (as fuliginosa, incorrect original spelling). Holotype or syntypes: Camiolia (formerly in Austria, since 1919 part of Italy and Yugoslavia), lost, see Rogenhofer und Dalla Torre, 1882:599. ! Neotype: 9, Austria: "Carinthia, Ostkarawanken, Ebriach, 580-750 m, 21-29. VII. 1964, G. van Rossem," present designation (CAS). — As probable synonym of figulus: vander Linden, 1829:42. — As synonym of figulus: subsequent authors. Trypoxylon figulus var. majus Kohl, 1 883:657, 9, 6 (as major. incorrect original spelling). Holotype or syntypes: 9, Swit- zerland: no specific locality (originally NHMV, Vienna, now lost). ! Neotype: 9, Switzedand, "P. 3 VIII 84" and "Cn. Toumier" (=Peney near Geneva, collection Tournier), pres- ent designation (MHNG). New synonym. — de Beaumont, 1945:477 (var. major): Bluthoen, 1951:234 (var. major): DE Beaumont, 1958:206 (forma major). 1959:30 (same); Wolf, 1959:15, \6 (figulus major): \alke\la, 1961:244 (var. /?!i2/or); DE Beaumont, 1964a:290, 1964b:84 (forma wa/or), 1965:56 (same), 1967:338 (same); Bohart and Menke, 1976: 346 (ssp. major): Lomholdt, 1976:267 (figulus major): Richards, 1980:45 (var. major). Trypo.xylon apicale W. Fox, 1 89 1 : 1 42, 9 (as apicalis, incorrect PULAWSKI: THE STATUS OF TRYPOXYLON FIGULVS. MEDIUM. AND MINI'S 127 original spelling). Lectotype: 9, Canada (ANSP, Philadel- phia), designated by Cresson, 1928:52. — Sandhouse, 1940: 156 (apicale). Synonymized by Pate, 1943:16. Trypoxylon figiihis barbaruin de Beaumont, 1957: 9. $. Ho- lotype: 3, Morocco: Marrakech (Mus. Zool. Lausanne). New synonym. — Bohart and Menke. 1976:346. Trypoxylon figulus yezo Tsuneki, 1956:29, 9, 6. Holotype: 9. Japan: Hokkaido [=Yezo]: Jozankei (K. Tsuneki collection, Mishima). New synonym. — Bohart and Menke, 1976: 346; Tsuneki, 1981:21 (summary of faunistic data). Trypoxylon fteuzet I Giner Mart, 1959:389, 6. ! Holotype: (5, Morocco: Fez (lEE, Madrid). Synonymized W\i\\ figulus bar- barum by Suarez, in Giner MarI, 1959:400. Comments on New Synonyms. — T. figulus barbarum was based mainly on the elongate male flagellomere XI. Because flagellomere XI varies in length (see Geographic Variation below) this subspecies is not recognized here. I also feel that a formal name for the Japanese populations (fi- gulus yezo) is unwarranted on morphological or other grounds. Comments on Neotypes. — The identity of Sphex fiiliginosus has never been satisfactorily established, because the original description is inadequate and the original material is lost (Ro- genhofer und Dalla Torre, 1882). Consequently, the name can only be defined by designation of a neotype. In selecting a specimen of Trypoxylon figulus as a neotype of Sphex fidiginosus I have followed the traditional interpretation of the last name. The original material of Trypoxylon majus cannot be found in the Vienna Museum (Dr. M. Fischer's letter of 21 October 1982) and must be lost. However, this name indicates a large body size, and figulus averages larger than either me- dium or minus. A neotype of majus has also been designated. Diagnosis. — Most specimens of figulus differ from medium and minus in having the meso- pleural setae around the scrobe slightly longer than the midocellar diameter. However, the setae length is slightly less than this diameter in some specimens from southern France and the Iberian Peninsula (as they are in the other two species). The anteroventral mesothoracic carina is sin- uate, curved posterad mesally, but unlike most minus it has no process. The free margin of the female clypeus (Fig. 2A) is usually sinuate be- tween orbit and the median projection (free mar- gin concave in medium, almost straight in mi- nus). However, the free margin is almost evenly concave in certain specimens from Spain (almost like medium, which is unknown from Spain), in a specimen from Zirbelwald, Austria, and one from Balderschwang, Federal Germany. The fe- male hindcoxal pit is mostly oblong (Fig. 2B-D) in western palearctic specimens, but occasionally it is nearly circular, as in medium and minus: it is evanescent in Japanese females. In the male, the maximum length of flagellomere X equals 0.65-0.8 of its width (the lowest ratios are ob- served is specimens in which flagellomere XI is short, and vice versa); the maximum length of flagellomere XI usually is 2.4-3.6 times the basal diameter instead of 2.0-2.2 in most medium, but only 2.2 times in occasional specimens (which differ from medium in having a longer meso- pleural vestiture and a shorter flagellomere X). Body length 9-12 mm in female, 7.5-10 mm in male. Geographic Variation. — In most males (in- cluding the two males seen from Portugal), the maximum length of flagellomere XI equals 2.4- 2.7 times its basal diameter, but in occasional specimens it is only 2.2 times (e.g., in a male from Wachseldommoos, Switzerland); it is 2.7- 3.2 times its basal diameter in Spanish individ- uals, and 3.3-3.6 times in Moroccan individuals. Life History. — Many specimens of figulus (voucher specimens examined by me) were reared from nests established in wood (Wolf. 1959). Six females and 1 7 males examined were reared by O. Lomholdt from nests in reed stems which had been used for thatching roofs at Tisvilde Hegn, Denmark. Geographic Distribution (Figs. 3, 4). — Most of the Palearctic Region between Great Britain and Japan, and also eastern North America (east- em Canada and northeastern USA). Records (Old World).— Algeria (1 3): El Harrach (as Maison Carree, apical flagellomeres missing, BMNH). Austria: (102 9, 50 S, NHMV if not indicated otherwise): Karnten: Afritzer See (WJP), Ebene Reichenau (RMNH), Ebriach in Ostkarawanken (WJP), Eisenkappel, Mallnitz (ZMB), Mauthen (ZMB). Notsch. Waidisch bei Ferlach (FSAG. JG). Niederosterreich: Bisamberg near Vienna (NHMV, CU), Buck- lige Welt S Vienna. Bruhl, Dombach (CAS, NHMV). Eichkogel near Vienna (RMNH), Guntramsdorf (DEI), Hainbach (FSAG), Hainburg an der Donau (ZMB). Herzogenburg. Herzograd (JG), Kalksburg near Vienna, Krumbach, Lobau near Vienna (NHMV. ZMH). Marchfeld (ZMB). Mistelbach (ZMH), M6- dling (ZMH), Oberweiden (DBB), Piesting, Purk (W Krems), Rappendorf bei Molk (AWE), Roggendorf bei Melk (JG), Rohr im Gebirge, Schneeberg, Stillfried (ZMH), Traismauer, Weid- lingsbach (ZMH), Wien (NHMV. FSAG. ZMH) including Donauauen, Kahlenberg and Tiirkenschanze. Oberosterreich: Frauenstein (JG), Gemeinde Reichenthal (AWE), Gutau (FSAG, JG), Hofkirchen (FSAG), Innerbreitenau (FSAG. JG). Kalten- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 Figure 2. Trypoxylon figulus: A — female clypeus, B — female hindcoxa ventrally, C — female hindcoxal pit, vertical view, D— same, oblique view, E— male genitalia (arrow: area shown in F). F— same, portion of gonoforceps. berg (AWE), Linz (HY, NHMV), Molln, Muhlviertel (JG), Neumarkt (JG), Oberwallsee bei Mullachen (JG), Riedegg bei Gallneukirchen (JG), Sankt Willibald (AWE), Ternberg (JG), Welserheide, Zeissberg bei Freistadt (FSAG, JG). Salzburg: Koppl bei Aschach (JG), Salzburg (BMNH). Steiermark: Leut- schach (JG), Sankt Ulrich (JG), Tragoss-Oberort (DBB), Wem- burg (FSAG). Tirol: Huben in eastern Tirol (ZMB), Innsbruck (NHMV, ZMB), Iselsberg (DBB). Lienz (RMNH), Obladis, Hopfgarten, Salvenberg (CAS), Zirbelwald near Obergurgl, 1 km SW Zwiselstein in Otztal. Voralberg: Ittensberg. Belgium (95 9, 63 <5, FSAG if not indicated otherwise); Bra- bant: Evere, Genval, Gistoux. Grez-Doiceau, Mont-Saint-Gui- bert, Nethen, Rhode-Sainte Agathe, Thorembais-Saint-Trond, Uccle, Waterloo. Hainaut: Aiseau, Athis, Barry, Binche. Bous- su, Bouvignies, Fleurus, Orcq, Seneffe, Taintignies, Velaines, Wanfercee. Liege: Acosse, Aubel, Barchon, Ben-Ahin, Beyne: ca 15 km SE Liege (BMNH), Beyne-Heusay, Cerexhe, Char- neux. Chevron, Clermont-sur-Berwinne, Fleron, Foret de Grunhault, Francorchamps, Henri-Chapelle, Hombourg, Ju- pille. La Calamine, La Reid, Lontzen, Montzen, Pepinster, Queue-du-Bois, Romsee, Spa, Xhendelesse, Welkenraedt. Limburg: Berg pres de Tongres, Bocholt, Godsheide, Tongres. Luxembourg: Amonines, Hotton, Les Epioux, Lomprez, Ozo, Saint-Medard. Sampont, Smuid, Waharday, Wibrin. Namur: Aische, Alle (RMNH). Andenne, Baillonville, Belgrade, Bidvre, Branchon, Champion, Eghezee. Emage, Feschaux. Gembloux, Gesves, Grand Leez, Ham-sur-Sambre, Lonzec, Mount-Gau- thier, Saint-Aubin, Saint-Gerard, Saint-Marc, Sorinnes, Sau- veniere, Sombreffe, Winenne. Bulgaria (1 <5): Rila Mts. (DEI). PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS. MEDIUM. AND MINUS 129 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 Figure 4. Trypoxylon figulus: geographic distribution in the New World. Czechoslovakia (1 9, 3 5): Jihomoravsky Kraj: Vranov W Znojmo (as Frain, NHMV). Vapadoslovensky Kraj: Sturovo (SMT). Denmark (41 9, 33 3, ZMK if not indicated otherwise): Al- borg: Vegger. Bornholm: Bastemose. Frederiksborg: Asserbo, Hillerod, Hundested, Jaegerspris Nordskov (tip of Horn- sherred peninsula), Sorte Mose near Farum, Store Karlsminde near Lynjes, Tisvilde Hegn (FSAG, ZMK). Holbsk: Kongsore. Maribo: Maribo. K£(benhavn: Holte. Odense: /Ebelo. Randers: Glatved Strand on Djursland Peninsula, Mols Bjerge. Ring- kflbing: Gindeskov, Kjelstrup (E Skjern). SBnderborg: Sonder- borg. Sor«: Agerso. Svenborg: Langeland Island: Hellenor. Tonder: Romo (VH), Stensbaek Plantage. Vejle: Klattrup. Finland: (47 9, 48 S, HY unless indicated otherwise): Abo: Lohja, Pernio (AKM), Rymattyla (AKM), Turku (AKM). Alandia: Eckero (AKM, H Y), Finstrom (AKM), Hammarland, Jomala (AKM), Saltvik (AKM). Karelia Borealis: Pyhaselka (Hammaslahti). Nylandia: Helsinki (CU, HY), Parvoo = Bor- gS, Pemaja. Ostrobotnia Australis: Koivulahti. Satakunta: Loimaa (AKM), Ylane (AKM). Tavastia Australis: Hattula, Hameenlinna, Janakkala, Nastola (AKM), Palkane (AKM, H Y), Somero (AKM), Urjala, Vanaja, Ylojarvi. France (35 9, 16 S): Alpes-Maritimes: Guillaumes (FSAG). Ariege: Ax-les-Thermes (ZMB). Basses-AIpes: Alios (FSAG), Annot (FSAG), Fugeret (FSAG), Les Dourbes (KMG), Mon- tagane de Lure (ZMK), Peyresq (FSAG), Saint-Andre-les-Alpes (FSAG). Bouche-du-Rhone: Marseille (FIS). Calvados: Lisieux (FSAG). Corse: Corte (KMG). C6tes-du-Nord: Saint-Rieul. Haute-Savoie: Dent d'Oche (MHNG), Mont Jorat (RMNH). Val de Charmy (RMNH). Haute-Vienne: Rochechouart (FSAG). Jura: Arbois. Loire-et-Cher: Blois (FSAG). Loire- Atlantique: Foret de la Roche Bernard (RMNH), Herbignac (RMNH). Saone-et-Loire: Uchizy (FSAG). Seine-et-Oise: Poissy (lEE). Var: Frejus (KMG), Gonfaron (FSAG), Montouroux (RMNH). Vaucluse: Carpentras (RMNH). Yonne: Foissy-sur-Vanne (FSAG). Germany, Democratic (81 9, 29 <5, DEI if not indicated oth- erwise): Berlin: Berlin (DEI, HD, ZMB, ZSM). Cottbus: Alt Dobem (ZMB), Muskau, Neu Zauchc (ZMB), Schlieben (ZMB). Dresden: Daubitz (SMT), Gersdorf near Kamenz (SMT). Er- furt: Erfurt (CU), Gotha (ZSM). Frankfurt: Biesental, Ebers- walde area. Gera: Blankenburg (ZMB), Jena (NHMV, ZMB). Halle: Gemrode (ZMH), Halle (DEI, ZMB), Kyffliauser, See- burg, Naumburg (TMB). Leipzig: Winkelmiihle. Magdeburg: Arendsee (SMT). Neubrandenburg: Faule Ort, Naturschutz- gebiet Muritzhof Potsdam: Furstenbcrg(TMB), Zechlin (ZMB), Zootzen. Rostock: Prerow, Rostock, Stralsund (DEI, ZMB). Rugen: Hiddensee Island (DEI, SMT), Rugcn Island: Monch- gut (SMT) and Ummanz, Schwerin: Campow (ZMB), Schwerin (ZMB), Wendeltorf near Schwerin. PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM. AND MINUS Germany, Federal (93 9. 51 S): Baden- VViirtemberg: Enz- klosterle (KS). Heidelberg (ZSM), Hochwacht (HW). Isny (NHMB), Kaiserstuhl (ZMB), Karlsruhe (KS, ZMH). Kussa- berg (KS), Radolfzell (ZMH), Schwarzwald (SMT), Tiengen in Wutach Valley (KS). Bayern: Abensberg(ZSM), Allach (ZSM). Aschaffenburg (FIS), Balderschwang (KS), Bamberg (ZSM), Ebenhausen (ZSM), Erdweg (ZSM), Horgersthausen near Moosburg (ZSM), Ingolstadt (ZSM), Kahl (FIS), Munchen (FSAG, ZSM), Numberg (ZSM), Rotwand area (ZSM), Schlier- see (ZSM), Tegemsee (ZSM). Hamburg (ZMH): Ochsenwarder, Warwisch. Hessen: Batlenfeld near Biederkopf (ZMH), Grun- dau E Frankfurt (HW, HY). Marburg (HW, WJP). Nieder- sachsen: 2 km NW Dotlingen (VH), Dorpen: 14 km SW Pa- penburg (VH). Elbe Islands (VH), 5 km S Oldenburg (VH), Pevestorf: 72 km SE Lauenburg (VH), Wobeck (ZMH). Nord- rhein-Westfalen: Ahaus (ZSM), Neheim (FSAG), Leverkusen (ZMH), Plettenberg (H W), Siegen (H W). Rheinland-Pfalz: low- er Ahr valley (FIS). Mainz (KS), Nattenheim (FSAG), Worms (FIS). Schleswig-Holstein: Amrum Island (VH), Eutin (KS). Ihlkathe 2 km SE Kiel (VH), Lutjenburg(KS), Ratzeburg (ZMH). Schierensee SW Kiel (VH), Schleswig (VH). Great Britain (92 2, 106 i: BMNH unless stated otherwise): Berkshire: Reading. Buckingham: Iver, Slough. Devon: Paign- ton. Dorset: Wareham. Essex: Brentwood, Colchester, Epping Forest. Gloucester: Chalford. Hampshire: Brockenhurst, Fleet, New Forest, Wickham. Isle of Wight: Sandown, Shanklin. Kent: Cobham, Darenth, Faversham (WJP), Goudhurst. London: Hampstead, Mill Hill. Mitcham Common, Norwood, Putney. Northampton: Ashton Wold (Oundle). Oxford: Goring, Ox- ford, Tubney near Oxford. Somerset: Dunster. Suffolk: Arger Fen, Bury St. Edmunds. Dunwich. Surrey: Byfieet, Esher, Hor- sell, Weybridge. Sussex: Midhurst: Ambersham Common. Greece (2 2, 4 S): Peloponnesus (de Beaumont, 1965): Mega Spilaion, Pirgos. and Taygetus. Sterea Ellas: Karpenission (KMG). Thessalia: Aspropotamos near Kalabaka (KMG). Hungary (13 2. 3 3): Bacs-Kiskun: Kalocsa (TMB). Tabdi (TMB). Gyor-Sopron: Neusiedlersee (NHMV). Somogy: Bal- atonszemes (TMB). Szolnok: Jaszbereny (TMB). Tolna: Si- montomya (NHMV). Veszprem: Tihany Peninsula on Balaton Lake (HD. TMB, WJP). Italy ( 1 9 2. 8 3): Emilia-Romagna: Cattolica (RMNH). Lom- bardia: Pavia: Cignolo Po (MSNM), Sondrio: Valtellina (KS). Piemonte: Alpignano (GP). Colle di Sestriere in Alpi Cozie (GP). Murazzano (GP). San Benedetto Belbo 20 km S Alba (GP). Val d'Angrogna in Alpi Cozie (WJP). Valle d'Aosta: Bresson near St. Vincent (GP). Venezia Giulia: Trieste (CU. NHMV). Trentino-Alto Adige: Bolzano (as Bozen, NHMV). Cavalese (BB). Collalbo (de Beaumont. 1959), Ortisei (NRS). Trafoi (NHMV). Japan (Tsuneki 1981): western Hokkaido (Esashi. Hakodate. Jozankei. Kamikawa) and central Hondo (Prefectures: Fukui. Ishikawa, Kyoto. Nagono. Niigata. Saitama, and Yamanashi). Specimens studied: 4 2, 4 <5: KT, TN. Morocco (1 2, 2 $): Asni. 1250 m alt. (paratype 2 oi figidus barbanun. RMNH). Fez (holotype offteuzeti, lEE), Marrakech (BMNH). Netheriands (108 2.73 3. RMNH if not indicated otherwise): Drenthe: Emmen. Erm. Havelte. Sellingen. Wijster. Zuidlaren. Gelderland: Apeldoom (FSAG). Bameveld. Hulshorst, Nij- kerk. Putten. Renkum. Vierhouten, Wageningen (GVR, RMNH). Groningen: Onnen. Limburg: Epen (GVR). Haelen. Heerien. Helden. Posterholt. Noord Brabant: Empel. Helen- aveen. Neerijnen. Udenhout. Ulvenhout. Noord Holland: Aer- denhout. Hilversum. Laren. Overijssel: Heino. Wezepe. Utrecht: Baarn. Bilthoven. de Bilt. den Dolder. Zuid Holland: Delft (GVR). Hillegersberg. Leiden. Leiderdorp, Loosduinen, Oeg- stgeest, Rijnsburg, Rotterdam, 's Gravenhage, Warmond. Norway ( 1 2, 1 3): Buskerud: Al (as Aal, DEI). Oppland: Lorn (RMNH). Poland (35 9. 24 3): Biafystok: Biatowieia (as Bialowies. FIS, ZMB). Leiikowo near Grajewo (ZMB). Bydgoszcz: Bydgoszcz (as Bromberg, SMT). Kielce: Gory Pieprzowe near Sandomierz (CAS), Kielce (TMB), Sandomierz (CAS). Lublin: Kazimierz on Wisla River (WJP). Warszawa: Podkowa Lesna (CAS). Wroclaw: Muszkowice near Henrykow (WJP), Osola: 25 km NW Wroclaw (WJP), Wroclaw (CAS. WJP). Szczecin: Bielinek (as Bellinchen. ZMB). Mi^dzyzdroje (as Misdroy. ZMB). Zie- lona Gora: Kostrzyn (as CUstrin, ZMB). Portugal (2 2. 2 3): Douro: Resende (RMNH). Estramadura: Lisboa (RMNH). Romania (8 2. 9 3, MGA if not indicated otherwise): Bacau: Lacul Bicaz 20 km W Piatra Neam^ (HD). Bra$ov: Sighi^oara, Tumisor near Sibiu. Bucure^ti: Brane§ti (WJP), Bude^ti. Con- stanta: Cri§an, Hagieni. Cri^ana: Ineu (as Borosjeno, TMB). Orasul Bucure§ti: Mogosoaia. Pantelimon. Timi^oara: Esel- ni^. Jupalnic near Mehedin^i. Mehadia (NHMV). Orsova (RMNH). Spain (53 2. 5 1 3): Alicante: Alicante (KMG). Orihuela (lEE) Avila (SFG): Becedas. Gilbuena. Puerto Castilla, Solana de Bejar, Tremedal. Barcelona: Canet de Mar (FSAG). Caceres (SFG): Bailos de Montemayor, Cabezuela del Valle, Gargan- tilla. La Garganta. Cadiz: Algeciras (NHMV). Ciudad Real: Ruidera (FJS). Granada: Salobrefla (VH). Huesca: Arguis (RMNH), Torla (FSAG), Valle de Ordesa. Lerida: Bohi, Lago di San Mauricio (FSAG). Logroilo: San Roman de Cameros (lEE). Madrid: El Escorial (lEE), Sierra de Guadarrama (lEE). Salamanca (SFG): Bejar, Cantagallo, Colmenar de Monte- mayor, El Cerro, La Cabeza de Bejar, Lagunilla, Montemayor del Rio. Navacarros, Palomaresde Bejar. Pefiacaballera, Pueb- la de San Medel, San Medel, Sorihuela, Valdehijaderos. Val- lejera de Rio Frio. Teruel: Albarracin (KMG, ZSM), Libros (FSAG). Toledo: Toledo (RMNH, WJP). Valencia: La Eliana (FSAG). Valladolid: Olmedo (FJS, WJP), Simancas (RMNH, WJP), Valladolid (FJS, WJP), Villa Baftez (FJS). Villalba de los Alcores (FJS). Zamora: Montamarta (FSAG). Sweden (43 2, 40 3. NRS if not indicated otherwise): Dalarna: Taktbo. Oland: Arontorp. Glomminge. Hogsrum. Halltorps hage, MorbylSnga, Repplinge, Vickleby. Ostergotland: St. Anna Korsnas, Simonstorp. Skine: Ahus, Halsingborg, Ha\ang. Hoor distr. (BMNH), Torekov (ZMK), Trolleholm, Vitemolla. Smi- land: Hagby, Jonkoping, Kalmar. KalmarSund (BIS Jungfrun). SoderSkra. Sodermanland: Haggenas. Malarhojden. Morto. Tullgam. Tyreso. Tyreso - Brevik. Uppland: Bjorko. HSbo - Tibbie. Osteraker. RSdmanso. Svartsjo. Varmdo. Vassunda. Switzeriand (78 2. 41 3. NHMB if not indicated otherwise): Bern: Adelboden. Bantiger(7 km NE Bern). Batterkinden. Bern. Biischiwald forest (5 km SSW Bern). Gadmen. Miirren. Signau. Spiez. Wachseldommoos bog (27 km SE Bern). Zweisimmen. Zwischenfluh (16 km SSW Thun). Geneve: Geneve (MHNG. NHMV), Genthod (NHMV), Peney (MHNG, NHMV). Grau- bunden: Chur (ZSM), Pare National Suisse (MHNG), Somvix, Versam. Luzern: Entlebuch Graben (CAS). Sankt Gallen: Vat- tis. Valais: Ausserbergd 1 km W Brig), Ayel -Zinal (RMNH), Berisal (BMNH, NHMB, NHMV). Chalet a Gobet (BMNH). Champery, Eusegne (as Usegne). Evolene. Hauderes. Inden. Lotschental. Martigny, Riederalp (8 km NNE Brig). Saas. Sankt Niklaus. Sierre (NHMV). Simplon (BMNH). Stalden. Verbier (BMNH). Vissoye. Zermatt. Vaud: Nyon (MHNG). 132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 Turkey (6 2, 3 6): Afyon: ^ay (FSAG). Ankara: Ankara (as Angora, TMB), Karagol (BMNH). Denizli: Saraykoy (as Seraj- Koj, TMB). Istanbul: Sile area (BMNH). Konya: Konya (JG). Kutahya: Karat Dagi (BMNH). Samsun: K6pruba§i (de Beau- mont, 1967). Tokat: Arguslu above Niksar (BMNH). Trabzun: Zigana Dagi (de Beaumont, 1967). USSR; Armenian SSR: Yerevan area (8 6. ZMMU). Azer- baydzhan SSR ( 1 9): Khanlar (as Helenendorf, NHMV). Ka- zakh SSR: Vostochnokazakhstanskaya obi.: foothills of Azu- tau Khrebet 18 km N Alekseyevka (1 d. Horeda. Jarsnas. Ljungarum. Sodermanland: Flisb\. Runmaro. Tullgam. Tyreso - Svartocken. Uppland: Harparbol. RSdmanso. Stockholm. Uppsala (HY). Vassunda. Vira bruk. Vasterbotten: Handene. Vastmanland: Sala. Switzeriand (21 2. 26 3. NHMB if not indicated otherwise): Bern: Batterkinden. Bern. Burgdorf Grauholz forest (up to 9 km NNE Bern). Wachseldoramoos bog (27 km SE Bern). En- gadin: Zuoz (.AWE). Geneve: Geneve Cologny (BMNH). Gen- thod (NHMV). Peney (MHNG). Graubunden: Pare National Suisse (de Beaumont. 1958), Somvix. Tessin: Lugano (FS.AG). Valais: Binntal (15 km NE Brig). Gnmentz (FS.AG. RMNH). St. Luc (RMNH). Simplon (BMNH). Zermatt (RMNH). Vaud: Nyon (MHNG). Zurich: Wadenswil. Turkey (1 2. 2 3): Artvin: Artvin. Berta (BMNH). Istanbul: Belgrat Orman (de Beaumont. 1967). Samsun: Bafra (BMNH). USSR: Georgian SSR: Bakuriani ( 1 2. ZMMU). Kazakh SSR: Chimkent. 2000 m (1 3. WJP); Razdolnyi area. 50°43N. 8 1°06'E ( 1 3. VLK). Zapovednik .Aksu-Dzhabagly ( 1 2. ZMMU). Kirgh- iz SSR: Arkit in Chatkalskiy Khrebet (1 3. ZMMU). Russian SSR: Karelian ASSR; ozero Tumas (as Kolatselka. 1 2. HY). Kemerovskaya oblast: 60 km SE Novokuznetsk (1 3, VLK). PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43, No. 10 Leningradskaya oblast: Kondratyevo W Vyborg (as Sakkijarvi, 1 $, H Y). Moskovskaya oblast: Mytishchi ( 1 6, ZMMU). Perm- skaya oblast: Nizhnyaya Kurya, 1 5 km W Perm (3 5, 2 S, WJP). Ukrainian SSR: Crimea: Krymskiy Zapovednik ( 1 $, ZMMU). Yugoslavia (4 9, 3 <5): Croatia: Plitvice (RMNH), Skrad (GP). Slovenia: Bled (RMNH), Cerknica (KS), Logatec (as Loitsch, ZMH), Opcina Mtn. on Italian border E Triest (NHMV). Ser- bia: Drazevac (FSAG). Trypoxylon medium de Beaumont Trypoxylon figidus var. medium de Beaumont, 1945:477, $, <5 (as media, incorrect original spelling). Holotype: 9, Swit- zerland: Martigny (Mus. Zool. Lausanne). — Bluthgen, 1951: 234 (var. media); de Beaumont, 1958:206 (forma media), 1959:30 (same); Wolf, 1959:15, 16 {figulus medium); Yalk- eila, 1961:144 (var. media); de Beaumont, 1964a:290, 1964b:84 (forma me^/a), 1965:56 (same), 1967:338 (same); Bohart and Menice, 1976:346 (ssp. medium); Lomholdt, 1976:267 {figulus media); Richards, 1980:45 (var. media); TsuNEKi, \9&\;\9 (medium). Trypoxylon figulus minus var. ruhi Wolf. 1959:15, 6. ! Lec- totype, 3, Federal Germany: Plettenberg (coll. H. Wolf Plet- tenberg), present designation. Synonymized by de Beaumont, 1964:291. Trypoxylon figulus minus: Wolf. 1959:fig. c. Diagnosis. — There is no single character by which medium can be distinguished from both figulus and minus. The anteroventral mesotho- racic carina is either straight or sinuate and curved posterad mesally, but unUke that of most mimis it is not expanded into a process. UnUke those of mosl figulus, the mesopleural setae around the scrobe are shorter than the midocellar diameter. In the female, the clypeal free margin is evenly concave between the orbit and median projec- tion (Fig. 7A), unlike that of minus and most figulus, and the setae of the hindcoxal pit form a curious channel-like structure visible only un- der high magnifications (Fig. 7C, D) and which has not been found in the other two species. The hindcoxal pit is circular (Fig. 7B) instead of ob- long (as it is in most figulus). The maximum length of male flagellomere X equals 0.75-0.9 of its basal diameter (the highest ratios are observed in specimens in which the flagellomere XI is the longest); the maximum length of flagellomere XI is usually 2.0-2.2 times its basal diameter (2.2-3.6 in figulus and minus). However, the fla- gellomere XI length is 2.4 times its basal di- ameter in some specimens, e.g., in a male from Horsell, England (KMG), or a male from Uilac, Romania; such specimens differ externally from figulus in having a shorter mesopleural vestiture and a longer flagellomere X. Gonoforceps with ventroextemal expansion at about midlength (Fig. 7E, F); expansion absent in figulus and minus. Body length 6.5-12 mm in female, 6.0-8.5 mm in male. Life History. — Several specimens of medium (seen by me) were reared from Rubus twigs (Wolf, 1959). A male was reared from an old gall of the chloropid fly Lipara lucens Meigen containing a vacated nest of the bee Hylaeus pectoralis Forster (England, Hampshire, Browndown, G. R. Else collector, BMNH). Geographic Distribution (Fig. 8). — Western and central Palearctic Region between Great Britain and eastern Kazakh SSR, north to south- em England and beyond the Arctic Circle in Fin- land, south to northern Mediterranean countries including Mallorca, Crete, and Cyprus, as well as Turkey, Syria, Israel, and northern Iran. Un- known from Iberian Peninsula and North Africa. Records. -Austria (30 9, 20 3, NHMV if not indicated oth- erwise): Burgenland: Andau (RMNH), Jois (JG), Donners- kirchen an Neusiedlersee (KS), Neusiedl (KS, NHMV, ZMB), Panzergraben an Neusiedlersee (JG), Winden (JG), Zumdorf (AWE, RMNH). Niederosterreich: Bisamberg near Vienna, Deutsch Altenburg (ZMB), Donauauen near Vienna (ZMH), Hamburg (ZMB), Herzograd (JG), Marchfeld (JG), Oberwei- den (DBB), Piesting, Schneeberg, Stammersdorf (DBB), Wien- Tiirkenschanze. Oberosterreich: Gutau (JG). Salzburg: Kat- chenberghohe (VH), Salzburg (BMNH). Tirol: Galtiir (BMNH), Huben (ZMB), Innsbruck, Kraspes (DEI), I km SW Zwiesel- stein. Voralberg: Bielerhohe. Belgium (II 9. 17 <5, FSAG): Antwerpen: Geel. Liege: Liege, Montzen, Spa, Wandre. Limburg: Bree, Eben, Lummen, Lux- embourg: Chatillon, Les Epioux, Torgny. Namur: Eprave, Fer- age, Mont-Gauthier. Bulgaria (1 9): Ruse (SMT). Cyprus (3 2, 6 6); Amathus (FSAG, RMNH), Paphos (KMG). Czechoslovakia (1 9, 2 6); Jihocesky Kraj: Prachatice (as Prachatitz, NHMV). Vapadoslovensky Kraj: Sturovo (SMT). Denmark (1 1 9, 8 3, ZMK, if not indicated otherwise): Born- holm: Amager. Frederiksborg: Hulerod, Humblebak (S Hel- singor), Tisvilde Hegii. Hjurring: Fauerholt Hede (ca 10 km W Frederikshavn), Skoven on Lsese Island. Maribo: Kris- tiansssde Skov on LoUand Island. Randers: Glatved Strand and Kale on Djursland Peninsula, Mols Bjerge. Ribe: Bors- mose. Ho Plantage. Svendborg: Keldsnor on Langeland Island. Thisted: Bagso at Nors (ca 7 km NW Thisted). T«nder: Rome Island (RMNH), Stensbjeck Plantage. Finland (18 9, 15 'giophore, which is complete. In the dorsal fin the situation is more complicated, and the fin ray proximal radial Figure 1 8. Thryssocypris smaragdinus. 53.2 mm, posteriormost fin rays and pterygiophores in dorsal fin (lateral view above, dorsal view below). ROBERTS & KOTTELAT: THRYSSOCYPRIS. A NEW CYPRINID GENUS homology of the bifurcate terminal radial with the proximal, medial, and distal radials is un- clear. In T. smaragdimis the bifurcate terminal radial constitutes by itself the terminal pteryg- iophore, articulating endochondrally only with the medial radial of the preceding pterygiophore. In T. tonlesapensis an additional element is pres- ent ventral to the bifurcate terminal radial; the dorsal margin of this element articulates with the ventral margins of the bifurcate terminal radial and of the preceding medial and proximal ra- dials. and its homology also is unclear. Intermuscular Bones. — Intermuscularbones well-developed, epineurals extending entire length of body, epipleurals commencing at vertebra 12. Proximal ends of anteriormost two or three epineurals attached to exoccipital in a small depression or fossa (shown in Fig. 8) near its posterolateral margin and on a level with fora- men magnum. Distal ends of posteriormost two or three epineurals and epipleurals multifid (mostly bifid or trifid). Epineurals and epipleur- als forked anteriorly with expanded laminae pos- teriorly. In chelines and cultrines the anteriormost epi- neurals tend to insert much higher on the cra- nium (Howes 1979) than in Thryssocypris. Caudal Skeleton (Fig. 19).— Generalized in nearly all respects, with a single epural; one or two uroneurals; parhypural fused to ural cen- trum, with an exceptionally large, elongate hy- purapophysis; six hypurals, separate from each other, with hypurals 1-2 fused to ural complex centrum. In most lower teleosts with the primitive prin- cipal caudal fin ray formula of 10 + 9, including all characoids and all Cyprinidae, the caudal fin skeleton tends to be highly conservative, retain- ing a relatively generalized or primitive mor- phology, including the primitive complement of six hypurals (exclusive of the parhypural). There is a maximum of three epurals, and perhaps in- variably at least one epural. Presumably three is the primitive number of epurals in teleosts in- cluding ostariophysans, but reductions to two and one and subsequent reversion to three seem to have occurred repeatedly, particularly often in ostariophysans including Cyprinidae. Thus the single epural in Thryssocypris is not likely to be helpful in assessing its relationships to other cyp- rinids. It should be noted that previous works on os- uroneural epural urostyle neural spine ural centrum hypurapophysis parhypural hemal spine hypurals 1-6 Figure 19. Thryssocypris smaragdimis. 53.2 mm. caudal fin skeleton (lateral view). tariophysan caudal skeletons report seven hy- purals (e.g., Weitzman 1962; Roberts 1969). Fol- lowing Monod (1968) the element previously considered as hypural one is here termed the parhypural. The parhypural bears the hypura- pophysis which serves as origin of the main muscles for adduction of the upper caudal fin lobe. Thus the hypurals formerly numbered 2-7 are now hypurals 1-6. This new nomenclature has been used by most authors dealing with os- tariophysan caudal skeletons subsequent to Monod (1968) including Lundberg and Baskin (1969) and Fink and Fink (1981). Conclusion The discovery of Thryssocypris provides another example of the extraordinary diversity of Cyprinidae, especially in Southeast Asia, and of the close biogeographic relationship between the Kapuas and Mekong basins. Upon first examining Thryssocypris we thought that it might be a specialized Barilius or at least closely related to that genus. Lacking osteological information on most other cyprinid genera we are not prepared to discuss its relationships at this time except to note that it does not seem to be related to the chelines (sensu Howes 1979). The specialized characters in Thryssocypris that seem most unusual or highly derived and there- 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43. Nc fore likely to indicate phyletic relationships have not been reported in any other cyprinids. These include the peculiar morphology of ceratobran- chials and absence of hypobranchials 2-3, shape of jaw bones, elongate anterior and posterior pro- jections of ischiac process, and bifid terminal radials in median fins. Acknowledgments For making available material of Thryssocy- pris in their care we wish to thank J. Daget and F. d'Aubenton and W. J. Rainboth. The study was also facilitated in various ways by the fol- lowing persons: G. J. Howes, Michael A. Hearne, W. N. Eschmeyer, Daphne Dunn, and Vincent Lee. The Kapuas ichthyological survey was sponsored by the Smithsonian Tropical Research Institute, Indonesian National Research Coun- cil, and Museum Zoologicum Bogorense. This paper is part of a project to report on the fishes of the Kapuas basin supported by National Sci- ence Foundation grant DEB77-24759. Literature Cited Fink, S. V., and W. L. Fink. 1981. Interrelationships of ostariophysan fishes (Teleostei). Zool. J. Linn. Soc. 72(4): 297-353. GosLiNE, W. A. 1975. The cyprinid dermosphenotic and the subfamily Rasborinae. Occ. Pap. Mus. Zool. Univ. Michi- gan, no. 673, 13 pp. Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Amer. Mus. Nat. Hist. 131(4):339-456. Hasslir, R. L. 1970. Studies on the osteology of catfishes. order Siluriformes. Ph.D. dissertation, Stanford Univ.. 133 pp. Howes, G. J. 1978. The anatomy and relationships of the cyprinid fish Luciohrama macrocephalus (Lacepfede). Bull. Brit. Mus. (Nat. Hist.), zool. ser. 34(1): 1-64. . 1979. Notes on the anatomy of A/afrac/!/>/c/!/M'5 macrochirus (Valenciennes) 1844, with comments on the Cultrinae (Pisces, Cyprinidae). Bull. Brit. Mus. (Nat. Hist.), zool. ser. 36(3): 147-200. . 1980. The anatomy, phylogeny and classification of bariliine cyprinid fishes. Bull. Brit. Mus. (Nat. Hist.), zool. ser. 37(3): 1 29-1 98. Lekander, B. 1949. The sensory line system and the canal bones in the head of some Ostariophysi. Acta Zool. 30:1- 131. Llindberg, J. G., and J. N. Baskin. 1969. The caudal skel- eton of the catfishes, order Siluriformes. Amer. Mus. Nov. no. 2399, 49 pp. MoNOD, T. 1968. Le complex urophoredesteleosteens. Mem. Inst. fr. Afr. noire 81:1-705. Ramaswami, L. S. 1955. Skeleton of cyprinoid fishes in re- lationship to phylogenetic studies: 7. The skull and Weberian apparatus of Cyprininae (Cyprinidae). Acta Zool. 36:199- 242. Roberts, T. R. 1969. Osteology and relationships of chara- coid fishes, particularly the genera Hepsetiis. Salminus. Ho- plias. Ctenolucius. and Acestrorhynchus. Proc. California Acad. Sci. 36(15):39 1-500. . 1973. Interrelationships of ostariophysans, pp. 373- 395 in P. H. Greenwood, R. S. Miles, and C. Patterson (eds.). Interrelationships of Fishes, London, Academic Press. Weitzman, S. H. 1962. The osteology of Brycon meeki. a generalized characid fish, with an osteological definition of the family. Stanford Ichth. Bull. 8:1-77. PROCEEDINGS OF THE Marine 8io?ogic?l Ishomtor^ 'UL 2 7 1984 CALIFORNIA ACADEMY OF SCIENCES Vol. 43, No. 12, pp. 159-177, 32 figs. c?::s Hcie. Mas- July 12, 1984 STUDIES ON NEBRIINI (COLEOPTERA: CARABIDAE), V. NEW NEARCTIC NEBRIA TAXA AND CHANGES IN NOMENCLATURE By David H. Kavanaugh Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Abstract: This paper introduces new Nearctic Nebria taxa and changes in nomenclature. Names are pro- vided for 10 new species and 3 new subspecies (type locality in parentheses): Nebria altisierrae (Olmsted Point, Yosemite National Park, California), A', campbelli (Mount Baker, Whatcom Count)', Washington), A^. wallowae (West Fork Wallowa River, Wallowa County, Oregon), N. jejfreyi (South Fork McCoy Creek, Harney County, Oregon), N. haida (Mount Needham, Graham Island, Queen Charlotte Islands, British Columbia), N. louiseae (Skedans, Louise Island, Queen Charlotte Islands, British Columbia), A', gebleri albimontis (Birch Creek, Mono County, California), N. labontei (West Fork Wallowa River, W allowa County, Oregon), A', calva (Mount Baldy, Apache County, Arizona), A^. sierrablancae (Sierra Blanca, Lincoln County, New Mexico), A. piute sevieri (Parowan Creek, Iron County, Utah), A', steensensis (South Fork McCoy Creek, Harney County, Oregon), and N. trifaria pasquineli (Lefthand Creek, Boulder County, Colorado). For each, diagnostic combination of characters and notes on geographical distribution are provided and distin- guishing features are illustrated. Changes in status are proposed for the following names (second name in each pair considered valid): Nebria intermedia Van Dyke = N. crassicornis intermedia Van Dyke; N. sonorae Kavanaugh = N. acuta sonorae Kavanaugh; N.fragilis Casey = A. arkansana fragilis Casey; N. trifaria piute Erwin and Ball = N. piute piute Erwin and Ball; and N. trifaria utahensis Kavanaugh = N. piute utahensis Kavanaugh. New synonymies proposed include: Nebria arkansana uinta Kavanaugh = N. arkansana fragilis Casey; N. fragilis teewinot Kavanaugh = N. arkansana fragilis Casey; and N. trifaria tetonensis Erwin and Ball = A', trifaria trifaria LeConte. Introduction For several years, I have been working on a monographic treatment of genus Nebria Latreille for the Nearctic Region. During that time, I have provided names for several new species and nu- merous new subspecies, designated lectotypes, and proposed certain nomenclatural changes (Kavanaugh 1979 and 1981). Validation of these new names and clarification of the status of ex- isting names were needed to permit their proper use in various other reports by the author and several colleagues. Since 1981, additional spec- imens and data have been acquired. Study of this new material has revealed 1 3 hitherto unknown taxa (10 species and 3 subspecies) as well as sev- eral nomenclatural problems with previously de- scribed taxa. The purpose of this report, which serves as a final presentation of nomenclatural matters pre- liminary to submission of the monographic treat- ment, is to provide names for the new taxa and to present formally the needed nomenclatural changes. The latter include both new synonymies and other changes in status of names. As before. [159] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 these names are needed immediately for use in other manuscripts; and data and discussions pre- sented for each name are Umited to httle more than the minimum required by the International Code of Zoological Nomenclature. More detailed information for all taxa, including those pre- sented here as new, will be provided in the mono- graph. Materials and Methods This report is based on examination of 1 1 36 adult Nebria specimens, representing taxa de- scribed here as new, and over 80,000 additional specimens, representing previously described Nebria taxa, used for comparative purposes. Recognition of new synonymies and changes in status of taxa are based on study of this total specimen resource pool. Following is a list of acronyms used in the text. These refer to collections from which specimens have been received and/or in which paratype specimens have been deposited. Curators re- sponsible for collections during the course of my study are also listed; and I here acknowledge with sincere thanks their assistance in providing spec- imens for study on loan. ANSP— Academy of Natural Sciences, Philadelphia, Pennsyl- vania 19103; W. W. Moss. BCPM — British Columbia Provincial Museum, Victoria, Brit- ish Columbia V8V 1X4; R. A. Cannings. CArm-C. Armin, 191 West Palm Avenue, Reedley, Califor- nia 93654 (specimens deposited in CAS). CAS— California Academy of Sciences, San Francisco, Cali- fornia 941 18; D. H. Kavanaugh, H. B. Leech. CNC— Canadian National Collection of Insects, Biosystem- atics Research Institute, Ottawa, Ontario KIA 0C6; R. de Ruette, A. Smetana. CUB -University of Colorado, Boulder, Colorado 80302; H. Rodeck. CUIC- Cornell University, Ithaca, New York 14850; L. L. Pechuman. DEUN— University of Nebraska, Lincoln, Nebraska 68503; B. C. Ratcliffe. EAMa— E. A. Martinko, University of Kansas, Lawrence, Kansas 66044 (specimens deposited in CAS). FMNH — Field Museum of Natural History, Chicago, Illinois 60605; H. Dybas, L. Watrous. JRU-J. R. La Bonte, 7 10 NW 1 1th, Corvallis, Oregon 97330. KSUC— Kansas State University, Manhattan, Kansas 66502; H. D. Blocker. LACM — Los Angeles County Museum of Natural History, Los Angeles, California 90007; C. L. Hogue. MCZ— Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, Massachusetts 02138; J. F. Lawrence, A. F. Newton, Jr. MSU— Michigan State University, East Lansing, Michigan 48823; R. L. Fischer. OSUC-Ohio State University, Columbus, Ohio 43210; C. A. Triplehom. PJJo-P. J. Johnson, 1408 28th Street SE, Auburn, Washing- ton 98002. QCIM— Queen Charlotte Islands Museum, Skidegate, British Columbia VOT ISO; N. Gessler and T. Gessler. ROM — Royal Ontario Museum, Toronto, Ontario M5S 2C6; G. B. Wiggins. RTBe-R. T. Bell, University of Vermont, Burlington, Ver- mont 05401. UAFA— University of Arkansas, Fayetteville, Arkansas 72701; R. T. Allen. U ASM — University of Alberta, Strickland Museum, Edmon- ton, Alberta T6G 2E3; G. E. Ball. UMMZ— University of Michigan, Ann Arbor, Michigan 48 104; R. D. Alexander. USNM — United States National Museum, Smithsonian Insti- tution, Washington, D.C. 20560; T. L. Erwin. UWEM — University of Wisconsin, Madison, Wisconsin 53706; L. J. Bayer. ZMLS— Zoological Institute, University of Lund, Lund, Swe- den; C. H. Lindroth, R. Danielsson. Methods used in the present study, including measurement and dissection techniques and cri- teria for ranking taxa as species or subspecies, have been described in a previous paper (Ka- vanaugh 1979). New Nebria Species and Subspecies The order of presentation of new species and subspecies below reflects a new classification of Nearctic Nebria, which will be provided in a monographic treatment of the genus now in preparation for publication. Except as noted, all specimens listed (by locality) in sections about geographical distribution have been designated as paratypes. Nebria altisierrae, new species (Figures 2, 15, 31) Nebria virescens; Kavanaugh 1978:345 (in part). HoLOTYPE, 6, in CAS, labelled: "U.S.A., California, Yosem- ite N. P., Sierra Nevada, Olmsted Point, 1.5 mi. sw. Tenaya Lk. on Tioga Rd., 2560m, 8 Nov. 1976 D. H. Kavanaugh"/ "D. H. Kavanaugh Collection" [orange labell/"Holotype Ne- bria altisierrae n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 14338." Paratypes: five (four 6 and one 9), also deposited in CAS. Type-Locality.— Olmsted Point, 2560 m. Sierra Nevada, Yosemite National Park, California. Diagnostic Combination. — Head uniformly dark, without pale spots on vertex; labium with paraglossae distinct as short, pointed lateral lobes on apical margin of ligula; pronotum (Fig. 2) semiovoid, relatively short and moderately wide, ratio pronotal width to elytral width less than or equal to 0.75, basal sinuation of lateral margin KAVANAUGH: NEW NEARCTIC NEBRIA 161 absent or short and very shallow, basal angles not or only moderately denticulate, midlateral seta present; elytra with silhouette subrectan- gular, not narrowed basally, humeri (Fig. 1 5) not or only faintly carinate; metepisternum impunc- tate; hind tarsus with all tarsomeres glabrous dorsally, fourth tarsomere truncate ventrally with medial and lateral apicoventral setae symmet- rical in length and position; specimen from lo- cality in Sierra Nevada of California (Fig. 31). Derivation of Taxon Name.— The species epithet is a combination of the Latin word for "high" {=altus) and a shortened form of "Sierra Nevada," in reference to the High Sierra region inhabited by members of this species. Geographical Distribution. — Figure 31. Known only from high elevations in the Sierra Nevada of California, from Tuolumne County south to Sequoia National Park. I have studied specimens from the following localities: United States of America California: Sequoia National Park, Alta Meadow ([2740 m]) [Aug.] (2; CAS); Tuolumne County, Blue Canyon Lake ([3110 m]) [July] (1; CAS); Yosemite National Park. Olmsted Point (1.5 miles SW of Tenaya Lake [2560 m]) [Nov.] (3; CAS). Nebria campbelli, new species (Figures 3, 16, 31) Nebria virescens; Kavanaugh 1978:345 (in part). HoLOTYPE, 3, in CAS, labelled: "U.S.A., Wash., Whatcom Co., Cascade Range, ne. slope Mt. Baker, Kulshan Ridge, 1460m-1520m, 11 Aug. 74 D. H. Kavanaugh"/ "D. H. Ka- vanaugh Collection" [orange label]/ "Holotype Nebria camp- belli n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 14339." Paratypes: two (one 3 and one 2) deposited in CAS and CNC. Type- Locality. — Mount Baker, 1460-1520 m. Cascade Range, Whatcom County, Washington. Diagnostic Combination. — Head uniformly dark, without pale spots on vertex; labium with paraglossae distinct as short, pointed lateral lobes on apical margin of ligula; pronotum (Fig. 3) semiovoid, relatively long and narrow, basal sin- uation of lateral margin short and moderately deep, basal angles markedly denticulate laterally, midlateral seta present; elytra with silhouette narrowed basally, nearly subovoid, humen (Fig. 16) markedly carinate; metepisternum impunc- tate; hind tarsus with all tarsomeres glabrous dorsally, fourth tarsomere truncate ventrally with medial and lateral apicoventral setae symmet- rical in length and position. Derivation of Taxon Name. — I take plea- sure in naming this species in honor of my friend and colleague J. Milton Campbell, who collected the only known female specimen of this species. Geographical Distribution. — Figure 31. Known only from the northern end of the Cas- cade Range and its flanking ranges, from Man- ning Provincial Park in southern British Colum- bia to Mount Baker in northern Washington. I have studied specimens from the following lo- calities: Canada BRITISH COLUMBIA: Manning Provincial Park (Three Brothers Mountain [2130 m]) [July] (1; CNC). United States of America WASHINGTON: Okanogan County, Pasayten Wilderness (Bunker Hill Lookout [2120 m]) (1; CAS); Whatcom County, Mount Baker (NE slope on Kulshan Ridge [1520 m]) [Aug.] (1;CAS). Nebria wallowae, new species (Figures 4, 17, 24, 31) Holotype, 3. in CAS, labelled: "U.S.A., Oregon, Wallowa County, Wallowa Mts., West Fork Wallowa River, 2070- 2130m, 10 July 1982 Stop #82-16 D.H. & J. L. Kavanaugh colls."/ "D. H. Kavanaugh Collection" [orange label]/ "Ho- lotype Nebria wallowae n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 1 4347." Paratypes: 87 (36 3 and 5 1 2), deposited in CAS, CNC, JRLa. UASM, and USNM. Type-Locality. -West Fork Wallowa River. 2070-2 1 30 m, Wallowa Mountains, Wallowa County, Oregon. Diagnostic Combination. — Pronotum (Fig. 4) with lateral explanation broad throughout, basal sinuation of lateral margin moderately deep, basal angles rectangular or slightly acute, mod- erately projected posteriorly, midlateral seta ab- sent; elytra with silhouette subovoid, slightly narrowed basally, humeral angles not markedly rounded or obtuse, humeral carinae moderately developed, slightly projected anteriorly; hind- wing full-sized, with reflexed apex distal to stig- ma; hind coxae bi- or plurisetose basally; middle tibiae dorsally concave or sulcate (at least near middle), with brush of dorsal setae moderately and densely developed; third to fifth visible ab- dominal sterna each with two or more pairs of posterior paramedial setae; median lobe of male (Fig. 17) long, very slender; bursa copulatrix of female (Fig. 24) with very small bursal sclerite; specimen from locality in Wallowa Mountains of Oregon (Fig. 3 1 ). Derivation of Taxon Name.— This species is named for the Wallowa Mountains. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 Geographical Distribution. — Figure 31. Known only from the Wallowa Mountains of northeastern Oregon. I have studied specimens from the following localities: United States of America Oregon: Wallowa County, Glacier Lake (meadow below [2440 m]) [Sep.] (2; JRLa), Mirror Lake area ([2350 m]) [July] (5; CNC), West Fork Wallowa River (0.25 to 0.5 miles NE of Frazier Lake [2040-2190 m], and small side stream [1980 m]) [July] (81; CAS, JRLa). Nebria jefTreyi, new species (Figures 5, 31) HoLOTYPE, S, in CAS, labelled: "U.S.A., Oregon, Harney County, Steens Mountains, South Fork McCoy Creek, 2290- 2560m, 14 July 1982, Stop #82-22, D. H. & J. L. Kavanaugh colls."/ "D. H. Kavanaugh Collection" [orange label]/ "adult specimen used in laboratory rearing of immature stages"/ "Ho- lotype Nebria jefTreyi n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 14342." Paratypes: 35 (1 1 <5 and 24 9), deposited in CAS, JRU, PJJo, UASM, and USNM. Type-Locality. -South Fork McCoy Creek, 2390-2560 m, Steens Mountains, Harney County, Oregon. Diagnostic Combination.— Pronotum (Fig. 5) with lateral explanation broad throughout, apical angles very broad and rounded, basal an- gles rectangular, basal sinuation of lateral margin moderately deep, midlateral seta absent; elytra with silhouette subrectangular, not narrowed ba- sally, humeral angles not markedly rounded or obtuse, humeral carinae absent or only slightly developed, not projected anteriorly; hind coxae bi- or plurisetose basally; middle tibiae dorsally concave or sulcate (at least near middle), with brush of dorsal setae moderately and densely de- veloped; third to fifth visible abdominal sterna each with two or more pairs of posterior para- medial setae; specimen from locality in Steens IVIountains of Oregon (Fig. 3 1 ). Derivation of Taxon Name. — I take great pleasure in naming this species in honor of my son, Jeffrey L. Kavanaugh, who assisted me in collecting the first known specimens of this species. Geographical Distribution. — Figure 31. Known only from the Steens Mountains of southcentral Oregon. I have studied specimens from the following localities: United States of America Oregon: Harney County, Little Blitzen River ([2560 m]) [July] (10; PJJo), South Fork McCoy Creek ([2390-2560 m]) [July] (26; CAS, JRLa). Nebria haida, new species (Figures 6, 31) HoLOTYPE, S, in CAS, labelled: "Canada, British Columbia, Queen Charlotte Islands, Graham Island, 1.8 km N of Mt. Needham, 700m-780m, 18 July 1981, Stop #81-37, D. H. Kavanaugh collector"/ "D. H. Kavanaugh Collection" [orange label]/ "Queen Charlotte Islands Expedition— 1981" [row of asterisks] "D. H. Kavanaugh Calif Acad. Sciences"/ "Holo- type Nebria haida n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 14341." Paratypes: 151 (68 3 and 83 9), deposited in BCPM, CAS, CNC, QCIM, UASM, and USNM). Type- Locality. -1.8 km N of Mount Needham, 700-780 m, Graham Island, Queen Charlotte Islands, British Columbia. Diagnostic Combination. — Size medium: standardized body length of male 9.0 to 10.5 mm, of female 9.4 to 10.8 mm; head relatively large and wide; pronotum (Fig. 6) relatively long and slender, with lateral explanation broad throughout, apical angles short, relatively broad and slightly rounded, basal angles rectangular, basal sinuation of lateral margin long, moder- ately deep, midlateral seta absent; elytra with silhouette subovoid, long and slender, distinctly narrowed basally, humeral angles not markedly rounded or obtuse, humeral carinae absent or only slightly developed, not projected anteriorly, intervals markedly convex; legs long, slender, femora and tibiae piceous; hind coxae bi- or plurisetose basally; middle tibiae dorsally con- cave or sulcate (at least at middle), with brush of dorsal setae moderately and densely devel- oped; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; specimen from locality in Queen Charlotte Islands, British Columbia (Fig. 31), from above treeline in alpine area. Derivation of Taxon Name.— This species is named in honor of the Haida people, tradi- tional inhabitants of the Queen Charlotte Ar- chipelago. Geographical Distribution. — Figure 31. Known only from high elevations in the Queen Charlotte Islands, British Columbia. I have stud- ied specimens from the following localities: Canada British Columbia: Queen Charlotte Islands: Graham Is- land, Mount Needham (1.0 km [790-910 m] and 1.8 km [700- 780 m] N) [July] (148; CAS); Moresby Island, Mount Moresby (northwest-facing cirque [910-1070 m]) [July] (4; CAS). Nebria louiseae, new species (Figures 7, 31) Holotype, 6, in CAS, labelled: "B. C, Q. C. I. Louise Is., KAVANAUGH: NEW NEARCTIC NEBRIA 163 Skedans 1 1. VI. 1981 R. A. Cannings"/ "Holotype Nebria lou- iseae n. sp. det. D. H. Kavanaugh 1 983" [red label]/ "California Academy of Sciences Type No. 15005." Paratypes: four S, deposited in BCPM, CAS, and CNC. Type-Locality. — Skedans, Louise Island, Queen Charlotte Islands, British Columbia. Diagnostic Combination. — Size large: stan- dardized body length of male 10.4 mm, of female 10.6 to 1 1.0 mm; head relatively large and wide; pronotum (Fig. 7) relatively long and slender, with lateral explanation broad throughout, apical angles moderate in length, relatively narrow and pointed, basal angles rectangular, basal sinuation of lateral margin long, moderately deep, midlat- eral seta absent; elytra with silhouette subovoid, long and slender, distinctly narrowed basally, hu- meral angles not markedly rounded or obtuse, humeral carinae absent or only slightly devel- oped, not projected anteriorly, intervals mark- edly convex; legs long, slender, femora and tibiae piceous; hind coxae bi- or plurisetose basally; middle tibiae dorsally concave or sulcate (at least at middle), with brush of dorsal setae moderately and densely developed; third to fifth visible ab- dominal sterna each with two or more pairs of posterior paramedial setae; specimen from lo- cality in Queen Charlotte Islands, British Colum- bia (Fig. 31), from upper sea beach area. Derivation of Taxon Name. — This species is named for Louise Island, on which the type locality is found. Geographical Distribution. — Figure 31. At present known only from Louise Island, Queen Charlotte Islands. I have studied specimens from the following locality: Canada British Columbia: Queen Charlotte Islands: Louise Island, Skedans [June] (5; BCPM). Nebria gebleri albimontis, new subspecies (Figures 1, 8, 18, 25, 32) HoLOTYPE, 3, in CAS, labelled: "U.S.A., California, Mono County, White Mts., Birch Creek, 3290m-3410m, 8 July 1980 D. Giuliani collector"/ "Collection of California Academy of Sciences, San Francisco, Calif"/ "Holotype Nebria gebleri al- bimontis n. ssp. det. D. H. Kavanaugh 1 983" [red label]/ "Cal- ifornia Academy of Sciences Type No. 14340." Paratypes: four (two & and two 2), also deposited in CAS. Type-Locality. -Birch Creek, 3290-3410 m. White Moun- tains, Mono County, California. Diagnostic Combination. — Head dark, with a pair of pale paramedial spots on vertex; anten- nal scape (Fig. 1) short, moderately thick; prono- tum (Fig. 8) with basal angles markedly acute, distinctly divergent posteriorly, margination of apical angles and anterior one-third of lateral margin very narrow, midlateral and basolateral setae present; elytra without metallic reflection, elytral silhouette subrectangular, hindwing full- sized; median lobe of male (Fig. 1 8) with pre- apical area markedly bulbous left dorsolaterally; bursa copulatrix of female (Fig. 25) with sper- mathecal chamber small, narrow in dorsal as- pect; specimen from locality in White Mountains of California (Fig. 32). Derivation of Taxon Name.— The subspe- cific epithet is a combination of the Latin words for "white" {=albus) and "mountain" {=mons), in reference to the White Mountains. Geographical Distribution. — Figure 32. Known only from the type locality in the White Mountains of eastern California. I have studied specimens from the following locality: United States of America California: Mono County, Birch Creek ([3290-3410 m]) [July] (5; CAS). Nebria labontei, new species (Figures 9, 32) Holotype, S, in CAS, labelled: "U.S.A., Oregon, Wallowa County, Wallowa Mts., West Fork Wallowa River, 2040- 2190m, 11 July 1982, D. H. & J. L. Kavanaugh Stop #82- 17"/ "D. H. Kavanaugh Collection" [orange label]/ "adult specimen used in laboratory rearing of immature stages"/ "Ho- lotype Nebria labontei n. sp. det. D. H. Kavanaugh 1983" [red label]/ "California Academy of Sciences Type No. 14343." Paratypes: 57 (29 3 and 28 9), deposited in CAS, JRLa, UASM, and USNM. Type-Locality. - West Fork Wallowa River, 2040-2 1 90 m, Wallowa Mountains, Wallowa County, Oregon. Diagnostic Combination. — Size very large: standardized body length of male greater than 11.5 mm, of female greater than or equal to 1 2.0 mm; head moderate in size, dark, with a pair of pale paramedial spots on vertex; pronotum (Fig. 9) broad, markedly cordate, midlateral and ba- solateral setae present; elytra with brilliant red metallic reflection, elytral silhouette distinctly subovoid, narrowed basally with lateral margins distinctly rounded, intervals flat; specimen from locality in Wallowa Mountains of Oregon (Fig. 32). Derivation of Taxon Name. — I am pleased to name this species in honor of my friend and fellow collector, James R. LaBonte, who col- lected the first known specimen of this extraor- dinary species. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43. No. 12 Geographical Distribution. — Figure 32. Known only from the Wallowa Mountains of northeastern Oregon. I have studied specimens from the following localities: United States of America OREGON: Wallowa County, Glacier Lake (meadow below [2440 m]) [Sep.] ( 1 ; JRLa), West Fork Wallowa River (0.25 to 0.5 miles NE of Frazier Lake [2040-2190 m]) [July] (57; CAS, JRLa). Nebria calva, new species (Figures 10, 19, 26, 32) HoLOTYPE, -S\J 36025). In this spec- imen breeding tubercles occur on the anal, pec- toral and pelvic fins, abdominal keel, and head. The strong lateral projections on the anterior face of the first nine branched anal fin-rays are en- tirely or almost entirely covered by a thickened, longitudinal band of thickened skin 9 mm long and 1.2 mm high. The surface of this spongy band of skin is covered with hundreds of small, overlapping, scale- or leaflike breeding tuber- cles, with their raised free margins projecting an- teriorly. There are about 1 2-20 of these tubercles in a vertical series. Discrete pads of similarly thickened skin covered with similar breeding tu- bercles extend obliquely posteroventrally on the basal third of the first five branched anal fin-rays. There are up to eight tubercles across each ray. The skin on the middle third of the same rays appears to be only slightly thickened and bears only a few, small widely spaced, low-lying round (not scalelike) tubercles. The distal third or branched portion of the first eight branched rays is covered with thick skin densely coated with scalelike tubercles. There are up to about eight tubercles across each ray-branch. The leading edge of the third (enlarged) simple anal fin-ray bears a thick, lamellar projection of skin, 1 1.5 mm long and up to 2.2 mm wide, covered with widely scattered, low-lying round tubercles without free margins. The midventral abdominal keel is also notably thickened, and covered with minute, closely spaced round or granular tubercles which extend for a short distance onto the abdomen and sides of the body just anterior to the anal fin. The pelvic and pectoral fins bear round tu- bercles dorsally and ventrally; these are most no- ticeable on the enlarged outermost pectoral fin- ray. The dorsal fin is slightly tuberculate, the adipose and caudal fins non-tuberculate. The dorsal, lateral, and ventral surfaces of the head bear irregularly scattered, round, low-lying tu- bercles without free margins. These are largest and most numerous on its ventral surface. The skin of the oral margin of the upper and lower jaws and gular margin of the lower jaw is thick- ened and tuberculate. Fine granular projections, which may be minute breeding tubercles, extend in a dorsomedian longitudinal band from the dorsal fin origin anteriorly halfway to the occi- put. In the two gravid females the skin is less modified, and although tuberculation is very much lighter, there are small, low-lying round tubercles on the anal, pelvic, and pectoral fins and on the head. In one of them the skin on the jaws is thickened as in the male; in the other it is not. The first female has the median abdominal fold somewhat thickened, suggestive of the more pronounced thickening of this fold seen in the male; the other female does not. Tubercles have not been observed in females of any other sa- langoid. In other salangoids breeding tubercles have been observed only on the anal fin of males. An 83.1 -mm male Salangichthys microdon (CAS 52033) has small scalelike breeding tubercles on 184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 the branched portion of the anteriormost branched anal fin-rays. These are arranged uni- serially on each fin-ray branch. A 47.5-mm male Neosalanx jordani (AMNH 51704) has very similar scalelike tubercles on thickened skin sur- rounding the lateral projection at the base and on the basal half of the first four branched anal fin-rays (which are enlarged); fin-rays 6-14, which are bent, each have three to four melanophores on the basal one-fourth of their length. (Similar coloration has been observed on the anal fin in occasional males of Salangichthys microdon.) Breeding tubercles apparently do not occur in Sundasalangidae, in which neither secondary sexual dimorphism nor dichromatism has been observed. Pigmentation The only pigmentation known to be exhibited by salangoids, apart from that of their eyes, is in melanocytes or melanophores, which tend to oc- cur as widely separated single cells or isolated clumps of relatively few cells. In life all, or almost all, salangoids (except Protosalanx) are trans- parent or translucent, except for the prominent eyes. The most constant pigmentary feature of the salangoids is a row of melanophores at the interface of the ventral myotomic musculature and the non-segmentally muscularized ventral abdominal wall. This series of melanophores, with a single cell at about the middle of the ven- tral end of each myotome, from the most anterior myotome to the anal fin origin, is present in near- ly all salangoid specimens examined. Usually these melanophores are longitudinally elongate, giving the appearance of a series of widely spaced thin black dashes. A second pigmentary feature found in many salangoids is a ventromedian row of widely spaced melanophores, one for each body segment. These melanophores tend to be den- dritic when expanded or round when contracted, and may extend the entire length of the abdomen; sometimes they are restricted to the preanal membranous keel. These two pigmentary fea- tures of salangoids occur in many teleost larvae and in adults of other neotenic teleosts. Some salangoids exhibit a row of melano- phores along the anal fin base, one between each anal fin-ray. This row of melanophores, lying deep in the body and median rather than paired, may be the continuation of the midabdominal row of melanophores described above. This row usually extends the length of the anal fin; some- times it continues beyond the anal fin onto the caudal peduncle near its ventral margin. Clusters of a few melanophores occur just an- terior to the bases of the pectoral and pelvic fins in most salangoids, at the tip of the snout and chin, especially in Salangichthys, and infre- quently on the dorsal surface of the head over- lying the fore- and hind-brain. In sexually mature (spawning?) males of Salanx and Salangichthys there may be a cluster of melanophores on the proximal portion of the middlemost anal fin- rays. The dorsal, anal, pectoral, and pelvic fins are otherwise usually devoid cf melanophores, but the caudal fin lobes frequently are dark or dusky due to numerous fine melanophores. The anal scales of the males are always entirely de- void of melanophores. In most salangoids the entire dorsal and most of the lateral body surfaces are devoid of mela- nophores. Protosalanx chinensis and Neosalanx andersoni provide notable exceptions. Young of Protosalanx and Neosalanx exhihil very few me- lanophores. Large and sexually ripe individuals of these two species, however, may have the dor- sal and lateral surfaces of the body with numer- ous melanophores. Those on the dorsal body sur- face are fine, exceedingly numerous, and generally scattered over the entire musculature, but those on the sides are few and peculiarly restricted along the course of the myotomal septae. About a doz- en melanophores lie on each myotomal septa; the melanophores of successive septae are more or less parallel to each other; the cells are oblique- ly elongate, conforming to the thinness and obliquity of the septae and thus forming a series of widely spaced thin black slashes. This pattern, sometimes barely evident or absent in P. chi- nensis, is very well developed in two gravid fe- males of 1 29-1 32 mm (USNM 1 20746). Wakiya and Takahasi (1937) show it well developed in female P. chinensis (not gravid?) and N. ander- soni (gravid); and relatively weakly developed in males of both species. It is present only on the upper part of the body in the relatively small male holotype of TV. andersoni (NRM 10287, 79 mm). Chyung (1961) shows it well developed in a gravid TV. andersoni. I have seen clupeomorphs but no osmeroids or other salmoniforms with similarly distributed melanophores. ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 185 Some Misconceptions Some misconceptions about Salangidae should be noted. These concern the reported presence of scales other than anal scales, presumed ab- sence of the swimbladder, and reputed oral brooding of eggs. The most persistent misinfor- mation concerns the occurrence of scales on the body other than the anal scales. Giinther (1866: 205) stated that the body is "naked or covered with small, exceedingly fine, deciduous scales (?)" and added in a footnote, "There is no trace of scales in specimens preserved in spirits for some time; but others, which we received lately, show scattered fragments of scales, without any regular arrangement." He was unaware of the anal scales of males. Regan (1908b:444), in diagnosing Sa- langinae (=Salangidae), stated simply "scales de- ciduous" but described the anal scales of males in a footnote. Fang ( 1 934a:239) stated body "na- ked or with a few exceedingly thin, large, scat- tered, deciduous scales, without any regular ar- rangement" in addition to the anal scales of males. Nichols (1944) referred to several species with "scales small, deciduous, little evident." Nelson (1976:104) cautiously stated "body generally scaleless" without referring to the anal scales of males. As noted by Wakiya and Takahasi (1937) all salangids are totally scaleless except for the anal scales of sexually mature males; as noted above, the anal scales are large and strongly ad- herent. Reports of scales on other parts of the body are all attributable to dislodged scales from other fishes. Various authors, including Gunther (1866: 205), Fang (1934a:239), and Nelson (1976:104) have stated that salangids lack a swimbladder. Wakiya and Takahasi (1937:268, fig. 1) reported a physostomous swimbladder in Protosalanx chinensis, Salanx ariakensis, S. prognathus, Neosalanx jordani, Salangichthys ishikawae, and S. microdon. In P. chinensis and S. ishikawae the swimbladder is depicted as relatively large and oval, and in the others as equally long but almost uniformly slender for its entire length. The condition of the swimbladder in Sundasa- langidae is unknown. Fang (1934a:238, 252, fig. 7) suggested that Salangidae are oral brooders. In a series of 6 1 males and 27 females identified as Hemisalanx {=Salanx) propnathus collected at Chinkiang in April 1933, Fang found 6 males and 19 females with 1-21 eggs in the mouth. He also reported one Protosalanx (sex not mentioned) with eggs in its mouth. I have also observed a few speci- mens of both sexes, especially of Salanginae, with small numbers of eggs in the mouth; this is at- tributable to rupture of the ovaries and spillage of eggs after the fish had been caught. There is no information indicating that salangoids prac- tice oral brooding or any other form of parental care. This introduction to salangoids concludes with a key for their identification. Key to Salangoidea la. Pelvic fin with 5 rays; adipose fin absent; pectoral fin rayless throughout life; sex- ually mature males without anal scales or enlarged anal fin; vertebrae 37-43; standard length to 22 mm (Sundasalan- gidae) 1 1 lb. Pelvic fin usually with 7 rays (rarely 6 or 8); adipose fin present; pectoral fin with rays except in larvae; sexually mature males with a row of large anal scales and enlarged anal fin; vertebrae 48-79; adults at least 35 mm in standard length (Sa- langidae) 2 2a. Teeth on palatal toothplate and lower jaw in two rows; teeth on tongue in two marginal rows or widely spread over ba- sihyal toothplate (Protosalanginae) Protosalanx chinensis 2b. All oral teeth in single rows 3 3a. Head extremely depressed; snout very elongate and relatively pointed; cranial fontanel entirely closed in juveniles and adults; premaxillae larger than maxillae, those of opposite sides meeting broadly in front of snout; premaxillary teeth rel- atively large; supramaxilla absent; ver- tebrae 66-79 (Salanginae) 4 3b. Head moderately depressed; snout mod- erately elongate and broadly rounded; cranial fontanel with anterior and pos- terior portions open throughout life, pre- maxillae smaller than maxillae, more or less separated from each other in front of snout; premaxillary teeth relatively small, tiny, or absent; supramaxilla pres- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 ent; vertebrae 48-65 (Salangichthyinae) 7 4a. Tongue with a median row of conical teeth (subgenus Leucosoma) Salanx reevesi 4b. Tongue toothless 5 5a. Head strongly pointed; lower jaw not projecting beyond upper jaw; presym- physeal fleshy appendage, bone, and teeth frequently present in adults; vertebrae 72-79 (subgenus Salanx) 6 5b. Head less strongly pointed; lower jaw projecting slightly beyond upper jaw; no presymphyseal fleshy appendage, bone, or teeth; vertebrae 70-73 (subgenus Hemisalanx) Salanx prognathus 6a. Presymphyseal bone usually present in specimens over 1 00 mm standard length, relatively elongate and with up to 1 7 teeth on each side; vertebrae usually 77-78 (rarely 76 or 79) Salanx cuvieri 6b. Presymphyseal bone usually absent, or relatively short and with no more than 6 teeth on each side; vertebrae 72-75 Salanx ariakensis 7a. Palatal toothplate with minute teeth; premaxilla with numerous small or mi- nute teeth, snout relatively elongate; ver- tebrae 59-65 {Salangichthys) 8 7b. Palatal teeth absent; premaxilla usually toothless or with 1-5 minute teeth; snout relatively short except in Neosalanx an- dersoni; vertebrae 48-65 (Neosalanx) 9 8a. Pectoral fin-rays 14-19 Salangichthys microdon 8b. Pectoral fin-rays 20-28 Salangichthys ishikawae 9a. Snout relatively short, standard length to 64 mm, males with 14-21 anal scales, vertebrae fewer than 60 1 9b. Snout relatively elongate, standard length to 100 mm, males with 20-28 anal scales, vertebrae 63-65 Neosalanx andersoni 10a. Vertebrae 55-59; standard length to 64 mm; total rakers on first gill arch 15-19 (mainland Asia) Neosalanx brevirostris 10b. Vertebrae usually 50-53, rarely 49 or 54; standard length usually less than 50 mm; total rakers on first gill arch 9-15 (main- land Asia) Neosalanx jordani 10c. Vertebrae 52-56, average 53.75 (after Wakiya and Takahasi 1937); standard length to 58 mm; total rakers on first gill arch 15 (known only from Ariake Bay, Kyushu, Japan) Neosalanx reganius 1 la. Horizontal diameter of eye less than 4% of standard length; ceratobranchial 5 with 0-3 small conical teeth; total rakers on first gill arch 0-2; vertebrae 41-43 Sundasalanx microps lib. Horizontal diameter of eye more than 5% of standard length; ceratobranchial 5 with about 8-10 large conical teeth; total rakers on first gill arch 10-12; vertebrae 37-4 1 Sundasalanx praecox Material Examined Salangoid specimens deposited in the follow- ing institutions have been examined for this study: American Museum of Natural History, AMNH: British Museum (Natural History), BMNH; Cal- ifornia Academy of Sciences, CAS, including specimens formerly deposited at Stanford Uni- versity, CAS-SU; Museum national d'Histoire naturelle, Paris, MNHN; Naturhistoriska Riks- museet, Stockholm, NRM: Museum of Zoology, University of Michigan, UMMZ; Smithsonian Institution, USNM; and Zoologisch Museum, Universiteit van Amsterdam, ZMA. A detailed list of material examined (including alcian-alizarin preparations) is given under each species in the systematic account. Skeletal Anatomy Salangoid skeletal anatomy cannot be ob- served adequately from alizarin preparations be- cause it is largely cartilaginous, and even ossified portions (including dermal bones) often fail to stain with alizarin. The only previous observa- tions of salangoid skeletal anatomy are brief and relatively uninformative. The only general ac- count, that of McDowall (1969:815), is limited to three paragraphs, one on the cranium, one on the jaws, and one on the remainder of the skel- eton emphasizing the median fins. Wakiya and Takahasi (1937) figured toothed portions of the jaws, palate, and tongue of various salangids. Nelson ( 1 970) described and figured the gill arch- es in Salanx reevesi and Neosalanx brevirostris ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 187 parietal dilatator fossa supraethmoid frontal nasal cranial fontanel trabecula communis ethmoid plate basioccipital lamina orbitonasalis parasphenoid 4 mm Figure 3. Dorsal and ventral view of cranium. Protosalanx chinensis, CAS-SU 6306, 158 mr (his Salanx chinensis and Salangichthys micro- don). He particularly noted the well-developed fourth hypobranchials, "which so far as known are absent from all other adult teleostean fishes." My own observations and drawings of salangid gill arches agree closely with Nelson's. Rosen (1974; figs. 16g, 26a & b) figured and com- mented briefly upon the caudal skeleton and por- tions of the gill arches of Neosalanx brevirosths (his Salangichthys microdon). PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 frontal supraethmoid parasphenoid 1 mm Figure 4. Dorsal and ventral view of cranium. Salanx cuvieri. CAS-SU 32454, 69.4 mr The advent of a technique for staining whole specimens with alcian and ahzarin (Dingerkus and Uhler 1977) made the present relatively ex- tensive observations possible but even so there have been difficulties. Some specimens stained well with alcian but not with alizarin, or vice versa, and in some specimens that otherwise stained well with both stains there are still por- tions of the skeleton which failed to take up no- ticeable amounts of either stain. Such difficulties could not always be made up for by staining additional specimens. In general, alizarin stains only bone. Alcian stains cartilage but also stains some skeletal fea- tures which are obviously bony and have no car- tilaginous precursors, such as fin-rays. Cartilag- inous structures, however, often stain much more deeply with alcian than such non-cartilaginous structures. Thus the salangoid hyopalatine is al- most always stained deep blue and the opercle ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 189 ethmoid plate Oi supraethmoid frontal parietal 2 mm Figure 5. Dorsal view of cranium and membrane bones on dorsal surface of cranium. Salanx prognathus. CAS 51439. 1 10 mm. appears variably pale blue and/or red. In a few of my figures such differences are indicated by the intensity of stippling, but in general the dis- tribution of stain is far too complex to permit its representation in black-and-white illustrations. Some idea of the difficulty involved may be gained from Figure 20 (pelvic girdle of Protosalanx), in which the distribution of stain is indicated. In the cranium the distribution is far more com- plicated and could be conveyed only by illustra- tions in full color. Cranium (Figures 3-8) The cranium of all salangoids is depressed, very strongly in Salanginae and almost as strong- ly in Protosalanginae, but relatively moderately in Salangichthyinae and Sundasalangidae. Some other features correlated with the cranial depres- sion are the peculiarly underslung maxilla, ven- trolateral eye position (especially in Salanginae). and perhaps the posterioriy recurved jaw teeth (especially in Salanginae and Protosalanginae). 190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 tectum synoticum auditory capsule cranial fontanel ethmoid plate supraethmoid taenia marginalis epiphyseal bar semicircular canals pila prooptica nasal recess lamina orbitonasalis trabecula communis basioccipital hyomandibular fossa parasphenoid 1 mm Figure 6. Dorsal and ventral views of cranium. Neosalanx jordani. CAS 52028, 38.3 mm. The development of the cranial fontanel ex- hibits considerable differences. The fontanel ap- parently remains open anterior and posterior to the epiphyseal bar throughout life in Salangich- thyinae and Sundasalangidae, although the an- terior portion may be greatly reduced in larger Salangichthyinae. In Protosalanginae the ante- rior portion closes while the posterior portion always remains open, albeit much reduced in the largest specimens examined. In Salanginae the cranial fontanel is entirely closed in all specimens in which skeletal preparations have been ex- amined. Young Osmeridae in which the cranium is still ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA auditory fenestra taenia marginalis lamina orbitonasalis parachordals trabecula communis hypophysial fenestra I 1 mm notochordal groove I Figure 7. Dorsal and ventral views of cranium. Sundasalanx microps, CAS 44220, 17 mm. cartilaginous have a median bar (taenia tecti me- dialis) separating the anterior and posterior por- tions of the cranial fontanel into left and right halves. Such a feature is usually but not invari- ably absent in salangoids. In a series often Neo- salanx jordani (39.7-45.7 mm), nine have the cranial fontanel entirely undivided, but one (4 1 .0 mm) has a median cartilaginous bar dividing both the anterior and posterior portions of the fon- tanel. The bar is slender posteriorly, but ante- riorly it is much wider, so that the anterior por- tion of the fontanel is represented by two widely PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 cranial fontanel ethmoid plate anterior myodome lateral fenestra pila prooptica trabecula communis lamina orbitonasalis pila prooptica Figure 8. Lateral view of cranium, (a) Protosalanx chinensis. CAS-SU 6306, 158 mm; (b) Salanx cuvieri, CAS-SU 32454, 69.4 mm; (c) Neosalanx jordani. CAS 52028, 38.3 mm; (d) Siindasalanx microps. CAS 44220, 17 mm. separated and relatively small openings. The epi- physeal bar in this specimen is also larger than usual. The condition of the cranial fontanel in this specimen closely resembles that observed in osmerid chondrocrania. In Protosalanx of 85- 89 mm, the anterior portion of the cranial fon- tanel is similarly divided into greatly reduced left and right openings, which become entirely closed in specimens slightly larger. The ethmoid plate is greatly enlarged in all salangoids. In Salangichthyinae and Sundasa- langidae it is broad and moderately elongate, while in Protosalanginae and Salanginae it is broad and extremely elongate. Ossification of the chondrocranium is relatively poor in all salan- goids but varies greatly. The greatest amount of cranial ossification is observed in the skulls of the largest Protosalanx, in which the supraeth- moid, frontals, parietals, parasphenoid, and basi- occipital are all stained more or less deeply with alizarin. In large Protosalanx the posterior por- tion of the parasphenoid has broad lateral wings and the basioccipital has small thin lateral wings (largely obscured by the overlying parasphe- noid). Neither of these features has been ob- served in other salangoids. In all other salangoids the basioccipital ossification is apparently re- stricted to the basioccipital centrum. In Protosalanginae the outline of the cranium is more irregular, suggesting a more primitive condition; while in Salanginae it is relatively smooth and streamlined, suggesting a more de- rived or specialized condition. The auditory cap- sules are most pronounced or laterally prominent in Salangichthyinae. The interorbital septum is relatively open in Sundasalangidae and Salangichthyinae, almost as open in Protosalanginae, but greatly reduced in Salanginae. In Salangichthyinae the anterior- medial portion of the orbit is occupied by very large pilae proopticae arising from the ventral surface of the taenia marginalis or anterior su- praorbital cartilage. In Sundasalangidae the pilae proopticae are rudimentary. A number of cranial features that occur in Sun- dasalangidae have not been observed in the other (mostly juvenile and adult) salangoids examined. Thus the lamina orbitonasalis, which appears as a single apparently simple entity in other sa- langoids, has two components in Sundasalanx: a dorsoanterior contribution from the taenia marginalis and a ventroposterior contribution from the trabecular communis or posteroventral portion of the ethmoid plate. The ethmoid plate is separated by the anterior myodome into dorsal and ventral portions; the anterior myodome ex- tends anteriorly almost to the tip of the snout. In other salangoids the anterior myodome lies much farther posterior, and the ethmoid plate is relatively thin and more or less greatly depressed (least so in Salangichthyinae). In Sundasalanx the base of the cranium is largely occupied by the hypophysial fenestra, a character of all developing teleost chondrocrania usually lost at an early stage. In all other salan- goids the hypophysial fenestra is closed off by cartilaginous growth and the area it once occu- pied may be overlaid by the parasphenoid. In Sundasalangidae the passage for the internal ca- rotid artery is represented by an anterolateral ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 193 hyopalatine premaxilla dentary Meckel's cartilage coronoid process 4 mm quadrate interopercle subopercle Figure 9. Lateral view of jaws, suspensorium, and opercular bones. Protosalanx chinensis. CAS-SU 6306, 158 mm. extension of the hypophysial fenestra; in all other salangoids the passage for this pair of arteries is isolated and widely separated. In Sundasalanx the base of the cranium ex- hibits a median groove on either side of which extends a slight ridge. This groove probably rep- resents the pathway of the embryonic cranial no- tochord before its absorption (complete in all other salangoids examined) into the basioccipital centnim. The ridges on each side may be rem- nants of the parachordal cartilages. Jaws (Figures 9-13) The jaws of salangoids are relatively general- ized, in that the jaw bones, their shape, and the distribution of teeth on them are similar to those in many lower teleosts. In all salangoids the max- illa is toothed and enters broadly into the gape. All salangoids have a single supramaxilla, except Salanginae, in which this element is lacking. In some Salanginae the bony tip of the lower jaw is formed not by the dentaries, but by a median presymphyseal bone (usually tooth-bearing). Due in part to poor quality of alcian-alizarin staining of the lower jaw in salangoids, the relationships of bones that constitute it have not been ade- quately observed. The premaxillae and maxillae are somewhat variable (see remarks in system- atic account). Suspensorium (Figures 9-13) The outstanding feature of the salangoid sus- pensorium is the union of the hyomandibula (hy- osymplectic) and pterygoquadrate, which are united into a single continuous cartilaginous ele- ment, here called the hyopalatine (=palatohyo- mandibuloquadrate of Roberts 1981). Only in Sundasalanx praecox is the hyopalatine divided into anterior and posterior portions, but the di- vision apparently is more anterior than the prim- itive division between hyomandibula (or hy- osymplectic) and pterygoquadrate. In developing vertebrates the rudimentary mandibular arch divides into two cartilages where it bends around the comer of the mouth: the pterygoquadrate bar (dorsal) and the mandibular bar or Meckel's cartilage (ventral). The rudi- mentary hyoid arch divides into the hyoman- dibular (dorsal) and hyoid bar (ventral). All sa- langoids except Sundasalanx praecox show the 194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 supraorbital hyopalatine pseudobranch , 1 mm . Figure 10. Lateral and medial views of jaws, suspensorium, and opercular bones. Salanx cuvieh, CAS-SU 32454, 69.4 mm. most unusual condition of having the dorsal por- tions of the mandibular and hyoid arches fused into a single element. This salangoid element has readily definable features corresponding to the palatine or pterygoid, quadrate, and hyoman- dibula of teleosts in which these elements are separate, but it is unclear whether a portion rep- resenting the symplectic is present. No separate symplectic has been detected in any salangoid; the symplectic may be represented by a thickening or ridge near the ventral margin of the quadrate portion of the hyopalatine. In Sundasalangidae and some Salanginae and Salangichthyinae the suspensorium consists sole- ly of the cartilaginous hyopalatine, but in other Salanginae and Salangichthyinae and in Proto- salanginae a number of perichondral, endochon- dral, or dermal ossifications develop on the sus- pensorium. The elements most often added are the mesopterygoid and an anterior palatal tooth- plate (=ectopterygoid?), which may or may not bear teeth. The suspensorium exhibits more os- sification in large Pwtosalanx than in any other salangoids examined: heavily toothed palatal toothplate, mesopterygoid, and partial ossifica- tion of quadrate and hyomandibula. Whether the dorsal portions of the mandibular and hyoid arches are similarly fused in any other fishes is unknown. In the few fishes for which the development of these arches has been adequately observed it would appear they are separate, in- cluding Salmo (DeBeer 1937), Elops (pers. obs.), Hepsetus (Bertmar 1959). In young salmoni- forms I examined (including Salmo, Galaxias, Lepidogalaxias, Hypomesus, and Spirinchus) cartilaginous pterygoquadrate and hyomandib- ular or hyosymplectic are always separate. Circumorbital Bones (Figure 12) A supraorbital bone is seen in all Salangidae but is absent in Sundasalangidae. The dermo- sphenotic or sixth infraorbital appears to be ab- sent in all salangoids. An isolated infraorbital (fourth or fifth?) is seen in some Salangichthyinae but is greatly reduced (Fig. 1 2). Gill Arches (Figures 14-17) The upper elements of the gill arches of sa- langoids are relatively generalized and, except in Sundasalangidae, so are the lower elements. Ex- cept for the upper and lower pharyngeal tooth- plates the salangoid gill arches apparently are entirely cartilaginous. Four basibranchials are probably present in all salangoids but in none are all of them separate. In Protosalanginae, Sa- ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA Figure Lateral view of jaws, suspensorium, and opercular bones. Salanx prognathus. CAS-SU 51439, 110 langinae, and Salangichthyinae basibranchials 2 and 3 are indistinguishably fused to each other, and in some Salanginae basibranchials 1, 2, and 3 may be so fused. All salangoids have four hypobranchials; hy- pobranchial 4 is always separate and relatively large. The basibranchial series in salangoids is en- tirely cartilaginous. Not only do the basibran- chials themselves not ossify, but basibranchial toothplates, a characteristic feature of many sal- moniforms including salmonids, osmerids, and galaxiids, are absent. The basic basibranchial ar- rangement in Salangidae appears to be basi- branchial 1 separate, basibranchials 2 and 3 fused, and basibranchial 4 separate. A basibranchial 5 is fused to basibranchial 4 in various salmoni- forms, and is apparently usually present in many salmonoids, osmeroids, and galaxioids (includ- ing Lepidogalaxias) as a thin cartilaginous shaft projecting posteriorly between the fifth cerato- branchials. In some instances there is a clear de- marcation between basibranchials 4 and 5, and they may be separate or at least not completely fused. Basibranchial 5, fused with basibranchial 4, is indicated in Salangidae by Nelson (1970), but in Salangidae I have examined there is no indication of a fusion or demarcation between the presumed basibranchial 5 and basibranchial 4. Basibranchial 5 does not project so far pos- teriorly nor is it slender and rodlike as in other Salmoniformes in which its presence is less doubtful. I therefore tentatively consider basi- branchial 5 absent in Salangidae. That it is absent in Sundasalangidae seems highly likely. Gill rakers are poorly ossifed (never stained with alizarin) and edentulous (frequently dentig- erous in salmonoids, osmeroids, esocoids). Those on the trailing (inner) face of the arches usually are fewer and smaller than those on the leading (outer) face (Figs. 14-17). Total number of gill rakers on leading face of first gill arch is 8-19 in Salangidae and 0-10 in Sundasalangidae (Table 2). Dentition The most complete and presumably most primitive dentition in salangoids is observed in infraorbital 1 mm Figure 12. Lateral view of jaws, suspensorium, and opercular bones. Neosalanx jordani, CAS 52058, 35.1 mm. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 PL HQ Figure 13. Lateral view of jaws, suspensorium, and opercular bones, (a) Sundasalanx praecox. CAS 52031, 17 mm; (b) Sundasalanx microps, CAS 44290, 17 mm. HQ = hyomandibula + quadrate, LJ = lower jaw or Meckel's cartilage, MX = maxilla, P = premaxilla, PHQ = hyopalatine cartilage, PL = palatine, OP = opercle, SO = subopercle. urohyal branchiostegal rays basibranchials ceratobranchials Figure 14. Dorsal and ventral views of hyoid and branchial arches. Protosalanx chinensis, CAS-SU 6306, 153 mr ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA ■basihyal interhyal epibranchials 197 infrapharyngobranchials upper pharyngeal toothplate lower pharyngeal toothplate Figure 15. Dorsal view of hyoid and branchial arches and ventral view of upper pharyngeal elements. Salanx cuvieri. CAS- SU 32454, 69.4 mm. 'c:? 1 mm Figure 16. Dorsal view of hyoid and branchial arches; ventral view of infrapharyngobranchial 4 and upper phar>ngeal toothplate. Neosalanx jordani, CAS 52058, 38.3 mm. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 basibranchial 1 + 2 + hypobranchial 1 + 2 basibranchial 3 + hypobranchial 3 hypobranchial 4 basibranchial 4 1 mm Figure 17. Dorsal view of hyoid and branchial arches. Sundasalanx microps. CAS 44220, 17 mm. Protosalanx, with numerous large, conical teeth on the premaxilla, maxilla, palatal toothplate (=ectopterygoid?), tongue (basihyal toothplate), and upper and lower pharyngeal toothplates. As in all Salangidae, there are only two pairs of pha- ryngeal toothplates: the upper, on infrapharyn- gobranchial 4, and the lower, on ceratobranchial 5; the branchial arches are otherwise entirely toothless. In Protosalanx the teeth on the palatal tooth- plate and lower jaw are in two rows, as in many other salmoniforms, but in all other salangoids the teeth on each tooth-bearing element except those in the pharynx are restricted to single rows. In largest specimens of Protosalanx the tongue teeth are more or less widely scattered over the surface of the basihyal (as in Fig. 9), but in small- er ones they are restricted to two marginal rows, as in salmoniforms generally. The only other sa- langoid with tongue teeth, Salanx (Leucosoma) reevesi, has them in a single median row on the basihyal toothplate, a unique specialization for salmoniforms. This character is diagnostic of the subgenus Leucosoma. The maxilla and lower jaw are well-toothed in all salangoids; the palate is toothless in Neosa- lanx and Sundasalanx. In Neosalanx the teeth on the premaxilla, maxilla, and lower jaw are very small, and frequently the premaxilla and lower jaw are entirely toothless. In Sundasalanx bony pharyngeal toothplates apparently fail to form, and the pharyngeal teeth appear to be di- rectly attached to the cartilaginous infrapha- ryngobranchial 4 and ceratobranchial 5. The only bony tooth-bearing elements in Sundasalanx ap- pear to be the premaxilla and maxilla; the lower jaw teeth are loosely attached to Meckel's car- tilage. Pectoral Girdle (Figures 18-19) All salangoids have a secondary pectoral girdle (connecting the primary girdle to the back of the cranium) consisting of three dermal bones: post- temporal, supracleithrum, and cleithrum. Post- cleithra are absent except in Salanginae, in which there is a single postcleithrum. In Salangidae, the primary shoulder girdle consists of the entirely cartilaginous paired scapulocoracoids and one or two series of radials. The basic number of pri- mary radials appears to be five in all Salangidae. The first primary radial, associated with the out- ermost (enlarged) pectoral fin-ray, is relatively simple; it is largest in males of Protosalanginae and Salanginae. The other primary radials are complex, with numerous deep divisions approx- imately corresponding in number to the fin-rays. These divisions are most numerous in Salangich- thyinae, particularly Neosalanx, but are well de- veloped in all Salangidae. Comparable divisions or fimbriae occur in the pectoral basal plate of ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 2 post temporal b 199 postcleithrum Figure 18. Left half of pectoral girdle, (a) Protosalanx chinensis, CAS-SU 6306, 1 58 mm (medial view); (b) Salanx cuvieri. CAS-SU 32454, 69.4 mm (lateral view); (c) Neosalanxjordani, CAS 52058, 43.1 mm (dorsal view); (d) Salangichthys ishikawae, CAS 6780, 74 mm (lateral view). the salmoniform Dallia pectoralis but are not present in other salmoniforms I have examined and do not seem to have been reported in any other teleosts. Secondary radials, more or less corresponding in number to the pectoral fin-rays, are small and simple. The mesocoracoid is lack- ing in all salangoids except that Protosalanx has a process on the median surface of the scapu- locoracoid that may represent the ventral portion of the mesocoracoid (Fig. 1 8a, medial process). In Sundasalangidae the primary pectoral girdle consists of a U-shaped median scapulocoracoid and a basal plate. Fin-rays are absent. Pelvic Girdle (Figure 20) The left and right halves of the pelvic girdle develop in the ventral myotomic wall, and, as the ventral myotomic progression is arrested in Salangidae while the myotomes are still widely separated, the pelvic girdle halves remain widely apart and fail to form any sort of ligamentous or cartilaginous connection between each other. As pointed out by Klyukanov (1975), in Salmoni- formes the two halves of the pelvic girdle are usually joined at least anteriorly for a short dis- tance by strong cartilaginous or ligamentous tis- AxiAL Skeleton (Figure 1) All salangoids have a pair of small dorsal car- tilages straddling the intervertebral disc between the basiocciptal and first vertebral disc; such car- tilages occur in many (perhaps most or all) Sal- moniformes. In all Salangidae the neural arches of vertebrae 1 and 2 are fused dorsally; this condition has not been observed in Osmeridae or any other sal- moniforms I have examined. In Sundasalangidae the neural arches of vertebrae 1 and 2 are sep- arate from each other and morphologically sim- ilar to those of the vertebrae immediately suc- ceeding them. In salangoids the mineralized portion of each centrum is relatively elongated and hourglass shaped, so that the intervertebral joints are nar- row and the notochord greatly constricted. In 200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 pp SCO 1 mm 1 mm Figure 1 9. Pectoral girdle, (a) Sundasalanx microps, CAS 44220, 1 7-mm adult (posterior view); (b) Elops hawaiiensis, CAS 52035, 30-mm leptocephalus larva (posterior view); (c) Sardina pilchardus, 20-30 mm (ventral view?, after Goodrich 1922); (d) Dallia pectoralis, (lateral view, after Starks 1904; apparently based on CAS-SU 12615, 125 mm, Nushagak River, Alaska); AP = ascending process, CL = cleithrum, F = fin margin, PP = posterior process, PT = posttemporal, R = basal plate, SCL = supracleithrum, SCO = scapulocoracoid. In (b) and (c) the first primary radial has pinched off from the basal plate. salmonids, osmerids, galaxiids, and other Sal- moniformes, especially in the young stages, the mineralized portion of each centrum tends to be relatively short and cylindrical, so that the in- tervertebral space is much larger and the verte- bral section of the notochord is entirely intact. A comparable condition is not present in any salangoid skeletal material I have examined. Ribs are absent or weakly developed and stain poorly. They are small, weakly stained with al- cian when present (Fig. lb). Gosline (1960) and others have pointed out that neural and hemal spines of most Salmoni- formes, especially posteriorly, may be flattened or laminar, even to the extent of resembling a continuous keel. The neural and hemal spines of salangoids are always relatively slender, espe- cially posteriorly. A round, oval, or elongate and splintlike adi- pose fin cartilage lies at the base of the adipose fin in all Salangidae. A survey of lower teleosts for the adipose fin cartilage by Matsuoka and Iwai (1983) revealed its presence in Salangidae, Osmeridae, Plecoglossidae, Myctophidae, and Neoscopelidae; it was not observed in other low- er teleosts with an adipose fin including Sal- ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 201 radials1-4 blue I 1 mm ) parapelvic cartilages 1 mm 1 mm Figure 20. Ventral view of left half of pelvic girdle, (a) Protosalanx chinemis. CAS-SU 6306, 158 mm; (b) Salanx cuvieri. CAS-SU 32454, 69.4 mm; (c) Neosalanx jordani, CAS 52028, 43.1 mm; (d) Sundasalanx microps. CAS 44220, 17 mm (with lateral view of pelvic girdle and parapelvic cartilages above). monidae, Retropinnidae, Prototroctidae, Aulo- podidae, Synodontidae, Chlorophthalmidae, Argentinoidei, Characoidei, or Siluriformes. The similar morphology of the adipose fin cartilages in Salangidae and Osmeridae, as noted by Mat- suoka and Iwai, is possibly indicative of rela- tionship between these two families. Caudal Fin Skeleton (Figure 21) The caudal fin is more or less deeply forked, and the upper and lower lobes are about equal. Principal caudal fin-rays are invariably 10 + 9; upper and lower procurrent caudal fin-rays are moderately numerous (to 14). The complex ural or hypural centrum apparently consists of three centra and uroneural 1 (sometimes also uro- neural 2?) fused into a single unit. The three cen- tra involved are the terminal centrum and post- terminal centra 1-2. according to the nomenclature of Gosline (1960), or preural cen- trum 1 and ural centra 1-2, according to Rosen (1974). In none of the skeletal material examined is there any indication of separate centra poste- rior to the complex hypural centrum. Epurals 0- 3. A separate uroneural 2 is sometimes present, but uroneural 1 is apparently always fused to complex hypural centrum. Free radial or pter\- gial cartilages are sometimes present, usually be- tween ray halves at the base of the anteriormost 2-3 upper or lower procurrent rays and the low- ermost upper and uppermost lower principal rays. Hypurals six. Six separate hypurals occur in Sa- langichthys micwdon (Rosen 1974, Fig. 26). Pro- tosalanx chinensis occurs with hypurals 1-2 and 5-6 separate, but with 3-4 fused near the base. The hypurals are more fused in Neosalanx, Sa- lanx, and Sundasalanx. In Salanx parhypural and hypurals 1-2 are fused near the base: hy- purals 1-2 and 3-4 are fused for their entire length except for oblong basal foramina where fusion evidently failed to complete. In Sundasalanx parhypural and hypurals 1-3 are evidently fused into a single element. Systematics In the present account the salangoids are rec- ognized as a salmoniform superfamily separate from osmeroids, which they superficially resem- ble. There are two families, Sundasalangidae, with 202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 opjsthural hypurals1-6 hemal spines hypurapophysis^ ^ parhypural Figure 21. Lateral view of caudal fin skeleton, (a) Protosalanx chinensis, CAS-SU 6306, 158 mm; (b) Salanx cuvieri, CAS- SU 32454, 61.7 mm; (c) Neosalanx jordani, CAS 52028, 43.1 mm (note: hypurals 2 and 3, normally separate from each other in all salangoids, are fused in this specimen); (d) Sundasalanx microps CAS 44220, 17 mm. In a-c left half of fin rays removed to facilitate observation of median structures. only a single genus and two species, and Salangi- dae. Salangidae is further divided into three subfamilies, four genera, and eleven species. The genus Salanx is further divided into three sub- genera; this taxonomic category is not employed in the other genera of salangoids. In addition to the new superfamily Salangoidea, the new subfamily Salangichthyinae is proposed for Neo- salanx and Salangichthys, leaving the subfamily Protosalanginae with only the genus Protosa- lanx. No new genera or species are proposed. Some previous workers, particularly Regan (1908b) and Fang (1934a, b) recognized far more species than I have, especially in the subgenus Salanx (genera Salanx and Parasalanx of Re- gan). This is attributable in part to their basing species on only one or a few type-specimens and utilizing characters such as cranial proportions, body depth, and relative position of dorsal and anal fins which vary considerably within the species. Neither Regan nor Fang utilized verte- bral counts, which I find extremely useful in dis- tinguishing species. My extensive data on ver- tebral counts of types and other material are presented in Table 2. My counts of vertebrae, fin-rays (except pelvic) anal scales, branch iostegal rays, and gill rakers are presented in Table 2. This table includes all species of salangoids herein recognized as valid except Neosalanx reganius, which I have not ex- amined. Pelvic fin-ray counts are excluded be- cause they are invariably 5 in Sundasalangidae and almost invariably 7 in Salangidae (6 in one observed specimen of Neosalanx Jordani, 8 in two specimens of Salangichthys microdori). Pre- vious authors have presented data on most of the species but have often lumped data from various localities (and frequently of two or more ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 203 species). In order to minimize this problem, my data are presented separately for each locality. Although Table 2 includes meristic data ob- tained from whole specimens, radiographs, and cleared and stained specimens, the stained spec- imens provide the best material for accurate count of fin-rays, teeth, and gill rakers. It is difficult to observe the jaw teeth and lowermost pectoral fin- rays in Neosalanx except in stained material. In dorsal and anal fin-ray counts the last "two" rays ("divided to base") are counted as one ray. In vertebral counts the basioccipital centrum is not counted and the hypural complex centrum is counted as one. In salangoids, especially in fe- males, the anteriormost anal-fin pterygiophore does not provide a ready basis for distinguishing abdominal and caudal vertebrae. In order to ob- tain additional meristic data from the vertebral column and at the same time obtain more precise data on the relative position of fin origin, I have taken data on the vertebrae parallel to the origins of the pelvic, dorsal, and anal fins. The number of vertebrae posterior to a vertical line through the base of the last anal fin-ray is also recorded. Radiographs are usually satisfactory for ob- taining vertebral counts of salangoids and some- times for fin-ray counts. Sometimes the verte- brae may show up very faintly but it is almost always possible to obtain a count repeatable to within one vertebra. Fin-rays, however, fre- quently cannot be accurately counted on radio- graphs, and I have only incorporated data on fin-ray counts taken from radiographs when the radiographs seemed reliable. Some characters utilized by other workers to distinguish species are not emphasized here be- cause they do not seem useful. This particularly applies to pectoral fin-ray counts in Neosalanx and to the elongation of the head or cranium, relative position of the dorsal- and anal-fin bases, and body depth, especially in Salanx. In salan- gids the number of pectoral fin-rays generally continues to increase slightly with growth, es- pecially so in those such as Neosalanx, in which the rays are exceptionally numerous. The elon- gation of the cranium (particularly its anterior portion) is extremely variable in Salanx, as not- ed also by Wakiya and Takahasi (1937:289). This variation is individual and is probably enhanced by growth. The position of the dorsal and anal fins relative to each other is also highly variable in salangids, subject to individual variation as well as sexual dimorphism. In defining species of Salanx too much reliance has been placed on slight differences in fin positions based on only one or two specimens. Salanx, Salangichthys, and other salangids vary enormously in body depth due to sex-related body changes and non- sexual factors of condition and preservation. In discussing salmonoid classification, Gosline (1971:1 19) stated: The suborder Salmonoidei as here recognized (Families Sal- monidae, Osmeridae, Plecoglossidae, Salangidae, Retropin- nidae, Aplochitonidae, and Galaxiidae) is a group of highly diverse inshore and freshwater salmoniform fishes. Though the included families no doubt should be divided into su- perfamily groupings, inadequate knowledge of the Salan- gidae and the Southern Hemisphere forms would seem to make any formal superfamily classification premature at the present time. Informally, the members may be divided between Northern and Southern Hemisphere forms. The diverse forms from the Southern Hemisphere seem to be most closely related to the northern osmerids. . . . The Northern osmeroids are represented by four quite distinct lines: Salangidae, Plecoglossidae, Osmeridae, and Salmon- idae. Rosen (1974) divided the suborder Salmo- noidei into two superfamilies, Salmonoidea— in- cluding the Southern Hemisphere families (ex- cept Retropinnidae) and Salmonidae — and Osmeroidea (with four families listed as incertae sedis: Osmeridae, Plecoglossidae, Retropinni- dae, and Salangidae). I have not investigated Retropinnidae or the highly aberrant Plecoglos- sidae but suspect that Retropinnidae (particu- larly Prototroctes) and Plecoglossus may indeed be closely related to each other and perhaps to Osmeridae. But I have not been able to find any good evidence (in the form of shared specializa- tions or derived characters) between Salangidae and any one or combination of these families. I have therefore designated the new superfamily Salangoidea, which is coequal with the superfam- ilies Osmeroidea and Salmonoidea (and Galax- ioidea, if this is also to be recognized). Salangoidea, New Superfamily This superfamily apparently differs from all other Pisces in having a suspensorium in which the cartilaginous palatine and pterygoid (of the mandibular arch) and quadrate and hyomandib- ular (of the hyomandibular arch) are fused into a single element, the hyopalatine. Gill arches with well-developed fourth hypobranchials— so far as known absent from all other adult teleosts (Nel- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 son 1970). Pharyngobranchials 4. Anterior pha- ryngobranchial modified as an elongate "suspen- sory pharyngeal"; only fourth pharyngobranchial bears teeth (teeth absent in Siindasalanx). Fifth ceratobranchial with well-developed teeth (ab- sent in osmeroids; McAllister 1963:4). Bran- chiostegal rays 2-5 (7-19 in salmonoids, 5-10 in osmeroids, 3-9 in galaxioids). Cranium mod- erately to excessively flattened (more so than in any other salmoniforms). Maxillary bone, bear- ing teeth for its entire length, with its posterior half abruptly curved medially beneath head (so that teeth on posterior half of maxillary are di- rected anteriorly rather than ventrally). Scales entirely absent except for a row of strongly ad- herent anal scales in adult male Salangidae. Dermosphenotic and circumorbital bones ab- sent, except for a single small troughlike bony element observed in Neosalanx, which may rep- resent a fifth or sixth circumorbital (not dermo- sphenotic). Supraocciptal bone absent (present in most other salmoniforms). Pectoral fins pedunculate throughout life (with pectoral radials in a fleshy pedestal separate from body). Pelvic fin-rays usually 5 or 7 (rarely 6 or 8; 8 in osmeroids). Principal caudal fin-rays in- variably 10 + 9 (as in most lower teleosts includ- ing salmoniforms with generalized caudal fins; galaxioids have fewer). Salangoids apparently have no laterosensory canals on the body. The cephalic laterosensory canals, although well developed, are superficial (i.e., not enclosed in bony tubules) and often dif- ficult to observe in their entirety. Those of Sa- lanx chinensis, illustrated by Nelson (1970, Fig. 15), do not exhibit any particularly unusual fea- tures for lower teleosts. There are preopercular, mandibular, supraorbital, infraorbital, and ex- trascapular canals. The mandibular is not con- tinuous with the preopercular. The supraorbital and infraorbital extend anteriorly only a short distance in front of the nostrils, i.e., not signifi- cantly onto the greatly depressed and enlarged snout. The infraorbital has 8 pores, the preoper- cular 6, and the mandibular 5. Alimentary canal a relatively simple, straight tube. Pyloric caecae absent. Gonads paired. Salangidae Jordan and Snyder, 1 902 Pelvic fin almost invariably with 7 rays (8 ob- served in one specimen of Salangichthys ishi- kawae and two S. microdon, 6 in one Neosalanx jordani). Pelvic girdle without parapelvic carti- lages. Pectoral fin-rays 8-34. Pectoral girdle with five proximal radials; distal ends of one or more proximal radials with more or less numerous branches; adult males with a series of anal scales and enlarged, modified anal fins; total vertebrae 49-79. The family Salangidae comprises three subfamilies: Protosalanginae, Salangichthyinae, and Salanginae. Protosalanginae Wakiya and Takahasi, 1937 This subfamily, here restricted to the mono- typic genus Pwtosalanx, differs from all other salangoids in having the premaxilla, palatal toothplate (=ectopterygoid?), and dentary with two rows of teeth instead of at most a single row; the basihyal toothplate of the tongue also has the teeth in two marginal rows (a primitive condition for salmoniforms) or irregularly scattered over its entire surface; the only other salangoid with basihyal teeth has them in a single median row. Pelvic fins relatively larger and more anterior than in any other salangoids (see Fig. 1, Table 2). Cranium strongly depressed (almost as much as in Salanginae); adults with anterior portion of cranial fontanel closed, posterior portion of cra- nial fontanel greatly reduced but remaining open throughout life (both portions closed in adult Salanginae, open throughout life in Salangich- thyinae and Sundasalangidae). Lower jaw weakly projecting beyond upper jaw; premaxillae pro- jecting anteriorly beyond snout tip as in Sa- langinae but failing to form a membrane-covered space through which symphyseal teeth of lower jaw project. Lower jaw without enlarged sym- physeal teeth (present in Salanginae), sometimes with a weakly developed fleshy presymphyseal process but without presymphyseal teeth or bony process. Adults attaining slightly greater stan- dard length (Table 1) and heavier-bodied than any other salangoids. Dorsal fin-rays 16-18 and anal fin-rays 30-32 (vs. 10-15 and 14-32 in all other salangoids); vertebrae 66-70 (Table 2). Protosalanx Regan, 1908 Eperlanus Bas\i.ewsky, 1855:242. Salanx Abbott, 1901:490. Protosalanx Regan, 1908b:444 (type-species, by monotypy, Salanx hyalocranius Abbott, 1901 = Eperlanus chinensis Basilewsky, 1855). Paraprolosalanx Fang, 1934a:246 (type-species, by mono- ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 205 typy, Protosalanx andersoni Fang, 1 934a (non Rendahl, 1923) = Protosalanx chinensis Basilewsky, 1855). Protosalanx chinensis (Basilewsky, 1855) Eperlanus chinensis Basilewsky, 1855:242 (type-locality "in sinu Tschiliensis habitat" [not "Pekin" as usually cited]). Salanx hyalocranius Abbott, 1901:3490 (type-locality Pei-ho at Tien-tsin). Protosalanx hyalocranius Regan, 1908b:445. Paraprotosalanx andersoni Fang, 1934a:246 (Figs. 4-6, text in part [non Paraprotosalanx andersoni Rendahl, 1923]). Protosalanx chinensis Chyung, 1961:163. Material Examined. -BMNH 1929.2.5.2.-3, 61-65 mm, Kiangyin; CAS 52026, 60:48.2-70.7 mm, no locality (pur- chased in San Francisco); CAS-SU 6306, 25:80.5-163 mm, Pei-ho at Tien-tsin, paratypes ofSalanx hyalocranius (7:85.5- 158 mm alcian-alizarin); CAS-SU 23639, 1:137 mm, Seoul; CAS-SU 36025, 3:120-136 mm, no locality; UMMZ 180096, 2:127-129 mm, Korea; USNM 120746, 2:129-132 mm, Ko- rea. Protosalanx appears to be the most primitive salangoid. There is no indication that it com- prises more than a single species. Although Ab- bott's account begins "'Salanx hyalocranius new species," it concludes "this species is probably identical with Eperlanus chinensis Basilewsky, from Pekin, but the name chinensis is already used for the 'whitebait of Makao' " (Abbott 1 90 1 : 490-491). In Abbott's time Salangidae were so poorly known it was reasonable for him to as- sume that his material might represent an un- described species, but even so it is clear from this statement that Abbott was really proposing a re- placement name. Now that Salangidae are better known it seems Basilewsky's account could only refer to this species, as explicitly recognized by Wakiya and Takahasi (1937), although they re- tained the name P. hyalocranius. The holotype of P. chinensis cannot be found (Barsukov, pers. comm. 1983). Since the "whitebait of IVlakao" has been referred to as Leucosoma or Salanx chinensis but never as Eperlanus or Protosalanx chinensis, the epithet chinensis is available for a species oi Protosalanx. As this is also the earliest name proposed it must replace hyalocranius, and the species should be known as Protosalanx chi- nensis. The only publication to come to my at- tention in which this name is correctly applied is by Chyung (1961). Wakiya and Takahasi (1937) correctly iden- tified Paraprotosalanx andersoni Fang, 1934a with this species. Fang's figures agree in every respect with P. chinensis. The fleshy presymphy- seal appendage, presumed by Fang to differen- tiate his Paraprotosalanx from Protosalanx, is also present in some of Abbott's type-specimens of 5. hyalocranius. Fang's figures presumably are based upon the single large male, "S. 4374," 153 mm (total length according to Table 4, but stan- dard length according to p. 247) from Nanking. All or almost all of the other specimens referred to Paraprotosalanx andersoni by Fang are prob- ably Neosalanx. It should be noted that small specimens in museum collections identified as Protosalanx diXt usually Neosalanx and that all or almost all pub- lished reports of smaller Protosalanx up to the present time are based on Neosalanx. For ex- ample, I find that all of the small specimens in Abbott's type-series of S. hyalocranius are Neo- salanx. Young P. chinensis are relatively rare in collections. Those I examined (smallest 48.2 mm) closely resemble the largest adults in every way except they lack the sexually dimorphic char- acters of adult males. The strongly pointed snout and large teeth arranged in two rows on the pal- ate, tongue, and lower jaw are easily observable. Neosalanx have no teeth on the tongue or palate, and the jaw teeth except on the maxillary are absent or minute and difficult to observe, while the males are sexually mature and provided with greatly enlarged anal fins and anal scales at rel- atively small size. The smallest male Protosalanx with anal scales is probably considerably larger than any Neosalanx. Protosalanx chinensis appears heavier-bodied at all sizes and to attain a greater size than any other salangoid. The 163-mm specimen is the largest that has been reported. Salanginae Regan, 1 908b Cranium and especially ethmoid plate very strongly depressed and elongate, more so than in any other salmoniforms. Adults with cranial fon- tanel entirely closed (posterior and sometimes also anterior portion of cranial fontanel open throughout life in all other salangoids). Upper and lower jaws with strongly pointed or project- ing tips. Teeth relatively large and few in num- ber. Premaxillae projecting beyond concave an- terior margin of ethmoid plate to form a membrane-covered space penetrated by enlarged symphyseal teeth of lower jaw. Lower jaw often with a fleshy or bony presymphyseal process and presymphyseal teeth (Wakiya and Takahasi 1937, pi. 20, figs. 31-34). Supramaxilla absent 206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43, No. 13 (present in all other salangoids). Pectoral fin-rays 7-1 1 (20-32 in all other Salangidae). Pectoral girdle with a single postcleithrum (absent in all other salangoids). Body extremely elongate, more so than in any other salmoniforms. Basal portion of neural and hemal arches expanded, frequently covering centrum laterally and fusing with each other. Distal portion of neural arches with an- terior and posterior projections, those of succes- sive vertebrae articulating with each other. Ver- tebrae 68-79 (37-70 in all other salangoids). The enlarged symphyseal teeth of the dentary and the membrane in the roof of the mouth through which they project presumably form a puncturing device to kill prey. Similar devices, with foramina in the roof of the mouth through which fanglike lower jaw teeth pass, occur in other piscivorous or predatory teleosts, including the characoids Hepsetus, Hop lias, and Acestro- rhync hus (Rohens 1969). In many respects Salanginae appear to be the most highly specialized members of the family. Even the low pectoral fin-ray count, which might be considered primitive, may be secondarily evolved, since primary pectoral radials 2-4 of Salanginae exhibit the distally fimbriate or dig- itate condition that is probably associated with the retention of pedunculate pectoral fins (prob- ably a neotenic character) and evolution of large numbers of pectoral fin-rays (a specialization shared by all other Salangidae). SalanxOken, 1817 "Les Salanx" Cuvier, 1817:185 (French vernacular; not avail- able for zoological nomenclature). Salanx Oken, 1817:1183 (Latinization of Cuvier's "Les Sa- lanx," and the earliest name available for zoological no- menclature; see ICZN Declaration 87, paragraph 12. Type- species, by monotypy, Salanx cuvieri Valenciennes, 1849). Leucosoma Gray, 1831:4 (type-species, by monotypy, Leu- coso ma reeves i Gray. 1831). Hemisalanx Regan, 1908b:444 (type-species, by monotypy, Hemisalanx prognalhus Regan, 1908b). Parasalanx Regan, 1908b:444 (type-species, by subsequent designation of Fang, 1934a:259, Parasalanx gracillimus Regan, 1908b = ?5'a/a«.v cwv/m Valenciennes, 1849). Reganisalanx Fang, 1934b:509 (type-species, by monotypy, Reganisalanx normani Fang, 1934b = Sa/a«.\- ariakensis Kjshinouye, 1901). Metasalanx Wakjya and Takahasi, 1937:293 (type-species, by monotypy, Metasalanx coreanus Wakjya and Takahasi, 1937, a nomen nudum). The four species herein recognized as consti- tuting the genus Salanx have been placed by other authors in three genera, Salanx, Hemisa- lanx, and Leucosoma. Wakiya and Takahasi (1937) even placed Hemisalanx in a subfamily of its own, Hemisalanginae, regarded by them as intermediate between Protosalanginae and Sa- langinae. Because these four species differ strik- ingly from all other salangids in several features of skeletal anatomy but agree closely with each other in conformation of the cranium and jaws, distribution and size of jaw teeth, number of pectoral fin-rays, and the peculiar modification of their neural and hemal arches and high ver- tebral counts, I prefer to recognize them as be- longing to three subgenera in the sole genus of the subfamily Salanginae. Salanx (Salanx) ariakensis (Kishinouye, 1901) Salanx ariakensis Kishinouye, 1 90 1 :359 (type-locality Ariake Bay, Kiushiu). Salanx acuticeps Regan, 1908a:360 (type-locality Lake Can- didius, Formosa). Parasalanx acuticeps Regan, 1908b:446. Parasalanx longianalis Regan, 1 908b:446 (type-locality Liao- ho, northern China). Parasalanx annitae van Dam, 1926:342 (type-locality Pei- taiho, China). Reganisalanx normani Fang, 1 934b:509 (type-locality Ichang. as herein restricted). Material Examined. -AMNH 10327, 7:125-147 mm, Hunan; BMNH 1 888.5. 1 5. 1 1-12, 2: 141-143 mm, Ichang (lec- totypeandparalectotypeof/?. normani): BMNH 1898.2.8.20- 23, 4:1 14-123 mm, Liao-ho, northern China (syntypes of P. longianalis): BMNH 1904.4.2835-36, 2:116-118 mm, Uke Candidius, Formosa (syntypes of S. acuticeps): BMNH 1927.3.26.3, 125 mm, Nanking; BMNH 1928.6.22.6, 1 15 mm. Wenchow; CAS-SU 8574, 2:99.1-104 mm, Ariake Sea (iden- tified by Kishinouye); CAS-SU 23103, 107 mm. Maruyama, Taihoku, Formosa; ZMA 1 12.587, 128 mm, Peitaiho, China (holotype of P. annitae). In vertebral counts and in all other respects so far as known the four syntypes of P. longianalis agree well with other material herein referred to as Salanx ariakensis, except for their consis- tently high anal fin-ray counts of 30-32 (reported by Regan 1908b:446). Most samples of 5. ari- akensis examined have only 26-29 anal fin-rays, but two specimens from Ariake Bay have 27 and 31. Reganisalanx normani is based primarily on the description by Regan (1908b) and supple- mentary notes by Fang (1934b: 509) of two spec- imens from Ichang (BMNH 1888.5.15, 11-12), identified by Regan (ibid.) as Salanx cuvieri. Fang declared that the specimens represented a dis- tinct genus but did not provide a proper generic ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA diagnosis or description; apparently he distin- guished it from Salanx based on the lack of a presymphyseal bone. In my opinion the char- acter cannot be used to split the genus Salanx. I have reidentified these specimens as S. ari- akensis, a species in which the presymphyseal bone may be present or absent. I have not seen the third specimen referred to R. normani by Fang (ibid.). It is clear from Fang's account that he did not compare this specimen directly with the two specimens from Ichang, and it might not be conspecific. In order to fix the identity of this nominal taxon, the 141 -mm undamaged speci- men from Ichang (BMNH 1888.5. 15. 11) IS here- by designated the lectotype. The 143-mm spec- imen, with the body damaged just behind the head and at mid-abdomen, is a conspecific para- lectotype (BMNH 1888.5.15.12). Fang (1934a) reported 1 1 specimens (as Para- salanx longianalis) with the following anal fin- ray counts: 28(5), 29(2), 30(3), 32(1). The ver- tebral counts are unknown for these specimens but it seems likely from Fang's account that they are all S. cuvieri. The holotype of P. annitae has the head rel- atively short and broad (for the subgenus Salanx) and in this respect is more like S. ariakensis than S. cuvieri. A presymphyseal bone is present, but it is short considering the large size of the spec- imen, and has only 2 teeth on each side. The premaxilla has 7 teeth, maxilla 1 2, and dentary about 10. Salanx (Salanx) cuvieri Valenciennes, 1849 Salanx cuvieri Valenciennes in Cuvier and Valenciennes, 1849:360 (type-locality unknown). IParasalanx gracillimus Regan, 1908b:446 (type-locality Shanghai). Parasalanx angusticeps Regan, 1908b:446 (type-locality China). Parasalanx cantonensis Herre, 1932:425 (type-locality Can- ton). Material Examined.-AMNH 51689, 3:88.6-106 mm. Canton; BMNH 1855.9.19.1539, 144 mm (holotype of P. a«- gusticeps); BMNH 1891.1.31.20, 1 1 1 mm, Shanghai (holotype of/'.^raa7/;mwj);BMNH 1936.10.7.13, 1 19 mm. Sharp Peak, Fukien; CAS 52057, 4:76.5-98.0 mm. Hong Kong ( 1 alizarin); CAS-SU 225732, 1 12 mm. Canton (holotype of P. cantonen- sis): CAS-SU 32454, 18:56-66 mm, Chuan Is. (4:61.7-69.4 mm aician-alizarin); CAS-SU 32943, 1 17 mm, near Pakhoi, SW Kwangtung; MNHN 9900, 1 1 2 mm, no locality (holotype). So far as I have been able to determine, vari- ation in the presymphyseal bone within each species, including its presence or absence and its length or amount of dentition, is correlated chief- ly with size and is not sexually dimorphic. Notes on Holotype. — The holotype (Fig. 2a) is dried but complete and in fair condition. The body immediately posterior to the head is badly damaged and fin-rays brittle, so it must be han- dled with care. Cranial width (at anterior margin of eyes) 3.5 in cranial length. Presymphyseal bone, 2.1 mm long, with 1-2 moderately large teeth basally and at least 2 minute teeth distally. Pre- maxilla considerably elongated anteriorly, with 7-8 teeth. Maxilla with about 7 teeth. Dentary with about 13 teeth of variable size. Palatal teeth 7, very small and in a single row. The following proportional measurements are expressed as times in standard length. Length of cranium about 7; length of head (to end of gill cover) 4.7; length from anterior midline of ethmoid plate (concave) to anterior rim of orbit 16; length from tip of upper jaw (premaxilla) to anterior rim of orbit 1 0; diameter of eye (slightly shrunken) approx- imately 28. Notes on Synonymy.— P. angusticeps is dis- tinguished by Regan primarily on the basis of its exceptionally elongate head: "head nearly 4 times as long as broad; snout a little longer than post- orbital length of head" versus head 3 times or a little more than 3 times as long as broad, and snout only as long as or a little shorter than post- orbital length of head in all other Parasalanx and Salanx (Regan 1908b:445-446). The den- tition of the holotype of P. angusticeps, a gravid female of 144 mm, is complete and undamaged. Presymphyseal bone elongate with 5-6 teeth on each side; premaxilla with 7 teeth; maxillary teeth 10 or 11; dentary with a small tooth anteriorly (just behind symphysis), then an enormous ca- nine tooth, followed by 7 small teeth and 6 mod- erately large teeth; palatine with 8 small teeth in a single straight row^ The holotype of P. gracillimus is in poor con- dition, dried, twisted, and slightly shrunken. Its body depth, reported as 18 times its length, is attributable to the poor condition (emaciation) of the specimen. Its dentition is as follows: pre- symphyseal bone with 3 teeth on each side, pre- maxillary 5, maxillary 8, dentary with 1 mod- erately large, 6 small, and 5 moderately large, and palatal 7 moderately large. The vertebral column is broken anteriorly, making all of the counts based on vertebrae doubtful. Wakiya and Takahasi (1937:288) tentatively placed P. gra- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 < s 3 M t: C n t^ o 2 C O 'S .So ^ s g 1 lit 1 5u g 11 ■* § -2 a 5 a &C K III ^^ S >> >, m o o o i C/3 C/5 CA o X o o X X —■ ^ ^ rj o T 1 1 1 1 1 2 oo -- 1 1 5 - ri> r!. oo ' ' ' ' Ov 1 1 so (^J r^ OJ <^J (N 8 S 9^ I I I I I I I I O I I O O Ov w-1 ",' O) Tf ■^ O '^' Ov o z ^ i^ E^ ^ o ^ o C/:j i^J ly^J "^r- cA c>2 t/0 £ < < < 2 U O U D c riooooovr-oot-- — ^ m oo o^ C^ ri n ir^ rn -^ OOOOOOvOONQOf^^ o o — o — 4, I I I O On O i_; _ ro _ <-0 f-,| r^ r*^ r*^ ^ r*^ _j^ m _i. ^, r- \o r-- — r- ::_:, I t/2 U. O r^' I O O ^ -:2 "i:; ^O >>^ 0^ f*^ r-~' 1 1 z z < CO s s - i;s ^ ^< >-. Z c/: < < s 112 U U N ,? £ .1 o 5u O — — r-) vo ^ 3. ^ c/p op c/p i 5 5 o^ c/'j ^ g s S < < < ■| < < u u o ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 209 1-1° o — — , \D r- '7 r-- -Si ■§ 2 5 ci C 3 3 3 O CO t^ X H U. U. U 1^ O lO T^f O r<-i OO 1/-1 Tj- o\ m T — O ^ r<-i tT ro r^ ■q- ro z z < < til i 11 = C O i/5 § '^ ^' ,^ o£ 1 X d SS 5 u^' ^' ;5 TS o o rsf u^ oo c r— r^ Qo' PJ — vO 0^ o w-l C/) !/2 z ^ ^ a I I !^ ^ ^ ^ < < < z 5 lis o o u s 1 < < o Ov o \0 o vD c;, '^ 3 - ci r7 C2 — ^ a o 1^1 D <: n D ~C; c/n C/0 cA t>0 <• u-i yj § « <• T3 < fgi^^ Vol. 43, No. 16, pp. 249-267, 6 figs., 6 tables December 11, 1984 THREE NEW SPECIES OF SEVEN-GILLED HAGFISHES (MYXINIDAE, EPTATRETUS) FROM THE PACIFIC OCEAN By Charmion B. McMillan and Robert L. Wisner Marine Biology Research Division, A-002, Scripps Institution of Oceanography, La Jolla, California 92093 Abstract: Three new species of hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean are described, and compared with E. cirrhatus. All four species have seven pairs of gill pouches and associated external openings. Of the new species, E. carlhubbsi is known from Molokai to Guam, north-central Pacific, E. laurahubbsi from off south-central Chile, and E. strahani from near Lubang Island, Philippines, South China Sea. Eptatretus cirrhatus occurs in the Australian-New Zealand area. Methods used in examination of hag- fishes are described, and sensory (lateral line) canals are delineated and discussed briefly. Introduction This study of seven-gilled hagfishes (genus Ep- tatretus) from the Pacific Ocean is one of a series resuhing largely from the specimens and data accumulated under direction of the late Carl L. Hubbs. Herein we describe three new species, present new data on E. cirrhatus (Bloch and Schneider 1801), offer suggestions for initial preservation of myxinids to provide good study material, and discuss methods useful in the taxo- nomic study of hagfishes. We also offer figures and a brief description of the sensory canals found in the ocular regions of two of the four species. Discussion Our examinations have shown that species of Eptatretus from the Pacific Ocean have six to fifteen pairs of gill pouches and corresponding external apertures. The three new species de- scribed below, with Eptatretus cirrhatus, com- prise a group having seven pairs of gill pouches. One aberrant specimen has eight pouches on each side, but with corresponding apertures arranged abnormally. Our rather limited counts (22 pairs) from the three new species may not reflect ex- tremes of variation, but the number of gill ap- ertures in Eptatretus cirrhatus appears to be con- stant—seven pairs in 48 specimens. In 44 counts from 22 specimens of the three new species, the only variation from seven was the specimen cited above (further discussed and figured below). Counts of six apertures for Eptatretus cirrhatus recorded in the literature apparently resulted from a confusion of species. Giinther (1870) stated that the species had "six or seven gill openings on each side," but he listed specimens from South Africa {E. hexatrema Miiller, 1834) and Japan (£". burgeri Temminck and Schlegel, 1850). Species from these areas commonly have six pairs of gill openings. Referring to Eptatretus cirrha- tus, Waite (1909) stated, "The gill-openings ap- pear to be seven in number, but I have seen an [249] 250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 example in which there were but six openings on one side, though seven were present on the other." A variation of one per side is common in species having ten or more gill pouches, such as E. deani (Evermann and Goldsborough 1907) and E. St out a (Lockington 1878). Also, it is possible that Waite had an abnormal specimen in which two pouches shared the same opening (see above). Strahan's (1975) finding of "seven (rarely six) pairs of branchial apertures" for Eptatretus cir- rhatus may have been influenced by Gunther's or Waite's accounts. Regan (1912) listed a species with "7 gill open- ings: on each side two rows of 8 teeth. Southern Pacific" as Heptatretus banksii, and placed in its synonymy Homea banksii Fleming 1822, and Bdellostoma heptatrema Mtiller (1834). Regan's total count of 32 teeth is much lower than that of any of the four species treated herein (Table 6), and may indicate an erroneous count or an undescribed species. Regan may have counted three fused median teeth (multicusps) on each row as one, thus reducing the count to 32 from a possible 40. This would have been much nearer our minimal count of 43 for Eptatretus cirrhatus, under which we synonymize the above three names. Species of Eptatretus having seven gill aper- tures are not restricted to the Pacific Ocean. Fern- holm and Hubbs (1981) Usted a species having seven apertures from the Caribbean Sea. Fern- holm (1982) has further described it as new. In general, we concur with Femholm and Hubbs in terminology, with but minor varia- tions. We believe the term "dental muscle" is more appropriate than "tongue," "lingual," or "club-shaped muscle" in reference to the firm elongate complex of muscles and cartilages which constitutes the feeding mechanism of myxinids. Apparently the term "tongue" was first used by MUller (1834), but we concur with Ayers and Jackson (1900) that the entire apparatus in no way resembles a tongue. They stated, "The ho- mology of this organ with the vertebrate tongue has never been discussed, nor do we know of any effort to determine the true nature of this organ." Dawson (1963:248, fig. 11) provided a detailed analysis and figure of the structure, and of the "teeth" and "jaw apparatus." She concluded (p. 253) that it was unwise to make any definite assumptions concerning homologies of the car- tilages and muscles. There are two pairs of anterior and posterior sets (series) of sharply pointed, laterally flattened, homy structures in the oral cavity which are embedded in a dental plate. These structures cut and scrape food into ingestible portions when everted and retracted by the dental muscle. Al- though the term "teeth" has been widely used in reference to these structures, they are unlike the teeth of other vertebrates, being composed en- tirely of keratin and devoid of calcification. Daw- son (1963:247) concluded that, "It is most likely that there is no phylogenetic connection between these teeth and calcified teeth, and that they are an individual adaptation to a parasitic mode of life."' For descriptive and statistical purposes, we prefer the terms unicusps and multicusps to differentiate between single and composite teeth— the latter formed by the fusion of two or three unicusps. We consider the number and arrange- ments of both the multicusps and unicusps to be a significant species character. Materials Collection data and disposition of specimens examined in this study are listed in the treatment of each species. Institutions which have fur- nished study material, or in which type speci- mens have been deposited, are: Bemice P. Bish- op Museum, Honolulu, Hawaii (BPBM); United States National Museum, Washington, D.C. (USNM); Scripps Institution of Oceanography, La Jolla, California (SIO); California Academy of Sciences, San Francisco (CAS); Museum Na- tional d'Histoire Naturelle, Paris, (MNHN); University of the Philippines Zoological Mu- seum, Diliman, Quezon City, Philippines (UPZM); Australian Museum, Sydney (AMS); Zoological Institute, Academy of Sciences, Len- ingrad (ZIN). Methods The methods of measuring and counting de- scribed herein represent original methods as well as some used by prior authors including Dean (1904), Nani and Gneri (1951), Richardson (1953), and Strahan (1975). Femholm and Hubbs (1981) reported many of these methods in their study of the eastem Atlantic Eptatretus. When ' Hagfishes are not parasitic; they scavenge dead or mori- bund fishes and invertebrates. McMillan and wisner: new species of pacihc haghshes 251 the senior author, in collaboration with the late Carl L. Hubbs, began work on the myxinids (in 1969), it was obvious that no standard criteria existed for the study of hagfishes, which lack the jaws, opercula, rayed fins, scales, gill rakers, and bones found in most fishes. Early workers ap- plied different names to the same anatomical characters, defining them differently or not at all, and often not mentioning the methods used in measuring and counting. Therefore, it was dif- ficult to correlate or compare data of different authors, and taxonomic confusion resulted. We hope that the methods proposed and defined be- low will provide future investigators with a stan- dard by which hagfish species and specimens may be readily compared and identified. Proper treatment immediately after capture is of particular importance in rendering specimens suitable for study. Often too many live hagfish are crowded in jars of preservative, resulting in coiled or bent bodies, usually heavily coated with slime (mucus) and difficult to measure or count. The copious secretion of slime, characteristic of the family Myxinidae, is dramatically curtailed by prompt immersion in fresh water, preferably warm. This rapidly kills the hagfish and prevents further extrusion of slime, which otherwise con- tinues for several minutes even in formalin. Any remaining slime may be removed with paper or cloth towels, and the specimens should then be laid straight in a suitably large container of for- malin until fixed. If a specimen is too large for a flat pan, it should be coiled smoothly in a 3- 5 -gallon container, taking care not to deform the snout or twist the body, and covered with for- malin. This treatment produces fairly straight specimens with a minimal coating of slime, and greatly facilitates accurate counts and measure- ments. Since fresh hagfishes deteriorate rapidly, pres- ervation should be prompt. Color photos or notes should be made to record pigmentation, and tis- sue or blood desired for biochemical or chro- mosomal studies should be taken prior to im- mersion in formalin. We find that initial freezing prior to chemical preservation may cause soft- ening of the tissue and collapse of eggs and in- ternal organs, but it may be preferable to crowd- ing into a too-small container. Due to the many body openings, we consider it unnecessary to slit the skin or to inject preservatives; hagfishes are so soft that the skin may tear and some under- lying tissues may come apart, causing difficulty in subsequent measures and counts. Abbreviations PCD: external opening of the pharyngocuta- neous duct; ordinarily confluent with the pos- teriormost left gill aperture, and much larger than all other apertures. GA: gill (branchial) aperture; external opening of the efferent duct leading from a gill pouch. GP: gill pouch; rounded, serially arranged structures along and posterior to the dental mus- cle. DM: dental muscle; the firm, elongate, cylin- drical complex of muscles and cartilages that moves the dental plates and sets of cusps during feeding. Posterior portions of DM are shown in Figure 3. VA: ventral aorta; the portion between the heart (ventricle) and where it branches to each side of DM. ABA: afferent branchial artery; one of the small blood vessels that lead to each gill pouch from VA or its branches. Measurements If the specimen is distorted due to preserva- tion, it should be moderately straightened to ap- proximate its normal form. Measurements are taken from the left side with the fish lying on a meter stick; dividers or dial calipers are advisable for shorter lengths. We arbitrarily divided the body into four major sections (Fig. 1): prebranchical, branchial, trunk, and caudal. These are particularly apropos to genera Eptatretus and Paramyxine, as each has more than one GA, thus a branchial section. In Myxine, Neomyxine, and Nemamyxine, there is only one GA on each side, that on the left being confluent with PCD. Synonymous terms appearing in the literature are: "head" or "pectoral" for prebranchial, "gill" for branchial, and "abdominal" for trunk. The term "mucus" has often been used for slime, "teeth" for cusps, "tongue" or "lingual muscle" for dental muscle, and "outer" and "inner" for posterior and anterior in referring to the series of cusps. Body measurements we have found particu- larly useful are: Total length (TL): snout (anterior tip of ros- trum, excluding barbels) to posteriormost mar- gin of tail or caudal fin. 252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 Figure 1. A-B: Sketches of an Eptatretus and a Myxine, showing regions of body used in study of myxinids: 1 to 5, total length; 1 to 2, prebranchial; 2 to 3, branchial; 3 to 4, trunk; 4 to 5, caudal. C: sketch of head region of a myxinid, showing barbel pairs 1,2, and 3, and nasopharyngeal opening, 4. Preocular length: snout to center of eyespot, unpigmented area (if present) marking the ocular region. Prebranchial length: snout to front of first, or only, GA. Branchial length: front of first to front of last GA (PCD). The anterior edge of the last GA is used because the posterior margin is often too vague and poorly defined to provide a definite reference point. Trunk length: front of PCD to origin of cloaca. Body width: maximum dimension about mid- way between rostrum and PCD. Body depth: maximum vertical depth in trunk region, including finfold if present; depth ex- cluding finfold should be taken at the same place. In both width and depth measurements the body should be molded into a seemingly natural shape if necessary. Depth at cloaca: vertical depth at origin of cloaca. Tail depth: maximum vertical depth of flat- tened tail, with any roll-up or fold of the thin tail margin uncurled and flattened. Barbel length: from center of base to tip of each barbel (Fig. 1). The distance between bases of each pair may be measured from the inside edge of each base. Barbels are often curled and difficult to measure accurately, but in certain species barbel length may be a significant char- acter, and is worth measuring. Dental muscle length (DM): snout to tip of DM, as revealed by a midventral incision in the prebranchial region. Dental muscle width: measured at a straight- sided portion well anterior to tapering end. Dental muscle depth: measured at same place as width measure. Rather than using the total McMillan and wisner: new species of PACinc haghshes length, we have found it convenient to compare the length (or width) with the unbranched por- tion of the VA with measurements of the DM. This is a significant ratio in certain species, but varies greatly between specimens of other species. Weight: may be taken, but we have not found it to be a reliable or useful character, principally because of the uncertainty in determining if all the entrapped fluid was drained, and because of dehydration of body fluids during preservation. Counts Ordinarily the branchial openings (GA) are the first items examined to ascertain the genus and possible species. The gill pouches are usually counted after the teeth (cusps) when the oral cav- ity incision is extended midventrally to the re- gion of the PCD. Before counting the slime pores, we gently scraped away any coagulated slime overlying the line of pores; an air jet greatly fa- ciUtated location of pores. Because so few spec- imens were available for this study, both sides were counted to obtain wider range of variation. Counts we have found particularly useful are: Slime pores: Prebranchial— from anteriormost slime pore to last one before first GA. Branchial— those pores in immediate associ- ation with (usually below and to the right of) each GA; often one less than GA count in Ep- tatretus, and much less, or absent entirely, in Paramyxine. There is usually no slime pore as- sociated with PCD, but this varies with species and individual specimens. In this study all species except E. strahani have a branchial pore count equal to or higher than the number of GA; the extra pores vary in location and number. Trunk— the series posterior to PCD and ter- minating anterior to end of cloaca, distinctly sep- arate from cloacal series. Cloacal— the pores distinctly before a vertical from posterior end of cloaca, usually starting somewhat anterior to and elevated from origin of cloaca. Caudal— from first pore distinctly behind a vertical from posterior end of cloaca to last pore on tail. For statistical purposes we combine counts of cloacal and caudal pores under the heading "tail pores" (Table 2). Cusps (teeth): We refer to a single "tooth" as a cusp, or unicusp, if it is not fused to one or more adjoining cusps. A unit of two or more cusps fused together at some point prior to its Figure 2. Cusps and basal plates, in excised and spread condition, of £. carlhubbsi, paratype USNM 233742, 955 mm TL. embedment in the cartilaginous dental plate is a multicusp. The two paired sets of cusps (the outer and inner rows of Femholm and Hubbs [1981] and Femholm [1982]) are examined from the ventral aspect. They are revealed by a midline incision from the base of the oral cavity through the car- tilaginous pharynx until the sets are free and eas- ily turned outward for viewing. There are dis- advantages to this method. It is easy to misjudge the midline (if the "face" has been distorted in preservation) and cut through the median teeth, making counts difficult; also, the resulting view presented to the observer is a reversed image of the actual arrangement. The inner left row ap- pears on the outer right side and vice versa. To avoid this confusion, the incision may be made from either side of the oral cavity to just under the third barbel, then extended laterally down- ward through the thin membrane, exposing the paired sets of cusps which, when spread apart, appear as shown in Figure 2. On most specimens the count of multicusps may be determined by placing a dissecting or air jet needle under the first two cusps and gently PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 L Figure 3. Ventral view of branchial region of: 1, E. carlhubbsi: 2-3, E. laurahubhsi, showing diversity in afferent branchial arteries (ABA) leading off from branches of ventral aorta (VA); 4, E. strahani; 5, E. cirrhatus. lifting; the multicusp usually lifts and separates from the adjacent unfused cusp. However, in the nine largest specimens examined by us {E. carl- hubbsi), lifting often raised the entire dental plate and set of cusps. Even if cusps are unquestion- ably fused, a line may extend among the fusion to the plate or "gum line;" in such instances per- haps the only valid criterion for separating mul- ticusps from unicusps is the distinctness of this line as seen under magnification. Such lines are in marked contrast to the condition shown by scanning electron microscopy of E. springeri (Femholm and Hubbs 1981: fig. 2), wherein no lines are evident in the multicusps. Gill apertures and pouches: In genera Myxine, Neomyxine, Nemamyxine, and Notomyxine, dissection is necessary to determine the number of gill pouches, since only one pair of efferent ducts leads to the exterior. A midventral incision is made from the single pair of GA anteriorly McMillan and wisner: new species of pacihc haghshes 255 until all pouches are revealed (Fig. 3). The cut should be deep enough to expose VA and ven- tricle, taking care not to sever branches of VA or any ABA, or to destroy the origin of the ventral finfold if it is present anterior to PCD. There are multiple, readily visible GA in gen- era Eptatretus (5-15 pairs) and Pammyxine (5- 7 pairs). Although the number of internal pouch- es ordinarily is the same as the external apertures, there may be variation; thus, it is desirable to count the pouches and examine the arrangement of the GP relative to DM and branched and un- branched portions of VA (Fig. 3). The arrange- ment is often of taxonomic importance, although variation occurs (see E. laurahubbsi). Sensory canals (lateral lines): Ayers and Wor- thington (1907:331, figs. 5-10), in a study of the skin-end organs of the trigeminal and lateralis nerves of Bdellostoma dombeyi {=Eptatretus stoutii [Lockington 1878]), described and figured lateral line canals, associated dermal grooves, and nerve endings. They showed the canals as short lines occurring only dorsally and somewhat lat- erally on the "head" and in two groups, one be- fore and one behind the eyespots. Plate (1924: 66, fig. 6 1 D) accepted the interpretation by Ayers and Worthington that the short lines constituted lateral line canals, but considered the dermal grooves to be artifacts. Ross (1963:155) cited both these studies and stated that the lateral lines had not been described in Myxine glutinosa. To our knowledge these are the only prior references to lateral line canals of hagfishes. We concur with Ayers and Worthington that the canals occur only on the head (in the ocular area of the prebranchial region). However, they are lateral only in that a few occur on the side of the head, with most on the dorsal surface (Fig. 4), and none at all on the rest of the body. As- suming that the canals are indeed sensory in function, we prefer the term "sensory" to "lat- eral." Sensory canals occur in only two of the four species discussed here (£". strahani and E. cirrhatus. Fig. 4), but not on all specimens, and are irregular in number and form. The erratic occurrence in location and in numbers of canals is intriguing, as is their total absence in two of the four species. Due to the limited number of specimens avail- able, it is difficult to draw any firm conclusions regarding the taxonomic value of sensory canals. Ayers and Worthington (1907) stated that these canals were difficult to find because they were \ ^'■ \ /• E. cirrhatus /'"'''^ '3;;\ /'ngeal orifice about equal to length of third barbels. First pair of barbels about 63% of length of third pair; sec- ond pair about 75%. Color notes were not taken at time of capture (December 1965); all specimens are now a uni- form yellowish color, no doubt a result of fading. The eyespots are no longer discernible, but pre- ocular measurements were recorded by the se- nior author in November 1973. Removal of in- tegument over the right eye of a 287-mm specimen shows the eye to be round, about 2.5 mm in diameter, with a small triangular pupil with its base dorsad and slanting forward at a slight angle to horizontal axis of body. No sen- sory canals are evident on any specimen. Despite the faded condition of all specimens, the branchial apertures and most slime pores have whitish borders. Usually one pore, plus an oc- casional extra one, occurs adjacent to each BA. Two pores are near the opening of PCD on four specimens; three have one pore, and one has none near PCD (as is the usual condition on other Figure 5. Tail shapes (to scale) and patterns of occurrence of the last four trunk pores and cloacal and tail slime pores of four species ofseven-gilledfpmrre/Mi from the Pacific Ocean: \-9 E.carlhubbsii(l = tan color. P = purplish color); 1— Holotype. SIO 68-473, 961 mm TL; 2-9 Paratypes: 2-SIO 68-473, 813 mm TL; 3-USNM 227440, 900 mm TL; 4-USNM 233742. 955 mm TL; 5-CAS 50705, 1064 mm TL; 6-BPBM 27848, 1 160 mm TL; 7-CAS 50706. 908 mm TL; 8-SIO 82-63. 1 125 mm TL; 9-BPBM 27851, 830 mm TL; 10-Holotype, E. laurahubbsi. SIO 65-643, 375 mm TL; 1 1 -Holotype, E. strahani. MNHN 1978-462, 520 mm TL; \1-E. cirrhatus. 655 mm TL. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 Eptatretus). Space between the last trunk pore and first cloacal pore is about equal to length of cloaca (Fig. 1). Two or three slime pores lie over cloaca in a straight line and equally spaced with caudal pores. Prebranchial pores in a fairly straight line; occasionally the first one to three slightly depressed. There is great variation in the arrangement of GP and afferent branchial arteries (ABA) with respect to the DM and branching of VA (Figs. 3-2 and 3-3). The number of GP along DM are far more variable than in the other three species, ranging from two to five. Also, length of VA varies notably; in six specimens VA averages 8.1% (6.3-9.3%) of length of DM, but in one 240-mm specimen the length of VA was 14.4% of DM, with three ABA leading off the left side and two off the right. This variation is in marked contrast to the regular arrangement of the branchial ap- paratus of the three other species discussed herein. The eight specimens off. laurahubbsi are un- usual within genus Eptatretus in the appearance of the multicusps, apparently having only two fused cusps in each series. Indeed, it is often a highly subjective decision as to whether any of the anteriormost cusps are fused as multicusps. One specimen appears to have three fused cusps in each of the anterior series and two in the pos- terior series. In two specimens it is questionable as to whether two or three cusps are fused in the posterior rows. In other Eptatretus known from the southern hemisphere the usual configuration is three fused cusps on each of the four multi- cusps; all Eptatretus known from the North American Pacific coast have three in the anterior and two in the posterior row, which are distinct and clearly seen even in juveniles. A juvenile (188 mm) E. stoutii (Lockington 1878) clearly shows hard, well-developed cusps with the pat- tern 3/2. Also, a 100-mm specimen of £■. poly- trema (Girard 1855) from Chile has three dis- tinctly fused cusps in each multicusp; adults of this species attain a total length of at least 550 mm. Thus, the presence of only two fused cusps in most specimens of E. laurahubbsi, and the uncertainty regarding the number fused in the others, is apparently not due to immaturity or small size. All of our study specimens are juveniles, but it is highly probable that adults exceed one meter in length. The longest (holotype), although 373 mm TL, contains minute eggs, seen with diflfi- culty under magnification. In another Eptatretus (undescribed) from the Gulf of California, a spec- imen of this size may have fully developed go- nads and large eggs. Gumersindo Revuelta, a for- mer student at the University of Chile, Valparaiso, in an unpublished thesis (1976), sketched and gave limited data on a very large hagfish, slightly exceeding a meter in length (from scale provided with sketch). He tentatively iden- tified it with the giant Eptatretus taken at Wake Island (from data sent to him by Hubbs), prob- ably because of its gigantic size compared to oth- er species from Chilean waters. Revuelta had at least two females, both apparently very large, for he sent to Hubbs (in 1976) two large eggs: one 68 by 16.5 mm from Valparaiso, and one 72 by 16.8 mm from "Juan Fernandez" (presumably the island). We presume these large specimens to be adults of E. laurahubbsi because Revuelta reported the multicusps as 2/2, and his limited data are in close agreement to those of our spec- imens taken in the same vicinity. Also, in E. carlhubbsi the ventral finfold is entirely absent, but Revuelta's sketch shows a ventral finfold originating a little behind the anterior third of the body. In our juveniles a pronounced finfold originates variably between the midpoint and an- terior third of the body; the exact point of origin is indeterminate because of wrinkling during preservation. Etymology.— We dedicate this unique species to our friend and and co-worker, Laura Clark Hubbs, who contributed in so many ways to the life and works of her husband, Carl Leavitt Hubbs. Eptatretus strahani new species Holotype. -MNHN 1978-462, female, 520 mm TL, con- taining eggs of about 4.5 mm in length, taken at 14°00'N, 1 20°1 8'.2"E, South China Sea near Lubang Island, Philippines, in a trap net at 189 meters. Station 22 Musorstom Expedition, 21-22 March 1976, 1800-0600 hours. Paratypes (and remaining material examined; all taken with the holotype).-MNHN 1981-722, female, 420 mm TL; SIO 81-116, female, 265 mm TL, male, 450 mm TL; USNM 227442, male, 465 mm TL. Additional Material. -UPZM 1981-809, 400 mm TL; UPZM 1981-811, 480 mm TL. Total lengths, comparisons, and identifications were made by Prof. Reynaldo de La Paz, University of the Philippines, Diliman, Quezon City, Philip- pines, based on methods and data provided by us. Diagnosis.— A seven-gilled Eptatretus having no eyespots, a well-developed ventral finfold, and three fused cusps on each of the four multicusps. Description. — Counts: Those of the holotype given first, followed by ranges in parentheses for McMillan and wisner: new species of pacihc hagfishes 263 all five specimens (both sides counted): gill ap- ertures 7, 7 (all); prebranchial slime pores 14. 16 (13-16); branchial pores 6, 6 (6-7); trunk pores 45, 47 (45-48); cloacal pores 4, 3 (3-4); caudal pores 7, 7 (6-8); total tail pores 11,11(10-12); total slime pores 76, 79 (76-80). Cusps on mul- ticusps 3, 3 (all); anterior unicusps 11, 11 (9-11); posterior unicusps 9, 9 (8-10); total cusps 52 (47- 52). Morphometry: Values for holotype given first, fiallowed by ranges for all five specimens, in thou- sandths of total length: preocular length (no eye- spots); prebranchial length 231 (210-231); bran- chial length 81 (69-83); trunk length 500 (500- 537); tail length 196 (174-202); body depth in- cluding finfold 115(101-117); excluding finfold 95 (94-105); body depth over anterior margin of cloaca 88 (77-94); tail depth 119 (109-125). Morphometry (Table 1) and counts (Tables 2- 6) are compared with similar data for other sev- en-gilled Eptatretus from the Pacific Ocean. Body deeper than wide, deepest at midsection. Ventral finfold well developed, extending from about midbody to cloaca, its length about 31% of TL. Tail margin quite thin posterior to cloaca, extending around tail to dorsal surface, ending at about a vertical from posterior end of cloaca. Ventral outline of tail forms a nearly straight line, ending with an abrupt curvature up and around end to dorsal aspect. This shape is in marked contrast to the gradual curvature of tails of the other three species treated herein (Fig. 5). Dorsal profile of head sloping steeply to snout; rostrum more rounded than in E. carlhubbsi or E. laurahubbsi; width of nasopharyngeal opening about 60-80% of length of first pair of barbels. First two pairs of barbels nearly equal in length; respectively, about 66% and 72% of length of third pair. First barbel, left side, of a 420-mm female is bifurcate to near base, with posterior branch shorter. Since we have occasionally seen this bifurcation in other hagfishes (usually near the tip, and always on only one barbel of the six), we assume this form is the result of an injury rather than some genetic malformation. No eyespots are visible on any specimen (about 30 months after capture). Since the body color is still fairly dark we have assumed that little or no fading has occurred, and that the unpig- mented eyespot area should still be visible if present in life. No notes regarding eyespots were made at time of capture. Removal of overlying integument on holotype shows eye to be ovate (3.4 by 2.4 mm) and slanted ventrodorsally at about a 45° angle; pupil more rounded (1.4 by 1.1 mm). Sensory canals are present in two groups on each side of the head before and behind the area where eyespots normally occur (Fig. 4). One group of sensory canals is found near the bases of the third pair of barbels, anterior to embedded eyes, another group slightly posterior to eyes. Anterior group consists of five more-or-less longitudinal lines 1-3 mm long; canals of posterior group both longitudinally and horizontally arranged, those on top of head tending to be more horizontal. No canals extend across dorso-medial line. Color of holotype (in preservative) a light brown, all paratypes a darker brown, the smallest the darkest. No discernible whitish rings around slime pores or GA on larger specimens, but the GA of the smallest one has distinctly pale mar- gins. Finfold anterior to cloaca is same color as body, but tail has a very narrow, pale margin extending a short distance forward on the dorsal surface. The line of the anterior prebranchial slime pores is straighter in this species than in the other three discussed; two specimens have only slight cur- vature, and no anterior pores are markedly ele- vated above adjacent ones in the prebranchial series. Space between last trunk and first cloacal pore about 65% of length of cloaca; spacing is variable with degree of slant or elevation of first cloacal pore (Fig. 5). Cloacal pores form a distinct dorsoventral slant on left side of holotype, but not on right; slanting is variable on paratypes. Most GA are shaped as slits, slanting ventro- dorsally; this shape could be an artifact of pres- ervation, but the GA may be made to assume a rounded form only by considerable pulling and squeezing of surrounding flesh; all the slime pores below GA are rounded. Three to five GP lie anterior to tip of DM (Fig. 3); two to four lie between that tip and branching of VA, and none posterior, although one GP of the smallest specimen (265 mm TL) lies just at the branching. Length of VA 6.4% (5.4-7.6%) of DM length; DM length 26% (25-27%) of TL, its width 15% (13-16%) of its length. Distance be- tween tip of DM and branching of VA 9% (7.2- 10.9%) of DM length. Etymology. — We are pleased to dedicate this new species to Ronald Strahan in acknowledg- ment of his important contributions to the study of Myxinidae. 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 Eptatretus cirrhatus (Bloch and Schneider, 1801) Petromyzon cirrhatus Bloch and Schneider, 1801:532 (orig- inal description fide Forster ms 11:24 [habits: New Zealand]). Hornea banksii Fleming, 1822:374 (South Seas [presump- tive]). Bdellostoma Forsteri Muller, 1834:71, 80 (anatomy; char- acters in key; reference to Petromyzon cirrhatus Bloch); Schneider, 1880:115 (status uncertain; based on a poor specimen). Bdellostoma heptatrema Muller, 1 834:7 (original description; New Zealand). Bdellostoma cirrahatum Gunther, 1870:511 (synonymy, in part; diagnosis, in part; distribution [New Zealand only]); HuTTON, 1872:87 (characters; color reddish brown, white around mouth; common Australia and South Africa [mis- identified with E. heptatrema]); Putnam, 1874:156 (in part; New Zealand; 7 gill slits); Schneider, 1880:115 (in part; doubts on status); Adam and Strahan, 1963:6 (6 or 7 pairs of gills; average length 480 mm; South Pacific, common off New Zealand). Homea cirrhata Garman, 1899:341, 345, 349, 419 (synony- my; nomenclature); Dean, 1 904:2 1 (in part; synonymy; New Zealand). Heptatrema cirrata [sic] Hutton, 1904:55 (listed; New Zea- land). Eptatretus cirrahatus Berg, 1906:173 (in part; New Zealand); Waite, 1909:2 (description; behavior; average length 680 mm; Timaru and Chatham Islands; New Zealand); Graham, 1965:67 (plentiful on North Otago Shelf New Zealand; often takes baited hooks); Heath and Moreland, 1 967:30 (shore to 1800 ft; more abundant south of Hawke Bay than else- where in New Zealand); Whitley, 1 968:4 (synonymy); Scott, Glover, and Southcott, 1 974: 1 9 (New Zealand, New South Wales, S.E. Australia); Fernholm, 1974:351 (in shallow water. New Zealand); Fernholm and Holmberg, 1975:253 (struc- ture of eye, comparative; Kaikoura, S. Island, New Zealand); Strahan, 1975:145 (key; description; ranges of counts and body proportions). Heptatretus banksii, Regan, 1912:534, 536 (comparisons; syn- onymy; diagnosis; D'Urville Is., Queen Charlotte Sound, New Zealand.). Material Examined (counts and measurements both tak- en).-SIO 81-105, two males, 488 and 655 mm TL, three females 48 1-636 mm TL, 42°24'S, 1 73°4 1 'E, no data on depth or date of capture, received from J. A. F. Garrick, Zoology Department, Victoria University of Wellington, New Zealand, 1 Nov. 1972; SIO 62-482-4A, two females, 577 and 580 mm TL, received from L. R. Richardson, Wellington, New Zealand, 25 March 1959, no data on depth of capture; ZIN 717-966, male, 595mmTL,40''19'S, 172°15'E, 160-172 meters, 18 Jan. 1965. Counts only taken: AMS I 15527-001, three males, 254- 452 mm TL, 26°32'S, 153°5rE, agassiz trawl, 175 fms (320 m), 27 July 1968; AMS Kapala Station 71-07-03, female, 505 mm TL, 33''33'-37'S, 152''0r-151°57'E, 205 fms (375 m), 21 April 1971; AMS Kapala Station 71-08-05, male, 265 mm TL, female, 552 mm TL, 33°1 1'S, 152°23'E, otter trawl, 310 fms (567 m), 29 April 1971; AMS Kapala Station 71-11-07, two females, 49 1 , 5 1 4 mm TL, 34''40'-35°0 1'S, 1 5 1°1 0'-07'E, otter trawl, 300 fms (549 m), 7 July 1971; AMS Kapala Station 71- 1 1-08, female, 410 mm TL, 34°56'-35''02'S, 151°06'-05'E, ot- ter trawl, 160 fms (194 m), 8 July 1971; AMS Kapala Station 71-11-10, male, 447 mm TL, 35°1 1 '-37'S, 1 50°45'-42'E, otter trawl, 230 fms (421 m), 8 July 1971; AMS Kapala Station 71- 12-04, female, 546 mm TL, 33°41'-49'S, 151°53'-47'E, otter trawl, 245-250 fms (448-457 m), 20 July 1971; AMS Kapala Station 71-12A-06, three females, 374-503 mm TL, 35°25'- 29'S, 150°50'-48'E, otter trawl, 300 fms (549 m), 2 Aug. 1971. Diagnosis.— A seven-gilled Eptatretus having a vestigial ventral finfold, small but prominent eyespots, white around the mouth, pale rings around branchial apertures and slime pores; three fused cusps on each multicusp; sensory canals may occur. Description. — Despite its being the first hag- fish described from the Pacific Ocean, the liter- ature contains minimal data on morphology and counts. Waite (1909) and Graham (1965) pro- vided descriptions and accounts of behavior, but no meristic data. Strahan (1975) listed only ranges of counts and percentages of total length for cer- tain body measurements for 13 specimens. We offer morphometry based on only the eight spec- imens available to us, but include counts on 22 specimens examined by Carl L. Hubbs in 1971 at the Australian IVIuseum, Sydney. Counts: Averages followed by ranges in pa- rentheses, both sides counted: gill apertures 7 (all); prebranchial slime pores 17-18 (16-20); branchial pores 7 (6-8); trunk pores 48-49 (46- 53); cloacal pores 3 (2-5); caudal pores 9 (6-1 1); total tail pores 12 (10-14); total sHme pores 86 (83-90). Three fused cusps on each of the four muhicusps; anterior unicusps 9 (8-1 1), posterior unicusps 8 (7-9); total cusps 46 (43-51). Morphometry: Averages followed by ranges in parentheses, in thousandths of TL, for eight spec- imens: preocular length 60 (52-67); prebranchial length 225 (214-239); branchial length 76 (69- 89); trunk length 550 (525-563); tail length 154 (135-168); maximum body depth including fin- fold 93 (84-102); excluding finfold 91 (81-102); body depth over cloaca 67 (57-74); tail depth 82 (77-9 1); body width at mid-prebranchial section 50 (46-55). Dorsal profile of head sloping gently down- ward, face sloping at about a 45° angle; naso- pharyngeal opening about equal to or slightly less than length of first barbel. Average length of first pair of barbels about 69% and second pair about 75% of the length of the third pair. Body rounded anteriorly, becoming more laterally compressed and deeper posteriorly; tail bluntly rounded, spatulate, its depth slightly less than greatest depth McMillan and wisner: new species of pacihc haghshes 265 of body (Table 1). Ventral finfold narrow and relatively short, its length about 30% of TL, ex- tending from well behind midbody to cloaca. Color of our specimens in preservative varies from light chocolate to dark brown; the most lightly colored one is strongly mottled anteriorly with small, irregular pale spots and patches. Waite (1909) stated that "the colour varies from blue to bluish violet [presumably fresh material]. Some examples show irregular white spots and mark- ings; the ventral finfold and the margin of the tail may also be white." On our material, pre- served at least ten years prior to our examination, the poorly developed finfold has a narrow pale margin on two specimens, but not on the other six; color is highly variable. Tail of one specimen is irregularly margined with pale areas; pale rings around most of the gill apertures and slime pores. Small but plainly visible eyespots are present on all specimens. Removal of integument cov- ering right eyespot of a specimen 577 mm TL reveals the embedded eye as elliptical (6.3 by 3.2 mm) and slanting forward ventrodorsally at about a 45° angle; pupil small, more rounded (about 1.9 by 1.4 mm). Sensory canals, only as posterior series (in the specimens available to us), are located close be- hind eyespots (Fig. 4). Canals are readily iden- tifiable on only three of eight specimens (488- 655 mm TL) and only on the left side of largest one; they are similar in size, form, and arrange- ment to those of E. strahani. On two specimens (580 and 597 mm TL) positive identification of canals is prevented by presence of much scarring. The anterior prebranchial slime pores on all specimens form a downsloping curve; a high in- cidence of irregular spacing of these pores occurs. On left side of four specimens, first or third pore is notably elevated or very closely spaced. On one specimen four pores form an almost-square pattern; all apparently lead from a single slime gland. In most Eptatretus from the Pacific Ocean, the usual number of branchial slime pores is one less than the number of GA, since ordinarily there is no pore associated with the PCD. In E. cirrhatus there is a high incidence of extra pores and irregular spacing of slime glands in the bran- chial region, especially near the PCD. Posterior two or three GA, left side, curve gently downward on seven of eight specimens, but only on the right side of one. On a 480-mm juvenile female the normally confluent openings of the seventh GA and PCD are distinctly separate; such separation also occurs infrequently in other species of Eptatretus. Usually five, occasionally four, GP lie anterior to tip of DM. Length of DM 27% of TL, its width 13% (12-14%) of its length; somewhat flattened posteriorly, depth is 80% (75- 88%) of its width. The shape of the tip of the DM varies somewhat from that of the other three species discussed herein (Fig. 3). Distance be- tween tip of DM and branching of VA about 7.4% (4.8-10.8%) of DM length; VA usually greater in width than in length, in contrast to that of the other three species. No ABA lead off"before branching of VA in the eight specimens available to us. A 655-mm TL female contains about 50 large eggs, ranging from 29 to 32 mm long by about 10 mm in diameter; this is an unusually large number of maturing eggs. All are still in the mes- entery which is attached to the body wall; ter- minal anchor filaments and hooks are not present on any egg. Acknowledgments We are deeply grateful to the late Carl L. Hubbs for his efforts in amassing much of the material on hagfishes, without which this study might never have been accomplished. Also, we are most grateful to the following persons and agencies: Jacques Forest, Museum National d'Histoire Naturelle, Paris, and Professor Reynaldo M. de La Paz, University of the Philippines, Diliman, Quezon City, for material of E. strahani; J. A. F. Garrick, Victoria University, Wellington, New Zealand, and L. R. Richardson, Wellington, for specimens of E. cirrhatus; and Richard H. Ro- senblatt, for the holotype and one paratype of £■. carlhubbsi. Other material of E. carlhubbsi was provided by Steven Kramer, National Marine Fisheries Service, Hawaii; Robert Moffitt, NMFS Field Station, Mangilao, Guam; Paul Strushaker, Honolulu; and John E. Randall and Arnold Su- zumoto, Bemice P. Bishop Museum, Honolulu. All specimens of £". laurahubbsi were taken by the junior author on Cruise 1 2, R/V Anton Bruun, Southeastern Pacific Biological Oceanography Program, sponsored by the National Science Foundation. The holotype and one paratype of E. carlhubbsi were taken on the Scripps Styx ex- pedition, sponsored by the National Science Foundation through a grant (GB-7596) to R. H. Rosenblatt and W. A. Newman. We greatly ap- 266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 predate the efforts of all persons involved. R. H. Rosenblatt critically read the manuscript. References Cited Adam, H., and R. Strahan. 1963. Notes on the habitat, aquarium maintenance, and experimental use of hagfishes. Chapter 1, pages 1-8 in The Biology of Myxine. A. Brodal and R. Fange, editors. Universitetsforlaget, Oslo. 588 pp. Ayers, H., and C. M. Jackson. 1900. Morphology of the Myxinidae 1. Skeleton and musculature. Bull. No. 1, Uni- versity of Cincinnati 185-224. , AND J. Worthington. 1907. The skin-end organs of the trigeminal and lateralis nerves of Bdellostoma dom- beyi. Am. J. Anat. 7:327-336. Berg, L. 1 906. Ubersicht der Marsipobranchii des Russisch- en Reiches. Bull. Acad. Sci. de St. Petersbourg, V' Ser., 24(3): 173-197. Bloch, M. E., and J. G. Schneider. 1801. Systema ichthy- ologiae iconibus ex illustratium. Post obitum auctoris opus inchoatum absolvit, correxit, i interpolovit Jo. Gotlob Schneider Saxo, Berlin: 1-584. Dawson, J. A. 1963. The oral cavity, the "jaws," and the homy teeth of Myxine glutinosa. Chapter 5, pages 231-255 in The Biology of Myxine. A. Brodal and R. FSnge, editors. Universitetsforlaget, Oslo. 588 pp. Dean, B. 1899. Onlhe emhryo\ogy of Bdellostoma stoutii. A general account of myxinoid development from the egg and segmentation to hatching. Fest. zum Siebensigsten Ge- burstag von Carl von Kuppfer, Jena. 221-276. . 1 903. Albinism, partial albinism, and polychromism in hagfishes. Am. Naturalist 37(437):295-298. . 1904. Notes on Japanese myxinoids. A new genus, Paramyxine, and a new species, Homea okinoseana, refer- ence also to their eggs. Jap. Coll. Sci., Imperial Univ., Tokyo 19(2):l-23. EvERMANN, B. W., and E. L. Goldsborough. 1907. The fishes of Alaska. Bull. U.S. Bur. Fish. 26(for 1906):2 19-360. Fernholm, B. 1974. Diurnal variations in the behavior of the hagfish, Eptatretus burgeri. Mar. Biol. 27:351-356. . 1982. Eptatretus caribbeaus: A new species of hagfish (Myxinidae) from the Caribbean. Bull. Mar. Sci. 32(2):434- 438. , AND K. Holmberg. 1975. The eye in three genera of hagfish {Eptatretus, Paramyxine, and Myxine)— & case of degenerative evolution. Vision Research 15:253-259. , AND C. L. HuBBS. 1981. Western Atlantic hagfishes of the genus Eptatretus (Myxinidae), with descriptions of two new species. Fish. Bull. 79(l):69-83. Fleming, J. 1822. The philosophy of zoology; or a general view of the structure, functions, and classification of animals. Fishes. Edinburgh 2:305-397. Garman, S. 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands in charge of Alexander Agassiz, by the U.S. Fish Commission SXt&mtr Albatross d\mng 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. XXVI. The Fishes. Mem. Mus. Comp. Zool. 24:1-431. GiRARD, C. F. 1855. Abstract of a report to Lieut. Jas. M. Gillis, U.S.N., upon the fishes collected during the U.S.N. Astronomical Expedition to Chile. Proc. Acad. Nat. Sci. Philadelphia 7(11): 197-1 99. Graham, J. 1965. The North Otago Shelf fauna. VI. Chor- data, subclass Cyclostomata. Trans. Royal Soc. New Zealand 6(6):67-68. GiJNTHER, A. 1870. Catalogue of the Physostomi, containing the families Gymnotidae, Symbranchidae, Muraenidae, Pe- gasidae, and of the Lophobranchii, Plectognathi, Dipnoi, Ganoidei, Chondropterygii, Cyclostomata, and Leptocardii, in the British Museum. Catalogue of the fishes in the British Museum, London 8:1-549. Heath, E., AND J. MoRELAND. 1967. Marine fishes of New Zealand. A. H. and A. N Reed, Wellington, Auckland, and Sydney. 1-56. HuTTON, F. W. 1872. Fishes of New Zealand: Catalogue with diagnosis of the species. Colonial Museum and Geological Survey Dept., Wellington. 1-93. . 1904. Index Faunae Novae Seelandiae. Dulau and Co., London. 1-372. Jensen, D. 1959. Albinism in the California hagfish, Epta- tretus stoutii. Science 130(3378):798. Jespersen, A. 1975. Fine structure of spermiogenesis in east- em Pacific species of hagfish (Myxinidae). Acta Zool., Stock- holm 56:189-198. LocKJNGTON, W.N. 1878. Walks around San Francisco. No. Ill -Lake Honda and Seal Rock. Am. Nat. 12:786-793. Muller, J. 1834. Vergleichende Anatomic der Myxinoiden, der Cyclostomen mit durchbohrtem Gauman. Erster theil. Osteologie und Myologie. Abhandlungen der Koniglichen Akademie der Wissenschaften zu Berlin, Jahre, 1834, 1836: 65-340. Nani, a., AND F. S. Gneri. 1951. Introduccion al estudio de los mixinoideos Sudamericanos. I. Un nuevo genero de "Ba- bosa de Mar," "Notomyxine" (Clase Myxini, Familia Myxinidae). Rev. Inst. Nac. Invest. Ciencias Nat. 1(4):183- 224. Plate, L. 1924. AUgemeine Zoologie und Abstammungs- lehre. Teil 2. Die Sinnesorgane der Tierre, Jena. 1-806. Putnam, F. W. 1874. Notes on the genus Bdellostoma. Proc. Boston Soc. Nat. Hist. 16:156-160. Regan, C.T. 1912. A synopsis of the myxinoids of the genus Heptatretus or Bdellostoma. Ann. Mag. Nat. Hist., ser. 8, 9: 534-536. Revuelta, G. 1976. Informe de datos y observaciones: Re- vision del genero Eptatretus en Chile (Agnatha: Myxinidae). Unpublished thesis. Departamento de Oceanologia, Uni- versidad de Chile, Valparaiso. Richardson, L. R. 1953. Neomyxine. n.g. (Cyclostomata) based on Myxine biniplicata Richardson and Jowett, 1951, and further data on the species. Trans. Royal Soc. New Zealand 31(3):379-383. Ross, D. M. 1963. The sense organs of Myxine glutinosa L. Chapter 2, pages 150-160 in The Biology of Myxine. A. Brodal and R. FSnge, editors. Universitetsforlaget Oslo. 588 pp. Schneider, A. 1 880. Uber die Arten von Bdellostoma. Arch. Naturgesch. 46(1):115-116. Scott, T. D., C. J. M. Glover, and R. V. Southcott. 1974. The marine and freshwater fishes of South Australia (2nd ed.). Gov't. Printer, S. Australia. 1-329. j Strahan, R. 1975. Eptatretus longipinnis, n.sp., anew hag- m fish (family Eptatretidae) from South Australia, with a key ) to the 5-7 gilled Eptatretidae. Australian Zool. 1 8(3): 137- 148. McMillan and WISNER: new species of PACIHC HACnSHES 267 Temminck, C. J., andH. Schlegel. 1842-1850. Pisces. Pages Government trawling expedition, 1 907. Rec. Canterbury Mus. 1-345 in Fauna Japonica. Siebold, Fr. de. A. Ams et Socios, 1(2): 1-26. Lugdini. Plates and Suppl. Batavorium {=Leiden). Whitley, G. P. 1968. A check-list of the fishes recorded Waite, E. R. 1909. Scientific results of the New Zealand from the New Zealand region. Australian Zool. 15:1-102. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 941 18 PROCEEDINGS OF THE *^ CALIFORNIA ACADEMY OF SaENCES Vol. 43, No. 17, pp. 269-282, 5 JCES I JAN 7 li, December 11, 1984 NEW AND NOTEWORTHY ADDITIONS TO THE MELASTOMATACEAE OF PANAMA By Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: Descriptions, discussions, diagnostic illustrations, and distributional notes are provided for five new species of Melastomataceae (Clidemia tetrapetala, C. trichosantha, Graffenrieda bella, Miconia arboricola, and Topobea calophylla). Based on a study of the first known flowering collections, Topobea elliptica is transferred to Blakea and provided with an amplified description. Notes are also presented on the size and delimitation of these genera with special reference to Panama. Introduction The first and only floristic treatment of the Panamanian Melastomataceae was published some 25 years ago (Gleason 1958). The manu- script for that study was completed several years before it appeared in print. Gleason's study is a very useful account that unfortunately suffers from a lack of adequate field study and a paucity of local material for many of the species attrib- uted to Panama. The many new collections of Melastomataceae generated by the heightened botanical exploration of Panama during recent years has underscored the need for an updated supplement. Such an undertaking is still pre- mature in view of the many undescribed taxa represented by collections that are too incom- plete to serve as a basis for formal descriptions. In this interim report I propose new species in Clidemia, Graffenrieda, Miconia, and Topobea, present a new combination in Blakea, and pro- vide brief notes on the size and delimitation of these genera in Panama. Blakea P. Browne Blakea and the closely related Topobea con- stitute the tribe Blakeae, which is characterized by baccate fruits, ovoid to pyramidal seeds, and axillary, 6-merous flowers that are individually subtended by two pairs of decussate bracts in- serted at the base of the hypanthium. These gen- era exhibit many parallel variations in foliar characters, floral bract morphology, and calyx development. The traditional distinction be- tween Blakea and Topobea is based on the an- droecial characters summarized in the following couplet: Anthers short, oval, oblong, or elliptic, blunt or broadly rounded at the summit with two typically well-separated (often minute) api- cal pores - Blakea Anthers linear-oblong to lanceolate or subu- late, the dorsally inchned apical pores ap- proximate and often confluent at anthesis Topobea [269] 270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 In addition to the three species of Blakea de- scribed by Almeda (1974, 1980) and the generic transfer presented below, Blakea tuberculata Donn.-Smith of Costa Rica has recently been collected in western and central Panama (Chi- riqui: Folsom et al. 4838, CAS; Code: Knapp 5334, CAS; Panamd: Sytsma 1154, CAS). These additions increase the number of described Pan- amanian species to thirteen. Blakea elliptica (Gleason) Almeda, comb. nov. Topobea elliptica Gleason, Fhytologia 3:353. 1950. Type. — Panama. Bocas del Toro, Robalo Trail, northern slopes of Cerro Horqueta, elevation 6000-7000 ft [1828-2133 m], 5-7 Aug. 1947, Allen 5001 (holotype MO!; photo CAS!). The type collection of this species consists of a branch with four leaves and four more or less mature berries (one of which is detached). When Gleason described this species from in- complete material he was impressed by its sessile, clasping leaves, which are reminiscent of those of Topobea brenesii Standley, a rare Costa Rican endemic. Two recent flowering collections from near the type locality in western Panama are an exact vegetative match for the species that Gleason placed in Topobea. The uniform, bluntly ovoid anthers of these new collections are characteristic of Blakea, thus necessitating the generic transfer proposed herein. Because all available descrip- tions of this taxon are based on the fragmentary type collection, an amplified description is presented below to facilitate rec- ognition of this little-known cloud forest epiphyte. Epiphytic shrub reportedly 2-4 m tall. Cauline intemodes glabrate, terete, becoming striate to furrowed or cracked in age. Distal branchlets and floral pedicels sparsely covered with a deciduous indument of slender, spreading, glandular tri- chomes and stoutly conic trichomes, the latter typically copious on distal nodes and vegetative buds. Mature leaves sessile, blades chartaceous to coriaceous, entire, somewhat revolute on drying, 5-1 1.5 cm long and 3-7.5 cm wide, el- liptic-ovate to obovate, obtuse to rounded api- cally and auriculate to cordate-clasping basally, 3-nerved with an additional but less conspicuous submarginal pair, essentially glabrous through- out or with a sparse scattering of conic and glan- dular trichomes at the base of the blade below. Flowers 6-merous, solitary or paired in the axils of distal branches, pedicels 2.5-4.3 cm long. Flo- ral bracts foliaceous, sparsely covered with a mixture of blunt conic and spreading glandular trichomes on the inner and outer surfaces; outer bracts 3-5-nerved, essentially free to the base, 12-14 mm long and 13-15 mm wide, broadly elliptic-ovate to suborbicular, obtuse to rounded or somewhat retuse apically; inner bracts (at an- thesis) connate basally for about 4-5 mm to form a collar that tightly envelops the ovary, the free lobes ± semicircular, 7 mm long and 12-13 mm wide basally between sinuses. Hypanthia (at an- thesis) glabrous, campanulate, 8-1 1 mm long to the torus. Calyx lobes triangular, ± entire, most- ly recurved, 5-6 mm long and 5 mm wide ba- sally, apically beset with a mixture of spreading glandular and sessile globular trichomes. Petals thin and ± translucent on drying, mostly gla- brous with a moderate but somewhat concen- trated abaxial patch of brown verrucose excres- cences, reportedly white with pink blotches, elliptic-ovate to obovate and rounded apically, entire to irregularly ciliolate, 20-23 mm long and 14-18 mm wide. Stamens isomorphic, free and declined to one side of the flower, filaments gla- brous, 4.5-6 mm long and 1.5 mm wide; anthers yellow, ovoid, laterally compressed, biporose and rounded apically, 5 mm long and 1 .5-2 mm wide, connective thickened dorsally and dilated basally about 0.5 mm above point of filament insertion into a short deflexed truncate appendage about 0.5-1 mm long. Style straight, glabrous, con- spicuously exserted, 11-14 mm long; stigma truncate. Berry 13-15 mm long to the torus and 14-15 mm in diameter, glabrous and subglobose. Seeds beige, cuneate to clavate, or narrowly pyr- iform, mostly 1 mm long. Additional Specimens Examined. — Panama. Chiriqui: Bo- cas del Toro border along Continental Divide NE of Cerro Pate Macho, above Palo Alto in windswept elfin forest. 8°47'N, 82''2 1 ' W, 2200 m, 1 5 Mar. 1 982, Knapp et al. 4233 (MO); end of road past Palo Alto NE of Boquete in forest along ridge; elev. 6200-6800 ft [ 1 890-2073 m], 8 Feb. 1979, Hammel 6048 (MO). Distribution. — Western Panama. Known only from the northern slopes of Cerro Horqueta and the region NE of Cerro Pate Macho above Palo Alto at elevations of about 1800-2100 m. This species is apparently rare and localized, as evidenced by the few specimens collected over the past three decades. It is among the most dis- tinctive of the Central American species by vir- tue of its sessile, clasping leaves, and deciduous indument of spreading glandular and blunt conic trichomes on distal branchlets, pedicels, floral bracts, and vegetative buds. The petals of this species are remarkably thin and translucent when dry and have a peculiar abaxial cluster of brown veniicose excrescences. Additional collections and field observations will be needed to deter- ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 271 mine whether this petal feature is a consistent diagnostic character or an artifact of pressing and drying. Clidemia D. Don Clidemia is one of approximately 30 genera constituting the tribe Miconieae. The generic classification of this complex tribe is difficult be- cause it contains species groups that intergrade across generic lines that have been defined on the basis of one or few characters. The evolu- tionary development of characters used in ge- neric delimitation of the Miconieae is unclear, and some of the genera as currently delimited are probably polyphyletic. Until a reassessment of generic Hmits can be accomplished, Clidemia is defined in the traditional sense to include those berry-fruited species with axillary inflorescences, more or less obtuse petals, linear to subulate an- thers, wingless hypanthia, and flowers that are not individually subtended by two pairs of de- cussate bracts. Of the approximately 165 species oi Clidemia recorded for tropical America, 30 are now known from Panama. This count includes the two new taxa proposed below. Clidemia tetrapetala Almeda, sp. nov. (Figure 1) Frutex 1-2 m. Ramuli teretes sicut petioli in- Jlorescentiafoliorum subtus venae primariae pilis stipitato-stellatis densiuscule ornati et pilis stel- latis sessilibus modice vel dense induti. Petioli 10-27 mm longi; lamina 5-9.5 x 2.1-3.9 cm ovato-elliptica apice gradatim acuminata basi rotundata vel subcordata, chartacea et Integra, 5-7 -plinervata. Inflorescentiae pauciflorae info- liorum superiorum axillis oppositis, axe plerum- que 1.5-2.5 cm longo; flores 4-meri plerumque subsessiles (pedicellis supra articulationem 0.5 mm longis), bracteolis 1-1.5 mm longis lineari- bus. Hypanthium (ad torum) 1.5 mm longum dense stellato-puberulum et pilis laevibus glan- duliferis patentibus 0.5-1 mm longis modice in- termixtus; lobis 1 mm longis triangular ibus, den- tibus exterioribus subulatis 3 mm eminentibus. Petala 5-6 x 1.5-2 mm oblonga glabra (apice late obtuso). Stamina isomorphica glabra; fila- menta 2 mm longa; antherarum thecae 1.5 x 0.5 mm oblongae, poro dorsaliter inclinato; con- nectivum nee prolongatum nee appendiculatum. Stylus 5.5 mm glaber; stigma truncatum; semina 0.5-1 mm pyriformia. Shrubs 1-2 m tall. Older cauUne intemodes glabrate and terete. Distal branchlets, vegetative buds, juvenile leaves, and inflorescences copi- ously covered with sessile, and stipitate-stellate trichomes, the distal cauline intemodes sparingly intermixed with spreading glandular hairs. Leaves of a pair slightly unequal in size, otherwise iden- tical in shape and vestiture. Leaf blades flat to somewhat bullate above when dry, chartaceous, entire, 5-9.5 cm long and 2. 1-3.9 cm wide, ovate- elliptic, acuminate apicafly and rounded to sub- cordate basally, 5-7-plinerved with a network of secondary and tertiary nerves below, sparsely pu- bescent above with a mixture of appressed sim- ple trichomes and spreading stalked stellate tri- chomes, lower leaf surfaces copiously stellate pubescent on the elevated primary nerves but with a moderate to sparse mixture of sessile and stalked stellate trichomes on and between the secondaries; petioles 10-27 mm long and 1-1.5 mm broad. Inflorescence axillary, typically a lax few-flowered modified dichasium 1.5-2.5 cm long; bracteoles sessile, 1 - 1 . 5 mm long and most- ly less than 0.5 mm wide, linear-lanceolate to linear-oblong, mucronate apically, glabrous above and sparsely stellate pubescent below. Pedicels terete, mostly less than 0.5 mm long and disar- ticulating at the tribracteolate node closely sub- tending each flower. Hypanthia (at anthesis) nar- rowly campanulate, 1.5 mm long to the torus, copiously covered with a mixture of stellate and spreading glandular trichomes. Calyx lobes (on fruiting hypanthia) widely spreading, persistent, triangular, entire, 1 mm long and 1 mm wide basally; exterior calyx teeth subulate, mostly 3 mm long, stellate pubescent and completely ob- scuring external surface of the calyx lobes. Petals 4, glabrous, reportedly white, linear-oblong, broadly obtuse apically, entire, 5-6 mm long, 1.5-2 mm wide. Stamens 8, isomorphic, erect at anthesis; filaments glabrous, ligulate but abruptly constricted distally, 2 mm long; anthers 1.5 mm long and 0.5 mm wide, yellow when dry. linear- oblong, truncate to rounded distally with a cir- cular dorsally inclined apical pore; connective simple, somewhat thickened dorsally but not di- lated or prolonged below the thecae. Ovary to- tally inferior, 4-celled, essentially glabrous at the summit but sparsely setose surrounding the sty- lar scar. Style 5.5 mm long; stigma truncate. Ber- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 7 Figure 1 . Clidemia tetrapetala Almeda. A, habit, x ca. %; B, representative leaf (lower surface), x ca. y^\ C, mature berry, x ca. 4; D, petal, x ca. 6; E, stamens, Va lateral view (left) and dorsal view (right), x9; F. seeds, x ca. 7. (A from Folsom 4871; B-Ffrom Folsom 6106.) ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 273 Figure 2. Clidemia trichosantha Almeda. A. habit, x ca. Vi; B, representative leaf (lower surface), x ca. V*; C, representative flower, buds, bracts and bracteoles, x ca. 4; D. petal, x 7; E, stamens, lateral view (left) and ventral view (right), x 9; F. seeds, X ca. 14. {A-E from the holotype; F from Hammel 965.) ry reportedly opaque green but turning translu- Type. — Panama. Bocas del Toro/Chiriqui Border: Cerro cent red at maturity, globose, 5-6 mm long to Colorado, along mtersection of Bocas Road with main ridge ^, ^ c /- J- ^oj 1 road, 1 1.8 km from Cham i along path headed into Bocas del the torus, 5-6 mm m diameter. Seeds narrowly ^^^;^ ^,^^^^.^^ ^^^^^^^^ ^ \l ^^^ ,,,, ^^^^^^ ,^,, pynform, brown, smooth, vemicose, and mostly (holotype: CAS!; isotype: mo). 0.5—1.0 mm long. Additional Specimens Ex-amined. — Panama. Bocas del 274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 Toro/Chiriqui Border: Cerro Colorado, 11.2 km along ridge road from main road to Escopeta, 1700 m, 16 Aug. 1977, Folsom 4871 (CAS, MO). Chiriqui: Cerro Colorado on Con- tinental Divide, 1400 m, 25 Jul. 1979, Antonio 1416 (CAS). Distribution. — Known only from Cerro Col- orado in western Panama at elevations of 1 400- 1700 m. All cited specimens are in flower; only the type has mature fruits. Among Panamanian species of Clidemia with 4-merous flowers, C. tetrapetala is distinguished by its lance-ovate, 5-7-plinerved leaves, linear- oblong petals, and mixture of spreading glan- dular and stellate hypanthial trichomes. The in- florescence of this species is also notable for its elongate distal peduncles that can readily be mis- taken for floral pedicels. The actual pedicels in this species are very short and disarticulate at tribracteolate nodes that closely subtend the flowers. On the basis of its 4-merous flowers, well de- veloped calyx teeth, and equal to subequal prin- cipal leaves in each pair, this species is provi- sionally placed in section Sagraea as defined by Cogniaux (1891). Among species of this section, C. tetrapetala bears a superficial resemblance to C. saltuensis Wurdack of Venezuela which is re- portedly known only from the type (Wurdack 1973). The latter species diflers most conspicu- ously in having ovate-elUptic, basally obtuse leaves, stellate-pinoid hypanthial trichomes, shorter (0.7 mm), erect calyx teeth, narrowly ob- long anther thecae, and abaxially mucronate pet- als. Clidemia trichosantha Almeda, sp. nov. (Figure 2) Frutex 1-2.5 m. Ramuli teretes primum sicut petioli laminae subtus inflorescentia hypan- thiaque dense setosi pilis l-3(-4) mm longis de- mum glabrati. Folia in quoque pari in dimen- sionibus disparilia (2:1); petioli 0.2-2.2 cm longi; lamina 3-14.3 x 1.5-5.8 cm elliptica apice acu- minata basi interdum obtuse vel rotundata vel paulo asymmetrica, chartacea et denticulata, 5- 7-plinervata, supra sparse vel modice strigosa. Inflorescentiae ca. 1.5-3 cm longae in foliorum superiorum axillis; Jlores 5-meri, pedicellis 1-1.5 mm longis, bracteolis 1.5-3 mm longis lanceo- latis amplexicaulibus. Hypanthium (ad torum) 2-2.5 mm longum; lobis interioribus late deltoi- deis, dentibus exterioribus setosis 1-2 mm emi- nentibus. Petala glabra 4-4.5 x 1.5 mm oblon- go-lanceata. Stamina isomorphica glabra; filamenta 2.5 mm longa; antherarum thecae 1.5 X 0.75 mm oblongae, connectivum nee pro- longatum nee appendiculatum. Stylus 4.5 mm glaber; stigma punctiforme; semina 0.5 mm cu- neata. Slender shrubs 1-2.5 m tall. Intemodes terete, glabrate at maturity, but vegetative buds, young leaves, and distal branchlets moderately to densely covered with smooth, eglandular, spreading trichomes mostly l-3(-4) mm long. Leaves of a pair usually unequal in size, other- wise identical in shape and vestiture. Leaf blades chartaceous, denticulate, elliptic, acuminate api- cally, acute varying to obtuse, rounded or con- spicuously oblique basally, 5-7-plinerved with a network of secondary and tertiary nerves, the innermost pair of primary nerves diverging from the median nerve (0. 6-) 1.0-1. 6 cm above the petiole/laminar junction, moderately to sparsely strigose or hirtellous above and moderately to copiously hirsute below. Larger leaf at each node 5.8-14.3 cm long and 2.5-5.8 cm wide with pet- ioles mostly 0.5-2.2 cm long. Smaller leaf 3-9.5 cm long and 1.5-4. 1 cm wide with petioles most- ly 2-4 mm long. Inflorescence an axiflary, mul- tiflowered, modified dichasium with flowers borne in pedunculate terminal glomerules; rachis terete, 1.5-3 cm long, moderately to sparsely hir- sute; bracts and bracteoles sessile, paired and often fused basally into an amplexicaul nodal collar, 1.5-3 mm long and 0.5-1.5 mm wide, lanceolate to naviculiform, glabrous but irregu- larly fringed with spreading setose trichomes. Pedicels hirsute, 1-1.5 mm long. Hypanthia (at anthesis) ± ovoid, 2-2.5 mm long to the torus, copiously covered with smooth spreading tri- chomes. Calyx lobes (on fruiting hypanthia) in- conspicuous, erect, persistent, broadly deltoid, entire, 1 mm long and 1.5 mm wide basally be- tween sinuses; exterior calyx teeth setiform, 1-2 mm long, sparsely hirsute. Petals 5, glabrous, reportedly translucent white, elliptic-lanceolate, acute apically, entire, 4-4.5 mm long, 1.5 mm wide. Stamens 10, isomorphic, apparently erect at anthesis; filaments glabrous, ± linear-oblong but constricted distally, 2.5 mm long; anthers 1 .5 mm long and 0.75 mm wide, linear-oblong, trun- cate distally; connective simple, not dilated or prolonged below the thecae. Ovary ca. % inferior, 5-celled. Style 4.5 mm long; stigma punctiform. Berry purple at maturity, globose, 4-6 mm long ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 275 and 4-5.5 mm in diameter. Seeds cuneate, ± smooth with vemiculose angles and a promi- nent ± translucent lateral raphe, 0.5 mm long. Type. — Panama. Code: sawmill above El Cope, in forest along stream east of sawmill on the Atlantic drainage, elevation 2300 ft (701 m), 27 July 1978, //aww£'/^;ii(holotype, CAS!; isotype, MO). Additional Specimens Examined. — Panama. Code: above El Petroso sawmill at Continental Divide, N of El Cope, 1 3 May 1981, Sytsma & Andersson 4624 (CAS); area surrounding Rivera sawmill, Alto Calvario, 7 km N of El Cope at Conti- nental Divide, 25 Nov. 1977, Folsom & Collins 6473 (CAS); near Continental Divide along lumbering road 8.4 km above El Cope, 19 Jan. 1978, Hammel 965 (CAS); 7 km N of El Cope around Rivera sawmill, 21 Dec. 1911, Folsom el al. 7093 (CAS); El Cope on Pacific side about V2 hour walk from the sawmill, 16 Oct. 1979, Antonio 2116 (CAS). Panam^: forested slope along El Llano-Carti road, 12 km from Panamerican Highway, 10 Sept. 1976, D'Arcy 10617 (CAS). San Bias: El Llano-Carti road, 12 mi from Panamerican Highway, 10 May 1981, Sytsma & Andersson 4493 (CAS). Veraguas: Cerro Tute, 30 Nov. 1979, Antonio 2928 (CAS). Distribution. — Presently known from wet forests and stream margins at elevations from 700-1300 meters in central Panama. Flowering specimens have been gathered in January, May, July, and September. The epithet for this species draws attention to its most remarkable feature— the dense covering of spreading, lustrous trichomes on the flowers and peduncles of the inflorescence. The tri- chomes, which are often basally flattened when dry, are so copious that it is difficult to examine the structure and posture of bracteoles and other floral parts without a dissection of hydrated ma- terial. Other characters that readily separate C. trichosantha from its congeners include the mod- ified dichasial inflorescence of pedunculate ter- minal glomerules, the distally constricted fila- ments, the cuneate seeds with vemiculose angles, and the sessile, paired bracts and bracteoles that are often fused into amplexicaul collars. A fruiting collection from Cerro Habu, San Bias {Sytsma et al. 2673, CAS) appears to be a variant or closely related, undescribed taxon but its essentially glabrous leaves, glabrate berries, and short (0.5 mm) calyx teeth fall outside the range of variation here attributed to C tricho- santha. The 5-merous, short-pedicellate flowers and anisophylly of the new species dictate placement into Cogniaux's section Calophysoides. Clidemia trichosantha is unlike any of the species currently placed in that section. I am, therefore, reluctant to include it with an assemblage that may prove to be artificial and can only suggest that no close relatives are apparent. The combination of smooth, spreading trichomes and elongate calyx teeth is vaguely suggestive of the copiously hir- sute variants of C. petiolaris (Schlecht. & Cham.) Schlecht. ex Triana which, otherwise, differ markedly in their lax, divaricately branched in- florescence, subulate, apically notched anthers, and pyriform, papillate seeds. GraflFenrieda DC. Graffenrieda, a genus of about 40 species, is one of five closely related merianioid genera oc- curring in Panama. Gleason (1958) reported one species (under the generic synonym Calyptrella Naud.) in his treatment of Panamanian Melas- tomataceae. The escalated botanical exploration of Panama in the past two decades has increased this total to about eight. In addition to the species proposed below, G. galeottii (Naud.) L. Wms., G. gracilis (Triana) L. Wms., and G. micrantha (Gleason) L. Wms. are known from Panama. Collections of about four additional entities probably represent undescribed taxa or range ex- tensions of South American species. These re- main unnamed because of incomplete material. In the field, the superficial resemblance of Pan- amanian species of Graffenrieda to Miconia is striking. Graffenrieda is readily distinguished by the combination of capsular fruits, calyptrate ca- lyx, dorsally spurred, arcuate anthers, and linear, pyramidate seeds. Graffenrieda bella Almeda, sp. nov. (Figure 3) Arbor parva 5 m. Ramuli glabri et teretes. Pe- tioli 0.5-1 .6 cm; lamina 4.7-8 x 2.1-4 cm ellip- tica apice breviter subabrupteque acuminata basi acuta vel cuneata vel biauriculata, supra glabra vel sparse lepidota, subtus modice lepidota, 3(-5)-nervata, coriacea et Integra. Panicula 5-7 cm longa multiflora; flores 5(-6)-meri subsessiles (vel pedicellis 0.5-1 mm longis) ad ramulorum apices glomerati (floribus ca. 3-6 in quoque glo- merulo), bracteolis 0.5-1 mm longis subulato- setosis evidenter mox caducis. Hypanthium (ad torum) 3 mm longum; calyx in alabastris ma- turis clausus 3-3.5 mm longus ad anthesim ir- regulariter supra torum dehiscens. Petala 7-9 x 4-6 mm elliptico-ovata glabra. Stamina isomor- phica glabra; filamenta 2.5-3 mm longa; an- 276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 Figure 3. Grajfenrieda bella Almeda. A, habit, ><%; B, representative leaves, lower surface (left) and upper surface (right), X ca. %; C, mature hypanthia, x ca. 4; D, calyptrate calyx, x ca. 8; E, petal, x ca. 5; F, stamen (lateral view), x ca. 10; G, seeds, x ca. 14. {A-B, D-F from the holotype; C& G from Mori el al. 7581.) therarum thecae 3-4.5 x 0.5 mm oblongo- subulatae; connect ivum vix (0.25 mm) prolon- gatum, dente dorsali 0.5 mm longo acuta. Stylus 7.5-9.5 mm glaber; stigma punct (forme; semina numerosa 1 mm longa, recta paulo angulata. Trees to 5 m tall. Cauline intemodes glabrous and terete; the nodes on defoliated branchlets somewhat swollen with age. Leaf blades coria- ceous, entire, 4.7-8 cm long and 2. 1-4 cm wide, broadly elliptic, apically acuminate basally acute to cuneate or biauriculate with well-developed reflexed marginal lobes ca. 2 mm above the pet- iole-laminar junction, glabrous to sparsely lepi- dote above and moderately lepidote below at ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 277 maturity, uniformly 3-nerved (excluding the in- conspicuous submarginal pair); petioles 0.5-1.6 cm long and 1-1.5 mm broad. Inflorescence a multiflowered panicle mostly 5-7 cm long (in- cluding the peduncle), the uppermost units com- monly glomerulate; bracteoles sessile, narrowly subulate to acicular, 0.5-1 mm long, apparently early deciduous and mostly absent at anthesis. Pedicels 0.5-1 mm long. Hypanthia (at anthesis) narrowly campanulate to subcylindric, 3 mm long to the torus. Calyx acute, apiculate and closed in bud, 3-3.5 mm long rupturing near the torus and falling away as a unit at anthesis. Calyx teeth (on mature hypanthia) evident as blunt tuberculi- form projections mostly 0.25-0.5 mm long. Pet- als mostly 5, but varying to 6 in some flowers, glabrous, white, elliptic to ovate, entire, acute apically, cuneate to clawed basally, 7-9 mm long and 4-6 mm wide. Stamens mostly 10 but some- times 12, isomorphic; filaments glabrous, ligu- late, 2.5-3 mm long and 0.5 mm wide; anthers 3-4.5 mm long and 0.5 mm wide, yellow, sub- ulate, the apical pore ± truncate; connective pro- longed ca. 0.25 mm below the thecae and mod- ified dorsally (at filament insertion) into an acute spur mostly 0.5 mm long. Ovary 'A inferior, 3-celled, ovoid, glabrate but copiously lepidote distally. Style straight to somewhat curved, glabrous, 7.5-9.5 mm long; stigma punctiform. Hypanthia (at maturity) prominently costate, 5 mm long and 3 mm wide. Seeds linear-pyram- idate, brown, mostly 1 mm long. Type. — Panama. Chiriqui: along road between Gualaca and the Fortuna Dam site at 10. 1 m; NW of Los Planes de Homito, elev. 1300 m, 8 Apr. 1980, Antonio 4078 (holotype: CAS!; isotype: MO). Additional Specimens Examined. — Panama. Chiriqui: Cerro Homitos, 40 km NW of Gualaca in cloud forest dom- inated by Quercus spp., Podocarpus, and Drimys. 26 Jul. 1975, Mori & Bollen 7486 (CAS). Veraguas: Cerro Tute, ca. 10 km NW of Santa Fe on ridgetop in cloud forest, 19 May 1975, Mori 6255 (CAS), 3 Aug. 1975, Mori et al. 7581 (CAS). Distribution.— Known only from cloud for- ests on Cerro Tute in Veraguas province and Cerro Homitos and vicinity in Chiriqui province at elevations from 1000-2238 m. Flowering specimens have been collected in April, May, and July. This species is reportedly (fide Mori 6255) one of the most common trees in an area about 10 km northwest of Santa Fe on Cerro Tute. Graffenrieda bella is most closely related to G. micrantha (Gleason) L. Wms. which differs in its longer leaf blades (1 3-20 cm) with five prom- inently elevated abaxial nerves, longer petioles (2.5-5 mm), shorter (2-3 mm) narrowly lanceo- late petals, and smaller fruiting hypanthia (2-2.5 mm). My circumscription of G. bella includes two morphological entities representing geo- graphically distinct populations. The collections from Chiriqui in western Panama have 5-merous flowers and foliar margins with bilaterally de- veloped reflexed auricles about 2 mm above the petiole/laminar junction. Collections from Vera- guas in central Panama have 5(-6)-merous flow- ers and lack the conspicuous foliar auricles. Ex- cept for these differences, the few available specimens from each area are identical and ex- hibit parallel ranges in foliar size, hypanthial and staminal length, and petal width. More collec- tions, especially from intervening areas, are needed to determine whether this bimodal vari- ation shows a consistent geographical correla- tion. Miconia Ruiz & Pavon As presently circumscribed, Miconia with ap- proximately 1 000 species is the largest genus in the family and one of the largest among flowering plants. It is generally characterized by its ter- minal inflorescences and apically rounded, ob- tuse, or retuse petals. Miconia occupies a central position in the complex of genera comprising the taxonomically difficult tribe Miconieae. Through its broad neotropical range Miconia displays great diversity in habit, foliar characters, infforescence organization, trichome and androecial mor- phology, and seed structure. Success in identi- fying species, however, inevitably requires ex- amination of anther structure because the current sectional classification reUes heavily on this char- acter. Miconia is greatly in need of monographic study to evaluate character variation, assess the status of satellite groups, and redefine sectional limits based on character correlations. Some 60 species of Miconia occur in Panama. These rep- resent all twelve currently recognized sections. In addition to the species proposed below, sev- eral new taxa await formal description pending collection of complete material. Miconia arboricola Almeda, sp. nov. (Figure 4) Caulis scandens primum sicut foliorum subtus venae primariae inflorescentia hypanthiaque dense vel modice pilis stipitato-stellatis 0.5-1 mm PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol, 43, No, 1 7 (^&- Figure 4. Miconia arboricola Almeda. A, habit, x ca. 'A; B, representative leaf (lower surface) with enlargement (left), x ca. V2; C, stipitate-stellate trichome, x ca. 17; D, mature berry, x ca. 5; E, representative flower showing young hypanthium, two petals, style, bracteoles, and pedicellar scars, x ca. 7; F, stamens, lateral view (left) and dorsal view (right), x ca. 9; G, petal, X ca. 7; H, seeds, x ca. 20. {A-C from the holotype; D, F, G from Knapp 5735\ E &H from Croat 49151.) latis indutus. Petioli 8-23 mm longi; lamina 5.5- 11.4 y^ 3.5-7.1 cm late ovata vel cordata, apice breviter acuminata basi subcordata vel cordata, 5-7-nervata, fir me coriacea et Integra. Panicula 11.5-25 cm longa multiflora; flores 5-meri sub- sessiles in glomerulis interruptis vel ad ramulo- rum apices aggregati, bracteolis 1-2.5 x 0.25- 0.5 mm anguste oblongis persistentibus. Hypan- thium (ad torum) 1.5-2 mm longum; calycis tu- bus non eminentibus, lobis interioribus 0.5 mm longis deltoideis, dentibus exterioribus crassis lo- bos interiores aequantibus. Petala 2-3 x 1.5-2 mm glabra obovata apice retuso. Stamina isomorphica glabra; filamenta 2.5 mm longa; ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA antherarum thecae 1.5 x 0.5 mm oblongo- cuneatae; connectivum nee pwlongatum nee ap- pendiculatum. Stylus 4 mm glaber; stigma eap- itatum; semina 0.5 mm galeiformia. Pendent woody vines or hemiepiphytes ad- hering to bark of host tree by adventitious roots. Older cauline intemodes terete and ± glabrate. Distal branchlets, petioles, vegetative buds, ju- venile leaves, and inflorescences moderately to copiously covered with ferrugineous stipitate- stellate trichomes. Leaves of a pair essentially equal in size; blades coriaceous at maturity, en- tire, 5.5-11.4 cm long and 3.5-7.1 cm wide, broadly ovate to cordate, short acuminate api- cally and subcordate to cordate basally, 5-7- nerved with a prominulous reticulate network of secondaries and tertiaries below, mostly glabrate and vemicose above at maturity or with stipi- tate-stellate trichomes persisting to varying de- grees in the furrows created by the impressed primary nerves, sparsely to moderately stipitate- stellate below; petioles 8-23 mm long and 2-2.5 mm wide. Inflorescence basically paniculiform with ultimate branchlets terminating in multi- flowered congested glomerules; bracts of the rachis nodes paired, linear-oblong, 2.5-5(-10) mm long, 1-1.5 mm wide, glabrous adaxially and stipitate-stellate to glabrate abaxially; bracteoles 3-5 per pedicel, sessile, linear-oblong, acute to obtuse or roimded apically, 1-2.5 mm long, 0.25- 0.5 mm wide, entire, glabrous adaxially and stip- itate-stellate abaxially. Pedicels 0.5 mm long but typically inconspicuous and concealed by con- gested glomerules. Hypanthia (at anthesis) sub- cylindric to narrowly campanulate, 1.5-2 mm long to the torus, moderately to copiously beset with short stalked-stellate trichomes and occa- sionally with a sparse scattering of spreading glandular trichomes on or near basal portions of the calyx teeth; adaxial rim of the torus minutely glandular-puberulent. Calyx lobes (on young fruiting hypanthia), glabrous, hyaline, and erect, deltoid, 0.5 mm long and 1 mm wide basally; calyx teeth bluntly subulate, copiously stellate, adnate to and ± equaling subtending calyx lobes. Petals 5, glabrous, white, obovate to oblong-ob- ovate, strongly reflexed, irregularly retuse api- cally, 2-3 mm long and 1.5-2 mm wide. Stamens 10, isomorphic, filaments straight to somewhat incurved distally, glabrous, subulate, 2.5 mm long; anthers 1.5 mm long and 0.5 mm wide distally, linear-oblong to narrowly cuneate, typ- ically recurved distally; connective thickened dorsally but not dilated or prolonged below the point of filament insertion. Ovary (young fruiting hypanthia) ca. 'A to 'A inferior, 3-celled, coarsely papillate to farinaceous, caducously glandular- puberulent along the bluntly fluted stylar collar that becomes depressed and inconspicuous on mature fruits. Style straight, glabrous, 4 mm long; stigma broadly capitate. Berry reportedly blue at maturity, globose, 3^ mm long and 3^ mm in diameter. Seeds galeiform, white, papillate to costate-papillate, 0.5 mm long. Type. — Panama. Chiriqui: slope NW of confluence of Rio Homito and Rio Chiriqui, ca. 8°44'N, 80°07'W, elev. 1050- 1100 m, 11 Nov. 1980, Stevens 18266 (holotype: CAS!; iso- type: MO). Additional Specimens Examined.— Costa Rica. Cartago: Pejibaye, 30 May 1924, Lankester 877 (US). Limon: Cerro Uren, Cordillera Talamanca, 1 Sept. 1898, Pittier 12683 (US). Panama. Code: hills N of EI Valle, E slope and ridges leading to Cerro Gaital, 8°40'N, 80°07'W, 27 June 1982, Knapp 5735 (MO); Alto Calvario, above sawmill on Continental Divide, 5.2 mi above EI Cope, 6 Dec. 1979, Croat 49151 (CAS). Distribution. — This little-collected cloud forest species ranges from the Caribbean slopes of the Cordillera Talamanca in south-central Costa Rica south and east to the hills surround- ing El Valle de Anton in central Panama at el- evations from 300-1100 m. Flowering speci- mens have been collected in June, September, and November. Labels on collections of this species describe it as a pendent woody vine. The notes on Croat 49151 describe it as a hemiepiphyte with the stem rooted in the ground but closely attached to a tree trunk. It seems likely that plants of this species are typically hemiepiphytes that ulti- mately become scandent vines requiring the sup- port of host trees. The specific epithet refers to this discordant habit'among species of Miconia. Mieonia arborieola is quite unlike all other species o{ Mieonia from Central America. In ad- dition to the unusual habit, it is distinctive in its coriaceous leaves, stipitate-stellate pubescence, paniculiform inflorescence, congested, polybrac- teolate flowers, and sculptured, galeiform seeds. As pointed out on the label of Pittier 12683, in Cogniaux's monograph, this species comes clos- est to M. mollieula Triana and M. cordata Triana in section Mieonia. In foliar shape, intemodal pubescence, and inflorescence structure, M. ar- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 Figure 5. Topobea calophylla Almeda. A, habit, x'/j; B. stamens, ventral view, xl; C, hypanthium with floral bracts removed, x ca. 1 Vr, D, cauline trichomes, x ca. 4; E, petal, x ca. 1 1/4; F, outer floral bract (outer surface), x ca. 1 ; G, inner floral bract (inner surface), x ca. 1 . (/I & Z» from Nee 987 3\ B. C & E from Mori & Kallunki 5353; F-G from the holotype.) ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA boricola resembles these two shrubby species. They differ most notably from the new species in having irregularly denticulate leaves, sessile stellate or stellate- furfuraceous hypanthial pu- bescence, and a differentiated androecium in which the larger stamens are inserted opposite the petals. Topobea Aublet The species proposed below represents the first new Topobea to be described from Panama since the appearance of Gleason's (1958) summary. The following description of T. calophylla, the transfer of T. elliptica to Blakea (also proposed herein) and the discovery of T. pittieri Cogn. in Chiriqui province (Cerro Colorado, Folsom & Collins 1770, CAS) now brings the number of Panamanian species to nine. Topobea calophylla Almeda, sp. nov. (Figure 5) Frutex epiphyticus. Ramuli primum obscure quadrangulati demum teretes sicut petioli lami- narum subtus venae primariae pedicelli bracte- aeque pilis barbellatis ca. (l-)3-9 mm longis in- duti. Petioli 2.2-5 cm longi; lamina J 4.5-37.5 x 8.6-17.8 cm, elliptico-ovata, apice abrupt e cau- dato-acuminato basi rotundata vel paulo corda- ta, supra ad maturitatem glabra, subtus modice setosa (pilis ca. 1-3 mm longis). Flores 6-meri infoliorum superiorem axillis oppositis plerum- que 2-4; bracteae liber ae; bract eae exter lores 1.7- 2.3 X 1.5-1.7 cm late ovatae; bracteae inter lores 1.6-1.9 X 1.3-1.7 cm elliptico-ovatae. Hypan- thium (ad torum) 1.1-1.3 cm longum extus dense strigosum pilis 2-4 mm longis, calycis lobis 14- 17 mm longis triangularibus. Petala glabra 2 x 1 cm spathulata apice hebeti-acuto vel obtuso. Filamenta 5 mm longa; antherae 8 x 1.5 mm subulatae inter se cohaerentes, ports binis ter- minalibus; connectivum ad basim dorsaliter mi- nute calcaratum. Stigma capitellatum vel trun- catum; stylus 11-14 mm glaber in ovarii collo 5- 6 mm immersus. Coarse spreading epiphytic shrub. Cauline in- temodes ± terete to subquadrangular. Distal branchlets, vegetative buds, pedicels, and floral bracts covered with a hirsute indument of rusty brown barbellate trichomes mostly 3-9 mm long. Mature leaves firmly chartaceous to coriaceous, inconspicuously dentate, 14.5-37.5 x 8.6-17.8 cm, elliptic-ovate, abruptly caudate-acuminate apically, rounded to subcordate basally, 5-7- nerved with conspicuous elevated secondaries, glabrous above, moderately hirsute below with barbellate trichomes mostly 1-3 mm long; pet- ioles 2.2-5 cm long and 3-5 mm diam. Flowers 6-merous, erect to widely spreading, paired or borne in clusters of three or four in axils of the distal branches; pedicels 2.8-4 cm long. Floral bracts foliaceous, entire, 3-5 -nerved free basally, each pair closely subtending one another or sep- arated on the pedicel by a distance of 3-4 mm, outer bracts 1 .7-2.3 x 1.5-1.7 cm, broadly ovate, acuminate apically; inner bracts 1.6-1.9 x 1.3- 1.7 cm, elliptic-ovate, acute to acuminate api- cally. Hypanthia (at anthesis) campanulate, 1.1- 1.3 cm long to the torus, copiously appressed- strigose with barbellate trichomes 2-4 mm long. Calyx lobes lance-triangular, often involute to uncinate apically when dry, entire, hirsute to hir- tellous, 14-17 mm long and 5-6 mm wide ba- sally between sinuses. Petals glabrous, reportedly pink, ± spathulate but acute to obtuse apically, entire to sparingly glandular-ciliate, 2 cm long and 1 cm broad distally. Stamens isomorphic and strongly declined to one side of the flower, filaments glabrous, connivent, 5 x 1.5 mm; an- thers linear-subulate, 8 x 1.5 mm, laterally co- herent but free distally, strongly incurved at an- thesis, each anther tipped with two somewhat dorsally inclined confluent pores; connective thickened dorsally near the point of filament in- sertion into a blunt callosity. Style ± straight, glabrous, 1 1 - 1 4 mm long and 1 mm wide; stigma capitellate to truncate, the surface appearing somewhat crateriform. Ovary inferior, 6-celled, distended apically into a glandular puberulent fluted cone and stylar collar mostly 7-9 mm long. Mature berry not seen. Type. — Panama. Veraguas: 5 mi W of Santa Fe on road past Escuela Agricola Alto Piedra on Pacific side of divide, elev. 800-1200 m, 18 Mar. 1973, Croat 23000 (holotype: CAS!; isotype: MO). Additional Specimens Examined. — Panama. Veraguas: 6- 7 km W of Santa Fe on new road past agriculture school, 18 Feb. 1974, Nee 9873 (CAS); NW of Santa Fe, 4.2 km from Escuela Agricola Alto de Piedra, 25 Feb. 1975, Mori & Kal- lunki 4833 (CAS); NW of Santa Fe, 2.7 km from Escuela Agricola Alto de Piedra along stream on road to Calovebora, 30 Mar. 1975, Mori & Kallunki 5353 (CAS). Distribution. — Known only from montane forests NW of Santa Fe in Veraguas province at elevations of 800-1200 m. Available material, 282 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 all of which is in flower, was collected in February and March. Like so many epiphytic species in the Blakeae that grow in wet forests, this species has a re- stricted distribution and does not appear to be closely related to any described taxon in Topo- bea. As emphasized by the specific epithet, T. calophylla has large, handsome leaves that are glabrous above and moderately hirsute below. It is also striking because of the copious indument of brown barbellate trichomes that gives distal branches, floral bracts, and hypanthia a coarse bristly appearance. Among the species of Topobea recorded for Panama by Gleason (1958), this species also dif- fers in the following combination of characters: its leaves are inconspicuously dentate; the inner and outer floral bracts are separate to the base, copiously pubescent abaxially, and essentially equal in length; and the ovary is elaborated api- cally into a prominent glandular-puberulent, fluted cone crowned by several setiform lobules that envelop the style basally for 5-6 mm. No information is available on the size of in- dividual plants in this species. Judging from fo- liar size and the coarseness of its branches, I suspect that T. calophylla can become an epi- phyte of massive proportions comparable to To- pobea dumndiana Cogn. and Blakea tuberculata Donn. -Smith, both of which can obscure and overtake the crowns of their host trees. Casual collectors unfamiliar with the propensity for epi- phytism in Topobea and Blakea frequently de- scribe the epiphytic species in these genera as trees. Labels for all except one of the few avail- able collections of T. calophylla indicate that it is an epiphyte. Although Nee 9873 is described as a small tree, additional field study is needed to confirm this observation. Acknowledgments I am grateful to Terry Bell for preparing the line drawings, to Colleen Sudekum for technical assistance, and to J. J. Wurdack for critical re- view of the manuscript. I also thank curators of the following herbaria who provided special loans and/or made their facilities available for study: BM, CR, DUKE, F, MO, US. Literature Cited Almeda, F. 1974. A new epiphytic Blakea (Melastomata- ceae) from Panama. Brittonia 26:393-97. . 1 980. Central American novelties in the genus Blak- ea (Melastomataceae). Rhodora 82:609-15. CoGNiAux, C. A. 1891. Melastomaceae. Pages 1-1256. In A. and C. de Candolle, eds., Monographiae phanerogama^ rum, vol. 7. G. Masson, Paris. Gleason, H. A. 1958. Melastomataceae. In R. E. Woodson, Jr. and R. W. Schery, Flora of Panama. Ann. Missouri Bot. Gard. 45:203-304. Wurdack, J. J. 1973. Melastomataceae. In T. Lasser, ed.. Flora de Venezuela 8:1-819. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES ^ Vol. 43, No. 18, pp. 283-300, 10 figs., 1 table. .COOS Hole, K'aSOecember 11, 1984 MARINE AND FRESHWATER STINGRAYS (DASYATIDAE) OF WEST AFRICA, WITH DESCRIPTION OF A NEW SPECIES By Leonard J. V. Compagno Tiburon Center for Environmental Studies, San Francisco State University, P.O. Box 855, Tiburon, California 94920 and Tyson R. Roberts California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Abstract: Three genera and 12 species of stingrays, all members of the family Dasyatidae, are known from West Africa. A key is provided for their identification. The two most common species in shallow coastal waters (including lagoons, estuaries, and river mouths) are the closely related Dasyatis margarita (Giinther, 1870) and Dasyatis margaritella new species, which differ in disc shape, meristic features, and adult size. Two species occur mainly or exclusively in fresh water: Dasyatis garouaensis (Stauch and Blanc, 1962) and Dasyatis ukpam (Smith, 1863). Dasyatis garouaensis, a small, thin-bodied species previously known only from the lower Niger and a tributary, the Benue, is reported from Lagos and the Cross River. It is closely related to D. margarita and D. margaritella. Dasyatis ukpam, a large thick-bodied species with a vestigial sting previously known only from fetal specimens obtained at Old Calabar (without precise information on habitat), is now reported from the Ogooue and the lower Zaire or Congo rivers. It is perhaps related to the genus Urogymnus. Introduction This study was undertaken to clarify the sys- tematics, distribution, and relationships of West African freshwater stingrays. Although widely distributed and familiar to local fishermen, sting- rays inhabiting the larger rivers of West Africa are poorly known scientifically. There are at least two species. One, Dasyatis ukpam, was de- scribed more than a century ago, but the two fetal type-specimens obtained at Old Calabar lacked precise habitat data, and the species was not reported again or recognized as valid until the junior author obtained specimens in the Ogooue basin in 1978. We have also identified a specimen of this species collected in the lower Zaire (Congo) River in 1937. The other West African freshwater dasyatid, D. garouaensis, was described originally as a species of Potamotrygon, a genus of the otherwise exclusively Neotropical freshwater family Po- tamotrygonidae. Evidence that it is actually a member of the family Dasyatidae was advanced by Thorson and Watson (1975). Our own ob- servations fully support this conclusion. Reid and Sydenham (1979) suggested that D. garouaensis may be identical with the coastal species D. mar- [283] 284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. li garita. Our studies, however, indicate that it is a vahd species, albeit a close relative of D. mar- garita and the previously undescribed D. mar- gahtella. Dasyatis gawuaensis was known only from the Benue and lower Niger, but the junior author collected a specimen in the Cross River, Cameroun, in 1980, and we have also identified a specimen from Lagos, Nigeria (habitat un- known). So far as we have been able to deter- mine, D. margarita and D. margaritella occur only in marine and estuarine habitats. The key below should permit identification of all stingrays now known from West Africa. Fol- lowing the key we present a definition of the genus Dasyatis and detailed descriptions of D. margarita, D. margaritella, D. garouaensis, and D. ukpam. Materials and Methods This paper is based on material deposited in the American Museum of Natural History (AMNH); British Museum (Natural History) (BMNH); California Academy of Sciences (CAS and CAS-SU); Institut fur Seefischerei, Hamburg (ISH); Museum National d'Histoire Naturelle, Paris (MNHN); Musee Royale de I'Afrique Cen- trale, Tervuren, Belgium (MRAC); and Smith- sonian Institution (USNM). Disc width (measured at widest point) is our standard measure of size, and proportional mea- surements (unless otherwise indicated) are ex- pressed as percent of disc width. Definitions or explanations of some other terms are as follows: Disc length — midline measurement from snout-tip to a transverse line parallel to poste- riormost extension of pectoral fins Disc depth— greatest height or depth of disc (usually at scapulocoracoid) Preorbital length— midline measurement from snout-tip to a transverse line parallel to anterior margin of eyes Prenarial length— midline measurement from snout-tip to a transverse line parallel to anterior border of nostrils Prebranchial length — midline measurement from snout-tip to a transverse line parallel to opening of first gill slits Head length — midline measurement from snout-tip to a transverse line parallel to opening of fifth gill slits Pectoral fin inner margin— from posterior in- sertion to posteriormost extension of pectoral fin Pelvic fin span— distance between apices of pelvic fins when maximally extended Upper and lower tooth rows — maximum number of tooth rows across upper and lower jaws Vertebral counts in stingrays are complicated by the extraordinary specialization of the ver- tebral column as a support for the pectoral fins, and by its termination in an elongate tail, which is frequently damaged. Anteriorly the column is fused into two synarcuals incorporating a vari- able number of centra. In Dasyatidae the ante- riormost 23-40 vertebrae are incorporated into the first synarcual. In most of these vertebrae the centra are completely fused, but their number can be determined by counting the spinal nerve foramina. The second synarcual frequently is separated from the first by a small number of intersynarcual vertebrae; in most of the Dasyatis herein reported, however, there is only a joint between the two synarcuals. In the second syn- arcual the centra, although fused, retain their form and are readily counted in radiographs. Some- times the posteriormost centrum in the second synarcual is sharply distinguished from the monospondylous trunk centra succeeding it. In specimens in which the end of the second syn- arcual cannot be determined, we give a combined count of second synarcual plus monospondylous trunk vertebrae. This is usual in late fetal or new- bom specimens with poor calcification and in heavily denticulated specimens in which this portion of the vertebral column is obscured in radiographs (e.g., in D. ukpam). Posteriorly the vertebral column ends in a long series of diplos- pondylous tail centra followed by a highly flex- ible, unsegmented rod (apparently consisting of the notochord and a heavily calcified notochor- dal sheath). The monospondylous and diplos- pondylous sections of the vertebral column are usually sharply demarcated in radiographs. For terminology and illustrations of dasyatid clasper morphology see Compagno and Roberts (1982). Family Dasyatidae Jordan, 1888 We follow Bigelow and Schroeder (1953) in restricting Dasyatidae to the whiptailed sting- rays, and tentatively recognize the following gen- era: Dasyatis, Himantura. Hypolophus, Taeni- ura, Urogymnus, and Urolophoides (see also Compagno and Roberts 1982). COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 285 Figure 1. Disc shape in West African Dasyatidae. (a) Dasyatis violacea (trapezoidal); (b) Dasyatis centroura (diamond- shaped); (c) Taeniura grabata (circular); (d) Urogymnus asperrimus (oval). lb. 2a Key TO Dasyatidae of West Africa Disc oval (Fig. Id); tail without dermal folds; sting invariably absent Urogymnus africanus (Bloch and Schneider, 1801) Disc variable in shape, tail with dermal fold or folds, sting usually present (ab- sent in some Dasyatis ukpam) 2 Ventral tail fold extending to end of tail; disc circular (Fig. 1 c) Taeniura grabata (E. Geoffroy Saint-Hilaire, 1817) 2b. Ventral tail fold ending far anterior to end of tail (Dasyatis) 3 3a. Disc trapezoidal or diamond-shaped (Fig. la-b) 4 3b. Disc oval or circular 9 4a. Disc trapezoidal, anterior margin broad- ly rounded, snout not projecting as an angular lobe from disc (Fig. la); upper and lower surfaces of disc dark D. violacea (Bonaparte, 1832) 4b. Disc diamond-shaped, anterior margin angular, snout projecting as an angular lobe from disc (Fig. lb); lower surface of disc light 5 5a. 5b. Upper surface of disc with a dark retic- ular pattern; ventral tailfold very short, about twice length of sting D. marmorata (Steindachner, 1892) Upper surface of disc plain; ventral tail- fold long, much more than twice sting length 6 6a. Entire dorsal surface of disc covered with small denticles; no middorsal row of en- larged denticles or thorns; adults with over 100 rows of teeth in each jaw; disc very broad, about 1.5 times as wide as long in adults D. rudis (GUnther, 1870) 6b. Dorsal surface of disc only partially cov- ered with small denticles, along middle of back, or naked except for a middorsal row of enlarged denticles or thorns; adults with much less than 100 rows of teeth in each jaw; disc narrower, 1.0-1.3 times as wide as long 7 7a. Anterior margin of disc anterior to spi- racles nearly straight behind snout-tip, with tip projecting; posterior parts of pel- vic fins projecting well rearward beyond rear tip of pectoral fins D. pastinaca (Linnaeus, 1758) 7b. Anterior margin of disc anterior to spi- racles slightly concave behind snout-tip, with tip not conspicuously projecting; posterior parts of pelvic fins extending slightly behind rear tips of pectoral fins 8 8a. Ventral tailfold high, about as deep as tail above it; a dorsal ridge present on tail behind sting; disc and tail in large juveniles and adults without enlarged, heavy, broad-based denticles, but with moderately enlarged middorsal and scapular denticles only D. ameri- cana (Hildebrand and Schroeder, 1928) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. li 8b. Ventral tailfold low, half as deep as tail above it; no dorsal ridge on tail behind sting; disc and tail in large juveniles and adults with scattered enlarged, heavy, broad-based denticles, forming a mid- dorsal row at center of disc and tail D. centwura (Mitchill, 1815) 9a. Anterior margins of disc broadly convex anterior to spiracles, with tip of snout not projecting from them; dorsal disc surface in juveniles to subadults com- pletely covered with denticles, with flat- tened large denticles on midbelt, small pointed denticles laterally, and large, conical, erect, sharp-cusped denticles on thorns scattered on disc and tail base; sting reduced or absent; no dorsal keel on tail; base of tail circular in cross sec- tion; ventral surface of disc light with a broad dusky marginal band D. ukpam (Smith, 1863) 9b. Anterior margins of disc concave ante- rior to spiracles, with tip of snout con- spicuously projecting from them; dorsal disc surface partially naked, with a mid- belt of flattened denticles and often a midscapular pearl spine, or naked; no large conical thorns on disc and tail base; sting large; a low dorsal keel on tail be- hind sting; base of tail horizontally oval in cross section; ventral surface of disc light without a broad dusky marginal band 10 1 Oa. Back flattened, without an enlarged mid- scapular pearl spine (sometimes a row of up to 4 moderately enlarged flattened spines in its place); midbelt of flattened denticles reduced or absent, disc some- times entirely naked; snout long, preor- bital length 2.8-3.2 times interorbital space (down to 2.3 in late fetuses or new- bom specimens) and 27-32% of disc width; disc very flat, thickness at scap- ulocoracoid 8-11% (usually less than 1 1%) of disc width; lateral prepelvic pro- cesses of pelvic girdle greatly expanded D. garouaensis (Stauch and Blanc, 1962) 10b. Back somewhat arched, with an enlarged midscapular pearl spine; midbelt of flat- tened denticles well developed in large juveniles and adults; snout shorter, preorbital length 1.5-2.4 times interor- bital space and 1 9-26% of disc width; disc thicker, 1 1-15% of disc width over scapulocoracoid; lateral prepelvic pro- cesses slightly expanded 1 1 1 1 a. Upper jaw strongly undulated, with teeth greatly enlarged on prominent lateral projections; teeth less numerous, in 26- 29/31-34 rows; snout more broadly pointed; pearl spine usually larger and circular, length about 5-6 mm; pectoral radials 133-135; size larger, adults to 65 cm D. margarita (Gunther, 1870) 1 1 b. Upper jaw moderately undulated, with teeth moderately enlarged on low lateral projections; teeth more numerous, in 36- 42/38-50 rows; snout usually more acutely pointed; pearl spine usually smaller and often axially oval, length 2- 4 mm; pectoral radials 116-127; size smaller, adults to 26 cm D. margaritella new species Dasyatis Rafinesque, 1810 Dasyatis Rahnesque, 1810:16 (type-species Dasyatis ujo Ra- finesque, ISIO [=Rajapastinaca Linnaeus, 1758], by mono- typy). For full generic synonymy of Dasyatis see Bigelow and Schroeder(1953). Diagnosis. — Dasyatidae with disc circular, oval, trapezoidal, or diamond-shaped (Fig. 1), its dorsal surface smooth or variably covered with small, flat or prickle-like denticles; large, sharp, spine- or plate-like denticles present or absent on dorsal surface; snout angular or broadly rounded, its projecting tip variably developed; pectoral fins rounded or angular; pelvic bar mod- erately arched; tail long, slender, with dorsal and ventral folds or ventral folds only; ventral fold not reaching tip of tail; sting usually present (re- duced or absent in Dasyatis ukpam). Teeth small, rhomboidal, thin-crowned. Dasyatis as here recognized is a large, heter- ogeneous assemblage of about 33 species and may be polyphyletic. Dasyatis margarita, D. margaritella, and D. garouaensis are not far re- moved morphologically from the generic type- species D. pastinaca. Dasyatis ukpam, however, is distinct, approaching Urogymnus Miiller and Henle, 1837 in general morphology, heavy den- COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 287 Figure 2. (a) Dasyatis margarita, lectotype, 200-mm immature female. West Africa (BMNH 1865.7.4:1); (b) Dasyatis margaritella, 226-mm mature male, Conakry, Guinea (ISH 183/63) ticulation, and sting reduction. Urogymnus species invariably lack the sting, while specimens of D. ukpam either lack the sting or have a very small one. Smith (1863) noted that D. ukpam seemed intermediate between Urogymnus and Trygon {=Dasyatis), at least in the nature of its sting, but included it in Hemitrygon MuUer and Henle, 1837 {= Dasyatis) because it has a short ventral tail fold and no dorsal tailfold. We retain D. ukpam in Dasyatis pending modification of the limits of Dasyatis and other dasyatid genera. The species is readily distinguished from known Urogymnus species in having a ventral tailfold, much longer tail (less than 1.5 times disc width in Urogymnus), a less thick, more circular disc, darker dorsal coloration (dorsal surface pale brown in all Urogymnus examined), a dark mar- ginal band on ventral surface of disc, smaller flat denticles on dorsal surface of disc, and in some specimens a small sting. Dasyatis margarita (Gunther, 1870) (Figure 2a) Trygon margarita Gunther, 1870:479 (type-locality West Af- rica). Dasyatis sp. Blache et al., 1970:53, fig. 117. Material Examined. — BMNH 1865.7.4:1, 200-mm im- mature female. West Africa (formerly syntype of Z). margarita; designated lectotype below); USNM 222589, 1 30-mm late fetal or newborn male. Sierra Leone; BMNH 1930.3.24:3, 21 2-mm immature male, Accra, Ghana; BMNH 1936.8.20:2-3, 216- mm and 3 1 5-mm immature females, Lagos, Nigeria; BMNH 1 899.2.20:35, 206-mm immature female. Banana, Congo Riv- er mouth, Zaire; AMNH 40408, 235-mm female, Angola. Lectotype Designation. — In the original description of D. margarita Gunther ( 1 870:479) listed two specimens from West Africa without indicating either as holotype: a. Disk 8'/2 inches long, tail 19 inches. Purchased of Mr. J. Wood. b. Young. From the collection of the Zoological Society. These two specimens are therefore syntypes, but our studies indicate they are not conspecific. Specimen a is BMNH 1865.7.4: 1 , a 200-mm immature female with a single, large, round pearl PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. U ii -J vn ^1 ^ ON C c 2 e 5:5 I C ro C o — 2 ^ a o 44 0\ O m ooor-oo — (Nr^TtoOOO — — — — Tt f<-l\0— 'OOO'/^Or^OOt^fN0va- O ^ rn vo r. I H-i u — O Ov ■* r~ — >o w-> 1^ l> ^o 1^ (N fl VO O ■* O ro — OV Ov O r^ [^ r^ . cen- troura). Trygon ukpam Gunther, 1870:480 (description after Smith, 1863, placed in Trygon = Dasyatis). Dasyatis margarita Lxjubens, 1964:1 1 (freshwater lakes south of Lambarene district, Ogooue basin; presumably no spec- imens preserved). Dasyatis ukpam Stehmann, 1 98 1 :4 (in key to marine Dasyatis of West Africa); Compagno and Roberts, 1982:321 (ref- erence). Material Examined. -BMNH 1874.5.23.1, syntype, 266- mm late fetal male. Old Calabar River, Nigeria; USNM 219780, 520-mm immature female. Lake Ezanga, Ogooue River sys- tem, Gabon; CAS 42761, 650-mm immature female. Lake Ezanga near Nzame-Akesile village, Ogooue River system, Ga- bon; MNHN 1979-244, 499-mm immature female, Booue, Ogooue River, Gabon; MRAC 55778, 361-mm immature fe- male, Binda, Congo (Zaire) River, Zaire. Diagnosis.— Z). ukpam is a very large and thick-bodied freshwater dasyatid, probably growing much bigger than our largest specimen, a 650-mm immature female. Newborn young, 266 mm, are as large as the largest known D. margaritella and far larger than newborn young of D. margarita and D. garouaensis. Entire dor- sal surface of disc covered with stout-spined den- ticles (smooth in newborn) (peripheral portions of disc without denticles in all other West African dasyatids except the very large marine species D. centroura, with a diamond-shaped disc, and Uro- gymnus, with an extremely thick disc). Sting greatly reduced in size or absent (sting invariably absent in Urogymnus but normally present and relatively large in all other West African dasy- atids). Disc oval, very thick. Medial lobe narrow- based and short, its anterior margin broadly con- vex; disc depth 13.3-15.7% of disc width, disc width 0.9-1.0 times disc length; dorsal surface of disc entirely covered with denticles at all free- living stages except possibly newborn; small, heart-shaped or circular denticles covering mid- dle of disc, small prickles on sides of disc, with scattered large, conical, stellate, sharp denticles (absent in newborn and fewer in small immatures than large) and usually 1-3 small to moderate- sized midscapular pearl spines on disc; no sting, or a single smaU, possibly vestigial one, only 6.5% of disc width in 520-mm specimen; eyes small and strongly elevated, eyebaU lengths 2.1-3.3 times in interorbital space; interorbital space 1.5- 1.9 times in preorbital length; spiracles large and high; floor of mouth with 4 or 5 elongated pa- pillae; total tooth rows 38-46/38-48; pelvic fins short, anterior margins 16-19% of disc width; tail uniformly tapering to a slender whiplash, its length from vent when intact 2.0-2.9 times disc width; base of tail circular in section; a very low ventral tailfold but- no dorsal fold or keel; disc and pelvic fins dark brown or gray-brown above, without spots or prominent markings, white be- low except for broad dark margins; tail blackish except for underside of its base; intestinal valve turns 19-20; total pectoral radials 142-148, total vertebral centra 108-122 and total segments 155 (in one specimen), vertebral centra ending in front of sting or extending to its origin. Proportional measurements (as percent of disc width) and counts are given in Table 1 . Preoral length 2.3-2.6 times intemarial width and 0.9- 1 .0 times width between first gill openings. Snout broad, angle in front of eyes 123-132°. Spiracle PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. li Figure 6. Dasyatis ukpam, 266-mm fetal male syntype, Old Calabar, Nigeria (BMNH 1874.5.23:1). Figure 7. Dasyatis ukpam, 650-mm immature female, Lake Ezanga, Ogooue basin, Gabon (CAS 42761). COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 297 Figure 8. Dasyatis ukpam, 520-mm immature female, Lake Ezanga, Ogooue basin, Gabon (USNM 219780). length 1.5-2.0 times eyeball length, 0.8-1.0 times intemarial width, and 1.8-2.2 times in distance between fifth gill openings. Internal gill openings with close-set transverse ridges on gill arches. Nasal curtain with a fringed, slightly concave or trilobate posterior margin. Mouth weakly arched, midline of lower jaw slightly indented; a shallow to deep, curved groove extending posteriorly from posterior nasal flap around comers of mouth. Skin on ventral surface of lower jaw more or less Figure 9. Dasyatis ukpam, 36 1 imature female, lower Zaire River at Binda, Zaire (MRAC 55778). PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. Figure 1 0. Pelvic girdle (dorsal view) in West African Das- yatis: (a) D. ukpam, 266-mm fetal male; (b) D. ukpam, 361- mm immature female; (c) D. garouaensis, 342-mm mature female; (d) D. garouaensis, 255-mm adult male; (e) D. mar- garita, 200-mm immature female; (f) D. margaritella, 190- mm mature male. Note prominent lateral prepelvic processes in D. garouaensis. Scale bar = 2 cm. corrugated and papillate. Palate behind fringed maxillary valve with three strong, short ridges, a medial longitudinal ridge and a pair of diagonal lateral ridges. Two pairs of long, low ridges be- hind the three palatine ridges. Floor of mouth with a transverse row of usually three moderately large, elongated oral papillae, a medial papilla (absent in syntype) and a lateral papilla behind each end of dental band; a pair of large papillae just behind and lateral to medial papilla. Upper jaw with about 4 and lower jaw with 6 functional tooth series. Teeth in quincunx pave- ment, close-set, with crowns closely adjacent to one another. Teeth similar in upper and lower jaws, uppers slightly larger than lowers at sym- physis, varying continuously lateral to symphy- sis, teeth largest and longest relative to width at or near symphysis and smallest and shortest at mouth comers. Upper dental band with a low knob of slightly enlarged teeth at symphysis, sep- arated from similar knobs on either side by depressions with smaller teeth; lower dental band with a weak symphyseal depression into which symphyseal knob of upper jaw fits; a pair of low lateral knobs fit into depressions in upper dental band. Teeth of both jaws in females with mod- erately high, broad-tipped, cuspless crowns shaped like mushroom caps, with a transverse, blunt keel or carina, strong transverse ridges on both labial and lingual sides, and a strong basal ledge and concave, inset basal groove separating crown and root. Roots moderately large, high, pedicellate, bilobed basally, with a transverse groove and nutrient foramen. Teeth of adult males unknown, but probably differ from those of fe- rnales. Dorsal surface of 266-mm late fetus covered with small flat denticles on middle third of disc and base of tail to its abbreviated sting, including dorsal surface of cranium, branchial region, scap- ular region, and abdominal area; in addition, specimen has two small, slightly elevated, en- larged rounded denticles or pearl spines in medi- scapular area. The larger, free-living specimens examined have outer two-thirds of disc, snout, and tail posterior to sting region with small to moderately large, conical, prickle-like denticles in addition to flattened denticles covering mid- belt; they also have massive, conical, erect, flut- ed, sharp denticles or thorns over much of dorsal surface of disc and tail base. Large thorns fewest on smallest (36 1 -mm) specimen, most numerous on largest (650-mm), suggesting that they be- come more numerous with growth. These thorns make dorsal surfaces of large D. ukpam ex- tremely rough, and, as noted by Smith (1863), difficult to handle. Free-living specimens ex- amined have one or two round, enlarged pearl spines on mediscapular region. Neurocranium observed on radiographs but it and other skeletal parts obscured by thickness of disc and heavy covering of denticles in this species. Cranium apparently similar to that of Himantura signifer as described in Compagno and Roberts (1982), but with a straighter anterior margin to its nasal capsules. Pelvic girdle (Fig. lOa-b) broadly arched, semicircular, relatively thick, with a medial an- terior angle but no medial prepubic process. Lat- eral prepubic processes low, rounded, and lobate; iliac processes well-developed; ischial processes short. Five obturator foramina present on one specimen. Claspers of mature male not available. Discussion It has been known for some time that stingrays identified as Dasyatis margarita represent two species (Daget and litis 1965; Blache et al. 1970; Stehmann 198 1). It appears that the first ichthy- ologist to become aware of this was the late J. Cadenat; he recognized that the two species differ greatly in weight, the one not exceeding 1 kg while the other attains easily 1 5-20 kg and per- haps much more (quoted in Daget and litis 1 965: 15). The small species is D. margaritella, the COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS large one D. margarita. Both are common and widely distributed in shallow water along the coast of West Africa. Most accounts of D. margarita are based on D. margaritella or on both D. mar- garitella and D. margarita. D. margaritella is more common than D. margarita in museum collections and perhaps also in nature. We are unaware of any difference in habitat preference. It is remarkable that such notable animals as freshwater stingrays remain so poorly known. We believe that they will be found in additional river basins in West Africa and that possibly additional freshwater species are present. We have heard that stingrays occur in the Sanaga basin in Cameroun, particularly in Lac Ossa, but have no material evidence for this. It is curious that no Dasyatidae have been reported from rivers west of Nigeria. This might be due to insufficient col- lecting. A stingray was reported from the Cross River at Mamfe Pool by Sanderson (1937), but the specimen was not preserved, and the account is so extraordinary that we hardly know what to make of it. According to Sanderson, the ray was "diamond-shaped, like all fish of this class, and measured from the tip of one lateral point to the tip of the other, four feet eight inches; from the snout to the base of the tail, five feet eleven inches; and from the base to the tip of the tail, which had no fin, five feet two inches. Emerging from the upper edge of the tapering whip-like tail near its base was a long, straight, sharp spine or sting, one foot seven inches in length." He went on to say that the arrival of this monster altogether un- hinged his sense of logic, so perhaps it also af- fected his ability to observe and record accu- rately. "That it was still alive and therefore undoubtedly caught in Mamfe Pool, as the na- tives stated, was almost incredible, because this bit of water was nearly three hundred miles from the sea. I therefore had to adjust myself to the idea that such things are true fresh-water animals indigenous to the great rivers of Africa. Why do not natural history books depict these fish in- stead of the everlasting crocodile?" Why not, in- deed? Sanderson provides as good an answer as any: "We didn't want the brute because we were not collecting fish, but we photographed him alongside sundry natives and inanimate objects and purchased the sting." Unfortunately the pho- tograph was not published and the sting had to be discarded: "When this sting got really dry it split longitudinally and opened like a star, re- vealing a clear crystaUine plug within. This sub- stance gradually broke up under the damp at- mospheric conditions; some of it dropped in water fizzed furiously. I could not find anything that would preserve it among our selection of trav- elling drugs and chemicals." We have seen dried stings of various rays with- out observing anything like the disintegration re- ported by Sanderson, and suspect he had a few chuckles and perhaps something else while con- cocting this giant sting. The fishermen at Mamfe, who regularly catch D. garouaensis, maintained that it is the only species of ray in the Cross River from Mamfe to the Nigerian border and that the example caught and preserved during the junior author's visit was nearly as large as the largest they had ever caught. When shown a photo- graphic print of D. ukpam they indicated that this species was unknown to them. Concerning sting size, the largest sting we have observed on a West African freshwater ray is under four inches. In Dasyatis centroura, which probably has the largest sting of any marine species in West Africa, the sting of a specimen with a disc width of four feet eight inches would be only about six or seven inches. We conclude that D. garouaensis is the only species of stingray in the Cross River in the vicinity of Mamfe. Identification of a specimen oi D. garouaensis from Lagos may indicate that the species occurs in Lagos Lagoon and in the rivers flowing into it. Unfortunately the specimen is not accom- panied by information on habitat, and we cannot rule out the possibility that it was caught in the Niger River and transported to Lagos. Although the title of the paper in which D. ukpam was described states that it lives in the Old Calabar River, and the text indicates that it Hves as much as 1 SO^niles upriver (Smith 1 863), this is based on hearsay. It should be noted that the vernacular name "ukpam" or "okpam" is a generical term for stingrays. At Mamfe this name is employed by present-day speakers of the Man- yu and Ejagam languages for D. garouaensis. The occurrence of an unidentified stingray in the Ogooue basin was first brought to our atten- tion by an ichthyological colleague, Dr. Jaques Gery, who related to the junior author that he had observed two rays in the Ivindo near Ma- kokou while collecting characins and other small fishes in 1 964. The Ivindo flows into the Ogooue PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 near Booue, where one of our specimens of D. ukpam was collected, but there are some for- midable waterfalls on the Ivindo below Mako- kou and the species there might be different. The Zaire locality for D. ukpam, Binda, is on a rel- atively narrow and swift-flowing portion of the lower Zaire (Congo) River about 1 00 km upri ver from the mouth of the river at Banana and 35 km downriver from the end of the mainstream rapids of the lower Zaire River near Matadi. No stingrays are known from the interior of the Zaire or Congo basin. So far as we have been able to determine D. garouaensis and D. ukpam are the only living freshwater stingrays known from Af- rica. Acknowledgments For providing information and facilitating ex- amination of specimens we wish to thank Al- wyne Wheeler, Mandy Holloway, and Oliver Crimmen, British Museum (Natural History); D. F. Thys van den Audenaerde, Musee Royale de I'Afrique Centrale, Tervuren, Belgium; and Thomas B. Thorson, School of Life Sciences, University of Nebraska. Fieldwork in Gabon was facilitated by the Centre National pour la Re- cherche Technique et Scientifique and particu- larly M. le Commissaire Nzoghe-Ngueme. We thank Jeanne Byloghe and Joseph Mebiaghe of Tame and Nzame-Akesile villages for catching Dasyatis ukpam, and Pierre Ville of Geri Con- sult, who also obtained a specimen of this species for us. Permission for fieldwork in Cameroun was kindly granted by the Office National pour la Recherche Scientifique et Technique. At Mamfe the junior author was assisted by S. T. Mbianyor of the Forestry Department, John Corrao and Alan Ferguson of the Peace Corps, and a number of excellent local fishermen. Literature Cited BiGELOw, H. B., AND W. C. ScHROEDER. 1953. Fishcs of the western North Atlantic. Sears Found. Mar. Res., Mem. No. 1(2). Blache, J., J. Cadenat, AND A. Stauch. 1970. Cles de de- termination des poissons de mer signales dans I'Atlantique oriental. Faune Trop. (ORSTOM, Paris), No. 18. Castello, H. 1973. Sobre la correcta posicion sistematica de la raya de agua dulce africana (Chondrichthyes, Dasy- atidae) (Republic Federal del Camerun). Trab. V Congr. La- tinoamer., Zool. 1:67-71. Castex, M. 1967. Fresh water venomous rays. Pages 167- 176 in Animal toxins. Pergamon Press, Oxford and New York. CoMPAGNo, L. J. v., AND T. R. ROBERTS. 1982. Freshwater stingrays (Dasyatidae) of Southeast Asia and New Guinea, with description of a new species of Himantura and reports of unidentified species. Environ. Biol. Fishes 7(4):32 1-339, 12 figs. Daget, J., AND A. Iltis. 1965. Poissons de Cote d'lvoire (eaux douces et saumStres). M6m. Inst, franc. Afr. noire no. 74. Daget, J., AND A. Stauch. 1963. Poissons de la partie Ca- merounaise du bassin de la Benou6. M6m. Inst, franc. Af- rique noire 68:85-107. Fowler, H. W. 1936. The marine fishes of West Africa. Bull. Amer. Mus. Nat. Hist. 70(1): 1-606. . 1969. Acatalogofworldfishes(XI). Quart. J. Taiwan Mus. 22(3-4):125-190. GuNTHER, A. W. 1870. Catalogueofthefishesin the British Museum. 8. Krefft, G. 1968. Knorpelfische (Chondrichthyes) aus dem tropischen Ostatlantik. Atlantide Rep. 10:33-76, pis. 3-6. LouBENs, G. 1964. Travaux en vue de developpement de la peche dans le bassin inferieur de I'Ogooue. Publ. Cent. Tech. Forest. Trop., Nogent-sur-Mame (Seine) 27:1-51. Rafinesque, C. S. 1810. Caratteri di alcuni nuovi generi e nuove specie di animali (principalmente di pesci) e piante della Sicilia, con varie osservazioni sopra i medisimi. Pa- lermo, 105 pp, 20 pis. Reid, G. M., AND H. Sydenham. 1979. A checklist of lower Benue River fishes and an ichthyogeographical review of the Benue River (West Africa). Sanderson, I. 1937. Animal treasure. Viking Press, New York, 325 pp. Smith, J. A. 1863. Notice of the ukpam, a large species (probably new) of sting ray (Trygon, Cuvier), found in the Old Calabar River, Africa. Proc. Roy. Phys. Soc. Edinburgh 1859-62,2:64-69. Stauch, A., AND M. Blanc. 1962. Description d'un s61acien rajiforme des eaux douces du Nord-Cameroun, Potamotry- gon garouaensis n. sp. Bull. Mus. Nat. Hist. Natr. (Paris) 34(2): 166-1 71. Stehmann, M. 1981. Dasyatidae. In Fischer, W., G. Bianchi, and W. B. Scott (eds.). FAO species identification sheets for fishery purposes. Eastern Central Atlantic. Fishing Area 34, 47(5): 1-5. Thorson, T. B., and D. E. Watson. 1975. Reassignment of the African freshwater stingray, Potamotrygon garouaensis, to the genus Dasyatis, on physiologic and morphologic grounds. Copeia 1975(4):701-712, figs. 1-3. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94 1 1 8 PROCEEDINGS OF THE ■ i ■ ■■- CALIFORNIA ACADEMY OF SCIENCES ■■'35 Vol. 43, No. 19, pp. 301-315, 12 figs., 1 table. December 11, 1984 REVISION OF EASTERN PACIFIC CATALUFAS (PISCES: PRIACANTHIDAE) WITH DESCRIPTION OF A NEW GENUS AND DISCUSSION OF THE FOSSIL RECORD By John E. Fitch Research Associate, California Academy of Sciences, San Francisco, California 94118 and Stephen J. Crooke California Department offish and Game, Long Beach, California 90802 Abstract: Four species of catalufas inhabit eastern Pacific waters: Cookeolus boops (Schneider, 1801), Heteropriacanthus cnientatus (Lacepede, 1801), Pseudopriacanthus serrula (Gilbert, 1891), and Priacanthus alalaua Jordan and Evermann, 1904. Each of these species is illustrated, and diagnostic characters, meristic data, morphometric measurements, maximum size, geographic range, depth distribution and other informa- tion also are presented. Heteropriacanthus is a new generic name for Priacanthus cnientatus, a cosmopolitan species that differs in numerous salient features from the species assignable to Priacanthus (i.e., alalaua, arenatus, hamrur, macracanthus, meeki and tayenus). An identification key is presented. Otoliths (sagittae) and scales of the four eastern Pacific species also are illustrated. The only reported priacanthid fossils are from the Eocene of Europe. Of the six species, Pristigenys substriata is known from skeletal remains and is unquestionably a priacanthid. Only two of the five species described from otoliths, Pristigenys bella and P. dentifer, appear to be priacanthids, but there is no assurance they can be assigned to Pristigenys, since none of the skeletal "imprints" of P. substriata contained otoliths, nor do all of the otoliths assigned to these two species appear to be correctly identified. Introduction partment of Fish and Game (DFG) biologists, For years, fishermen aboard long-range sport- skippers and crew members of these vessels com- fishing boats that operated out of San Diego menced fishing for and saving miscellaneous sought only such large game species as yellowfin small fishes caught at the Revillagigedo Islands, and bigeye tuna ( Thunnus albacares and T. obe- Alijos Rocks, and other fishing spots off southern sus), wahoo {Acanthocybium solanderi), yellow- Baja California. tail (Seriola lalandi), giant sea bass {Stereolepis Among the first of these incidentally caught gigas) and several kinds of large serranids {Epi- species turned over to DFG personnel were a nephelus spp. and Mycteroperca spp.). During half-dozen catalufas that appeared to represent 1978, however, at the urging of California De- three species of Priacanthus. A literature search [301] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 Figure 1. Type specimen ofAnthias macrophthalmus Bloch, 1792 (ZMB 8156). Photo courtesy of Hans-J. Paepke, Museum fur Naturkunde der Humboldt-Universitat zu Berlin. and examination of all the priacanthids in the fish collections at Scripps Institution of Ocean- ography (SIO), Natural History Museum of Los Angeles County (LACM), University of Califor- nia, Los Angeles (UCLA) and California Acad- emy of Sciences (CAS) revealed that, until then (Dec. 1978), any priacanthid collected in the eastern Pacific that was not judged to be either Pseudopriacanthus or Cookeolus, automatically had been relegated to Priacanthus cruentatus (see also Fitch and Schultz 1978). Once it became obvious that we were dealing with several species, the differences in scale counts, pelvic fin pigmentation, eye diameter, otoliths, and gas bladder morphology became equally obvious. During the succeeding four years, in an effort to determine exactly what species we were observing, we examined several hundred priacanthids from throughout the world. As a result, we arrived at the conclusion that there were four species of catalufas in the eastern Pa- cific: Cookeolus boops, Priacanthus alalaua, Priacanthus cruentatus and Pseudopriacanthus serrula. In the course of our investigation, we found that otoliths (sagittae) and gas bladders of Pria- canthus alalaua were so radically different from otoliths and gas bladders of P. cruentatus that a new generic name was needed for one of the two. This necessitated determining the condition of the otoliths and gas bladder of Bloch's (1792) Anthias macrophthalmus, since it was the type- species for Priacanthus Oken, 1817. Fortunately, Bloch's type specimen, a skin from the right side of the fish, still exists in the Museum fur Natur- kunde der Humboldt-Universitat zu Berlin (ZMB8156), and Hans-J. Paepke, Curator of Fishes, sent us an excellent photograph of this specimen (Fig. 1). Wayne Stames (pers. comm.), to whom we sent a copy of the photograph, has confirmed that Bloch's Anthias macrophthalmus is conspecific with Sciaena hamrur Forsskai, 1775. Key to Eastern Pacihc Priacanthidae la. Pored scales in lateral line 35 to 40; dorsal soft rays 1 1 (rarely 10 or 12); anal rays 10; dorsal profile turns abruptly downward under base of soft portion of second dorsal fin; ventral profile turns abruptly upward HTCH & CROOKE: REVISION OF EASTERN PACIHC CATALUFAS Figure 2. Pseudopriacanthus serrula. Photo by Jack W. Schott. under anal fin base; ctenii on scales in par- allel species, presenting a corrugated or waffled appearance 3a. Pseudopriacanthus serrula lb. More than 50 pored scales in lateral line; 1 2 (rarely) or more dorsal soft rays; 1 2 or more anal rays; dorsal and ventral profiles tapering gradually to caudal peduncle; ctenii on scales not as above 2 2a. Eighteen to 20 scale rows between 8th dor- sal spine and lateral line; more than 20 rows on dorsum of caudal peduncle; anal rays 13; peduncle depth about equal to horizontal eye diameter; dorsal, anal and 3b. caudal fins yellow, edged with black; ctenii thick and bristlelike, in rows Cookeolus boops 2b. Eight to 10 scale rows between 8th dorsal spine and lateral line; fewer than 1 5 scale rows on dorsum of caudal peduncle; anal rays 14; peduncle depth fits about 1 Va times into horizontal eye diameter; dorsal, anal and caudal fins never with yellow or tipped with black; ctenii thin and filamentous 3 Second dorsal, anal and caudal fin mem- branes red, without spotting; pelvic fin membranes jet black; gas bladder with an- teriorly projecting "ears" that extend to the otic bullae, and posteriorly projecting horns that reach to above end of anal fin; otoliths (sagittae) with a ventral keellike blade and centrally positioned pronglike rostrum; ctenii filamentous, in rows; pre- opercle completely scaled Priacanthus alalaua Second dorsal, anal and caudal fin mem- branes with rust-colored spotting (dusky in formalin-preserved specimens); pelvic fin membranes lightly pigmented to clear; gas bladder contained entirely within body cavity, without ears and no anterior or posterior projections; otoliths (sagittae) lack keellike blades, oval in outline, with normal, anteroventral rostrum; ctenii fil- 304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 amentous, only on scale margin, sparse; posterior margin of preopercle without scales Heteropriacanthus cruentatus new genus Species Accounts Pseudopriacanthus serrula (Gilbert, 1891) (Figure 2) Diagnostic Characters. — 35 to 40 pored scales in lateral line; 1 to 12 (typically 1 1 ) dorsal soft rays; 1 anal rays; ctenii on scales in a par- allel series (Fig. 3d) presenting a corrugated or waffled appearance; dorsal profile turns abruptly downward, becoming nearly vertical, under sec- ond dorsal fin base; ventral profile becomes near- ly vertical under anal fin base. Meristic Data.-D. X, 10-1 2; A. 111,10; P. 17; GR 6-7 + 15-18 = 21-25; pored lateral line scales 35-40; vertebrae 10 + 13 = 23. Twenty- three of the 25 specimens we examined for me- ristic data had 1 1 dorsal soft rays, one had 1 0, and one had 12. Maximum Size.— The largest individual we observed was a female 274 mm SL (353 mm TL) that weighed 1 300 g. This was one of 24 indi- viduals caught in gill nets in "30-40 fm" (55- 73 m) off Magadalena Bay, Baja CaUfomia, dur- ing March 1976. Range. — Monterey Bay, California, to Talara, Peru (Fitch and Lavenberg 1975) and at most offshore islands from the Coronados to the Ga- lapagos in 3.6 to more than 100 m (DFG, un- published data). Remarks. — Morphometric data were taken on only 1 3 of the more than 60 individuals we ex- amined (Table 1). The smallest of these (34 mm SL, LACM 22796) had a relatively short pelvic fin (41% of SL) as compared with the Eocene fossil Pristigenys substriata (Fig. 1 1) and pelagic stages of the extant Cookeolus boops (Fig. 5, Ta- ble 1). Fritzsche and Johnson (1981) considered Pseudopriacanthus a junior synonym of Pristi- genys, but for reasons given later, we believe both genera are distinct and valid. Of perhaps 20 individuals that have been caught or observed in Califomian waters, one each was from Monterey Bay and off San Luis Obispo, the rest have been from south of Pt. Dume. Scuba divers report that P. serrula is un- afraid and can be picked up by hand when en- countered in its natural surroundings. Cookeolus boops {Schneider, 1801) (Figure 4) Diagnostic Characters.— 1 8 to 20 scale rows between eighth dorsal spine and lateral fine; more than 20 rows of scales on dorsum of caudal pe- duncle; ctenii on scales thick and bristlelike (Fig. 3a), in rows; anal rays 12-13; dorsal, anal and caudal fins yellow, edged with black; peduncle depth about equal to horizontal eye diameter; pelvic fins of pelagic juveniles comprise 50-70% of SL or more. Meristic Data.-D. X,13; A. 111,12-13; P. 18-19; GR 6-8 + 17-18 = 23-26; pored lateral line scales 53-61; vertebrae 10 + 13 = 23. One of the 35 specimens we examined for meristic data had 12 anal rays, the rest had 13. Maximum Size.— The largest individual we observed was a female 397 mm SL (507 mm TL) that weighed 2725 g. It was caught by a sport- fisherman at Alijos Rocks, Baja California, Mex- ico, in 1974 in "18 fm" (33 m) of water. This fish (LACM 34253) was erroneously identified by Fitch and Schultz (1978) as Priacanthus cruentatus. In the western North Atlantic, a C. boops has been reported that was 507 mm SL and weighed 5.2 kg (Anderson et al. 1972). Range.— Worldwide in tropical and subtrop- ical seas; in the eastern Pacific from Alijos Rocks, Mexico (24°57'N, 115°45'W) to 10°N 98'W (LACM 30506-1). Although adults have been caught in water as shallow as 30 m in the eastern Pacific, they are most commonly hooked at depths "exceeding 40 fm" (73 m) and have been taken as deep as "75 fm" (137 m). Pelagic juveniles have been captured at the surface at scattered offshore localities between the Tres Marias Is- lands and the Gulf of Tehuantepec. In the west- em North Atlantic, C. boops has been taken at depths exceeding 365 m (Anderson et al. 1972). Remarks.— Although Cookeolus was not rec- ognized from the eastern Pacific until Fritzsche (1978) reported upon six pelagic juveniles rang- ing from 148 to 226 mm SL, adults had been taken at Alijos Rocks as early as 1970, but were erroneously identified as Priacanthus cruentatus (Fitch and Schultz 1978). Not until 1978, when long-range sportfishing boats started bringing in fair numbers of adult C boops from Alijos Rocks, the Revillagigedo Islands and Hurricane Bank (16°52'N, 117°28'W) were they recognized for what they were. Most of the confusion in iden- tification had resulted from the relatively shorter HTCH & CROOKE: REVISION OF EASTERN PACIHC CATALUFAS 305 Figure 3. Scales of eastern Pacific Priacanthidae (scale height and length, in mm. in parens): a) Cookeolus boops (4.6 by 3.5); b) Priacanthus alalaua (3. 1 by 3.1); c) Heteropriacanthus cruentatus (\ .9 by 2.4); d) Pseudopnacanthus serrula (3.6 by 4.0). SEM photos by Richard Huddleston. pelvic fins of the bottom-dwelling adults and the lack of spots and blotches which are so typical of the pelagic juveniles (Fritzsche 1978). Although pelvic fin length commences to shorten when Cookeolus takes up a bottom- dwelling existence, six of the specimens we used in obtaining morphometric data (Table 1) still had pelvic fins that exceeded 40% of SL. All six of these fish, ranging from 197 to 267 mm SL (Fig. 5), had been hooked on the bottom. On the other hand, two bottom-dwelling specimens (222 and 257 mm SL) had pelvic fins that had short- 306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 Figure 4. Cookeolus boops. Photo by Lee Stockland. PERCENT OF STANDARD L E N G T t- PELVIC FIN LENGTH T 1 1 1 1 1 1 1 1 1 1 I I 160 180 200 220 240 260 280 300 320 340 360 380 400 STANDARD LENGl Figure 5. Pelvic fin length (as percent of SL) plotted against SL for 32 specimens of Cookeolus boops. HTCH & CROOKE: REVISION OF EASTERN PACIHC CATALUFAS 307 .^4\^ Figure 6. Juvenile Cookeolus boops, 211 mm SL, taken on hook and line in "15 fms" at Alijos Rocks. Photo by Paul Gregory. Table 1 . Comparative Measurements for the Four Eastern Pacific Priacanthids (in percent standard length). Species and number of specimens measured Pseudopriacanthus Cookeolus Phacanthus Heteropriacanthus Measurement serrula boops alalaua cruentatus Number of specimens 13 32 14 50 Standard length (mm) 34-261 151-392 215-261 81.5-245 Head length 36.1-41.2 31.2-37.4 29.4-33.3 28.6-36.4 Horizontal eye diameter 14.3-18.0 9.6-13.9 12.5-14.1 10.2-15.2 Snout length 9.0-11.4 9.8-11.7 '9.5-10.5 9.0-10.1 Maxillary length 20.0-21.6 16.6-19.2 14.7-16.1 15.5-17.1 Bony interorbital width 7.9-10.2 8.2-10.5 8.0-9.1 8.4-10.1 Snout to 1 St dorsal fin insertion 33.7-36.4 28.0-33.3 30.0-32.3 29.7-32.3 Snout to pectoral fin insertion 37.6-40.8 31.7-37.9 30.9-34.8 30.1-32.9 Snout to pelvic fin insertion 44.8-46.8 34.0-44.0 33.3-38.8 35.4-38.3 Snout to anal fin insertion 70.9-75.6 58.1-67.9 55.3-60.9 51.3-59.0 Dorsal fin insertion to pelvic insert 48.4-53.6 37.7-48.8 36.4-39.1 35.0-40.3 Depth perpendicular to AS, 48.8-54.2 37.1-49.2 35.6-38.6 34.3-39.8 Caudal peduncle depth 12.8-14.0 9.6-12.3 7.7-8.2 8.3-10.5 Pectoral fin length 19.3-23.2 18.9-24.8 20.8-23.2 17.0-21.0 Pelvic fin length 25.6-35.4" 27.0-70.2^ 25.3-31.4 19.7-26.7 Longest gill raker 5.9-8.8 5.0-7.2 4.1-5.4 4.4-5.2 Pelvic fin length of 34 mm specimen (41.2%) not included, remaining specimens 172 to 261 mm SL. See Figure 5. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 Figure 7. Priacanthus alalaua. Photo by Lee Stockland. ened to 34% of SL (Figs. 5 and 6). The largest pelagic juvenile reported by Fritzsche (1978) was 226 mm SL. Based upon these lengths, in the eastern Pacific, Cookeolus remains in the pelagic environment until it reaches a size of approxi- mately 200-250 mm SL. Interestingly, the otoliths (sagittae) of Cook- eolus also change with adulthood (Fig. 8c, d). Those of pelagic juveniles are almost perfectly oval in outline with a ratio of height into length of about 1:1.25. In adults, the otolith becomes more elongate (height into length ratio 1:1.6) and the posteroventral border becomes slightly con- cave as the marginal ornamentation (lobules) spreads and the notches deepen. We do not know of any other perciform in which such differences in otolith morphology occur with age. Priacanthus alalaua Jordan and Evermann, 1 904 (Figure 7) Diagnostic Characters. — Second dorsal, anal and caudal fin membranes red; pelvic fin membranes jet black; gas bladder with anteriorly projecting "ears" that extend beyond the body cavity to the otic bullae, and posteriorly pro- jecting horns that reach to above the end of the anal fin (Fig. 9); otoliths (sagittae) with a ventral keellike blade and centrally positioned pronglike rostrum (Fig. 8e); preopercle completely scaled; ctenii filamentous, in rows (Fig. 3b). Meristic Data.-D. X,13; A. 111,14; P. 18- 19; GR 4-6 + 14-17 = 19-23; pored lateral line scales 61-66; vertebrae 10 + 13 = 23. Only one of the 20 specimens we examined for meristic data had 1 9 elements in the pectoral fin; the rest had 18. Maximum Size.— The longest individual we observed was a female 26 1 mm SL (335 mm TL) from San Benedicto Island, Revillagigedos. The heaviest individual was a 257 mm SL female from Socorro Island, Revillagigedos, which weighed 574 g. Gosline and Brock (1960) report that in Hawaiian waters P. alalaua "reaches 1 4 inches in length" (357 mm), but they apparently did not examine any specimens of this species. Range. — Hawaiian Islands and eastern north Pacific. In the eastern North Pacific, P. alalaua has been taken at Alijos Rocks and all of the Revillagigedo Islands (San Benedicto, Socorro, Roca Partida and Clarion). In the Revillagigedos, where sportfishermen catch fair numbers on oc- nXCH & CROOKE: REVISION OF EASTERN PACIHC CATALUFAS Figure 8. Otoliths (sag