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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 94118 


PRINTED IN THE UNITED STATES OF AMERICA 
BY ALLEN PRESS, INC., LAWRENCE, KANSAS 


Acanthocybium solanderi 301 
Acarina 14 
Acestrorhynchus 206 
Acridium 
coloratum 49-50 
xanthopterum 44, 51 
Ahlia 60 
egmontis 64 
Albula chinensis 210 
?Albula chinensis 210 
Alloeocarpa 239 
sp. 239 
Allopora 245 
Alopias 
superciliosus 89 
vulpinus 89 
Alopiidae 94-96 
Alytes obstetricans 217 
Amazonsprattus 317-321 
scintilla 317-321 
Ambassis 314 
Anchoviella 317, 321 
Andricus kollari 132 
Anthias macrophthalmus 302 
Apius 126 
figulus 126 
Aplidium californicum 244 
Aplochitonidae 203, 215 


Arctocephalus doriferus 229-230 


Argentinoidei 201 
Ascidiacea 239 
Atherinops 314 
Atractoscion nobilis 232 
Aulopodidae 201 


Balanus 
glandula 244 
nubilis 244-245 

Balitora 68 

Baltimora recta 49 

Barilius 151, 156-157 

Bdellostoma 
cirrhatum 264 
dombeyi 255 
Forsteri 264 
heptatrema 250, 264 

Benthenchelys 60 
cartieri 64 

Berycidae 311 

Beryx 311 


INDEX TO VOLUME 43 


(Compiled by Lillian J. Dempster) 


New names in boldface type 


Blakea 269-270, 281-282 


elliptica 270-271 
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 
Cestrum 


lorentziana 49 
parqui 49 
strigillatum 49 


Cetengraulis juruensis 317 
Cetorhinidae 94-96 
Cetorhinus 95-96, 109-110 


maximus 96, 232 


Chaenogobius sp. 181 
Chaetodon 311 


striatus 311 
substriatus 311 


Characoidei 201 

Chela 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 


[323] 


colorata 44-51, 54 

colorata group 45, 47, 51 
colorata-miles group 50 
icterus 44, 46-48, 50, 54-56 
miles 44-48, 51, 55 

minuta 43-47, 51 

nuptialis 44-48, 53 


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 44-45, 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 
Cladocera 317, 319 
Clavilithes macrospira 314 
Clidemia 269, 271, 274 
petiolaris 275 
saltuensis 274 
tetrapetala 269, 271-274 
trichosantha 269, 273-275 
Clupeidae 218, 317, 321 
Cnemidocarpa 247 
Coleoptera 159 
Colomesus 15 
Compositae 49 
Cookeolus 302, 304-305, 308, 312, 314 
boops 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 20-21 
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-abbotti series 247 
(Styelopsis) 239, 246-247 
(Styelopsis) abbotti 239-248 
uniplicata 246-247 

Diplomystes 154-155 


Diptera 317 
Drimys 277 


Eleotrididae 321 
Elopomorpha 218 
Elops 194, 217-218 
hawaiiensis 200, 218 
Engraulididae 317, 321 
Enhydra lutris 229, 231 
Eperlanus 204 
chinensis 204-205 
Epiactis prolifera 244 
Epigonus 314 
Epinephelus spp. 301 
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 
stoutii 250, 255, 262 
strahani 249-267 
undescribed 262 
Eremopodes sp. 41 
Esomus 148 
Eugomphodus 94 
tricuspidatus 94 
Eurystomella bilabiata 244 
Exoglossum 143 


Fugu 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 112-113, 115, 119 
becki 112-113, 115, 119 
chathamensis 112-113, 115, 119 
chilensis 119 
darwini 112-113, 115, 119 
elephantopus 112-113 
ephippium 111-113, 115-120 
galapagoensis 113 
guntheri 111-113, 115-120 
hesterna 119 
hoodensis 112-115, 119 
macrophyes 113 


INDEX 


microphyes 112-115, 119 
nigra 113 
nigrita 111-120 
phantastica 112-115, 119 
porteri 113-114 
sp. 114 
vandenburghi 112-115, 119 
vicina 112-116, 119 
vosmaeri \11 
wallacei 112-114, 119 
Ginglymostoma 109 
Glaucosoma 314 
Glenoglossa 59-66 
wassi 59-66 
Graffenrieda 269, 275 
bella 269, 275-277 
galeottii 275 
gracilis 275 
micrantha 275, 277 
Gryllus 
locusta miles 51 
speciosus 47, 51 
Gymnomuraena marmorata 22 
Gymnothorax 18, 20 
enigmaticus | 7-24 
leucacme 17, 22 
petelli 17, 20, 22 
““petelli’ form 20-21 
reticularis 20-21 
rueppelliae 17, 19-22 
rupelli 20 
ruppeli 17, 20 
“ruppelli”’ 21 
signifer 17, 21-22 
undulatus 22 
waialuae 17, 22 


Hemisalanginae 206, 210 
Hemisalanx 206, 210 
prognathus 185, 206, 208, 210 
Hemitrygon 287 
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 
signifer 290, 292, 294, 298 
Homaloptera 75 
Homea 
banksii 250, 264 
cirrhata 264 
Hoplias 206 
Hylaeus pectoralis 136 
Hymenoptera 27, 123 


325 


Hypolophus 284 
Hypomesus 194 
olidus 215 


Tlisha 317 

Tsistius 110 
brasiliensis 110 

Tsurus spp. 233 


Jenkinsia 317 


Lactarius 314 
Lamnidae 94-96 
Lamniformes 89, 93 
Leguminosae 49 
Lepidogalaxias 194-195, 214-217 
salamandroides 215-216 
Leptasterias pusilla 244 
Leucosoma 206, 210 
chinensis 205, 210 
reevesi 206, 210 
Leuresthes 314 
Lipara lucens 136 
Lovettia 215 
Luciobrama 152-153 
Luciosoma 143 
Lycium 49 
cestroides 49 
Lycodontis 20 


Macrochirichthys 144, 151 
Macrocystis pyrifera 232 
Mallotus 215 
Medicago sativa 49 
Megachasma 87-110 

pelagios 87-110 
Megachasmidae 87-110 
Melastomataceae 269-282 
Metasalanx 206 

coreanus 206, 210 
Miconia 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 110 
Mixodigmatidae 110 
Monotreta 2, 15 
Muraena 

interrupta 17, 22 

umbrofasciata 17, 21 

xanthopterus 22 


326 


Muraenichthys 60, 62 
Muraenidae 18 
Mycteroperca spp. 301 
Myctophidae 200 
Myliobatis californica 230-232 
Myrophinae 59-60 
Myrophis 60 
Jrio 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 167 
fragilis teewinot 159, 167 


gebleri 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 sevieri 159, 164-165, 168, 170, 172, 174, 


7/7 
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 164 
trifaria coloradensis 165 


trifaria pasquineli 159, 165-166, 170, 172, 175, 


9/7) 
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 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43 


parvipectoralis 64 
spp. 64 
Nemamyxine 251-254 
Neomysis sp. 181 
Neomyxine 251, 254 
Neosalanx 182, 184, 186, 198, 201-205, 210-212, 214, 
219 
andersoni 179, 181-182, 184, 186, 209, 211-212, 
219 
brevirostris 179, 181, 186-187, 209, 211-212, 219 
hubbsi 212 
Jordani 179-181, 183-186, 190-192, 195, 197, 199, 
201-202, 204, 209-212, 219 
regani 212 
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 

(Pagellus?) gregarius 313 

(Sparidarum) gregarius 313 

(Sparidarum) rutoti 313 
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 


longianalis 206-208 
(Salanx) 202 
?Parasalanx gracillimus 207 
Parexoglossum 143 
Pellona 317 
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 
meeki 301, 310 
tayenus 301, 310 
Prionace glauca 228 
Pristigaster cayana 317 
Pristigenys 301, 304, 311-314 
bella 301, 310, 314 
caduca 310, 313 
dentifer 301, 310, 314 
macrophthalmus 311 
rutoti 310, 312-313 
spectabilis 310, 314 
spp. 314 
substriata 301, 304, 310-312, 314 
substriatus 311 
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 212 
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 211 
?Protosalanx tangkahkeii 212 
Prototroctes 203 
Prototroctidae 201 
Pseudocarcharias 105 
kamoharai 94 
Pseudocarchariidae 94, 96 


B27] 


Pseudomyrophis 59-60, 64-65 
atlanticus 65 
micropinna 65 
nimius 65 
spp. 64-65 
Pseudopriacanthidae 313 
Pseudopriacanthus 302, 304, 311-313 
altus 312 
niphonius 312-313 
serrula 301-305, 307, 309, 312 
Psilorhynchidae 67 
Psilorhynchus 67-70, 73, 75 
balitora 67-76 
balitora-like 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 73-74 
variegatus 68, 74 


Quercus spp. 277 


Raja pastinaca 286 
Rasbora 151 
Reganisalanx 206 
normani 206-208, 210 
?Reganisalanx brachyrostralis 210 
Retropinnidae 201, 203, 215 
Rhiniodon 109-110 
typus 89, 96 
Rhinosardinia amazonica 317 
Rhomalea 
icterus 55 
latipennis 44, 53 
miles 52 
miles Var. C 52 
nuptialis 53 
opulenta 44, 55-56 
pedes 44, 49-50 
peruviana 56 
speciosa 52 
stolli 44, 51 
trogon 53 
Romalea psittacus 54 
Romaleidae 43 
Romaleinae 43, 47 
Rubus 136 


Sagraea 274 
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 213 
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 211 
ariakensis 181, 185-186, 206-208, 210, 213 
brachyrostralis 210 
chinensis 181, 187, 204—205, 210, 216 
cuviert 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 
reevesil 210 
(Salanx) 186, 202, 207, 210 
(Salanx) ariakensis 179, 206 
(Salanx) cuvieri 179, 207 
?2Salanx 
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 301 
Sideria chlevastes 17, 22 
Siluriformes 201 
Solanaceae 49 
Solanum 49 
argentinum 49 
elaeagnifolium 49 
verbascifolium 49 
Somniosus 
microcephalus 89, 105 
pacificus 89, 105 
(Somniosus) 105 
Sparidarum rutoti 312 
Sphecidae 27, 123 


Sphex 
figulus 125-126 
fuliginosa 126 
fuliginosus 123, 126-127 
Sphyrna mokarran 89 
Spirinchus 194 
Spirobis 244 
Squalus 228 
acanthias 231-232 
Stereolepis gigas 301 
Styela 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 27-42 
acanthophorus 27, 29, 39-41 
alayoi 29 
apricus 27, 29-31, 34 
arizonac 27, 29-30, 32, 35 
armatus 27, 29, 40-41 
ashmeadii 31 
belfragei 36, 41 
bohartorum 27, 29-31 
brevicornis 29 
brulli group 28-29, 36, 39-40 
crenulatus 35-36 
fulvitarsis 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 41 
menkei 27, 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 30-31 
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 


INDEX 


tarsatus 34 
terminatus 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, 15 
modestus 2 
Tetraddon 
(Arothron) modestus 7-8 
modestus 2 
naritus 2 
Tetraodontidae 1, 14-15 
Tetraodontiformes 14 
Thryssocypris 141-158 
smaragdinus 141-158 
tonlesapensis 141-158 
Thunnus 
albacares 301 
obesus 301 
Thysanopoda pectinata 108 
Topobea 269-270, 281-282 
brenesii 270 
calophylla 269, 280-282 
durandiana 282 
elliptica 269-270, 281 
pittieri 281 
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 126 
figulus var. major 136 
figulus var. majus 126 
figulus media 136 
figulus forma media 136 
figulus var. media 136 
figulus medium 132, 136 
figulus var. medium 136 
figulus minor 132 
figulus forma minor 132 
figulus var. minor 132 
figulus minus 132, 136 
figulus var. minus 132 
figulus minus var. rubi 136 
figulus yezo 123, 127 
majus 123, 127 

medium 123-140 

minus 123-140 

rubi 123, 139 


Umbridae 217 
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 


329 


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CONTENTS OF VOLUME 43 


RosertTSs, TYSON R. The Southeast Asian freshwater pufferfish genus Chone- 
rhinos (Tetraodontidae), with descriptions of new species. Published June 15, 


McCoskeER, JOHN E., AND JOHN E. RANDALL. Synonymies of Indian Ocean eels, 
with the description of Gymnothorax enigmaticus, a moray previously known 
asaonnel. Published june sl 5:01 982 2 8 6 cs ne 


PuLAwskKI, WoyjciecH J. New species of North American Tachysphex wasps 
(igmenoptera., Sphecidae).. Published June, 15, 1982.2 


Roserts, H. RADCLYFFE, AND CARLOS S. CARBONELL. A revision of the grass- 
hopper genera Chromacris and Xestotrachelus (Orthoptera, Romaleidae, Roma- 
leinae)y BublishedtNovember 4.1982... ns 


McCoskeEr, JOHN E. A new genus and two new species of remarkable Pacific 
worm eels (Ophichthidae, subfamily Myrophinae). Published November 4, 


RAINBOTH, WALTER J. Psilorhynchus gracilis, a new cyprinoid fish from the 
Ganeeric lowlands: Published July 65 1983-2 eee 


Le Boeur, 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. 


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 


CRUMLY, CHARLES R. The cranial morphometry of Galapagos tortoises. Pub- 
lished January 17, 1984 


PuLAwskKI, WojciecH J. The status of Trypoxylon figulus (Linnaeus, 1758), 
medium De Beaumont, 1945, and minus De Beaumont, 1945 (Hymenoptera: 
Salccidac)sublished January 17.1984. 08 ee ee 


ROBERTS, TYSON R., AND Maurice Kortte.atT. Description and osteology of 
Thryssocypris, a new genus of anchovylike cyprinid fishes, based on two new 
species from Southeast Asia. Published January 17, 1984 00 


KAVANAUGH, Davin H. Studies on Nebriini (Coleoptera: Carabidae), V. New 
Nearctic Nebria taxa and changes in nomenclature. Published July 12, 1984... 


ROBERTS, TYSON R. Skeletal anatomy and classification of the neotenic Asian 
salmoniform superfamily Salangoidea (icefishes or noodlefishes). Published July 


TrICcAS, TIMOTHY C., AND JOHN E. McCosker. Predatory behavior of the white 
shark (Carcharodon carcharias), with notes on its biology. Published July 12, 


NewBerryY, ANDREW Topp. Dendrodoa (Styelopsis) abbotti, sp. nov. (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, 


17-24 


27-42 


43-58 


59-66 


67-76 


77-85 


87-110 


111-121 


123-140 


141-158 


159-177 


179-220 


221-238 


239-248 


No. 20. 


McMILLAN, CHARMION B., AND RoBerT L. WISNER. Three new species of seven- 
gilled hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean. Published De- 
Cemiber Nil; 1984 xe eo at ee ee 


ALMEDA, FRANK. New and noteworthy additions to the Melastomataceae of 
Panama..,Published December jis 1984. ee eee 


CoMPAGNO, LEONARD J. V., AND Tyson R. Roserts. Marine and freshwater 
stingrays (Dasyatidae) of West Africa, with description of a new species. Pub- 
lished: December UU 9840925 OSL Bee SOOO 2) ER eee, eee 


Fitcu, JOHN E., AND STEPHEN J. CROOKE. Revision of Eastern Pacific catalufas 
(Pisces: Priacanthidae) with description of a new genus and discussion of the 
fossilosrecord.: Published December, 1 1984.22 ee ee 


RoBERTS, TYSON R. Amazonsprattus scintilla, new genus and species from the 
Rio Negro, Brazil, the smallest known clupeomorph fish. Published December 


IndexstoeVolume 43h. 2s Tie Ore’ A A OE ee ee 


Instructions: tov AUTHORS)... <0 ee ee 


liv] 


Pages 


249-267 


269-282 


283-300 


301-315 


317-321 


323-329 
331-332 


siti al i OT AY OS IS Fae i aa i nts 


PROCEEDINGS 
OF THE 


CALIFORNIA ACADEMY OF SCIENGE! 


Vol. 43, No. 1, pp. 1-16, 10 figs. 


JU 


r:rorx ny 
ee ’ * 


4 June 15, 1982 


Woo0s Hole Mass. | 
THE SOUTHEAST ASIAN- FRESHWATER PUFF ERFISH 
GENUS CHONERHINOS (TETRAODONTIDAE), WITH 
DESCRIPTIONS OF NEW SPECIES 


By 


Tyson R. Roberts 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


ABSTRACT: The tetraodontid pufferfish genus Chonerhinos, restricted to fresh water in Southeast Asia, 
comprises five species, four of which are described as new. The species differ in adult size, coloration, orientation 
of squamation, depth of caudal peduncle, size of nasal organ, food habits, and geographical distribution. The 
most widely distributed, C. nefastus n.sp., occurs in southern, western, and northern Borneo, the Malay 
Peninsula, Thailand, Vietnam, Kampuchea, and Laos; it feeds mainly on fish fin rays and scales, and has a 
slender caudal peduncle and the smallest nasal organ. Chonerhinos modestus (Bleeker, 1850), in western Borneo 
and Sumatra, with perhaps the most varied diet, is the largest species and has the deepest caudal peduncle. 
The distinctively colored C. amabilis n.sp., with the largest nasal organ, occurs in western Borneo and Sumatra 
and feeds almost exclusively on large aquatic insects. The two new species C. silus, with a moderately deep 
caudal peduncle, and C. remotus, with a slender caudal peduncle, have varied diets including insects, and are 


known only from northern and northeastern Borneo. 


INTRODUCTION 


The freshwater pufferfish genus Chonerhinos 
currently includes a single species, C. modestus 
(Bleeker, 1850), reported from localities 
throughout much of Southeast Asia. The nomi- 
nal species C. africanus Boulenger, 1909, 
known only from the holotype supposedly col- 
lected in the interior of the Congo basin, has 
been identified as a junior synonym of C. mo- 
destus with incorrect locality data (Roberts 
1981; herein). The species formerly known as C. 
naritus (Richardson, 1848), from marine, brack- 
ish, and perhaps freshwater habitats along the 
coasts of the South China Sea and eastern Indian 
Ocean, has been placed in a monotypic genus, 
Xenopterus (Fraser-Bruner 1943; Tyler 1980; 
herein). 

I undertook this revision because three 
species of Chonerhinos were obtained during 


[1] 


my ichthyological survey of the Kapuas basin in 
western Borneo (Kalimantan Barat, Indonesia) 
in 1976. 


MATERIAL EXAMINED AND METHODS 


More than 250 specimens of Chonerhinos 
from throughout the range of the genus were 
examined during this study. These are deposited 
in the British Museum (Natural History), Lon- 
don, BMNH: California Academy of Sciences, 
San Francisco (CAS), including material for- 
merly deposited at Stanford University, Stan- 
ford (SU); Field Museum of Natural History, 
Chicago (FMNH); Museum Geneve, Geneva 
(MG); Muséum National d’Histoire Naturelle, 
Paris (MNHN); Museum Zoologicum Bogo- 
rense, Bogor, Indonesia (MZB); Musée Royal 
de l'Afrique Centrale, Tervuren (MRAC); Nat- 
ural History Museum, Basel (NHMB); Riks- 


io) 


museum van Natuurlijke Historie, Leiden 
(RMNH); University of Michigan Museum of 
Zoology, Ann Arbor (UMMZ); U.S. National 
Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. (USNM); and Zoolog- 
ical Museum, Universiteit van Amsterdam, Am- 
sterdam (ZMA). 

Length of specimens is given as standard 
length unless total length is expressly indicated, 
and all proportional measurements are given as 
times in standard length (SL). Vertebral counts 
were made from radiographs prepared in the 
Department of Ichthyology, CAS. Sections of 
skin anterior and ventral to the pectoral fin were 
removed with a scalpel and stained in alizarin to 
facilitate illustration of the scales. Orientation of 
the scales is also obvious in radiographs and can 
be observed in whole specimens without special 
preparation. 


Chonerhinos Bleeker 


Chonerhinos BLEEKER, 1854:259-260 (type-species Tetraodon 
modestus Bleeker, 1850, by subsequent designation of Fra- 
ser-Bruner 1943:16). 

Chonerhinus BLEEKER, 1865:213 (unjustified spelling change). 


DESIGNATION OF TYPE-SPECIES.—Fraser- 
Bruner (1943) is apparently the first author to 
have properly designated a type-species for 
Chonerhinos. The original description of the ge- 
nus is as follows: ‘‘Chonerhinos Blkr [is 
gekenmerkt] door trechtervormige verdieping 
ter plaatse der neusopeningen met verhevene 
randen, lange rug- en aarsvinnen, zigtbare zijlijn 
en onegekielden rug .. . van Chonerhinos 2 t. 
w. Chonerhinos modestus Blkr = Tetraodon 
modestus Blkr olim (van Borneo, Sumatra), 
Chonerhinos naritus Blkr = Tetraodon naritus 
Richds (van Borneo).’’ Thus, Bleeker included 
two species in his original account of Chone- 
rhinos and did not indicate a type-species. Hol- 
lard (1857) defined Xenopterus (type-species X. 
bellengeri = X. naritus, by monotypy) in such 
a way that it excludes Chonerhinos, which, 
however, he did not mention by name. Gill 
(1892) discussed the nomenclatural history of 
Chonerhinos (and Xenopterus) at length but 
oddly did not mention the lack of a type-species. 
Jordan (1919:256) incorrectly stated that Tetrao- 
don modestus Bleeker is the ‘‘orthotype’’ of 
Chonerhinos, meaning that Bleeker (1854) indi- 
cated or distinctly implied that this species is the 
type-species. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


D1aGNosis.—Chonerhinos and its close rela- 
tive Xenopterus differ from all other tetraodon- 
tids in having three lateral line canals on side of 
body instead of one, two, or none; dorsal fin 
with 22 or more rays; anal fin with 18 or more 
rays; at least 24 vertebrae; and prefrontal bones 
absent (Tyler, 1980). Chonerhinos differs from 
Xenopterus in its smaller adult size, less exten- 
sive squamation, less exposed olfactory lamel- 
lae, and fewer fin rays and vertebrae. The largest 
Chonerhinos | have examined is 106 mm; Xen- 
opterus attains at least twice this size. In Cho- 
nerhinos the scales are relatively small and re- 
stricted to the head and body ventral to the level 
of the pectoral fin; in Xenopterus the scales are 
relatively large and extend dorsally to the pec- 
toral fin. In Chonerhinos the olfactory lamellae 
are largely covered by nasal flaps in broad con- 
tact; in Xenopterus the nasal flaps are greatly 
reduced and the olfactory lamellae are conse- 
quently almost entirely exposed. Chonerhinos 
has 22-28 dorsal-fin rays, 18-22 anal-fin rays, 
13-17 pectoral-fin rays, and 24-28 vertebrae; the 
same counts in Xenopterus are 32-38, 28-29, 
18-19, and 29-30. 

REMARKS.— Tyler (1980) stated that Chone- 
rhinos and Xenopterus are highly specialized 
tetraodontids which have secondarily increased 
the number of dorsal- and anal-fin rays and ver- 
tebrae, elaborated the lateral line system, in- 
creased the number and size of the olfactory la- 
mellae, and increased the size of at least some 
of the scales; and that the greater numbers of 
vertebrae and fin rays in Xenopterus as well as 
the structure of the skull indicates that it is the 
more specialized of the two. In Chonerhinos, 
according to Tyler, apart from the absence of 
the prefrontal bones, the skull is not markedly 
different from that in many species of the tet- 
raodontid genera Monotreta, Chelonodon, and 
Tetraodon, whereas in Xenopterus the frontals 
are much more laterally expanded and thickened 
than in Chonerhinos, forming a large plate over 
most of the dorsal surface of the skull, and the 
supraoccipital crest is wider and heavier; in 
large specimens the two frontals may become 
indistinguishably fused to each other in the mid- 
dle of their lengths (Tyler 1980:340, fig. 274). I 
have examined two X. naritus from Sarawak, 
BMNH 1894.1.19.86-87, 71.2 and 108 mm. Ra- 
diographs reveal that the frontal bones, supra- 
occipital crest, supraneural bone, anteriormost 


ROBERTS: FRESHWATER PUFFERFISH 


fate 


FIGURE |. 


: Pf = 


ata 


[esas 


(hv 


isi 


Scales on side of body immediately anterior and ventral to pectoral fin (each square = 5 x 5 mm): (a) Xenopterus 


naritus, 71.2 mm, BMNH 1894.1.19.86; (b) Chonerhinos modestus, 48.4 mm, USNM uncatalogued; (c) Chonerhinos silus, 48.5 
mm, FMNH 68815; (d) Chonerhinos remotus, 49.9 mm, FMNH 68475; (e) Chonerhinos nefastus, 48.3 mm, CAS 49507; (f) 


Chonerhinos amabilis, 48.7 mm, MZB 3973. 


anal-fin pterygiophore, and posteriormost neural 
and haemal spines are enormously thickened or 
hypertrophied, far out of proportion to neigh- 
boring bony elements. They appear to be hy- 
perosteotic (and in the case of the frontal bones, 
partially synosteotic), and therefore, | am du- 
bious about their phylogenetic significance and 
their being used as characters to distinguish 
Xenopterus from Chonerhinos. Other differ- 
ences between the two genera, cited above and 
in Tyler (1980), are sufficient to merit their sep- 
aration. 

Chonerhinos is known only from fresh water. 
Xenopterus, so far as I have been able to deter- 
mine, is marine or estuarine. There do not seem 
to be any museum specimens of Xenopterus 
with locality data from fresh water, and state- 
ments in the literature that Xenopterus occurs 
in fresh water (e.g., Cantor 1850:384; Weber and 
de Beaufort 1962:373) appear to be based at least 
partly on misinformation or confusion with Cho- 
nerhinos. 

In Chonerhinos and Xenopterus, as in many 
other tetraodontids, each scale has a spinelike 


distal portion which projects more or less 
straight out from the skin when erected, as usu- 
ally occurs when the fish inflates itself. When 
the scales are not erect, they are partially or 
wholly retracted beneath the skin, and the 
spines may be oriented dorsally, dorsoposte- 
riorly, or posteriorly, depending upon the 
species (Fig. 1). 

Size and shape of the jaw-teeth appear to be 
nearly identical in all species of Chonerhinos. 
One or two specimens of each species were dis- 
sected to permit observation of the gill rakers; 
all of the species have about 8-10 total gill rakers 
on each gill arch (sometimes fewer on the first 
arch). I have not attempted to distinguish the 
species by differences in the pathways of the 
lateral line canals. These are difficult to observe 
in many specimens, and they seem to be highly 
variable among individual specimens, often 
being irregularly interrupted or running into 
each other (Tyler 1980:fig. 223) and frequently 
differing in their courses on opposite sides of a 
specimen. Neither have I attempted to distin- 
guish the species by counts of olfactory lamel- 


4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


TABLE |. PROPORTIONAL MEASUREMENTS IN Chonerhinos (expressed as times in standard length). 

C. amabilis C. modestus C. nefastus C. remotus C. silus 
n 20 13 54 31 33 
SL mm 35.6-70.4 46.8-106 19.0-70.7 32.8-61.4 32.7-81.8 
Eye 7.8-11.2 9.4-14.1 7.2-11.7 8.6-12.0 8.8-12.9 
Nasal organ length 10.1-17.9 14.7-25.3 17.3-27.4 12.5—20.1 11.2-20.0 
Snout length 6.0-7.2 6.47.5 5.8-7.1 6.6-8.7 6.3-8.1 
Interorbital width 5.1-6.1 4.6-6.8 4.5-7.0 5.2-7.0 4.9-6.6 
Pectoral-fin base length 10.5—12.2 9.5-11.5 9.9-13.5 8.9-11.1 9.1-12.4 
Caudal peduncle depth 7.3-8.3 6.7-7.4 7.8-9.9 7.6-9.9 7.2-8.2 
Caudal peduncle length 5.0-6.5 5.2-6.4 4.45.9 4.6-6.3 4.6-6.5 


lae, the number of which seems to be highly 
variable within each species, as is the size of the 
nasal organ itself (Table 1). 


PROPORTIONAL MEASUREMENTS; 
MERISTIC FEATURES 


Proportional measurements, in most instances 
broadly overlapping and of little help in distin- 
guishing species, are presented in Table |. Fre- 
quencies of counts of fin rays and vertebrae, 
diagnostic for the genus but differing slightly 
among species and of little or no help in identi- 
fying individual specimens, are presented in Ta- 
bles 2-3. Except in a few instances when counts 
or measurements are particularly useful for def- 
inition of species, these data are not repeated in 
the text. 


KEY TO SPECIES OF Chonerhinos 


la. Scales on side of body anterior and ven- 
tral to pectoral fin with spines directed 
postentonly (Bigs lie—f) eee 2 

lb. Scales on side of body anterior and ven- 
tral to pectoral fin with spines directed 
dorsally or dorsoposteriorly (Fig. 1b-d) 


2a. 


3a. 


3b. 


A roundish dark spot in middle of caudal 
peduncle; dorsal and anal fins always with 
angulated margins; upper lip not project- 
ing beyond lower lip; exposed portion of 
eye round; nasal organ relatively large, its 
length 10.1—17.9 (times in SL) 


C. amabilis 


. No spot on caudal peduncle; dorsal and 


anal fins usually with rounded margins; 
upper lip usually projecting beyond lower 
lip; exposed portion of eye usually hori- 
zontally oval, especially in larger speci- 
mens; nasal organ relatively small, its 
length 17.3—27.4 C. nefastus 


Depth of caudal peduncle 6.7—7.4; upper 
and lower lips about equally projecting or 
lower lip slightly protruding; snout gently 
sloping; scales on side of body anterior 
and ventral to pectoral fin, very close-set 
with spines directed dorsally (Fig. 1b); 
anal-fin rays 20-22, modally 22 (Table 2); 
adult size to 106 mm 


Depth of caudal peduncle 7.2-9.9; lower 
lip usually projecting beyond upper lip; 
snout strongly sloping; scales on side of 


FREQUENCIES OF FIN RAY Counts IN Chonerhinos. 


TABLE 2. 

Dorsal fin 
22 23 24 25 26 27 28 
C. amabilis  — - | 12 6 1 - 
C. modestus — - - 4 12 8 ] 
C. nefastus - 3 23 16 8 4 = 
C. remotus 5 30 40 8 2 = = 
C. silus - 1 8 18 27 3 ] 


Anal fin Pectoral fin 
20 21 22 13 14 15 16 17 
a 14 6 = - 3 16 1 - 
= 1 7 il7/ = 1 9 14 l 
| 19 31 3 5 28 20 l = 
37 3 = 9 52 = 
2 19 34 3 = 2 36 1 1 


ROBERTS: FRESHWATER PUFFERFISH 


tn 


FIGURE 2. Chonerhinos amabilis, 45.2 mm, MZB 3972 (holotype). 


body anterior and ventral to pectoral fin Chonerhinos amabilis new species 
not as close-set and with spines directed (Figure 2) 
dorsoposteriorly (Fig. 1c-d); anal-fin rays Chonerhinus naritus WEBER AND DE BEAUFORT, 1962:374 
18-22, rarely 22, modally 19 or 20 (Table (specimens reported from ‘‘Labang hara, soengei Serawai’’). 
2); adult size to 82 mm 4  Chonerhinus modestus WEBER AND DE BEAUFORT, 1962:fig. 
a5. pe 84. 

4a. Caudal peduncle moderately deep, its HoLotyPe.—MZB 3972, 45.2 mm, Kapuas R. 6 km w of 
depth 7.2-8.2; dorsal-fin rays 23-28, av- Putussibau, Kapuas Ichthyological Survey, 9 Aug. 1976. 
Snide De Sr ere C. silus PARATYPES.—CAS 49504, 45.0 mm, same data as holotype; 


. MZB 3973, 48.7 5 Ix in, i Landok at Nga- 
4b. Caudal peduncle slender, its depth 7.69.9; minty Kapuas Dasin, SungarLandok ab Nes 
Bana bang, 83 km ENE of Pontianak, Kapuas Ichthyological Survey, 


dorsal-fin rays 22-26, average 23.6 ______- 15 July 1976; MZB 3974, 41.8 mm, Kapuas basin, Sungai Pi- 
2 ee a A ee es 5 C. remotus noh 20-60 km upstream from Nangapinoh, Kapuas Ichthyo- 


TABLE 3. FREQUENCIES OF VERTEBRAL COUNTS IN Chonerhinos. 


C. amabilis C. modestus C. nefastus C. remotus C. silus 


9+ 15 = 24 (1) 


9? + 16 = 25? (1) 9+ 16 = 25 (2) 9+ 16 = 25 (1) 9? + 16 = 25? (1) 
10 + 15 = 25 (3) 10 + 15 = 25 (2) 

9+ 17 = 26 (1) 9 + 17 = 26 (1) 
10 + 16 = 26 (9) 10 + 16 = 26 (3) 10 + 16 = 26 (1) 10 + 16 = 26 (8) 10 + 16 = 26 (8) 
10 + 17 = 27 (6) 10 + 17 = 27 (1) 


11? + 16 = 27? (1) 11? + 16 = 27? (2) 11? + 16 = 27? (1) 


6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


rap as 


2 hen Rid 


FIGURE 3. 
unknown; (b) 78.7 mm, RMNH 26931 (neotype). 


logical Survey, 22-26 July 1976; MZB 3975, 38.3 mm, Kapuas 
R. near Kampong Nibung, 7 km NE of Selimbau, Kapuas Ich- 
thyological Survey, 5-6 July 1976; MZB 3976 and USNM 
230359, 2:35.9-36.8 mm, Kapuas R. 53 km w of Putussibau, 
Kapuas Ichthyological Survey, 6-7 Aug. 1976; MZB 3977 
and FMNH 94255, 2:35.6-46.0 mm, Kapuas R. about 23 km 
wsw of Putussibau, Kapuas Ichthyological Survey, 8-9 Aug. 
1976; MNHN 91.216, 36.9 mm, Kapuas basin, M. Chaper, 
1890; RMNH uncat., 2:40.9-41.2 mm, Kapuas basin, Sintang, 
July 1894; RMNH 7935, 4:55.5—68.1 mm, Kapuas basin, Raun, 
Mar.—May 1894; ZMA 108.912, 3:56.3-70.4 mm, Kapuas ba- 
sin, Soengai Serawai, Lebang Hara, Witkamp, no date; 
UMMZ 171708, 2:36.2-38.3 mm, Sumatra, Moesi R. at Moera 
Klingi, A. Thienemann, 1913. 


D1AGNosis.—Chonerhinos amabilis is readily 
distinguished from all other members of the ge- 


Chonerhinos modestus: (a) as illustrated in Bleeker 1865; length, locality, and present disposition of specimen 


nus by its highly distinctive coloration, almost 
all elements of which are visible in all specimens 
examined, including some century-old speci- 
mens which may have been dead for some time 
before being preserved. These unique features 
include a roundish dark spot in middle of caudal 
peduncle, visible in all specimens; a large, dis- 
tinctively shaped dark mark on dorsal surface of 
head extending uninterrupted from just behind 
upper lip to well behind the eyes, set off by pale 
coloration on the upper lip, sides of snout, nasal 
flaps, and skin dorsal to orbits; pale white or 
milky coloration on ventral and lateral surfaces 
of body extending very far dorsally; dark col- 


ROBERTS: FRESHWATER PUFFERFISH 


FIGURE 4. 


oration on dorsal surface of body markedly en- 
hanced around base of dorsal fin; and a small 
dark or dusky oval spot with indistinct margins 
near tip of chin (very faint or absent in some 
specimens). In addition, C. amabilis tends to 
have the largest nasal organ of any Chonerhi- 
nos, and thus of any tetraodontid (Tyler 
1980:290); relatively large dorsal and anal fins 
with angulated (rather than rounded) margins; 
and scales on side of body anterior and ventral 
to pectoral fin relatively small, few in number, 
and with spines directed posteriorly (Fig. 1f). 
ETYMOLoGy.—Latin amabilis, lovely. 


Chonerhinos modestus (Bleeker) 
(Figures 3-5) 


Tetraddon (Arothron) modestus BLEEKER, 1850:16 (type-lo- 
cality ‘‘Banjermassing, in fluviis’’). 

Chonerhinos modestus BLEEKER, 1854:260. 

Chonerhinus africanus BOULENGER, 1909:201 (type-locality 
‘“‘riv. Sankuru, a Kondué Kasai, Congo’’). 


NEOTYPE.—RMNH 26931, 78.7 mm, Kapuas basin, Sang- 


gau, Westenenk, 1894. 
ADDITIONAL MATERIAL EXAMINED.—RMNH uncat., 


—! 


Chonerhinos modestus, 64.6 mm, CAS 49505. 


2:49.2-59.2 mm, same data as neotype; RMNH 7934, 
3:50.0-58.9 mm, Kapuas basin, Sintang, July 1894; CAS 49505 
and MZB 3978, 2:64.6-106 mm, Kapuas R. about 23 km wsw 
of Putussibau, Kapuas Ichthyological Survey, 8-9 Aug. 1976; 
MZB 3979 and USNM 230360, 2:46.8-48.4 mm, Kapuas 
R. at Silat, Kapuas Ichthyological Survey, 17 Aug. 1976; 
BMNH 1846.6.22.75, 86.1 mm, Borneo, Frank Collection, no 
date; BMNH 1867.11.28.125, 87.3 mm, Borneo, Bleeker Col- 
lection, no date; RMNH 12004, 3:66.6-81.1 mm, Sumatra, 
Lahat, Bleeker Collection, 1850-60; NHMB 822-824, 
3:44.7-73.5 mm, Sumatra, Indragiri, H. A. von Meckel, 1895; 
RMNH 7344 (part only), 8:47.9-62.0 mm, no locality data, 
Bleeker Collection, no date; MRAC 15306, 52.5 mm, ‘*‘Congo, 
Sankuru River, Kasai’’ (holotype of C. africanus). 


SELECTION OF NEoTyYPE.—Identification of 
C. modestus presented a difficult and taxonom- 
ically important problem which I have resolved 
by selecting a neotype. The holotype is lost or 
at least it cannot be positively identified, and the 
original description fits all five species of Cho- 
nerhinos about equally well. In order to facilitate 
the following discussion the original description 
(Bleeker 1850:16) is reproduced here in its en- 
tirety: 


FIGURE 5. 


Tetraddon (Arothron) modestus Blkr. 

Tetradd. corpore oblongo compresso, altitudine 4 circiter 
in ejus longitudine, latitudine 2 in altitudine; vertice, dorso, 
lateribus caudaque laevibus, pectore genisque scabris; ca- 
pite obtuso; lineo rostro-dorsali convexa; maxilla superiore 
paulo prominente; oculis paulo superis; tentaculis nasalibus 
2 conicis obtusis loco narium; linea laterali inconspicua; 
saccO pneumatico parvo; ano ante pinnam dorsalem sito; 
pinnis dorsali et anali obtusis angulatis angulis rotundatis, 
pectoralibus emarginatis, caudali truncata vel leviter emar- 
ginata 5 in longitudine corporis; colore corpore supra viridi 
infra argenteo, pinnis hyalino-viridescente. 

D. 5/20. P. 2/12. A. 3/20. C. 9 vel 11 et lat. brev. 

Habit. Banjermassing, in fluviis. 

Longitudo speciminis unici 60°”. 


Bleeker almost invariably recorded the length of 
his specimens as total length in millimeters 
(pers. commun. M. Boeseman, RMNH). Thus, 
the last two lines of the description indicate that 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


Chonerhinos modestus: (a) 52.5 mm, MRAC 15306 (holotype of C. africanus); (b) 48.4 mm, USNM 230360. 


it was based on a single specimen, the holotype, 
total length 60 mm, from riverine habitat at 
Bandjarmasin, southeastern Borneo (Barito ba- 
sin). Bleeker obtained in all 59 specimens which 
he identified as C. modestus, total lengths 
46-126 mm, from Palembang (=Lahat?), Su- 
matra, and Sambas, Pontianak, and Sintang as 
well as Bandjarmasin in Borneo (Bleeker 
1865:78). All extant ‘‘C. modestus’’ from Bleek- 
er’s collection apparently are deposited in the 
BMNH, RMNH, and ZMA. The BMNH and 
ZMA each have a single Bleeker specimen, both 
of which are too large to be the holotype: 
BMNH 1867.11.28.125, 87.3 mm, Borneo, ex- 
amined by me, and ZMA 102.263, 104 mm, Bor- 
neo, examined for me by H. Nijssen. The 
RMNH has two lots, RMNH 12004, 3:66.6-81.1 


ROBERTS: FRESHWATER PUFFERFISH 


mm, Sumatra, Lahat (=Palembang?), and 
RMNH 7344, 52:29.0-74.0 mm (total lengths 
37-88 mm), without locality data. If the holotype 
still exists, it presumably is in RMNH 7344. 
Among the 52 specimens are 4 which approxi- 
mate 60 mm in total length; thus, on the basis 
of length alone, the holotype cannot be identi- 
fied. Moreover, each of the four specimens dif- 
fers by one or two fin rays in at least two of the 
three counts reported by Bleeker for the dorsal, 
anal, and pectoral fins of the holotype. In my 
opinion, none of these specimens can reason- 
ably be identified as the holotype, and since their 
locality data are lost, a neotype should not be 
selected from among them. Unfortunately, I 
have been unable to find any specimen of Cho- 
nerhinos with locality data from Bandjarmasin 
or the Barito and do not know which of the 
species occur(s) there. 

As noted above, the original description of C. 
modestus fits all five species of Chonerhinos 
about equally well. All species of Chonerhinos 
normally have 11 caudal-fin rays, and all species 
are represented by specimens with 25 dorsal-fin 
rays and 14 pectoral-fin rays. On the other hand, 
none of the more than 250 specimens examined 
have 23 anal-fin rays. The highest number of 
anal-fin rays observed, 22, is usually found in 
the species herein identified as C. modestus, but 
also occurs in C. nefastus and C. silus. Color- 
ation and its variation in the species of Chone- 
rhinos are too poorly known at present to be of 
much help in their identification, and Bleeker’s 
description of coloration of the holotype cannot 
be accepted without reservation since he did not 
collect the specimen himself and could not have 
observed it until it had been in preservative for 
many days or weeks. Bleeker (1865:pl. 213, fig 
8) published an excellent figure of a specimen 
which he identified as C. modestus. The length, 
locality, and date of collection of the specimen 
figured are not recorded, but it is not the holo- 
type. It is evidently a much larger specimen, 
with lateral line canals on the body plainly vis- 
ible, and differs also in fin-ray counts from the 
holotype as described by Bleeker. I have not 
tried to match up the figure with an extant spec- 
imen, although it may well be part of RMNH 
7344. The figure does, however, show a number 
of features characteristic of the largest species 
of Chonerhinos, with which I unhesitatingly 
identify it. These features include its large size 


(indicated by the large size of the published il- 
lustration as well as by the relatively small eye); 
scales with dorsally oriented spines; relatively 
high counts of dorsal- and anal-fin rays; and 
deep caudal peduncle. All four specimens of to- 
tal length 60 mm in RMNH 7344 also belong to 
this species. Thus, there is every reason to identi- 
fy it as C. modestus, although we cannot be sure 
that this is the same species obtained for Bleeker 
at Bandjarmasin. In the absence of specimens 
with locality data from Bandjarmasin or the Ba- 
rito, a specimen from the Kapuas basin has been 
selected as neotype. This specimen bears a 
strong resemblance to Bleeker’s figure of C. 
modestus (Figs. 3a—-b). 

DIAGNOSIS.—Chonerhinos modestus, attain- 
ing at least 106 mm, apparently is the largest 
species of Chonerhinos and has the deepest cau- 
dal peduncle. Depth of caudal peduncle 6.7—7.4 
(vs. 7.2-9.9 in all other Chonerhinos). Scales 
relatively large and close-set, those on body an- 
teroventral to pectoral fin with spines directed 
dorsally, as in Xenopterus (vs. spines directed 
dorsoposteriorly or posteriorly in all other Cho- 
nerhinos). Upper and lower lips about equally 
projecting or lower lip slightly protruding. Ex- 
posed portion of eye round. Snout gently slop- 
ing. Nasal organ moderately large, its length 
14.7—25.3. Dorsal-fin rays 25-28; and anal-fin 
rays 20-22 (generally fewer in other Chonerhi- 
nos). 

REMARKS ON SYNONYMY.—Most records of 
C. modestus in the literature other than those 
cited in the synonymy above refer in whole or 
in part to other species of Chonerhinos. 

Chonerhinos africanus was described briefly 
(and without a figure) on the basis of a single 
specimen supposedly obtained together with 
other fish specimens by E. Luja in the Sankuru 
River, Kasai, Congo basin, in 1908. No addi- 
tional specimens of Chonerhinos have been 
found in Africa, and the holotype has not been 
compared previously to Chonerhinos from 
Southeast Asia. I have examined the 52.5-mm 
holotype (Fig. Sa), comparing it directly with 
specimens of all five species of Chonerhinos, 
and conclude that it is conspecific with C. mo- 
destus. It has 26 dorsal-fin rays; 22 anal-fin rays; 
15 pectoral-fin rays; 10 + 16 vertebrae; scales 
relatively large, those on sides of body antero- 
ventral to pectoral fin with spines directed dor- 
sally; lower lip slightly protruding; snout gently 


10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


FIGURE 6. Chonerhinos nefastus, 47.0 mm, MZB 3980 (holotype). 


sloping; eye 10.7; olfactory organ 18.7; snout 
7.2; interorbital width 5.25; depth caudal pedun- 
cle 7.4; length caudal peduncle 5.9; and pectoral- 
fin base 10.8. The caudal peduncle depth and 
anal-fin ray count, while not conclusive, agree 
best with C. modestus. Presumably, the speci- 
men originated somewhere in Southeast Asia 
and somehow became mixed with Luja’s mate- 
rial when it was sent on loan to the BMNH for 
Boulenger to study. 


Chonerhinos nefastus new species 
(Figure 6) 


Chonerhinos modestus D’ AUBENTON AND BLANC, 1966:561 
(Mekong basin, Kampuchea); Taxi 1974:199-200, fig. 187 
(Mekong basin, Laos); IMAKI et al. 1978:29, pl. 18 (Kapuas 
R. at Sintang); TYLER 1980 (fig. 2237). 


HoLotyPe.—MZB 3980, 47.0 mm, Kapuas R. 29 km w of 
Putussibau, Kapuas Ichthyological Survey, 11 Aug. 1976. 

PARATYPES.—CAS 49506 and MZB 3981, 4:34.3-43.3 mm, 
Kapuas R. near Kampong Nibung, 7 km NE of Selimbau, Ka- 
puas Ichthyological Survey, 5—6 July 1976; BMNH 1982.3.29. 
254-5 and MZB 3982, 3:36.6-43.7 mm, Kapuas R. 53 km w 
of Putussibau, Kapuas Ichthyological Survey 6-7 Aug. 1976; 
IRSNB 632, MZB 3983, ROM 38601, and USNM 230361, 
6:32.9-60.2 mm, Kapuas R. about 23 km wsw of Putussibau, 
Kapuas Ichthyological Survey, 8-9 Aug. 1976; MZB 3984, 


51.7 mm, Kapuas basin, small tributary of Sungai Mandai 17 
km wsw of Putussibau, Kapuas Ichthyological Survey, 10 
Aug. 1976; MZB 3985, 64.9 mm, Kapuas basin, Sungai Mandai 
Kechil, 18 km wsw of Putussibau, Kapuas Ichthyological Sur- 
vey, 11 Aug. 1976; CAS 49507 and MZB 3986, 3:36.7-57.8 
mm, Kapuas basin, Sungai Tawang near Danau Pengembung, 
Kapuas Ichthyological Survey, 14-15 Aug. 1976; RMNH 
7936, 61.8 mm, Kapuas basin, Sibau, June 1894; RMNH un- 
cat., 25.5 mm, Kapuas basin, Sintang, July 1894; ZMA 
110.220, 65.8 mm, Kapuas basin, Bunut, H. A. Lorentz, 26 
June 1909; FMNH uncat. 3:38.2—50.1 mm, Sarawak, Niah R., 
T. Harrisson, | Apr. 1963; FMNH uncat., 2:64.8-70.7 mm, 
Sarawak, Niah, T. Harrisson, no date; FMNH uncat., 
3:36.9-41.8 mm, Sarawak, Rejang basin, Baleh R. between 
Sungai Mujong and Sungai Gaat, R. F. Inger, 3 Aug. 1956; 
RMNH 7933, 2:56.6-68.0 mm, Mahakam basin, Tepoe, A. W. 
Nieuwenhuis, 1896-97; MG 2058.94, 34.9 mm, Kalimantan 
Tengah, Mentaya basin near Sampit, Pfeuffer, May 1980; 
UMM Z uncat., 50.1 mm, Sumatra, Moesi R. at Moera Klingi, 
A. Thienemann, 1913; SU 36040, 41.7 mm, Malay Peninsula, Per- 
ak, Chandra dam, A. W. Herre, 18 Mar. 1923; UMMZ 197038, 
43.7 mm, Thailand, Songkhla Lake off Patalung, K. F. Lagler, 
6 Jan. 1965; UMMZ uncat., 48.0 mm, Thailand, Mekong ba- 
sin, Ubon Ratchtani, Huay Phai, 16 Oct. 1975; UMMZ uncat., 
38.9 mm, Thailand, Mekong basin, Ubon Ratchtani, Huay 
Kwang, | Oct. 1976; UMMZ uncat., 42.5 mm, Thailand, Me- 
kong basin, Huay Kwang s of Khong Chiam, Arden, 7 Oct. 
1975; UMMZ uncat., 30.9 mm, Thailand, Mekong basin, Mun 
R. at Khong Chiam, Songrad and Buskirk, 19 July 1975; 
UMMZ uncat., 3:15.4-32.2 mm, Thailand, Mekong R. and 


ROBERTS: FRESHWATER PUFFERFISH 


Chonerhinos remotus 


FIGURE 7. 


tributaries from Ban Dan to Nakon Phanom, Mekong fish sur- 
vey, Mar.—Apr. 1975; MNHN 1966.55-S6, 9:21.6-48.1 mm, 
Kampuchea, Mekong basin, Prek Tasom, F. d’Aubenton, 5 
June and 9 Nov. 1961; MNHN 1966.57, 12:19.0-47.5 mm, 
Kampuchea, Mekong basin, Prek Andor, F. d’Aubenton, 2 
Dec. 1961. 


D1aGNosis.—Chonerhinos nefastus differs 
from all other species of Chonerhinos in having 
upper lip usually projecting beyond lower lip; 
nasal organ relatively small (Table 1); and ex- 
posed portion of eyeball usually horizontally 
oval rather than round or vertically oval. It dif- 
fers from all other species except C. amabilis in 
having scales on side of body anterior and ven- 
tral to pectoral fin usually with spines directed 
posteriorly (Fig. le), and from all except C. re- 
motus in its slender caudal peduncle (Table 1). 
Body usually without distinct color marks ex- 
cept for a slightly darkened spot on dorsal sur- 
face of head posterior to eyes. 

COMMENTS.—The exposed portion of the 
eyeball is distinctly horizontally oval in more 
than half of the specimens examined. It is usu- 
ally round in very small specimens, however, 
and sometimes round in large specimens (in- 
cluding the holotype). Most specimens have the 
scales on the side of the body anterior and ven- 
tral to the pectoral fin with the spines directed 
posteriorly, as in Figure le. This character is 
variable, however, and in a few specimens the 
spines are directed posterodorsally or almost 
dorsally. This is most noticeable in the sample 
of 12 specimens from Prek Andor, 4 of which 


, 52.7 mm, FMNH 68475 (holotype). 


have the spines more dorsally directed than is 
usual in C. nefastus. The rest of the specimens 
in the sample have the spines directed poste- 
riorly or posterodorsally. Specimens from the 
Mekong River differ from C. nefastus from oth- 
er localities in having a dark transverse mark on 
the dorsal surface of the snout between the up- 
per lip and the nostrils. 

ETYMOLOGY.—Latin nefastus, wicked, 
abominable, in reference to the food habits (see 
below). 


Chonerhinos remotus new species 
(Figure 7) 


Chonerhinos modestus HERRE, 1940:55 (Sandakan District, 
Sungei Segaliud and Sungei Sibuga); INGER AND CHIN 
1962:190-191, fig. 101 (Kinabatangan District). 


HoLotyPeE.—FMNH 68476, 52.7 mm, Kinabatangan basin, 
mouth of Sungai Deramakot, R. F. Inger and P. K. Chin, 27 
Apr. 1956. 

PARATYPES.—FMNH uncat., 9:32.8-54.4 mm, same data as 
holotype; CAS 49743 and FMNH 68475, 61:29.1-56.8 mm, 
Kinabatangan R. below mouth of Malubok R., R. F. Inger and 
P. K. Chin, 25 Apr. 1956; FMNH 68474, 3:47.2-54.4 mm, 
Kinabatangan R. at Deramakot camp, R. F. Inger and P. K. 
Chin, 24 Apr. 1956; FMNH 44931, 38.3 mm, Kinabatangan 
District, N. Borneo Fisheries Dept., 20 Jan. 1949; SU 33487, 
2:60.5-61.4 mm, Sandakan District, Sibugal R. (=Sungai Si- 
buga), A. W. Herre, 19 Apr. 1938; SU 33563, 10:30.5-40.4 
mm, Sandakan District, Segaliud R., A. W. Herre, 4 Feb. 
1937. 


DIAGNosiIs.—Chonerhinos remotus is most 
similar to C. silus, from which it differs in hav- 
ing a more slender caudal peduncle (Table 1); 
fewer dorsal- and anal-fin rays on the average 


12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


FIGURE 8. 


(Table 2); an even more strongly sloping snout; 
lower lip almost always strongly projecting be- 
yond upper lip (vs. lower lip slightly projecting 
or equal to upper lip); and eye vertically oval or 
round (vs. usually round). Scales anterior and 
ventral to pectoral fin moderately large and 
close-set, with spines directed posterodorsally 
(Fig. Id). No distinctive color marks except for 
a well-defined dark blotch on dorsal surface of 
head posterior to eyes. 

ETyYMOLoGY.—Latin remotus, remote, in ref- 
erence to the type-locality. 


Chonerhinos silus new species 
(Figure 8) 

HoLotyPeE.—FMNH 68477, 44.9 mm, Sarawak, Rejang ba- 
sin, Sungai Baleh between Sungai Mujong and Sungai Gaat, 
R. F. Inger, 3 Aug. 1956. 

PARATYPES.—CAS 49744, FMNH uncat., 36:35.0-60.1 mm, 
same data as holotype; FMNH 62987, 44.0 mm, Sarawak, 
Niah R., Niah, Lord Medway, 22 Aug. 1959; FMNH 68813, 
81.8 mm, Sarawak, Niah, T. Harrisson, no date; FMNH 
68814; 2:44.1-68.7 mm, Sarawak, Niah, Niah R., Pengkalan 
Lobang, T. Harrisson, 2-11 Nov. 1960; FMNH 68815, 
16:37.1-72.9 mm, Sarawak, Niah R., T. Harrisson, | Apr. 
1963; SU 33610, 32.7 mm, Sarawak, 16 miles [ca. 26 km] E of 
Kuching, A. W. Herre, 16 Feb. 1937. 


DIAGNosis.—Chonerhinos silus is most simi- 
lar to C. remotus and C. modestus. Differences 
between C. silus and C. remotus are set forth 
above in the diagnosis of C. remotus. It differs 
from C. modestus in attaining smaller adult size 
(largest specimen examined 82 mm vs. 106 mm); 
snout more strongly sloping; lips equally pro- 


Chonerhinos silus, 44.9 mm, FMNH 68477 (holotype). 


jecting, or lower lip variably protruding, fre- 
quently much more so than in C. modestus; 
scales anterior and ventral to pectoral fin with 
spines projecting dorsoposteriorly (Fig. Ic) rath- 
er than dorsally (Fig. 1b); and caudal peduncle 
relatively slender, its depth 7.2-8.2 (vs. 6.7-7.4). 
C. silus tends to have fewer dorsal-, anal-, and 
pectoral-fin rays than C. modestus (Table 2), but 
the counts are broadly overlapping and of little 
help in identifying individual specimens to 
species. 
ETyMoLoGy.—Latin silus, pugnosed. 


COLORATION IN LIFE 


Most of the specimens of Chonerhinos col- 
lected during the 1976 Kapuas Ichthyological 
Survey were caught at night and preserved be- 
fore their coloration in life could be properly 
observed. Colors of the 106-mm C. modestus, 
gill-netted at night and removed the next morn- 
ing, are recorded in my field notes and in a 35- 
mm Kodachrome slide. It was pale blue dorsal- 
ly, white on the sides and abdomen, and with a 
reddish eye. It is my impression that the three 
smaller C. modestus collected during the survey 
were similarly colored. C. amabilis is described 
in my field notes as lime-green dorsally, with a 
darkened area along the base of the dorsal fin, 
and a reddish eye; the round spot on the caudal 
peduncle, so evident in preserved specimens, 
was not observed during life (at least it is not 
recorded in my field notes, and I do not recall 


ROBERTS: FRESHWATER PUFFERFISH 


having seen it in the live specimens). I suspect 
that some C. amabilis were blue dorsally but 
this is not recorded in my field notes. My 
impression is that all C. nefastus caught during 
the survey were pale green dorsally; at least this 
was So in several specimens observed during the 
day. I doubt that any of them were blue dorsally. 
D’ Aubenton and Blanc (1966) reported color- 
ation of C. nefastus (as C. modestus) from the 
Mekong basin in Kampuchea as green on the 
back and white on the flanks and belly, while 
Taki (1974) reported specimens from the Me- 
kong in Laos as having ‘‘back and upper surface 
of head and body olivaceous golden, underside 
pale yellow to white. Dorsal and caudal fins 
greenish yellow; anal fin pale yellow; pectoral 
fins hyaline.” 


SEXES 


Secondary sexual dimorphism is unknown in 
Chonerhinos. | have examined ripe males and 
gravid or ripening females in all five species. 
Ovaries of the left and right sides are about 
equally well developed. The following approxi- 
mate counts of eggs and measurements of egg 
diameters contained in the right ovary were 
made; C. amabilis, 57.4 mm, 180 eggs, 1.1-1.9 
mm; C. modestus, 106 mm, 800 eggs, 1.5—2.1 
mm; C. nefastus, 56.5 mm, 100 eggs, 1.4-1.5 
mm, 57.8 mm, 80 eggs, 1.3 mm, and 64.9 mm, 
230 eggs, 1.3-1.6 mm; C. remotus, 54.2 mm, 85 
eggs, 1.9-2.3 mm; and C. silus, 58.7 mm, 200 
eggs, 1.5-2.1 mm. All of these specimens are 
gravid except the three C. nefastus, which are 
nearly ripe. In C. remotus I observed two gravid 
females, 54.2 and 54.4 mm, and three spent fe- 
males, 48.8, 51.4, and 52.7 mm (the holotype, 
Fig. 7), with genitoanal areas much swollen. 
Such swelling, perhaps present only in females 
just before or after spawning, has not been ob- 
served in other species. 


Foop HABITS 


Food habits of Chonerhinos, determined by 
complete or partial examination of gut contents 
in more than 100 specimens, may be summa- 
rized as follows: C. amabilis feeds almost ex- 
clusively on large aquatic insects; C. modestus 
feeds mainly on terrestrial insects, shrimps, 
seeds, and to a less extent on whole fish, fin 
rays, or scales; C. nefastus feeds mainly on fish 
fin rays and scales, and to a lesser extent on 


insects (aquatic and terrestrial); C. remotus and 
C. silus feed mainly on insects aquatic and ter- 
restrial), but also ingest vegetable matter and 
other items. No fish remains were found in C. 
amabilis, C. silus, or, excepting a single fish 
scale in one specimen, C. remotus. Pieces of 
clam flesh and gills were found in several C. 
silus, and numerous small, whole clams in a sin- 
gle C. nefastus, but otherwise molluscs were 
absent. The food of the five species may be de- 
scribed in more detail as follows. 

In C. amabilis, 18 of 20 specimens contained 
more or less abundant remains of insects, mainly 
large aquatic forms; partial examination of the 
gut contents of these specimens failed to reveal 
any other food items. Of the remaining two 
specimens, one contained moderate amounts of 
an unidentified flocculent material, and one had 
empty guts. This species is noteworthy in that 
nearly all individuals had much food in their 
guts, and in being the most stenophagic of any 
species of Chonerhinos. In C. modestus, guts 
were examined in 10 specimens, half of which 
had empty guts. Of the remaining five, four con- 
tained moderate to large amounts of insects 
(mainly terrestrial), two had prawns, two had 
seeds, two had fish scales, one had fish fin rays, 
and one had the remains of a small whole cobitid 
fish (identified by its Weberian apparatus). The 
last C. modestus, the 106-mm specimen, is of 
particular interest because of its large size and 
because of the circumstances of its capture. It 
was gill-netted together with a large catfish, 
Pangasius polyuranodon (Fig. 9), which had 
much of its abdominal wall and portions of its 
anal and caudal fins and caudal peduncle bitten 
away. I suspected that part of the damage may 
have been done by the C. modestus, but careful 
examination of its gut contents failed to reveal 
any material from the Pangasius. While the C. 
modestus may have regurgitated, its stomach 
did contain other food items, and it seems more 
likely that the Pangasius was ravaged by some 
other predator, possibly C. nefastus. Of 31 C. 
nefastus in which the gut contents were exam- 
ined, 11 had more or less substantial amounts of 
fish fin rays, six had fish scales, three contained 
small pieces of fish flesh, six had small to mod- 
erate amounts of insects (terrestrial and aquat- 
ic), two had unidentified debris or detritus, one 
had numerous small, whole bivalves, and one 
had a large amount of sand and grit; seven had 


14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 


Figure 9. An 106-mm Chonerhinos modestus gill-netted together with an 80-cm Pangasius polyuranodon catfish ravaged 
by an unknown predator, possibly C. modestus or C. nefastus (Kapuas River near Putussibau). 


empty guts. The Latin name nefastus refers to 
the predominantly pterygophagous and lepido- 
phagous habits of this species. Inger and Chin 
(1962:191) reported gut contents of 11 C. re- 
motus (as C. modestus) as follows: bits of leaves 
(6); parts of terrestrial insects (6); Plecoptera 
nymphs (3); Trichoptera larval cases (1); un- 
identified insect larvae (3); Acarina (2); unspec- 
ified parts of fishes (2). Of 21 C. remotus I ex- 
amined, 18 had guts containing food items: 14 
with insects (aquatic and terrestrial), 4 with 
parts of higher plants, | with a mite, | with a 
fish scale, and several with unidentified debris 
or detritus. In 33 C. silus, 22 had guts containing 
insects (aquatic and terrestrial), 6 contained 
higher plant material (fine rootlets, leaf, seeds, 
or seed pulp?), | had several pieces of a large, 
spinulose oligochaete, and | had chunks of spiny 
or hairy flesh (mammalian?); the remainder had 
empty guts. 


INTRASPECIFIC BITING 


Intraspecific biting, although infrequently 
documented, probably occurs in many members 
of the family Tetraodontidae. In Fugu niphobles 
(Jordan and Snyder, 1901), biting is an integral 
part of spawning behavior: egg laying occurs on 
the beach at high tide after a female has been 
bitten on the sides by two to four males (Uno 
1955). Many of the specimens of Chonerhinos 
examined exhibited characteristically shaped 
bite marks on the flanks and, even more fre- 
quently, had portions of the median fins bitten 
off. I suspect that much of the biting, at least in 


C. nefastus, is inflicted by conspecifics. More 
than half of the specimens examined of this 
species had bite marks on the flanks or had por- 
tions of the dorsal, anal, or caudal fins missing. 
In many specimens these fins appear to have 
been bitten repeatedly, as evidenced by scar tis- 
sue and imperfect regeneration of fin rays. It is 
noteworthy that this species feeds predominant- 
ly upon fish fin rays (see above under Food Hab- 
its). C. modestus and C. silus, both of which 
occur sympatrically with C. nefastus, also ex- 
hibit high frequencies of specimens with bite 
marks and bitten fins, but it is unclear whether 
this is a result of intraspecific attacks, attacks 
by C. nefastus, or a combination of both. In all 
three species the bite marks and fin damage ap- 
pear to be about equally distributed between the 
sexes, and between gravid and nongravid fe- 
males. None of the specimens of C. amabilis 
and C. remotus examined exhibited bite marks 
on the flanks, and their fins were relatively un- 
damaged, with little or no indication of fin-nip- 
ping. Perhaps the generally pterygophagous and 
lepidophagous feeding behavior of C. nefastus 
was preceded by the evolution of an exception- 
ally aggressive intraspecific biting and fin-nip- 
ping behavior. 


GEOGRAPHICAL DISTRIBUTION 


Tetraodontidae is the only one of the nine 
families of the large order Plectognathi or Te- 
traodontiformes which has representatives that 
occur in fresh water. About 25 of the approxi- 
mately 140 described tetraodontid species are 


oe 


ROBERTS: FRESHWATER PUFFERFISH 


70° 80° 90° 100° 
\ \ \ \ 
a: 2 aS 
s pies 


amabilis 
modestus 
nefastus 
remotus 
Silus 


! 1 ! 


110° 
\ 


—20° 


Ficure 10. Geographical distribution of species of Chonerhinos. 


endemic to fresh water. Carinotetraodon and 
Chonerhinos, both from Southeast Asia, are the 
only tetraodontid genera restricted to fresh 
water. Other genera with freshwater species in- 
clude Tetraodon or Monotreta in India, South- 
east Asia, and New Guinea; Tetraodon in Afri- 
ca; and Colomesus in South America. Two 
features of the geographical distribution of 
freshwater Tetraodontidae merit comment. 
First, although marine tetraodontids extend into 
high latitudes in the Northern and Southern 
hemispheres, freshwater species occur only 
within tropical latitudes. Second, the tropical 
rivers with endemic tetraodontids generally 
have rich ichthyofaunas dominated by primary 
freshwater fishes. 

Geographical distributions of the species of 
Chonerhinos, based mainly on material exam- 
ined in this study, are illustrated in Figure 10. 
Two of the species, C. amabilis and C. modes- 


tus, have distributions lying within the hydro- 
graphic limits of the ancient Central Sundaland 
River basin, now fragmented by the Java and 
South China seas. I suspect that C. modestus 
also occurs in Thailand but have not examined 
specimens from there. The most widely distrib- 
uted species, C. nefastus, occurs throughout the 
area occupied by the Central Sundaland River 
basin; it also occurs in northern and southern 
Borneo and in the Mekong basin. Whether the 
Mekong River once also formed part of the Cen- 
tral Sundaland drainage is a matter under inves- 
tigation. C. silus and C. remotus, in northern 
and northeastern Borneo, have restricted distri- 
butions entirely outside the limits of the Central 
Sundaland drainage area. C. amabilis, C. mo- 
destus, and C. nefastus occur sympatrically in 
the Kapuas River and probably also in some 
rivers in Sumatra including the Indragiri and 


16 He otra 
4 y Far, 7 * 
4 she api DOU id 
1 J ‘ 5 
Moesi. C. nefastus and C. silus occur sympatri- 


cally in Sarawak (Rejang and Niah basins). 


ACKNOWLEDGMENTS 


It is a pleasure to thank the following individ- 
uals for their help durihg this study; Oliver 
Crimmen, Gordon Howes, and Alwyne Wheel- 
er, BMNH; Marie-Louise Bauchot, Jaques Dag- 
et, and Martine Desoutter, MNHN; Marinus 
Boeseman, Peter van Helsdingen, and M. J. P. 
van Oijen, RMNH; Han Nissen, ZMA; Volker 
Mahnert, MG; Donald J. Stewart, FMNH; Wal- 
ter Rainboth, UMMZ,; Dirk Thys van den Au- 
denaerde, MRAC; Maurice Kottelat, Universite 
de Neuchatel; and Lillian Dempster, Madeleine 
Graham, W. I. Follett, Michael Hearne, and 
James Jackson, CAS. Photography is by AIl- 
phonse Coleman, Museum of Comparative Zo- 
ology, Harvard University, and Orrin Moon, 
The Darkroom, San Rafael. 

The ichthyological survey of the Kapuas basin 
was sponsored by the Museum Zoologicum Bo- 
gorense, Indonesian National Research Council, 
and Smithsonian Tropical Research Institute. 
Soetikno Woerjoatmodjo, Leo Poerwadi, and 
Rajali assisted in the field. Research was done 
during visits to the BMNH, MNHN, RMNH, 
and ZMA, and at the California Academy of Sci- 
ences and Tiburon Center for Environmental 
studies, and was supported by National Science 
Foundation grant DEB77-24759. 


LITERATURE CITED 


BLEEKER, P. 1850. Bijdrage tot de kennis der ichthyologische 
fauna van Borneo, met beschrijving van 16 nieuwe soorten 
van zoetwatervisschen. Nat. Tijdschr. Ned. Ind. 1:1-16. 

. 1854. Vijfde bijdrage tot de kennis der ichthyolo- 

gische fauna van Celebes. Nat. Tijdschr. Ned. Ind. 

7:225-260. 


ant PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 
_ ‘ {i a . 


. 1865. Atlas ichthyologique des Indes Orientales Neéér- 
landaises, vol. 5. 

BouLENGER, G. A. 1909. Catalogue des poissons du Congo 
du Musée d’Histoire naturelle de Luxembourg. Faune de 
Sankuru a Kondué (Collection Ed. Luja). Monatsber. Ge- 
sell. Luxemburg. Naturf., n. ser., 3:189-202. 

CANTOR, T. E. 1850. Catalogue of Malayan fishes. J. Thomas, 
Calcutta. xii + 461 p., 24 pls. 

D’ AUBENTON, F., AND M. BLANC. 1966. Poissons tétraodon- 
tiformes du Cambodge. Bull. Mus. Natl. Hist. Nat. ser. 2, 
38:554-561. 

FRASER-BRUNER, A. 1943. Notes on plectognath fishes.— 
VIII. The classification of the suborder Tetraodontoidea, 
with a synopsis of the genera. Ann. Mag. Nat. Hist. ser. 11, 
10:1-18. 

GILL, T. N. 1892. Note on the genus Chonerhinos or Xe- 
nopterus. Proc. U.S. Natl. Mus. 14:696-699. 

Herre, A. W. 1940. Additions to the fish fauna of Malaya 
and notes on rare or little known Malayan and Bornean 
fishes. Bull. Raffles Mus. 16:27-61. 

HOoLiarpb, H. 1857. Etudes sur les Gymnodontes et en par- 
ticulier sur leur ostéologie et sur les indications qu’elle peut 
fournir pour leur classification. Ann. Sci. Nat. (Paris), zool., 
ser. 4, 8:275-328. 

IMAkI, A., A. KAWAMOTO, AND A. SuzuKI. 1978. A list of 
freshwater fishes collected from the Kapuas River, West 
Kalimantan, Indonesia. The Institute for Breeding Re- 
search, Tokyo University of Agriculture, 50 p. 

INGER, R. F., AND P. K. CHIN. 1962. The fresh-water fishes 
of North Borneo. Fieldiana: Zool. 45:1—263. 

JORDAN, D. S. 1919. The genera of fishes, 2. Stanford Univ. 
Publ., univ. ser., i-x + 163-284 + 1-xiii p. 

Roserts, T. R. 1981. Identification of the presumed African 
freshwater fishes Micracanthus marchei (Belontiidae) and 
Chonerhinos africanus (Tetraodontidae). Cybium, ser. 3, 
5:91-92. 

Taki, Y. 1974. Fishes of the Lao Mekong basin. USAID 
Mission to Laos, Agric. Div., vi + 232 p. 

TyLer, J. C. 1980. Osteology, phylogeny, and higher classi- 
fication of the fishes of the order Plectognathi (Tetraodon- 
tiformes). NOAA Tech. Rep. NMFS Circ. 434, 422 p. 

Uno, Y. 1955. Spawning habit and early development of a 
puffer, Fugu (Torafugu) niphobles (Jordan et Snyder). J. 
Tokyo Univ. Fish. 41:169-183. 

WEBER, M., AND L. F. DE BEAUFORT. 1962. The fishes of 
the Indo-Australian Archipelago, vol. 11. E. J. Brill, Lei- 
den. ix + 481 p. 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


ar 


DINGS) shor aiory 
OF MME: AR Y 


CALIFORNIL agin hi sash eee S 


| 


Vol. 43, No. 2, pp. 17-24, 7 figs. 


| Woods Hole, Mass. 


June 15, 1982 


SYNONYMIES OF INDIAN OCEAN EELS, WITH THE 
DESCRIPTION OF GYMNOTHORAX ENIGMATICUS, 
A MORAY PREVIOUSLY KNOWN AS G. RUPPELI 


By 


John E. McCosker 


Steinhart Aquarium, California Academy of Sciences, Golden Gate Park, 
San Francisco, California 94118 


and 
John E. Randall 
Bernice P. Bishop Museum, Honolulu, Hawaii 96818 


ABSTRACT: 


The common, banded Indo-Pacific morays called Gymnothorax petelli (Bleeker, 1856) and G. 


ruppeli (McClelland, 1845) by recent authors are recognized as G. rueppelliae (McClelland, 1845) and G. 
enigmaticus n.sp., respectively. They are separable on the basis of coloration, vertebrae, and morphology and 
have different geographic ranges. G. signifier Bliss, 1883, is placed in the synonymy of G. rueppelliae, along 
with Muraena umbrofasciata Ruppell, 1852; M. interrupta Kaup, 1856; Sideria chlevastes Jordan and Gilbert, 
1883; G. leucacme Jenkins, 1904; and G. waialuae Snyder, 1904. The moray Uropterygius xanthopterus Bleeker, 
1859, is recognized as distinct from U. marmoratus (Lacepede, 1803), and U. alboguttatus Smith, 1962, is 
synonymous with it. Ophichthus retifer Fowler, 1935, from Durban, South Africa, is a synonym of O. erabo 
(Jordan and Snyder, 1901), an ophichthid also known from Hawaii, Japan, and Taiwan. 


INTRODUCTION 


In preparation for the publication of the eel 
section of the revised Sea Fishes of Southern 
Africa (McCosker and Castle, Ms), we assign 
several poorly known taxa to synonymy and 
provide a description for a common, conspicu- 
ously banded Indo-Pacific moray, Gymnothorax 
ruppeli of earlier authors, which lacks a holo- 
type and scientific name. 


METHODS 


Measurements are straight-line, made either 
with a 300-mm ruler with 0.5-mm gradations (for 
total length, trunk length, and tail length) and 
recorded to the nearest 0.5 mm, or with dial cal- 
ipers (all other measurements) and recorded to 
the nearest 0.1 mm. Body length comprises head 
and trunk lengths. Head length is measured from 


[17] 


the snout tip to the posterodorsal margin of the 
gill opening; trunk length is taken from the end 
of the head to mid-anus; maximum body depth 
does not include the median fins. Vertebral 
counts (which include the hypural) were taken 
from radiographs. Materials used in this study 
are housed at the following institutions: Acad- 
emy of Natural Sciences of Philadelphia 
(ANSP); Bernice P. Bishop Museum (BPBM); 
British Museum of Natural History (BMNH); 
California Academy of Sciences (CAS); U.S. 
National Museum of Natural History (USNM); 
Museum of Comparative Zoology, Harvard Uni- 
versity (MCZ); J. L. B. Smith Institute of Ich- 
thyology, Rhodes University (RUSI); Natur- 
Museum Senckenberg (SMF); and the Scripps 
Institution of Oceanography (SIO). Paratypes of 
the new species will also be deposited at the 


18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2 


Ficure |. Paratype of Gymnothorax enigmaticus n.sp., BPBM 9491, 335 mm TL, from Palau. 


Museum National d’Histoire Naturelle, Paris 
(MNHN), the BMNH, and the USNM. 


FAMILY MURAENIDAE 


Gymnothorax enigmaticus, new species 
(Figure 1) 


Holotype —CAS 48815, 303.2 mm total length, a male (?) 
collected with rotenone in a 0-1-m tidal flat, off southern cor- 
ner Ngatchab Beach, Angaur I., Palau, Western Caroline Is., 
by H. DeWitt, Sumang, and Sengich, 21 Oct. 1957. 

Paratypes.—Collected in shallow coral reef flats and tide- 
pools (0-3 m) using rotenone ichthyocides. PALAU: CAS 48823 
(8; 51-299 mm), Angaur I., H. DeWitt et al., 22 Oct. 1957. 
CAS 48817 (293 mm), Auluptagel I., H. Fehlmann et al., 10 
Aug. 1955. CAS 48816 (335.5 mm), Ngethil I., Sumang and R. 
Johaness, 13 July 1959. CAS 48822 (2; 238-257 mm), Aulong 
I., Sumang, 5 Nov. 1959. CAS 48826 (301.5 mm), Urukthapel 
I., H. Fehlmann et al., 19 Aug. 1955. BPBM 9491 (335 mm), 
Malakal Harbor, A. Emery, 21 Apr. 1970. KAPINGAMARINGI: 
CAS 48818 (163.8 mm), Thokotaman, R. Harry, 12 July 1954. 
IFALUK ATOLL: CAS 48819 (174 mm), Falarik Islet, R. Harry, 
26 Sep. 1953. GuAM: CAS 48820 (214.4 mm), N of Cocos Is., 
Nangauta and H. Fehlmann, 8 Oct. 1958. ENEWETAK ATOLL: 


CAS 42377 (144 mm), Runit I., R. Nolan and L. Taylor, Jr., 
23 Feb. 1974. BPBM 8184 (127 mm), Enewetek I., J. Randall, 
1 Dec. 1967. BPBM 22339 (2; 219-233 mm), Enjebi I., J. Ran- 
dall et al., 27 Apr. 1978. Bikin1 ATOLL: BPBM 12354 (310 
mm), Eman I., V. Brock et al., 18 June 1947. LINE Is.: CAS 
48825 (302 mm), Palmyra I., E. Herald et al., 16 Aug. 1951. 
BPBM 7715 (2; 310-393 mm), Cooper I., J. Randall, 13 Nov. 
1968. HonG Kona: CAS 48821 (3; 79-88 mm), Santa Cruz Is., 
Vanikoro I., R. Bolin, 30 Sep. 1958. INDONESIA: BPBM 20890 
(2; 103-383 mm), Bali, Sanur Beach, J. Randall, 18 July 1977. 
THAILAND: BPBM 22827 (460 mm), Similan I., Ko Miang, J. 
Randall, 14 Feb. 1979. PHILIPPINES: CAS 48824 (2; 508—518.5 
mm), Negros Oriental, D. Empero, 28 July 1958. 


DIAGNosIs.—A moderate-length species of 
Gymnothorax with anus before midbody; tubu- 
lar anterior nostrils; uniserial jaw and vomerine 
teeth; and cream body coloration with 17—21 dis- 
tinctive brown bands encircling head and body 
and extending onto fins. 

DESCRIPTION OF HOLOTYPE (followed paren- 
thetically by mean and range of the condition of 
holotype and nine paratypes).—Greatest depth 


McCOSKER & RANDALL: INDIAN OCEAN EELS 


FIGURE 2. 


of body 16.8 (19.0; 15.4—22.7) times in total 
length (TL). Tail longer than body, its length 
ie7euGl- 76; 167 1—1:82) in Tie: Head, 7.94: (7.69; 
7.19-8.19) and trunk 3.26 (3.30; 3.19-3.55) in 
TL. Dorsal fin low, its origin ahead of gill open- 
ings, arising above fourth vertebra. Snout 6.37 
(5.76; 5.29-6.37), upper jaw 3.01 (2.78; 2.65-3.01) 
times in head length (HL). Eye 9.5 (9.4; 
8.3-10.4) in HL and 1.5 (1.63; 1.41.9) in snout, 
closer to rictus than to tip of snout. Fleshy in- 
terorbital width 7.8 (8.4; 7.7-9.9) in HL. Gill 
openings nearly horizontal, their centers slightly 
below midbody, their length about equal to di- 
ameter of eye. 

Anterior nostril tubular, elongate, slightly less 
than eye diameter in length. Posterior nostril a 
hole above eye, beginning in a line with eye. 

Jaws subequal, the mouth closing completely. 
Teeth in jaws uniserial, stout, pointed and slight- 
ly retrorse. Six pairs of intermaxillary canines 


Gymnothorax rueppelliae, BPBM 18412, 339 mm TL, from Enewetak. 


form a U-shaped margin around three central 
canines, the third the largest. Approximately six 
uniserial, small vomerine teeth. About 12 upper 
jaw teeth pairs, 18 lower jaw pairs; 3 pairs of 
depressible canines behind mandibular sym- 
physis. 

Number of vertebrae 130 (129.7; 128-131), 
50.5 (50.8; 50-51.5) before anal fin. First dorsal 
pterygiophore arises above fourth vertebra. 

Head pores present but not obvious. A single 
pore anterior and proximal to, and a second pore 
below base of anterior nostril. Six pores along 
the mandible, the second through fifth the larg- 
est. Four equally spaced pores along upper jaw, 
the first beneath nostril base, the last beneath 
rear of eye. A single pore between anterior and 
posterior nostrils. 

Color in isopropyl alcohol cream, overlain 
with 17—21 distinctive brown bands which com- 
pletely encircle head and body and extend onto 


20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2 


FIGURE 3. 


median fins. The light interspacing of large in- 
dividuals becomes mottled. The first brown 
band begins mid-snout, ends at the hind margin 
of the eye, and is continuous with lower jaw. 
The second begins at the rictus. Tail tip and an- 
terior nostrils not dark, like body coloration. 
Bleeker’s Atlas (1864: pls. 177 and 183, as G. 
reticularis) contains excellent illustrations 
showing juvenile and adult colorations. 

DISTRIBUTION.—On the basis of our speci- 
mens, field records, and valid literature records, 
the new species is known to range widely within 
the central and western Pacific (excluding Ha- 
waii and Australia) and Indian oceans to Aldabra 
and the Seychelles. Red Sea records (Gunther 
1910; Fowler 1956) are based on Klunzinger’s 
(1871) misidentification of specimens of G. ruep- 
pelliae. 

EtyMoLoGy.—Named enigmaticus, from the 
Latin aenigma, in the light of the nomenclatural 
confusion surrounding this species. 

REMARKS.—Some recent authors have divid- 


Gymnothorax reticularis, BPBM 18734, 492 mm TL, from Hong Kong. 


ed Gymnothorax into Lycodontis for those mo- 
rays without serrated lateral jaw teeth, and 
Gymnothorax for those with serrated lateral jaw 
teeth. We conservatively recognize only Gymno- 
thorax because species intermediate in this con- 
dition exist. 

Two common Indo-Pacific species of Gymno- 
thorax share a distinctive brown banding. 
These were treated by most recent authors as 
G. petelli (Bleeker, 1856) and G. rupelli (Mc- 
Clelland, 1845).* On examining the holotypes, 
the junior author (Randall 1973) discovered that 
they were both the G. “‘petelli’’ form. Thus, G. 
petelli was placed in the synonymy of G. ruep- 
pelliae, and the species described herein lacked 
a name. Following Schultz (in Schultz et al. 
1953), Randall (1973) suggested that G. reticu- 


* McClelland’s species was named Dalophis Riipelliae, 
properly rueppelliae (fide McCosker and Rosenblatt 1975), but 
has been variously and improperly emended to rupelli, rup- 
peli, ruppelli, and ruppellii. 


McCOSKER & RANDALL: INDIAN OCEAN EELS 


Ficure 4. Underwater photograph of adult Gymnothorax enigmaticus taken at night by J. E. Randall, depth ca. 1 m, 
Sumilon Island, Philippines. 


laris Bloch, 1795, was the next available name, 
but McCosker and Rosenblatt (1975) pointed out 
that it is a valid and different species. We have 
subsequently examined the 395-mm holotype 
(SMF 151) of Dalophis Riipelliae and compared 
it to other Indo-Pacific specimens of the ‘‘pefel- 
li’ form. It is in excellent condition and clearly 
conspecific. It has 135 vertebrae, with 50 preanal 
and 4 before the first dorsal pterygiophore. 

Dr. Wolfgang Klausewitz has brought our at- 
tention to Muraena umbrofasciata Ruppell, 
1852, an obscure species which has not been 
reported since its description. On the basis of 
McClelland’s type-specimen of Dalophis Riipel- 
liae and two specimens from India (SMF 2870, 
SMF 7346) received by Ruppell in 1845 from M. 
Boissenau, Ruippell (1852:33) described M. um- 
brofasciata, which we herein recognize as a syn- 
onym of Gymnothorax rueppelliae. 

Smith’s (1962:434) suggestion that Gymno- 
thorax signifer Bliss, 1883, was a synonym of 
G. “‘ruppelli’> prompted us to examine the ho- 


lotype, a 180-mm specimen from Mauritius. Its 
chin, throat, and anterior trunk bands are dis- 
continuous, and it is clearly a specimen of the 
‘‘petelli’’ form, G. rueppelliae. The holotype of 
G. signifer (MCZ 6147) has 132 vertebrae, 51 
preanal; the vertebral range of six specimens of 
G. rueppelliae was 132-135, with a mean of 
133% 7. 

The new species is closely related to G. ruep- 
pelliae (Fig. 2), and might also be confused with 
G. reticularis (Fig. 3). It differs from G. ruep- 
pelliae in possessing light, rather than dark, an- 
terior nostrils, in having continuous chin and 
throat bands, in lacking a black mark at the jaw 
corner, and in attaining a considerably smaller 
adult size. It differs from G. reticularis, which 
is heavily spotted in the head and throat regions 
and lacks the three prominent intermaxillary 
fangs. 

The ‘“‘petelli’’ form, G. rueppelliae, is known 
from shallow water collections from Hawaii, 
throughout the Pacific and Indian oceans, and 


N 
Ne 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 


tN 


FIGURE 5. 


Uropterygius xanthopterus, CAS 35254, 245 mm TL, from Kapingamaringi, displaying white cephalic puncta- 


tions. Arrows indicate the location of the anterior lateral line pores. 


the Red Sea. Its synonyms also include Mu- 
raena interrupta Kaup, 1856; Sideria chlevastes 
Jordan and Gilbert, 1883; Gymnothorax signifer 
Bliss, 1883; G. leucacme Jenkins, 1904; and G. 
waialuae Snyder, 1904. 

Whereas previous literature has suggested 
that most morays are nocturnal, it now appears 
that many, and possibly the majority of, moray 
species are diurnal but rarely observed due to 
their secretive nature (fide Chave and Randall 
1971; Hobson 1974). On the basis of material 
collected and our observations while diving, we 
presume that G. enigmaticus is a shallow-water, 
nocturnally active piscivore (Fig. 4). It is note- 
worthy that G. rueppelliae is also a nocturnal 
predator (Hobson 1974, as G. petelli), as is G. 
undulatus (our observations), both of which are 
also strongly banded species. 


Uropterygius xanthopterus Bleeker, 1859 


Uropterygius xanthopterus Bleeker, 1859, has 
had a sketchy taxonomic history. We have lo- 
cated the type-specimen, recognize it as a valid 
species, and include U. alboguttatus Smith, 
1962, in its synonymy. 

Weber and de Beaufort (1916:397), without 
comment, included U. xanthopterus in the syn- 
onymy of Gymnomuraena marmorata La- 
cepede, 1803, a wide-ranging, elongate Indo-Pa- 
cific species of Uropterygius which possesses a 
single anterior lateral line pore and lacks white 
spotting on its head. Schultz (in Schultz et al. 
1953:154) and Gosline (1958:226), on the basis 


of central Pacific specimens, recognized U. xan- 
thopterus as a distinct small species (the largest 
of 213 specimens from 76 CAS rotenone collec- 
tions in the Indian and central Pacific oceans 
which we examined was 345 mm) which pos- 
sesses two anterior lateral line pores and white 
cephalic punctations (Fig. 5). Smith (1962:427) 
again synonymized U. xanthopterus with U. 
marmoratus and described U. alboguttatus on 
the basis of Indian Ocean and Schultz’s central 
Pacific specimens. In describing U. kamar 
McCosker and Randall, 1977, we considered U. 
alboguttatus to be a possible synonym of U. 
xanthopterus. One of us (JEM) has subsequently 
examined the complete type-series of U. albo- 
guttatus and was unable to find differences in 
coloration, meristic features, or morphometry. 

The type-specimen of U. xanthopterus has 
not been clearly identified; however, through 
correspondence with Alwyne Wheeler, we have 
located the 275-mm specimen in the British Mu- 
seum (cat. no. 1867.11.28.271) received from 
Bleeker and labeled *‘Muraena xanthopterus.” 
In that no specimen similar to Bleeker’s type 
exists in the Rijksmuseum (M. Boeseman, 
in litt.), we presume that this is the type, and 
the specimen which Bleeker illustrated and 
described in his Atlas (1864:pl. CLXIV, fig. 
4). A radiograph of the British Museum speci- 
men clearly indicates that it is not U. marmo- 
ratus, a species which possesses obvious, large 
intramuscular bones. 


McCOSKER & RANDALL: INDIAN OCEAN EELS 


FIGURE 6. 


FAMILY OPHICHTHIDAE 
Ophichthus retifer Fowler, 1935 


Fowler (1935) described and illustrated Oph- 
ichthus retifer on the basis of a 718-mm speci- 
men from Durban, Natal. Eugenie Bohlke has 
kindly examined the holotype (ANSP 63915) for 
us and compared it with a syntype (ANSP 26224) 
of O. erabo (Jordan and Snyder, 1901) from Ja- 
pan. They do not significantly differ in color- 
ation or proportions, yet there is a vertebral dif- 
ference. A radiograph of the holotype of O. 
retifer shows 143 vertebrae, with 73 before the 
anal opening. McCosker (1979) reported that six 
specimens of O. erabo from Japan, Hawaii, and 
Taiwan had 152-155 vertebrae (x = 154). Fow- 
ler (1935) suggested that O. retifer was ‘‘greatly 
like Microdonophis fowleri Jordan and Ever- 


Uropterygius marmoratus, BPBM 12336, 701 mm, Nuku Hiva, Marquesas. 


mann 1903’? (=O. erabo fide McCosker 1979) 
“‘and its synonym Ophicthys garretti Gunther 
1910” (a valid species). We consider O. retifer 
to be conspecific with O. erabo, and account the 
vertebral difference to clinal variation. 


ACKNOWLEDGMENTS 


We thank the following individuals: Susan 
Middleton for photographic assistance; Michael 
Hearne for the preparation of the radiographs; 
M. Boeseman (Rijksmuseum van Natuurlijke 
Historie), Eugenie and James Bohlke (ANSP), 
William Fink (MCZ), W. Klausewitz (SMP), 
Margaret M. Smith (RUSI), and Alwyne Wheel- 
er (BMNH), for assistance with museum speci- 
mens and records; Lillian Dempster and W. I. 
Follett (CAS) for nomenclatural advice; and the 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2 


: 


. ete, e 
j 3.8 17 /@ 3 


FIGURE 7. 


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.-Indié 19:329— 
B52) 

. 1864. Atlas ichthyologique des Indes Orientales Néer- 
landaises. Vol. 4. Amsterdam. 132 p. 

Biss, R. 1883. Descriptions of new species of Mauritian fish- 
es. Trans. Soc. Roy. Arts Sci., Maurice 13:45-63. 

Biocu, M. E. 1795. Naturgeschichte der Auslandischen 
Fische. Vol. 9. 

CHAVE, E. H., AND H. A. RANDALL. 1971. Feeding behavior 
of the moray eel, Gymnothorax pictus. Copeia 1971 
(3):570-574. 

Fow _er, 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. 

GosLINnE, W. A. 1958. Central Pacific eels of the genus Urop- 
terygius, with the descriptions of two new species. Pac. Sci. 
12(3):221—228. 

GUNTHER, A. 1910. Andrew Garrett’s Fische der Siidsee, 
... Heft IX. J. Mus. Godeffroy, Hamburg 17:389-515. 
Hosson, E. S. 1974. Feeding relationships of teleostean fish- 
es on coral reefs in Kona, Hawaii. Fish Bull., U.S. 

72(4):915—-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. 


Adult Ophichthus erabo, from Jordan and Snyder (1901). 


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. 
22(1):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. Apodal fishes of 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(11):167—214. 

RUPppPELL, 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 Indo-Australian Archipelago. Vol. 3. Leiden. 455 p. 


Arch. Naturges. 


PROCEEDINGS 
é Maries. Filegical Latioratory 


CALIFORNIA ACADEMY ¢ OF SCIEN 
| JUL 9 


| Méoscs bohe: Mass. { 


Vol. 43, No. 3, pp. 27-42 


LipRaRY 
ES 
1982 


June 15, 1982 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX WASPS 
(HYMENOPTERA, SPHECIDAE) 


By 
Wojciech J. Pulawski 


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 Junction), krombeiniellus (Florida: Levy County), /amellatus (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), verticalis (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; EEG, 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, 
B.x~ Wiliams} HKC. A. Ky Coun2JeHe Isc. 
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] 


28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


M. S. Wasbauer; PDH, P. D. Hurd; PMM, P. 
M-eMarsh:sPPL, Pe: E> forchio- ROB Re: CC: 
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. Lawgence, 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. Kurezews- 
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 (see Pulawski 1971, 1974, 1977), but 
only four of them are represented in North 
America. They are: the pompiliformis, termi- 
natus, brullii, and julliani groups. The species 
described in this paper belong to the pompili- 


formis and brullii 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 mirandus). By comparison, in 
the julliani 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 brullii 
group, the apical female tarsomeres are simple 
(see that group for details). The pompiliformis 
group is cosmopolitan. Its species prey upon 
acridid nymphs, but the Palearctic species ful- 
vitarsis collects tettigonids. The following new 
species are members of the pompiliformis 
group: apricus, arizonac, bohartorum, idiotri- 
chus, 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 angulate 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. 
ETYMOLOGyY.—The specific name apricus is 


a Latin word meaning exposed to the sun. 
D1aGnosis.—Tachysphex apricus differs from 


29 


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 2, 60 d, 31 Mar. to 3 July, 10 and 31 Aug., 
9 Sep. Specimens for which institution is not indicated below 
are all in UCD. 


UNITED STATES OF AMERICA 


Arizona. Cochise: 6 mi. N Apache, collector unknown (1 
3, NYSU). Coconino: 4.5 mi. E Moenkopi, JMD & MAC (1 
2, ASU). Maricopa: 10 mi. E Gila Bend, GDB (2 ¢); 3 mi. 
sw Wickenburg, PFT & GEB (1 6, USU). Mohave: 4 mi. w 
Chloride, PFT, GEB, FDP (1 6, USU); 8 mi. E Mesquite 
(Nevada), FDP & PFT (1 2, USU). Pima: Organ Pipe Cactus 
National Monument, J. L. Sperry (1 2, | ¢); Tucson, W. 
Benedict (1 2, NYSU), Bryant (1 2, 1 6, CAS), FDP, LAS 
(1 2,246; 12, WJP). Pinal: w Stanfield, GDB & FGW (1 
2). Yavapai: Bloody Basin, collector unknown (1 2); 10 mi. 
Nw Congress, FDP & LAS (1 2). 

California. Imperial: Glamis, RMB (1 2), FDP (1 ¢); Palo 
Verde, ROS (1 2); Pinto Flat, FXW (1 6, CAS). Inyo: An- 
telope Springs, HKC (1 2, 5 ¢); Big Pine Cr., RMB (1 2), 
FDP (1 do); 2 and 5 mi. E Big Pine, EEG (1 2, 1 G); Little 
Lake, BV (1 2); 3 mi. Ww Lone Pine, RMB (2 ¢); Tuttle Cr. 
(2 mi. sw Lone Pine), JAP (1 2, CIS). Kern: Kernville, T. R. 
Haig (1 ¢). Riverside: 18 mi. w Blythe, RMB (1 ¢); 3.5 and 
4 mi. s Palm Desert, MEI, S. Frommer & R. M. Worley (2 
2 , UCR); San Andreas Canyon, RMB (1 ¢); Shavers Summit, 
MSW (1 2, UCD); San Timoteo Canyon, MSW, R. McMaster 
(1 2, CSDA). San Bernardino: | mi. s Adelanto, MEI (1 ¢d); 
Colton Hills, TG (2 2, TG), Kramer, MSW (2 2,3 6, CSDA); 
3 mi. s Kramer Junction, MEI (1 2); sand dunes 7 mi. sw 
Kelso, MSW & J. S. Wasbauer (1 2, CSDA), Mitchells Cav- 
erns, TG (1 2, TG), 36 road mi. E Twentynine Palms, TG (1 
3, TG). San Diego: Borrego Valley, RMB (4 d; 1 2,2 4, 
USNM), JCH (1 6), PDH (4 3), G. A. Marsh (1 2), EIS (2 
3; 26, WJP), MSW (1 6, CIS), MSW, J. Slansky (1 9, 1 
3, CSDA), FXW (9 3, CAS); Scissors Crossing, J. C. Down- 
ey (1 2), H: & M. Townes (1 6, HKT). Ventura: Sespe Can- 
yon, R. W. Sporne (1 <G). 

Nevada. Clark: Jean, GEB (3 ¢); 30 mi. s Searchlight, PFT, 
R. Rust, Youssef (1 ¢, USU). 


30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


New Mexico. Dona Ana: Las Cruces, RMB (1 2, UCD). 
Otero: Alamogordo, collector unknown (1 2); Alamo Canyon 
near Alamogordo, MEI (1 6, UCR). 

Texas. Brewster: Big Bend National Park (Nine Point 
Draw), W. R. M. Mason (1 2, CNC). 


MExICcOo 


Lower California. San Angel, N. Leppla, JMD, J. Bigelow, 
M. Bentzien, W. Fox, S. William, MAC (1 2, ASU); 16 mi. 
N Puertocitos, MEI (1 2, UCR). 


Tachysphex arizonac sp.n. 


ETyMOLOGY.—The specific name arizonac is 
an Indian word meaning little spring. 

DIAGNOSIS.—Tachysphex arizonac is charac- 
terized by the well-defined mesopleural punc- 
tures, median scutal setae transversely oriented, 
and sternum I with a horizontal depression api- 
cally. The females of arizonac and lamellatus 
have a peculiar clypeus whose free margin is 
undulate; they can scarcely be distinguished 
from each other. A useful character is the hy- 
postomal carina which is low in arizonac, but 
high in many lamellatus. Furthermore, in some 
arizonac the middle projection of the clypeal lip 
is markedly larger than the sublateral one (pro- 
jections about equal in /Jamellatus). The male of 
arizonac has a peculiar clypeus: bevel semilu- 
nate, lobe forecorner prominent (acutely in 
some specimens), lip usually with obtuse pro- 
jection mesally. The clypeus is somewhat simi- 
lar in fexanus, but in that species setae are ap- 
pressed or nearly so beneath the mesopleural 
scrobe (setae suberect in arizonac). 

GEOGRAPHIC DISTRIBUTION.—Southern Utah, 
Arizona, and adjacent areas of California; So- 
nora State in Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: 6, Arizona, Cochise 
Co., 2 mi. NE Portal, 26 June 1964, J. M. Puckle, M. A. Mor- 
tenson, M. A. Cazier (UCD). 

PARATYPES: 13 2, 17 36, 5 May to 17 July. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Huachuca Mts., FXW (1 2, CAS; | 2, 
3. ¢, UCD). Maricopa: 3 mi. sw Wickenburg, PFT & GEB 
(1 2, UCD). Pima: Brown Canyon, Baboquivari Mts., K. W. 
Radford & W. Patterson (1 2, 2 6, UAT); Sabino Canyon, 
Santa Catalina Mts., RMB (1 3, UCD), collector unknown 
(1 2, WJP). 

California. Imperial: Experimental Farm and no specific lo- 
cality, J. C. Bridwell (3 2, 1 6, USNM; 2 2, 1 3d, WIP). 
Inyo: Big Pine, N. J. Smith (1 ¢, UCD); Deep Springs, H. 
Nakakihara (1 ¢, UCR). Riverside: Upper Deep Canyon at 
Horsethief Cr., MEI (1 6, UCR). 

Utah. Washington: Leeds Canyon, G. F. Knowlton, W. J. 
Hanson, T. H. Hsiao (2 2, 2 6, USU; 1 2, WJP); Santa 


Clara, FDP & PFT (1 6, USU); Toquerville, G. F. Knowlton, 
W. J. Hanson (1 ¢, USU). . 


MEXICO 


Sonora. Cocorit, FDP & LAS (1 6, UCD); La Aduana, 
FDP & LAS (1 6, UCD). 


Tachysphex bohartorum sp.n. 


ETYMOLOGY.—This species is dedicated to 
Dr. G. E. Bohart (Logan, Utah) and Dr. R. M. 
Bohart (Davis, California) who have made ex- 
tensive contributions to our knowledge of North 
American Tachysphex, and who have also 
helped me in my studies. 

D1aGNosis.—Like pechumani, the female of 
bohartorum has brassy golden frontal vestiture. 
The frontal vestiture is also golden in psilocerus 
which has a peculiar upper metapleuron. Unlike 
pechumani, the gaster of female bohartorum is 
all red, the flagellum is all black, the clypeal lip 
is deeply indented laterally (shallowly in pechu- 
mani), and the mesothoracic venter is peculiar 
(the posterior, horizontal part is shorter than the 
anterior, oblique part); the densely punctate- 
throughout tergum V is a subsidiary recognition 
feature. T. bohartorum is known from Califor- 
nia, Oregon, and Nevada, while pechumani oc- 
curs in New Jersey and Michigan. 

The male of bohartorum can be recognized by 
the closely punctate terga (punctures mainly 
subcontiguous) combined with the apicomesally 
impunctate and glabrous sterna II-IV. 

Supplementary diagnostic characters of both 
sexes are: propodeum not ridged (at most the 
hindface has a few, inconspicuous ridges be- 
low); propodeal dorsum with setae appressed 
mesally, oriented obliquely anterad. 

GEOGRAPHIC DISTRIBUTION.—Montane areas 
of southern Oregon and northern California, also 
Sierra Nevada. 


MATERIAL EXAMINED.—HOLOTYPE: & , California, Nevada 
Co., Boca, 11 July 1961, R. M. Bohart (UCD). 

PARATYPES: 90 2 , 231 ¢, 24 Apr., 8 June to 14 Aug., mainly 
UCD, also BMNH, CAS, CIS, CNC, CSDA, LACM, MCZ, 
OSU, TG, UCR, USNM, WIP. 

California. Alpine: Carson Pass, RMB (1 ¢); Hope Valley, 
PDH (1 3), P. C. Hutchinson (1 ); 15 mi. NE Red Lake, EIS 
(2 3); Winnemucca Lake, RMB (1 9, 2 3), N. J. Smith (2 
2, 14); Woodfords, W. W. Middlekauf (3 3). Del Norte: 
Little Grayback (NE part of county), JAP (4 2, 4 6, CIS). 
Eldorado: Meyers, RMB (1 6d); Tahoe, FXW (3 2, 3 3); 
Strawberry Valley, E. C. VanDyke (1 2); Echo Lake, W. W. 
Middlekauf (1 2, 1 6); Echo Portal, P. H. Arnaud (1 2); Echo 
Pass, C. A. Downing (1 2). Humboldt: Red Cape Lake, TG 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX 


(1 2, 14, TG). Inyo: Big Pine Cr., 7500 ft, RMB (1 @, 1 
3); near Mono Pass, 12,000 ft, C. D. MacNeill (1 2, CIS). 
Lassen: Bridge Cr. Camp, RCB, RMB, G. Schaefers (6 2, 1 
3); Summit Camp, JWMS (1 2). Mariposa: Sentinel Dome 
Cutoff (Yosemite National Park), T. N. Seeno (1 2). Modoc: 
Cedar Pass in Warner Mts., JWMS (1 2, 2 d, CIS), 6000 ft, 
collector unknown (1 2); Warner Mts. 2 mi. NNW Fort Bid- 
well, D. C. Rentz & C. D. MacNeill (1 6, CAS). Mono: 11 mi. 
N Bridgeport, RMB, PMM (3 3); East Walker R., 13 mi. NE 
Bridgeport, MSW & J. S. Wasbauer (2 ¢); Leavitt Meadow, 
R. L. Usinger (1 ¢, CIS). Nevada: Boca, RMB, MEI, PMM, 
WJP (8 2, 11 ¢); Sagehen Cr. near Hobart Mills, RMB, R. 
C. Blaylock, R. L. Brumley, M. A. Chambers, R. H. Good- 
win, EEG, D. S. Horning, PDH, MEI, JAP, WJP, G. Schae- 
fers, R. L. Westcott (15 2, 98 3). Placer: Carnelian Bay (Lake 
Tahoe), RMB, FDP, WJP, BV (3 @, 21 d); Tahoe, FXW (1 
do). Plumas: Bucks Lake, EIS (1 d); Burks, FXW (1 2); Lake 
Almanor, E. G. Linsley (1 6); Meadow Valley, W. H. Nelson 
(1 2); 14 mi. w Quincy, W. Turner (1 2, 2 6, WSU). Shasta: 
Lake Eiler, C. H. Spitzer (1 2); Lassen Peak, 7500 ft, JWMS 
(1 ©). Sierra: Independence Lake, RMB, R. D. Moon (16 
2,516), BV (1 2); Kyburz Flat, RMB (1 6); Sierra Buttes, 
F. J. Montgomery (1 2); Sierra Valley, RMB (1 ¢); Sierra- 
ville, RMB (1 6); Yuba Pass, MEI (1 ¢, UCR). Siskiyou: 
McBridge Springs (3 mi. NNE Mt. Shasta City), C. D. MacNeill 
(3 3); Mount Shasta, JAP (2 2), 1 mi. se Salmon Mt., TG (2 
3, TG). Trinity: Coffee Cr. Ranger Sta., A. J. Mueller (1 
2), JWMS (1 2, CIS). Tuolumne: Chipmunk Flat, ASM (1 
2), JWMS (1 2, CIS); Dardanelle, EIS (1 2); Sonora Pass, 
9-10,000 ft, S. M. Kappos, JWMS (1 2,2 32). 

Nevada. Douglas: Spooners Lake N junction Hwy. 28, P. 
Adams (2 3). Washoe: Mount Rose, 6500 ft, RMB (1 3). 

Oregon. Jackson: 8 mi. SE Butte Falls, R. L. Westcott (1 
2, OSDA). Klamath: 15 mi. NE Bly, Schuh & Denning (1 °, 
OSU); Eagle Ridge near Klamath Lake, C. L. Fox (1 2); Lake 
of the Woods, H. A. Scullen (2 2). Lake: Warner Pass, DRM 
(1 3). 


Tachysphex idiotrichus sp.n. 


ETYMOLOGyY.—The specific name idiotrichus 
is derived from the Greek words idios, own, pe- 
culiar, and trix (tricho-), hair; with reference to 
the peculiar vestiture. 

D1aGnosis.—lTachysphex idiotrichus differs 
from other North American species of the pom- 
piliformis group by the unusually long setae on 
the head, thorax, and femora (the vertex setae, 
for example, equal 2.2—3.0 midocellar diame- 
ters); the large punctures on the middle section 
of the female clypeus (also basally); the pres- 
ence of graduli on sterna III-V of the female; 


and sparsely punctate (except apex) male ter- 


gum VII. Subsidiary diagnostic characters are: 
sparsely punctate vertex; and in the male: vertex 
width more than twice length (like pechumani); 
presence of graduli on sterna III-VI (like ash- 
meadii, glabrior, irregularis, and verticalis); and 
compressed forefemoral notch (as in apricus). 


31 


GEOGRAPHIC DISTRIBUTION.—LoW moun- 
tains of southwestern Texas, New Mexico, 
southern Arizona, and southern California south 
to Jalisco State, Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: 2, Arizona, Cochise 
Co., Southwest Research Sta., 5 mi. w Portal, 20 Oct. 1962, 
Vincent Roth (UCD). 

PARATYPES: 25 2, 11 5; 19 Feb. to 30 Nov. 


UNITED STATES OF AMERICA 


Arizona. Cochise: 3 mi. E Apache, Rozen & Schrammel (1 
2, UCD); 2 mi. w Chiricahua National Monument, A. L. 
Steiner (1 2, WJP); Nw Portal, O. W. Richards (2 9, 
BMNH); 2 mi. sw Portal, A. L. Steiner (1 2, UAE); same 
data as holotype, except | 2, 19 Feb. 1963 (3 9; UCD, 
USNM, WJP); same locality, MAC & Ordway (1 6 AMNH): 
Sulphur Springs Valley, G. Munson (1 2, UCD); 14 mi. w 
Tombstone, RMB (1 2, AMNH;4 2,26, UCD; 1¢,USNM; 
1 6, WJP); same locality, FDP (1 2, 2 6, UCD); 1 mi. sE 
Willcox, G. I. Stage (1 2, CAS). Pima: Tucson, M. L. Noller 
(1 2, UAT). Yavapai: Cottonwood, R. C. Miller (2 2, 1 6, 
UCD); 7 mi. N Cottonwood, R. C. Miller (1 ¢6, UCD). 

California. San Bernardino: Mid Hills (9 mi. ssE Cima), 
5400-5500 ft, TG (2 2, TG). 

New Mexico. Hidalgo: Rodeo to Road Forks (1 2, UCD). 
Otero: Alamogordo, collector unknown (2 2 , UCD). Socorro: 
10 mi. w Socorro, U. N. Lanham (1 2, CSU). 

Texas. Pecos: no specific locality, RMB (1 ¢, UCD). 


MExIco 


Jalisco. Lagos de Moreno, R. C. Bechtel & EIS (1 d, 
UCD). 


Tachysphex irregularis sp.n. 


ETYMOLOGyY.—The specific name irregularis 
is a Latin word for irregular, with reference to 
the mesopleural sculpture. 

D1aGnosis.—lachysphex irregularis differs 
from other members of the pompiliformis group 
in having a rugose or punctatorugose mesopleu- 
ron. The subsidiary recognition features are: the 
almost impunctate venter of the female trochan- 
ters, and in the male: presence of graduli on ster- 
na III-VI, sternal punctures about as large as 
those on the mesothoracic venter. 

GEOGRAPHIC DISTRIBUTION.—Western U.S., 
eastwards to Wyoming and New Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: 2, California, Lassen 
Co., Hallelujah Junction, 12 July 1954, G. A. Schaefers 
(UCD). 

PARATYPES: 41 2, 29 5; 16 Apr. to 11 Sep. 

Arizona. Cochise: 8 mi. NE Apache, PDH, E. G. Linsley 
(1 2,14, CIS); 14 mi. w Tombstone, FDP (1 ¢, UCD). Pinal: 
8 mi. SE Olberg, M. A. Mortenson, JMD, MAC (1 2, UCD). 

California. Inyo: Bishop, F. P. VanDuzee (2 2, UCD, 
WJP). Lassen: Hallelujah Junction, RMB, J. E. Gillaspy, C. 
J. Horning, G. A. Schaefers, F. Morishita, EIS, R. H. James, 


32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


R. L. Sisson (11 2, 10 6; AMNH, MCZ, UCD, USNM, 
WJP). Modoc: 5.5 mi. E Cedarville, V. L. Vesterby (1 ¢, 
UCD); Hot Cr., RMB & GEB (1 2, UCD). Mono: Benton 
Crossing, F. G. Andrews (2 2, CSDA). Nevada: Boca, RMB 
(5 2, 1 6, UCD), MEI (1 36, UCD); Sagehen Cr., PDH (1 
3, UCD). Sierra: Sattley, RMB (2 ¢, UCD). Siskiyou: Mac- 
doel, J. Schuh (2 2, UCD), Hatfield, J. Schuh (1 6, UCD); 
Mt. Shasta City, JAP (1 2, UCD); Red Rock, J. Schuh (1 
2, UCD). Tuolumne: Sonora Pass, R. W. Thorp (1 2, UCD). 

Colorado. Jackson: 10 mi. N junction of roads 14 & 40, R. 
& K. Dreisbach (1 2, UCD). Mineral: South Clear Cr., TG 
(1 2, TG). Routt: 7 mi. E Hayden, J. S. Buckett (1 ¢, UCD). 

Idaho. Canyon: Nampa, GEB (1 2, UCD). Cassia: 4 mi. SE 
Malta, R. L. Westcott (1 2, UIM); 5 mi. N Malta, R. A. 
Mackie (1 2, UIM). Custer: 2 mi. E Leslie, D. S. Horning (1 
2, UCD). Elmore: 4 mi. E Orchard, A. J. Walz (1 6, UIM). 
Owyhee: Silver City, A. R. Gittins (1 2, UIM); 17 mi. w Silver 
City, A. R. Gittins (1 2, 1 d, UIM). 

Nevada. Elko: Cobb Cr., 6 mi. sw Mountain City, P. H. 
Baldwin (1 2, UCD). Humboldt: Orovada, MEI (1 6, UCD), 
FDP (1 2, UCD); Paradise City, P. H. Baldwin (1 6 , UCD); 
15 mi. E Paradise City, collector unknown (1 ¢, WJP). Wash- 
oe: Reno Hot Springs, C. N. Slobodchikoff (1 2, CAS), Sky 
Ranch near Reno, EIS (1 ¢, UCD). 

New Mexico. Dona Ana: Las Cruces, Arabella Sanchez (1 
2, UCD). Hidalgo: Rodeo, PDH (1 6, UCD), H. V. Weems 
(1 6, FSCA). 

Oregon. Klamath: Lower Klamath Lake, J. Schuh (1 °, 
UCD). 

Wyoming. Fremont: Shoshoni, GEB & PFT (1 d, USU). 


Tachysphex lamellatus sp.n. 


EtTyMOLoGY.—The specific name lamellatus 
is a Latin adjective derived from /amella, with 
reference to the unusually high hypostomal ca- 
rina. 

D1aAGnNosis.—Tachysphex lamellatus has a 
punctate mesopleuron, median scutal setae 
transversely oriented, and sternum I apically 
with a horizontal depression. Several other 
species share these features. Males of lamella- 
tus and sonorensis have a triangular or subtrian- 
gular clypeal lip and a nondentate inner mandib- 
ular margin; unlike sonorensis, flagellomeres II 
and IV of /amellatus are about equal in length. 
Females of /amellatus and arizonac have a pe- 
culiar, undulate-free margin of the clypeal lip, 
but they cannot be distinguished with certainty 
from each other. The hypostomal carina is un- 
usually high in some females and most males of 
lamellatus, and the gena adjacent to the hypo- 
stoma is ridged in most males. Both features are 
unique to /amellatus, and when present, distin- 
guish the species from all other Tachysphex. 

GEOGRAPHIC DISTRIBUTION.—New Mexico 
to southern California, north to southwestern 
Utah, and Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: ¢, Mexico, Sonora, 
Alamos, 25 Feb. 1963, P. H. Arnaud, Jr. (CAS Type No. 
13465). 

PARATYPES: 76 2,45 ¢, 2 Jan., 25 Feb. to 15 May (Mexico), 
10 Mar. to 25 Oct. (USA). Specimens for which institution is 
not indicated below are kept in UCD. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Paradise, LAS (1 2); Portal, JMD (1 
2); 5 mi. sw Portal, C. W. Sabrosky (1 2, USNM); same 
locality, M. Stathem (1 2); 5 mi. w Portal, V. Roth (1 6, 
NYSU); same locality, collector unknown (1 6, NYSU); 
Ramsey Canyon, Huachuca Mts., R. F. Sternitzky 3 6, 
CNC). Gila: Gila R. 3 mi. sw Christmas, FGW (1 6, UAT). 
Pima: Baboquivari Mts., Bryant (1 2, CAS), A. L. Melander 
(22,16), K. W. Radford & W. Patterson (7 2,2 d, UAT); 
Elkhorn Ranch, Baboquivari Mts., M. L. Noller, K. Roever 
(2 2, UAT); Madera Canyon, 4400 ft, Santa Rita Mts., HKC 
(1 2), ASM & WJP (1 2, WJP), FGW (3 2, | db, UAT); 
Madrona Ranger Sta., Rincon Mts., M. L. Noller, J. C. Be- 
quaert, H. Elton (2 2, | 6, UAT); Molino Camp, Santa Cat- 
alina Mts., R. S. Beal (1 ¢, CIS); Sabino Canyon, Santa Cat- 
alina Mts., RMB (1 2), R. H. & L. D. Beamer, W. LaBerge, 
C. Llang (1 2, KU), A. D. Telford (1 2, 1 36); Santa Catalina 
Mts., FGW & GDB (1 2); Tucson, Bryant (1 2, CAS), FGW, 
Malaise trap (2 2, UAT); 5 mi. N Tucson, FGW & GDB (1 
2). Pinal: Superior, Boyce Thompson Arboretum, FGW, J. 
Bequaert (1 ¢ , UAT). Santa Cruz: Florida Canyon (Santa Rita 
Mts.), D. P. Levin (1 2, UIM); Madera Canyon (Santa Rita 
Mts.), D. P. Levin (1 ¢, UIM); Patagonia, FGW & GDB (1 
2); Sycamore Canyon, Tumacacori Mts., A. & H. Dietrich 
(1 2, NYSU). Yavapai: Irving Power Sta., w Strawberry, O. 
Flint & ASM (1 6, USNM). Also Atascasa Mts. [a mistake 
for Atasco Mts.?], R. H. Crandall (1 2, 1 6, USNM). 

California. Inyo: Darwin Falls, JAP (1 ¢, CIS). Orange: 
Laguna Canyon, MEI (1 2, UCR). San Bernardino: Cajon 
Pass, J. E. Gillaspy (1 2); Mid Hills (9 mi. ssE Cima), TG (2 
2,26, TG). 

New Mexico. Catron: Glenwood, RMB (1 2). Luna: 6 mi. 
Nw Florida, J. S. Buckett, M. R. & R. C. Gardner (2 2). 

Utah. Washington: Leeds Canyon, Malaise trap (2 °, | 
3, USU). 


MExIco 

Chihuahua. 6 mi. s Encinillas, MEI (1 2, UCR). 

Hidalgo. Jackala, L. D. Beamer (1 2, KU). 

Puebla. 3 mi. NW Petlalcingo, FDP (1 3). 

Sinaloa. 9 mi. E Chupaderos, FDP, LAS (1 2, 2 d); 54 mi. 
s Culiacan, MEI (7 2, 2d, UCR). 

Sonora. Alamos, same data as holotype (9 2, 17 ¢6; CAS, 
UCD, WJP); La Aduana, FDP, LAS (8 2, 1 6; 4 2, UCR), 
LAS (4 3). 

Tres Marias Islands. Maria Magdalena I., H. H. Keifer (1 
Ns EZNS))- 


Tachysphex mirandus sp.n. 


ETYMOLOGY.—The specific name mirandus 
is a Latin word meaning wonderful, strange, sin- 
gular. 

DriAGNosis.—Tachysphex mirandus can be 
recognized by its peculiar sculpture. In both 


NEW SPECIES OF NORTH AMERICAN TACH YSPHEX 


sexes, the punctures are larger on the lateral 
clypeal section than on the adjacent frons, al- 
though the difference is slight in some speci- 
mens. The midfemur (also female forefemur) is 
alutaceous basoventrally, with only a few, 
sparse punctures. In the female, terga II—-V are 
alutaceous, glabrous (except somewhat pubes- 
cent laterally). The male can also be recognized 
by the mesally nonpubescent, largely glossy and 
sparsely punctate sterna II-VI combined with 
the nondentate inner mandibular margin, trian- 
gular clypeal lip, and the nonangular clypeal free 
margin between the lip and the lateral section. 
Subsidiary recognition features of both sexes 
are: mesopleural setae suberect (almost as in 
semirufus); horizontal part of mesothoracic ven- 
ter in most specimens glossy, sparsely punctate, 
contrasting with dull, strongly microsculptured 
mesopleuron. 

GEOGRAPHIC DISTRIBUTION.—Xeric areas of 
Nevada, Arizona, southern California, and 
Lower California. 


MATERIAL EXAMINED.—HOLOTYPE: 2, California, San 
Bernardino Co., Palm Springs, 11 Feb. 1958, A. Melander 
(USNM). 

PARATYPES: 17 2, 6 6, 20 Jan. to 14 Apr. 


UNITED STATES OF AMERICA 


Arizona. Mohave: 4 mi. s Hoover Dam, A. R. Gittins (1 
3, UIM). 


California. Fresno: Pinoche, 29 Mar. 1930, E. C. Van- 
Dyke (1 2, UCD). Imperial: 9 mi. w Coyote Wells, 26 Mar. 
1961, PDH (1 6, CIS); Ocotillo, 22 Mar. 1966, PDH (2 2, 1 
3, CIS); Yuha Desert, 15 Feb. 1948, C. D. MacNeill (1 2, 
CAS). Kern: 3 mi. NW Indian Wells, 12 Apr. 1954, J. M. 
Linsley (1 2, CIS); Short Canyon (7 mi. Nw Inyokern), 15 
Mar. 1955, PDH (1 d, CIS). Los Angeles: Little Rock, 28 Mar. 
1971, RMB (1 2, UCD), 22 Mar. 1941, Don Wasem (1 2, 
UCD). Riverside: Whitewater, 14 Apr. 1958, H. R. Moffitt (1 
2, UCD). San Bernardino: 5 mi. s Essex, 26 Mar. 1970, EEG 
& R. F. Denno (1 2, UCD); 14 mi. E Newberry, A. E. Mich- 
elbacher, 31 Mar. 1964 (1 2, CIS). San Diego: Borrego Valley, 
2 & 11 Apr., RMB (2 2; UCD, WJP); Borrego Springs, 30 
Mar. 1960, MSW (1 6, UCD). San Luis Obispo: Cuyama Val- 
ley, 30 mi. w Maricopa, 21 Mar. 1931, E. P. VanDuzee (1 
2, UCD). Tulare: Kaweah Power House, 20 Jan. 1972, F. G. 
Andrews (1 2, CSDA). 

Nevada. Nye: Mercury, N. T. S. [collector ?], 24 Apr. 1961 
(1 2, USNM), 28 Mar. 1962 (1 6, USNM). 


MExIco 


Baja California Norte. Diablo Canyon, E face of Sierra San 
Pedro Martir, 6 Apr. 1973, J. Donohoe (1 2, CIS); Upper 
Cantillas Canyon (Sierra Juarez), 19 Mar. 1967, JAP (1 °, 
CIS). 


33 


Tachysphex musciventris sp.n. 


ETYMOLOGY.—The specific name musciven- 
tris is derived from the Latin words muscus, 
moss, and venter; with reference to the dense 
pubescence on the female mesothoracic venter 
and male sterna. 

D1AGNosis.—The peculiar mesothoracic ven- 
ter of the female of musciventris is unique 
among Tachyspex: the mesothoracic venter is 
deeply sunken and densely pubescent along the 
midline on the posterior (horizontal) half. The 
mesally notched clypeal lip and the usually red 
hindleg are additional diagnostic characters. 

The males of musciventris and an undescribed 
species have a triangular clypeal lip, ill-defined 
mesopleural punctures, and velvety pubescent 
sterna III-VI (integument all or largely con- 
cealed). Unlike the other species, the vertex 
width in musciventris is 1.6—2.0 times its length, 
the vestiture is appressed on the hypoepimeral 
area, and sternum II is velvety pubescent. 

GEOGRAPHIC DISTRIBUTION.—California to 
southwestern Texas, north to southwestern 
Utah, south to northwestern Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: &, California, San 
Diego Co., Borrego, 2 Apr. 1973, R. M. Bohart (UCD). 

PARATYPES: 225 2, 216 6; 25 Mar. to 15 June, and 16-22 
Nov.; mostly UCD, but also AMNH, BMNH, CAS, CIS, 
CNC, CSDA, OSU, TG, UCR, UIM, USNM, USU, WIP. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Dragoon Mts., GDB & FGW (1 2). Gila: 
Globe, GEB (1 2). Maricopa: 8 mi. s Buckeye, MEI (3 6, 
UCR); Gila Bend, GDB & FGW (1 2); 18 mi. s Gila Bend, S. 
A. Gorodenski, JMD, MAC (1 2); 5 mi. N Mesa, A. R. Gittins 
(1 2, UIM). Mohave: 8 mi. E Mesquite, Nevada, FDP & PFT 
(3 2); 16 mi. N Wikieup, GDB & FGW (1 3). Pinal: w Stan- 
field, GDB & FGW (1 ¢). Santa Cruz: Tubac, PFT (1 3). 
Yavapai: 18 mi. N Aguila, FDP (1 2, USU). Yuma: Dateland, 
GDB & FGW (1 @); Ligurta, RMB (1 2); 15 mi. N Yuma, 
MEI, FDP (2 2); 22 mi. N Yuma, S. A. Gorodenski, JMD, 
MAC (1 9). 

California. Amador: 5 mi. E Jackson, W. E. Simonds (1 
3). Fresno: 10 mi. w Coalinga, RMB (1 ¢). Imperial: Choc- 
olate Mts., Ogilby Road, 3 mi. s junction Hwy. 78, MSW (1 
2, CSDA); Fish Cr. Mts., D. F. Hardwick (1 2); Glamis, 
RMB, PMM, FDP (4 2, 5 3); 3 mi. N Glamis, MJW (5 9); 
Palo Verde, MSW & J. S. Wasbauer, PDH (3 2); 3 mi. s Palo 
Verde, G. Tamaki (1 2 , CIS); 8 mi. s Palo Verde, C. A. Toschi 
(1 3, CIS). Inyo: Wildrose Canyon, ASM (1 ¢). Lake: Lu- 
cerne, D. J. & J. N. Knull (1 2). Kern: 14 mi. N Blackwells 
Cor., C. D. MacNeill (2 3, CIS); Iron Canyon (El Paso Mts.), 
C. A. Toschi (1 2 , CIS); 19 mi. w Shafter, RMB (1 2). Lassen: 
Bridge Cr. Camp, J. E. Gillaspy (1 ¢); Summit Camp, PDH, 
E. G. Linsley (4 ¢). Los Angeles: 2.5 mi. Pearblossom, R. W. 
Brooks (1 2); Little Rock, E. P. VanDuzee (1 2). Modoc: 


34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


Cedar Pass, D. L. Dahlsten (1 3). Monterey: Monterey, FDP 
(1 2). Riverside: Boyd Desert Research Center (4 mi. s Palm 
Desert), C. A. Toschi (2 2, CIS); 18 mi. w Blythe, RMB, D. 
S. Horning, MEI, FDP, ROS (11 2, 28 3); 10 mi. Nw Cot- 
tonwood (Joshua Tree National Monument), PDH (1 ¢, CIS); 
Deep Canyon (3.5 mi. s Palm Desert), JCH (1 2, UCR), C. 
Wilkinson (1 2); 5 mi. s Hemet, RMB (1 o); Hwy. 
74 x Strawberry Cr., EIS (1 2, CIS); Hopkins Well, PDH, 
E. G. Linsley, JAP (6 2, 11 3); Indio, J. Wilcox (1 2, OSU); 
Keen Camp in San Jacinto Mts., EIS (1 36); Millard Canyon, 
MEI (1 3); 7 mi. w North Palm Springs, TG (1 d, TG); Palm 
Canyon, EIS (1 6, UCR); Palm Springs, C. Dammers, 11 
Nov. (4 2); Pinon Flat in San Jacinto Mts., E. S. Ross (1 
2); Riverside, JCH (1 2, UCR), TG (1 36, TG); 4 mi. s Riv- 
erside, RCB, EIS (2 2, 3 ¢); Shavers Summit, MSW (2 @); 
Strawberry Canyon, W. A. Hunt (1 ¢, UCR); Thousand 
Palms, RMB, EEG, MEI, PMM, DRM, FDP, W. R. Richards 
(22 2, 51 3); 10 mi. E Whitewater RMB (1 2, 3 do). San 
Bernardino: 13 mi. E Amboy, E. G. Linsley, JWMS (13 2, 7 
3); Baker, F. G. Andrews (1 2, UCR); 4 mi. s Baker, MSW 
& J. S. Wasbauer (1 2, | 3); 3 mi. w Barstow, Brown & 
Lundgren (1 2, CAS); 2 mi. w Cajon Pass, EIS (1 3); Colton, 
E. P. VanDuzee (1 2); 5 mi. SE Hesperia, Rozen & Schrammel 
(2 2); sand dunes 7 mi. SE Kelso, MSW (1 3); Needles, JWMS 
(1 3); 2 mi. w Phelan, EIS (1 3); Red Mountain, D. F. Hard- 
wick (1 ¢); Vidal Junction, PDH (1 @). San Diego: Alpine, 
FXW (1 3, CAS); Borrego, RMB, C. Goodpasture, EEG, 
PDH, H. L. McKenzie, A. L. Melander, FDP, R. Snelling, 
MSW (97 2, 56 3); Coyote Cr. (Borrego Valley), FDP (1 
3); Coronado, F. E. Blaisdell (1 2); Scissors Crossing, R. R. 
Pinger (1 3). San Luis Obispo: 5 mi. w Nipomo, RMB, C. 
Goodpasture (5 3); 10 mi. Ww Simmler, PDH, R. W. Thorp, 
C. A. Toschi (4 2,6 3, CIS). Ventura: Hungry Valley (5 mi. 
s Gorman), PDH, JAP (7 2,73). 

Nevada. Clark: 20 mi. w Glendale, FDP & PFT (4 @). 

New Mexico. Dona Ana: 4 & I1 mi. N Las Cruces, 2. W. 
Richards (2 2); Mesilla, GEB (1 6, UCD). 

Texas. Hudspeth: Sierra Blanca, J. O. Martin (1 2). 

Utah. Washington: Santa Clara, FDP & PFT (1 2, USU). 


MEXICO 


Lower California. 7 mi. s Guadalupe, MEI (1 ¢, UCR), San 
Quintin, FXW (1 2, CAS). 

Sonora. 60 mi. £ San Luis, G. R. Ballmer (1 2, UCR), 23 
km sw Sonoita, B. & C. Durden (1 2, AMNH). 


Tachysphex occidentalis sp.n. 


ETYMOLOGY.—The specific name occidenta- 
lis is a Latin adjective meaning western; with 
reference to the geographic distribution of the 
species. 

DIAGNOsIs.—Sternal punctures of male occi- 
dentalis are well defined, about as large as those 
of the mesothoracic venter (sometimes mark- 
edly larger). Sternal punctures are similar in 
some other species (e.g., irregularis, tarsatus), 
but the clypeal lobe of occidentalis is contrast- 
ingly rounded, nonangulate laterally, and the in- 
ner mandibular margin is nondentate (or at most 
with a rudimentary tooth). The usual presence 


of a foretarsal rake is an additional recognition 
feature. 

The female of occidentalis is less distinctive. 
It is primarily characterized by the combination 
of the impunctate mesopleuron, nonridged or 
only finely ridged propodeal side, the suberect 
setae on the hypoepimeral area, and the evenly 
arcuate clypeal lip (neither emarginate mesally 
nor indented laterally). Females of several other 
species share these features: apricus, idiotri- 
chus, some krombeini, mirandus, many semi- 
rufus. Most of them have various prominent di- 
agnostic features which are absent in 
occidentalis. Furthermore, occidentalis and 
some undescribed species have the clypeal free 
margin less concave between the lobe and orbit 
than do the other species. Subsidiary recogni- 
tion feature of female occidentalis is the red 
hindfemur (partly or all) and hindtibia. 

GEOGRAPHIC DISTRIBUTION.—Xeric areas 
west of the Rocky Mountains, north to Oregon 
and Idaho, south to Arizona and Lower Califor- 
nia. 


MATERIAL EXAMINED.—HOLOTYPE: 6, California, Inyo 
Co., 12 mi. E Lone Pine, 19 May 1970, R. M. Bohart (UCD). 

PARATYPES: 53 2, 96 6, 4 Apr. to 23 July, 3 Aug., 28 Sep. 
Specimens for which institution is not indicated below are all 
in UCD. 


UNITED STATES OF AMERICA 


Arizona. Apache: Lukachukai, L. Burroughs, J. Bigelow, 
MAC (1 2, ASU). Coconino: 4.5 mi. E Moenkopi, J. H. & J. 
M. Davidson, MAC (2 2, ASU). Mohave: 8 mi. NE Mesquite 
(in Nevada), FDP & PFT (1 2, USU). Navajo: Jadito Trade 
Post, same collectors (2 2, ASU). 

California. Fresno: Jacolitos Canyon, RMB (1 2). Inyo: 
Antelope Springs, MEI (1 2, UCR), 15 mi. s Big Pine, EEG 
(1 3); Deep Springs, D. Giuliani, MSW & J. S. Wasbauer, 
MSW & J. Slansky (4 2, 13 6, CSDA; 1 3, WJP); same 
locality, RMB, BV (6 3); Lone Pine, N. W. Frazier (1 2); 3 
mi. N Lone Pine, JAP (1 3, CIS); 7.3 mi. w Lone Pine, Ball- 
mer & Bath (1 2); 12 mi. NE Lone Pine, RMB, EEG (1 2, 6 
3). Kern: 14 mi. w Shafter, RMB (1 2). Lassen: Hallelujah 
Junction, MEI (2 ¢6, UCR). Los Angeles: 1 mi. w Little Rock, 
E. G. & J. W. McSwain (1 3). Mono: Chalfant, BV (1 3); 7 
mi. Sw Lee Vining, A. D. Telford (1 2). Monterey: Arroyo 
Seco Camp, PT (1 6, USU). Plumas: Chilcoot, N. B. & W. 
M. Elliott (1 6, NYSU). Riverside: Anza, RMB (1 ¢). San 
Bernardino: Cronise Valley, JAP (1 6); Cronise Wash 15 mi. 
E Baker, WJP (1 36, WJP); Four Corners, R. W. Thorp (1 
2, CIS); Kramer Hills, PDH, G. A. Marsh, ROS (3 6d; 1 4, 
WJP); Yermo, collector unknown (1 2, CAS). San Diego: Bor- 
rego Valley, RMB (1 d6), WJP (1 2, CAS; 1 3d, WJP); Borrego 
State Park, MSW & J. S. Wasbauer (1 2, CSDA); Scissors 
Crossing, F. G. Andrews (1 2, UCR). San Luis Obispo: 10 
mi. wW Simmler, PDH (2 6, CIS). 

Idaho. Cassia: 2.5 mi. s Malta, R. L. Westcott (1 2, UIM). 
Franklin: Preston, GEB (1 6, USU). Fremont: 6 mi. Nw St. 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX 


Anthony, D. S. Horning (1 2, 3 d, UIM), R. L. Westcott (4 
3, UIM); St. Anthony Sand Dunes, L. S. Hawkins (1 6, 
UIM), R. L. Westcott (1 ¢, UIM), N. B. & W. M. Elliott (5 
2,43, NYSU; 1 2.14, WJP). Lincoln: 4.5 mi. £ Dietrich, 
R. L. Westcott (1 2, UIM); Shoshone, R. W. Haegele (1 o, 
UIM). 

Nevada. Churchill: 12 mi. NE Stillwater, FDP (3 ¢; 1 2, 2 
3, USNM). Humboldt: Orovada, MEI (1 ¢); 10 mi. N Win- 
nemucca, RMB (1 2). Lyon: Fernley, T. R. Haig (1 2); Yer- 
ington, R. W. Lauderdale (1 2). Washoe: Nixon, RMB, R. J. 
Gill, MEI, FDP, J. E. Slansky (8 2, 4 d), MEI (1 2, UCR), 
R. L. Westcott (2 2, 1 6, LACM); Patrick, FDP (1 2); 15 mi. 
E Reno, RMB, MEI, FDP (9 3); Wadsworth, FDP (2 2, | 
3); 2.8 mi. W Wadsworth, G. I. Stage (2 3, CAS). 

Oregon. Harney: 21.5 mi. NW Fields sand dunes, R. L. 
Westcott (1 2, OSDA). 


Utah. Duchesne: 5.5 mi. w Roosevelt, R. W. Thorp (1 6, _ 


CIS). Emery: Goblin Valley, FDP (2 2, USU). Grand: 25 mi. 
s Moab, GEB & R. Brumley (1 6; 3 6, USU; 1 3, WIP). 
Millard: 6 mi. N Delta, S. M. Hogue (1 2, UIM); 15 mi. N 
Delta, PFT (1 d, USU); 12 mi. Nw Fillmore, FDP & Vincent 
(2 6, USU). Washington: Santa Clara, GEB (1 2, USU). 
Wyoming. Sweetwater: 20 mi. w Farson, PFT (1 ¢d, USU). 


MExICcOo 


Lower California. Punta de Cabras (12 mi. w of km 180 s 
San Thomas), S. & S. Frommer (1 ¢). 


Tachysphex papago sp.n. 

ETyMOLOGY.—Named after the Papago In- 
dians of Arizona. 

Di1AGnNosis.—Like psilocerus, papago has the 
punctate mesopleuron, metapleuron, and pro- 
podeal side (propodeal side impunctate in some 
psilocerus), apicomesally unsculptured terga I 
and II (smooth part contrasting with remaining 
surface), and the largely brown or red female 
flagellum. Unlike psilocerus, the upper meta- 
pleuron of papago is simple, the metapleural 
flange is narrow, the malar space is absent, and 
the hindwing base is broad. The presence of 
erect setae on the midfemoral venter in papago 
(setae length about | DOA) is a subsidiary di- 
agnostic feature. 

GEOGRAPHIC DISTRIBUTION.—Southern Ari- 
zona. 


MATERIAL EXAMINED.—HOLOTYPE: 2, Arizona, Santa 
Cruz Co., Nogales, Apr. 1937, R. C. L. Perkins (BMNH). 

PARATYPES: Arizona, same data as holotype (1 ¢d, BMNH). 
Cochise: 5 mi. w Portal, 7 May 1977, collector unknown (1 
3, NYSU); 6 mi. w Portal, 7 & 12 May 1973, A. L. Steiner 
(2 2, UAE, WIP). 


Tachysphex solaris sp.n. 


ETyMOLOGy.—The specific name solaris is 
Latin adjective meaning of the sun; it refers to 
the sunny habitats in which the species occurs. 


ie) 
nA 


DIAGNOsISs.—Most solaris can be recognized 
by the uniformly yellowish humeral plate of the 
forewing base (the median plate is usually con- 
trastingly dark), but in some specimens the plate 
is partly dark. In other species the humeral plate 
is all dark or with a dark spot at the middle. 
Subsidiary diagnostic features of solaris are: 
small size (body length 5—7.5 mm); clypeal free 
margin shallowly concave between lobe and or- 
bit; vestiture largely concealing mesopleural in- 
tegument; setae oriented mainly transversely on 
propodeal dorsum; and densely pubescent male 
sterna. 

GEOGRAPHIC DISTRIBUTION.—Xeric areas of 
southern California and adjacent areas of Ne- 
vada and Arizona. 


MATERIAL EXAMINED.—HOLOoTYPE: 2, California, San 
Diego Co., Borrego Valley, 20 Apr. 1957, R. C. Bechtel 
(UCD). 

PARATYPES: 59 2, 26 d, 2 Apr. to 17 June. 

Arizona. Mohave: 8 mi. E Mesquite (Nevada), FDP & PFT 
(1 2, USU). Pinal: 5 mi. Nw Coolidge, A. D. Telford (2 6, 
UCD). Yuma: 6 mi. se Parker, S. A. Gorodenski, JMD, MAC 
(1 2, ASU); 8 mi. se Parker, J. H. & J. M. Davidson, MAC 
G 2, UCD); 1simi- = Yuma, PEL, EDP; GEB (1 2, WSU): 
18 mi. NE Yuma, FGW & GDB (1 3, UCD). 

California. Imperial: Glamis, F. G. Andrews (1 2, CSDA; 
1 2, WJP); Pinto Wash, FXW (3 3d, CAS); Palo Verde, RMB 
(1 6, UCR), MSW (1 2, 1 ¢6, CSDA; 1 3d, WJP). Inyo: 13 mi. 
s Death Valley Junction, LAS & ASM (1 ¢, UCD); Lone 
Pine, RMB (1 2, UCD); 2 mi. E Lone Pine, RMB (1 3, UCD), 
15 mi. s Panamint Springs, PDH (2 2, 2 6, UCD); s end 
Owens Lake, M.S. & J. S. Wasbauer (3 2 , CSDA). Riverside: 
18 mi. w Blythe, FDP (2 2, UCD), WJP (23 2, 3 3, CAS); 
Hopkins Well, PDH (1 2, UCD); Thousand Palms, FDP (1 
3, UCD). San Bernardino: Bagdad, JAP (2 36, CIS); Colton 
Hills, TG (1 2, TG), Cronise Valley, FDP (2 2, UCD); Kelso, 
N. J. Smith (1 2, UCD); 7 mi. sw Kelso, M. S. & J. S. 
Wasbauer (1 6, CSDA); 10 mi. E Twentynine Palms, collector 
unknown (1 3d, CSDA). San Diego: Borrego Valley, RCB, 
UClst, Tals IRS Moin, IIS (@ 2. il 5 WCDE 2B ©, WSINIME Y 
OIL 5 VKAID PE OW @ O. PD Gin CAS), 

Nevada. Clark: Glendale, FDP & PFT (1 2, 1 6, USU); 20 
mi. Ww Glendale, FDP & PFT (1 2, USU). 


Tachysphex spatulifer sp.n. 


ETYMOLOGY.—The specific name spatulifer 
is a combination of the Latin word spatula, a 
broad, flat tool for stirring or mixing, and the 
suffix -fer, a bearer; with reference to the female 
clypeus. 

DIAGNosis.—The female of spatulifer has a 
distinctive clypeus: the lip is broadened mesally, 
usually variably, obtusely dentate (including one 
median tooth). The clypeus is somewhat similar 
in crenulatus, musciventris, arizonac, and la- 


36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


mellatus, but unlike the former two species the 
clypeal lip of spatulifer is not emarginate me- 
sally, and unlike the latter two the middle scutal 
setae are oriented posterad and the mesopleuron 
is impunctate. 

The male of spatulifer shares with crenulatus 
the following diagnostic combination of char- 
acters: clypeal lip triangular, middle scutal setae 
oriented posterad, mesopleuron impunctate, 
sterna evenly punctate, with nonvelvety pubes- 
cence. Unlike crenulatus, the trochanteral 
punctures of spatulifer are subcontiguous, the 
lip corners are usually closer to the orbit than 
to each other, and often the propodeal side is 
coarsely ridged, the gastral apex is black, and 
the frontal vestiture is golden. 

GEOGRAPHIC DISTRIBUTION.—Washington to 
California, east to southern Idaho and northern 
Utah. 


MATERIAL EXAMINED.—HOLOTYPE: 2, California, Mon- 
terey Co., Arroyo Seco Camp, 11 May 1959, A. S. Menke 
(UCD). 

PARATYPES: 53 2, 85 d, 10 Apr. to 25 June (California, 
Idaho, Oregon), 5-11 Aug. (Utah, Washington). Specimens 
for which institution is not indicated below are kept in UCD. 

California. Alameda: | mi. E Mission Peak, W. W. Middle- 
kauf (1 2, 6 6; 1 d, CIS). Amador: Volcano, RMB (1 <¢). 
Contra Costa: Mt. Diablo, RMB (4 ¢; 1 ¢d, WJP), J. G. Rozen 
(1 3), collector unknown (1 6; 1 6, USU); Las Trampas 
Ridge (w Danville), FXW (1 2, 1 ¢6, CAS). Eldorado: Pla- 
cerville, GEB (1 2), D. J. Burdick (1 ¢, CIS). Fresno: Deer 
Cove Cr., TG (1 2, TG); Watts Valley, B. J. Adelson (1 9, 
CIS). Kern: Glennville, RMB (2 2); Tejon Canyon, E. C. 
VanDyke (1 2); 1 mi. E Woody, JAP, C. A. Toschi (2 ¢, 
CIS). Lake: N. Fork Cache Cr. x Hwy. 20, MEI (1 3). Mar- 
iposa: E] Portal, RMB (1 ¢); Indian Flat, RMB (1 2). Men- 
docino: Mendocino, S. F. Cook (1 2, CIS). Monterey: Arroyo 
Seco Camp, RCB, RMB, D. J. Burdick, HKC, C. Goodpas- 
ture, FDP, LAS, PFT, BV (8 2, 21 6; 2 2, 5 6, UCR; 2 
2,53, USNM; 4 6, USU; 3 2,3 6, WJP); Monterey, FDP 
(1 2,3 4). Napa: Samuel Springs, RCB (1 ¢). Placer: 4 mi. 
s Rocklin, MSW, P. Adams (1 2, 2 6, CSDA). Riverside: 
Pinyon Flat, San Jacinto Mts., E. C. VanDyke (1 2). Sacra- 
mento: Folsom, RMB (1 2), MSW (1 6, CSDA); 10 mi. NE 
Folsom, MSW (3 6); N Sacramento, PDH (7 6, CIS). San 
Diego: Sorrento, JAP (1 ¢, CIS). San Luis Obispo: 2.5 mi. 
Creston, C. A. Toschi (1 2, CIS); La Panza Camp, 12 mi. NE 
Pozo, JAP, R. W. Thorp (2 2, CIS); Pozo, PDH, JAP, C. A. 
Toschi (2 9, 2 3, CIS); 3 mi. E Pozo, S. W. Thorp (1 6, CIS); 
5 mi. E Santa Margarita, W. S. & E. S. Ross (1 2, CAS). 
Santa Clara: Mt. Hamilton, collector unknown (1 2); Los 
Gatos, J. A. Kusche (1 @). Siskiyou: Windy Camp, TG (1 
3, TG). Solano: Mix Canyon, R. B. & L. S. Kimsey (1 3). 
Stanislaus: 3.2 mi. Ww Hwy. 120 on Evergreen Road, R. W. 
Brooks (1 @). Tulare: Camp Wishon, TG (2 9, TG); Sequoia 
National Park, “‘Ash Mt. R.,’’ EIS (1 6); Tule River Indian 
Reservation, R. P. Allen (1 2, CAS); Wood Lake, N. W. 
Frazier (1 2). Tuolumne: 4 mi. E Sonora, J. G. Rozen (1 @). 


Yolo: Bear Cr. and Cache Cr. junction, ASM (1 2); Davis, 
FDP, EIS (3 2, 1 5d); Rumsey, ASM (1 @). 

Idaho. Oneida: Black Pine Canyon, Malaise trap (1 2, 1 
3, USU). 

Oregon. Umatilla: Athena, K. Gray & J. Schuh (1 2). 

Utah. Cache: W. Hodges Canyon, Knowlton & Hanson (1 
2, USU). 

Washington. Pacific: Nahcotta, collector unknown (1 °@, 
CIS). 


Tachysphex verticalis sp.n. 


ETYMOLOGY.—The specific name verticalis is 
derived from vertex, which is unusually narrow 
in this species. 

DIAGNosIs.—Tachysphex verticalis resem- 
bles many species of the brullii group (such as 
belfragei or mundus) in having a longer than 
wide vertex (as long as wide in some females) 
and a fine thoracic sculpture. Unlike these 
species, the propodeal dorsum setae of verti- 
calis are inclined obliquely anterad, and the fe- 
male tarsomere V is simple (apicoventral margin 
not produced into a lobe, claws short, not pre- 
hensile). 7. verticalis differs from other species 
of the pompiliformis group in having a uniformly 
microareolate, impunctate mesopleuron and 
propodeal side, and the contrastingly ridged pro- 
podeal hindface. The markedly convex middle 
clypeal section is also distinctive. Like crenu- 
latus and glabrior, the lip of the female clypeus 
has two lateral incisions on each side, but unlike 
these species the vertex has a shiny, median sul- 
cus which extends posterad from the postocellar 
impression. The male differs from other species 
of the group by its longer than wide vertex; like 
glabrior, idiotrichus, and irregularis, sterna II- 
VI have gradull. 

GEOGRAPHIC DISTRIBUTION.—Northern 
Mexico, southern Arizona and southwestern 
New Mexico, California and adjacent areas of 
northern Nevada; also isolated in Idaho and 
Utah. 


MATERIAL EXAMINED.—HOLoTyYPE: 2, California, River- 
side Co., 9 mi. w Beaumont, 29 July 1957, J. E. Gillaspy 
(UCD). 

PARATYPES: 78 2, 165 3, 24 May to 5 Oct., mostly UCD, 
but also AMNH, CAS, CIS, CSDA, HKT, KU, LACM, 
MCZ, NYSU, UAT, UCR, UIM, USNM, W/JP. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Box Canyon (Chiricahua Mts.), RMB (1 
2,66); Portal, H. & M. Townes (1 3); Skelton Canyon, 6 
mi. SE Apache, PDH (1 2). Gila: Gila R. 3 mi. sw Christmas, 
FGW (2 2, 1 3). Pima: Baboquivari Mts., O. C. Poling (1 
3); Gates Pass, D. Graham (1 2); 2 mi. E Robles Pass, D. 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX 


Graham (2 2, | 3d); Sabino Canyon, GEB, RMB, FDP, LAS 
(2 2, 5d); Tanque Verde, FGW (1 3); Tucson, FDP, LAS 
(2 d). 

California. Alameda: Tesla, J. E. Gillaspy (3 2). Butte: | 
mi. NE Pulga, R. L. Langston (2 ¢, CIS). Inyo: Antelope 
Springs (8 mi. sw Deep Springs), RMB, HKC, T. H. Ganten- 
bein, EEG, PMM, JAP (1 2, 8 6, CAS; 1 6, CIS; 1 , 10 
3), JAP, G. I. Stage (3 5, CIS); 3 mi. w Big Pine, LAS (1 
2); Big Pine Cr., RMB, ROS (2 2); 10 mi. N Bishop, RMB 
(1 5); Lone Pine Cr., RMB, EEG (2 2), Westgard Pass, H. 
B. Leech (1 36, CAS); 4 mi. w Westgard Pass, H. V. Daly (1 
2, CIS). Kern: 2 mi. w Frazier Park, JAP (1 6, CIS); Kern- 
ville, D. E. Hardy (1 6, KU). Lassen: Hallelujah Junction, 
RMB (1 ¢). Los Angeles: Camp Baldy, W. A. McDonald (1 
3, LACM); Crystal Lake Road, RMB (1 2); Elizabeth Can- 
yon, EIS (1 6); La Crescenta, RMB (1 3); Mount Wilson 
Road, RMB (1 2, | d); Monrovia, R. Rosay (1 2); Sangus, 
collector unknown (1 2, NYSU); Santa Susana Pass, MEI 
(2 6); Tanbark Flat, RMB (3 6), FXW (1 2, 2 6, CAS). 
Mariposa: Jerseydale, P. D. Levin (1 ¢, UIM). Monterey: Mill 
Cr. (Santa Lucia Mts.), EIS (1 ¢); Paraiso Springs, L. S. 
Slevin (2 2); 4 mi. s San Ardo, P. E. Paige (2 d); 4 mi. w 
Soledad, G. R. Ballmer (1 2, UCR). Orange: Upper Trabuco 
Canyon, E. M. Fisher (1 2). Placer: Lake Tahoe, collector 
unknown (1 2). Riverside: Anza, RCB (1 2); Banning, JCH 
(1 2); 9 mi. W Beaumont, J. E. Gillaspy, JCH, H. R. Moffitt 
(3 2,96); Pinyon Flat, P. H. Arnaud (1 6, CAS); Riverside, 
JCH (2 2, UCR); San Timoteo Canyon, MSW, R. McMaster, 
A. Hardy, J. Slansky (5 2, 12 d); Whitewater, M. J. Stebbins 
(1 3). San Bernardino: Cajon, RCB, RMB, H. R. Moffitt (5 
3); Cajon Junction, D. Burnett (1 2); Hole-in-the-Wall (Proy- 
idence Mts.), TG (1 ¢, TG); Mill Cr. Canyon, JCH (2 2); Oak 
Glen, TG (1 2, 1 ¢6, TG); Upper Santa Ana R., A. L. Melan- 
der (1 2); Wildwood Canyon, H. R. Moffitt (1 d); 3 mi. sE 
Yucaipa, TG (1 d, TG). San Diego: Laguna Mts. road, FXW 
(1 6, CAS); La Jolla, K. Corwin, P. Adams (1 2, 2 d), J. C. 
Bridwell (1 d); Julian, H. R. Moffitt (2 ¢), FXW (1 6, CAS); 
Poway, F. Blaisdell (1 ¢); San Diego, H. E. & M. A. Evans 
(2 2), H. A. Hill (1 3); Scissors Crossing, EIS (1 3); Sorren- 
to, JAP (1 2, CIS); 9 mi. s Warner Springs, RMB (3 3). San 
Luis Obispo: Creston, L. E. Guenther (1 ¢); Nacimiento Dam, 
JAP (1 2, CIS); 3 mi. Nw Paso Robles, R. L. Langston (1 
2, CIS). Santa Barbara: Bluff Camp (San Rafael Mts.), PMM, 
FDP (3 3); 3 mi. w Cachuma Lake, RMB, P. E. Paige, FDP, 
J. R. Russel (4 3), W. A. Steffan (2 2, CIS); Los Prietos, J. 
S. Buckett (1 ¢d), JAP (1 3, CIS); Santa Ynez Mts., RMB, 
PMM, ASM, FDP (6 o). Santa Clara: San Antonio Valley, 
3.5 mi. N Del Puerto Canyon road, JAP (1 2, CIS). Shasta: 
Hat Cr. P.O., E. E. Lindquist (1 2 , CIS); Redding, T. R. Haig 
(1 d). Stanislaus: Del Puerto Canyon, RMB, N. J. Smith (3 
2,36), BV (3 2,4 3). Trinity: Junction City, T. R. Haig 
(1 2,164), 3 mi. Ww Weaverville, DRM (1 6, UCD). Tulare: 
Three Rivers, H. R. Moffitt (1 ¢, UCR). Ventura: Foster 
Park, RMB, ASM (2 d); Sespe Canyon, PMM, FDP, RMB 
(3.3), W. A. Steffan (1 ¢, CIS). Also: County unknown, Oak 
Grove, J. Wilcox (1 d, OSU). 

Idaho. Owyhee: 2 mi. sw Murphy, D. S. Horning, DRM (1 
3). 

Nevada. Douglas: 3 mi. s Genoa, RMB (1 2, 3 ¢); Minden, 
RCB, RMB (7 2, | d). Storey: Geiger Summit, W. H. Lange 
(1 3d). Washoe: 54 mi. NW Gerlach, FDP (4 3). 

New Mexico. Hidalgo: Granite Gap, 18 mi. N Rodeo, RMB 
22 2,36, AMNH; 3 2, 2 6, UCD). 


37 


Utah. Grand: Moab, GEB (1 2). Washington: Leeds Can- 
yon, G. F. Knowlton (1 2, USU). 


MExICcOo 


Lower California. 10 mi. E Bahia San Quintin, FXW (1 °, 
CAS); 4 mi. s La Rumorosa, MEI (1 2, 1 6, UCR); 4 mi. 
wsw Miraflores, J. Slansky, M. K. & C. Wasbauer (1 2, 2 
3); San Quintin, FXW (1 3, CAS). 

Sonora. Cocorit, FDP (1 ¢, UCD). 


Tachysphex yolo sp.n. 


ETYMOLOGY.—Named after Yolo County, 
California, where the holotype was collected. 

D1aGnNosis.—lTachysphex yolo is character- 
ized by the transversely oriented middle scutal 
setae, sternum I with a horizontal depression at 
apex, and the usually punctate mesopleuron 
(punctures indistinct in some individuals). Other 
species share this combination of characters, but 
the female of yolo has a distinctive clypeus and 
sternum II: the dense clypeal punctation attains 
the lip base laterally, so that the sparsely punc- 
tate, apical area does not extend laterad to the 
lip corner level (its width is about 0.5—0.8 of the 
lip foremargin); the micropunctation of sternum 
II is absent along the midline from the base (or 
near base) to apex. In the other species, the 
dense punctation does not attain the lip base, 
and the sparsely punctate, apical area is as wide 
as the lip or nearly so; and the micropunctation 
of sternum II is usually absent only from an ap- 
icomedian, triangular area. The male resembles 
lamellatus and sonorensis in having nonvelvety 
sternal pubescence, but unlike these species its 
clypeal lip is not triangular. Unlike most /amel- 
latus, the hypostomal carina of yolo is not la- 
melliform, and unlike the male of sonorensis, 
the flagellomeres III and IV are about equal in 
length. Subsidiary recognition features are: me- 
sopleural punctures subcontiguous, ridges of 
propodeal side evanescent in many specimens. 


MATERIAL EXAMINED.—HOLOTYPE: 6, California, Yolo 
Co., Davis, 4 June 1961, F. D. Parker (UCD). 

PARATYPES: 312 2, 414 3, 1 Apr. to 14 Oct. Specimens for 
which institution is not indicated are all in UCD. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Bowie, A. D. Telford (3 2); Willcox, 
RMB. PDE DD tmsdales(@39eiGIS- 10R? 4 ace) 2 
USNM); | mi. s Willcox, FGW, E. Erickson (1 2, UAT); 3.5 
mi. s Willcox, E. G. Linsley (1 2, CIS). Coconino: Moenkopi, 
RMB (1 2); 3 mi. SE Moenkopi, ASM & WJP (1 d, WJP); 4.5 
mi. E Moenkopi, J. M. & M. A. Davidson & MAC (3 Q, 
ASU). Graham: s side of San Carlos Reservoir, D. & J. Schuh 
(1 36, OSU). Maricopa: Gila Bend, GDB (2 2); 5 mi. w Gila 


38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


Bend, GDB & FGW (1 2); 20 mi. s Gila Bend, R. H. & E. 
M. Painter (1 6, UAT); Sentinel, GDB (3 2). Mohave: 4 mi. 
w Chloride, GEB, PFT, FDP (3 2, 5 6, USU); 9 mi. E Oat- 
man, same collectors (5 2, 1 6, USU); 16 mi. N Wikieup, 
FGW & GDB (1 ¢). Navajo: Jadito Trade Post, J. H. & J. M. 
Davidson & MAC (1 2, ASU). Pima: Tucson, GDB (2 2, | 
3, UAT), G. & A. Ferguson (1 2), R. X. Schick (1 2, 
LACM). Pinal: 3 mi. w Oracle, FDW, Bequaert, Holler (1 
3, UAT); Sacaton, A. D. Telford (1 2, 1 d). Yavapai: 8 mi. 
N Aguila, FDP (1 2, USU). Yuma: — Aztec, FGW & GDB 
(2 2); Dateland, GDB & FGW (1 ¢); Parker, J. M. & J. H. 
Davison (1 2, 2 ¢); 8 & 18 mi. se Parker, J. H. & J. M. 
Davidson & MAC (3 2, ASU); 8 mi. sE Parker, S. A. Goro- 
denski & JMD (1 6, ASU), J. Heddock (1 2, CIS); 12 mi. s 
Parker, MAC & M. A. Mortenson (1 2, CIS); Nw Vicksburg, 
FGW & GDB (2 2, | 3); Yuma, FGW & GDB (1 3); 15 mi. 
—E Yuma, PFT, FDP, GEB (1 2, USU); 21 mi. N Yuma, MEI 
(il @)y. 

California. Alameda: Arroyo Valle, W. J. Turner (1 2, CIS). 
Colusa: 2 mi. E Colusa, MSW (1 2). Eldorado: Chile Bar, L. 
W. Quate (1 2). Fresno: 25 mi. E Fresno, PFT (2 6, USU). 
Imperial: Chocolate Mts., Ogilby road, 3 mi. s junction Hwy. 
78, MSW (9 2, CSDA); Glamis, FDP (1 ¢); 20 mi. E Glamis, 
FDP (1 2); Palo Verde, RMB, EEG, ROS (3 2, 2 6), MSW 
(1 2, 14, CSDA); 3 mi. s Palo Verde, C. A. Toschi (1 2, 
CIS); Pinto Flat, FXW (1 2, CAS); Pinto Wash, FXW (1 d, 
CAS); 20 mi. w Yuma, PFT & FDP (1 2, USU). Inyo: An- 
telope Springs (8 mi. sw Deep Springs), RMB, HKC, T. H. 
Gantenbein (6 2, 3 6; 6 2, 11 d, CAS), PDH (6 2, 13 d, 
CIS), JWMS (1 2, CIS), JAP (1 ¢, CIS); Big Pine, C. D. 
Michener (1 6, KU); Darwin Falls, RMB, EEG (1 @, 1 3); 
3.5 mi. s Death Valley Junction, N. L. Rump (1 2); 13 mi. s 
Death Valley Junction, LAS (1 3); Deep Springs, RMB, BV 
(2 2,336), MSW, MSW & J. Slansky (2 2, 13 6, CSDA; | 
3, WJP); Eureka Valley, D. Giuliani (1 2, CSDA); | mi. N 
Lone Pine, MEI (1 2, 1 d, UCR); 2 mi. E Lone Pine, RMB, 
EEG (2 2, 9 6); Owens Lake, MSW & F. G. Andrews (2 
2, CSDA); Panamint Springs, PMM (1 2), FDP (1 3); West- 
gard Pass, H. V. Daly (1 ¢, CIS); Wyman Canyon (White 
Mts.), HKC (1 3, CAS). Kern: Johannesburg, EIS (1 2); 
Kernville, T. R. Haig (1 2). Lake: N. Fork Cache Cr. x Hwy. 
20, D. Q. Cavagnaro (1 2). Lassen: Hallelujah Junction, MEI 
(2 3, UCR). Los Angeles: Huntington Park, A. Bauman (1 
2); 8 mi. N Llano, JCH, EIS (3 2). Mendocino: Navarro, N. 
B. & W. M. Elliott (4 6, NYSU); Navarro R. x Hwy. 128, 
Hendy Groves State Park, MSW (9 2, 13 6, CSDA; 1 2, 2 
3, WJP); Robinson Cr. (4 air mi. sw Ukiah), TG (3 6, TG). 
Monterey: Fort Ord, HKC (1 6, UCD); Monterey, FDP, L. 
S. Slevin (3 3); Soledad, RMB (3 6). Modoc: Adin Pass, T. 
R. Haig (1 2, CSDA). Mono: Benton Inspection Sta., RMB 
(1 3); Paradise Camp, FDP (1 ¢). Plumas: Chilcoot, N. B. 
& W. M. Elliott (1 2,5 ¢, NYSU); Halsted Campground (E. 
Branch N. Fork Feather R.), P. H. Arnaud (1 2, 1 ¢d, CAS). 
Riverside: Andreas Canyon, RMB (2 6); Anza, RMB (2 &, 2 
3); 8 mi. E Banning, R. R. Snelling (1 6, LACM); 9 mi. w 
Blythe, JWMS (1 2, CIS); 18 mi. w Blythe, RMB, JCH, D. 
S. Horning, FDP (1 2, 4 d; 2 2, WJP), WJP (1 6, CAS); 5 
mi. N Desert Center, C. D. MacNeill (2 2, CIS); 17 mi. E 
Desert Center, Rosen & Schrammel (1 2); 5 mi. s Hemet, 
RMB (1 2); Indio, PDH (1 6), MEI (1 2, UCR); Joshua Tree 
National Monument, PFT (2 6, USU); Palm Springs, JWMS 
(1 2); 2 mi. E Palm Springs, EIS (1 9); Riverside, JCH (1 
3), EIS (1 ¢, UCR); Temecula, EIS (1 3); Thousand Palms, 
RMB, EEG, H. R. Moffitt, FDP (3 2,9 3), W. R. Richards 


(1 2, CNC); Whitewater, JWMS (1 2); Wiley Well, RMB (1 
3). Sacramento: Grand I., MSW (1 2°, CSDA); Sacramento, 
RMB, FDP (3 3), MSW (4 36, CSDA); Sacramento (Sacra- 
mento R. Levee), MSW (1 3), MSW & F. G. Andrews (2 
6, CSDA). San Benito: Pinnacles, PDH (1 6, CIS). San Ber- 
nardino: | mi. s Adelanto, MEI (1 d); 10 mi. s Adelanto, J. 
A. Froebe (1 d); Bagdad, JAP (1 d, CIS); 4 & 14 mi. s Baker, 
M.S. & J. S. Wasbauer (2 6 , CSDA); Cottonwood Wash, TG 
(2 2, TG); Cronise Valley, FDP (1 2°); Cronise Wash (15 mi. 
E Baker), WJP (2 2, 5 ¢, CAS); 12 mi. sE Ivanpah, PDH (1 
2, 13); Joshua Tree National Monument, TG (1 3, TG); 
Kelso Dunes, TG (4 2, TG); Kelso Mts., TG (1 36, TG); Kra- 
mer Hills, G. A. Marsh (1 ¢, CIS), ROS (1 2); 3 mi. s Kramer 
Junction, MEI (2 3); 14 mi. s Kramer Junction. JWMS (1 
2); 20 mi. w Landers, PFT & N. Youssef (1 6, USU); 12 mi. 
ESE Tecopa, MSW, T. Eichlin (1 ¢, CSDA); 22 mi. N Manix, 
G. A. Marsh (1 2); 2 mi. w Phelan, EIS (2 6); 14 mi. w Rice, 
C. D. MacNeill (1 ¢, CIS); Twentynine Palms, R. R. Pinger 
(1 6, CSDA); 32 road mi. E Twentynine Palms, TG (1 2, 1 
6, TG); Vanwinkle Spring, G. E. Wallace (1 ¢); 11 mi. E 
Yermo, JWMS (1 2, CIS). San Diego: 8 mi. E Banner, JAP 
(2 3, CIS); Borrego Valley, RMB, EEG, PDH, FDP, EIS, 
MSW (19 2, 29 36;2 4, WJP), MEI (2 56, UCR), G. A. Marsh 
(1 2, CIS), MSW, J. Slansky, Adams (1 2, 4 6, CSDA), 
MSW (1 2,4 6, CIS), FXW (2 2, 29 6, CAS): Del Mar, C. 
H. Frady (1 2, OSU); | mi. s Del Mar, PDH (2 &, CIS); 
between Ocotillo & Borrego, A. R. Moldenke (2 6, LACM); 
Scissors Crossing, EIS (2 2), MSW (1 2, CSDA); Sorrento, 
JAP (1 2,76, CIS); 2 mi. N Warner Springs, B. M. Bartosh, 
RCB (3 2, 2 d). San Luis Obispo: Black Lake Canyon, RMB 
(1 2, 14); 10 mi. w Simmler, PDH (2 6, CIS). San Mateo: 
10 mi. sw San Francisco, W. Bohart (1 ¢, USU). Santa Bar- 
bara: 3 mi. Ww Cachuma Lake, P. E. Paige (1 2), Los Prietos, 
J. S. Buckett (1 3d), JAP (1 2, CIS); 2 mi. E Solvang, JAP 
(1 2, CIS). Santa Clara: San Jose, PFT (1 2 , USU). Siskiyou: 
between Hawkinsville & Lona Gulch, BV (1 2, CSDA). Sut- 
ter: Nicolaus, MSW (1 2, 17 6, CSDA). Trinity: Hayfork 
Agricultural Inspection Sta., JAP (2 ¢, CIS); Junction City, 
T. R. Haig (2 2, 1 6, CSDA). Ventura: Foster Park, J. L. 
Bath (1 2, UCR), J. R. Russell (1 2); Ventura, J. R. Russell 
(1 2). Yolo: Capay, R. E. Rice (1 2); Davis, RMB, C. G. 
Moore, C. R. Kovacic, ASM, L. R. Nault, FDP, WJP, LAS 
(12 2, 28 6;2 6, USNM; 2 2, 3 bd, WJP); 3 mi. s Davis, R. 
R. Snelling (1 6, LACM); Putah Canyon, FDP (1 2, 2 3); 
Rumsay, RMB (1 3), AMS & LAS (1 2, 1 d, LACM). 

Idaho, Franklin: Preston (1 2, USU). Fremont: St. Anthony 
Sand Dunes, N. B. & W. M. Elliott (1 2, 1 ¢6, NYSU). Lin- 
coln: 6 mi. NE Shoshone, A. R. Gittins (6, UIM). 

Nevada. Churchill: 23 mi. £ Fallon, E. G. Linsley (1 2); 3 
mi. w Hazen, ASM (1 2), 4 mi. E Hazen, MEI (1 6, UCR); 
Sand Mt. (9 mi. Frenchman), J. Doyen (1 2, CIS). Clark: 
Jean, GEB (1 3); 9 mi. sw Mesquite, RCB (10 2, 6 d); 30 
mi. s Searchlight, PFT, Rust, Youssef (1 ¢, USU); Valley of 
Fire, PFT, FDP, GEB (1 ¢6, USU). Humboldt: 10 mi. N Win- 
nemucca, EEG (1 2). Lyon: Weeks, FDP (1 °). Mineral: Lu- 
ning, R. F. Denno & DRM (2 3); 3 mi. sE Schurz, FDP (1 
3). Pershing: Woolsey, T. R. Haig (2 2, CSDA). Washoe: 
Nixon, RMB, FDP (1 2,4 3), MEI (1 6, UCR); Patrick, FDP 
(il Ger) 

New Mexico. Dona Ana: Las Cruces, RMB (1 @, 2 6), R. 
H. Beamer (1 2, KU); 4 mi. E Mesilla Park, PDH (1 ¢, CIS). 
Lincoln: 5 mi. s Oscuro, R. L. Westcott (1 2, UIM). Otero: 
Alamogordo, collector unknown (1 ¢, CU); White Sands Na- 
tional Monument, H. V. Weems (1 2, FSCA). Socorro: La 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX 


Joya, 20 mi. N Socorro, H. E. Evans (6 ?, 3 6, CSU). 
Valencia: Acoma Pueblo, ASM & WJP (1 2, WJP). 

Oregon. Deschutes: Smith Rock State Park, M. B. & W. M. 
Elliott (8 2, 15 ¢, NYSU). Josephine: 8 mi. w Grants Pass, 
R. L. Westcott (1 2, ODA). Klamath: Bonanza, J. Schuh (1 
2). Morrow: Boardman, G. R. Ferguson (2 ¢). Umatilla: Hat 
Rock State Park, EIS (1 @). 

Texas. Hudspeth: McNary, H. E. Evans (2 2, 1 d¢, CSU); 
Sierra Blanca, RMB (1 2). Presidio: 5 mi. E Presidio, D. S. 
Horning (1 ¢). 

Utah. Cache: Cornish, GEB & PFT (5 2, 2 d, USU; 2 @, 
WJP). Emery: Goblin Valley, FDP (1 6, USU), 2 air mi. w 
Little Gilson Butte, TG (1 6, USU). Juab: 12 mi. s Eureka, 
JWMS (1 2, CIS); White Sand Dunes (25 mi. sw Eureka), W. 
F. Barr (1 ¢, UIM). Millard: 15 mi. N Delta, PFT (1 ¢ , USU); 
Pahvant (near Flovell), GEB & E. A. Cross (1 6, UCD). San 
Juan: 6 mi. s La Sal Junction, R. W. Thorp (1 ¢); 25 mi. s 
Moab, GEB, R. Brumley (2 2, 1 6; | 2, USU). Washington: 
Leeds Canyon, G. F. Knowlton, W. J. Hanson, T. H. Hsiao 
(i 2,26, USU). 


MExICO 


Lower California. 10 mi. E Bahia San Quintin, FXW (1 6, 
CAS); Descanso, RMB (1 2); El Pescadero, MSW, J. Slansky 
(3 2,66, CSDA); La Paz, FXW (2 6, CAS); Los Barmiles, 
MSW (3 2, CSDA); 20 mi. N Mesquital, Ross & Bohart (1 
2); Progreso, Sierra Juarez, FXW (1 2, 3 d, CAS); 38 km s 
Rosarito (114°), E. Fischer, R. Westcott (1 2, CAS); San Car- 
los, H. E. Evans, W. Rubink & D. Gwynne (4 2, CSU); 3 
mi. N San Felipe, MEI (3 2, 1 6, UCR); 15 mi. N San Ignacio, 
Ross & Bohart (1 2); 10 mi. s San Quintin, J. Slansky, M. 
& K. Wasbauer (1 2, CSDA); San Vicente, C. H. Frady (1 
2, OSU), JAP (1 &, CIS). 

Chihuahua. Moctezuma, JWMS (1 6, CIS); Samalayuca, 
RMB (2 @). 

Sinaloa. 8 mi. s Elota, LAS (1 ¢). 

Sonora. Cerro Pinacate, McDougal Crater, GDB (1 ¢, 
UAT); Guaymas, E. P. VanDuzee (1 2); 39 mi. s Puerto Pe- 


nasco, M. Leppla, J. Bigelow, MAC, J. Davidson (1 2, ASU); 


‘*Sonora, 85 km. so.,’’ A. L. Melander (1 2, 6 3); Tepoca 
Bay, E. P. VanDuzee (1 @). 


Tachysphex yuma sp.n. 


ETYMOLoGy.—Named after the Yuma Indi- 
ans of Arizona. 

DIAGNOosIS.—Tachysphex yuma is character- 
ized by the punctate mesopleuron (punctures 
shallow), middle scutal setae oriented postero- 
laterad or (some males) posterad, and sternum 
I with an apical depression. Several other 
species share this combination of characters, but 
they have a uniformly ridged propodeal side 
(ridges evanescent in many yolo). In yuma, the 
propodeal side is either coarsely ridged poste- 
riorly and microridged along the metapleural 
sulcus; or (most specimens) nonridged along the 
metapleural sulcus and ridged along the dorsal 
margin (or dorsal and posterior); or (some 
males) all nonridged. Furthermore, the flagellum 


39 


of yuma is somewhat longer; for example, the 
length of flagellomere IV is 3.64.2 (female) and 
2.0—2.4 (male) times its width, and up to 3.2 and 
2.0 times, respectively, in the other species. In 
the male, the unusually broad clypeal lobe is 
distinctive (corners markedly closer to orbits 
than to each other) and the velvety sternal pu- 
bescence is an additional recognition feature. 

GEOGRAPHIC DISTRIBUTION.—Idaho, Ore- 
gon, California, Arizona, southern Texas, north- 
ern Mexico (Sonora, Lower California). 


MATERIAL EXAMINED.—HOLoTYPE: ¢, Mexico, Baja Cal- 
ifornia Sur, La Paz, 10-12 Oct. 1954, F. X. Williams (CAS 
Type No. 13966). 

PARATYPES: 8 ¢, 25 5, 2 Mar. (Sonora), 8 Apr. to | June, 
14 July (Oregon), 9-15 Oct., 11 Nov. 


UNITED STATES OF AMERICA 


Arizona. Cochise: Canelo, A. & H. Dietrich (1 2, NYSU); 
5 mi. w Portal, collector unknown (2 6, NYSU); 6 mi. w 
Portal, A. L. Steiner (2 ¢, UAE, WJP). Maricopa: Wicken- 
burg, PFT & GEB (1 6, WJP). Coconino: Grand Canyon Na- 
tional Park, 15 mi. NE Phantom Ranch, J. E. Slansky (1 6, 
UCD). 

California. Imperial: 20 mi. E Glamis, FDP (1 ¢, UCD). 
Inyo: Darwin Falls, ASM (1 ¢, UCD); Panamint Springs, MEI 
(1 6, UCD); Surprise Canyon, FDP (2 ¢, UCD, WJP). Riv- 
erside: Boyd Desert Research Center, 4 mi. s Palm Desert, 
PDH (1 2, 1 6, CIS; 1 2, WJP); Deep Canyon, MEI (3 6, 
UCD, UCR, WJP), EIS (1 6, UCD); Salton Beach, A. L. 
Melander (1 2, UCD); Thousand Palms Canyon, RMB (1 
6, UCD); Whitewater, D. J. R. (1 d, CIS). San Bernardino: 
3 mi. N Crossroads, C. D. MacNeill (1 ¢, CAS). Shasta: Red- 
ding, DRM (1 2°, UCD). Stanislaus: Empire, E. I. Beamer (1 
OPN): 

Idaho. Twin Falls: Rock Cr. Canyon (19 mi. s Hansen), R. 
L. Westcott (1 2, UIM). 

Oregon. Malheur: 4 mi. N Juntura, H. A. Scullen (1 d, 
UCD). 

Texas. Brewster: 20 mi. NNW Marathon, M. Masters (1 6, 
CU). 


MExICcO 


Baja California Norte. 65 mi. s San Felipe, R. D. Gehring 
(1d, GIs). 

Baja California Sur. 3 mi. s Ignacio, MEI (2 6, UCR, WJP), 
Rancho El Cayote, Maynard & Honey (1 ¢, LACM). 

Sonora. Bahia San Carlos, P. H. Arnaud (1 2, CAS). 


SPECIES OF THE BRULLII GROUP 
Tachysphex acanthophorus sp.n. 


ETYMOLOGY.—The specific name acantho- 
phorus is derived from the Greek words acan- 
thos, aspine, and phorein, to bear, a spine-bear- 
er; with reference to the tarsal spines. 

DiaGnosis.—The female of acanthophorus 
has a distinctive tarsal feature: one or two sub- 
apical spines on each lateral margin of tarsomere 


40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


V, and usually central spines on its venter; un- 
like armatus, the tarsomere V lacks basoventral 
spines. The lateral, subapical spines on tarso- 
mere V occur also in most western females of 
mundus, but unlike that species the mesopleural 
vestiture is dense (partly obscuring sculpture) in 
acanthophorus. Tarsomere V is similar in acan- 
thophorus and the South American species spi- 
nulosus, but in the latter the gaster is black and 
the mid- and hindfemora are red. 

The males of acanthophorus and armatus dif- 
fer from other species of the brullii group in 
lacking a clypeal bevel and graduli, and in hav- 
ing a very narrow clypeal lip and a transverse 
sulcus on sterna III—VI (the sulcus is visible only 
when segments are fully extended). Unlike ar- 
matus (in which the sternal surface is flat), ster- 
num II of acanthophorus is somewhat swollen 
along the foremargin of the apical depression. 
Furthermore, the mesopleural sculpture is ob- 
scured by vestiture or hardly visible in acantho- 
phorus, but easily visible in armatus. 

GEOGRAPHIC DISTRIBUTION.—Xeric areas of 
southwestern U.S. and northern Mexico. 


MATERIAL EXAMINED.—HOLOTYPE: @, Arizona, Cochise 
Co., Willcox, 14 Aug. 1958, P. D. Hurd (UCD). 

PARATYPES: 134 2, 180 4, | intersex; May to 7 Nov. Spec- 
imens for which institution is not indicated are all in UCD. 


UNITED STATES OF AMERICA 


Arizona. No specific locality (1 2). Cochise: 1 mi. E Apache, 
J. G. Ehrenberg (2 6, AMNH); Bowie, A. D. Telford (1 °, 
WJP); 7 mi. sE Dos Cabezas, D. S. Chandler (1 ¢, UAT); 
Douglas, G. L. Ballmer (2 2); 2 mi. E Douglas, R. W. Brooks 
(1 2); Portal, K. V. Krombein (28 9, 22 ¢; BMNH, KVK, 
WJP); 2 mi. E Portal, J. H. Puckle, M. A. Mortenson & MAC 
(2 3); 2 mi. NE Portal, M. & T. M. Favreau (1 3d, AMNH); 
Willcox, RMB (2 2,2 ¢), D. D. Linsdale (1 2,5 36; AMNH, 
UCD), PDH (1 2, 2 6, CIS; 1 2, USNM). Maricopa: 5 mi. 
N Aguila, GEB & PFT (1 2, USU); Tempe, collector un- 
known (1 6, MCZ); 5 mi. se Wickenburg, PDH (1 2, CIS). 
Pima: 30 mi. sE Ajo, C. R. Kovacic (1 2); Continental, MEI 
(2 3); Sabino Canyon (Santa Catalina Mts.), GDB & FGW 
(1 6, UAT), FDP & LAS (1 2); Tucson, C. L. Crow (1 2), 
M. L. Lindsay (1 6, UAT), FDP (1 3), FDP & LAS (1 @). 
Pinal: 5 mi. Nw Coolidge, A. D. Telford (1 ¢); Picacho Pass, 
DRM & J. E. Lauck (1 2); collector unknown (1 2, UCR); 
Superior (Boyce Thompson Arboretum), GDB (1 ¢, UAT). 

California. Imperial: Brawley, GEB (1 @). Inyo: Antelope 
Springs (8 mi. Sw Deep Springs), HKC (13 2, 11 6; AMNH, 
UCD), PMM (3 2), DRM (1 @, | od), JAP, G. I. Stage (11 
2, 48 3, CIS); Deep Springs, H. Nakakihara (1 2, UCR). 
Riverside: 12 mi. N Blythe (also 18 mi. w), RMB (2 @); 20 mi. 
w Blythe, JWMS (3 2, 1 ¢, CIS); Indio, MEI (1 6, UCR); 
3.5 mi. s Palm Desert, S. Frommer & B. Morley (1 2, 16, 
UCR). San Bernardino: no specific locality, D. W. Coquillett 
(1 ¢, USNM). San Diego: Borrego, A. L. Melander (1 @). 


Tulare: Lemon Cove, J. C. Bradley (3 2, NYSU; 4 6, CU); 
Three Rivers, collector unknown (1 6, CU). 

Colorado: Bent: Hasty, H. E. Evans (9 9, 2 6; BMNH, 
CSU, WJP). 

Nevada. Clark: Sandy, RCB (1 3). Mineral: Luning, R. F. 
Denno & DRM (1 2). Nye: Mercury (1 ¢, USNM). 

New Mexico. Dona Ana: Las Cruces, RMB (1 2). Eddy: 
15.5 mi. w Artesia, V. E. Romney (1 ¢, USNM). Grant: 25 
mi. E Lordsburg, H. A. Scullen (1 3). Hidalgo: 21 mi. s An- 
imas, J. G. & B. L. Rozen (1 2, AMNH); 22 mi. s Animas, 
J. Rozen & M. Favreau (1 6, AMNH); Cienaga Lake, J. H. 
& J. M. Davidson & MAC (2 6); Granite Gap (17 mi. N Ro- 
deo), F. G. Andrews (1 2, CSDA); Granite Pass area (20 mi. 
N Rodeo), Hwy. 80, MSW, J. Slansky & C. Freeberg (1 2, 
CSDA); 1 mi. N Rodeo, J. H. Puckle, M. A. Mortenson & 
MAC (1 3); 4 mi. sw Rodeo, J. G. Rozen (1 2, AMNH). 
Otero: White Sands National Monument, RMB (1 ¢). Socorro: 
La Joya (20 mi. N Socorro), W. Rubink (1 2, CSU; 2 ¢, 
USNM). Quay: Tucumcari, RMB (2 ¢). 

Texas. Bexar: no specific locality, H. B. Parks (3 2; UCD, 
WJP). Brewster: Big Bend National Park (Nine Point Draw), 
W.R. M. Mason (4 2, 22 6; BMNH, CNC, WJP); Big Bend 
National Park (Boquillas), W. R. M. Mason (1 intersex, CNC); 
Glenn Spring, F. M. Gaige (1 3). El Paso: Sierra Blanca, col- 
lector unknown (1 6, CU). Hudspeth: Fort Hancock, CU 
Exped. (1 2, NYSU); McNary, H. E. Evans (1 2, 1 6; 
BMNH, CSU). Presidio: 3 mi. E Presidio, H. E. Evans (2 
2, MCZ), J. E. Gillaspy (1 2). 

Utah. Garfield: Shootaring Canyon, D. Vogt (1 2, 1 6, 
USU). Washington: Leeds Canyon, G. F. Knowlton, W. 
J. Hanson, T. H. Hsiao (2 2, 1 6, USU); St. George, GEB 
(1 3); Toquerville, G. F. Knowlton, W. J. Hanson, T. H. 
Hsiao (2 6, USU). 


MExICcOo 


Baja California Sur. 4 mi. wsw Miraflores, J. Slansky, M. 
K. & C. Wasbauer (1 2, 4 6, CSDA). 

Chihuahua. 15 mi. s Chihuahua, H. E. Evans (2 9, 4 6; 
BMNH, MCZ, WJP). 

Sinaloa. Culiacan, H. E. Evans (1 ¢, CU); s Lorenzo, GEB 
& RMB (2 2, USU); Mazatlan, W. R. M. Mason (1 2, CNC); 
2.5 mi. N Mazatlan, MSW (1 2, CIS); 5 mi. N Mazatlan, MSW 
& J. Chemsak (1 ¢, CIS). Sonora. Alamos, RMB (1 6d), W. 
J. Hanson & T. L. Whitworth (1 ¢, USU); 10 mi. sw Alamos, 
FDP & LAS (1 3); Desemboque, C. & P. Vaurie (1 d); 19.4 
& 20 mi. s Estacion Llano, MEI, EIS, P. A. Rauch (3 6, 
UCR); Guaymas, RMB (1 @); 5 mi. s Magdalena, FDP & LAS 
(1 3); Minas Nuevas, C. & P. Vaurie (1 ¢); 10 mi. E Navajoa, 
W. L. Nutting & FGW (2 ¢, UAT); San Carlos, RMB (1 @, 
lS) 


Tachysphex armatus sp.n. 


ETYMOLOGY.—The specific name armatus is 
a Latin word for armed, with reference to the 
ventral spines of female hindtarsomere V. 

DIAGNosis.—The female of armatus is unique 
among the North American Tachysphex in hav- 
ing basoventral spines on the tarsomere V. Oth- 
erwise it is very similar to acanthophorus with 
which it shares other structures of the tarsomere 
V: one to several medioventral spines and one 


NEW SPECIES OF NORTH AMERICAN TACHYSPHEX 


or two preapical spines on each lateral margin. 
The lateral spines are also found in most western 
specimens of mundus. 

The male of armatus is very similar to acan- 
thophorus. See that species for differences. 

GEOGRAPHIC DISTRIBUTION.—Desert areas 
between southwestern Texas and southern Cal- 
ifornia, also Lower California. 


MATERIAL EXAMINED.—HOLOTYPE: 2, Nevada, Clark 
Co., Sandy, 24 July 1958, R. C. Bechtel (UCD). 
PARATYPES: 17 2, 9 5; May to 5 Sep. 


UNITED STATES OF AMERICA 


Arizona. Maricopa: 10 mi. E Gila Bend, GDB (1 2, UCD); 
Phoenix, R. H. Crandall (1 2, UCD); 5 mi. sE Wickenburg, 
PDH (1 2, CIS), P. H. Timberlake (1 ¢ , UCR). Pima: Tucson, 
F. M. Carpenter (1 2, UCD). 

California. San Diego: San Diego, F. E. Blaisdell (1 2, 
UCD). 

Nevada. Clark: Sandy, RCB (1 ¢, WJP). Lincoln: Alamo, 
FDP (1 2, WJP). 

Texas. Brewster: Big Bend National Park, R. W. Strandt- 
mann (1 2,1 d¢6;USNM, UCD); Big Bend National Park (Nine 
Point Draw), W. R. M. Mason (1 2, CNC). Hudspeth: 
McNary, H. E. Evans (2 2, 3 6; CSU, WJP). 

Utah. Washington: Leeds Canyon, G. F. Knowlton, W. J. 
Hanson, T. H. Hsiao (1 2, 3 6, USU). 


MEXICO 


Lower California. 7 mi. sw La Paz, J. A. Chemsak (1 2, 
CIS), 220 km s Tijuana, FXW (5 2, CAS). 


Tachysphex krombeiniellus sp.n. 


ETyMOLoGy.—Named after K. V. Krombein 
as a mark of friendship and gratitude. 

DIAGNosis.—Tachysphex krombeiniellus is 
similar to belfragei in having a bicolored gaster 
and short thoracic vestiture which does not con- 
ceal mesopleural sculpture. Unlike that species, 
the propodeal dorsum of krombeiniellus is 
evenly microareolate, and at least the apical 
third of the hindfemur is red. It differs from 
maurus and mundus by the shape of the clypeus 
(female lip broadened, male middle section 
slightly longer than wide). It can also be distin- 
guished from marurus and most mundus by its 
basally red gaster combined with the partly or 
all red hindfemur. 

GEOGRAPHIC DISTRIBUTION.—Mainly central 
U.S. between northern Texas and North Da- 
kota, west to 105th meridian, but also South 
Carolina, Arkansas, and Florida. 


MATERIAL EXAMINED.—HOLOTYPE: @ , Florida, Levy Co., 
no specific locality, 3 June 1954, H. V. Weems (USNM). 


41 


PARATYPES: 22 2, 14 ¢; June to Sep. Specimens for which 
institution is not given below are all in UCD. 

Arkansas. Mississippi: no specific locality, J. C. Nickerson 
Ge): 

Colorado. Yuma: Yuma, collector unknown (1 ¢d; | 6, 
WIP). 

Florida. Gadsden: Quincy, Malaise trap (2 2, UFG, WJP). 
Levy: no specific locality, H. V. Weems (1 2, CU;2 ¢, FSCA; 
12,16, USNM; 2 6, WIP). 

Kansas. Graham: Hill City, RRD (1 2). Pottawatomie: 
Blackjack, H. E. & M. A. Evans, C. S. Lin, C. Yoshimoto 
(2 6, MCZ). 

Minnesota. Scott: Barden (between Savage & Shakopee), 
C. E. Mickel (1 29, UMSP). Goodhue: Cannon Falls, C. E. 
Mickel (1 2, UMSP). 

Nebraska. Blaine: Halsey & Dunning, RRD (2 2). Box 
Butte: Alliance, RRD (1 2). Dawson: Gothenburg, RRD (1 
3). Douglas: Omaha, collector unknown (1 2). Hall: 6 mi. w 
Cairo, C. W. Rettenmeyer (1 d). Lancaster: Lincoln, collector 
unknown (1 2). Lincoln: North Platte, R. K. Schwab (1 9; | 
2, WJP). Nance: Genoa, R. M. Barnes (1 2, INHS). Thomas: 
Thedford, RRD (3 2; 1 2, WJP). 

North Dakota. Richland: 11 mi. w Walcott, J. R. Powers 
2, Elis) 

South Carolina. Aiken: New Ellenton, A. Hook (1 °, 
UGA). 

Texas. Potter: 5 mi. N Amarillo, D. R. Miller (1 ¢d, USNM). 

Wisconsin. Vernon: Genoa, collector unknown (2 °, 
MPM). 


Tachysphex menkei sp.n. 


ETyMOLoGy.—Dedicated to A. S. Menke as 
a mark of gratitude for his help. 

D1aGNosis.—lachysphex menkei can be eas- 
ily recognized by the woolly setae of the head 
and thorax; the finely, sparsely punctate scu- 
tum, scutellum, mesopleuron, and propodeal 
side; the largely impunctate, red gaster; and gla- 
brous male sterna III-VI. 

NATURAL History.—A female paratype is 
pinned with her prey, a young nymph of a long- 
horned decticine grasshopper, probably Ere- 
mopedes sp., det. A. B. Gurney. 

GEOGRAPHIC DISTRIBUTION.—Desert areas 
from southwestern Texas to southern California. 


MATERIAL EXAMINED.—HOLorTyPE: 2, California, San 
Diego Co., Borrego Valley, 19 Apr. 1957, R. M. Bohart 
(UCD). 

PARATYPES: 8 2, 32 d, Apr. to June. 

Arizona. Graham: 18 mi. E Stafford, FGW & GDB (2 6, 
UCD). Maricopa: 30 mi. £ Gila Bend, R. F. Smith (1 ¢, UCD). 
Yavapai: 10 mi. Nw Congress, FDP & LAS (1 2, UCD). 
Yuma: 5 mi. sE Bouse, S. A. Gorodenski, JMD, MAC (1 6, 
ASU). 

California. Imperial: 30 mi. NE Glamis, R. R. Pinger (2 ¢, 
CSDA). San Bernardino: Adelanto, MEI (1 ¢, UCD). San 
Diego: Borrego Valley, H. R. Moffitt, EIS (2 ¢, UCD). Riv- 
erside: Palm Springs, RMB (1 ¢, UCD); Andreas Canyon, 
RMB, HKC (10 ¢, UCD, USNM, WJP). 


42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3 


a’ 
e a * 
—_ 


“i 
New Mexico. Otero: Alamogordor€olle¢ arittichawis Gina 
3, ANSP). Socorro: Bernardo, A. ‘a 2, ‘USU). 


o> LITERATURE CITED 

Bouart, R. M., AND A. S. MENKE. 1976. Sphecid wasps of 
the world. A generic revision. University of California 
Press, Berkeley, Los Angeles, London. | color pl., ix + 
695 p. 

PULAWSKL, W. 1971. Les Tachysphex Kohl (Hym., Spheci- 
dae) aia région paléarctique occidentale et centrale. Pan- 
stwowe Wydawnictwo Naukowe, Wroctaw. 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. 


Texas. Brewster: Alpine, J. Gillaspy (1 6, MCZ); ‘Big Bend 
National Park (Nine Point Draw), R. Mason, J. F. McAlpine 
(3 2,74; CNC, WJP); Big Bend National Park (Santa‘Elena 
Canyon), J. F. McAlpine (1 6, CNC); Big Bend National Park 
(near Doughout Well), B. J. Adelson (2 5, UCD). El Paso: 
Tornillo, H. E. Evans & Rubink (1 2, WJP). Hudspeth:)+ . 
McNary H. E. Evans (1 2, CSU). Presidio: 3 site 
H. E. Evans (1 2, MCZ). Werte 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENC 


Vol. 43, No. 4, pp. 43-58, 27 figs. 


ES _ 


296 atory 
YF 4 
November 4, 1982 


el 


' 


amt. A 
- & F 


> a ale 2 
JC 
' 


| 


A REVISION OF THE GRASSHOPPER GENERA 
CHROMACRIS AND XESTOTRACHELUS. ~ 
(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 


The grasshopper species of the genus Chro- 
macris are of a striking color, usually a glossy 
green with yellow markings, and red or yellow 
wings. They occur in the humid areas of the 
American tropics from Mexico to Argentina. As 
a general rule, but one species occurs at any one 
locality. Adults are usually seasonal in their ap- 
pearance, so that some months of the year a 
species may appear to be absent from a local 
fauna. Because most of the taxa of the genus 
have been poorly defined or understood, the ap- 
plication of a number of their names has been 
uncertain, and there has been no comprehensive 


* Bolsista, Conselho Nacional de Desenvolvimento Cienti- 
fico e Tecnologico, Brazil. 


[43] 


treatment of the genus, a revisionary study ap- 
peared to be needed for this common and wide- 
spread group of grasshoppers. The monotypic 
genus Xestotrachelus of southern South Amer- 
ica is included in this study because of its sim- 
ilarity to Chromacris and because it is the only 
closely related genus. 

The subfamily Romaleinae currently includes 
about 48 genera. Rehn and Grant (1959) erected 
16 tribes in this subfamily and proposed the tribe 
Chromacrini for the genera Chromacris and 
Xestotrachelus. Because 10 of their tribes in- 
clude but one or two genera each, and they give 
no distinguishing characters for these tribes, it 
does not appear useful to recognize the tribe 
Chromacrini and other such tribes of the 
subfamily. 


44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


é = 
/ = 
a& ; 
J 4 
j é 4 
a ges 
\ Wo 
4 
4 
\ 1 
% 
4 fy ,_ 10mm —— 


FIGURE 1. 


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. psittacus 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, Muséum 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- 


Chromacris speciosa 


Chromacris speciosa, Casupa, Florida, Uruguay. 


camps and Christiane Amedegnato, MNHN;; Dr. 
Kurt K. Gtinther, 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 
Chromacris minuta n.sp. 
Chromacris miles (Drury) 
Chromacris speciosa (Thunberg) 


Acridium xanthopterum Hahn 
Rhomalea stolli Pictet and Saussure 


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 

0 


WwW 


‘ni 


. Chromacris psittacus pacificus n.subsp. 

. Chromacris icterus (Pictet and Saussure) 
Rhomalea opulenta Gerstaecker 

11. Chromacris peruviana (Pictet and Saussure) 

12. Xestotrachelus robustus (Bruner) 


Xestotrachelus hasemani Bruner 


ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS 45 


2(1): 


FIGURE 2. 


KEY TO SPECIES OF CHROMACRIS 


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 
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 


Yellow on part of antennae. Stridulat- 
ing structures weakly developed (Fig. 
HS) tae ee Ee BO 3 
Antennae entirely black. Stridulating 
structures well developed (Figs. 16— 
7, ea = Mee 6 Ls oa et a nS ele eee 4 


Basal portion of antennae yellow 
(Mexico to Costa Rica) ___ C. colorata 
Distal portion of antennae yellow 
(Acapulco, Mexico) C. minuta 


. Membrane of tegmina dark brown to 


black with strongly contrasting straw- 
colored veins (inland south central 
nazi eee Ak C. nuptialis 
INGt aS abOVewme- ke. eee... 5 


10mm 


Xestotrachelus robustus 


5(4’). 


907): 


Xestotrachelus robustus, Chapada dos Guimaraes, Mato Grosso, Brazil. 


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 
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) 


Bee eeee 6 8 Sone ne eae eee een C. miles 
. Antennae-entirely black. === - | 
Antennae with yellow tips _____________- 10 
No! bands-:onthind tibiae = 8 
Yellow bands on hind tibiae __-____- 9 


Yellow bands lacking on all legs (Cos- 
ta Rica) jpesee 2 eee a! C. trogon trogon 
Yellow bands on hind femora (Guate- 
mala, Belize, and Honduras) 
C. trogon intermedia 


Proximal yellow band on hind femora 
interrupted on outer ventral portion 
(Costa Rica to northern Colombia and 
Venezuela) == C. psittacus psittacus 
Proximal yellow band on hind femora 


46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


3 


C. colorata 


4 


C. minuta 


8 


C. psittacus 


5 


C. miles 


9 


C. icterus 


6 


C. speciosa 


10 


C. peruviana 


X. robustus 


Ficures 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 (11), all 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; (8) psittacus Cabima, Panama; 
(9) icterus Tabatinga, Amazonas, Brazil; (10) peruviana Satipo, Junin, Peru; (11) X. robustus Cerro Cora, Amambay, Paraguay. 


ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS 47 


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.—I wo groups of Chroma- 
cris can be recognized. One group including fro- 
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 (1A) and second (2A) 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, 


48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


TaainT erat! 
Reh WAS 


Zar TTT ere 


Sami ae] K 
[ |*Rs 

ae 

eel 


e277 


ideal 


i Was 


YD GD LD ED BY LD 
EGGS GSe 


TA 

SV-K 

TA 
-Cul 
-1A 
2A 
m2Aa 


Figures 12-14. 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. 


Ficures 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; 1A, 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 49 


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. 

Foop PLANTs.—Chromacris speciosa prefers 
solanaceous plants. Turk and Barrera (1976) re- 
ported its feeding on Cestrum parqui, C. stri- 
gillatum, C. lorentziana, Lycium cestroides, 
Solanum 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 
Solanum 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. ] 


DriaGNosis.—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 


50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


19 


C. colorata 


20 


C. speciosa 


21 


X. robustus 


0.5mm 


Ficures 19-25. 


25 C. peruviana 


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 robustus 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), 65, 32. 22 mi. [35 km] Nw Poptun, 15 Jul. 1974 
(J. C. Lee), 16, 32. 

Belize. Rio Grande, Aug. 1931 (J. J. White), 3°. 

Nicaragua. MANAGUA: Sep. 1955 (E. Morales-Agacino), 
Mein, WE (CNCe 

Costa Rica. GUANACASTE: Canas, Sep. 1965 (C. H. F. Row- 
ell), 16, 12, CHFR. 


COMMENTS.—Specimens from Volcan Colima 
(Jalisco, Mexico) and Guanacaste (Costa Rica) 
agree with Serville’s description of Acridium 
coloratum 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 51 


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), 7d, 12. Same data as 
for holotype, 6d, 82, 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 
Rio 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, pl. 42, fig. 2. [Holo- 
type, 2, 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. 1960 (P. M. Litchfield), 1¢, 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, 2, 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, d, 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, 


52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


TABLE |. 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 0 
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 
Bolivia 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), 86, 3°; (3 lack proximal tibial band). 

Venezuela. CARABOBO: San Esteban, Nov.—Dec. 1939 (P. 
Van Duse), 6d, 72. CosEDES: 26 km s jct. Rts. 8 and 13 on 
Rt. 8, forest, 13 Jul. 1981 (Otte et al.), 1d, 3°. 

Guyana. Bartica, Dec. 1912 (H. S. Parish), 30d, 35°. 

Brazil. AMAPA: Rio Puxaca, Mazagao, Feb. 1961 (J. C. M. 
Carvalho), 32, UMMZ. Eighteen specimens from Belém, 
Santarém, Obidos, and Manaus on the Amazon are similar to 
the Guyana series. PARA: Jacareacanga, 6°16’S, 57°44’W, 
Dec. 1968 (Alvarenga), 3d, 92, UMMZ. Banta: 100 km Nw 
Feira de Santana, 13 Mar. 1981 (Roppa, Carbonell, Roberts), 
2d, 2; Itabuna, research center, 22 Nov. 1974, cacao forest 
(Roberts, Carbonell), 12; Mucuri, Aug. 1977 (Roppa, Beck- 
er), 2d (one yellow wings, other orange wings). Espirito 
SANTO: Itapemirim falls, 1-6 km E BR. 101, edge of forest and 
marsh, 5 Dec. 1974 (Roberts, Carbonell), 42 (lack usual yel- 
low median spots on lateral lobe of pronotum); 3 km s Lin- 
hares, cacao forest, 1 Dec. 1974 (Roberts, Carbonell), 2d, 12 
(lack usual yellow spots on lateral lobe of pronotum). R1o DE 
JANEIRO: Floresta de Tijuca, Jan. 1981, 4d, 62 (yellow 
wings), 56, 62 (orange wings); BR. 101, 1 Feb. 1974 (D. 
Otte), 2d (yellow wings, tegmina lack red tinge); Petropolis, 
12 Apr. 1913 (M. Burr), 2d (yellow wings, tegmina lack red 
tinge). MINAS GerRalIs: Vicosa, 9 Aug. 1938 (B. T. Snipez), 
22 (yellow wings, tegmina lack yellow tinge); 46 km sE Ita- 
juba, 1400 m, 21 Mar. 1980 (Roppa, Carbonell, Roberts), 3d, 
32. GotAs: betw. Sao Simao and Jatai, 5 Mar. 1980 (Roppa, 
Carbonell, Roberts), 1¢, 22; 15-30 km E Mineiros, 7-9 Mar. 
1980 (Roppa, Carbonell, Roberts), 12. SAo PAULO: 10 km w 
Sao Joao de Boa Vista, 19 Mar. 1980 (Roppa, Carbonell, Rob- 
erts), 30d; Franca, Jan. 1911 (E. Garbe), 1d, 12; Salto 


Grande, Feb. 1911 (H. Luderwaldt), 1¢; Cubatao (Alin), 2d, 
12 (orange wings, tegmina lack red tinge); Piracicaba, 1d 
(orange wings). PARANA: 24°38'S, 54°07’'W, 500 m, Mar. 1965, 
‘virgin deciduous forest (no Araucaria) with many palms 
(Euterpe etc.), no grass, under growth of ferns and other 
plants’ (F. Plaumann), 4°, 192, UMMZ; Curitiba, 13 Feb. 
1941 (J. R. Bailey), 1d, 12, UMMZ (male has orange wings, 
lacks red tinge on tegmina). SANTA CATARINA: Nova Teuton- 
ia, 27°11'S, 52°23'W, 6 km sw Seara, 300-500 m, 1961-1964 
(F. Plaumann), 66, 52, UMMZ; Corupa, Jan.—Mar. 1956— 
1962 (Anton Maller), 3¢,92, UMMZ (1d, 52 have red wings, 
2¢ lack red tinge on tegmina, 2d, 42 have orange-yellow 
wings and lack red tinge on tegmina); Rio Capivari, 1889 
(Fruhstorfer), 22 [gift of Dr. H. Saussure, ANSP, labelled R. 
miles Drury and Rhomalea speciosa, probably what Pictet 
and Saussure considered to be R. miles Var. C, as it has 
yellow wings; there are two Rio Capivari’s in eastern lowlands 
of Santa Catarina]; Pinhal 700 m, Apr. 1959 (Anton Maller), 
12 (yellow wings, lacks red tinge on tegmina). RIO GRANDE 
po SuL: Aratinga, Feb. 1964 (Carbonell, Mesa, Monné), 1d 
(yellow wings). MATo Grosso: 40 km E Rodonopolis, 11 Mar. 
1980 (Roppa, Carbonell, Roberts) 1d; 30 km Nw Alto Ara- 
guaya, 750 m, 10 Mar. 1980 (Roppa, Carbonell, Roberts) 1d; 
Chapada near Cuiaba, 446, 32 (1d lacks distal band on hind 
femur); Corumba, Urucum, 22-29 Dec. 1919 (R. G. Harris), 
123, 122 (proximal median bands weak, distal band strong 
on hind femur, proximal tibial band usually absent); Tres La- 
gos, 6-10 Dec. 1919 (Harris), 49 (hind legs fully banded, and 
one of these with yellow markings strongly tinged with red), 
12 (distal band on hind femur and hind tibia only). Mato 
Grosso SuL: 30-60 km E Aquidauana, 16 Mar. 1981 (Roppa, 
Carbonell, Roberts), 2d. 

Uruguay. Whole country, Dec.—Apr., large series, CSC 
(yellow markings tinged with red). 

Ecuador. Balzapamba (R. Haensch), 1d (weak proximal 
band on hind femur). Putumayo Dist., La Sombra to El En- 
canto, 23 Aug. 1920, 1d (strong proximal and weak median 
band on hind femur). 

Peru. JUNiN: Satipo, 15 Nov. 1945 (P. Paprzycki), 1d, 89; 
Satipo, 650 m, Jul. 1940 (Schunke) 1°; Col. Perené, El] Cam- 
pamento, 22 Jul. 1920, 22; Chanchamayo, 1d, 1°; Vilcanota, 
12; Puerto Yessup, Feb. 1930 (M. A. Carriker), 12. Cuzco: 
Valle de Urubamba, Sahayaco, 800 m, 7 Dec. 1947 (Wey- 
rauch), 4d; Prov. La Convencion, Sangobatea, Jan. 1976 (J. 
S. Escalante), 3d, 32, JAE; Prov. Paucartambo, Salvacion, 
Oct. 1968 (F. Carrasco), 12, FCZ. All Peruvian specimens 
have moderate to well-developed proximal bands, weak to no 
median bands, and no distal bands on hind femur; no bands 
on hind tibia and other legs. 

Bolivia. SANTA CRUZ: Province of Sara, 450 m, Jan. 1918 
(J. Steinbach), 19¢, 272 (35 had no tibial banding, 11 had 
weak distal yellow bands; on hind femur all had weak to strong 
proximal bands, 14 had no median bands, 22 had weak to 
strong distal bands, and 24 had no distal bands) [Note: this 
previously recognized Province of Sara, bounded in part by 
the Rio Grande or Guapay and the Rio Mamoré or Ichilo, is 
the region where Steinbach did much of his collecting, and 
included the town of Buena Vista (where his relatives lived 
and where a niece presently operates a small restaurant, store, 
and inn), Portachuelo, and the city of Santa Cruz, that is, 
Santa Cruz de la Sierra.]; Buena Vista, May 1917 (Steinbach), 
23, 12; between Buena Vista and San Carlos, 350 m, cacao 
forest, 21 Feb. 1976 (Ronderos, Roberts) 2d, 12; between 
Buena Vista and Portachuelo, 20 Feb. 1976, 12 (specimens 
from last. three localities similar in variation to those from 


ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS 


Prov. of Sara; Santa Cruz de la Sierra, Feb. 1922 (J. Stein- 
bach), 5d, 62; 18 km sw Santa Cruz, 400 m, 16 Feb. 1976 
(Ronderos, Roberts), 3d, 22 (of last 16 specimens, 15 had a 
strong distal band on hind femur, Id lacked this band); Prov. 
of Nuflo de Chavez, Ascencion, 15°42'S, 63°05'W, 500 m, 
Nov. 1963 (Walz), 4d, 52 (5 had distal and 4 had no distal 
band on hind femur). See Table | for a summary of this band- 
ing. 

Paraguay. 26 specimens from Villa Rica, Jan., Feb.; Sapu- 
cay, Jan._Apr.; Horqueta, Dec., Jan. (yellow markings 
strongly tinged with red; hind femur with three bands, hind 
tibia with two bands). 

Argentina. 102 specimens from 24 localities in the provinces 
of Jujuy, Feb.; Salta, Mar.; Chaco, Feb.; Misiones, Dec., 
Feb.; Tucuman, Mar.; Catamarca, Mar.; Cordoba, Mar.; 
Entre Rios, Mar.; La Rioja, Feb.; Mendoza Feb.—Apr.; San 
Luis, Jan.; Buenos Aires, Feb. As in Paraguay and Uruguay, 
yellow markings are strongly tinged with red; hind femur with 
three bands, hind tibia two bands. Adults may be found De- 
cember to April. 


COMMENTS.—Unlike other species of the ge- 
nus, no consistent differences have been found 
to distinguish various geographical develop- 
ments. Comparing specimens, for example, 
from Carabobo, Venezuela, with those from 
Santa Cruz, Bolivia, or Corupa, Brazil, it is ev- 
ident that considerable geographic differentia- 
tion occurs, but it does not seem possible or 
practical with our present evidence to recognize 
subspecific elements of the species. 


5. Chromacris nuptialis (Gerstaecker) 


Rhomalea nuptialis GERSTAECKER, 1873:185. [Holotype, ¢, 
bearing label with number 2008, and 2 allotype, Berlin Mu- 
seum; Saltogrande (Sellow). The locality of Salto Grande of 
Sellow is uncertain. Sellow visited Salto Grande on the 
Uruguay River (Department of Salto, Uruguay), but none 
of the species he labelled Saltogrande has ever been found 
in Uruguay or the adjacent Brazilian state of Rio Grande do 
Sul. Sellow’s Saltogrande is very probably that on the Par- 
anapanema River in the state of Sao Paulo. Types exam- 
ined. ] 


Rhomalea latipennis PicTET AND SAUSSURE, 1887:351. [Holo- 
type, d , Geneva Museum; Brazil. It does not have a locality 
label but bears the number 477-56, which in the museum 
records corresponds to Brazil, collected by Ferrier circa 
1856. We have been unable to trace the collector’s route in 
Brazil. Holotype examined. New synonym.] 


DIAGNosIs.—Coloration highly variable, in- 
cluding individuals with pale yellow and pale red 
wings. Body and legs with yellow or red mark- 
ings (irrespective of wing color). Antennae black. 
Pattern of hind wings as in C. speciosa. Tegmina 
very characteristic, membrane dark brown to 
black, strongly contrasting straw-colored veins. 
Fore and middle legs without transverse bands, 
ground color variable from greenish yellow to 
reddish brown; longitudinal series of black spots, 
sometimes coalescing into black streaks. Hind 


n 
eS) 


femora black to dark brown, with longitudinal! 
carinae and fishbone pattern of a lighter color, 
variable from reddish brown to straw-yellow; 
transverse bands absent or very slightly marked, 
only exceptionally, plainly visible, proximal one 
on upper half only, median one may be complete, 
distal one always absent. Proximal and median 
bands always visible on inner and lower surfaces 
of hind femur as conspicuous yellow or red areas, 
no trace of distal one. Hind tibia generally dark 
colored, especially on inner side, transverse 
bands generally absent, sometimes faintly 
marked, the distal one, proximal one, or both 
may be visible in different specimens. Prosternal 
tubercle relatively slender, long, and curved rear- 
wards. 

DISTRIBUTION.—Inland south-central Brazil, 
including southern Goias, western Minas Gerais, 
western Sao Paulo, and northwestern Parana. 

SPECIMENS.—Brazil. D.F.: Brasilia, Nov. 1963 (N. Tanger- 
ini), 2¢ (yellow wings), CACS. GorAs: rodovia Anapolis— 
Brasilia, km 63, 17 Feb. 1964 (H. M. Canter), 1¢ (yellow 
wings), MZSP; betw. Sao Simao and Jatai, 5 Mar. 1980 
(Roppa, Carbonell, Roberts), 12 (red wings), CACS; Min- 
ieros, Feb. 1975 (Roppa, Silva), 12 (red wings), CACS. 
MINAS GerRaIs: Uberaba, Feb. 1979 (Roppa, Silva), 1d (red 
wings), CACS; Diamantina, Mar. 1956 (D. Albuquerque), 1 2 
(red wings), CACS; Lagoa Santa, Jul. 1965 (M. S. Morgante), 
1d (yellow wings), MZSP. PARANA: Vila Velha, Jan. 1975 
(C. Valle), 16, 12 (yellow wings), MZSP. 

COMMENTS.—Individuals of this species are 
highly variable in color, and therefore difficult 
to identify. However, the only other species 
known from the area is C. speciosa, from which 
it can be separated by the peculiar coloration of 
its tegmina, the very different color on pronotum 
and legs, and the form of its pronotal tubercle. 
The species appears to be uncommon, being al- 
ways represented by one or two specimens from 
each locality, which is unusual for species of this 
genus. Possibly, this taxon is a highly aberrant 
variation of C. speciosa in the middle of whose 
territory it occurs, but the constancy of some of 
its characters seems to indicate that it is a dis- 
tinct species. 


6. Chromacris trogon trogon (Gerstaecker) 


Rhomalea trogon GERSTAECKER, 1873:186. [Holotype, 1°, 
Berlin Museum; Costa Rica. (Gerstaecker noted that hind 
legs were lacking. A bright yellow marked leg was later 
attached and now removed. Holotype examined. ] 
DiAGNosis.—Hind wings yellow to orange- 

yellow, and lack black on anterior basal half of 

first two plaits (Fig. 7). Antennae entirely black. 

No yellow banding on legs, although often faint 

pale green bands present on hind femora (Table 


54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


TABLE 2. 


DIAGNOSTIC CHARACTERS FOR THE SPECIES AND SUBSPECIES OF THE TROGON GROUP OF GENUS Chromacris. A 


band that does not extend entirely around the hind femur is listed as a half band. 


Basal 2 Pronotal 
Name Range Antenna of wing metazona Hind femur Hind tibia 

t. intermedia N Guatemala all black all yellow no yellow 3 half bands no bands 
to Honduras 

t. trogon Costa Rica all black all yellow no yellow no bands no bands 

p. psittacus Costa Rica to all black all yellow no yellow 2 and % bands 2 bands 
N Colombia, 
N Venezuela 

p. pacificus w Colombia, all black all yellow no yellow 3 bands 2 bands 
w Ecuador 

icterus E Colombia tip yellow all yellow yellow patches 3 bands 1 band 
to NE Peru, 
Nw Brazil 

peruviana Peru tip yellow plait 1 black yellow patches 3 bands 2 bands 


2). Reproductive structures and prosternal tuber- 
cle similar to those in C. psittacus and other 
yellow-winged species of the trogon group. 

DIsTRIBUTION.—Costa Rica and questionably 
Nicaragua. 


SPECIMENS.—Costa Rica. Pozo Azul, Rio Perris or Parrita 
(forested foothills of Pacific coastal plain), May 1902 (M. A. 
Carriker), 2d, 12; 22 Aug. 1927 (Lankester and Rehn), 36, 
12. Between La Union and Buenos Aires, Terraba Valley, 
5500 ft. [1670 m], May 1935 (Lankaster), 12. Juan Vinas, 3300 
ft. [1000 m], Mar. 1902 (L. Beamer), 1d, 12; 27 Jun. 1909 (P. 
P. Calvert), 1d, 12. Peralta, 8 Aug. 1909 (Calvert), 1¢; May 
1923 (Lankester), 1d. La Emelia near Guapiles, Aug. 1923, 
Sep. 1927 (Rehn), 4d. Cariblanca, 600 m (Lankester), 1°. 
Parisimina, 5 m, 26 Jul. 1928 (M. Valerio), 12. PUNTARENAS: 
Rio Cataratas, near Brujo, Sep. 1979 (Rowell), 1d, 12. ALa- 
JUELA: 5 km s San Lorenzo, Sep. 1979 (Rowell), 1d, 12, 
CHER. 


Nicaragua. CHONTALES: (Janson), 1d, 12 (poorly pre- 
served but appears to be this species). 


COMMENTS.—Although we have no records 
of C. trogon trogon and C. psittacus occurring 
together, it seems possible that they do. For ex- 
ample, we have this species from near Guapiles 
and C. psittacus from Siquirres about 30 km dis- 
tant in similar lowland forest country. 


7. Chromacris trogon intermedia n.subsp. 


DIAGNOsIS.—Wing orange with no black on 
basal half of first two plaits, or first paired fold 
as in C. colorata. Antennal segments all black. 
Hind femur with yellow bands that may be 
weakly or strongly developed. Hind tibia with 
no bands. Prominent wide yellow stripe along 
ventral margin of lateral lobe of pronotum that 


extends onto cheek of head. Prosternal tubercle 
short, tapering rapidly to a point. Distinguished 
from C. psittacus by lack of banding on hind 
tibia, and from the nominate subspecies of C. 
trogon by banding on hind femur (Table 2). 

DISTRIBUTION.—Northern Guatemala, Be- 
lize, and Honduras. 


SPECIMENS.—Holotype: 6, ANSP; Honduras, Lancertilla 
near Tela, Dept. Atlantica, 100-800 ft. (30-250 m), rain forest, 
8 Nov. 1930. 

Paratypes: Same data as for holotype, 32. Belize. 50 mi. [80 
km] s El Cajo, Mountain Pine Ridge road, 17 Aug. 1960 (P. 
N. Litchfield), 16, 22, UMMZ. Guatemala. Piedras Negras, 
600-800 ft. [180-240 m], 30 Jun. 1933 (D. W. Amran), 1°. 


COMMENTS.—This subspecies is intermediate 
between C. trogon trogon and C. psittacus in 
the reduction in banding of the hind leg, which 
might suggest that they should be treated as 
three subspecies, but C. trogon trogon and C. 
psittacus occur close together in Costa Rica, 
though as noted under the subspecies, C. trogon 
trogon, they have not been recorded from the 
same locality. Specimens from. Nicaragua and 
Honduras are needed to help clarify the prob- 
lem. 


8. Chromacris psittacus psittacus (Gerstaecker) 


Romalea psittacus GERSTAECKER, 1873:185. [Lectotype, d, 
among four male syntypes with same data, one marked 
‘‘typus’’ and here designated; Berlin Museum; Bogota, Co- 
lombia. Species of Chromacris probably do not occur in the 
vicinity of Bogota. ANSP has specimens of C. icterus also 
labelled Bogota, and this species actually occurs at lower 
elevations on the eastern slopes of the Andes.] 


ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS 55 


FIGURES 26 and 27. 


Internal male genitalia of Chromacris miles from Quintana Roo, Mexico (Fig. 26), and Xestotrachelus 


robustus from Thi, Caaguazu, Paraguay (Fig. 27). A, phallus, lateral; B, cingulum, lateral; C, endophallus, lateral; D, phallus, 
dorsal; E, cingulum, dorsal; F, endophallus, dorsal; G, epiphallus, dorsal; H, epiphallus, frontal; 7, epiphallus, lateral, left side. 


DIAGNOsIs.—Wings yellow to orange-yellow, 
lacking black on anterior basal half (Fig. 8). An- 
tennae all black. No yellow on dorsum of me- 
tazona of pronotum. Hind femur with three yel- 
low bands, but proximal band not entire. Hind 
tibia with two bands. For comparison with other 
species see Table 2. 


SPECIMENS.—Costa Rica. Siquirres, 3 Jul. 1903 (M. A. Car- 
riker), 7d, 12. Ujarass de Terraba, 10 Sep. 1907, 1d. Monte 
Verde, “‘summer,”’ 1928 (F. G. Wallace), 1d, 12. Castilla 
Farm, lower Rio Reventazon, 29 Jul. 1936 (C. W. Dodge), 1d. 

Panama. Gatun, Jul._Aug. 1916 (D. E. Harower), 8d, 32. 
Cabima, 24 May 1911 (August Busck), 2d, 1 juv. Barro Col- 
orado Island, C.Z., 22 Jul. 1933 (H. H. Hood), 1°. 

Colombia. CUNDINAMARCA: Las Mesitas, Sep. 1915 (A. 
Maria), 1¢, 32. 

Venezuela. ZULIA: Kasmera, Rio Yasa, Sierra de Perija, 250 
m, 19 Sep. 1961, 26, 32, Universidad Central, Instituto de 
Zoologia Agricola, Maracay. 


COMMENTS.—Evidently seasonal. One of us 
(H.R.R.) visiting Costa Rica for several years in 
the Pacific and Caribbean lowlands during Feb- 
ruary and March never encountered this species 
or C. trogon. Most of our records suggest that 
it occurs commonly May to September. 


9. Chromacris psittacus pacificus n.subsp. 


DIAGNOsIs.—Similar to C. psittacus psittacus 
except proximal yellow band on lower portion 


of hind femur entire rather than interrupted. See 
Figure | and Table 2. 


SPECIMENS.—Holotype: d, ANSP; Ecuador, Dos Puentes, 
below Naranjapata along the Guayaquil-Quito railway in 
Chanchan River valley, 1750 ft. [S30 m], 15 Mar. 1931 (W. J. 
Coxey). [Additional information on location of Dos Puentes 
is added here from Coxey 1927:10.] 

Paratypes: Colombia. EL VALLE: Jiménez, 1600 ft. [486 m], 
19 Mar. 1907 (M. G. Palmer), 2¢, 12; Choco (M. G. Palmer), 
12. ANTIOQUIA: Andagoya, 1°; Cordillére, “‘vers occid. Rio 
Yurumaqui,’’ 1933 (E. Aubert de la Rue), 1d, 12, MNHN. 
NARINO; ‘“‘entre Guayacana et el Diviso,’’ 80 m, Nov. 1968 
(M. Descamps), 1d. 

Ecuador. CHIMBORAZO: Dos Puentes, 1750 ft. [S33 m], 11 
Jan. 1921 and 15 Mar. 1931 (W. J. Coxey), 5d, 22; Ventura, 
1400 ft. [469 m], 10-13 Apr. 1922, 36, 32. Guayas: Bucay, 
900 ft. [274 m], 19 Mar. 1922 (G. H. Tate), 12. TUNGURAHUA: 
Ambato, 16, MNHN; Balzapamba, near Ambato (R. 
Haensch S.), 1d, Berlin Museum. 


CoMMENTS.—The slight but consistent differ- 
ence in the form of the proximal yellow band on 
the hind femur of these specimens warrants sub- 
specific recognition. 


10. Chromacris icterus (Pictet and Saussure) 


Rhomalea icterus PICTET AND SAUSSURE, 1887:353. [Lecto- 
type, 2, so labelled and here designated, Geneva Museum; 
Quito, Ecuador.] 

Rhomalea opulenta GERSTAECKER, 1889:32. [Holotype, 2, 
Zoological Museum, University of Greifswald; Sao Paulo 


56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4 


de Olivenca, Amazonas, Brazil. Holotype examined. New 
synonym. ] 


DIAGNosIs.—Tip of antenna yellow. Anterior, 
basal portion of hind wing entirely yellow (Fig. 
9). Yellow patches on dorsum of metazona of 
pronotum. Three yellow bands on hind femur. 
Single proximal band on hind tibia distinguishes 
species from all others (see Table 2). A large 
species with relatively long wings; males 30-38 
mm, females 40-55 mm from fastigium to wing 
tips. 

DISTRIBUTION.—Southeastern Colombia, 
eastern Ecuador, northwestern Brazil and 
northeastern Peru. 


SPECIMENS.—Colombia. BoGora: (so labelled but probably 
in error) (A. Maria), 26, 42. CUNDINAMARCA: Susumuco, 
Feb., Sep. 1916, 1917 (A. Maria), 3d, 22. Meta: Villavicen- 
cio, May, Jun. 1919 (A. Maria), 2d, 22. PuTUMAyo: “‘bord 
riviére Mocoa,’’ 800 m, Nov. 1968 (M. Descamps), 1d, 
MNHN. 

Ecuador. MORONA-SANTIAGO PRov.: s of Méndez, 800 m, 
19-21 Oct. 1977 (L. E. Pena), 1°. 

Peru. Loreto: Putumayo District, La Chorrera to La Som- 
bra, 21 Aug. 1920, 1°; Iquitos, 8 Jan. 1920 (H. S. Parish), 1°. 

Brazil. AMAZONAS: ‘‘Hyntanahan’’ probably Huitanaa, Rio 
Purus, Jan. 1922 (S. M. Klages), 12; Tabatinga, Sep.—Dec. 
1977 (L. C. Pereira, B. Silva), long series both sexes, CACS; 
Eirunepe, Jun. 1950 (J. C. Carvalho), 2d, 12, CACS; Atalaya 
do Norte, Nov. 1977 (B. Silva), long series, CACS. 


11. Chromacris peruviana (Pictet and Saussure) 


Rhomalea peruviana PICTET AND SAUSSURE, 1887:352. [Lec- 
totype, 2, so labelled by C.S.C. and here designated from 
among Id and 32 syntypes, Geneva Museum; Peru.] 


DiaGNosis.—Similar to C. icterus in having 
yellow wings and yellow antennal tips, but dif- 
fers by having two yellow bands on hind tibiae 
rather than one (Table 2); differs from all other 
yellow-winged species by having the black on 
anterior portion of first plait of hind wings ex- 
tend to base of wing (Fig. 10). Tegmina relative- 
ly long, narrow, and greatly surpassing ends of 
hind femora; anterior and posterior margins bor- 
dered by yellowish or pale green areas. This teg- 
minal coloration is unique for genus. 


SPECIMENS.—Peru. JUNiN: Chanchamayo, a district around 
La Merced in valley below Tarma, 2000-3000 ft. (610-914 m), 
13; Satipo, near Huancayo, 1650 m, Jul. 1844 (Schunke), 1d; 
Mar., Jun. 1944 (P. Paprzycki), 32; Puerto Yessup, Feb. 1930 
(M. A. Carriker), 12. HUANuco: Leonpampa, 110 km E 
Huanuco, Dec. 1937 (Felix Woytkowski), 3d , 12; Tingo Mar- 
ia, 670 m, Sep. 1946 (Weyrauch), 32; same locality, 2 Dec. 
1954 (E. D. Schlinger, E. S. Ross), 1°; Divisoria, Cordillera 
Azul, 1500 m, 1d, 12. Loreto: Rio Aguaytia, between Tingo 
Maria and Pucallpa, 400 m, Feb. 1961, 16, 22. SAN MARTIN: 
Prov. Huallaga, Rio Mixiollo, 1200 m, 7 Aug. 1900 (C. A. 


Baer), 12. Cuzco: Paucartambo, Pilcopata, Nov. 1968 (F. 
Carrasco), 1d, 12; Paucartambo, Atalaya, May 1976 (Des- 
camps, Carbonell), 1d, 12, CACS. 

Venezuela. ARAGUA: Nov. 1942, 32. 


COMMENTS.—The Venezuelan specimens are 
undoubtedly this species. However, C. icterus 
occurs between this and the Peruvian localities, 
suggesting the need to confirm the correctness 
of the locality. 


Xestotrachelus Bruner 


Xestotrachelus BRUNER, 1913:469. [Type-species Xestotrach- 
elus hasemani Bruner (=X. robustus) by original designa- 
tion.] 


D1iaAGNosis.—Red and black pattern of hind 
wings (Fig. 11) closely resembles red-wing 
species of Chromacris, but head and thorax are 
much more robust, and tegmina may extend well 
short of, or only slightly beyond, end of hind 
femora (Fig. 2). Head, pronotum, and other 
parts of body and appendages may be contrast- 
ingly marked with black, pale olive-yellow, and 
red. Hind tibiae usually red. Phallic structures 
described under X. robustus. 

DisTRIBUTION.—Known from Maranhao in 
northeastern Brazil, Bahia, Espirito Santo, 
Mato Grosso; Paraguay; and eastern Bolivia. 


12. Xestotrachelus robustus (Bruner) 


Zoniopoda robusta BRUNER, 1911:60. [Lectotype, ¢, here 
designated, ANSP; Chapada dos Guimaraes, Mato Grosso, 
Brazil. The type-series consisted of a male and female, each 
labelled as the type.] 

Xestotrachelus hasemani BRUNER, 1913:470. [Holotype, 2, 
ANSP; labelled as from Galhao, not Calhao as reported by 
Bruner, Rio Sapao, western Bahia, Brazil. Actually Galhao, 
10°35’S, 46°15’W, is in Goias on the Rio Galhao, and the 
Rio Sapao is nearby in Bahia. New synonym.] 


DIAGNosIs.—See diagnosis of genus. Extent 
of black on prozona and mesozona of pronotum 
variable, may be entirely black, or divided to 
form two black transverse bands. Extent of red 
on sides and undersides of abdomen also vari- 
able. Easily distinguished from species of Chro- 
macris by red hind tibia and lack of banding on 
hind femur. 

Phallic structures generally similar to those in 
Chromacris. Aedeagal valves (Fig. 21) strongly 
sclerotized, much larger, and sculptured. Rami 
of cingulum (Fig. 27B) narrow, partly surround- 
ing base of aedeagus, strongly bent inward and 
ventrad near their end, and outer surface at bend 
covered with small spines. Epiphallus (Fig. 


ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS 57 


27G-I) similar to that in Chromacris except that 
the ancorae are well developed. 
DISTRIBUTION.—See distribution of genus. 


SPECIMENS.—Brazil. MARANHAO: Barra do Corda, Feb. 
1955, 12, CACS. Banta: Maracas, Feb. 1963 (F. M. Oliv- 
eria), 32, CACS. Espirito SANTO: Linhares, Mar. 1981 (B. 
Silva), 1¢, CACS. GotAs: in addition to the type-locality of 
X. hasemani, 60 km w Mineiros, 10 Mar. 1980 (Roppa, Car- 
bonell, Roberts), 1d. Mato Grosso: in addition to the type 
of X. robustus, Corumba, Urucum, 23-29 Dec. 1919 (R. G. 
Harris), 4 juv., 18¢, 142. Mato Grosso SUL: 30-60 km E 
Aquidauana, 16 Mar. 1980 (Roppa, Carbonell, Roberts), 1d. 

Bolivia. SANTA CRUZ: prov. of Sara, 450 m, Jan.—Feb. 1922 
(J. Steinbach), 8d, 162; Buena Vista, 500 m, 3 Feb. 1922 (J. 
Steinbach), 1°. 

Paraguay. CAAGUAZU: near [hu, Mar. 1965 (Carbonell, 
Mesa, Monne), 1d, 22, CACS. AMAmMBay: Cerro Cora, Jan. 
1972 (Descamps, Ronderos, Carbonell), 8d, 52, 1 last instar 
nymph, CSC. 


COMMENTS.—Individual geographic variation 
in relative tegminal length is evident. In the 
series recorded from eastern Bolivia, tegmen 
shorter than or about as long as hind femur. In 
the series from Corumba, tegmen slightly or de- 
cidedly longer than femur. Tegmen of male from 
near Aquidauana 19 mm and hind femur 15.5 
mm in length, whereas in the male from Minei- 
ros, Goias, tegmen 16 mm and femur 17 mm. 
Tegmen of female holotype of X. hasemani is 
26 mm, hind femur 18 mm. This specimen has 
a decidedly longer tegmen in proportion to the 
hind femur than do specimens to the south. It 
seems possible, however, that populations to be 
found between this type-locality and those pres- 
ently observed to the south will be intermediate 
in relative tegminal length. The aedeagus of the 
male from Espirito Santo is relatively shorter 
and therefore appears to be somewhat broader 
in lateral view than others examined. The red 
patch near the end of the hind wing in the longer- 
winged specimens from Linhares and Maranhao 
connects narrowly with the large red field of the 
wing. Conceivably, this modification is the re- 
sult of the lengthened wing. With more evidence 
the longer-winged hasemani form might be rec- 
ognized as a subspecies. 

Noteworthy is the last instar nymph from Cer- 
ro Cora, Paraguay, recorded above, the color of 
which is well preserved and matches the color 
pattern of the adult. Unlike the species of Chro- 
macris which have a bright but relatively cryptic 
coloration in the adult stage, X¥. robustus has a 
much more striking coloration in the adult, 
which may well serve as a warning to predators. 


Assuming this is correct, then there would be no 
need to develop a different coloration in the 
nymphal stages as in Chromacris. 


LITERATURE CITED 


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ASTACIO-CABRERA, O. 1975. Notas sobre algunas acridioi- 
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BARRERA, M., AND S. Z. TurK. 1977. Acridios del NOA. II. 
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BRUNER, L. 1911. South American Acridoidea. Annals of the 
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Coxey, W. J. 1927. Impressions of Ecuador. Year Book 
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Drury, D. 1773. Illustrations of natural history, wherein are 
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GAUGLIUMI, P. 1973. Pragas de cana-de-acucar, nordeste do 
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GERSTAECKER, A. 1873. Acridiodea nonulla nova insigniora. 
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. 1889. Charakteristik einer Reihe bemerkenswerther 
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en Verein Neu-Pommern und Riigen in (Greifswald) Berlin 
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Haun, C. W. 1835. Icones Orthoptorum. Nurnberg. 3 p., 4 
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KirBy, W. F. 1910. A synonymic catalogue, Orthoptera Sal- 
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PicTeET, A., AND H. DE SAUSSURE. 1887. Catalogue 
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PRETTO-MALCA, R. 1968. Estudios del ciclo biologica, mor- 
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RaGGE, D. R. 1955. The wing-venation of the Orthoptera Sal- 
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SERVILLE, M. A. 1839. Histoire Naturelle des Insects Or- Estudios biologicos, morfométricos y aspectos ecolégicos 
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CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


PROCEEDINGS 
OF THE 


, CALIFORNIA ACADEMY OF SCIENCES 


Vol. 43, No. 5, pp. 59-66, 5 figs., 1 table. 


November 4, 1982 


A NEW GENUS AND TWO NEW SPECIES OF 
REMARKABLE PACIFIC WORM EELS»)... 
(OPHICHTHIDAE, SUBFAMILY MYROPHINAE) | 


By 


John E. McCosker 


Steinhart Aquarium, California Academy of Sciences, Golden Gate Park, 
San Francisco, California 94118 


ABSTRACT: Glenoglossa wassi gen. et sp.nov., described from Samoa, differs from all other myrophine 
ophichthids in its elongate tongue decorated with a lure, and in certain osteological and cephalic pore con- 
ditions. Neenchelys daedalus sp.nov., is described from midwater-captured juveniles and adults from off New 
Guinea and the central Pacific and represents the second known midwater worm eel; it differs from its 
congeners in its extreme elongation and vertebral number. The status of the species of Pseudomyrophis and 
Neenchelys is discussed. An analytical key to the genera of the ophichthid subfamily Myrophinae is provided. 


INTRODUCTION 


The snake eels and worm eels of the family 
Ophichthidae are the most diverse and inventive 
of true eels. The more than 220 species distrib- 
uted among more than 53 genera inhabit all trop- 
ical and subtropical oceans and seas, and have 
invaded the intertidal zone, coral reefs, shallow 
substrates, and even the midwater realm. To this 
array of astounding evolutionary forays, I add 
two remarkable new western Pacific species, one 
representing a distinct new genus of worm eels, 
of the subfamily Myrophinae (sensu McCosker 
1977). The first, Glenoglossa wassi gen. et sp.nov., 
is unique among eels in having modified its glos- 
sohyal into a lure, not unlike that of a urano- 
scopid stargazer or an antennariid. The other, 
Neenchelys daedalus sp.nov., represents a sec- 
ond, independent invasion of the midwater realm 
by an ophichthid. 


[59] 


MATERIALS AND METHODS 

Measurements are straight-line, made either 
with a 300 mm ruler with 0.5 mm gradations 
(for total length [TL], trunk length, and tail length) 
recorded to the nearest 0.5 mm, or with dial 
calipers (all other measurements) and recorded 
to the nearest 0.1 mm. Body length comprises 
head and trunk lengths. Head length (HL) is mea- 
sured from the snout tip to the posterodorsal 
margin of the gill opening; trunk length is taken 
from the end of the head to mid-anus; maximum 
body depth does not include the median fins. 
Vertebral counts, which include the hypural, were 
taken from radiographs. Stained and cleared 
specimens were prepared using the Taylor (1967) 
trypsin technique. Institutional abbreviations of 
material examined are explained in the 
Acknowledgments. 


60 


2a. 


2b. 


3a. 


3b. 


4a. 


4b. 


Sa. 


Sb. 


6a. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5 


ANALYTICAL KEY TO THE GENERA OF 
OPHICHTHIDAE, SUBFAMILY MYROPHINAE 


All branchiostegal rays originate either in 
association with hyoid or before level of 
epihyal tips; free rays, when present, fewer 
than attached; tail tip a hard or fleshy fin- 
less point; gill opening midlateral to 
entirely ventral, unconstricted _..... 
SSC nae subfamily Ophichthinae 


. Accessory branchiostegal rays originate 


behind ends of epihyal, free rays more 
numerous than attached; caudal fin rays 
conspicuous, confluent with dorsal and 
anal, tail tip flexible; gill openings mid- 
lateral, a constricted opening (subfamily 
Myrophinae) 


Anterior nostril non-tubular, posterior 
nostril before eye; eye large, ca. 6 times 
in head length; pectoral fin moderately 
developed _. Benthenchelys Fowler, 1934 
Anterior nostril tubular, posterior nostril 
either before eye, along upper lip, or within 
mouth; eye smaller, 10 or more in head; 
pectoral fin may be absent 3 


Posterior nostril before eye, above the lip 
and not covered by a flap; pectoral fin 
present, but may be reduced to a small, 
barely noticeable flap in posterodorsal 
cormeroleillopening a eee 4 
Posterior nostril labial, either within lip 
and opening into mouth, or along lip and 
covered by a flap; pectoral fin either pres- 
ent and well developed or absent 


Dorsal fin origin in anterior trunk region; 
snout conical; pectoral fin well developed, 
=~ snout; third preoperculomandibular 
POLE POPs) ra SCM bee nes 
ss be en ee Neenchelys Bamber, 1915 
Dorsal fin origin in posterior trunk region; 
snout broad, tumid; pectoral fin minute, 
=<VeVeEsPOP) DIeSeiit, a ee see ee eee 
ih SACS. dis Pseudomyrophis Wade, 1946 


Pectoral fin well developed; pleural ribs 
absent behind 15th—20th vertebrae _. 6 
Pectoral fin absent; pleural ribs present on 
allitrunksventebrace — Nee eee 7 


Dorsal fin origin above or behind anus; 
maxilla stout, not tapering posteriorly, and 
abutting pterygoid; vomerine teeth absent 
Cerda eeenneee Ahlia Jordan and Davis, 1891 


6b. Dorsal fin origin anterior to mid-trunk 
region; maxilla thin and tapering poste- 
riorly, not closely associated with ptery- 
goid; vomerine teeth present _. 
elle ae cee eS Myrophis Liitken, 1851 


7a. Tongue elongate, extending well beyond 
mouth and decorated with a fleshy appen- 
dage; inner edge of lips and palate deco- 
rated with fleshy lappets; teeth conical and 
UnISehial = ee Glenoglossa novum 
Tongue not elongate, not extending out- 
side of mouth, lacking a fleshy appendage 
at its tip; inner edge of lips and palate 
smooth; teeth either conical or blunt, uni- 
serial or multiserial — eee 8 


7b. 


8a. A prominent median toothed groove on 
ventral side of snout, bordered by dermal 
folds, extends anteriorly to anterior nos- 
trils; anterior nostrils elongated tubes equal 
to eye in length __ Ea 
Brena Sk Schismorhynchus McCosker, 1970 
Ventral side of snout without a prominent 
median groove bordered by dermal folds; 
anterior nostrils less than eye in length __ 9 


8b. 


9a. Teeth absent on vomer, absent or embed- 
ded on intermaxillary, those on maxillary 
and dentary minute or villiform; dorsal 
fin origin behind anus 
a TAS Sbe seeS Te, I, Schultzidia Gosline, 1951 
Teeth present on intermaxillary, maxil- 
lary, dentary, and vomer; dorsal fin origin 
either before or behind anus _.. 


Se Muraenichthys Bleeker, 1853 


9b. 


Glenoglossa McCosker, gen.nov. 


Type-species. — Glenoglossa wassi McCosker, sp.nov. 


D1AGnosis.—Body moderately elongate, lat- 
erally compressed posteriorly; snout conical, 
grooved on underside; anterior nostril within a 
tube, posterior nostril within a short tube at outer 
edge of lip, directed ventrally; dorsal fin origin 
slightly before anus; pectoral fin absent; pop’ 
absent; tongue elongate, extends beyond mouth, 
decorated with fleshy appendage; inner edge of 
lips and palate decorated with fleshy lappets; teeth 
conical, uniserial, absent on vomer; gill arches 
reduced, third hypobranchial and second infra- 
pharyngobranchial absent, third and fourth upper 
pharyngobranchial tooth plates weakly fused; 
suspensorium nearly vertical, pterygoid slender 


McCOSKER: NEW WORM EELS 


61 


FiGure 1. 
and anal fins. 


and reduced; cleithrum and supracleithrum 
reduced to thin slivers. Other characters those of 
the single species. 

ETyMOLoGy.—From the Greek yAjuvoo (gle- 
nos), a thing to stare at, and yA@ooa (glossa, fem- 
inine), tongue. 


Glenoglossa wassi McCosker, sp.nov. 
(Figures 1-3) 

Ho.totyre.—CAS 47049, 153.5 mm TL, collected using 
rotenone over sand at base of large coral head, 40 m depth, 
Larsen Bay, Tutuila Island, American Samoa; R. Wass and G. 
Yamasaki, 18 Nov. 1975. 

Paratype. —CAS 47048, 88.8 mm TL, collected using rote- 
none along sloping sand bottom at base of coral and lava cliff, 
40 m depth, Steps Point, Tutuila Island, American Samoa; R. 
Wass and R. Lubbock, on 4 Mar. 1975. This specimen was 
cleared and stained. 


COUNTS AND MEASUREMENTS. — The condition 
of the holotype is parenthetically followed by 
that of the paratype: TL 153.5(88.8); head length 
20.8(12.8); trunk length 54.7(30.0); tail length 
78.0(46.0); body depth at gill openings 5.6(3.7); 
body width at gill openings 3.0(1.9); origin of 
dorsal fin 67.1(39.6); snout length 3.5(2.3); upper 
jaw length 6.7(4.4); eye diameter 1.1(0.7); inter- 
orbital distance 2.0(1.0). Total vertebrae 
127(126), preanal vertebrae 54(53). 

DESCRIPTION. — Body moderately elongate, 
depth at gill oepnings 24—26.5 in TL, tapering 
and laterally compressed posteriorly. Head and 
trunk 2.03-2.07 and head 6.9-7.4 in TL. Snout 
acute; lower jaw included, tip slightly in advance 
of anterior nostril base. Anterior nostril tubular; 
posterior nostril at edge of lip, entirely outside 
of mouth, within small tube. Eye slightly in 


Holotype of Glenoglossa wassi McCosker, sp.nov., CAS 47049, 153.5 mm TL. Arrows indicate origin of dorsal 


advance of midpoint of upper jaw. Gill opening 
mid-lateral, a constricted opening. 

Median fins low, lying partially within a groove, 
meeting each other and extending noticeably 
beyond caudal tip. Dorsal fin arises less than a 
head length in advance of anus. 

Head pores minute, difficult to discern. Single 
temporal and interorbital pores. Four pores along 
mandible. Two preoperculomandibular pores. 
Lateral line pores difficult to identify in pre- 
served specimens; approximately 11 pores before 
the gill opening. 

Tongue extends from mouth. A fleshy appen- 
dage, differing slightly in each specimen (Figs. 2— 
3), extends beyond the slender glossohyal. The 
inner edge of lips, floor of mouth, and palate 
flanked by fleshy tissue (Fig. 2). 

Teeth small, conical, uniserial in jaws. An 
intermaxillary chevron of four teeth, followed by 
two medial teeth. Vomerine teeth absent. Nine 
teeth along maxilla, 14 along mandible. 

Body color in isopropyl alcohol uniformly tan. 
Numerous, minute brown punctations in mouth, 
along head and dorsal body surface. Fins pale. 
Base of lure has a dark spot. Eyes dark blue. Color 
of paratype in life, recorded by R. Wass, “‘light 
greenish-yellow with tiny purple brown specks. 
Lure transparent with black ‘eye.’ ”’ 

EtyMoLoGy.— Named in honor of Richard C. 
Wass, collector of these and many other impor- 
tant fishes from Samoa. 

REMARKS.— This myrophine is remarkable in 
the development of its tongue which, because of 
its length and appearance, serves as a lure to 
attract small fish. It is the only eel known to use 


62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5 


Figure 2. Open mouth of holotype of Glenoglossa wassi 
McCosker, sp.nov., CAS 47049. 


such a feeding strategy, and is certainly no less 
remarkable than those of certain uranoscopids, 
ceratioids, or the alligator snapping turtle. The 
“eyes” and appendages of the lure would indicate 
that it resembles a small crustacean. In the stom- 
ach of the holotype was a partially digested goby- 
like fish nearly 2 cm in length. The other oral 
decoration provided by the fleshy labial lappets 
suggests that the eel lies buried in the sand with 


Ficure 3. Tongue lure of paratype of Glenoglossa wassi 
McCosker, sp.nov., CAS 47048. 


its head exposed, luring prey by flicking its glos- 
sohyal. 

Glenoglossa wassi is most closely related to 
species of Muraenichthys and Schismorhynchus. 
It is easily separable from them on the basis of 
its tongue development, its absence of vomerine 
teeth, and certain other osteological characters. 
It is most like Schismorhynchus in its general 
facies, the development of its snout groove, 
reduced pterygoid, and gill arch reductions, con- 
ditions which probably relate to the feeding 
behavior of the species involved. It is clearly a 
specialized worm eel whose ancestry is in the 
sharp-snouted, conical-toothed species group of 
Muraenichthys (sensu McCosker 1977) that gave 
rise to Schismorhynchus and may have shared a 
common ancestor with it. 


Neenchelys Bamber, 1915 


Type-species. — Neenchelys microtretus Bamber, 1915. 


DiAGNosis.— Body moderately to extremely 
elongate, laterally compressed behind head; tail 
much longer than head and trunk; snout conical, 
anterior nostril in a tube, posterior nostril an 
elongate slit entirely before eye; eye moderate; 
dorsal fin origin mid-trunk; pectoral fin moder- 
ately developed, girdle limited to reduced clei- 
thrum and supracleithrum; gill opening reduced, 
aconstricted hole; pop? absent; teeth few, conical 
and slender, uniserial on jaws and vomer; max- 
illary attachment at mid-vomer; gill arches 
reduced; suspensorium nearly vertical, slightly 


McCOSKER: NEW WORM EELS 


63 


Ficure 4. Holotype of Neenchelys daedalus McCosker, sp.nov., AMS 1.19690-012, 341.5 mm TL. 


inclined anteriorly; pterygoid slender and 
reduced; neurocranium rounded, lacking a crest; 
color uniform. 


Neenchelys daedalus McCosker, sp.nov. 
(Figures 4-5) 

Hototyre.—AMS I.19690-012 (field no. JP 69-29), held in 
trust for Papua New Guinea National Fish Collection, 341.5 
mm TL, a female with eggs, captured in Astrolobe Bay, s of 
Madang, Papua New Guinea (5°24’S, 145°52.5’E), 6-ft Isaacs- 
Kidd midwater trawl (IKMT), 0-140 fms [0-256m] over a 
“rough peak 500+ fms” bottom, 1850-2100 h, 7 Oct. 1969. 
In that same collection were numerous myctophids, gono- 
stomatids, and chauliodontids. 

PARATYPE.—CAS 50708, 272.9 mm TL, collected with the 
holotype. 


COUNTS AND MEASUREMENTS (in mm).—The 
condition of the holotype is parenthetically fol- 
lowed by that of the paratype: TL 341.5(272.9); 
head length 23.5(21.4); trunk length 65(53.5); tail 
length 253(198); body depth at gill openings 
6.6(5.9); body width at gill openings 4.7(4.0); body 
depth at anus 6.2(5.0); body width at anus 
4.2(3.5); origin of dorsal fin 49.5(42); length of 
pectoral fin 5.2(5.0); snout length 3.9(3.8); upper 
jaw length 6.2(5.2); eye diameter 1.2(1.2); inter- 
orbital distance 2.05(1.8). Total vertebrae 
235(225); preanal vertebrae 59(58). 

DESCRIPTION. — Body extremely elongate, depth 
at gill opening 46.3-51.7 in TL, tapering slightly 
and laterally compressed posteriorly. Tail much 
longer than head and trunk, 1.34-1.38 in TL. 
Head and trunk 3.64—-3.86 and head 12.8-14.7 
in TL. Snout moderately acute; lower jaw 
included, its tip extends to anterior base of ante- 
rior nostril. Top and sides of snout and edge of 
lower jaw covered with numerous small papillae 


(presumably sensory in function), slightly smaller 
than cephalic pores. Anterior nostril tubular, 
directed ca. 45° laterally, posterior nostril an 
elongate slit above upper lip, nearly as wide as 
orbit. Eye begins behind midpoint of jaw, mod- 
erate in size, 4.3—5.2 in jaw length. Gill openings 
low on side, each a constricted opening. 

Pectoral fin well developed, nearly as long as 
gape. Median fins well developed and obvious, 
not lying in a groove as in most myrophines. 
Anal fin larger than dorsal, nearly as deep as 
body. Dorsal fin arises about mid-trunk, the pre- 
dorsal distance 6.49-6.89 in TL. 

Head pores small but apparent (Fig. 5). Single 
temporal and interorbital pores. Five mandib- 
ular pores, and two over preopercle. Two post- 
orbital pores. Lateral line pores small but obvious; 
14 on head, 61 before anal opening, not discern- 
ible in posterior tail region. 

Teeth few, slender, conical, of moderate size 
for a myrophine. A single premaxillary tooth, 
flanked by a pair of retrorse teeth, followed by 
two medial teeth, then a pair of teeth, at which 
point the maxillae attach, each possessing 6-7 
uniserial teeth. Vomer has three teeth that end 
about midway along toothed portion of maxilla. 
Lower jaw teeth uniserial, 17-18 on each side. 

Gill arches reduced; first basibranchial absent, 
third and fourth infrapharyngobranchial tooth 
plates weakly fused. Branchiostegal rays numer- 
ous, unbranched; eight attached to hyoid (1 along 
the ceratohyal, 7 along the epihyal), 25 unat- 
tached, on each side. 

Body color in isopropyl alcohol uniform tan, 
except belly which is dark brown to black. Fins 
colorless. Eye dark blue. 


64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5 


Ficure 5. Head of holotype of Neenchelys daedalus McCosker, sp. nov., AMS I.19690-012. 


EryMoLocy.—Named daedalus, a noun in 
apposition, in honor of the Greek artisan who 
escaped from his Earth-bound prison and 
ascended into heaven. 

REMARKS.—AIll specimens of Neenchelys dae- 
dalus have come from midwater. It therefore 
seems likely that it is a midwater eel, although 
the possibility exists that juvenile and subadult 
specimens were merely transformed leptocephali 
that were captured just prior to settlement, and/ 
or the adults were benthic eels that were captured 
en route to a surface spawning event. Both pos- 
sibilities are unlikely in that the size range of the 
specimens is too great and their condition is too 
“uniform” to have been captured during trans- 
formation, and none appear to have retained lar- 
val conditions. Furthermore, the morphometric 
changes in eye size undergone by surface-mi- 
grating benthic myrophines, such as Ahlia 
egmontis (see Cohen and Dean 1970, and 
McCosker 1977), are absent. Further evidence 
for a midwater habitat is provided by the par- 
tially digested crustacean material in the gut of 
the cleared-and-stained specimen (which sug- 
gests that it had fed prior to capture and not while 
in the net) as well as the darkened vent and pres- 
ence of sensory papillae on the snout. 

The evidence thus suggests that N. daedalus is 
the second ophichthid known to have left the 


substrate to adopt a midwater life style. This 
adaptation has been independently achieved, 
however, in that its closest relatives are benthic, 
fossorial species. The other midwater ophichthid, 
Benthenchelys cartieri Fowler, lives pelagically 
at 100-250 m over deep water in the central 
Indo-Pacific (Castle 1972) and displays many 
similar adaptations, such as enlarged median fins, 
sensory papillae, an elongate tail, and slender 
teeth. 

The new species is more similar in appearance, 
owing to its extreme elongation, to the Atlantic 
species of Pseudomyrophis than to its congeners. 
However, its osteology, pore condition, and fin 
size and placement are in agreement with 
Neenchelys microtretus Bamber, 1915, N. bui- 
tendijki Weber and de Beaufort, 1916, and the 
sketchily described N. parvipectoralis Chu, Wu, 
and Jin, 1981. Castle (1980) has illustrated the 
larvae of Neenchelys and commented upon their 
distribution. The new species may be separated 
from its congeners using the characters in Ta- 
ble 1. 

In an earlier publication (McCosker 1977), I 
cautiously recognized the generic distinction 
between Neenchelys and Pseudomyrophis. My 
subsequent examination of additional osteolog- 
ical preparations of Neenchelys spp. and Pseu- 
domyrophis spp. have further substantiated those 


TasLe 1. VERTEBRAE AND Bopy PRroportIons (in thousandths of TL) oF THE Species OF NEENCHELYS. 


Head 

length 
N. microtretus 114! 
N. buitendijki 117-127! 
N. parvipectoralis ~95 
N. daedalus 68-78 


' From Nelson 1966. 
? From Castle 1980. 
3 Calculated from Chu, Wu, and Jin 1981. 


Tail Body Total 
length depth vertebrae 
580 38! 151? 
565-643 31-51! 142-148? 
~6153 ~ 60 - 
725-746 19-22 225-235 


McCOSKER: NEW WORM EELS 


differences. In an independent study, Mark M. 
Leiby (in litt., Florida Dept. of Natural Resources, 
14 Jan. 1982) has compared the leptocephali of 
species of those genera and concluded that they 
are trenchantly different. The two Pseudomyro- 
phis species that are similar in body elongation 
to Neenchelys daedalus, P. nimius Bohlke, 1960, 
and P. atlanticus Blache, 1975, inhabit mud and 
sand substrates in deep water. Dean (1972) and 
Mark Leiby (in litt.) have concluded that Myro- 
phis frio Jordan and Davis, 1891, and an unde- 
scribed Atlantic species are congeners of Pseu- 
domyrophis nimius. Two species are known from 
the eastern Pacific: P. micropinna Wade, 1946, 
the type-species; and an undescribed species 
ranging from Costa Rica to Baja California. 

Eleven other Pacific specimens, captured by 
midwater trawls, were tentatively identified as 
Neenchelys daedalus. They are all smaller spec- 
imens and appear identical in proportions to the 
new species. They differ considerably, however, 
in total vertebrae numbers: the holotype and 
paratype have 235 and 225, respectively, whereas 
eight of the others had 251-274 (X = 266.8) ver- 
tebrae. I am unable to account for such a large 
mean difference and broad range in vertebral 
number for conspecifics in such close geograph- 
ical proximity, and therefore have not made them 
type-specimens. 


COMPARATIVE MATERIAL. — Neenchelys daedalus (non-para- 
types): AMS I.19707-017 (field no. JP 69-53), 5(172-187), 
Manus Island, Papua New Guinea, 4°15’S, 145°11’E, 6-ft IKMT, 
0-125 m, over 750+ m depth, John E. Paxton aboard FRV 
TaGcuta, 0120-0320 h, 22 Oct. 1969. (Many gonostomatids 
and myctophids were captured in the same collection.) From 
the same collection: CAS 50709, 2(187—225); CAS 50710, 1(190, 
cleared and stained); and ANSP 149295, 2(175-185). SIO 77- 
171 1(144), Banda Sea, 105 km sw of Buru Is., 04°30.5’S, 
125°34.6'E, 0-1500 m over 3600 m, Jim Coatsworth, 26 Aug. 
1976. 

Neenchelys buitendijki: ZMA.102.171, 1(218), syntype, 
“probably from Moluccos,” Indonesia. UH uncat., 2(1 18-123, 
specimens dissected), Bombay City, India (specimens from 
Mohamed 1958, reported on by Nelson 1966). 

Neenchelys microtretus: BMNH 1915.10.25.1, 1(183), holo- 
type, Red Sea. 

Pseudomyrophis micropinna: LACM 21557, 1(139), holo- 
type, Isla Ladrones, Gulf of Chiriqui, Panama. SIO 60-72, head 
and trunk only, cleared and stained. 

Pseudomyrophis nimius: USNM 186274, 1(319), holotype, 
Gulf of Mexico. ANSP 110150, 1(350, cleared and stained), 
Gulf of Mexico. 

Pseudomyrophis atlanticus: MNHN 1971-40, 1(259) holo- 
type, Pointe-Noire. MNHN 1971-41, 1(241), paratype, Pointe- 
Noire. 


65 


ACKNOWLEDGMENTS 


Specimens and assistance were generously pro- 
vided by the staffs of many institutions. In par- 
ticular, I thank: John Paxton and Doug Hoese, 
Australian Museum, Sydney (AMS); Eugenia 
Bohlke and the late James B6hlke, Academy of 
Natural Sciences of Philadelphia (ANSP); the staff 
of the British Museum (Natural History) 
(BMNH); John E. Randall, Bernice P. Bishop 
Museum (BPBM); William N. Eschmeyer and 
staff, California Academy of Sciences (CAS); 
Robert Lavenberg, Natural History Museum of 
Los Angeles County (LACM); Jacques Blache 
and Marie-Louise Bauchot, Muséum National 
d’Histoire Naturelle, Paris (MNHN); Richard H. 
Rosenblatt, Scripps Institution of Oceanography 
(SIO); William Bussing, University of Costa Rica 
(UCR); Leighton R. Taylor, Jr., Waikiki Aquar- 
ium and University of Hawaii (UH); the staff of 
the United States National Museum of Natural 
History (USNM); and Hans Nissen, ZoGdlogisch 
Museum Amsterdam (ZMA). I also thank: Beth 
A. Meinhard for the preparation of Figures 1-3; 
Kathy Smith for the preparation of Figures 4-5; 
Michael Hearne for the preparation of radio- 
graphs; Lillian Dempster for nomenclatural 
assistance; and Mark Leiby, Peter Castle, and 
Eugenia Bohlke for their stimulating discussions 
concerning the relationships of worm eels. 


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yo 


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sis. Pac. Sci. 24(4):506—-516. : 

. 1977. The osteology, classification and relationships 
of the eel family Ophichthidae. Proc. Calif. Acad. Sci., ser. 
4, 41:1-123. 

Mouamen, K.H. 1958. On the occurrence of the eel Neenche- 
lys buitendijki Weber & deBeaufort in Indian waters. J. Bom- 
bay Nat. Hist. Soc. 55:511-517. 

NE son, G. J. 1966. Osteology and relationships of the eel 
Neenchelys buitendijki. Copeia 1966:32 1-324. 

Taytor, W. R. 1967. An enzyme method of clearing and 
staining small vertebrates. Proc. U.S. Natl. Mus. 122(3596): 
1-17. 

Wape, C. B. 1946. Two new genera and five new species of 
apodal fishes from the eastern Pacific. Allan Hancock Pac. 
Exped. 9(7):181-213. 

Weber, M., AND L. F. DEBEAUFORT. 1916. The fishes of the 
Indo-Australian Archipelago. III. Ostariophysi: II Cypri- 
noidea, Apodes, Synbranchi. E. J. Brill, Leiden. 455 p. 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


PROCEEDINGS 
: OF THE 


CALIFORNIA ACADEMY OF S 


Vol. 43, No. 6, pp. 67-76, 3 figs., 3 tables. 


July 6, 1983 


PSILORHYNCHUS GRACILIS, A NEW CYPRINOID FISH 
FROM THE GANGETIC LOWLANDS 


By 
Walter J. Rainboth 


Department of Biology, University of California, Los Angeles, 
Los Angeles, California 90024 


Asstract: Psilorhynchus gracilis is described from 147 specimens collected at 10 lowland localities in 
Bangladesh. Of the four previously described species of Psilorhynchus (balitora, homaloptera, pseudecheneis, 
and sucatio), the new species resembles P. balitora most closely. The new species is compared with P. balitora 
and P. sucatio, both of which are syntopic species of the Gangetic plains. P. sucatio may be distinguished 
from P. balitora and the new species by its reduced dorsal ray count and numerous shape and coloration 
differences. P. balitora differs from the new species in lateral line and circumferential scale counts, number 
of unbranched pectoral rays, coloration, and shape of the head and mouth parts. The preferred habitat and 
distribution of the three lowland species are compared, and a key to the genus Psilorhynchus is provided. 


INTRODUCTION 


Fishes of the genus Psilorhynchus McClelland 
are known to occur primarily in the Gangetic 
drainage of southern Asia, where four described 
(Menon 1974) and one undescribed species are 
found. Most ichthyologists (following Hora 1925) 
have recognized Psilorhynchus as the sole genus 
in the family Psilorhynchidae. A recent re-anal- 
ysis of the relationships of Psilorhynchus has been 
published by Chen (1980), who believes the genus 
to belong to the Cyprinidae. I have not located 
a copy of that publication and must reserve com- 
ment at this time. 

Of the five known species, two (Psilorhynchus 
homaloptera Hora and Mukerji, 1925, and P. 
pseudecheneis Menon and Datta, 1964) have rel- 
atively smaller scales and greater numbers of 
simple pectoral rays than the remaining three 
species. These small-scaled species also prefer 
high-gradient streams located in the eastern 
Nepalese Himalayas (P. pseudecheneis) and the 


Naga Hills of the Assam-Burma border (P. hom- 
aloptera). The subspecies P. homaloptera rowleyi 
Hora and Misra, 1941, of the Chindwin River 
(Irrawaddy) is one of the two species of this genus 
from Burma (Fig. 1). These taxa from high-gra- 
dient streams are not known to present any taxo- 
nomic problems. 

Two species from easily accessible lowland 
areas were described by Hamilton in 1822 as 
Cyprinus sucatio and C. balitora. While collect- 
ing in the People’s Republic of Bangladesh in 
1977 and 1978, I obtained three species from 
Gangetic lowland streams. Two of the species 
were those described by Hamilton and one is 
described, herein, as new. 

Hamilton’s (1822) somewhat brief original 
descriptions of two lowland species now referred 
to Psilorhynchus were not accompanied by fig- 
ures, although he had prepared figures for them 
during his stay in India. Hamilton’s figures are 
of prime importance because he kept no pre- 


[67] 


68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6 


hs 


FiGure 1. 


Distribution of the known species of Psilorhynchus. Inset map indicates region displayed. Darkened circles represent 


P. gracilis localities, with triangle marking the type-locality. Open circles are localities for P. balitora, with question marks in 
outlying non-verified reports. Two localities have both P. gracilis and P. balitora. Vertical hatching covers the range of P. 
pseudecheneis. Diagonal hatching shows the range of P. homaloptera. Stippling highlights the known range of P. sucatio. 


served material. However, only half of the figures 
(and neither of the Psilorhynchus figures) were 
published in Hamilton’s original monograph. The 
remaining figures were published subsequently 
by several authors including McClelland (1839), 
who published Hamilton’s figures of Psilorhyn- 
chus. The reasons for this were explained par- 
tially by McClelland (1839) and more fully doc- 
umented by Gudger (1924) and Hora (1929). 
Although he had seen but a single individual of 
one of the species, McClelland placed both species 
in the new genus Psilorhynchus and presumed 
that his single specimen was P. balitora (Ham- 
ilton), for which he coined a new specific name, 
variegatus, to replace balitora—a local term 
meaning sand-digger. McClelland preferred the 
use of classical Latin or Greek to the use of local 
dialects; for example, he changed the homalop- 
terid generic name Balitora Gray, 1834, to Pla- 
tycara. Such replacement of names is inadmis- 
sible under the present rules of nomenclature. 
In Hamilton’s figures (McClelland 1839:pl. 50), 
Psilorhynchus sucatio is easily recognized; how- 
ever, I collected two species which resemble the 
illustration of P. balitora. Because McClelland’s 
P. variegatus refers to an individual ofa balitora- 
like species, the possibility that his species may 


have been distinct from Hamilton’s will also be 
explored. 


MATERIALS AND METHODS 


Measurements were made to the nearest 0.1 
mm with dial calipers. Standard length was mea- 
sured from the snout tip to the base of the caudal 
fin. All pre-fin lengths extend from the tip of the 
snout to the base of the first unbranched ray at 
fin origin. Dorsal and anal fin lengths are mea- 
sures of the longest simple ray, whereas the paired 
fins are depressed and measured from insertion 
to posteriormost extension. Caudal fin length is 
measured from the posterior end of the urocen- 
trum to the tip of the normally spread upper fork. 
Body depth is measured from dorsal fin origin 
to pelvic fin insertion. The snout to occiput length 
(head length = HL) is the distance to the poste- 
rior margin of the supraoccipital bone. The snout 
to preopercle length is taken on a horizontal. 
Snout to maxilla length is measured to the pos- 
terior end of the maxilla. Orbital measurements 
are taken to the bony margin. Mandible length 
is distance from the symphysis to articulation 
with the quadrate. Gape width is the distance 
between the two articulation points of upper and 
lower jaw. 


RAINBOTH: NEW PSILORHYNCHUS 


Fin-ray counts are expressed with lower case 
Roman numerals signifying unbranched rays and 
Arabic numérals for branched rays. The deeply 
divided final branched ray in both the dorsal and 
anal fin is counted as one. Caudal counts list 
procurrent rays of the upper fork as Roman 
numerals with Arabic numerals for principal rays 
of the upper/lower forks followed by Roman 
numerals giving lower-fork procurrent rays. Lat- 
eral line scales were counted for the body and 
tallied separately from those on the caudal fin 
base. Lateral transverse counts include the 
median scale at the dorsal fin origin, record the 
lateral line with a slash (/), and include the median 
ventral row before the anal fin. Circumferential 
counts encircle the body on the scale row imme- 
diately anterior to the dorsal and pelvic fins. Cir- 
cumpeduncular counts include all scales around 
the peduncle at its narrowest region. The number 
of anal scales refers to median scale rows between 
the anus and anal fin. Belly scale rows include 
all complete, free-edged midventral scale rows 
crossing anterior to the pelvic fin insertion. Counts 
given for lateral blotches include all distinct mid- 
lateral blotches whether or not they are perfectly 
bisected by the lateral line. 

Body measurements are summarized as per- 
cent standard length (% SL), head measurements 
as percent head length (% HL). 

Most of the material was collected by the author 
and is housed at the Museum of Zoology, The 
University of Michigan, Ann Arbor (UMMZ) or 
at the Chandpur Freshwater Fisheries Research 
Station, Chandpur, Bangladesh (CFRS). Other 
specimens are from the American Museum of 
Natural History, New York (AMNH), and the 
Stanford University (SU) collection now housed 
at the California Academy of Sciences, San Fran- 
cisco. Paratypes of the new species have been 
deposited at all aforementioned institutions plus 
the Field Musuem of Natural History, Chicago 
(FMNH), the Academy of Natural Sciences of 
Philadelphia (ANSP), and the United States 
National Museum, Washington, D.C. (USNM). 

Distribution maps are part of drainage maps 
of the southern half of the Asian continent drafted 
by the author from the most recent world-wide 
series of 1:5,000,000-scale topographic maps 
prepared by the U.S. Defense Mapping Agency. 


Psilorhynchus McClelland 


Psilorhynchus MCCLELLAND, 1839:300, 428 (type-species 
Cyprinus sucatio Hamilton, by subsequent designation of 
Jordan 1919). 


69 


D1aGnosis.— Body arched dorsally and flat- 
tened ventrally. Anteriorly depressed, becoming 
cylindrical with lateral compression posteriorly. 
Ventral surface of head markedly flattened. 
Mouth small, inferior, and transverse with a pro- 
jecting snout; devoid of barbels. Pharyngeal teeth 
uniserial. Gill membranes joined broadly to isth- 
mus with aperture extending ventrally to base of 
pectoral fin. Paired fins inserted horizontally. 
Breast naked. Scales moderate to large, 31 to 50 
pored scales on the complete lateral line. Fin-ray 
P,1i/7. Anus very close to pelvic fin; at least 8 
scale rows separate anus from anal fin. 


Key to the Species of Psilorhynchus 


las Branched dorsal finsrays 7 22 ee 2 
1b. Branched dorsal rays 8 (or 9) _. 4 


2a. Abdomen fully scaled; simple pectoral rays 
Ae Re atl cence ee A sucatio (Hamilton) 
2b. Abdomen naked; simple pectoral rays 7- 
(pee ree ire ld 2 a Sek ee ST 3 


3a. Total lateral line scales 42-44; simple pec- 
toral rays 7 or 8; abdomen smooth _.... 
LEC Nag Rte! homaloptera Hora and Mukerji 

3b. Total lateral line scales 48—50; simple pec- 
toral rays 10; abdomen with three trans- 
VETSEstOl SO fis Kelty ee ene ee ae eee 


4a. Gape width greater than mandible length; 
simple pectoral rays 6—7 (rarely 5) or more; 
circumiferentialiscales 1G). sa 
i OR naa balitora (Hamilton) 

4b. Gape width much less than mandible 
length; simple pectoral rays 4—5; circum- 
ferentiall'scales 16) gracilis sp.nov. 


Psilorhynchus gracilis sp.nov. 
(Figures 2 and 3) 


Ho.otyre.—UMMZ 205342 (adult female, 50.5 mm SL), 
Jabuneswari River at Badarganj, Rangpur Dist., Bangladesh, 
3 Apr. 1978. 

ParatyPes (all from Bangladesh).—UMMZ 205343 (26 
specimens, 29.4-49.9 mm SL), and CFRS uncat. (9 spec.) same 
collection data as holotype. UMMZ 205337 (2 spec., 25.7- 
29.7), Sangu River at Bandarban, Chittagong Hill Tracts, 25 
Dec. 1977; UMMZ 205345 (8, 34.1-40.7), Ghaghat River at 
Rangpur, Rangpur Dist., 3 Apr. 1978; UMMZ 205348 (31, 
26.6-51.2), USNM 231693 (5), AMNH 43097 (5), CAS 50011 
(5), and ANSP 148729 (5), and FMNH 94285 (5), Mahananda 
River at Tetulia, Dinajpur Dist., 5 Apr. 1978; UMMZ 205351 
(15, 27.6-46.4), Keratoya River at Bhajanpur, Dinajpur Dist., 
6 Apr. 1978; UMMZ 205353 (2, 34.3-39.0), Tangam River 
at Thakurgaon, Dinajpur Dist., 6 Apr. 1978. 

OTHER MATERIAL EXAMINED (all from Bangladesh). -UMMZ 


70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6 


Ficure 2. Young specimens of three lowland species of Psilorhynchus from Bangladesh. (top) P. gracilis, 34 mm SL, 
Jabuneswari River. (middle) P. balitora, 20 mm SL, Mahananda River. (bottom) P. sucatio, 64 mm SL, Rangapani Creek. 


205340 (18, 10.4-19.0), Dharla River at Kurigram, Rangpur 
Dist., 2 Apr. 1978; CFRS uncat. (3, 33.8-—35.4), Muhuri River, 
10 km ne of Feni, Noakhali Dist., 2 Feb. 1978; CFRS uncat. 
(1, 40), Halda River at Daulatpur, 40 km n of Chittagong, 
Chittagong Dist., 24 Feb. 1978; CFRS uncat. (6) Halda Creek 
at Khaia Bazaar, 53 km n of Chittagong, Chittagong Dist., 24 
Feb. 1978. 


Diacnosis.— Lateral line scales 33 to 36 on 
body plus | or 2 pored scales on caudal fin, to- 


talling 35 to 37; pectoral fin with 4 or 5 simple 
rays; branched dorsal rays 8 (rarely 9); 2 distinct 
dorsal spots anterior to dorsal fin origin with a 
third at the origin; midventral region scaleless 
anteriorly, with 2 to 5 complete, free-edged scale 
rows immediately anterior to pelvic fin insertion; 
pectoral fin short, extending beneath dorsal fin 
origin, but never reaching pelvic insertion; eye 


RAINBOTH: NEW PSILORHYNCHUS 


71 


FiGure 3. 
balitora, and right is P. gracilis. In ventral aspect (A), P. balitora, 39 mm SL, Siliguri, and P. gracilis, 47 mm SL, Jabuneswari 
River. In dorsal aspect (B), P. balitora, 27 mm SL, Mahananda River, and P. gracilis, 32 mm SL, Jabuneswari River. 


in upper part of head, not visible from below; 
no spots on anal or paired fins, although mem- 
brane between first 5 pectoral rays slightly dark- 
ened in some adults. 

DESCRIPTION. —Predorsal scales 10 or 11; cir- 
cumferential scales 16 (rarely 15 or 17); circum- 
peduncular scales 10; lateral transverse scale-rows 
4/3; scale rows between anus and anal fin 8 or 
9. Scales one row above lateral line immediately 
posterior to dorsal fin have 6 or 7 radii in adults 
of 50 mm SL. 

Paired fins inserted horizontally; pectoral with 
4 or 5 unbranched rays and 9-12 branched rays, 
totalling 14-17 elements, extending as far as dor- 
sal fin origin when depressed, with distal margin 
separated from pelvic fin insertion by at least 2 
scale rows; pelvic fin inserted slightly posterior 
to dorsal fin origin, with 2 simple and 7 branched 
rays. Dorsal fin closer to snout tip than to caudal 
base, with 3 simple and 8 (rarely 9) branched 
rays. Anterior and posterior rays of equal exten- 
sion in depressed dorsal fin, which when nor- 
mally expanded has an oblique and concave pos- 
terior margin. Anal fin short and somewhat 


Comparison of ventral (A) and dorsal (B) aspects in Psilorhynchus balitora and P. gracilis. In each pair, left is P. 


falcate, its posterior margin well in advance of 
caudal fin, with 3 simple and 5 branched rays. 
Caudal fin deeply and evenly forked, possessing 
19 (10/9) principal rays preceded by 6 or 7 pro- 
current rays above and 5 or 6 below. Body elon- 
gate, arched above and generally flat below; 
greatest depth at dorsal fin origin and greatest 
width at pectoral fin insertion; shape somewhat 
depressed anteriorly, gradually becoming cylin- 
drical, then compressed posteriorly. 

Head depressed, ventral mouth small and 
transverse. Upper lip separate from upper jaw 
by a deep groove and from snout by a shallow 
groove. Ventral surface of snout separated from 
lateral surfaces by deep rostral grooves. Upper 
lip joined to lower lip at corner of mouth by a 
fairly prominent flap of skin. Lower lip thick and 
followed on chin by a number of large, globular 
papillae which decrease in size posteriorly. Lower 
jaw long, its length much greater than width of 
gape. Gill membranes broadly joined to isthmus; 
gill aperture extends dorsally from pectoral fin 
base. Eye large, upper margin level with flat inter- 
orbital space; width of orbit approximating inter- 


72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6 


orbital width. Tip of snout somewhat pointed if 
viewed from dorsal aspect. 

Nuptial tubercles evident on large males, ran- 
domly distributed on head. Heaviest tubercle 
accumulation below eye on lower part of cheek 
where eruption takes form of tight clusters rather 
than usual single pattern. Anterior body scales 
have tubercles bordering their free margins. 
Tubercle distribution on body progressively 
sparser posteriorly. Pectoral fins on breeding 
males show single row of tubercles ornamenting 
dorsal surface of each ray, with tubercles on ante- 
rior rays largest. Pelvic fins with same pattern 
but with smaller tubercles. Large females have 
same tubercle distribution as males but with slight 
differences. Females have lower tubercle density 
than males, lack tubercles on paired fins, and do 
not display tubercles in the clumped suborbital 
pattern of males. 

COLORATION.—Generally recognizable in 
specimens greater than 15 mm SL, although vari- 
able. Laterally, a series of 7-10 dark blotches 
with posteriormost extending onto caudal fin. 
First lateral blotch midway between lateral line 
and base of pectoral fin in small specimens. This 
spot enlarges with age, to cover lateral line. Lat- 
eral line always bisects remaining large lateral 
spots. Middorsal spots discontinuous with lateral 
spots; dorsal spots on same transverse scale rows 
as lateral blotches, having identical counts on all 
specimens. Where fish has unequal lateral spot 
counts, some dorsal spots obliquely cross mid- 
dorsal line, and on each side are still on same 
scale rows as lateral spots. Two distinct dorsal 
spots anterior to dorsal fin origin, with another 
at origin and a fourth under posterior dorsal fin 
rays. Four or five spots between dorsal fin and 
caudal fin. All scales of back and upper side have 
marginal melanophores connecting to create net- 
work ventrally to just below lateral line. Network 
darkens with age, but always distinct. Head dark 
dorsally with light median-longitudinal streak 
flecked with large discrete melanophores from 
snout to posterior edge of occiput. A dark band 
projects from each nostril to coalesce anteriorly 
at tip of snout. Pigment lacking on fins of small 
specimens, appearing gradually at 30 mm SLand 
increasing substantially with size. Caudal fin ulti- 
mately has 2 black blotches per lobe, with prox- 
imal spot often joining median spot at caudal 
base. Dorsal fin has apical spot which runs along 
anterior 2 or 3 rays. No blotches on paired fins, 


although large specimens have darkened mem- 
branes between first 4 or 5 pectoral rays. Peri- 
toneum dense black dorsally, becoming an irreg- 
ular medium to dark gray below. 

EtyMoLoGy.—The Latin adjective gracilis 
(gender masculine) was chosen because of the 
fish’s slender shape, which immediately distin- 
guishes the new species from Psilorhynchus bal- 
itora, the most similar species. 


COMPARISON OF SPECIES 


Several counts aid in distinguishing the three 
lowland species. The diagnostic counts (Table 1) 
demonstrate resemblances between P. gracilis and 
P. balitora in dorsal rays and total pectoral fin 
elements. However, lateral line and circumfer- 
ential counts are closest between P. gracilis and 
P. sucatio. Belly squamation is distinctive in P. 
gracilis, which never has more than two to five 
midventral scale rows anterior to the pelvic fins. 
P. sucatio always has a fully scaled belly and 
sympatric specimens of P. balitora never exhibit 
fewer than eight midventral prepelvic scale rows. 
Specimens from the Chindwin River, Burma, and 
the Rapti River, Nepal have naked breasts and 
bellies, but are typical P. balitora in all other 
respects. Both P. homaloptera and P. pseude- 
cheneis have scaleless bellies. Two AMNH 15767 
paratypes of P. homaloptera from the Naga Hills 
were incorrectly identified. One is P. balitora with 
typical counts and measurements which have 
been included in all four tables. The other spec- 
imen is a loach (genus Noemacheilus). 

Several proportional measurements show 
notable differences. However, overlap in pro- 
portional measurements may occur when juve- 
niles are included, even though adult proportions 
display pronounced differences. For instance, of 
seven measured P. balitora, four adults were over 
40 mm SL and three juveniles were less than 30 
mm SL. The robust adults have a body depth of 
25.5% SL, whereas the juveniles have a body 
depth of 19.4% SL (Table 2). Juvenile propor- 
tions of P. balitora overlap with those of adults 
of the other two species. Other proportions dis- 
tinguishing P. gracilis from P. balitora at any size 
are those for the anal, pectoral, and pelvic fins. 
Proportions (taken as % HL) differing consis- 
tently in these two species are interorbital width, 
gape width, and mandible length (Table 2). A 
single perfectly discriminating character is gape 
width, which is considerably less than the man- 


RAINBOTH: NEW PSILORHYNCHUS 


73 


Taste 1. DiaGNostic COUNTS FOR THE THREE LOWLAND Species OF Ps/LoRHYNCHUS. Number counted in parentheses. 
P. balitora P. gracilis P. sucatio 
Dorsal fin rays iii/8 (17) 111/8—9 (18) 1/7 (18) 
Pectoral fin rays v—vili/7—9 (41) iv—v/9-12 (60) iv/8-9 (18) 
Caudal fin rays v—vi, 9/8, iv—v (10) vi-v1i, 10/9, v—vi (18) ili-iv, 9/9, iii (18) 
Lateral line scales 30-34 + 1 or 2 (41) 33-36 + | or 2 (60) 34-35 + 1 or 2 (18) 
Circumferential scales 17-20 (41) 15-17 (60) 16-18 (18) 
Anai scales 9-10 (17) 8-9 (18) 8-11 (13) 


dible length in P. gracilis, whereas the size re- 
lationship is reversed in P. balitora. 

Psilorhynchus gracilis differs from P. sucatio 
in the depth of the caudal peduncle and several 
head-measurement proportions, most notably the 
interorbital width: approximating the orbital 
width in P. gracilis and nearly doubling the orbital 
width in P. sucatio. Also, in P. sucatio the ante- 
rior dorsal rays show greater extension when 
depressed, whereas P. gracilis and P. balitora 
have equal or greater posterior extension of the 
last rays. 

Color patterns are most similar between P. 
gracilis and P. balitora in younger specimens. 
Therefore, sub-adults have been illustrated (Fig. 
2). Juveniles of P. sucatio have a continuous black 
midlateral stripe extending from opercle to cau- 
dal fin. The stripe fades with age, and gradual 
coalescence of blotches gives the adult color pat- 
tern. Remnants of the stripe can be seen on the 
figured specimen as discrete melanophores 


between the lateral spots. Dorsally, P. sucatio 
exhibits darkening of entire scales rather than 
the reticulated network found in P. gracilis and 
P. balitora. Both P. gracilis and P. balitora always 
have a series of lateral blotches with smaller and 
more numerous spots on P. gracilis (Table 3). 
The lateral spots on P. gracilis and P. balitora 
are on the same diagonal scale rows as the dorsal 
spots. On P. gracilis dorsal and lateral spots are 
totally distinct, whereas P. balitora exhibits a 
faint continuous band between the lateral and 
dorsal spots. The predorsal spot pattern (Fig. 3) 
for P. gracilis is two distinct blotches with a third 
at the dorsal fin origin, and for P. balitoraa single 
blotch, with a second beginning at the dorsal 
origin. In adult P. balitora the dorsum gradually 
darkens causing the pattern to become somewhat 
obscure, although it still persists. P. gracilis retains 
a distinct reticulated pattern throughout adult 
life. 

A recently described taxon Psilorhynchus 


TABLE 2. PROPORTIONAL MEASUREMENTS FOR THREE SPECIES OF Ps/LoRHYNCHUS. Characters 2 through 9 expressed as %SL. 


Characters 10 through 15 expressed as %HL (snout to occiput). 


P. balitora P. gracilis P. sucatio 
Range (mm) ESD Range (mm) X+sD Range (mm) X+spD 
1. Standard length 22.7-47.8 (n = 7) 28.2-51.2 (n = 10) 26.4-64.3 (n = 13) 
2. Snout to dorsal fin 48.1-52.8 S10).3) Se 21! 44.6-47.9 46.8 + 1.1 43.9-48.9 46.3 + 1.6 
3. Snout to pectoral fin 22.4-23.9 DED OES 18.8-22.7 DONE IES 19.7-23.1 ZALO) =a 15 
4. Body depth 18.5-26.9 7S) ae 3h5) 18.0-21.4 WET) se is) 15.7-21.4 11823) = 1k9 
5. Peduncle depth 8.2-9.4 8.9 + 0.4 7.5-8.4 8.0 + 0.3 6.1-7.3 7.0 + 0.4 
6. Pectoral fin length 25.9-28.7 Aye ae | (0) 21.0-23.4 22.4 + 0.8 17.3-22.7 20.6 + 1.6 
7. Pelvic fin length 20.7-22.9 Piles) ae (03! 16.9-19.9 18.4 + 1.0 16.4-19.3 18.4 + 1.0 
8. Anal fin height 15.4-18.0 17.0 + 0.9 14.2-15.2 14.8 + 0.4 12.1-14.4 13.0 + 0.9 
9. Snout to occiput 23.0-24.8 24.1 + 0.6 20.3-24.0 2210 == les 20.4-23.9 PS se 10) 
10. Snout to maxilla 32.3-35.6 3316) es 23.6-31.0 27.8 + 2.4 25.5-32.2 31.0 + 0.9 
11. Orbit width 31.3-35.4 33.4 + 1.5 30.3-33.8 31.8 + 1.0 25.2-30.5 PUT) asl) 
12. Interorbital width 36.5-41.6 38.6 + 2.0 29.8-36.3 3317 == 210 42.8-56.5 50.0 + 4.7 
13. Gape width 26.8-31.6 28.4 + 2.0 19.1-24.7 D2 EG 23.3-28.2 23162 eG 
14. Mandible length 19.6-23.6 D4 15 29.4-42.5 33.9 + 4.1 25.0-32.7 Pfs) as P)3\ 
15. Head depth at pupil 46.4-55.3 SOEs 39.4-47.3 43.8 + 2.8 33.6-42.0 38:6) 22 227 


74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6 


TABLE 3. DISTRIBUTION OF VALUES FOR CHARACTERS 
DISTINGUISHING PsILORHYNCHUS GRACILIS AND P. BALITORA. 
Counts for P. gracilis holotype are underlined. 


Total lateral line scales 32 3304S Ono, 


P. gracilis 0:91 Oke 10544 0m, Sel 
P. balitora 3 Al 4 1 0 


Simple pectoral rays 


P. gracilis 82 0 0 0 
P. balitora 0 5 19 16 1 
Circumferential scales 1S Gl Si ol O20 
P. gracilis Ser 2) 0) ping 0 
P. balitora 0 0 AD 26, 33 8 
Lateral blotches 5 6 7 8 9 10 
P. gracilis 0) ® is sky 9 2. 
P. balitora 23 «16 D 0 0 0 


sucatio var. damodarai David, 1953, appears to 
be a local race of P. sucatio with slightly larger 
pectoral fins than those found on individuals from 
Bangladesh. The Damodar River variety is also 
listed as having “8” branched dorsal rays as 
opposed to “‘7—8”’ for Gangetic specimens. I have 
not seen any specimens with eight branched dor- 
sal rays, although the deeply divided last ray could 
be erroneously counted as such. 


DISCUSSION 


Of the characteristics given for identification 
in Hamilton’s description (1822), few would be 
of use in distinguishing species as similar as P. 
gracilis and P. balitora. However, Hamilton does 
state that there are approximately 12 rays in each 
pectoral fin. This count is closer to what I found 
for P. balitora (minimum 13) and fewer than in 
the new species (Table 3). 

Hamilton’s figure of the dorsal aspect repro- 
duced in McClelland (1839) shows six simple 
rays in each pectoral fin, and anterior-dorsum 
and head color patterns identical to my obser- 
vations for P. balitora. Hamilton’s figure dis- 
plays the anterior part of the dorsum on P. bal- 
itora with one blotch midway between the occiput 
and the dorsal fin, and another at the dorsal fin 
origin on both lateral and dorsal views. The dor- 
sal coloration of the head has two black spots 
separated by a transverse white line in the inter- 
orbital space. There is no longitudinal white streak 
from the snout to occiput as on the new species 


(Fig. 3). There are 34 total scale rows in longi- 
tudinal series, which better describes P. balitora 
(Table 3). Although the lateral blotches on Ham- 
ilton’s illustration of the lateral aspect might cor- 
respond to the new species, six lateral spots are 
commonly seen in P. balitora, which also exhibits 
a lengthened anterior blotch similar to that in 
Hamilton’s figure. The total dorsal spots illus- 
trated in dorsal aspect might be high for P. bal- 
itora, however, even though both lateral and dor- 
sal views are presumably taken from the same 
fish, the blotches do not match (the lateral view 
has one fewer dorsal spot on the peduncle). 
Because the lateral view shows an equal number 
of lateral and dorsal blotches, that would pre- 
sumably be a better indicator of the dorsal spot 
pattern on the caudal peduncle than the illus- 
trated dorsal aspect. From these characters it 
would seem that Hamilton’s Cyprinus balitora 
is synonymous with my Psilorhynchus balitora 
rather than with the new species. 

Psilorhynchus variegatus McClelland remains 
as a potential name for the new species. 
McClelland (1839:430) stated that his single 
specimen differed little from Hamilton’s balitora 
except for a few features. McClelland listed 17 
rays in the pectoral fins and 33 scales in the lateral 
line. The total of 17 pectoral fin elements occurs 
in both species as a maximum, but 33 lateral line 
scales is the mode for P. balitora and two scales 
below the minimum total count found in P. gra- 
cilis (Table 3). Therefore, it appears that P. var- 
iegatus McClelland is indeed a synonym of P. 
balitora (Hamilton) and that P. gracilis is a new 
species. 

Other accounts in Day (1878) and Shaw and 
Shebbeare (1937) apply to P. balitora. No pub- 
lished account appears to have included the new 
species under the name P. balitora, which is rather 
surprising in view of its abundance and appar- 
ently widespread occurrence in the Gangetic low- 
lands. 


DISTRIBUTION AND HABITAT 
PREFERENCE 


Thus far the new species has been collected 
only in Bangladesh. However, the extent of its 
preferred habitat would suggest a much wider 
distribution throughout the lower reaches of the 
Ganges and Brahmaputra rivers. The southern- 
most collections of Psilorhynchus gracilis in 
Bangladesh are from rivers that currently have 


RAINBOTH: NEW PSILORHYNCHUS 


independent exits into the Bay of Bengal (Fig. 1). 
The species is also fairly common in sandy 
streams of northwest Bangladesh. Psilorhynchus 
gracilis was taken in the same collections with 
P. balitoraand P. sucatio in the Mahananda River 
at Tetulia and the Keratoya River at Bhajanpur, 
both in Dinajpur District. P. sucatio was taken 
at each of my collection localities for P. gracilis 
except one site from the Muhuri River (Feni River 
drainage) of Noakhali District in southeast Bang- 
ladesh. However, P. sucatio was taken in two 
other collections from the Feni River drainage. 

Psilorhynchus gracilis is found over small peb- 
bles in shallow running waters where the bottom 
is primarily sand. In this regard it resembles P. 
balitora which is sometimes found in the same 
habitat but which is always closely associated 
with hard substrates. P. gracilis is generally free- 
swimming and occasionally rests on its spread 
paired fins. P. balitora prefers to maintain close 
fin contact with the substrate, often not moving 
unless strongly disturbed. Psilorhynchus gracilis 
may be caught easily on pebble outcrops having 
both species present because of its greater ten- 
dency to leave the bottom. A seine pulled under 
or through the gravel of the same outcrop catches 
P. balitora, which will be taken out along with 
the substratum. P. sucatio differs from the other 
lowland species in being taken primarily along 
the edges of sandy streams, and seems to be most 
abundant near emergent or overhanging vege- 
tation. I have not observed Psilorhynchus bur- 
rowing, although several species of loaches were 
seen to burrow into sand within inches of Psi- 
lorhynchus under observation. Attempts to elicit 
burrowing by disturbing the fishes were not suc- 
cessful. 

The P. balitora taken at the same localities as 
P. gracilis were all small, about half to two-thirds 
adult size. This possibly indicates that fully grown 
individuals occur upstream in areas of higher 
gradient. It is also consistent with the physical 
and behavioral characteristics of the species. P. 
balitora has much larger pectoral fins with more 
simple rays, and a wider and higher body pre- 
dorsally. It depresses its head when positioned 
in an area of strong current, and is forced down 
onto its fins. This shape is common among Asian 
hillstream fishes which attach themselves to hard 
substrata in high-gradient streams (e.g., Garra, 
Homaloptera, Gastromyzon). This somewhat 
passive posturing for increased friction is a much 


15 


more efficient method than constant swimming 
for maintaining position in the current of tor- 
rential streams. Thus, it would appear that north- 
ern Bangladesh may be the southernmost region 
of the Gangetic plain occupied by P. balitora, a 
species adapted to higher gradients than either 
P. gracilis or P. sucatio. 


COMPARATIVE MATERIAL EXAMINED 


Psilorhynchus balitora—Inv1a: SU 28701 (2 specimens, 42.8— 
45.2 mm SL), Siliguri, North Bengal, no date given; SU 32627 
(1, 40.2) Siliguri, North Bengal, Apr. 1937; AMNH 15767 (1, 
47.8), Keleki Stream at Emilioni, Naga Hills, Assam (paratype 
of Psilorhynchus homaloptera Hora and Mukerji). NepaAt: 
UMMZ 207678 (7, 33.1-37.2), Rapti River at Chitawan Val- 
ley, Apr.-May 1975. Burma: AMNH 13811 (14, 32.5-42.8), 
Upper Burma, Chindwin drainage. BANGLADESH: UMMZ 
205347 (19, 15.6—26.7) Mahananda River at Tetulia, Dinajpur 
Dist., 5 Apr. 1978; CFRS uncat. (15) same data, UMMZ 205350 
(11, 14.1-18.6), Keratoya River at Bhajanpur, Dinajpur Dist., 
6 Apr. 1978. 

Psilorhynchus sucatio—BANGLADESH: UMMZ 205338 (5 
specimens, 20.2—22.6 mm SL), Sangu River at Bandarban, 
Chittagong Hill Tracts, 25 Dec. 1977; UMMZ 205339 (32, 
46.3-73.3), Rangapani Creek, 6 km nnw of Jaintapur, Sylhet 
Dist., 19 Feb. 1978; UMMZ 205341 (5, 12.5-17.5) Dharla 
River at Kurigram, Rangpur Dist., 2 Apr. 1978; UMMZ 205344 
(5, 15.5-55.6), Jabuneswari River at Badarganj, Rangpur Dist., 
3 Apr. 1978; UMMZ 205346 (57, 13.8—-58.8), Ghaghat River 
at Rangpur, Rangpur Dist., 3 Apr. 1978; UMMZ 205349 (42, 
15.7-41.8), Mahananda River at Tetulia, Dinajpur Dist., 5 
Apr. 1978; UMMZ 205352 (22, 11.1-50.7), Keratoya River 
at Bhajanpur, Dinajpur Dist., 6 Apr. 1978; UMMZ 205354 
(37, 12.7-45.7), USNM 231694 (5), AMNH 43096 (5), and 
50010 (5), ANSP 148728 (5), and FMNH 94284 (5), Tangam 
River at Thakurgaon, 6 Apr. 1973; CFRS uncat. (19, 34.2- 
71.9), Koilla Creek, 13 km w of Ramgarh, Chittagong Dist., 
3 Feb. 1978; CFRS uncat. (8, 25-40), Feni River at Ramgarh, 
Chittagong Hill Tracts, 3 Feb. 1978; CFRS uncat. (3, 46.4— 
62.5), Tangam River at Kestapur, Dinajpur Dist., no date; 
AMNH 19648 (2) Sevoke River, Darjeeling. 


ACKNOWLEDGMENTS 


My sincere thanks and appreciation go to Karl 
F. Lagler, The University of Michigan, and John 
R. Snell, A. Q. Chowdhury, and Kermit E. Sneed, 
Irrigation Fishery Development Project, Bang- 
ladesh, for help in obtaining and shipping the 
specimens to The University of Michigan. I owe 
a great deal to my close friend and associate Ataur 
Rahman and many others in the Directorate of 
Fisheries, Government of Bangladesh, who 
greatly facilitated the field work. William N. 
Eschmeyer (CAS) and Donn E. Rosen (AMNH) 
loaned specimens. Reeve M. Bailey, Gerald R. 
Smith, Barry Chernoff, and Michael L. Smith of 
The University of Michigan offered helpful sug- 


76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6 


gestions pertaining to many aspects of manu- 
script preparation. Tyson Roberts kindly donated 
several specimens of P. balitora from Nepal to 
the UMMZ. Mark Orsen and Karna Steelquist 
of The University of Michigan gave much- 
needed advice on illustrations. This work was 
supported in part by the UCLA Department of 
Biology Fisheries Program. 


LITERATURE CITED 


Cuen, Y. 1980. Acta Hydrobiologica Sinica (not seen). 

Davip, A. 1953. Onsome new records of fish from the Damo- 
dar and Mahanadi river systems. J. Zool. Soc. India 5(2): 
243-254. 

Day, F. E. 1878. The fishes of India, etc. 2 vols. London. 

Gray, J.S. 1834. Illustrations of Indian zoology of new and 
hitherto unfigured Indian animals from the collection of 
General Hardwicke. 2 vols. London. 

Gupcer, E. W. 1924. The sources of the material for Ham- 
ilton-Buchanan’s fishes of the Ganges, the fate of his collec- 
tions, drawings and notes, and the use made of his data. J. 
Proc. Asiatic Soc. Bengal 19(4):121-136. 


HamILTon (formerly BUCHANAN), F. 1822. An account of the 
fishes found in the river Ganges and its branches. Edinburgh. 
405 p., 39 pls. 

Hora, S. L. 1925. Notes on fishes in the Indian Museum. 
XII—The systematic position of the genus Psilorhynchus 
McClelland. Rec. Indian Mus. (Calcutta) 27:457-460. 

. 1929. An aid to the study of Hamilton Buchanan’s 

“Gangetic Fishes.”” Mem. Indian Mus. (Calcutta) 9(4):169- 

192. 


, AND K. S. Misra. 1941. Fishes collected by the 
Vernay-Hopwood upper Chindwin Expedition, 1935. J. 
Bombay Nat. Hist. Soc. 42(3):478-482. 

, AND D. D. Mukersi. 1935. Fishes of the Naga Hills, 
Assam. Rec. Indian Mus. (Calcutta) 37:381-404. 

MCCLELLAND, J. 1839. Indian Cyprinidae. Asiat. Res. 19: 
217-471. 

Menon, A. G. K. 1974. A check-list of fishes of the Hima- 
layas and the Indo-Gangetic plains. Inland Fisheries Society 
of India, Spec. Publ. 1. 

, AND A. K. Datta. 1961. Zoological results of the 
Indian Cho-Oyu Expedition (1958) in Nepal. 7— Pisces. Psi- 
lorhynchus pseudecheneis a new cyprinid fish from Nepal. 
Rec. Indian Mus. (Calcutta) 59:253-255. 

SHaw, G. E., AND E. O. SHEBBEARE. 1937. The fishes of 
Northern Bengal. J. Asiat. Soc. Bengal, Science 3:1-137. 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


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PROCEEDINGS 
OF THE 


a 


q 


<. 


Mariné Biolegical Laboratory 


LTRRARY 
JuL 15 1983 


CA LIFORNIA ACADEMY OF SCIENGESods Hole, 98S. 


Vol. 43, No. 7, pp. 77-85, 1 fig., 5 tables. 


July 6,183 


SIZE AND DISTRIBUTION OF THE CALIFORNIA 
SEA LION POPULATION IN MEXICO 


By 


Burney J. Le Boeuf,' David Aurioles,? Richard Condit,' 
Claudio Fox,’ Robert Gisiner,’ Rigoberto Romero,* 


and Francisco Sinsel’ 


Asstract: California sea lions were censused during the 1979 and 1981 breeding seasons on rookeries and 
hauling grounds in the Gulf of California and the Pacific coast of Mexico. Correcting for underestimate bias 
and uncensused islands yielded estimates of approximately 20,000 sea lions in the Gulf with an annual 
production of 7,000 pups, and 63,000 sea lions on the Pacific coast with an annual production of 29,000 pups. 
Counts on seven major rookeries in the Gulf in 1979 were 35% higher than in 1966. Sixteen percent of the 
California sea lion population in the United States and Mexico (estimated at 145,000 animals) resides in the 


Gulf and 46% inhabits the Pacific coast of Mexico. 


INTRODUCTION 


The California sea lion, Zalophus californi- 
anus, along with numerous species of birds and 
whales, is a top trophic-level predator in the pro- 
ductive waters of the Gulf of California. In the 
southern part of the Gulf, it feeds predominantly 
on a variety of fishes and occasionally on squid 
(Aurioles, Fox, and Sinsel 1981). The full impact 
of this pinniped on Gulf of California fisheries 
is unknown because few systematic censuses of 
these animals have been conducted. 

Long before California sea lions in the Gulf of 
California were counted, they were exploited. 
Early inhabitants of Baja California, Sonora, and 
Sinaloa killed California sea lions for their meat. 


' Department of Biology and Center for Coastal Marine 
Studies, University of California, Santa Cruz, California 95064. 

> Centro de Investigaciones, Biologicas de Baja California, 
La Paz, Baja California, Mexico. 

3571 West B Street, Dixon, California 95620. 


[77] 


Between 1860 and 1870, whalers killed thou- 
sands of sea lions for their oil and skins. Later, 
the animals were hunted for their “trimmings” — 
vibrissae and genitalia which were sold in the 
Orient as aphrodisiacs. Local slaughter for oil 
and meat continued until the early 1960’s on 
some islands in the Gulf, e.g., Puerto Refugio on 
Angel de la Guarda (Lluch 1969), and poaching 
is still going on today. 

The first systematic census of the sea lions in 
the Gulf was conducted by Lluch (1969). He esti- 
mated 6,027 total animals on eight islands from 
counts made in 1963, 1964, and 1966 at the peak 
of the breeding season. Orr, Schonewald, and 
Kenyon (1970) censused several islands between 
1960 and 1968, but with the exception of Los 
Islotes, no rookery was censused during the 
breeding season. Mate (1977) estimated 9,428 
sea lions in the Gulf of California from aerial 
censuses taken in June 1975, but his data are 
difficult to compare with other censuses because 


78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7 


i N 
x= Is. San \ 
PS Benito YS) 
\ AiCedros 
4 


Natividad® 
\\ RE 


\ 
°s 
Ignacio 

, Farall6n 


Ficure 1. The route of R/V Ellen B. Scripps on expedi- 
tions in 1979 and 1981 from San Diego, California, to Cali- 
fornia sea lion rookeries along the Pacific coast of Baja Cali- 
fornia, Mexico, and in the Gulf of California. The itineraries 
were similar in both years except for deviations taken in 1979 
which are noted by solid lines. The numbers refer to rook- 
eries not censused (1-6) and islands supporting nonbreeding 
aggregations (7-12). The code is: 1 = Los Coronados; 2 = Isla 
San Ger6énimo; 3 = Punta Rosarito; 4 = Isla Magdalena; 5 = 
Isla Lobos; 6 = Farall6n at Mazatlan; 7 = Isla Salsipuedes; 8 = 
Cabo Haro; 9 = Roca Vela; 10 = Isla Patos; 11 = Cabo Lobos; 
12 = Punta Lobos. 


they are categorized by latitude rather than by 
island. 

The main purpose of this report is to present 
recent data on the number and distribution of 
California sea lions in the Gulf of California dur- 
ing the breeding season and to compare these 
data to earlier counts and to counts obtained on 
islands on the west coast of Baja California, Mex- 
ico. In addition, we report weights and mea- 
surements of sea lion pups obtained from Gulf 
and Pacific rookeries. 


METHODS 


CENSUSES.— The data in this report were col- 
lected on two expeditions aboard the Scripps 
Institution of Oceanography research vessel Ellen 
B. Scripps. Inclusive dates were 21 June to 21 
July 1979 and 1 to 30 June 1981. The track 
records are shown in Figure 1. 


Both expeditions were timed to coincide with 
the breeding season of California sea lions and 
to facilitate counting sea lions in the Gulf. At 
Los Islotes, the peak number of breeding females 
occurs at the end of Jine, and the maximum 
number of pups are observed on about 10 July 
(Aurioles, Romero, and Fox 1978). We were aided 
in locating all suspected rookeries in the Gulf by 
previous investigations and by information 
obtained from local fisherman. The only rook- 
eries not censused in the Gulf were Isla Lobos 
and the farall6n at Mazatlan. We did not census 
the following rookeries on the outer Pacific coast 
of the peninsula: Isla Natividad, Los Coronados, 
Isla San Geronimo,* and Punta Rosarito. Esti- 
mates of the sea lion population on these Gulf 
and Pacific rookeries are available from other 
sources (e.g., Mate 1977). 

Censuses were conducted from a dinghy or 
launch, from the ship, and on foot. Most counts 
were made from a 4-m Avon or Zodiac inflatable 
dinghy powered by a 15- or 25-hp outboard 
motor. One person operated the boat and one or 
two people censused. Small islands were circum- 
navigated; on large islands, a dinghy was launched 
after sea lion aggregations were sighted from the 
ship. In ideal conditions, the boat was run slowly 
within 10 m of the shoreline. This did not alarm 
the sea lions and accurate counts were obtained. 
When waves or wind were strong, or extensive 
kelp beds surrounded an island, the boat was 
operated 30 to 70 m offshore. In bad weather, 
binoculars were used from the research vessel 
from 0.5 to | km offshore. This was necessary 
at Isla Santa Margarita and three small non- 
breeding aggregations on Isla Magdalena, Salsi- 
puedes, and Cabo Haro in 1981. 

We censused sea lions in six categories: adult 
males, subadult males, females, juveniles, pups, 
and miscellaneous. Adult males are the largest 
animals with large dorsal head crests and thick 
necks. Subadult males are smaller, have thinner 
necks, and little crest development. For most 
analyses, these two age categories are combined. 
Females and juveniles are distinguished from 
males by their smaller size, thinner neck, and 
light brown color compared to dark brown in 
males. Juveniles are the smallest class in this 
group and include one- to two-year-olds, and 
possibly some three-year-olds. The miscella- 


4 Also spelled Isla San Jer6nimo. 


CALIFORNIA SEA LION POPULATION IN MEXICO 


neous category encompasses any animal except 
pups that could not be readily categorized because 
of high animal density or poor censusing con- 
ditions. Pups are the unmistakable, but easily 
concealed, young of the year. 

VALIDITY AND RELIABILITY ESTIMATES. — 
Validity tests were performed using ground counts 
and boat counts of the same rookery. After a 
census was conducted from a dinghy, one person 
went ashore, counted adults from a hiding place, 
and then frightened the adults into the water and 
quickly counted the pups. This was done at six 
places in 1981. In addition, at two sites we com- 
pared counts from the ship with counts from a 
dinghy. Estimates of reliability were obtained by 
having two censusers make simultaneous counts 
at all rookeries in 1979 and at five rookeries in 
1981. For these areas, the mean of the two cen- 
suses conducted is presented. 

WEIGHTS AND MEASUREMENTS.—A crew of 
three to six people weighed and measured 111 
pups on six rookeries, one in the Pacific and five 
in the Gulf. Landing near a rookery caused adults 
to flee into the water leaving their pups behind. 
The procedure consisted of capturing a one- to 
three-week-old pup by the hindflippers and 
restraining it on a flat rock. Curvilinear length 
(tip of nose to tip of tail) was measured along the 
contour of the dorsal midline. The pup was then 
placed in a light leather dog harness and weighed 
with a hand-held spring scale (25-kg capacity + 
0.5 kg). After extracting a blood sample, the pup 
was tagged with an “‘All-Flex” cattle ear tag in 
the trailing edge of a foreflipper, its sex was deter- 
mined, and it was carried back to where it was 
captured. Mothers retrieved their pups shortly 
after pups were released or soon after the launch 
and crew departed from the rookery. 


RESULTS 


WEATHER. — Weather conditions in 1979 were 
ideal for censusing in the Gulf; seas were calm; 
winds of 0-5 knots were variable in direction. 
Conditions were fair for censusing Pacific coast 
islands except at Isla Santa Margarita where high 
surf prevailed. Wind and choppy water made 
censusing in the Gulf difficult in 1981, especially 
on the windward sides of Islas San Esteban, San 
Pedro Martir, Granito, and San Ignacio Farallon; 
on the Pacific side, heavy surf made landings or 
censusing by dinghy impossible at Islas Mag- 


79 


dalena and Santa Margarita and on the west side 
of Isla Cedros. 

GULF OF CALIFORNIA.— More than 15,000 sea 
lions were counted in the Gulf in 1979 and more 
than 14,000 in 1981 (Table 1). Major rookeries 
in the Gulf were found on islands in the center 
or midriff area: San Esteban and Angel de la 
Guarda (Los Machos, Granito, and Los Can- 
tiles). In 1979 and 1981, 76 and 71% of the sea 
lions counted were observed in the area from the 
north end of Angel de la Guarda (29°35’'N) south 
to San Pedro Nolasco (27°58'N). Most of the sea 
lions in the north end of the Gulf were located 
on Isla San Jorge. Relatively few sea lions breed 
in the southern end of the Gulf, and these were 
concentrated on Los Islotes and San Ignacio Far- 
allon. 

No pups were observed in the small groups of 
sea lions present on Isla Salsipuedes, Cabo Haro, 
Roca Vela, Isla Patos, Cabo Lobos, and Punta 
Lobos. No sea lions were observed during careful 
surveys of Isla Coronado, Tortuga, Raza, Mejia, 
Cabo Tepopa, Los Frailes, Cabo San Lucas, and 
the eastern half of Islas San Lorenzo, Las Ani- 
mas, and Espiritu Santo. A few males or juveniles 
were Observed on Islas Carmen, San Lorenzo, 
and Partida. 

Eleven Gulf rookeries produced a minimum 
of 3,422 pups in 1979, 2,277 in 1981 (Table 1). 
The largest category of the animals counted on 
rookeries were female (48% in 1979 and 60% in 
1981). The mean operational sex ratio (adult 
males:adult females) for rookeries was 1:8.30 
(N = 9, sp = 5.61) in 1979 and 1:9.41 (N = 10, 
sD = 3.83) in 1981. The overall operational sex 
ratio for rookeries in these years were 1:5.84 and 
1:8.94, respectively. 

Sea lions in the Gulf occupied cobblestone 
coves or the boulder-strewn shoreline. Few 
breeding groups were seen on sand or gravel 
beaches, which tended to be occupied by non- 
breeding males. Most territorial males were 
observed “patrolling” the water in front of the 
females. Females and pups occupied a narrow 
zone near the water’s edge and rarely were 
observed 10 m or or more inland. In some loca- 
tions, rafts of females floated in the shallows and 
the females occasionally vocalized to their pups 
onshore. Even on islands supporting numerous 
sea lions, the population was broken up into small 
groups, and a small number of animals took up 
a large part of the shoreline. 


80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7 
TABLE 1. CENSUSES OF CALIFORNIA SEA LIONS IN THE GULF OF CALIFORNIA, 29 JUNE TO 8 JuLy 1979 (top figures) AND 8-21 
JUNE 1981 (bottom figures, in italics). ““—’’ denotes that no census was taken. Abbreviations for census methods: G = ground, 


L = launch, S = ship. 


Census 

Islands methods Males Females Pups Juv Misc. Totals 

Los Islotes G,L 18 88 28 4 138 
G 11 26 fl 0 44 

San Ignacio Farall6n — — — - _ — — 
G, L 10 133 64 36 80 323 

Cabo Haro L 1 32 16 49 
S 2S BS 

San Pedro Nolasco L 76 382 232 133 23 846 
L 82 696 125 201 ae, 1,126 

San Pedro Martir L 150 903 321 167 93 1,634 
G, L 134 1,061 252 120 125 1,692 

San Esteban G,L 504 2,044 820 325 268 3,961 
G,L, S$ 295 BSP 705 239 102 3,593 

Salsipuedes L 8 34 56 98 
S 19 NS) 34 

Roca Blanca L 8 121 40 38 207 
Patos L 454 19 473 

Angel de la Guarda 

Los Machos L 35 709 168 404 1,316 
SS: 84 762 226 247 70 1,389 

Granito L 87 436 337 242 1,102 
G, L 73 658 161 79 47 1,018 

Los Cantiles L 163 969 446 219 13 1,810 

L 9] 688 270 102 229 1,380 

Roca Vela L 3 50 30 83 
Cabo Lobos L 50 48 72 170 
San Jorge L 167 1,398 1,030 632 26 3,253 
L 165 2,034 457 667 21 3,344 

Rocas Consag — — — — —- -— — 
L 18 295 10 98 421 

Totals 1979 1,724 7,214 3,422 2,357 423 15,140 
Totals 1981 988 8,639 Dr DYY 1,789 696 14,389 


Approximately 10-20% of the females 
observed were still nursing yearlings, and some 
small groups were made up almost entirely of 
such pairs. 

PaciFic CoAst.—Approximately 23,000 sea 
lions were counted on Pacific coast rookeries in 
1979, a greater number than in 1981 (Table 2). 
However, some islands censused in 1979 were 
not censused in 1981, and vice versa. A better 
estimate of the total number of animals associ- 
ated with the rookeries shown in Table 2 is 
27,895—obtained by adding the 1981 census 
count for Islas Cedros (4,730) and Magdalena 
(47) to the 1979 total census. 

At least 6,529 pups were produced on the 


islands censused in 1979. If the number of pups 
counted on Isla Cedros in 1981 (2,138) is added 
to that, annual pup production was at least 8,667 
pups. 

As in the Gulf, females were in the majority 
(53% of animals counted in 1979 and 55% in 
1981). The mean operational sex ratio of rook- 
eries in 1979 was 1:6.87 (N = 6, sp = 5.83), the 
overall sex ratio was 1:3.58. Less than 2% of the 
females were observed nursing yearlings. 

Large aggregations of sea lions were observed 
on each rookery visited with the exception of Isla 
de Guadalupe. The females were hauled out in 
clumps well above the surf line, usually on sandy 
beaches. Territorial males stationed themselves 


CALIFORNIA SEA LION POPULATION IN MEXICO 


81 


TABLE 2. CENSUSES OF CALIFORNIA SEA LIONS ON THE PactiFic Coast OF BAJA CALIFORNIA, 13-20 JuLy 1979 (top figures) 
AND 3-5 JUNE 1981 (bottom figures, in italics). Numbers in parentheses are calculated from total number of animals censused; 
“> denotes that no census was taken. Abbreviations for census methods: G = ground, L = launch, S = ship. 


Census 
Islands method Males Females 
San Martin G 25 9 
Guadalupe L,S 5 85 
San Benito 
Oeste Gal 355 502 
Gale 21 HIS) 
Centro G,L 817 3,718 
G 79 1,985 
Este L 183 1,070 
IL (97) (441) 
Cedros - — — 
Gale 270 1,880 
Asuncion G,L 1,793 4,183 
G (105) (2,780) 
Magdalena — — _ 
S 
Santa Margarita ILS (G 265 2,676 
S (108) (1,086) 
Punta Lobos L 6 35 
Totals 1979 3,449 12,278 
Totals 1981 680 8,287 


on land with the females. Nonbreeding males 
were observed considerable distances inland on 
nearby rocks or on unoccupied beaches. In smaller 
groups that were sometimes seen in coves or on 
rocky ledges, territorial males were either on land 
or in the water near the females. 

Eighteen sea lions were observed at sea in the 
Pacific and in the Gulf in 1981. They were sol- 
itary and most were within a few kilometers of 
a rookery. Three exceptions were one sea lion 
seen near Cabo Pulmo, one east of Cabo San 
Lucas, and one near Isla Coronados. 

Census METHOD BiAs.—Comparison counts 
using different census methods are shown in Table 
3. Ground counts of pups exceeded those taken 
from a launch, with the underestimate from a 
launch being greatest when the animals were dis- 
tributed in rocky areas. Launch and ground counts 
of adults were similar; the slightly higher counts 
from a launch were partly because launch counts 
always preceded ground counts. The latter were 
conducted in the morning when the temperature 
was rising and some adults were entering the 
water to cool off. Ship counts of adults were lower 


Pups Juv Misc. Totals 
40 74 

7 8 100 

283 13 1,153 

1 150 287 
2,560 88 7,183 
1,185 38 3,287 
900 59 DAN 
(304) (10) 852 
2,138 282 160 4,730 
1,582 506 8,064 
(406) (608) 3,899 
47 47 

1,202 114 4,257 
(652) (46) 1,892 
14 55) 

6,529 842 23,098 
4,686 984 357 14,994 


than those taken from the launch, and pups were 
most difficult to see from the large vessel. 

Two measures of inter-observer agreement 
were obtained. In 1979, independent counts of 
two censuses on 13 rookeries were correlated. 
Correlation coefficients were high and positive 
for total animals (0.976), females (0.935), and 
pups (0.926). Correlation coefficients were lower 
for juveniles (0.777) and males (0.712), animals 
which are more difficult to categorize, 1.e., sub- 
adult males or juveniles can be confused with 
adult females. In 1981, two independent cen- 
suses were conducted on parts of five rookeries 
in the Gulf. The results, shown in Table 4, are 
similar to those obtained in 1979 in that agree- 
ment is usually high for total animals and females 
and relatively low for juveniles. Independent 
counts of pups in 1981 were more discrepant 
than in 1979. 

Pups WEIGHTS AND MEASUREMENTS. — The 
mean weight and curvilinear length of newborn 
pups is shown in Table 5. On each rookery, the 
mean weight and mean length of males is greater 
than that of females. For all rookeries combined, 


82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7 


TABLE 3. 
ground, L = launch, S = ship. 


A COMPARISON OF CENSUS METHODS ON SHORT SECTIONS OF RookertEs. Abbreviations for census methods: G = 


Census 

Island Weather and terrain method Adults Pups 
Cedros Calm sea; animals packed tightly on sandy beach with L — 55 
some in arroyo behind beach. G — 80 
San Pedro Martir Calm sea, overcast sky; animals on pebbly beach. L — 25 
G = 42 
Granito (A.G.) Calm sea on protected side; animals distributed on L 269 45 
small cobblestones & large boulders backed by ver- G 231 117 

tical cliffs. 
San Ignacio Calm sea; animals on rocky shoreline containing a L 65 4 
Farallon large open cave, flat shelves & large boulders. G 58 52 
San Esteban Slight chop; animals on long sandy spit, in rock stub- S 264 0 
ble at base of vertical cliffs & in shallow sea caves. I, 388 2 
Los Machos (A.G.) Calm sea; animals on sandy beaches, rock tables & S 347 0) 
among boulders at base of cliffs. L 488 179 


males were significantly heavier (t = 5.42, df= 
109, P = < 0.05) and longer (t = 4.88, df= 109, 
P = <0.05) than females. The mean weights of 
pups of both sexes from Cedros Island, the only 
Pacific rookery represented, are lower than that 
of pups from any Gulf rookery, but these differ- 
ences are not statistically significant. 


DISCUSSION 


We counted 15,140 California sea lions in the 
Gulf of California in 1979. This number is 61% 
higher than an aerial census in July 1975 by Mate 
(1977) and 151% higher than a launch and ground 
census in 1963-1966 by Lluch (1969). Is our 
higher count due to increased censusing effort, 
the census methods employed, or to a genuine 
increase in the population? Each explanation has 
some validity. 

Our census was more complete than Lluch’s. 
Fifty-four percent of the animals we counted in 
1979 were on rookeries and resting places Lluch 
did not visit; Los Islotes, San Esteban, Roca 
Blanca, Angel de la Guarda (Los Machos and 
Los Cantiles), Cabo Haro, Islas Salsipuedes and 
Patos, and Cabo Lobos. If sea lions were found 
on these islands in the mid-1960’s in the same 
relative proportions as in 1979, the total number 
of animals would have approximated 13,000, 
16% fewer than we counted in 1979. 

Terrain and climatic conditions in the Gulf are 
such that an aerial census like that conducted by 
Mate (1977) from a small airplane underesti- 


mates the number of animals present more than 


counts from a launch or on foot. Mate explains 


that his count of 9,428 was low because of the 
difficulty of counting and photographing animals 
from the air over the dark, rocky terrain, with 
many animals in the water by mid-morning. An 
indication of the different results of these two 
censusing methods is that Mate counted only 122 
pups in the entire Gulf compared to our 3,422. 

Because of discrepancies in method and loca- 
tion of censusing, entire censuses are not very 
useful for determining population change. How- 
ever, comparisons of individual rookeries indi- 
cate that the population in Mexico has increased. 
For seven rookeries, our counts can be compared 
with Lluch’s (1969). Both the dates and method 
of censusing were similar. He counted 5,977 ani- 
mals and we counted 7,662 and 8,091 animals 
in our two censuses. These represent increases 
of 28 and 35% over a period of 13 to 16 years. 
Our counts were higher on San Jorge (77%), San 
Pedro Martir (56%), and Rocas Consag (181%), 
and lower on Granito (40%), Roca Vela (81%), 
and San Ignacio Farallon (alias Topolobampo) 
(24%). Counts on San Pedro Nolasco were sim- 
ilar in 1966 and 1979 but increased by 32% in 
1981. Finally, it is not clear if the population 
decreased from 1979 to 1981, because the two 
censuses were conducted at slightly different times 
during the breeding season. The 1981 census was 
made prior to peak season, accounting for the 
lower count. 

There are numerous difficulties in simply 
counting large groups of sea lions, leading to 
underestimates of animals present. Counts vary 
with the census method used, weather condi- 


CALIFORNIA SEA LION POPULATION IN MEXICO 


83 


TABLE 4. COMPARISON OF INDEPENDENT CENSUSES TAKEN AT SEA LION ROOKERIES IN THE GULF OF CALIFORNIA IN JUNE 1981. 


Abbreviations: G = ground, L = launch, RC = R. Condit, FS = 


F. Sinsel, CF = C. Fox. 


Census Weather, light, 
Rookery method terrain Censuser Males Females Pups Juv Totals 

Los Islotes G Clear sky, fading light; RC 11 26 7 0 44 
rough terrain, boulder FS 16 14 6 6 42 
& cobblestones. 

San Pedro Martir IL, Windy, overcast; choppy RC 69 925 72 124 1,190 
to heavy seas; 2 sandy FS 134 1,186 252 120 1,692 
beaches, cobble coves, 
rock tables, large boul- 
ders backed by steep 
cliffs. 

Los Machos (A.G.) L Sunny, flat sea; most RC 84 832 226 247 1,389 

(partial) females & pups on ES 67 1,069 293 109 12535 
sandy beaches, juve- 
niles & subadult males 
on rock tables & in 
boulder fields. 

Granito (A.G.) IL, s side calm; N side very RC 60 621 163 72 916 
choppy, poor census- ES/CE 87 789 159 87 122: 
ing conditions; 1 sandy 
beach on each side 
bordered by irregular 
rocky coastline backed 
by steep cliffs. 

San Esteban IU, Calm sea nN & w sides; RC 234 1,865 369 500 2,968 
steep cliffs with caves FS PAV 1,824 545 160 2,741 
& rocky shelves; long 
sandy or cobbly 
beaches. 

Totals € 458 4,269 837 943 6,507 

FS/CF 516 4,882 1E255 482 fale 


tions, terrain, the experience and reliability of 
censusers, and the size, age, and sex composition 
of groups being counted. Moreover, the number 
of sea lions on land varies with time of year and 
the time of day. A special difficulty for estimating 
breeding females is that some will always be at 
sea feeding during the breeding season. Asa result, 
counts provide only minimum estimates of the 
number of animals present. Empirically based 
correction factors must be employed to estimate 
actual population numbers. 

We reason that the actual number of sea lions 
counted in the Gulf of California underestimates 
the number of animals and that a more valid 
estimate is 20,144 animals. This estimate is 
derived by applying three correction factors to 
the 1979 total count of 15,140 animals (the 1979 
census is selected over the 1981 census because 
it is closer to the peak of the breeding season). 
This count is augmented by: (a) 744 animals, 


representing known rookeries not censused in 
1979; San Ignacio Farallon (323) and Rocas Con- 
sag (421); (b) 3,496 pups, assuming that pups 
counted reflect only 50% of the pups present; (c) 
764 females, assuming that 10% of the females 
are at sea feeding (see Bonnell et al. 1978). We 
believe that 20,144 total animals in the Gulf and 
an annual production of approximately 7,000 
pups are reasonable, conservative estimates. 
Using the same assumptions, we can estimate 
the number of sea lions on the Pacific coast of 
Mexico. Before doing this, it should be noted 
that previous censuses from launches or on foot 
are of limited use for estimating population size 
because they are only partial counts of the area 
or they were conducted outside the breeding sea- 
son, e.g., Bartholomew and Hubbs (1952), Rice, 
Kenyon, and Lluch (1965), Orr, Schonewald, and 
Kenyon (1970), Brownell, DeLong, and Schrei- 
ber (1974). Aerial censuses of sea lions on the 


84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7 


TABLE 5. WEIGHTS AND MEASUREMENTS OF CALIFORNIA SEA LION Pups OBTAINED 11-28 JUNE 1981. For each island listed, 
the mean weight (in kilograms) or mean length (in centimeters) is shown plus or minus one standard deviation. N is in parentheses. 


Weight Length 
Rookeries Males Females Males Females 

San Pedro Martir 8.75 + 1.89 7.50 + 1.96 76.42 + 4.58 71.43 + 8.94 
(6) (4) (6) (4) 

San Esteban 10.17 + 1.54 8.00 + 1.06 74.93 + 3.00 74.08 + 5.03 
(6) (9) (6) (9) 

Granito (A.G.) 9.19 + 1.04 7.75 + 0.97 fol += 2162) 71.48 + 3.14 
(16) (13) (16) (13) 

San Jorge 8.86 + 0.92 7.71 + 1.14 75.01 + 2.30 72.55 + 3.16 
(9) (12) (9) (12) 

San Ignacio Farallon 8.80 + 1.10 8.20 + 1.30 76.20 + 0.92 70.34 + 3.31 
(5) (5) (5) (5) 

Cedros 8.54 + 1.45 7.29 + 1.93 76.79 + 3.37 72.82 + 3.91 
(12) (14) (12) (14) 

Totals 9.01 + 1.32 7.60 + 1.45 75.71 + 2.93 72.34 + 4.18 
(54) (57) (54) (57) 


Pacific coast of Mexico by Mate (1977) (see Bon- 
nell et al. 1978) yielded a direct count of 45,872 
animals. We estimate that the total number of 
sea lions on the Pacific coast of Mexico is approx- 
imately 63,020 animals, a number which includes 
the annual production of 29,000 pups. We start 
with the direct count of 23,098 animals obtained 
in 1979. We increase this number by: (a) 4,777 
representing two rookeries we counted in 1981 
but not in 1979 (Islas Cedros and Magdalena, 
see Table 2); (b) 12,682 representing the follow- 
ing rookeries we did not census but which were 
censused by Mate (1977) (see also Bonnell et al., 
1978): Los Coronados (297), San Gerdnimo 
(1,113), Punta Rosarito (2,722), Natividad 
(5,785), and Punta Tosca (2,765); (c) 8,667 pups, 
assuming that 50% of the pups counted from 
launches were missed; (d) 11,445 pups, to com- 
pensate for the failure to count pups from aerial 
censuses; and (e) 2,351 females, assuming that 
10% of the females were at sea. Augmentations 
in categories b, d, and e are based on Mate’s aerial 
census in 1975 (Mate 1977). Since he did not 
categorize animals by age or sex, we added pup 
and female counts to his numbers using ratios 
derived from our censuses. 

From these estimates and censuses conducted 
in the United States, we can estimate the size of 
the population. Bonnell et al. (1978) estimated 
50,000 animals in southern California waters 
from aerial censuses and 4,000 in the northern 
part of the nonbreeding range. Thus, the total 


number of California sea lions in the United States 
and Mexico is approximately 145,000 animals. 
Of this total, 16% are in the Gulf of California, 
46% are on the Pacific coast of Mexico, 35% are 
in southern California, and 3% range as far north 
as Vancouver Island, British Columbia (Han- 
cock 1970; Bigg 1973). Our estimate of the United 
States and Mexico population of this species 
exceeds the figure of 125,000 estimated by Bon- 
nell et al. (1978). Both estimates do not include 
the geographically separated subspecific popu- 
lations on the Galapagos Islands and in Japan. 
The distribution of breeding animals in the 
Gulf differs from that most commonly observed 
on Pacific coast rookeries. In the Gulf, male ter- 
ritories are predominantly aquatic, fronting on 
small groups of females and pups distributed in 
a narrow zone along the water’s edge on cobble- 
stone beaches, among boulders, or on rocky 
ledges. On Pacific coast rookeries in Mexico and 
California, females are most often hauled out in 
large groups well above the surf line on sandy 
beaches or on flat rock outcroppings sloping into 
the sea (Peterson and Bartholomew 1967; Odell 
1975; Bonnell et al. 1978); male territories are 
semi-aquatic or usually terrestrial. Higher tem- 
peratures in the Gulf may account in part for 
these differences in behavior and distribution. 


ACKNOWLEDGMENTS 


We thank Drs. Felix Cordoba Alva and Carlos 
de Alba Perez for help in obtaining permits and 


CALIFORNIA SEA LION POPULATION IN MEXICO 


making logistical arrangements in Mexico; George 
Shor, Robert Haines, and others at Scripps Insti- 
tution of Oceanography for helping to make the 
R/V Ellen B. Scripps available to us; Captains 
Hansen and Whitman and their crews for safe 
and efficient conduct; and Dr. Leo Ortiz, Dr. 
James Estes, Dr. Martha Field, Dr. Robert Brow- 
nell, Jr., Steve Davenport, Edward Keith, Kathy 
Panken, John Peterson, Joanne Reiter, Marianne 
Riedman, and Keith Skaug for field assistance. 
These expeditions were supported in part by 
National Science Foundation grant DEB 77- 
17063 AO! to B. Le Boeuf. 


LITERATURE CITED 


AuRIoLes, D., C. Fox, AND F. SINsEL. 1981. Species of fishes 
identified with the otoliths found in scats of sea lions at “Los 
Islotes” Island, B.C.S., Gulf of California, Mexico. Abstract, 
4th Biennial Conference on the Biology of Marine Mammals, 
Dec. 14-18, 1981, San Francisco, California. 

, R. ROMERO, AND C. Fox. 1978. Censos poblaciones 
de lobos marinos (Zalophus californianus) por edades y sexos 
en la bahia de la Paz y sus alrededores. Pages 139-154 in 
Informe General de Labores, 1978. Centro de Investiga- 
ciones Biologicas de Baja California, A.C. 

BARTHOLOMEW, G. A., AND C. L. Husss. 1952. Winter pop- 
ulations of pinnipeds about Guadalupe, San Benito, and 
Cedros Islands, Baja California. J. Mammal. 33:160-171. 

Bicc, M.A. 1973. Census of California sea lions on southern 
Vancouver Island, British Columbia. J. Mammal. 54:285- 
287. 

BonneELL, M. L., B. J. Le Boeur, M. O. Pierson, D. H. DETTMAN, 


85 


AND G. D. Farrens. 1978. Pinnipeds of the southern Cal- 
ifornia Bight. Vol. III, Part 1, 535 p. in K. S. Norris, B. J. 
Le Boeuf, and G. L. Hunt, Jr., eds., Marine Mammal and 
Seabird Surveys of the Southern California Bight Area, 1975— 
1978. Bureau of Land Management, Dept. of Interior Con- 
tract AAS50-CT7-37, Government Printing Office, Wash., 
DIC 

BROwneELL, R. L., Jk., R. L. DE LoNG, AND R. W. SCHREIBER. 
1974. Pinniped populations at Islas de Guadalupe, San Be- 
nito, Cedros and Natividad, Baja California, in 1968. J. 
Mammal. 55:469-472. 

Hancock, D. 1970. California sea lion as a regular visitant 
off the British Columbia coast. J. Mammal. 51:614. 

Liucn, B. D. 1969. El lobo marino de California (Zalophus 
californianus californianus Lesson, 1828) (Allen, 1880), en 
dos mamiferos marinos de Baja California. Inst. Mex. Rec. 
Nat. Renovables., 3-69. 

Mate, B. 1977. Aerial censusing of pinnipeds in the Eastern 
Pacific for assessment of population numbers, migratory dis- 
tributions, rookery stability, breeding effort, and recruit- 
ment. Report to Marine Mammal Commission Contract 
MMS5AC001, 28 Feb. 1977. 68 p. 

Ope.t, D. K. 1975. Breeding biology of the California sea 
lion, Zalophus californianus. Rapp. P.-V. Reun. Cons. Int. 
Explor. Mer 169:374-378. 

Orr, R. T., J. SCHONEWALD, AND K. W. Kenyon. 1970. The 
California sea lion: skull growth and a comparison of two 
populations. Proc. Calif. Acad. Sci., ser. 4, 37:38 1-394. 

Peterson, R. S., AND G. A. BARTHOLOMEW. 1967. The nat- 
ural history and behavior of the California sea lion. Spec. 
Pub. No. 1, Amer. Soc. Mammal. 79 p. 

Rice, D. W., K. W. Kenyon, AND D. Ltucu. 1965. Pinniped 
populations at Islas Guadalupe, San Benito, and Cedros, 
Baja California, in 1965. Trans. San Diego Soc. Nat. Hist. 
14:73-84. 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


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CALIFORNIA ACADEMY OF SCIE 


Vol. 43, No. 8, pp. 87-110, 15 figs. 


Woods Rbdle, Mass. 


July 6, 1983 


MEGAMOUTH-—A NEW SPECIES, GENUS, AND FAMILY OF 
LAMNOID SHARK (MEGACHASMA PELAGIOS, FAMILY 
MEGACHASMIDAE) FROM THE HAWAIIAN ISLANDS 


By 


Leighton R. Taylor 
Waikiki Aquarium, 2777 Kalakaua Ayve., Honolulu, Hawaii 96815 


L. J. V. Compagno 


Tiburon Center for Environmental Studies, San Francisco State University, 
P.O. Box 855, Tiburon, California 94920 


and 
Paul J. Struhsaker 
1050 Koloa St., Honolulu, Hawaii 96816 


Asstract: Megachasma pelagios, a new genus and species of lamnoid shark assigned to the new family 
Megachasmidae, is described and defined from a single adult male, 4.46 m total length. The holotype and 
only known specimen was collected approximately 42 km NE of Oahu, Hawaii. Structure and habitus distinctly 
differ from other lamnoid sharks, particularly in head and tooth morphology and in mesopelagic filter feeding. 


INTRODUCTION 


On 15 November 1976, the research vessel 
AFB-14 of the Naval Undersea Center (now the 
Naval Ocean Systems Center), Kaneohe, Hawaii, 
was conducting oceanographic research in waters 
about 42 km northeast of Kahuku Point, Oahu, 
at about 21°51’N and 157°46’W. From 1015 to 
1415 Hawaiian Standard Time the ship had de- 
ployed two large parachutes as sea anchors at a 
depth of about 165 m in water with a bottom 
depth of approximately 4600 m. When the para- 
chutes were hauled to the surface, using a small 
winch with an 180 kg pull shut-off, one of them 
had entangled in it a large adult male shark 4.46 
m (14.6 ft) long and 750 kg (1653 Ibs) (Figs. 1- 


[87] 


4). Crew members of the AFB-/4 realized that 
the shark was unusual and brought it aboard with 
much difficulty. The shark was shipped to the 
Kaneohe Bay facility of the Naval Undersea Cen- 
ter and tied alongside the dock overnight. 

The senior author inspected it the next morn- 
ing. Preliminary examination indicated that it 
represented a very distinct, undescribed species, 
and it was decided that it should be preserved 
intact. Accordingly, the shark was winched out 
of the water by the tail using a Navy crane, but 
the caudal fin broke off and the shark fell into 
the water and had to be retrieved by divers. The 
shark was quick-frozen at Hawaiian Tuna Pack- 
ers, Honolulu, while a large preservation tank 


| Laboratery 


RICHARD ELLIS -1977 


Ficure 1. Artist’s conception of Megachasma pelagios in its natural habitat, slowly swimming with open mouth and feeding 
on planktonic animals. (From a painting by Richard Ellis.) 


88 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 89 


was constructed. On 29 November 1976, the 
shark was transported frozen to the Kewalo dock 
site of the National Marine Fisheries Service for 
thawing and injection with formalin. Subsequent 
examination of the shark by the authors and col- 
leagues indicated that it is a lamniform shark 
(order Lamniformes of Compagno 1973a) that 
is not assignable to any known genus or family 
and is herein described as Megachasma pelagios, 
new genus and species, and placed in the new 
family Megachasmidae. 

The discovery of the novel shark was widely 
reported in newspapers (e.g., Anonymous | and 
2, 1976, and Dunford 1976). It was dubbed the 
‘““Megamouth shark” in reference to its unusually 
large oral cavity. This common name has since 
been adopted by several authors (Compagno 
1977, 1979, and 1981; Taylor 1977; Tinker 1978; 
Faughnan 1980; Clark 1981), and we suggest that 
it be considered as the accepted common name 
for the species. 

This strange shark is extraordinary in its dis- 
tinctness from other sharks and its great size. 
Most sharks are small, less than 2 m long at 
maturity (Compagno 1981). The new shark joins 
the company of the few giant sharks commonly 
reaching total lengths over 4 m, including the 
broadnose sixgill shark (Hexanchus griseus), Pa- 
cific sleeper shark (Somniosus pacificus), Green- 
land shark (.S. microcephalus), whale shark (Rhi- 
niodon typus), great white shark (Carcharodon 
carcharias), tiger shark (Galeocerdo cuvier), and 
great hammerhead (Sphyrna mokarran). The 
common thresher (Alopias vulpinus) and bigeye 
thresher (A. superciliosus) also reach total lengths 
over 4 m, but these sharks have greatly elongated 
caudal fins and hence are relatively small-bodied 
in comparison to the giant species. Although new 
species of small sharks are discovered fairly fre- 
quently, giant sharks are not, and almost all of 
the great species were described in the 18th and 
19th centuries. 

Because the only known specimen of Mega- 
chasma pelagios is an adult male, and because 
it is very common for female sharks to reach a 
somewhat larger size than males, it is reasonable 
to expect larger specimens of this species. 


METHODS 


On 30 November 1976, the thawed shark was 
placed in a large, above-ground plastic pool filled 
with seawater. Comprehensive measurements of 
the shark were recorded, following the proce- 


dures of Bigelow and Schroeder (1948). Skin 
samples were taken from the mouth, tongue, pec- 
toral fin, caudal fin, back below first dorsal fin, 
and gill-rakers for later examination using a 
Cambridge S410 Stereoscan electron micro- 
scope. 

Skin samples from the mouth lining and tongue 
were sectioned and stained using standard his- 
tological techniques. A short incision, approxi- 
mately 30 cm long, was made on the ventral 
surface to gain access to the stomach and val- 
vular intestine, and stomach contents were re- 
moved. The valvular intestine was removed, slit 
medially to count the ring valves and to remove 
intestinal worms for parasitologists (Dailey and 
Vogelbein 1982), and separately preserved. Sam- 
ples of muscle tissue and liver were taken for 
electrophoretic analysis. Extensive sets of still 
photos were made of the preservation process by 
the authors and Mr. Paul Meyers of the Naval 
Undersea Center, who also made 16 mm movies 
of these techniques. 

The body cavity and musculature of the shark 
were injected with 25 | of 100% formalin (40% 
aqueous formaldehyde gas solution). The shark 
was then lifted by crane and cargo sling into a 
4 x 3.5 X 1 m fiberglass box and covered with a 
40% seawater-formalin solution. After six months 
in formalin, the specimen was deposited in the 
fish collection of the Bernice P. Bishop Museum, 
Honolulu, where it was rinsed for 30 days in 
water and then placed in 55% isopropyl alcohol. 
Tooth samples were removed for examination 
of their morphology, and one tooth was sectioned 
and stained for tooth histology. A “peel” dissec- 
tion was made on the right side of the head to 
examine the neurocranium and jaw structure of 
the shark, and similar dissections were made on 
the right pectoral fin and right clasper. Vertebrae 
were excised from the base of the caudal fin and 
from beneath the first dorsal fin and sectioned 
to examine their calcification patterns. 

Terminology for descriptive morphology of 
Megachasma pelagios follows Bigelow and 
Schroeder (1948) and Compagno (1970, 1973a, 
1973b, and 1979). 


Megachasmidae, new family 


Type-GeNus.— Megachasma Taylor, Compagno, and Struh- 
saker, new genus. 
FAMILY DESCRIPTION. —Giant neoselachian 
sharks of the order Lamniformes (as defined by 


90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


Ficure 2. Holotype of Megachasma pelagios, within 12 hours of its capture. Note the extreme protrusibility of the jaws 


and the gill filaments visible in the first gill opening. 


Compagno 1973a) reaching at least 4.46 m length 
when adult. Trunk cylindrical but not highly fu- 
siform, tapering rearward from the head. Caudal 
peduncle short, stout, slightly compressed, and 


without lateral keels or ridges; a shallow, longi- 
tudinally oval upper precaudal pit present, but 
no lower pit. Head broad, very large and long, 
and not pointed, length greater than abdomen 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 91 


Ficure 3. Frontal view. Note Navy research vessel and winch which retrieved shark in background. (Official U.S. Navy 
photograph.) 


between pectoral and pelvic bases. Snout very Nostrils small, widths about 4, internarial 
short, depressed, and broadly rounded, not con- _ width, with short, low anterior nasal flaps; nos- 
ical or bladelike. Eyes lateral on head, length less _ trils lateral and opposite the first fourth of mouth. 
than one-fourth length of longest gill openings. Gill openings moderately large, not expanded 


92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


onto dorsal surface of head; internal gill openings 
with numerous gill-rakers of a unique type, 
formed as elongated, slender, cartilage-cored 
dermal papillae covered by imbricated denticles 
(Fig. 6). Mouth terminal and very large, broadly 
arched, extremely long, and extending far behind 
eyes when jaws are not protruded; jaws strongly 
protrusible, capable of extension well in front of 
snout. No true labial furrows or labial cartilages, 
but with inner labial grooves present along edges 
of mouth corners. 

Teeth similar in upper and lower jaws, weakly 
differentiated, with moderately long, broad, flat- 
tened roots, very short labial root lobes, and very 
long, broad, expanded lingual protuberance; 
moderately strong basal ledges and grooves on 
the labial crown face; a broad, enameloid-free 
neck on the crown foot; a strong, narrow, lin- 
gually hooked cusp with cutting edges confined 
to its tip, no striations or ridges on the cusp. 
Teeth not compressed and bladelike, relatively 
small, and very numerous, over 100 rows in each 
jaw and in three or four functional series. Tooth- 
less spaces on symphyses of jaws extremely broad, 
especially on upper jaw. Teeth in each jaw half 
apparently continuously varying and without 
discrete row groups; no gap or reduced inter- 
mediate teeth between teeth in anterior and lat- 
eral positions in upper jaw. 

Lateral trunk denticles with broad, teardrop, 
or wedge-shaped, flattened crowns, not erect, 
hooked, or directed anteriorly or dorsoventrally; 
pedicels of trunk denticles low and broad (Fig. 
11). Wavy grooves of naked skin on the pectoral, 
pelvic, and caudal fin webs. Pectoral fins rela- 
tively narrow, long and blunt-tipped, length from 
origin to free rear tip about half as long as pec- 
toral anterior margin. Origins of pectoral fins un- 
der fourth gill openings. Pectoral fins more than 
three times area of first dorsal fin, with anterior 
margins more than three times length of pelvic 
anterior margins. Pectoral fin skeleton plesodic, 
with pectoral radials extending into the distal fin 
web nearly to its edge; ceratotrichia reduced along 
distal fin margin and not extending proximally 
to radial musculature of fin. Pectoral fins very 
small, angular, smaller than first dorsal fin but 
larger than second dorsal, with an aplesodic fin 
skeleton. Claspers moderately slender and elon- 
gated, with attenuated tips and external spurs 
(Fig. 11). First dorsal fin moderately large, angular 
and relatively low, with a narrowly rounded apex 


and an aplesodic fin skeleton; origin of first dorsal 
much closer to pectoral fin bases than pelvic bas- 
es, and free near tip, well in front of pelvic origins. 
Second dorsal fin less than one-third area of first 
dorsal and slightly less than half as high, angular 
and broad-based, with its origin about over the 
pelvic fin insertions. Neither second dorsal nor 
anal bases pivoted. Anal fin about half area of 
second dorsal, angular and broad-based, with its 
origin about opposite free rear tip of second dor- 
sal and its free rear tip well in front of ventral 
caudal origin; insertion of anal separated from 
ventral caudal origin by space greater than base 
or anal. Caudal fin with a long dorsal lobe nearly 
half length of rest of shark, a long ventral lobe 
about % as long as dorsal lobe, a deeply notched 
postventral caudal margin, a weak subterminal 
notch, and no undulations or ripples on the dor- 
sal or preventral caudal margins; caudal fin not 
lunate or crescentic, dorsal caudal vertebral axis 
moderately elevated at an angle to body axis (het- 
erocercal). 

Neurocranium (Fig. 13) with tripodal rostrum 
formed of a small, moderately elongated, medial 
rostral cartilage originating from expanded in- 
ternasal plate and pair of basally enlarged, tri- 
angular lateral rostral cartilages that taper an- 
teromedially to fuse with medial rostral cartilage 
and form a narrow, flattened, unfenestrated ros- 
tral node. Base of medial rostral cartilage ele- 
vated by dorsally arched internasal septum above 
level of bases of lateral rostral cartilages and nasal 
capsules, so that medial rostral cartilage arches 
anteroventrally to meet rostral node. Rostrum 
short, less than half nasobasal length of cranium. 
Nasal capsules small, greatly compressed, far lat- 
eral to each other and separated by flattened in- 
ternasal septum. Entire anterior surfaces of nasal 
capsules forming bases of lateral rostral carti- 
lages. Broad subethmoid fossa not extending an- 
terior to nasal capsules. Cranial roof very broad 
and flat, not arched above the orbits, with a huge 
transverse anterior fontanelle; basal plate broad- 
ly arched. Orbits with low preorbital processes, 
complete supraorbital crests, and broad, low 
postorbital processes. A deep pit on each side of 
ventral surface of cranium between base of sub- 
orbital shelf and basal plate in front of stapedial 
fenestrae, for orbital processes of palatoquad- 
rates. Otic capsules broad and relatively long, 
without elongated pterotic horns. 

Jaws very long and stout, much longer than 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 93 


cranium, extending from the cranial rostral node 
to well behind the occiput when retracted. Pal- 
atoquadrates with long, stout palatine processes 
lacking dental bullae; strong, low, knoblike, car- 
tilaginous orbital processes, and low, strong 
quadrate processes. Orbital processes articulat- 
ing on ventral surfaces of suborbital shelves and 
basal plate below orbits, quadrate processes far 
below postorbital processes of cranium and not 
contacting them. Anterior ends of Meckel’s car- 
tilages extending below level of anterior ends of 
palatoquadrates, no “‘overbite” of palatoquad- 
rates on Meckel’s cartilages. Vertebral column 
with well-developed cartilaginous centra sepa- 
rated by broad bands of notochordal sheath, but 
with primary and secondary calcification vir- 
tually absent. Intestinal valve with 24 turns. 

CLASSIFICATION. —Compagno (1973a, 1977) 
divided the living elasmobranch fishes, or neo- 
selachians, into four superorders, of which the 
Galeomorphii or galeomorph sharks clearly 1n- 
cludes the new family Megachasmidae and genus 
Megachasma. Megachasmidae has the following 
galeomorph characters: head and body not great- 
ly depressed and not expanded laterally; spiracles 
without valves; five pairs of laterally situated gill 
openings; denticles covering almost entire body, 
not absent ventrally, nor enlarged on midline of 
back, and not enlarged on pectoral fins in adult 
males; pectoral fins without anteriorly expanded 
triangular lobes covering gills or fused to sides 
of head above them; propterygium of pectoral 
fin skeleton not anteriorly expanded; pectoral fins 
not modified into propulsive organs; pectoral gir- 
dle not articulating with vertebral column; vent 
confluent with pelvic fins; two dorsal fins and an 
anal fin present; caudal fin heterocercal, with a 
subterminal notch on the dorsal caudal lobe and 
with ventral lobe shorter than dorsal lobe; neuro- 
cranium with strong suborbital shelves, no antor- 
bital cartilages, ectethmoid processes, or en- 
larged ectethmoid chambers on the nasal capsules, 
rostrum not trough-shaped, no basal angle on 
basal plate, no lateral commissures on otic cap- 
sules, and with incomplete postorbital walls; no 
palatobasal articulation of palatoquadrates with 
neurocranium; hyoid arch complete, no pseu- 
dohyoids; vertebral column without synarcuals, 
and vertebral centra without concentric calcifi- 
cations. 

The Galeomorphii of Compagno (1973a, 1977) 


was subdivided into four orders, Heterodonti- 
formes, Orectolobiformes, Carcharhiniformes, 
and Lamniformes; of these, the family Mega- 
chasmidae falls in the order Lamniformes or 
lamnoid sharks. Lamnoid characters of Mega- 
chasmidae include its simple nostrils of the or- 
dinary shark type, entirely separate from the 
mouth, with small anterior nasal flaps, diagonal 
incurrent and excurrent apertures, and no peri- 
nasal folds and grooves, anterior barbels, or na- 
soral grooves; a long mouth extending behind 
the eyes when jaws are retracted; no supraorbital 
and subocular ridges; eyes circular and laterally 
without nictitating eyelids, subocular pouches, 
or postorbital eyelid muscles; osteodont teeth (Fig. 
9) with weak basal ledges; posterior teeth not 
enlarged and formed into molariform crushers; 
claspers with external spurs on the T-3 cartilage 
and with elongated, tubular, expanded marginal 
cartilages; dorsal fins spineless, with segmented 
basal cartilages; cranium with a tripodal rostrum, 
nasal capsules not anteroposteriorly elongated and 
trumpet-shaped; no ethmopalatine grooves for 
the articulation of the palatoquadrate orbital 
processes, complete preorbital walls, separate fo- 
ramina for superficial ophthalmic nerves in or- 
bits and for hyomandibular nerves on otic cap- 
sules, and relatively long otic capsules; jaws long, 
extending posterior to the occiput; mouth gape 
not restricted anteriorly, labial cartilages, folds 
and grooves reduced or absent; pectoral fin skel- 
eton with a small propterygium, moderately large 
mesopterygium, and large metapterygium; me- 
sopterygium and metapterygium not elongated 
parallel to the axes of their radials, and not proxi- 
mally shaftlike, distally expanded and without a 
fenestra between them; preorbitalis or levator 
labii superioris muscles relatively small and 
anteroposteriorly positioned on the jaws, with 
origins on posteroventral surfaces of the nasal 
capsules, fibers nearly horizontal when jaws are 
retracted, and insertions far posterior on the ad- 
ductor mandibulae muscles at the jaw angles; 
adductor mandibulae muscles anteriorly notched; 
levator palatoquadrati muscles simple, not sub- 
divided into anterior constrictor and spiracular 
muscles; no craniomandibular or mandibulo- 
cutaneous muscles; and an elongated, ring-valve 
intestine. 

RELATIONSHIPS TO OTHER LAMNOIDS. — Within 
the Lamniformes, the family Megachasmidae 


94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


represents a very distinct and singular taxon, 
well separated from all other families. Other 
lamnoids of the families Odontaspididae, Mitsu- 
kurinidae, Pseudocarchariidae, Alopiidae, Ceto- 
rhinidae, and Lamnidae all differ from the Me- 
gachasmidae in having a more elongated, more 
narrowly rounded, conical or bladelike snout; no 
papillose gill-rakers (denticle gill-rakers present 
in Cetorhinidae); mouth subterminal on head and 
less enlarged; tongue smaller; upper anterior and 
lateral teeth separated by a gap that may or may 
not have reduced intermediate teeth; tooth rows 
either less than 60 in each jaw, or more than 200 
(Cetorhinidae); toothless space on upper sym- 
physis relatively narrow; no wavy grooves of na- 
ked skin on the pectoral, pelvic, and caudal fins; 
lateral rostral cartilages narrow-based and only 
covering part of the dorsal surfaces of the nasal 
capsules or the preorbital processes; base of me- 
dial rostral cartilage well below bases of lateral 
rostral cartilages and with shaft of cartilage below 
rostral node; nasal capsules nearly spherical, not 
compressed, and with ventral nasal apertures; 
cranial roof narrow to only moderately expanded 
anteriorly, with anterior fontanelle varying from 
moderate to greatly reduced; orbital processes 
more or less reduced on palatoquadrates, artic- 
ulating with the suborbital shelves where present; 
jaws shorter, beginning well behind the snout tip 
when retracted; and with primary calcification 
of the double cones and secondary radii well de- 
veloped in their vertebral centra. 

Members of the family Odontaspididae (in- 
cluding the genera Eugomphodus and Odontas- 
pis) further differ from Megachasma and the Me- 
gachasmidae in having prominent, transverse 
precaudal pits; labial folds, furrows and carti- 
lages present (with the possible exception of E. 
tricuspidatus), nostrils in front of the mouth; teeth 
with strong labial root lobes, moderate lingual 
protuberances, narrow necks on the crown, and 
labiolingually diagonal attachment surfaces; 
symphyseal, anterior, lateral, intermediate, and 
posterior tooth-row groups well differentiated 
along dental bands, with anteriors and laterals 
enlarged; pectoral fins smaller, shorter, broader, 
less elongated, and not falcate, and with aple- 
sodic fin skeletons; pectoral fin origins behind 
fifth gill openings; claspers stouter and blunt- 
tipped, with blunt clasper spurs; origin of first 
dorsal fin well posterior to pectoral insertions; 


second dorsal fin more than half as high as first 
dorsal; caudal fin shorter, less than half as long 
as rest of shark; subterminal notch of caudal fin 
deep; ventral caudal lobe shorter, dorsal caudal 
margin with rippled edges; rostral node com- 
pressed, with vertical fenestra and strut; cranial 
roof narrow and arched above orbits; and otic 
capsules with strong pterotic horns. 

The family Pseudocarchariidae, which like 
Megachasmidae has a single, oceanic, highly dis- 
tinct species (Pseudocarcharias kamoharai), dif- 
fers from Megachasma in many characters, in- 
cluding its more slender body and shorter head; 
slender, cylindrical caudal peduncle with low lat- 
eral keels and upper and lower transverse, cres- 
centic precaudal pits; much larger eyes; nostrils 
anterior to mouth; more elongated gill openings, 
extending onto dorsal surface of head; teeth with 
strong labial root lobes, moderate lingual pro- 
tuberances, a narrow neck on the crown, and 
labiolingually diagonal attachment surfaces; an- 
teriors, intermediates, and lateroposteriors well 
differentiated in dental bands; anteriors and an- 
terior-laterals enlarged, pectoral fins smaller, 
broader, less elongated, and not falcate, with 
aplesodic fin skeletons; origins of pectoral fins 
behind fifth gill openings; anal fin with a narrow 
base and pivotable; caudal fin with a shorter dor- 
sal and ventral caudal lobe; rostrum longer, with 
appendices, a compressed rostral node, and ver- 
tical fenestrae and struts; basal plate and cranial 
roof extremely narrow, with narrow, slotlike, 
vertical anterior fontanelle; orbits of cranium very 
large; pterotic horns present and well developed 
on otic capsules; palatine processes of palato- 
quadrates enlarged and forming large dental bul- 
lae, articulating with the orbital notches of the 
cranium; and quadrate processes of palatoquad- 
rates elevated and contacting postorbital pro- 
cesses. 

The benthopelagic family Mitsukurinidae also 
has a single living, strongly distinct species (Mit- 
sukurina owstoni). The Mitsukurinidae differs 
from the Megachasmidae in lacking precaudal 
pits; having a greatly elongated, flattened, blade- 
like snout; smaller eyes; a very narrow, elongated 
mouth; lower labial furrows; teeth with strong 
labial root lobes, moderate lingual protuber- 
ances, a narrow neck and striations on the crown, 
and labiolingually diagonal attachment surfaces; 
symphyseals, anteriors, laterals, and posteriors 
well differentiated in dental bands, with anteriors 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 95 


and laterals enlarged; lateral trunk denticles with 
narrow, hooked, semierect crowns; pectoral fins 
smaller than pelvic fins, shorter, broader, not 
elongated and falcate, and with aplesodic fin skel- 
etons; pectoral origins behind fifth gill openings; 
first and second dorsal fins equal-sized, smaller 
than pelvic and anal fins; anal fin large, broadly 
rounded, and separated from lower caudal origin 
by a narrow notch; anal fin origin about opposite 
or close behind second dorsal origin; no ventral 
caudal lobe; rostrum of cranium greatly elon- 
gated, longer than nasobasal length of cranium, 
with a compressed, extremely long rostral node; 
subethmoid fossa extending anterior to the nasal 
capsules; supraorbital crest reduced to separate 
preorbital and postorbital processes; and with 
palatine processes of palatoquadrates deflected 
ventrally, with prominent bullae. 

The three highly specialized lamnoid families 
Alopiidae, Lamnidae, and Cetorhinidae have 
numerous additional differences from the Me- 
gachasmidae. The Alopiidae further differs from 
the Megachasmidae in having a shorter head; 
crescentic upper precaudal pits; larger eyes; nos- 
trils anterior to mouth; shorter gill openings; a 
much smaller mouth and less highly protrusible 
jaws; teeth with weaker lingual protuberances, 
stronger labial root lobes, and differentiated an- 
teriors, lateroposteriors, and (variably) inter- 
mediates and symphyseals; claspers very slender, 
without spurs; pelvic fins plesodic; first dorsal fin 
higher and plesodic, with its origin well posterior 
to the pectoral insertions; second dorsal much 
smaller relative to first dorsal, with a narrow, 
pivotable base; anal fin smaller, with narrow, 
pivotable base; caudal fin about as long as rest 
of shark, with a rippled dorsal margin; rostral 
node of rostrum compressed, with a vertical fe- 
nestra and strut; internasal septum narrow and 
high; subethmoid fossa very narrow; cranial roof 
narrow, flat or strongly arched; orbits large to 
gigantic; and palatine processes of palatoquad- 
rates with small dental bullae. 

The Lamnidae differs from the Megachasmi- 
dae in the following additional characters: trunk 
more fusiform; caudal peduncle greatly de- 
pressed, with strong lateral keels, and with trans- 
verse, crescentic, upper and lower precaudal pits: 
nostrils anterior to the mouth; gill openings long- 
er, extending partway onto dorsal surface of head: 
jaws less protrusible; teeth with low lingual pro- 
tuberances, enlarged anteriors, laterals, and in- 


termediates; pectoral fin origins behind fifth gill 
openings; second dorsal much smaller relative to 
first dorsal, with a narrow, pivotable base; anal 
fin slightly larger than second dorsal, with a nar- 
row, pivotable base; caudal fin shorter, less than 
half length of rest of shark, nearly symmetrical 
and lunate in Lamnidae, with a relatively shorter 
dorsal lobe, ripples in dorsal margin, and a longer 
ventral lobe; cranial roof narrow and arched; otic 
capsules with elongated pterotic horns; palatine 
processes of palatoquadrates with prominent 
dental bullae articulating with underside of eth- 
moid region of cranium; and quadrate processes 
of palatoquadrates very high. 

Finally, the family Cetorhinidae with the only 
other filter-feeding lamnoids of the genus Cetor- 
hinus, differs from the family Megachasmidae in 
the following particulars: trunk more fusiform; 
caudal peduncle somewhat depressed, with strong 
lateral keels and transverse, crescentic upper and 
lower precaudal pits; nostrils anterior to mouth; 
gill openings much larger, expanded onto dorsal 
and ventral surfaces of head; jaws little protru- 
sible; pectoral fins with their origins behind fifth 
gill openings; lateral trunk denticles with erect, 
hooked, narrow crowns, directed anteriorly and 
dorsoventrally as well as posteriorly; claspers 
stout, with broad tips and heavy spurs; first dor- 
sal fin with its origin far posterior to pectoral 
insertions, and midbase closer to pelvic bases 
than to pectoral bases; caudal fin shorter, less 
than half length of rest of shark, nearly sym- 
metrical and lunate, with a shorter dorsal lobe 
and longer ventral one; medial rostral cartilage 
very broad, platelike, and ventrally excavated by 
the broad anterior expansion of the subethmoid 
fossa; lateral rostral cartilages joining each other 
posterior to their junction with the rostral node, 
and extending anterior to that junction as a me- 
dial rod; cranial roof moderately broad, highly 
arched above orbits; supraorbital crests fenes- 
trate basally; and jaws very slender and weak. 

The phenetic comparisons between Mega- 
chasmidae and other lamnoids presented above 
are not intended to be exhaustive, but serve to 
demonstrate the separation of Megachasmidae 
from related families. They do not broach the 
question of the relationship of the megamouth 
shark to other lamnoids. A detailed account of 
lamnoid interrelationships is beyond the scope 
of this paper, but suffice it to note here that many 
of the characters of Megachasma pelagios, such 


96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


as its snout and jaw structure, gill-rakers, dermal 
grooves on fins, reduced vertebrae, and ethmoid 
morphology are evidently unique derived char- 
acters of this shark that do not offer a clue to its 
relationships. Its teeth are superficially similar 
to those of the basking shark (Cetorhinus max- 
imus), but this may be parallel evolution of ves- 
tigial structures in two very different lamnoid 
filter-feeders (as suggested by the superficial sim- 
ilarity of the teeth of the orectoloboid whale shark, 
Rhiniodon typus). Megachasma most resembles 
members of the Odontaspididae (especially the 
genus Odontaspis) and Pseudocarchariidae in its 
body shape, fin shape, relative fin sizes (except 
for the pectoral fins), fin positions, and relation- 
ships of interspaces between fins to fin size. By 
comparison with the derived families Alopiudae, 
Cetorhinidae, and Lamnidae, these similarities 
between Megachasmidae, Odontaspididae, and 
Pseudocarchariidae may prove to be common 
primitive characters not of importance in dem- 
onstrating phyletic relationships among these 
families. Megachasma shares the derived char- 
acter state of plesodic pectoral fins with the Al- 
opiidae, Cetorhinidae, and Lamnidae, but pres- 
ently appears to have little else in common with 
these derived families. 

On the other hand, two characters of Mega- 
chasma, if correctly interpreted as primitive, sug- 
gest that Megachasmidae is the sister-group of 
all other living lamnoids. The absence of differ- 
entiated anteriors, laterals, and intermediates (or 
a toothless gap between anteriors and laterals) in 
Megachasma may indicate that it is primitive in 
lacking them, and that all other lamnoids (in- 
cluding Cetorhinus) can be united by the pres- 
ence of these tooth-row groups as a shared de- 
rived character. However, the unusually broad, 
toothless space at the upper symphysis of Me- 
gachasma suggests another possibility, that it is 
derived in having lost these row groups, at least 
in the upper jaw; and that the simple gradient 
monognathic heterodonty in the dentral bands 
is secondary and correlated with the evolution 
of gill-rakers as the primary feeding structures in 
Megachasma. 

The second character is the well-developed or- 
bital processes on the palatoquadrates of Me- 
gachasma, which suggest a primitive condition 
by comparison with other, non-lamnoid sharks. 
The reduced (Alopiidae, Odontaspididae, Mit- 
sukurinidae, and Cetorhinidae) or apparently 


nonexistent (Lamnidae, Pseudocarchariidae) or- 
bital processes of other lamnoids would by this 
interpretation represent a shared derived char- 
acter of lamnoids other than Megachasma. A 
detailed assessment of these characters and oth- 
ers, grouping the various lamnoid genera and 
families will be considered in detail elsewhere 
(Compagno, in preparation). 

A possible fossil relative of M. pelagios is rep- 
resented by isolated small teeth (2-15 mm high) 
known since the 1960’s from early Miocene de- 
posits in the southeastern San Joaquin Valley of 
California (Shelton P. Applegate, pers. comm.), 
and subsequently found in other localities in the 
late Oligocene or early Miocene of northern Cal- 
ifornia (Phillips et al. 1976) and central Oregon 
(Bruce J. Welton, pers. comm.). The shark rep- 
resented by these teeth has never been named, 
but is known from abundant tooth material from 
southern California. Its affinities have been much 
debated among palaeoichthyologists, but it ap- 
pears most likely to be a lamnoid because of its 
osteodont tooth histology and external tooth 
morphology. Dr. Bruce J. Welton is preparing a 
paper describing this shark, and will compare it 
with M. pelagios, of which it is possibly a fossil 
congener but is distinctly more primitive. 


Megachasima, new genus 

Type-species. —Megachasma pelagios Taylor, Compagno, and 

Struhsaker, new species. 

DERIVATION OF NAME.— mega, from Greek, 
large, great; chasma, yawning hole, open mouth. 

GENERIC D1IAGNOsIs.—Characters of the new 
genus are those of the new family Megachas- 
midae (see above). 


Megachasma pelagios, sp.nov. 
MEGAMOUTH SHARK 


Hotoryre.—An adult male, 4460 mm total length, Bernice 
P. Bishop Museum, Honolulu, Oahu, Hawaii, BPBM 22730. 

TyYPE-LOCALITY.— Hawaiian Islands, about 42 km NE Ka- 
huku Point, Oahu, 21°51’N, 157°46'W, at about 165 m depth 
in water about 4600 m deep. 

DERIVATION OF SPECIES NAME. — pelagios, from 
Greek, of the open sea. 

MEASUREMENTS AND PROPORTIONS. — These are 
given below as measurements in millimeters, fol- 
lowed by their proportions as percentages of total 
length and precaudal length, given in that order 
in parentheses. 

Total length: 4460 mm (100% total length, 
144.3% precaudal length). 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 97 


Ficure 4. Lateral-view drawing of the holotype of Megachasma pelagios, with jaws in retracted position. Drawn by L. J. 


V. Compagno. 


Precaudal length (snout to upper caudal ori- 
gin): 3090 (69.3, 100). 

Tip of snout to: upper symphysis, 66 (1.5, 2.1); 
nostrils, 100 (2.2, 3.2); orbits, 240 (5.4, 7.8); spi- 
racles, 450 (10.1, 14.6); Ist gill openings, 850 
(19.1, 27.5); 2nd gill openings, 920 (20.6, 29.8); 
3rd gill openings, 1020 (22.0, 33.0); 4th gill open- 
ings, 1150 (25.8, 37.2); Sth gill openings (head 
length), 1180 (26.5, 38.2); pectoral origins, 1110 
(24.9, 35.9); pelvic origins, 2270 (50.9, 73.5); Ist 
dorsal origin, 1540 (34.5, 49.8); 2nd dorsal ori- 
gin, 2530 (56.7; 81.9); anal origin, 2830 (63.5, 
Sieo)eavent, 2295.(5 1.5; 74:3). 

Distance between: vent and caudal tip, 2165 
(48.5, 70.1); 1st and 2nd dorsal origins, 625 (14.0, 
20.3); Ist and 2nd dorsal bases, 590 (13.2, 19.1); 
2nd dorsal and upper caudal origins, 428 (9.6, 
13.9); 2nd dorsal base and upper caudal origin, 
395 (8.9, 12.8); pectoral and pelvic origins, 510 
(11.4, 16.5); pelvic and anal bases, 330 (7.4, 10.7); 
anal and lower caudal origins, 315 (7.1, 10.2); 
anal base and lower caudal origin, 230 (5.2, 7.4). 

Eyes (palpebral apertures or fleshy orbits): 
length, 56 (1.3, 108); height, 54 (1.2, 1.7); width 
across anterior corners (interorbital), 370 (8.3, 
12.0); eyeball diameter, 84 (1.9, 2.7). 

Nostrils: width, 30 (.07, 1.0); internarial space, 
340 (7.6, 8.8). 

Spiracles: diameter, 6 (0.1, 0.2); space between 
spiracles and eyes, 176 (3.9, 5.7). 

Mouth (jaws in retracted position): length, 273 
(6.1, 8.8); width, 827 (18.5, 26.8); width across 
outer edges of jaws, 1025 (23.0, 33.2); length of 
lower jaw, 820 (18.4, 26.5). 

Gill opening widths (heights): Ist, 265 (5.9, 
8.6); 2nd, 258 (5.8, 8.4); 3rd, 264 (5.9, 8.5); 4th, 
256:(5.7 28:3); Sth: 234.(5.2, 7:6). 


Head height: at spiracles, 500 (11.2, 16.2); at 
Ist gill openings, 625 (14.0, 20.2); at Sth gill 
openings, 630 (14.1, 20.4). 

Trunk height: at Ist dorsal origin, 640 (14.3, 
20.7); at pelvic origins, 515 (11.5, 16.7); at pelvic 
insertions, 440 (9.9, 14.2). 

Girth: at Ist dorsal origin, 1800 (40.4, 58.2); 
at 2nd dorsal origin, 1140 (25.6, 36.9). 

Caudal peduncle height: at 2nd dorsal inser- 
tion, 341 (7.6, 11.0); at upper caudal origin, 237 
6357): 

Caudal peduncle width: at 2nd insertion, 146 
(3.3, 4.7); at upper caudal origin, 109 (2.4, 3.5). 

Pectoral fins, length of: anterior margin, 837 
(18.8, 27.1); posterior margin, 615 (13.8, 19.9); 
base, 262 (5.9, 8.5); origin to free rear tip, 453 
(10.1, 14.7); inner margin, 190 (4.3, 6.1). 

Pelvic fins, length of: anterior margin, 264 (5.9, 
8.5); posterior margin, 181 (4.1, 5.9); base, 207 
(4.6, 6.7); origin to free rear tip, 245 (5.5, 7.9): 
inner margin, 38 (0.8, 1.2); height, 255 (5.7, 8.3); 
origin to rear tip of clasper, 575 (12.9, 18.6). 

Claspers: inner length from vent to tip, 550 
(12.3, 17.8); outer length from clasper base to 
tip, 355 (8.0, 11.5); width at outer pelvic base, 
AT (et S15): 

lst dorsal fin, length of: anterior margin, 415 
(9.3, 13.4); posterior margin, 265 (5.9, 8.6); base, 
404 (9.1, 13.1); inner margin, 82 (1.8, 2.7); height, 
DLE Bally 18) F 

2nd dorsal fin, length of: anterior margin, 198 
(4.4, 6.4); posterior margin, 158 (3.5, 5.1); base, 
191 (4.3, 6.2); inner margin, 80 (1.8, 2.6); height, 
104 (2.3, 3.4). 

Anal fin, length of: anterior margin, 196 (4.4, 
6.3); posterior margin, 80 (1.8, 2.6); base, 159 


98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


FiGure 5. (a) Schematic head drawing of Megachasma pe- 
lagios with jaws protruded. (b) Photograph of fresh shark with 
jaws protruded. (Upper arrow points to anteriormost edge of 
neurocranium; lower arrow to upper jaw and teeth.) 


(3.6, 5.1); inner margin, 67 (1.5, 2.2); height, 78 
(eT 2e5): 

Caudal fin, length of: dorsal margin, 1443 (32.3, 
46.7); preventral margin, 625 (14.0, 20.2); lower 
postventral margin, 377 (8.5, 12.2); upper post- 
ventral margin, 1220 (27.4, 39.5); subterminal 
margin, 57 (1.3, 1.8); terminal margin, 96 (2.2, 
3.1); terminal lobe or sector, 139 (3.1, 4.5); width 
of dorsal lobe at postventral notch, 471 (10.6, 
15.2); width of ventral lobe at postventral notch, 
DIS (Galeeses): 

Intestinal valve: length, 690 (15.5, 22.3); di- 
ameter, 145 (3.3, 4.7); thickness of broadest an- 
terior ring, 25 (0.6, 0.8). 

DESCRIPTION (based on the holotype and only 
known specimen).— Head length from snout tip 
to 5th gill openings, 26% of total length and 1.6 
times distance between pectoral and pelvic fin 
bases. Head broad, cylindrical, and approxi- 
mately circular in transverse section at eyes, but 
somewhat laterally expanded and oval in section 


over jaws when jaws are retracted; not depressed. 
Outline of head in lateral view nearly straight 
dorsally, except for bluntly convex snout, strong- 
ly convex ventrally along edges of lower jaws and 
nearly straight beneath gills; in dorsoventral view, 
anteriorly rounded and convex and tapering pos- 
teriorly to gills. Snout length from tip to edge of 
mouth about 12.5 times in mouth width. Snout 

roadly rounded in dorsal view, with lateral mar- 
gin slightly indented anterior to nostrils; in lateral 
view, convex dorsally and concave ventrally to 
fit the front of the retracted upper jaw (Fig. 5). 
External eye opening (palpebral aperture) or fleshy 
orbit without anterior or posterior notches, length 
about 21 times in head length. Irises of eyes black, 
nearly filling orbits. Eyeballs large, diameter 14 
times in head length. Spiracles small, their lengths 
about !/10 orbit length, located about 3 orbit 
lengths behind eyes and about opposite ventral 
margins of eyes. Gill openings of nearly equal 
length, the longest (1st and 3rd) about 4.5 in head 
length and 4.7 times eye length, the smallest (Sth) 
about °/10 length of longest. Edges of gill openings 
nearly straight, not incised, and with filaments 
not exposed when jaws are retracted. Gill open- 
ings with upper ends falling below level of eyes, 
and midheight of head at gill openings. Internal 
gill openings with numerous gill-raker papillae 
arranged in about 4 rows on their anterior and 
posterior edges, including both anterior and pos- 
terior edges of Ist gill cavity between hyoid and 
lst branchial arches and posterior edge of 5th 
gill cavity on anterior edge of Sth gill arch. Gill- 
raker papillae small, about 10-15 mm long, 
densely packed, slender, tapering to blunt point, 
arranged with tips pointing anteromedially into 
pharynx, with thick epidermis and dermis cov- 
ering hyaline cartilage core layered with flat- 
tened, imbricated denticles (Fig. 6). Nostrils with 
large lateral incurrent aperture, anterior nasal flap 
with an undulated, truncated posterior edge, and 
low keel on dorsal surface, but no distinct meso- 
narial flap, small medial excurrent aperture with 
low posterior nasal flap on its rim. Nostrils lateral 
to mouth edge and 2.4 times closer to snout tip 
than to eyes. Nostril width 1.8 in orbit length, 
8.8 times in longest gill opening. Inner labial 
grooves at mouth corners on both upper and 
lower jaws just lateral to dental bands and medial 
to vertical fold of skin sheathing adductor man- 
dibulae muscles. Mouth width when jaws are 
retracted about 1.4 in head length; mouth length 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 99 


b 


Ficure 6. Gill-raker papillae of Megachasma pelagios. (a) Drawing of a group of gill-raker papillae. (b) Scanning electron 
micrograph of the tip of a single papilla, showing the closely imbricated denticles (20X magnification). (c) Scanning electron 


micrograph of denticles from 5b, at higher magnification (51x). 


about 3 times in width. A broad fold of skin 
forming a deep pocket on dorsal surface of upper 
jaws below snout, and a vertical fold of skin en- 
closing anterior edges of adductor mandibulae 
muscles at each mouth corner. Tips of upper jaws 
can extend at least 6 orbit lengths in front of 
snout tip, with mouth corners passing anterior 


to eyes. Tongue extremely large, broadly round- 
ed and thick, enclosing greatly enlarged basihy- 
oid cartilage; tongue almost entirely filling mouth 
cavity when jaws are closed. Deep pocket under 
front of tongue, freeing it anteroventrally; pocket 
about 4 orbit lengths deep from anterior tongue 
edge to its basal attachment to mouth. Maxillary 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


100 


Illustrations of (a) upper symphysis and (c) lower symphysis of Megachasma pelagios, showing bare, toothless 


patches and mesial ends of dental bands. (b) Close-up of rows of teeth from upper jaw. 


FiGure 7. 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 101 


Ficure 8. Illustration of a lower tooth of Megachasma 
pelagios from about midlength on the lower left dental band 
in (A) labial; (B) lingual; (C) distal; and (D) basal views. Ab- 
breviations: AS, attachment surface of root; BG, basal groove; 
BL, basal ledge; CLGF, centrolingual foramen; CR, crown; CU, 
cusp; DCE, distal cutting edge; DRL, distal root lobe; DSH, 
distal shoulder; LAF, labial foramina; LGP, lingual protuber- 
ance; MRL, mesial root lobe; MSH, mesial shoulder; NK, neck; 
RT, root; TG, transverse groove. Drawn by L. J. V. Compagno. 


valve of upper jaw arcuate and relatively narrow, 
width less than an eye diameter. No enlarged 
buccal papillae in mouth cavity, but with scat- 
tered circular organs of undetermined function 
on tongue and mouth. 

Teeth very small and relatively numerous, in 
56 rows in left upper, 59 rows in left lower, 52 
rows in right upper, and 69 rows in right lower 
jaw halves, or 56-52/59-69; total tooth-row 
counts 108/128. Teeth not arranged in diagonal 
files. Symphyseal toothless space about 4 orbit 
diameters wide in upper jaw and less than one 
eye-length wide in lower jaw (Fig. 7). Dental bands 
of upper and lower jaws show strong gradient 
monognathic heterodonty; starting from small 
teeth at symphysis, teeth increase in size to about 
10 mm high in about 10 tooth rows distal to 
symphysis, then begin to gradually decrease in 
size and increase in width relative to height to 
distal ends of dental bands. Teeth (Fig. 8) have 
no cusplets, narrow crown shoulders, partial 
transverse groove on linguobasal attachment 
surface of root, large centrolingual foramen, and 
scattered labial foramina below basal ledge. A 
sectioned tooth (Fig. 9) shows thick osteodentine 


a 


\ 
eg ged 


penny 


FiGure 9. Diagrammatic sagittal section of a lower tooth 
of Megachasma pelagios from about midlength on the lower 
left dental band, lingual protuberance broken off. Abbrevia- 
tions: CR, crown; CU, cusp; END + ORD, enameloid + or- 
thodentine; OSD, osteodentine; RT, root. The teeth of M. 
pelagios are of the “‘osteodont” type, with a crown having a 
core of osteodentine and no pulp cavity or canal. Drawn by 
L. J. V. Compagno. 


core in crown, surrounded by relatively thin lay- 
ers of pallial orthodentine and enameloid, and 
no pulp canal or cavity; crown osteodentine con- 
tinuous with that of root, which forms its sole 
component. 

Body stout, trunk circular or vertically oval in 
section at first dorsal base. Length of head and 
trunk from snout tip to vent 50% of total length. 
Trunk relatively short, length from 5th gill open- 
ing to vent 1.1 times head length. No predorsal, 
interdorsal, or postdorsal ridges on midline of 
back and precaudal lobe; no lateral ridges on 
body. Precaudal lobe from vent to upper caudal 
origin short, 19.2% of total length. Height of cau- 
dal peduncle at insertion of second dorsal 2.3 
times its width there and 1.2 times in distance 
from insertion of second dorsal to upper caudal 
origin; height of caudal peduncle at upper caudal 
origin 2.2 times its width there and 1.7 times in 
distance from insertion of second dorsal to upper 
caudal origin. Upper precaudal pit not transverse 
and crescentic. 

Dermal denticles on body very small and flat- 
tened, giving skin a smooth texture. Denticles on 
sides of trunk below first dorsal fin (lateral trunk 
denticles) loosely spaced, not closely imbricated 
(Fig. 10), with a strong medial ridge and a pair 
of strong lateral ridges running entire length of 
crown, strong medial cusp, but with lateral cusps 
absent or hardly developed. Denticles on dorsal 
surfaces of pelvic fins (Fig. 11) similar to lateral 


102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


Ficure 10. Scanning electron micrographs of dermal den- 
ticles of Megachasma pelagios. (top) Lateral trunk denticles 
from just below the base of the first dorsal fin (60 magnifi- 
cation). (bottom) Denticles from the surface of the tongue (67). 


trunk denticles, except for having lower ridges 
and being closely imbricated. Denticles from 
tongue are wide-spaced but broader and more 
transversely oval than lateral trunk denticles. 
Distal webs of upper surfaces of pectoral and 
pelvic fins, and dorsal caudal lobe, have con- 
spicuous dark wavy lines, often parallel, which 
are channels of bare skin between areas of den- 
ticulate skin (Fig. 11). 

Pectoral fins moderately broad basally but dis- 
tally elongated, tapering, falcate, and broad- 
tipped. Anterior margins of pectoral fins mod- 
erately convex, apices broadly angular, posterior 
margins slightly convex, and free rear tips and 
inner margins smoothly rounded and broadly 
convex. Length of pectoral fin from origin to free 
rear tip 1.9 times in its anterior margin length. 


Apex of pectoral posterior to its free rear tip when 
fin is appressed to body. 

Pectoral fin skeleton with all radials except last 
5 on metapterygium greatly elongated, with 
broad, flattened tips. Radials with numerous seg- 
ments, the longest with 10; distalmost segments 
elongated but only about 4 length of each radial. 
Pectoral fin propterygium supporting one radial, 
mesopterygium with 5 radials, metapterygium 
with 8 radials on basal segment and 8 on axis. 
Propterygium small and slightly elongated dis- 
tally. Mesopterygium moderately elongated dis- 
tally, fairly broad and wedge-shaped with radials 
inserted on distal end at an angle to axis of elon- 
gation. Metapterygium diagonally elongated 
across fin base with radials inserted at an angle 
to long axis. Metapterygial axis of 5 segments, 
about 7% as long as basal metapterygium. Basal 
and radial cartilages of pectoral fins not highly 
calcified; fins rather flexible, despite having ple- 
sodic skeletons. 

Pelvic fins with anterior margin slightly con- 
cave anteriorly but convex posteriorly, apex very 
narrowly rounded, and inner margins slightly 
concave. Inner margins, posterior margins, and 
free rear tips of pelvics forming broad triangle. 

Claspers relatively slender, width at base 7.6 
times outer length from pelvic bases to tips, inner 
length from vent to tip 12.3 percent of total length. 
Rear tips of claspers reaching almost to midbase 
of anal fin when claspers are horizontal. Clasper 
tip elongated, forming a very narrow, slender 
process (Fig. 11), glans anterior to elongated tip 
slightly spatulate and flattened, shaft cylindrical. 
Clasper groove open, with edges not fused dor- 
sally; no pseudopera or lateral clasper groove and 
fold. Small, sharp-tipped, hardened clasper spur 
on ventral lobe, lateral to groove. Large, large- 
mouthed, prominent pseudosiphon on the dorsal 
clasper lobe. 

First dorsal fin with anterior margin slightly 
concave anteriorly and convex posteriorly, pos- 
terior margin nearly straight, free rear tip acute 
and slightly attenuated, and inner margin slightly 
concave. Origin about opposite or slightly pos- 
terior to pectoral fin insertions, midpoint of dor- 
sal base about 2.6 times closer to pectoral inser- 
tions than pelvic origins, dorsal fin insertion 
anterior to pelvic origins by about 0.8 times first 
dorsal base, and free rear tip about 2.9 times 
dorsal inner margin anterior to pelvic origins. 
Posterior margin slanting posteroventrally from 
dorsal apex, insertion well posterior to level of 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 103 


ee 


bi 


aD Mant 


Ficure 11. Right pelvic fin and clasper of Megachasma pelagios, showing channels of naked skin on dorsal surface of pelvic 
fin. (a) Close-up photograph of the channels. (b) Scanning electron micrograph of denticles from the anterior edge of the pelvic 


fin (54X magnification). (c) Drawing of the pelvic fin and clasper. 


dorsal apex. Base 1.5 times in interdorsal space 
and 3.6 times in dorsal caudal margin, height 1.8 
times in base, and inner margin 2.8 times in 
height. 

Second dorsal fin low, height 0.46 times first 
dorsal height, base 0.47 times first dorsal base. 


Anterior margin nearly straight, apex narrowly 
rounded, posterior margin slightly concave, free 
rear tip angular and attenuated, and inner margin 
slightly concave. Free rear tip about over anal 
fin origin. Posterior margin of second dorsal 
slanted posteroventrally from apex, insertion 


104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


Sagittal Section 


CCA 


VRA NOS 


Transverse Section 


FiGure 12. Monospondylous precaudal vertebrae of Me- 
gachasma pelagios, in (top) sagittal section, and (bottom) trans- 
verse section. Abbreviations: Sagittal section: CCA, central 
canal; CCE, cartilaginous centrum; NO, notochord; NOS, no- 
tochordal sheaths; VRA, vestigial radii. Transverse section: 
CCA and VRA, as above; BD, basidorsal; BV, basiventral; DI, 
dorsal intermedial; LIN, lateral intermedial; NAR, neural arch; 
VI, ventral intermedial. (From drawings by L. J. V. Com- 
pagno.) 


posterior to apex. Base 2.1 times in distance from 
insertion to upper caudal origin, height 1.8 times 
base, and inner margin 1.2 times height. 

Anal fin low, height 0.8 times second dorsal 
height, base length 0.8 second dorsal base. An- 


terior margin concave anteriorly but convex pos- 
teriorly, apex broadly rounded, posterior margin 
moderately concave or notched, free rear to acute 
and attentuated, and inner margin slightly con- 
cave. Posterior margin of anal fin slanted pos- 
terodorsally from apex, with anal apex just below 
insertion. Base 1.4 times in distance from inser- 
tion to lower caudal origin, height 2.0 times in 
base, and inner margin 1.2 times in fin height. 

Caudal fin relatively asymmetrical, with ba- 
sally broad dorsal lobe, and short terminal lobe. 
Length of dorsal margin 2.1 times in precaudal 
length, of preventral caudal margin 2.3 times in 
dorsal caudal margin, and of terminal lobe from 
caudal tip to subterminal notch about 10.3 times 
in dorsal caudal margin. Dorsal caudal margin 
slightly but continuously convex in lateral view, 
preventral margin almost straight dorsally but 
becoming more convex ventrally. Tip of ventral 
caudal lobe broadly angular, lower and upper 
postventral margins slightly convex, notch be- 
tween postventral margins broadly angular, sub- 
terminal notch shallowly concave, subterminal 
margin slightly concave, and terminal margin 
slightly convex. Subterminal margin length 0.6 
times terminal margin length. Ventral lobe of 
caudal fin aplesodic, not supported by hypural 
radials but by ceratotrichia and connective tissue 
only. 

Vertebrae (Fig. 12) examined from beneath 
first dorsal fin (monospondylous precaudal ver- 
tebrae) and at base of caudal fin (diplospondylous 
caudal vertebrae). These found to have ex- 
tremely reduced calcification, both of the pri- 
mary double cone of vertebral centra (which is 
almost entirely formed of uncalcified cartilage 
and connective tissue in M. pelagios), and of 
intermedial areas between basidorsals and bas- 
iventrals. Vertebral centra consist of biconic or 
bioconcave discs of cartilage, separated by broad 
bands of unchondrified notochordal sheath and 
spherical cavities containing notochordal tissue. 
Calcification in monospondylous precaudal cen- 
tra restricted to some irregular calcification on 
lateral centrum body, a layer on ventral part of 
neural canal, a layer on midventral groove on 
underside of centrum, and paired thin zones partly 
bounding intermedial areas between basals, in- 
cluding 2 dorsals, 2 ventrals, and 2 pairs of lat- 
erals. These intermedial calcifications resemble 
radii of other lamnoids, but differ in being only 
partially developed across intermedial areas and 
in not forming discrete longitudinal plates. These 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 105 


Dorsal View 


PT. 


SRI 
OR 


Ventral View 


STF 


Lateral View 


Ficure 13. 


Neurocranium of Megachasma pelagios, in dorsal, ventral, and lateral views. Abbreviations: Dorsal view: AF, 


anterior fontanelle; CR, cranial roof; LR, lateral rostral cartilage; MR, medial rostral cartilage; NA, nasal aperture; OR, opisthotic 
ridge; OT, otic capsule; PF, parietal fossa; PR, preorbital process; PRF, profundus foramen; PT, postorbital process; RN, rostral 
node; SC, supraorbital crest; SF, supraorbital fenestra; SRI, sphenopterotic ridge. Ventral view: LR, MR, and OT as above; AP, 
articular pit; BP, basal plate; ECF, ectethmoid foramen; HF, hyomandibular facet; ICF, internal carotid foramen; INS, internasal 
septum; NC, nasal capsule; SC, suborbital crest; SS, suborbital shelf; STF, stapedial fenestra. Lateral view: O, orbit; ORF, orbital 
fissure; SC, suborbital crest; SCA, sphenopterotic capsule; all others as above. (From drawings by L. J. V. Compagno.) 


intermedial calcified zones interpreted as rep- 
resenting vestigial radii, greatly reduced in Me- 
gachasma but probably well developed in its pre- 
cursors. Basal caudal centra similar to 
monospondylous precaudal centra, except for 
having intermedial calcifications even more re- 
duced to a set of dorsal and ventral pairs only. 
The poorly calcified vertebral centra of Mega- 
chasma recall the septate vertebral columns of 
large species of Somniosus (subgenus Somniosus, 
for S. pacificus and S. microcephalus) and some 
other squaloids (see Compagno 1977), with re- 
duction of form and calcification of centra and 
hypertrophy of notochordal tissue in between 
centra. The lamnoids Mitsukurina and Pseudo- 
carcharias have extremely simple centra with 
double cones and radii reduced to 8 slightly 
branched plates (2 bounding each intermedial 
area), but Megachasma goes far beyond these 


genera in reduction of its centra, in calcification, 
and in intrusion of notochordal tissue. Mitsu- 
kurina and Pseudocarcharias retain normal, close- 
set double cones, despite their simple radi. 
Neurocranium (Fig. 13) dissected on one side 
only, and reconstructed bilaterally. Cranium rel- 
atively large, extremely broad and moderately 
flat; nasobasal length (from base of medial rostral 
cartilage to occipital condyles) about 8.9 percent 
total length and 12.8 percent precaudal length; 
greatest width of cranium across preorbital pro- 
cesses about equal to nasobasal length, and great- 
est height from cranial roof to ventral edges of 
suborbital shelves 0.4 times in nasobasal length 
and greatest cranial width. Rostrum relatively 
short but very broad, length of medial rostral 
cartilage from its base to anterior edge of rostral 
node about 26 percent nasobasal length; width 
across outer bases of lateral rostral cartilages 2.2 


106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


Neurocranium 


sc O 


LR 


Ficure 14. Jaw suspension of Megachasma pelagios, in lateral view, jaws retracted. Abbreviations: H, hyomandibular, MC, 
Meckel’s cartilage; OP, orbital process; PP, palatine process; PQ, palatoquadrate; QP, quadrate process; VC, vertebral column; 
all others as in Figure 12. (From drawings by L. J. V. Compagno.) 


times length of medial rostral cartilage. Entire 
ethmoid region of cranium, including rostrum, 
nasal capsules, and internasal septum, and the 
anterior basal plate molded dorsally around en- 
larged palatine processes of palatoquadrates (Fig. 
14). Lateral rostral cartilages with broad bases 
that cover entire anterior surfaces of nasal cap- 
sules; diagonally compressed from dorsomedial 
to ventrolateral, and extending anteromedially 
as tapering triangular bars to meet rostral node 
separately on either side. Medial rostral cartilage 
and its base on internasal septum deflected up- 
ward over symphysis of palatoquadrates (jaws in 
retracted position), so that base originates at a 
level slightly above lateral rostral cartilages and 
the moderately depressed, narrow, barlike shaft 
arches anterodorsally and anteroventrally to ros- 
tral node. Rostral node a simple, depressed, 
narrow plate, not vertically or horizontally fe- 
nestrated, anteriorly expanded, vertically com- 
pressed, nor with rostral appendices. 

Nasal capsules extraordinarily modified, high- 
ly compressed, platelike, wedge-shaped struc- 
tures with nasal fenestra mainly on their lateral 
faces. Plane of compression of nasal capsules 
congruent with large-based lateral rostral carti- 


lages, together forming a lateral wall to expansion 
cavity enclosing palatine processes. Ectethmoid 
foramen present on dorsomedial surface of each 
nasal capsule. Large subethmoid fossa on ventral 
surface of depressed, laterally expanded inter- 
nasal septum, extending anteriorly beneath ros- 
trum and medially to nasal capsules, and pos- 
terolaterally to merge on either side with large 
orbital process cavity in basal plate. Foramina 
for nasal canals laterally situated in cranial cavity 
(not anterolateral), with canals running antero- 
laterally to nasal capsules. 

Basal plate very broad, width across orbital 
notches about 68 percent nasobasal length, 
broadly arched over rear ends of palatoquadrate 
palatine processes (when retracted) but relatively 
flat posterior to internal carotid foramina. Entire 
ventral surface of suborbital shelves, basal plate, 
and internasal septum padded with thick, soft, 
spongy connective tissue, probably to cushion it 
from palatoquadrates. Basal plate with pair of 
internal carotid foramina located about 59 per- 
cent nasobasal length behind medial rostral car- 
tilage, separated by a convex space with width 
80 percent nasobasal length and 1.1 times in dis- 
tance between internal carotid foramina and sta- 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 107 


pedial fenestrae. Stapedial fenestrae small, width 
about 3 percent nasobasal length, apertures about 
1.6 times closer to internal carotid formina than 
to lateral edges of suborbital shelves. Stapedial 
fenestrae apparently without greatly convoluted 
arteries or a rete mirabile elaborated from effer- 
ent spiracular arteries. Basal plate nearly hori- 
zontal posteriorly, without medial keels. 

Orbits nearly circular in lateral view, with large 
optic nerve foramen slightly dorsal to its center. 
Orbits moderately large, with horizontal diam- 
eters about 43 percent of nasobasal length. Su- 
praorbital crests broad, not fenestrate basally, 
only moderately concave in dorsal view. Preor- 
bital processes not strongly exserted from su- 
praorbital crests, and extending ventrally to pos- 
terior edges of nasal capsules. Small preorbital 
canal fenestra for superficial ophthalmic nerves 
present between broad preorbital process and 
cranial roof on each side; profundus nerve fo- 
ramen just mesial to fenestra. Postorbital pro- 
cesses ventrally produced almost to level of optic 
nerve foramen, bifurcate distally. Foramina of 
orbital wall not examined in detail but including 
foramina for superficial ophthalmic nerve an- 
terior cerebral veins, optic nerve, and large, deep, 
trigeminofacialis chamber or orbital fissure. Sub- 
orbital shelves nearly vertical, large, thick basally 
but distally thin, arcuate, and with sides nearly 
parallel in ventral view. 

Otic capsules large and subquadrate, with 
lengths about 36 percent nasobasal length and 
width about 82 percent nasobasal length. Hyo- 
mandibular facets huge, ventromedially incised, 
and broadly arcuate, extending along entire length 
of otic capsules from otic processes anteriorly to 
partway onto bases of suborbital shelves, but not 
exserted posteriorly from occiput. Hyomandib- 
ular nerve foramina just below opisthotic ridges 
and about midway along their lengths on otic 
capsules. Sphenopterotic ridges arching postero- 
medially in dorsal view, ending posteriorly in a 
bluntly rounded corner. Opisthotic ridges on 
dorsal surface of hyomandibular facets low and 
curved posteroventrally. Occiput flat and not 
exserted rearwards, with glossopharyngeal and 
vagus nerve foramina. 

Jaws (Figs. 5, 14) poorly calcified; length of 
palatoquadrates about 16 percent total length, 
Meckel’s cartilages 18.4 percent total length. Pal- 
atine processes of palatoquadrates articulating at 
symphysis and extending for about % of pala- 


toquadrate length to orbital processes. Meckel’s 
cartilages huge, ventrally arcuate, dorsally nearly 
straight, thick, and compressed, with long pos- 
terior extensions from their mandibular articu- 
lations with palatoquadrates. Meckel’s cartilages 
articulating closely at mandibular symphysis. 

Manipulation of the jaws of the fresh-caught 
Megachasma pelagios suggested that the jaws are 
highly protrusible, but not necessarily as a mech- 
anism to quickly eject them outward to capture 
prey, as in some other lamnoids (most notably 
Mitsukurina), nor to bring the upper teeth to bear 
on prey items, as in Carcharodon carcharias. The 
jaws may be protruded forward and outward to 
expand the mouth aperture and form a hoop-net 
for capturing plankters, though we do not know 
the exact shape of the jaws deployed in this con- 
figuration without photographic documentation 
ofa live M. pelagios feeding. The basking shark 
is able to deploy its much slimmer jaws almost 
in a circle while feeding and has been photo- 
graphed many times with jaws expanded (but not 
protruded); however, the exact shape of the mouth 
opening in a living, feeding basking shark would 
be somewhat difficult to work out from a dead, 
preserved specimen. The jaw structure of M. pe- 
lagios suggests that the jaws move downward, 
anteriorly, and outward at the mouth corners, 
and the distal ends of the hyomandibulae swing 
anterolateroventrally as protrusion occurs. The 
mechanism of jaw protrusion is poorly under- 
stood with the limited dissection possible during 
preparation of this description (the desire to limit 
damage to the specimen prior to making a cast 
of it prohibited a thorough investigation of the 
jaw mechanism and the hyobranchial skeleton 
and musculature), but the large, straplike, diag- 
onal preorbitalis muscles may help to pull the 
jaws forward. 

The jaw musculature was not investigated in 
detail, but sufficient information was collected 
to determine that the jaw muscles are similar to 
those in other lamnoids. Levator palatoquadrati 
muscle simple, originating on sphenopterotic 
ridges of otic capsules and running posteroven- 
trally to insert on quadrate processes of palato- 
quadrates. Adductor mandibulae muscles mod- 
erately large but small and weak compared to the 
huge jaws, and limited anteriorly by mouth cor- 
ners. Levator hyomandibuli muscles broad and 
relatively large. 

The viscera were not examined in detail, ex- 


108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


Ficure 15. 


cept to note that the liver is relatively small 
(though very oily) and that the valvular intestine 
is an elongated ring-valve type rather similar to 
that of other lamnoids (Fig. 15). Each ring has a 
densely fimbriated surface, apparently to in- 
crease the absorptive surface, with a maximum 
thickness of 23-25 mm at the pyloric end of the 
intestine but becoming increasingly thin towards 
the rectum. 

Color. When preserved, dorsal surface of head, 
trunk and tail, dorsal surfaces of pectoral and 
pelvic fins, dorsal fins, center of anal fin, and 
caudal fin dark gray to blue-black; sides of body 
lighter, underside of body and fins light gray, 
except for underside of head and lower jaw which 
are dark gray and mottled with black, undersur- 
face of pectoral fins which have an abruptly black 
anterior margin about an eye diameter wide, and 
underside of pelvic fins with a dusky anterior 
margin. Tips and posterior margins of dorsal sur- 
faces of pectoral and pelvic fins abruptly white, 
as are posterior margins of dorsal and anal fins 
and postventral caudal margins. Tissue of oral 
cavity and tongue blackish in preservative, but 
with oral lining silvery when fresh. 


FEEDING HABITS AND BIOLOGY 


The stomach contents were a thick reddish 
soup abundantly stocked with the euphausiid 
shrimp Thysanopoda pectinata, a species attain- 
ing a median length of 31 mm (Hu 1978). Ac- 
cording to Hu (1978), 7. pectinata off the west 
coast of Oahu (21°15’-20'N, 158°15’-30'W) 
shows a moderate day and night migration pat- 
tern. During the day most are caught between 
350 and 750 m depth, with some ranging up to 
300 m and down to 1100 m, but at night the 


Drawing of the valvular intestine of Megachasma pelagios with the dorsolateral quadrant removed to show the 
ring valves with highly fimbricated edges (anterior to the right). 


bulk are between 150 and 500 m depth, with 
some up to 75 m and down to 525 m. Apparently, 
when captured, Megachasma pelagios would 
have been in the upper depths (165 m) where 
these euphausiids are commonest at night, and 
quite possibly might have been feeding on them 
when it became entangled in the parachute. 

The megamouth shark unites an eclectic com- 
bination of habitus characters that (along with 
its apparent epipelagic habitat and filter-feeding 
habits) suggests an unusual mode of life. Deep- 
water epibenthic and epipelagic sharks often show 
a decrease in specific gravity and increase in hy- 
drostatic support by the enlargement of their ab- 
dominal cavity and liver volume to produce a 
large, oily, hepatic “‘float.”” M. pelagios, in con- 
trast, has reduced specific gravity in the form of 
extremely poor calcification; a soft, almost en- 
tirely hyaline cartilage skeleton; very soft, loose 
skin; and flabby, loose connective tissue and 
muscles. These features, and its soft, rubbery pre- 
caudal fins; lack ofa keel on the caudal peduncle, 
weak precaudal pit; lack of dorsal caudal ripples; 
and highly flexible, asymmetric caudal fin suggest 
that M. pelagios is a slow, weak swimmer. 

It is interesting to compare M. pelagios with 
the other two species of large, filter-feeding sharks: 
the basking shark and the whale shark. The bask- 
ing shark is the only lamnoid filter-feeder besides 
megamouth, but in contrast has many adapta- 
tions for a higher activity level and sustained 
powerful swimming, including a strongly calci- 
fied skeleton, firm muscles, stiff fins, dense skin, 
and tough connective tissue; a huge, oily liver 
and elongated body cavity; a more fusiform body, 
lunate caudal fin, strong caudal keels and pre- 
caudal pits, and huge gill openings. The filter 


a 


TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH—A NEW SHARK 109 


apparatus of Cetorhinus, with its vast gill cavities 
and slender, smooth, streamlined gill-raker den- 
ticles, is clearly adapted for a higher rate of water 
flow than is possible with the smaller gill cavities, 
more restricted internal gill apertures, and less 
streamlined gill-raker papillae of Megachasma. 
The basking shark is a slow but strong swimmer, 
which has often been observed and photo- 
graphed while feeding at or near the surface with 
its mouth distended to form a circular scoop. 
Although its mouth is relatively smaller than that 
of megamouth, the basking shark is probably a 
much more efficient dynamic filterer because of 
its stronger swimming abilities and high-flow fil- 
ter apparatus. The prey of the basking shark is 
far smaller than what is known for megamouth, 
consisting entirely of microscopic crustaceans 
(especially copepods). The basking shark prefers 
cool to cold coastal waters rich in nutrients and 
plankton. 

The whale shark resembles the basking shark 
in its strong swimming adaptations, except that 
it has a less fusiform body, flattened anteriorly; 
a shorter body cavity and much smaller liver; 
and much smaller external gill openings (but larg- 
er than those of megamouth). The filter appa- 
ratus of Rhiniodon differs from that of Mega- 
chasma and Cetorhinus in not being confined to 
the margins of the internal gill openings; instead, 
the gill filter elements of Rhiniodon cross and bar 
these openings. They are compressed, triangular, 
cartilage-cored, connective-tissue-covered, par- 
allel plates that transversely bridge the internal 
gill openings and connect adjacent holobranchs. 
The plates have highly lobulated pharyngeal 
margins that form an interconnected network, or 
dense filter grid, and are divided into paired dor- 
sal and ventral groups of plates or screens over 
each internal gill opening. The dense screens of 
Rhiniodon are obviously efficient filters, but are 
incapable of sustaining a high flow of water 
through them. However, this filter apparatus, 
combined with a broad but very short, transverse 
mouth; very long, broad, low pharynx, and rel- 
atively small gill openings apparently adapts the 
whale shark to a combination of suction feeding 
(as in Ginglymostoma and other orectoloboids) 
and filter-feeding not found in Megachasma and 
Cetorhinus. The bellowslike pharynx and filter 
screens of the whale shark may provide it with 
a more versatile feeding apparatus than in Ceto- 
rhinus (and presumably Megachasma) by allow- 
ing it to suck in and filter out a wide variety of 


prey animals, independent of the shark’s forward 
movement. Although the whale shark can ingest 
small crustaceans, it also eats squid and com- 
monly takes small schooling fishes such as an- 
chovies and sardines, and even small albacore 
and tuna (Bigelow and Schroeder 1948). It is not 
known whether Rhiniodon can filter out crus- 
tacean prey as small as the copepods favored by 
Cetorhinus, but almost certainly the euphausiids 
eaten by Megachasma are in the prey-size range 
of the whale shark, which is a warm-temperature 
to tropical, coastal to oceanic, slow but strong- 
swimming shark, often seen basking or cruising 
at the surface and feeding on schools of fishes. It 
often positions itself vertically beneath a school 
of prey, unlike the horizontal attitude Cetorhinus 
maintains while feeding at the surface. 

The soft, flabby body and fins, low-flow bran- 
chial filter apparatus, and small gill openings sug- 
gest that Megachasma is less active and possibly 
a less efficient filter-feeder than Cetorhinus or 
Rhiniodon. Nevertheless, this species has a spe- 
cialized, presumably efficient mechanism for 
capturing smail oceanic animals in its oversized 
jaws which are enlarged to increase the diameter 
of its “net”? and thickened to provide adequate 
support from its rubbery hyaline cartilage. The 
greatly distensible mouth and pharynx, closely 
packed gill-raker papillae, and large tongue prob- 
ably help to expel water from the pharynx when 
it closes its mouth. Megachasma can be imag- 
ined as slowly swimming through schools of eu- 
phausiid shrimp and possibly other prey with 
jaws widely opened, occasionally closing its 
mouth and contracting its pharynx to expel water 
and concentrate its prey before swallowing it. 

Inspection of the mouth of megamouth 24 hr 
after capture revealed a bright silvery lining 
punctuated by small circular porelike structures. 
At the time it was speculated that these might 
be bioluminescent organs, but we have no evi- 
dence of this. Histological sections of mouthlin- 
ing were made but were problematical because 
of the deteriorated state of the tissue. 

That Megachasma may not be a more active 
filter-feeder such as Cetorhinus or Rhiniodon may 
be related to its tropical deepwater oceanic hab- 
itat, which has a relative paucity of nutrients and 
prey in comparison to the cool coastal surface 
waters favored by Cetorhinus and the tropical 
coastal waters preferred by Rhiniodon. Various 
mesopelagic teleosts have reduced skeletal and 
other tissues as adaptations to a nutrient-poor 


110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8 


environment, and Megachasma may be \imilar 
ly limited to a reduced level of tissue develop- 
ment and hence a low activity léVel for a filter- 
feeding shark, far less than is possible in the 
habitats frequented by Cetorhinus and Rhinio- 
don. 


Two distinctive scars, one on the throat, 
another behind the mght pectoral fin, suggest that 
megamouth may be the only known selachian 
victim of Jsistius brasiliensis, the “‘cookie-cut- 
ter” shark, that is believed responsible for similar 
marks found on tuna, porpoise, and billfish caught 
in Hawaiian waters (Jones 1971). The soft skin 
and midwater habitat of megamouth may make 
it vulnerable to /sistius attacks. 

Megachasma pelagios, itself the representative 
of a new family of sharks, is the host of a new 
family of tapeworms Mixodigmatidae (order 
Trypanorhynchida), described by Dailey and 
Vogelbein (1982) for the new genus and species 
Mixodigma leptaleum. These parasitic tape- 
worms from the valvular intestine presented 
taxonomic problems over placement in existing 
trypanorhynch families comparable to the difh- 
culties encountered in attempting to place Me- 
gachasma pelagios in an existing lamnoid shark 
family. 


ACKNOWLEDGMENTS 

Many people contributed to the study of this 
interesting animal. We acknowledge Lt. Linda 
Hubble, John Hobbs, and Rick Kahakini for their 
recovery of the animal and their recognition of 
the find; Hawaiian Tuna Packers, National Ma- 
rine Fisheries Services Honolulu dockside staff, 
Phillip Motta, Bruce Carlson, Captain Gary Naf- 
tel and the crew of the R.V. Easy Rider, Mathew 
James, Les Matsuura, Marge Awai, and Arnold 
Suzumoto for curatorial assistance; Dr. Arthur 
Popper, Dr. James Margolis, and Karen Margolis 
for preparation of scanning electron micro- 
graphs; Dr. James Kendall for histological prep- 
arations of epithelial tissues; Drs. Thomas Clarke 
and K. Gopalakrishnan for identification of 
stomach contents; Drs. Carl L. Hubbs, Richard 
Rosenblatt, John McCosker, Bruce Welton, 
Murray Dailey, Scott Johnson, Tyson Roberts, 
and John Randall, and Mr. Richard Ellis, for 
discussions relating to nomenclature and the sci- 
entific importance of the animal; Allan Hart, Re- 
becca Brown, and Mary Morioka for scientific 
illustrations; Ruth Naftel for organizational sup- 


port; and Pam Miike for typing the manuscript. 
Particular thanks go to Richard Ellis and John 
McCosker for preparation of a preliminary 
manuscript which was of great help in the pro- 
duction of this final paper. 


LITERATURE CITED 


Anonymous 1. 1976. A novel variation of jaws. Science 
News 110(25 and 26):385-416. 

ANonyMous 2. 1976. The big one that got away. Smithsonian 
7(12):12. 

BiGELow, Henry B., AND W. C. SCHROEDER. 1948. Sharks. 
Fishes of the western North Atlantic. Memoir 1, Sears Foun- 
dation for Marine Research, Pt. 1, vol. 1:59-576. 

Criark, EuGente. 1981. Sharks: magnificent and misunder- 
stood. National Geographic 160(February): 138-187. 

CompaGno, L. J. V. 1970. Systematics of the genus Hemi- 
triakis (Selachii: Cafcharinidae), and related genera. Pro- 
ceedings of the California Academy of Sciences, ser. 4, 38(4): 
63-98, 8 figs. 

1973a. Interrelationships of living elasmobranchs. 

Pages 15-61 in Interrelationships of fishes. P. H. Green- 

wood, R. S. Miles, and C. Patterson, editors. Zoological 

Journal of the Linnean Society of London 53(suppl. 11). 

1973b. Gogolia filewoodi, a new genus and species 

of shark from New Guinea (Carcharhiniformes: Triakidae), 
with a redefinition of the family Triakidae and a key to 
triakid genera. Proceedings of the California Academy of 

Sciences, ser. 4, 39(19):383-410, figs. 1-7, table 1. 

1977. Phyletic relationships of living sharks and 

rays. American Zoologist 17(2):303-322, 15 figs. 

1979. Carcharhinoid sharks: morphology, system- 

atics, and phylogeny. Ph.D. Thesis, Stanford University, 

932 p. 


1981. Legend versus reality: the Jaws image and 
shark diversity. Oceanus 24(4)(Winter 1981/82):3-16. 

DaiLey, Murray D., AND WOLFGANG VOGELBEIN. 1982. 
Mixodigmatidae, a new family of cestode (Trypanorhyncha) 
from a deep sea, planktivorous shark. Journal of Parasitology 
68(1):145-149, 8 figs. 

Dunrorp, B. 1976. Huge shark may be new species. Ho- 
nolulu Star-Bulletin 65(322), p. 1, 2 pictures. 

FAUGHNAN, Victor. 1980. National shark-o-pedia. Under- 
sea Resources, Ltd., Honolulu, Hawaii. 

Hu, Vernon J. H. 1978. Relationships between vertical mi- 
gration and diet in four species of euphausids. Limnology 
and Oceanography 23(2):296-306, 5 figs. 

Jones, E.C. 1971. Isistius brasiliensis, a squaloid shark, the 
probable cause of crater wounds on fishes and cetaceans. 
National Marine Fisheries Service Fishery Bulletin 69(4): 
791-798. 

Puiues, F. J., B. WELTON, AND J. WELTON. 1976. Paleon- 
tologic studies of the Middle Tertiary Skooner Gulch and 
Gallaway Formations at Point Arena, California. Pages 137- 
154 in Neogene symposium; Society of Economic Paleon- 
tologists and Mineralologists Pacific Section, Annual Meet- 
ing, April 1976. 

TayLor, LEIGHTON R. 1977. Megamouth, a new family of 
shark. Oceans Magazine 10:46-47, 5 pls. 

TINKER, SPENCER W. 1978. Fishes of Hawaii. Hawaiian Ser- 
vice, Inc., Honolulu. 532 p. 


PROCEEDINGS 
OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


Vol. 43, No. 9, pp. 111-121, 5 figs., 8 tables. 


JAN 27 1984 


ical Laboratory | 
LIBRARY 


January 17, 1984 


—_— 


THE CRANIAL EEN va OF GALAPAGOS TORTOISES 


Wogds Hole, Mass. 


roe se we 


Charles R. Crumly' 


Department of Zoology and Physiology, Rutgers— The State University, 
Newark, New Jersey 07102 


AsstTRAcT: Saddlebacked tortoises have smaller and slightly broader skulls than non-saddlebacked tortois- 
es. Unlike the two types of shells, the skulls of saddlebacked and non-saddlebacked tortoises are difficult to 
distinguish, even as large adults. Factor analysis, although suggestive of different growth trends, does not 
delineate Geochelone ephippium, G. guntheri, and/or G. nigrita. Discriminant function analysis easily distin- 
guishes these species. Geochelone ephippium is discriminated from the other two species on the basis of 
overall size and G. nigrita has a smaller exposed basisphenoid than G. guntheri. Species represented by small 
samples were compared to Geochelone guntheri, G. ephippium, and G. nigrita by means of a discriminant 
function analysis classification procedure. The results suggest that skull variation does not parallel shell 


variation. 


INTRODUCTION 


The classification of Galapagos tortoises (ge- 
nus Geochelone) has changed over the years, de- 
pending on the prevalent philosophy pertaining 
to closely related forms. These insular tortoises 
have been considered different species (Van Den- 
burgh 1914) or different subspecies (Wermuth 
and Mertens 1961, 1977; Crumly 1980, 1982; 
MacFarland et al. 1974a, b). Perhaps the best a 
priori taxonomic strategy was employed by Fritts 
(in press), who considered each geographically 
isolated population a separate entity until more 
detailed analysis could be completed. These dif- 
ferent philosophies, compounded by the fre- 
quent lack of accurate locality data, are reflected 
in the confusing nomenclatural history of Ga- 
lapagos tortoises (Table 1). 


' Present address: Division of Reptiles and Amphibians, De- 
partment of Vertebrate Zoology, National Museum of Natural 
History, Smithsonian Institution, Washington, D.C. 20560. 


Although the nomenclatural status of these 
various populations remains changeable, it is clear 
that all Galapagos tortoises are more closely re- 
lated to each other than to other tortoises. This 
interpretation is supported by morphologic anal- 
yses (Crumly 1980, 1982; Fritts in press) and 
electrophoretic studies (Marlow and Patton 
1981). 

Despite their close relationships, Galapagos 
tortoises exhibit great structural diversity. The 
shells best reflect this diversity and are of two 
basic types: domed, like those of most other tor- 
toise species; and saddlebacked, resembling an 
ancient Moroccan saddle. The saddlebacked shell 
type seems derived from the domed type, but 
Fritts (in press) has noticed subtle differences be- 
tween saddlebacked forms that suggest this mor- 
phology evolved more than once. Marlow and 
Patton (1981) corroborate Fritts’s suggestion. 
Furthermore, the saddlebacked shell, long con- 
sidered unique to certain Galapagos tortoises, 
appears independently in Geochelone vosmaeri 


(111 


112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9 


Fernandina 
Ph° 


Isabela 


» % 


Saito Gru 
» San Cristobal 


santa Fe 


Archipielago de Colon 


Ficure 1. 


Espanola 
Santa Mari => 


Present distribution of Geochelone species in the Galapagos Islands. A = abingdoni, B = becki, C = chathamensis, 


D = darwini, El = elephantopus, Ep = ephippium, G = guntheri, H = hoodensis, M = microphyes, N = nigrita, Ph = phantas- 
ticus, Va = vandenburghi, Vi = vicina, W = wallacei, ? = unnamed form, | = domed, 2 = intermediate, 3 = saddlebacked, * = 


species probably invalid (from MacFarland et al. 1974a). 


of Rodrigues Island in the Indian Ocean (Arnold 
1979). 

Fritts (in press) showed that island (or volcano) 
topography is a reliable predictor of shell shape 
in Galapagos tortoises. Based on captive breed- 
ing data, he also suggested that shell differences 
are genetically determined. The topographic, cli- 
matic, and morphologic information combined 
in Fritts’s model suggests that these tortoises pos- 
sess great adaptive plasticity. This plasticity, re- 
flected in differences in shell shape, has not been 
documented for other anatomical regions. 

The purpose of this study is to determine 
whether variation in cranial morphometry par- 
allels variation in the two shell types, first pro- 
posed by Van Denburgh (1914) and corroborated 
by Fritts (in press). 


MATERIALS AND METHODS 


Sixteen measurements (Fig. 2, Table 2) were 
recorded from over 100 skulls, listed below. (The 
museum acronyms used are those recommended 
by Duellman et al. 1978.) 


G. abingdonii: CAS 8112; USNM 29269. 

G. becki: CAS 8120. 

G. chathamensis: CAS 8127, 8128, 8130, 8131, 
8133; USNM 29255. 

G. darwini: CAS 8106, 8108, 39601. 

G. ephippium: AMNH 93383; CAS 8299, 8313, 
8358, 8377-8380; MCZ 11068; USNM 29309, 
29251. 

G. guntheri: CAS 8225, 8267, 8413, 8406, 8401, 
8399, 8400, 8396, 8415, 8256, 8408, 8405, 
8199, 8194, 8210. 


PS) 


CRUMLY: TORTOISE SKULLS 


‘epiqey UO A[[eINJeU PILINIIO JOADU A[QeGOId SIS10}10} ISNLI9q PI[BAUI SI 129D//DM VY] UMOYS A[BUIDUTAUOD DARY SIOYING SOY] y% 
*S9SIOLIO] SOBedEIeH JO AWOUOXE] PUP UOMPLOYISSE]D IY} SUISTADI JO SS9d01d JY) UI SI (Ssa/d U7) SN 


(eueo1O[4) 


sIsug0spdp] D3 sisuavospdp]p3 sisuaosvdnjps DAS 1U sndojunydaja sisuaosndnjp3s uiniddiyda — payaeqo|ppes sajiey) Ble eUeS 
poweu jou 1nq 
1a]40d 1a]40d DIMS1U DASIU 1aj4od 1a]40d dATIOUTISIP powoqd a[qesnejopuy Zn1I. blues 
DILUBIU 10 (o3enUuUes) 
1UIMADp 1UIMADP 1UIMADP DLS 1U 1UIMADP 1UIMADP sndojunydaja d}VIPIUIIIU] sower JOpeales ues 
powieu jou 1nq 
SISUIUUDYIDYI SISUIUDYJOYI SISUIWUDYJDYI sadydosou SISUIUDY JDYI SISUIUIDY JOY dATOUTISIP 9] PIPIULII]U] weyley) [eqolsliy ues 
129D]]DM 4% 129D]]DM 19ID]]0M DAS IU 1090]]DM 19ID]]OM d1PIPIUI9}U] SIAJOf epiqey 
winiddiyda uimiddiyda wuiniddiyda sndojunydaja uiniddiyda uiniddiyda poyorqo[ppes uvounqd uoZzuUld 
1uopsulgo 1uopsuiqo nuopsuigo sndojunydaja 1uopsulgo muopsuiqo Nuopsulgd —- paydeqoa|Ppes uopsuIqy PUI 
poeweu 10U 1nqG 
1YSANGUIPUDA 143ANGQUIPUDA sndojunydaja dATIOUTISIP powoqd UW Ag[MoD 
(QA0Z vuen3y) 
DUIIIA DUIJIA sndojunydaja sadydosoiut DUIIIA DUIJIA DUIIIA dLIPIWII}U] dPIeUIDg|V “S 
(A [EWR TILA) 
1dYJUns May]uns sndojunydaja SaAYdOsIIU May Uns 1ayjUuns 91 RIPSW191U] apleWsg|y “A'S 
sady dos d}VIPIULII}U] 
sadydosdiu sad ydosoud sndojupydaja —-% sadydosrvud sadydosonu sadydosriu dA0Z snse] 
(Agjaysiog ode)) 
9[FeWoqg|y “N 
1400q 1499q sndojunydaja sadydosonu 1492q 1499q poyoeqgo|ppes op1eWdq|y BOQges| 
pouspjunyd pouspjunyd poijspjunyd paijspjunyd poyoeqo|ppes ysno1oqien euIpUueUIO-Y 
sISUapooy sisuapooy SISUBPOOY sadydosoiu SISUapooy SISUaPOOY Sadydosniut — paydeqa[Ppes pooyH pjouedsq 
6L61 pL6l |e Ie LL61 1961 L161 bI6l LO6I C161 £061 LL8I SL8I odA} ysi[3uq ystueds 
prey, purer sudLIO UBULIED) ysinquoqd ue, ‘TO61 ‘1061 JayUuny sovdeie) ieee ears 
pue yINULIA Plryosy1OYy eae 
4 SASIOLYO | SODVdVIV) OL GAlIddy SAWVN SIdIOadg “| ATAV 


114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9 


FiGuRE 2. 


G. hoodensis: CAS 8121, 8122. 

G. microphyes: 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. vicina: CAS 8179, 8193, 8177; 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, 8411, 
8409, 8402, 8377, 8407, 8410, 8403, 8414, 
8397, 8412, 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, 2933829305, 29254, 
DOZS2Z NONI OS S93 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 


Measurements taken from Galapagos tortoise skulls (see Table 2 for explanations of abbreviations). 


(Van Denburgh 1914) 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.) 


Variable— 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 of jugal 
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 premaxillae 
PC—Length of sagittal contact of prefrontals 


CRUMLY: TORTOISE SKULLS 


TABLE 3. 


115 


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. guntheri 


(N = 9) (N = 15) 
Variable X SD x SD 
B 96.7 11.4 128.0 21.4 
WAT UBS) SN) 106.6 19.9 
WO 35) Sel 35.4 VES 
HN 12ES 2.0 18.6 3.4 
WN 17.0 DED. Pal 4.4 
LB I3e3 3 18.7 4.2 
WB 14.6 Dee 19.1 4.3 
WZ 9.3 3.6 14.3 4.4 
WP 7.0 2D 9.5 23) 
WS 7.3 2:3 12.0 3.0 
DPV 322 0.8 4.2 0.8 
bP 14.1 Za 21.0 5.6 
WFS 10.0 2.0 16.0 6.1 
PW 19.2 1.8 BS) 3). 
APW 10.5 1.8 [522 2.4 
IO 8.6 Nes) 10.6 4.0 


G. nigrita G. vicina G. chathamensis 
(N = 18) (N = 4) (N = 6) 
xX SD x SD xX SD 
RS 39:2 109.0 49.2 98.1 27,3 
98.4 31.8 86.0 38.9 80.4 25.0 
37.0 13.3 28.4 12.6 28.4 Teil 
18.6 6.0 16.1 U3) 13.9 4.1 
235 Ui D3) 9.2 18.5 4.7 
14.7 4.3 18.1 8.9 14.7 5.6 
lea 4.6 15.8 Ted 13.8 . 3.4 
13.3 4.7 12.6 6.8 10.1 3.5 
9.0 3.5) 8.8 4.5 Wed) 3.0 
9:5 4.0 9.6 Dal 7.9 33531 
4.2 IES 3-7) 2.4 3p 1.0 
18.1 6.5 14.8 6.6 1522 By) 
12.8 6.1 8.9 6.3 12.5 6.2 
26.1 8.4 21.9 8.6 19.0 4.0 
14.1 4.9 11.8 8.0 10.5 3.0 
13.3 5.4 8.5 3.9 8.2 3.9 


populations were hampered by incomplete lo- 
cality data; 50 of 116 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 Fritts (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 15 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- 
wint [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 
weil 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 IT ORTOISES REPRESENTED BY SPECIMENS OF FIVE SPECIES 
AND NON-SADDLEBACKED TORTOISES REPRESENTED BY 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 xX SD xX SD 

B 98.9 15.9 116.5 Byal 
WAT 75.8 13.0 96.0 30.4 
WO 26.4 4.6 32.9 We? 
HN 1322 2.9 17.1 5.6 
WN 17.5 2.4 22.6 ied: 
LB 13.8 32 16.0 Ssi/ 
WB 14.5 2.0 Ni! 55) 
WZ 9.5 3.4 13.1 5:3 
WP el 29) 8.8 3.5 
WS 7-3 DES 9.9 3.9 
DPV 2.9 (le 3.9 1.4 
ee 14.4 Dei 18.4 6.2 
WFS 10.1 1.8 1337 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 ee 


116 


TABLE 5. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9 


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. 


Variable B WAT WO HN WN LB WB WZ WP WS DPV LP WFS PW APW 
WAT 98 

WO 94 94 

HN 96 94 93 

WN Oy 96 93 9)5) 

LB .88 .86 sf) .84 .88 

WB Roy) wy 87 .88 BOD 87 

WZ 84 .86 .86 81 87 78 .84 

WP 87 .86 .88 83 87 19 .80 .88 

WS Sil 2 87 .88 a9) .86 89 84 .84 

DPV 69 .69 69 64 P .60 .68 .64 .66 .66 

LP AO) 94 91 8) 94 85 Sil .86 84 I .63 

WFS 90 89 87 86 .89 .84 85 718 aH 87 5) 91 

PW a a9 90 90 aH 718 .88 85 .83 .83 .69 90 .82 

APW $3} 93 .89 91 94 .84 89 82 .83 .89 .68 86 82 85 

RE .60 56 55) .63 .60 38 .49 .43 .48 48 .49 55/5) 50 54 SY 


through the Office of Computer Services (OCS) 
at the Smithsonian Institution. Simple descrip- 
tive statistics, linear regression, factor analysis, 
and stepwise discriminant analyses were used to 
summarize observed cranial variation. 


RESULTS 


Geochelone ephippium appears to have the 
smallest skull and G. guntheri the largest skull 
of Galapagos tortoises (Table 3), but when max- 
imum basicranial lengths (mean plus two stan- 
dard deviations) are compared, G. nigrita ap- 
pears to possess the largest skull (B,,,,. = 171 mm 
for G. guntheri, 200 mm for G. nigrita). The 
efficacy of this procedure is in some doubt since 
the B,,,,. 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. ephippium, 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 11.74, 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 16 variables correlate 
most highly with factor one. Varimax rotation 


ne 


CRUMLY: TORTOISE SKULLS 


Ww 
e 


Facior 3 


FiGurR_E 3. 


® 


A plot of factor scores for factors two and three. Geochelone nigrita (solid circles), G. guntheri (cross-hatched 


circles) and G. ephippium (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. nigrita), the points for G. ephippium and G. guntheri 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 1n cra- 


TasBLeE 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 l 2 3 


Eigenvalue 13.19 0.47 0.28 
% Variation 94.6 3.4 2.0 
Summarization of WFS 0.82 WP 0.63 PG OMS 
Factor Matrix LB 0.80 DPV0.60 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 1n 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. guntheri. 

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 1s good 


118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9 


Ficure 4. A plot of the discriminant scores derived from 
an analysis that included Geochelone guntheri, G. nigrita, 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. StTAtTistics 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 1) and low Wilks’s lambdas (near 0) 
indicate that factors are good discriminators. 


Discriminant 


function l 2 

Eigenvalue 7.06 5.82 
% Variation 54.8 45.2 
Canonical correlation 0.94 0.92 
Wilks’s lambda 0.02 (P= 0.003) 0.15 (P=0.02) 
Pooled within groups ISUN| =O.27 13} 0), 

correlations bet. ILI (0), 240) WB —0.17 

canonical WN —0.19 

discr. fnets. & WAT-—0.19 

discr. variables APW—0.18 

WO —0.18 

Groups delineated G. ephippium G. nigrita 

from others from G. 


guntheri 


] 
i 


Ficure 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 1s 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 are available upon 
request. 

The third discriminant analysis compared 


CRUMLY: TORTOISE SKULLS 


TABLE 8. 


CLASSIFICATION RESULTS OF A DISCRIMINANT ANALYSIS CLASSIFICATION PROCEDURE. Individual specimens were 


classified to one of three species: G. ephippium (a saddlebacked species), G. guntheri (an intermediate form) or G. nigrita (a 
domed form). Asterisk indicates invalid taxon (fide MacFarland et al. 1974a). 


Shell type Trivial name Sex 
domed vandenburghi 2 
intermediate chathamensis ? 
intermediate chathamensis ? 
intermediate chathamensis 2 
intermediate chathamensis male 
intermediate chathamensis 2 
intermediate chathamensis 2 
intermediate darwini female 
intermediate darwini male 
intermediate microphyes male 
intermediate vicina male 
intermediate vicina female 
intermediate vicina ? 
intermediate vicina male 
intermediate wallacei* male 
saddlebacked abingdonii ? 
saddlebacked abingdonii male 
saddlebacked becki female 
saddlebacked hoodensis male 
saddlebacked hoodensis female 
saddlebacked phantastica male 


small samples of tortoise species to large sam- 
ples. Small samples were classified by the dis- 
criminant function 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 


Mus. no. Classified as 
CAS 8141 ephippium 
CAS 8133 ephippium 
CAS 8131 ephippium 
USNM 29255 ephippium 
CAS 8127 ephippium 
CAS 8130 ephippium 
CAS 8128 ephippium 
CAS 8106 ephippium 
CAS 8108 guntheri — 
CAS 8158 guntheri 
CAS 8179 ephippium 
CAS 8193 ephippium 
USNM 129247 ephippium 
CAS 8177 guntheri 
CAS 8134 guntheri 
USNM 29269 guntheri 
CAS 8112 guntheri 
CAS 8120 ephippium 
CAS 8121 ephippium 
CAS 8122 ephippium 
CAS 8101 guntheri 


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 individual’s 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. 


120 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 1S 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 guntheri 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 1n- 
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 1n- 
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. 


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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. 10:119. 

1915. The gigantic land tortoises of the Galapagos 
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Tuorpe, R.S. 1976. Biometrical analysis of geographic vari- 
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VAN DenBURGH, J. 1907. Expedition of the California Acad- 
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1978. Mu- 


CRUMLY: TORTOISE SKULLS 


I: Preliminary descriptions of four new races of gigantic land 
tortoises from the Galapagos Islands. Proc. Calif. Acad. Sci. 
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1914. Expedition of the California Academy of Sci- 
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gigantic land tortoises of the Galapagos Archipelago. Proc. 
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WermutH, H., AND R. MertTeENS. 1961. Schildkroten, kro- 


kodile, und bruckenechsen. Gustav Fischer Verlag, Jena. 
422 p. 


121 


1977. Liste der rezenten Amphibien un Reptilien. 
Testudines, Crocodylia, Rhyncocephalia. Das Tierrich 190: 
1-174. 

ZANGERL, R. 1969. The turtle shell. Pages 31 1-339 in Gans. 
C., A. d@A. Bellaris, and T. S. Parsons. Biology of the Rep- 
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, AND R. G. JoHNson. 1957. The nature of shield 


abnormalities in the turtle shell. Fieldiana: Geol. 10:341— 
62. 


CALIFORNIA ACADEMY OF SCIENCES 
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San Francisco, California 94118 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OESG | 


Vol. 43, No. 10, pp. 123-140, 8 figs., 1 table 


THE STATUS OF 


LIBRARY 
JAN 27 1994 


January 17, 1984 


(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 figulus, and figulus koma Tsuneki, 1956 is newly synonymized with minus. Neotypes are 
designated for Sphex fuliginosus Scopoli, 1763, and Trypoxylon majus Kohl, 1883, both synonyms of 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 (1945, 
1964a) and Richards (1980) called them vari- 
eties, Bliithgen (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 /figu/us 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 /i- 
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] 


124 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 Liége, Belgium, by A. Jacob-Remacle, 
and examined by me, all are minus (7 2, 5 6 reared 
in 1976, 44 9, 35 6 reared in 1981). This result 
contradicts Valkeila’s conclusions: If the three 
phena really are variants of one species, then one 
would expect some variation of phena in the 
Liége trap nest material. 

3. Geographic distribution. All three species 
are largely sympatric, but only figu/us and me- 
dium have been found in Great Britain, only 
figulus and minus in the Iberian Peninsula, and 
only figu/us 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 Liége, 
Belgium, in 1980 and 1981 (and examined by 
me), 4 are medium and the remainder are minus 
(202 2, 223 4). Not a single figu/us 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 /i- 
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 fiir Pflanzenschutzforschung der Akademie der 
Landwirtschaftswissenschaften der DDR, Zweigstelle Ebers- 
walde, Abteilung Taxonomie der Insekten (formerly 
Deutsches Entomologisches Institut), Eberswalde-Finow (Dr. 
J. Oehlke) 

FIS: Forschungsinstitut Senckenberg, Frankfurt am Main, Fed- 
eral Republic of Germany (Dr. J.-P. Kopelke) 

FJS: Sefior Francisco J. Suarez, Almeria, Spain 

FSAG: Faculté 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) 

IEE: Instituto Espafiol 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 TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 12 


TABLE |. 


Character 


. Thoracic pilosity 

Anteroventral mesothoracic process 
. Free margin of female clypeus 

. Female hindcoxal pit 

Sete of hindcoxal pit 

. Male apical flagellomere 

. Gonoforceps process 


absent 


absent 


MHNG: Muséum d’ Histoire Naturelle de Genéve, Switzerland 
(Dr. Cl. 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) 

NHMYV: Naturhistorisches Museum, Vienna, Austria (Dr. M. 
Fischer) 

NRS: Naturhistoriska Riksmuseet, Stockholm V, Sweden (Mr. 
S. Erlandsson) 

RMNH: Rijksmuseum van Natuurlijke Historie, Leiden, The 
Netherlands, including collections of J. P. van Lith and P. 
M. F. Verhoeff (Dr. K. van Achterberg) 

SFG: Dr. Severiano Fernandez Gayubo, Departamento de 
Zoologia, Universidad de Salamanca, Spain 

SMT: Staatliches Museum fiir Tierkunde, Dresden, German 
Democratic Republic (Dr. Regine Eck) 

TMB: Természettudomanyi Muzeum, Budapest, Hungary (Dr. 
J. Papp) 

TN: Mr. Toshiaku Nambu, Yorii-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, Alma Ata, USSR 

WJP: Wojciech J. Pulawski, San Francisco, California 

WSU: Washington State University, Department of Ento- 
mology, Pullman, Washington 

ZMB: Museum fiir 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 Burlington House, London, was possible ow- 


Ancestral 


shorter (as in medium) 


straight or sinuate 
circular 

evenly distributed 
shorter (as in medium) 


Wn 


ANCESTRAL AND DERIVED CHARACTER STATES OF THREE SPECIES IN THE GENUS 7 RYPOXYLON. 


Derived 


longer (as in figu/us) 
present 

concave 

oblong 

channel-like structure 
longer (as in figulus) 
present 


ing to the kind assistance of Michael C. Day of 
the British Museum (Natural History). North 
American specimens of figu/us 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 7ry- 
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. |. Tree B is the 
most probable, since no single derived character 
state 1s 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 figu/us 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 


A 


FiGure 1. 


B Cc 


Hypothetical phylogenetic trees of Trypoxylon figulus (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 midiength (Fig: 7E, F)2.. =. 
Er Be eee RS 3 medium de Beaumont 
— Female: clypeal free margin (Fig. 2A, 5A) 
sinuate to nearly straight, except concave 
in occasional western European figu/us 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 (hice JE) ee ee yD, 


2. Mesothorax without anteroventral 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 minus without meso- 
thoracic process. I cannot find characters for distinguishing 
them. 


2B-D) or (some specimens) circular, eva- 
nescent in Japanese specimens... 
Pee ae epee) Why Fs ee Se 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 
(Ese) DS Gi) a minus (de Beaumont) 


Trypoxylon figulus (Linnaeus) 


Sphex figulus Linnaeus, 1758:570. ! Holotype: 2, Sweden, 
Uppsala (Linnean Society, London). — Day, 1979:62. — In 
Trypoxylon: LATREILLE, 1802:79; TsuNEKI, 1981:15 (rede- 
scription, geographic variation). — In Apius: JURINE, 1807: 
142. 

Sphex fuliginosus Scopo.t, 1763:292 (as fuliginosa, incorrect 
original spelling). Holotype or syntypes: Carniolia (formerly 
in Austria, since 1919 part of Italy and Yugoslavia), lost, 
see ROGENHOFER UND DALLA Torre, 1882:599. ! Neotype: 
2, Austria: “Carinthia, Ostkarawanken, Ebriach, 580-750 
m, 21-29. VII.1964, G. van Rossem,” present designation 
(CAS). — As probable synonym of figu/us: VANDER LINDEN, 
1829:42. — As synonym of figu/us: subsequent authors. 

Trypoxylon figulus var. majus KOHL, 1883:657, 2, 6 (as major, 
incorrect original spelling). Holotype or syntypes: 2, Swit- 
zerland: no specific locality (originally NHMV, Vienna, now 
lost). ! Neotype: 2, Switzerland, “P. 3 VIII 84” and “Cn. 
Tournier”’ (=Peney near Geneva, collection Tournier), pres- 
ent designation (MHNG). New synonym. — pE BEAUMONT, 
1945:477 (var. major); BLUTHGEN, 1951:234 (var. major); 
DE BEAUMONT, 1958:206 (forma major), 1959:30 (same); 
Wo tr, 1959:15, 16 (figulus major); VALKEILA, 1961:244 (var. 
major); DE BEAUMONT, 1964a:290, 1964b:84 (forma major), 
1965:56 (same), 1967:338 (same); BOHART AND MENKE, 1976: 
346 (ssp. major); LomHo.LpT, 1976:267 (figulus major); 
RicHArps, 1980:45 (var. major). 

Trypoxylon apicale W. Fox, 1891:142, 2 (as apicalis, incorrect 


PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 127 


original spelling). Lectotype: 2, Canada (ANSP, Philadel- 
phia), designated by Cresson, 1928:52. — SANDHOUSE, 1940: 
156 (apicale). Synonymized by Pate, 1943:16. 

Trypoxylon figulus barbarum DE BEAUMONT, 1957: 2, 6. Ho- 
lotype: ¢, Morocco: Marrakech (Mus. Zool. Lausanne). New 
synonym. — BOHART AND MenkE, 1976:346. 

Trypoxylon figulus yezo TsuNEK1, 1956:29, 2, 6. Holotype: 2, 
Japan: Hokkaido [=Yezo]: Jozankei (K. Tsuneki collection, 
Mishima). New synonym. — BoHART AND MENKE, 1976: 
346; TsUNEKI, 1981:21 (summary of faunistic data). 

Trypoxylon fieuzeti GINER Mari, 1959:389, 6. ! Holotype: é, 
Morocco: Fez (IEE, Madrid). Synonymized with figulus bar- 
barum by SuAREz, in GINER Mari, 1959:400. 


COMMENTS ON New Synonyms.—T7. figulus 
barbarum was based mainly on the elongate male 
flagellomere XI. Because flagellomere XI varies 
in length (see Geographic Variation below) this 
subspecies 1s 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 NeEotypes.—The identity of 
Sphex fuliginosus 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 7rypoxylon 
figulus as a neotype of Sphex fuliginosus I have 
followed the traditional interpretation of the last 
name. 

The original material of 7rypoxylon 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. 

D1aGnosis.— Most specimens of figu/us 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 Wachseldornmoos, Switzerland); it is 2.7-— 
3.2 times its basal diameter in Spanish individ- 
uals, and 3.3—3.6 times in Moroccan individuals. 

Lire History.—Many specimens of figulus 
(voucher specimens examined by me) were reared 
from nests established in wood (Wolf, 1959). Six 
females and 17 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- 
ern Canada and northeastern USA). 


Recorps (Old World). — Algeria (1 4): El Harrach (as Maison 
Carrée, apical flagellomeres missing, BMNH). 

Austria: (102 2, 50 4, 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). 
Niederésterreich: Bisamberg near Vienna (NHMV, CU), Buck- 
lige Welt S Vienna, Briihl, Dornbach (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- 


128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 


~ 


i vet fh 
i yh, Ni 


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, NHMYV), Molln, Miihlviertel (JG), 
Neumarkt (JG), Oberwallsee bei Miillachen (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), Tragéss-Oberort (DBB), Wein- 
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 2, 63 6, 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, 


Wanfercée. Liege: Acosse, Aubel, Barchon, Ben-Ahin, Beyne: 
ca 15 km SE Liége (BMNH), Beyne-Heusay, Cerexhe, Char- 
neux, Chevron, Clermont-sur-Berwinne, Fléron, Forét de 
Grunhault, Francorchamps, Henri-Chapelle, Hombourg, Ju- 
pille, La Calamine, La Reid, Lontzen, Montzen, Pepinster, 
Queue-du-Bois, Romsee, Spa, Xhendelesse, Welkenraedt. 
Limburg: Berg prés de Tongres, Bocholt, Godsheide, Tongres. 
Luxembourg: Amonines, Hotton, Les Epioux, Lomprez, Ozo, 
Saint-Médard, Sampont, Smuid, Waharday, Wibrin. Namur: 
Aische, Alle (RMNH), Andenne, Baillonville, Belgrade, Biévre, 
Branchon, Champion, Eghezée, Ernage, Feschaux, Gembloux, 
Gesves, Grand Leez, Ham-sur-Sambre, Lonzée, Mount-Gau- 
thier, Saint-Aubin, Saint-Gérard, Saint-Marc, Sorinnes, Sau- 
veniére, Sombreffe, Winenne. 
Bulgaria (1 4): Rila Mts. (DEI). 


129 


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PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 


130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 


FiGure 4. 


Czechoslovakia (1 2, 3 4): Jihomoraysky Kraj: Vranov W 
Znojmo (as Frain, NHMV). Vapadoslovensky Kraj: Sturovo 
(SMT). 

Denmark (41 ¢, 33 6, ZMK if not indicated otherwise): Al- 
borg: Vegger. Bornholm: Bastemose. Frederiksborg: Asserbo, 
Hilleré6d, Hundested, Jaegerspris Nordskov (tip of Horn- 
sherred peninsula), Sorte Mose near Farum, Store Karlsminde 
near Lynes, Tisvilde Hegn (FSAG, ZMK). Holbek: Kongsore. 
Maribo: Maribo. Ksbenhayn: Holte. Odense: belo. Randers: 
Glatved Strand on Djursland Peninsula, Mols Bjerge. Ring- 
kebing: Gindeskoy, Kjelstrup (E Skjern). Sénderborg: Sonder- 
borg. Sors: Agers6. Svenborg: Langeland Island: Hellenor. 
Tender: Romo (VH), Stensbek Plantage. Vejle: Klattrup. 

Finland: (47 2, 48 6, HY unless indicated otherwise): Abo: 
Lohja, Pernié (AKM), Rymattyla (AKM), Turku (AKM). 
Alandia: Eckeré (AKM, HY), Finstr6m (AKM), Hammarland, 
Jomala (AKM), Saltvik (AKM). Karelia Borealis: Pyhaselka 
(Hammaslahti). Nylandia: Helsinki (CU, HY), Parvoo = Bor- 
ga, Pernaja. Ostrobotnia Australis: Koivulahti. Satakunta: 
Loimaa (AKM), Ylane (AKM). Tavastia Australis: Hattula, 
Hameenlinna, Janakkala, Nastola (AKM), Palkaine (AKM, HY), 
Somero (AKM), Urjala, Vanaja, Yl6jarvi. 

France (35 2, 16 4): Alpes-Maritimes: Guillaumes (FSAG). 
Ariege: Ax-les-Thermes (ZMB). Basses-Alpes: Allos (FSAG), 
Annot (FSAG), Fugeret (FSAG), Les Dourbes (KMG), Mon- 


Trypoxylon figulus: geographic distribution in the New World. 


tagane de Lure (ZMK), Peyresq (FSAG), Saint-André-les-Alpes 
(FSAG). Bouche-du-Rho6ne: Marseille (FIS). Calvados: Lisieux 
(FSAG). Corse: Corte (KMG). Cétes-du-Nord: Saint-Rieul. 
Haute-Savoie: Dent d’?Oche (MHNG), Mont Jorat (RMNH), 
Val de Charmy (RMNH). Haute-Vienne: Rochechouart (FSAG). 
Jura: Arbois. Loiré-et-Cher: Blois (FSAG). Loire-Atlantique: 
Forét de la Roche Bernard (RMNH), Herbignac (RMNH). 
Saéne-et-Loire: Uchizy (FSAG). Seine-et-Oise: Poissy (IEE). 
Var: Fréjus (KMG), Gonfaron (FSAG), Montouroux (RMNH). 
Vaucluse: Carpentras (RMNH). Yonne: Foissy-sur-Vanne 
(FSAG). 

Germany, Democratic (81 9, 29 6, DEI if not indicated oth- 
erwise): Berlin: Berlin (DEI, HD, ZMB, ZSM). Cottbus: Alt 
Débern (ZMB), Muskau, Neu Zauche (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: Gernrode (ZMH), Halle (DEI, ZMB), Kyffhauser, See- 
burg, Naumburg (TMB). Leipzig: Winkelmihle. Magdeburg: 
Arendsee (SMT). Neubrandenburg: Faule Ort, Naturschutz- 
gebiet Miiritzhof. Potsdam: Fiirstenberg (TMB), Zechlin (ZMB), 
Zootzen. Rostock: Prerow, Rostock, Stralsund (DEI, ZMB). 
Rigen: Hiddensee Island (DEI, SMT), Riigen Island: Ménch- 
gut (SMT) and Ummanz. Schwerin: Campow (ZMB), Schwerin 
(ZMB), Wendeltorf near Schwerin. 


————— 


PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 131 


Germany, Federal (93 2, 51 4): Baden-Wiirtemberg: Enz- 
klésterle (KS), Heidelberg (ZSM), Hochwacht (HW), Isny 
(NHMB), Kaiserstuhl (ZMB), Karlsruhe (KS, ZMH), Kiissa- 
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), H6rgersthausen near 
Moosburg (ZSM), Ingolstadt (ZSM), Kahl (FIS), Miinchen 
(FSAG, ZSM), Niirnberg (ZSM), Rotwand area (ZSM), Schlier- 
see (ZSM), Tegernsee (ZSM). Hamburg (ZMH): Ochsenwarder, 
Warwisch. Hessen: Battenfeld near Biederkopf (ZMH), Griin- 
dau E Frankfurt (HW, HY), Marburg (HW, WJP). Nieder- 
sachsen: 2 km NW D6tlingen (VH), Dérpen: 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 (HW), Siegen (HW). Rheinland-Pfalz: low- 
er Ahr valley (FIS), Mainz (KS), Nattenheim (FSAG), Worms 
(FIS). Schleswig-Holstein: Amrum Island (VH), Eutin (KS), 
Thlkathe 2 km SE Kiel (VH), Liitjenburg (KS), Ratzeburg (ZMH), 
Schierensee SW Kiel (VH), Schleswig (VH). 

Great Britain (92 2, 106 ; 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: Byfleet, Esher, Hor- 
sell, Weybridge. Sussex: Midhurst: Ambersham Common. 

Greece (2 2, 4 3): Peloponnesus (de Beaumont, 1965): Mega 
Spilaion, Pirgos, and Taygetus. Sterea Ellas: Karpenission 
(KMG). Thessalia: Aspropotamos near Kalabaka (KMG). 

Hungary (13 2, 3 4): Bacs-Kiskun: Kalocsa (TMB), Tabdi 
(TMB). Gyoér-Sopron: Neusiedlersee (NHMV). Somogy: Bal- 
atonszemes (TMB). Szolnok: Jaszbéreny (TMB). Tolna: Si- 
montornya (NHMYV). Veszprem: Tihany Peninsula on Balaton 
Lake (HD, TMB, WJP). 

Italy (19 2, 8 4): 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, 
NHMY). Trentino-Alto Adige: Bolzano (as Bozen, NHMYV), 
Cavalese (BB), Collalbo (de Beaumont, 1959), Ortisei (NRS), 
Trafoi (NHMYV). 

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 4; KT, TN. 

Morocco (1 2, 2 4): Asni, 1250 m alt. (paratype 2 of figulus 
barbarum, RMNH), Fez (holotype of fiewzeti, IEE), Marrakech 
(BMNB). 

Netherlands (108 2, 73 6, RMNH if not indicated otherwise): 
Drenthe: Emmen, Erm, Havelte, Sellingen, Wijster, Zuidlaren. 
Gelderland: Apeldoorn (FSAG), Barneveld, Hulshorst, Nij- 
kerk, Putten, Renkum, Vierhouten, Wageningen (GVR, 
RMNH). Groningen: Onnen. Limburg: Epen (GVR), Haelen, 
Heerlen, 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, Riynsburg, Rotterdam, ’s Gravenhage, Warmond. 

Norway (1 2, 1 6); Buskerud: Al (as Aal, DEI). Oppland: Lom 
(RMNH). 

Poland (35 2, 24 3): Biatystok: Bialfowieza (as Bialowies, FIS, 
ZMB), Lenkowo near Grajewo (ZMB). Bydgoszcz: Bydgoszcz 
(as Bromberg, SMT). Kielce: Gory Pieprzowe near Sandomierz 
(CAS), Kielce (TMB), Sandomierz (CAS). Lublin: Kazimierz 
on Wista River (WJP). Warszawa: Podkowa Lesna (CAS). 
Wroctaw: Muszkowice near Henrykow (WJP), Osola: 25 km 
NW Wroctaw (WJP), Wroclaw (CAS, WJP). Szczecin: Bielinek 
(as Bellinchen, ZMB), Miedzyzdroje (as Misdroy, ZMB). Zie- 
lona Gora: Kostrzyn (as Ciistrin, ZMB). 

Portugal (2 2, 2 6): Douro: Resende (RMNH). Estramadura: 
Lisboa (RMNH). 

Romania (8 2, 9 4, MGA if not indicated otherwise): Bacau: 
Lacul Bicaz 20 km W Piatra Neamt (HD). Brasov: Sighisoara, 
Turnisor near Sibiu. Bucuresti: Branesti (WJP), Budesti. Con- 
stanta: Crisan, Hagieni. Crisana: Ineu (as Borosjendé, TMB). 
Orasul Bucuresti: Mogosoaia, Pantelimon. Timisoara: Esel- 
nita, Jupalnic near Mehedinti, Mehadia (NHMV), Orsova 
(RMNH). 

Spain (53 2, 51 4): Alicante: Alicante (K MG), Orihuela (IEE). 
Avila (SFG): Becedas, Gilbuena, Puerto Castilla, Solana de 
Béjar, Tremedal. Barcelona: Canet de Mar (FSAG). Caceres 
(SFG): Bafios de Montemayor, Cabezuela del Valle, Gargan- 
tilla, La Garganta. Cadiz: Algeciras (NHMV). Ciudad Real: 
Ruidera (FJS). Granada: Salobrefia (VH). Huesca: Arguis 
(RMNH), Torla (FSAG), Valle de Ordesa. Lerida: Bohi, Lago 
di San Mauricio (FSAG). Logroto: San Roman de Cameros 
(IEE). Madrid: El Escorial (IEE), Sierra de Guadarrama (IEE). 
Salamanca (SFG): Béjar, Cantagallo, Colmenar de Monte- 
mayor, El Cerro, La Cabeza de Béjar, Lagunilla, Montemayor 
del Rio, Navacarros, Palomares de Béjar, 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 Banez (FJS), Villalba de 
los Alcores (FJS). Zamora: Montamarta (FSAG). 

Sweden (43 2, 40 6, NRS ifnot indicated otherwise): Dalarna: 
Taktbo. Oland: Arontorp, Glomminge, Hégsrum, Halltorps 
hage, Morbylanga, Repplinge, Vickleby. Ostergotland: St. Anna 
Korsnas, Simonstorp. Skane: Ahus, Halsingborg, Having, Héér 
distr. (BMNH), Torekov (ZMK), Trolleholm, Vitemélla. Sma- 
land: Hagby, Jonk6ping, Kalmar, Kalmar Sund (Bla Jungfrun), 
Séderakra. Sédermanland: Haggenas, Malarhojden, M6rto, 
Tullgarn, Tyres6, Tyres6 — Brevik. Uppland: Bj6rk6, Habo — 
Tibble, Osteriiker, Radmansé, Svartsj6, Varmdé, Vassunda. 

Switzerland (78 2, 41 6, NHMB if not indicated otherwise): 
Bern: Adelboden, Bantiger (7 km NE Bern), Batterkinden, Bern, 
Biischiwald forest (5 km SSW Bern), Gadmen, Miirren, Signau, 
Spiez, Wachseldornmoos bog (27 km SE Bern), Zweisimmen, 
Zwischenfliih (16 km SSW Thun). Genéve: Genéve (MHNG, 
NHMY), Genthod (NHMV), Peney (MHNG, NHMYV). Grau- 
biinden: Chur (ZSM), Parc National Suisse (MHNG), Somvix, 
Versam. Luzern: Entlebuch Graben (CAS). Sankt Gallen: Vat- 
tis. Valais: Ausserberg (11 km W Brig), Ayel — Zinal (RMNH), 
Berisal (BMNH, NHMB, NHMV), Chalet a Gobet (BMNH), 
Champery, Eusegne (as Usegne), Evoléne, Haudéres, Inden, 
Létschental, 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 °, 3 6): Afyon: Cay (FSAG). Ankara: Ankara (as 
Angora, TMB), Karagol (BMNH). Denizli: Saraykéy (as Seraj- 
K6j, TMB). Istanbul: Sile area (BMNH). Konya: Konya (JG). 
Kiitahya: Karat Dagi (BMNH). Samsun: K6priibasi (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 2): Khanlar (as Helenendorf, NHMY). Ka- 
zakh SSR: Vostochnokazakhstanskaya obl.: foothills of Azu- 
tau Khrebet 18 km N Alekseyevka (1 3, VLK); Kalbinskiy 
Khrebet, 20 km SE Leninka (1 2, VLK); 15 km NW Ust’- 
Kamenogorsk (1 6, VLK). Latvian SSR (2 2, 8 6): Wezkukkul, 
20 km S Jaunjelgava (ZMB). Lithuanian SSR (1 2, 2 2): Ignalina 
(DEI). Russian SSR: Belgorodskaya oblast: Valuyki (2 9, 
NHMYV). Chelabinskaya oblast: Ilmenskiy Zapovednik (1 8, 
ZMMU). Gorno-Altayskaya Avtonomnaya oblast: Uymen 
River (2 6, ZMMU). Ivanovskaya oblast: Kineshma (1 9, 
ZMMU). Karelian ASSR (2 2, 1 6, HY): Salmi, Sortavala. 
Kuybyshevskaya oblast: Mirnyi (2 2°, VLK). Leningradskaya 
oblast: Solnechnoye (as Ollila, 1 ¢, HY), Streltsovo (as Muola, 
1 2, HY), Suursari Island (as Hogland, | 6, HY). Karachayevo- 
Cherkesskaya oblast: Teberda-Dzhamagat (2 2, HD). Moskov- 
skaya oblast: Krylatskoe near Moskva (4 2, 1 6, ZMMU), Mosk- 
va (1 2°, ZMMU), Mytishchi (1 6, ZMMU), Zavety IV’icha 
(1 6, ZMMU). Orenburgskaya oblast: Kargala near Orenburg 
(1 °, ZMB). Ukrainian SSR: Otuzy Valley in Crimea (2 8, 2 4, 
ZMMU), Podgortse E Lvov (as Podhorce, 1 2, NHMYV), Se- 
bastopol area (1 6, ZMMU). 

Yugoslavia (11 2, 6 4): Croatia: Plitvice (RMNH), Senj (VH). 
Kosovo: Pec (TMB). Macedonia: Mavrovska Valley (RMNH), 
Ohrid - Resen (RMNH). Slovenia: Begunje near Postojna (KS), 
Kranjska Gora (as Weissenfels, NHMV), Portoroz (FIS), Rad- 
enciin Mura Valley (near Austrian border, as Radein, NHMV), 
Vipava (as Wippach, NHMV). 

Recorps (New World) (Krombein 1951:955, 1979:1643; 
Finnamore 1982:115).—Quebec: Duchesnay (1 2°, USNM), Ile 
Perrot (2 2, LEM), Lakeside (1 °, LEM), Levis (3 °, LEM), 
Montreal (1 2°, CU; 2 6, LEM), Ste. Anne de Bellevue (3 2, 6 
6, LEM), Ste. Annes (1 9, 3 6, LEM), St. Hilaire (5 °, LEM), 
St. John’s Co. (1 2, LEM). 

Ontario: Ottawa (2 2, 2 6, CNC), St. Anne’s (4 2, CNC). 

Maine: Bangor (1 2°, MCZ). 

New Hampshire: Glen House (1 6, USNM). 

Massachusetts: Cambridge (1 2, USNM), Dorchester (1 2, 
MCZ), Fall River (1 2, USNM), no specific locality (1 2¢, WSU). 


Trypoxylon minus de Beaumont 


Trypoxylon figulus var. minus DE BEAUMONT, 1945:478, 2, 6 
(as minor, incorrect original spelling). Holotype: 2, Switzer- 
land: Cologny near Geneva (Mus. Zool. Lausanne). — 
BLUTHGEN, 1951:234 (var. minor); DE BEAUMONT, 1958:206 
(forma minor), 1959:30 (same); Wolf, 1959:15, 16 (figulus 
minus); VALKEILA, 1961:144 (var. minor); DE BEAUMONT, 
1964a:290, 1964b:84 (forma minor), 1965:56 (same), 1967: 
338 (same); BOHART AND MENKE, 1976:346 (ssp. minor); 
RicHARps, 1980:45 (var. minor). 

Trypoxylon figulus koma Tsunek1, 1956:28, 2, 6. Holotype: 8, 
Korea: Mt. Kodai (K. Tsuneki collection, Mishima). New 
synonym. — BOHART AND MENKE, 1976:346; TsuNEKI, 1981: 
20. 

Trypoxylon figulus medium: Wo F, 1959:fig. b. 


Di1AGnosis.— Most minus can be recognized 
by the presence of a flat, median process emerg- 


ing from the anteroventral mesothoracic carina 
(Fig. 5B, C). However, the process is absent in a 
female and two males from Linz, Austria, and 
also one of the females from Gonfarons, France, 
which otherwise do not differ from the remaining 
minus examined. Unlike most figulus, the me- 
sopleural setae around scrobe are shorter than 
the midocellar diameter. In the female, the me- 
dian clypeal projection is shorter than in medi- 
um; unlike figulus and medium, the clypeal free 
margin is scarcely sinuate or evenly arcuate be- 
tween the orbit and the projection (Fig. 5A). The 
female hindcoxal pit is circular or nearly so (Fig. 
5D-F), unlike most figu/us. The maximum length 
of male flagellomere X is 0.5—0.7 times its width; 
flagellomere XI is longer than in most medium: 
its maximum length equals 2.2—3.1 times its bas- 
al diameter. Body length 6-9 mm in female, 
5-7.5 mm in male. 

VARIATION. —In most females, the free margin 
of the clypeal projection is emarginate, but it is 
entire in a female from Finland, one from France 
and one from Sweden. 

In most males, the maximum length of fla- 
gellomere X is 0.6—-0.7 times its width, and the 
maximum length of flagellomere XI is 2.2—2.4 
times the basal diameter. These ratios are 0.8 
and 2.6, respectively, in some males from Bel- 
gium, 0.6 and 2.6 in the single male examined 
from Sardinia, 0.8 and 3.0 in the single male 
from Corsica, and 0.5 and 3.1 in the single male 
from Gerona, Spain. 

Lire History.—A female from Elender Wald, 
Austria (NHMV), was reared from a gall of An- 
dricus kollari (Hartig). I was unable to find this 
locality. 

GEOGRAPHIC DISTRIBUTION (Fig. 5).— Palearc- 
tic Region between western Europe and Korea, 
north to about 64°N in Norway and Sweden and 
about 67°N in Finland, south to northern Med- 
iterranean countries including Sardinia and Sic- 
ily, northern Turkey and southern Kazakh SSR. 
Unknown from Great Britain and North Africa. 
The absence of this species in Great Britain sug- 
gests that it reached northwestern Europe only 
recently, after separation of the British Isles from 
the continent. 


Recorps.— Austria (41 9, 52 6, NHMV if not indicated oth- 
erwise): Burgenland: Breitenbrunn (JG), Donnerskirchen on 
Neusiedlersee (KS), Mogersdorf (JG), M6rbisch: 5 km S Rust 
(RMNH). Karnten: Bleiberg (RMNH), Mallnitz (ZMB), Wai- 
disch bei Ferlach (FSAG). Niederésterreich: Anninger, Bis- 
amberg near Vienna, Briihl, Dornbach, Hainbach, Hainsburg 


PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 133 


FiGure 5. 
female hindcoxa ventrally, D—female hindcoxal pit, vertical view, E—same, oblique view. 


Trypoxon 


(ZMB), Kahlenberg near Vienna, Neunkirchen (FSAG, JG), 
Piesting, Purgstall (JG), Schneeberg, Traismauer, Wien (Pra- 
ter). Oberésterreich: Almsee (NHMV), Gutau (JG), Inner- 
breitenau (JG), Kremsmiinster (FSAG), Linz (HY, JG, NHMV), 
Rutzing bei Hérsching (JG), Sarleinsbach, Zeissberg bei Fre1- 
stadt (FSAG, JG), Zellhof bei Bad Zell (JG). Salzburg: Salzburg 
(BMNH, FSAG). Steiermark: Admont (FSAG), Gleichenberg 
(JG), Riegersburg (JG), Tragdss-Oberort (DBB), Weinburg (JG). 
Tirol: Huben (ZMB), Innsbruck, Kals (RMNH), Stubai (WJP), 
Wenns (BMNH), Zwieselstein in Otztal. 


minus: A—female clypeus, B—mesothoracic venter, C—anteroventral mesothoracic process, D— 


Belgium (348 °, 379 4, FSAG if not indicated otherwise): 
Brabant: Berchem-Sainte-Agathe, Groenendael, Monstreux, 
Rhode-Saint-Genése, Rixensart, Tilly, *t Roth (RMNH), Wo- 
luwe-Saint-Pierre. Hainaut: Gilly, Gosselies, Lobbes-Calvaire, 
Quaregnon. Liége: Bleyberg, Chaudfontaine, Clermont-sur- 
Berwinne, Dalhem, Embourg, Esneux, Forét de Grunhault, 
Fouron-Saint-Pierre, Hergenrath, Herstal, Jupille, Li¢ge, Queue- 
du-Bois, Remersdael, Spa, Xhendelesse, Wandre. Limburg: 
Bassenge, Eben, Kanne, Sint Pietersberg (RMNH), Tongres, 
Wonck. Luxembourg: Bodange, Chatillon, Les Epioux. Namur: 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 


134 


“UOTINGLISIP I1ydess0a8 “snuIU uojAXOdA T. 


‘9 aUNOL] 


PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 13 


Beauraing, Couvin, Felenne, Gembloux, Gesves, Jambes, La 
Plante, Malonne, Mont-Gauthier, Namur, Saint Marc, Wil- 
lerzie, Y voir. West-Vlaanderen: Sint Pieter. 

Bulgaria (1 6): Chepalare Pass in Rhodope Mts. (ZMT). 

Czechoslovakia (4 2, 4 4): Jihomoraysky Kraj: Vranov W 
Znojmo (as Frain, NHMV). Vapadoslovensky Kraj: Stirovo 
(HD, SMT). 

Denmark (2 2, ZMK): Maribo: Lolland Island: Roden Skov. 
Vejle: Klattrup. 

Finland (105 2, 124 4, HY if not indicated otherwise): Abo: 
Pernid (AKM), Rymattylé (AKM), Sarkisalo (AKM), Turku 
(AKM). Alandia: Eckeré6 (AKM, HY, ZMK), Finstr6m (AKM), 
Geta (ZMK), Hammarland, Lemland. Karelia Australis: V1- 
rolahti. Karelia Borealis: Herajoki on Pielinen Lake (ZMK), 
Kesalahti, Kitee (AKM, HY), Polvyarvi (AKM), Tohmajarvi. 
Karelia Ladogensis: Parikkala, Simpele. Kuusamo: Pera Posio. 
Lapponia Kemensis: Sodankyla. Nylandia: Helsinki, Pernaja 
(as Parna). Ostrobottnia Australis: Koivulahti. Ostrobottnia 
Borealis: Ravasjaérvi, Rovaniemi. Ostrobottnia Media: Siika- 
joki. Satakunta: Loimaa (AKM), Suoniemi. Savonia Australis: 
Lappeenranta, Rautjdrvi. Tavastia Australis: Aitolahti, Ha- 
meenlinna, Hattula, Janakkala, Kalvola, Kangasala, Lahti 
(AKM), Lammi, Luopioinen (AKM), Palkane, Pirkkala (as 
Birkkala), Somero (AKM), Tyrvanté, Vanaja, Yl6jarvi. 

France (24 2, 17 4): Basses-Alpes: Allos (FSAG), Annot 
(FSAG), Colmars (FSAG), La Javie (FSAG), Peyresq (FSAG), 
Saint-André-les-Alpes (FSAG). Corse: Bonifacio (KMG), Col 
de Sorba (KMG). Drome: Dieulefit (RMNH). Haute Savoie: 
Chamonix (RMNH). Haut-Rhin: Rouffach (FSAG). Loiret: 
Lorris (RMNH). Lozere: Sainte Enimie (BMNH). Nieévre: 
Nevers (RMNH). Puy-de-Dome: Beaune (FSAG, ZMK), Besse 
(BMNH). Pyrenees-Orientales: Vinca (VH). Var: Gonfarons 
(FSAG). 

Germany, Democratic (33 2, 20 4, DEI if not indicated oth- 
erwise): Berlin: Berlin-Borgsdorf (HD). Dresden: Freital (SMT), 
Gersdorf near Kamenz (SMT), Moritzburg (SMT). Erfurt: 
Nordhausen area (HD). Frankfurt: Eberswalde area (DEI, HD), 
20 km NW Eisenhiittenstadt (HD), Sch6nwalde near Berlin 
(HD). Gera: Beutnitz bei Jena. Halle: Halle (DEI, ZMB), 
Kyffhauser (DEI, ZMB). Karl-Marx-Stadt: Freiberg. Leipzig: 
Dormreichenbach near Wurzen, Leipzig, Reglitz bei Schkeu- 
ditz. Magdeburg: Haldensleben, Stapelburg (ZMH). Neubran- 
denburg: Serrahn near Neustrelitz (HD), Teterower See (HD). 
Rostock: Rostocker Heide. Schwerin: Perleberg, Schwann near 
Giistrow. 

Germany, Federal (59 9, 42 4): Baden-Wiirttemberg: Enz- 
klésterle (KS), Grétzingen (KS), Heidelberg (BMNH), Kai- 
serberg (ZMH), Kaiserstuhl (ZMB), Karlsruhe (KS, ZMH), 
Kiissaberg (KS), Tiibingen (KS), Windenreute near Emmen- 
dingen (ZMB). Bayern: Aschaffenburg (VH), Balderschwang 
(KS), Karlstadt am Main (KS), Lohr am Main (KS), Main- 
franken (FIS), Miltenberg (FIS), Miinchen (ZSM), Oberau in 
foothill of Ammer Mts. (KMG), 32 km S Wiirzburg (RMNH). 
Hessen: Dillenburg (HW, WJP), Fulda (VH), Marburg (HW, 
WJP), Taunus (KS). Rheinland-Pfalz: Bad Miinster am Stein 
(KS), Burgen an der Mosel (TMB), Donnersberg, Mainz (KS). 
Niedersachsen: Sage: 25 km S Oldenburg (VH). Schleswig Hol- 
stein: Liitjenburg (KS), Russee near Kiel (VH), Schierensee SW 
Kiel (VH). Westfalen: Plettenberg (HW), Siegen (HW). 

Greece (2 2, 2 4): Ionian Islands: Isle de Levkas (as Lefkas, 
BMNH). Sterea Ellas: Timfristos (KMG). 

Hungary (1 2, 1 3): Balaton (HD). 

Italy (23 2, 14 4): Abruzzi: Scanno (K MG). Emilia-Romagna: 
Bologna: Pracchia (RMNH), Ronzano (NHMV): Modena: 


nN 


Zocca: Montetortore (NHMY). Friuli-Venezia Giulia: Ge- 
mona (JG), Prosecco (NHMYV), Tarvisio(NHMV). Lombardia: 
Varese: Mercallo (MSNM). Piemonte: Borgomale (GP), Cas- 
telnuovo in Asti (GP), Chianale (GP), Condove (GP), San 
Benedetto Belbo ca 20 km S Alba (GP), Val d’Angrogna in 
Alpi Cozie (1 2, det. J. de Beaumont, WJP), Vinovo 10 km § 
Torino (GP). Sardegna: Aritzo (BMNH), Desulo (BMNH). 
Sicilia: Piano Battaglia (KMG). Trentino-Alto Adige: Colle 
Isarco (MSNM), Merano (as Meran, FSAG). Valle d’Aosta: 
Buthier (ZMB). Veneto: Padova (de Beaumont, 1959). Venezia 
Giulia: Trieste (NHMY). 

Korea: North Korea: Mt. Kodai near 38th parallel (Tsuneki, 
1956, 1981), Mt. Sombo near 38th parallel (1 2, 1 4). South 
Korea: Seoul (Tsuneki, 1956, 1981). . 

Netherlands (5 2, 9 6, RMNH unless indicated otherwise): 
Limburg: Elkenrade, Epen, Maastricht, Slenaken, Vijlener Bosch 
(GVR). Noord Brabant: Empel. Zuid Holland: Leiden. 

Norway (11 2, 19 4): Akerhus: Oslo (as Kristiania, ZMUB). 
Hordaland: Granvin (ZMUB). Nord Tréndelag: Rora (FSAG). 
Oppland: Lom (RMNH). Sogn og Fjordane: Balestrand 
(ZMBU). Vestagder: Kristiansand (ZMUB). Vestfold: Herstad 
(ZMUB). 

Poland (7 2, 13 4): Biatystok: Biatowieza (as Bialowies, ZMB). 
Krakow: Ojcow (SMT), Pieniny Mts. (WJP). Wroctaw: Kar- 
konosze Mts. (as Riesengebirge, ZMB), Osola: 25 km NW 
Wroctaw (WJP), Roscistawice (WJP), Sobdtka: 35 km §S 
Wroctaw (WJP), Wroctaw (CAS), 10 km NE Wroctaw (WJP). 
Rzeszow: Jarostaw (CAS). 

Portugal (1 2): Douro: Resende (RMNH). 

Romania (6 2, 3 6, MGA if not indicated otherwise): Bu- 
curesti: Ciolpani, Peris, Videle (WJP). Constanta: Babadag. 
Ploesti: Valea Longa-Gorgota. Timisoara: Eselnita, Ogradina, 
Orsova. 

Spain (2 2, 4 4): Gerona: Gerona (BMNH). Huesca: Sierra 
de Oroel (FSAG), Valle de Ordesa (FIS). Salamanca: Horcajo 
de Montemayor (SFG), Valdehijaderos (SFG). 

Sweden (34 2, 44 4, NRS if not indicated otherwise): Bo- 
huslan: Ljung. Dalarna: Ludvika, Nas, Stora Kopparberg, 
Taktbo. Gastrikland: Sandviken. Gotland: Fardume Trask E 
Larbro (ZMK), Visby, St. Karls6. Jamtland: Undersaker. Narke: 
Orebro. Oland: Halltorps hage, Hégsrum, Rapplinge. Skane: 
Hoor distr. (BMNH), Skaralid, northern Skane (BMNH). Os- 
tergotland: Kvarsebo, Norrképing, Simonstorp. Smaland: Bar- 
keryd, Horeda, Jarsnas, Ljungarum. Sodermanland: Flisby, 
Runmaro, Tullgarn, Tyresé - Svart6cken. Uppland: Harparbol, 
Radmans6, Stockholm, Uppsala (HY), Vassunda, Vira bruk. 
Vasterbotten: Handene. Vastmanland: Sala. 

Switzerland (21 2, 26 6, NHMB if not indicated otherwise): 
Bern: Batterkinden, Bern, Burgdorf, Grauholz forest (up to 9 
km NNE Bern), Wachseldornmoos bog (27 km SE Bern). En- 
gadin: Zuoz (AWE). Genéve: Genéve Cologny (BMNH), Gen- 
thod (NHMV), Peney (MHNG). Graubiinden: Parc National 
Suisse (de Beaumont, 1958), Somvix. Tessin: Lugano (FSAG). 
Valais: Binntal (15 km NE Brg), Grimentz (FSAG, RMNH), 
St. Luc (RMNH), Simplon (BMNH), Zermatt (RMNH). Vaud: 
Nyon (MHNG). Ziirich: Wadenswil. 

Turkey (1 2, 2 4): Artvin: Artvin, Berta (BMNH). Istanbul: 
Belgrat Orman (de Beaumont, 1967). Samsun: Bafra (BMNH). 

USSR: Georgian SSR: Bakunani (1 °¢, ZMMU). Kazakh SSR: 
Chimkent, 2000 m (1 4, WJP); Razdolnyi area, 50°43’N, 81°06’E 
(1 6, VLK), Zapovednik Aksu-Dzhabagly (1 2, ZMMU). Kirgh- 
iz SSR: Arkit in Chatkalskiy Khrebet (1 6, ZMMU). Russian 
SSR: Karelian ASSR: ozero Tumas (as Kolatselka, | 2, HY). 
Kemerovskaya oblast: 60 km SE Novokuznetsk (1 4, VLK). 


136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10 


Leningradskaya oblast: Kondratyevo W Vyborg (as Saékkyarvi, 
1°, HY). Moskovskaya oblast: Mytishchi (1 6, ZMMU). Perm- 
skaya oblast: Nizhnyaya Kurya, 15 km W Perm (3 , 2 6, WJP). 
Ukrainian SSR: Crimea: Krymskiy Zapovednik (1 6, ZMMU). 

Yugoslavia (4 2, 3 6): Croatia: Plitvice (RMNH), Skrad (GP). 
Slovenia: Bled (RMNH), Cerknica (KS), Logatec (as Loitsch, 
ZMH), Op¢ina Mtn. on Italian border E Triest (NHMV). Ser- 
bia: Drazevac (FSAG). 


Trypoxylon medium de Beaumont 


Trypoxylon figulus var. medium DE BEAUMONT, 1945:477, 2, 
6 (as media, incorrect original spelling). Holotype: °, Swit- 
zerland: Martigny (Mus. Zool. Lausanne). — BLUTHGEN, 1951: 
234 (var. media); DE BEAUMONT, 1958:206 (forma media), 
1959:30 (same); WoLtF, 1959:15, 16 (figulus medium); VALK- 
EILA, 1961:144 (var. media); DE BEAUMONT, 1964a:290, 
1964b:84 (forma media), 1965:56 (same), 1967:338 (same); 
BOHART AND MeENnkKE, 1976:346 (ssp. medium); LOMHOLDT, 
1976:267 (figulus media), RicHARDS, 1980:45 (var. media); 
TsuNEKI, 1981:19 (medium). 

Trypoxylon figulus minus var. rubi WoirF, 1959:15, 6. ! Lec- 
totype, 4, Federal Germany: Plettenberg (coll. H. Wolf, Plet- 
tenberg), present designation. Synonymized by DE BEAUMONT, 
1964:291. 

Trypoxylon figulus minus: Wo Ff, 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 1s either straight or sinuate and curved 
posterad mesally, but unlike that of most minus 
it is not expanded into a process. Unlike those 
of most figu/us, 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 figu/us). 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 
ventroexternal expansion at about midlength (Fig. 
7E, F); expansion absent in figul/us and minus. 


Body length 6.5—12 mm in female, 6.0-8.5 mm 
in male. 

Lire 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- 
ern 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. 


Recorps.— Austria (30 2, 20 4, 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), Zurndorf 
(AWE, RMNH). Nieder@sterreich: Bisamberg near Vienna, 
Deutsch Altenburg (ZMB), Donauauen near Vienna (ZMH), 
Hainburg (ZMB), Herzograd (JG), Marchfeld (JG), Oberwei- 
den (DBB), Piesting, Schneeberg, Stammersdorf (DBB), Wien- 
Tiirkenschanze. Oberésterreich: Gutau (JG). Salzburg: Kat- 
chenberghohe (VH), Salzburg (BMNH). Tirol: Galtiir (BMNH), 
Huben (ZMB), Innsbruck, Kraspes (DEI), | km SW Zwiesel- 
stein. Voralberg: Bielerhdhe. 

Belgium (11 2, 17 6, FSAG): Antwerpen: Geel. Liége: Liége, 
Montzen, Spa, Wandre. Limburg: Bree, Eben, Lummen. Lux- 
embourg: Chatillon, Les Epioux, Torgny. Namur: Eprave, Fer- 
age, Mont-Gauthier. 

Bulgaria (1 2): Ruse (SMT). 

Cyprus (3 2, 6 4): Amathus (FSAG, RMNH), Paphos (KMG). 

Czechoslovakia (1 2, 2 4): Jiho¢esky Kraj: Prachatice (as 
Prachatitz, NHMV). Vapadoslovensky Kraj: Starovo (SMT). 

Denmark (11 2, 8 6, ZMK, if not indicated otherwise): Born- 
holm: Arnager. Frederiksborg: Hulerod, Humblebek (S Hel- 
singor), Tisvilde Hegn. Hjérring: Fauerholt Hede (ca 10 km 
W Frederikshavn), Skoven on Lese Island. Maribo: Kris- 
tianssede Skov on Lolland Island. Randers: Glatved Strand 
and Kalo on Djursland Peninsula, Mols Bjerge. Ribe: Bors- 
mose, Ho Plantage. Svendborg: Keldsnor on Langeland Island. 
Thisted: Bagso at Nors (ca 7 km NW Thisted). Ténder: Rome 
Island (RMNH), Stensbeck Plantage. 

Finland (18 2, 15 6, HY if not indicated otherwise): Abo: 
Pernié (AKM), Rymiattyla (AKM). Alandia: Finstrém (AKM), 
Hammarland. Karelia Australis: Virolahti. Karelia Borealis: 
Nurmes. Kuusamo: Kuusamo (ZMK). Lapponia Inarensis: Iva- 
lo. Nylandia: Hyvinkadad. Ostrobottnia Media: Paavola. Tavas- 
tia Australis: Hameenlinna, Hattula, Janakkala, Palkane, So- 
mero (AKM), Vanaja. 

France (24 2, 8 6, FSAG if not indicated otherwise): Aisne: 
Liesse. Alpes Maritimes: Aurons (AWE). Aube: bois de Lignié- 
res. Aveyron: Creissels. Basses-Alpes: Annot, Montagne de Lure 
(ZMK), Peyresque. Dordogne: Paunat. Drome: Dieulefit 
(RMNH). Haute-Loire: Pont de Suméne. Haut-Rhin: Rouf- 
fach. Hautes-Alpes: Saint-Véran. Hérault: Canet (BMNH). 


PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 


FiGur_E 7. 


Trypoxylon medium: 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, process of gonoforceps. 


Landes: Mont-de-Marsan (MHNG), Parentis (K MG), St. Gi- 
rons - Plage (RMNH). Moselle: Orny, Sierck. Pyrenées-Ori- 
entales: La Llagonne - Mont Louis (VH). Var: Saint-Tropez, 
Valescure (KMG). Vaucluse: Carpentras (RMNH). 

Germany, Democratic (15 2, 6 4): Berlin: Berlin (BMNH, 
DEI, ZMB), Spandau (ZMB). Cottbus: Luckau (DEI). Dresden: 
Gersdorf near Kamenz (SMT). Frankfurt: Buckow (ZMB), 
Eberswalde area (DEI), St6ritzsee near Berlin (DEI). Halle: 
Bad Frankenhausen near Kyffhaéuser (HD), Halle (DEI), Kat- 


tenburg near Kyffhaéuser (HD), Kyffhauser (DEI). Potsdam: 
Gross Machnow (ZMB). 

Germany, Federal (25 2, 34 6): Baden-Wiirttemberg: Fed- 
ersee (KS), Kaiserberg (ZMH), Kaiserstuhl (VH, ZMB), Karls- 
ruhe (ZMH), Radolfzell (ZMH). Bayern: Balderschwang (KS), 
Kreuth in Oberbayern (ZMB), Mainfranken (FIS), Miltenberg 
(FIS), Obersdorf in Allgau (ZMB), Sondershausen (ZMH). 
Hessen: Dillenburg (HW, WJP), Marburg (HW). Niedersach- 
sen: Dérpen: 14 km SW Papenburg (VH), Norderney Island 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43. No. 10 


138 


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PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS 139 


(VH). Nordrhein-Westfalen: Krefeld (DEI), Plettenberg (in- 
cluding lectotype and 2 paralectotypes of rubi, HW, WJP), 
Siegen (HW). Rheinland-Pfalz: Bad Miinster am Stein (KS), 
Mainz (KS). 

Great Britain (53 2°, 44 6, BMNH if not indicated otherwise): 
Buckingham: Slough. Devon: Bovey Tracey, Braunton. Dorset: 
Chideock, Wareham. Gloucester: Stroud (Rodborough). 
Hampshire: Basingstoke, Bramdean, Browndown, Fareham, 
Hurn, Stockbridge. Isle of Wigt: Alum Bay, Sandown, St. He- 
len’s. Kent: Cobham, Darenth, Pluckley (KMG). London: Mill 
Hill, Ruislip (KMG), Uxbridge. Oxford: Goring. Surrey: 
Chobbham (KMG), Ewell (DBB), Horsell. Sussex: Findon, 
Midhurst (Ambersham Common), Singleton (The Trundle). 

Greece (1 2, 1 6): Crete: Kato Metokhi (ZMB). Peloponnesus: 
Mega Spilaion (de Beaumont, 1965). Sterea Ellas: Kifissia 
(KMG). 

Hungary (2 2, 1 4): Pest: Ocsa (TMB). Tolna: Simontornya 
(NHMV). 

Iran (1 2): Gorgan, Shaskola Forest (JG). 

Israel (3 2): Baniass (KMG), Eshtaol: Kesalon Valley (KMG). 

Italy (19 2, 7 6): Emilia-Romagna: Modena: Zocca: Monte- 
tortore (NHMV). Lazio (Frosinone): Pontecorvo (NHMYV), 
Roma (4 2, det. J. de Beaumont, WJP). Liguria: Genova 
(MSNM), San Remo (GP). Lombardia (Varese): Mercallo 
(MSNM). Piemonte: San Benedetto Belbo ca 20 km S Alba 
(GP). Trentino-Alto Adige: Bolzano (as Bozen, NHMY), Ca- 
valese (BB), Merano (as Meran, ZSM), Predazzo (ZMB). Ve- 
neto: Colli Euganei: Montegrotto Terme (WJP), Garda (FSAG), 
Jesolo (FSAG), Lido di Venezia (de Beaumont, 1959). Venezia 
Giulia: Trieste (ZMH). 

Netherlands (14 2, 10 4, RMHN if not indicated otherwise): 
Drenthe: Emmen, Exloo, Mantinge. Gelderland: Ede (GVR), 
Kootwijk, Hulshorst. Limburg: Heerlen. Noord Brabant: 
Griendtsveen, Helenaveen, Udenhout, Waalwijk. Noord Hol- 
land: Hilversum. Utrecht: den Dolder. 

Norway (4 2, 5 6): Hordaland: Granvin (ZMUB). Sogd og 
Fjordane: Balestrand (ZMUB). Sér-Tréndelag: Osen district 
(FSAG). 

Poland (1 4): Szezecin: Miedzyzdroje (as Misdroy, ZMB). 

Romania (9 2, 18 6, MGA if not indicated otherwise): Brasov: 
Uilac. Bucuresti: Valea Rosie near Oltenita. Constanta: Ba- 
badag (RMNH), Caraorman, Crisan, Gorgova in Danube Del- 
ta (HD), Periprava, Valu lui Traian. Oradea: Oradea (RMNH). 
Ploesti: Valea lui Bogdon near Sinaia. 

Spain (1 2): Mallorca (DEI). 

Sweden (22 2, 10 6, NRS): Asele Lappmark: Saxnas. Ble- 
kinge: Ronneby. Bohuslan: Ljung. Dalarna: Falun. Gotland: 
Fardume, Faro, Stanga. Halsingsland: Jarvs6. Jamtland: Stor- 
lien. Oland: Hégby, Mérbylanga, Vickleby. Skane: Iv, Sten- 
shuvud. Smaland: Kalmar Sund (Bla Jungfrun). Uppland: Bo- 
gesund, Danderyd. Vastergétland: Osterplana hed. 
Vastmanland: Skinnskatteberg. 

Switzerland (20 2, 20 6, NHMB if not indicated otherwise): 
Bern: Bern, Biel. Geneve: Bois d’?Onex (MHNG), Genthod 
(NHMB, RMNH), Peney (MHNG). Graubiinden: Chur (ZSM), 
Parc National Suisse (de Beaumont, 1958), Scanfs, Somvix. 
Valais: Binntal (15 km NE Brig), Grimentz (RMNH), Lét- 
schental, Martigny (BMNH), Saar Fee (BMNH), Sierre 
(NHMY), Simplon (BMNH), Zermatt (RMNH). Vaud: Nyon 
(MHNG). 

Syria (1 4): Damascus (NHMV). 

Turkey (4 2, 9 4): Amasya: Amasya (BMNH). Antakya: An- 
takya (de Beaumont, 1967). Antalya: Antalya (BMNH), Finike 


(de Beaumont, 1967), Finike-Kas road (BMNH). Bursa: Ulu- 
dag (BMNH). Kastamonu: Kastamonu area (BMNH). Mersin: 
G6ézne (BMNH), Mut (JG). Mugla: near Kéycegiz (BMNH), 
Marmaris (BMNH). Samsun: Samsun area (BMNH). 

USSR: Armenian SSSR: Noyembryan (3 6, ZMMU). Geor- 
gian SSR (2 2°, ZMMU): Leselidze, Yermolovskoye near Gagra. 
Kazakh SRR (7 2, 14 4, VLK if not indicated otherwise): Al- 
maatinskaya oblast: 12 km W Alma Ata; Ili river 60 km E Ili 
(now Kapchagai); Kargalinka, 5 km SW Alma Ata; Malaya 
Almaatinka River in Zailiyskiy Alatau (ZMMU), 25 km S 
Turgen village in Zailiyskiy Alatau. Semipalatinskaya oblast: 
30 km SE Georgyevka; Tarbagatai Khrebet foothills 6 km N 
Irinovka (which is 47°09'N, 81°53’E); 12 km N Zharma. Vos- 
tochnokazakhstanskaya oblast: Baighym Canyon in Narym- 
skiy Khrebet; Kalbinskiy Khrebet 20 km SW Leninka; Ken- 
dyrlik river 15 km E Zaysan; 5 km N Oktyabrskiy in Ulbinskiy 
Khrebet; 15 km NE Ust’-Kamenogorsk; 15 km SSW Ziry- 
anovsk; 28 km SSE Ziryanovsk. Russian SSR: Bashkirskaya 
Avtonomnaya oblast: Kazmash (1 6, ZMMU). Belgorodskaya 
oblast: Valuyki(1 4, NHMYV). Leningradskaya oblast: Primorsk 
(as Koivisto, | 2, AKM). Saratovskaya oblast: Kuznetsk (1 9, 
ZMMU). Ukrainian SSR: Crimea (1 2, 2 6, ZMMU): Karadag, 
Sebastopol. 

Yugoslavia (6 2, | 4): Croatia: Pore¢ (FSAG), Puli (as Pola, 
NHMYV), Rab Island (as Arbe, TMB). Kosovo: Brezovica in 
Sar Mts. (TMB), Peé (TMB). Serbia: Drazevac (FSAG), Sid 
(FSAG). Slovenia: Portoroz (FIS). 


LITERATURE CITED 


Bouart, R. M., AND A. S. MENKE. 1976. Sphecid wasps of 
the world: a generic revision. University of California Press, 
Berkeley, Los Angeles, London. i-ix + 695 pp. 

BLUTHGEN, P. 1951. Neues oder Wissenswertes iiber mittel- 
europaische Aculeaten und Goldwespen II. (Hym.). Bonner 
Zool. Beitr. 2:229-34. 

Cresson, E. T. 1928. The types of Hymenoptera in the Acad- 
emy of Natural Sciences of Philadelphia other than those of 
Ezra T. Cresson. Mem. Amer. Ent. Soc. 5:1-90. 

Day, M. 1979. The species of Hymenoptera described by 
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. 1957 (1956). Hyménoptéres récoltés par une mission 
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Ges. 18:11-16. 


my aw: _— 
Gut io oi ha => Gor 


i > 


Marine ea: 


; LIBRARY 


CALIFORNIA ‘ACS DEMY OFS IENCES 


Vol. 43, No. 11, pp. 141-15 


SOV EnCIe, Mass. 


January 17, 1984 


DESCRIPTION AND OSTEOLOGY OF THRYSSOCYPRIS, A NEW 
GENUS OF ANCHOVYLIKE CYPRINID FISHES, BASED 
ON TWO NEW SPECIES FROM SOUTHEAST ASIA 


By 


Tyson R. Roberts 


California Academy of Sciences, Golden Gate Park, 
San Francisco, California 94118 


and 
Maurice Kottelat 


Naturhistorisches Museum, Augustinerstrasse 2, 
CH-4001, Basel, Switzerland 


ABSTRACT: 


Thryssocypris new genus comprises two new species, the generic type-species 7. smaragdinus 


from the Kapuas River in Kalimantan Barat (Indonesian western Borneo) and 7. tonlesapensis from the lower 
Mekong basin in Kampuchea and Vietnam. Extremely compressed and anchovylike, Thryssocypris differs 
from all other cyprinids in having a non-protrusible upper jaw with the posterior half of its margin formed 
solely by the maxilla, and a pair of elongate, slitlike supraethmoidal laterosensory trenches. Osteological 
study reveals features of the Weberian apparatus, gill arches, pelvic girdle, and terminal radials of the dorsal 
and anal fins that have not been reported previously in cyprinids. The two new species differ in proportions 
and fin positions, in numbers of anal fin rays, scales, and vertebrae, and in coloration. The relationships of 


the new genus to other cyprinids require further study. 


INTRODUCTION 


Two recently discovered, anchovylike, insec- 
tivorous cyprinid fishes, one from the Kapuas 
River in western Borneo (Kalimantan Barat, In- 
donesia) and the other from the lower Mekong 
basin in Kampuchea and Vietnam, are closely 
related to each other and represent a highly spe- 
cialized new genus. We do not know of any pre- 
viously described species referable to this genus. 

Five specimens of the Bornean species were 
obtained at a single locality during an ichthyo- 
logical survey of the Kapuas River in 1976, and 
we have been unable to locate any additional 
material of this species. Nine specimens of the 


Mekong species were obtained by F. d’Aubenton 
during a survey of the Tonle Sap in 1961, and 
at least one hundred specimens by W. J. Rain- 
both during a University of Michigan fisheries 
survey in the Mekong Delta in 1974. This species 
also does not seem to have been collected pre- 
viously. The species differ so markedly in head 
size and other characters that they were not im- 
mediately recognized as congeneric. However, 
closer comparison, including osteological study, 
revealed that they agree in numerous characters 
not found, or at least not reported, in any other 
cyprinids and that they clearly do belong in the 
same genus. 


[141] 


142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


Cyprinidae is the largest family of freshwater 
fishes, and we are very far from a phyletic clas- 
sification of the genera or even a reasonable di- 
vision of the family into taxonomic categories 
above the generic level. A great deal of work 
remains to be done in order to provide adequate 
definitions for cyprinid genera, many of which, 
as currently understood, are polyphyletic and 
must be split up before a phyletic classification 
can be achieved. Since many distinctive cyprinid 
genera are relatively rare, and live specimens vir- 
tually unobtainable, classification must be based 
mainly on information that can be extracted from 
preserved specimens. Osteological accounts of 
distinctive genera, especially if extensively illus- 
trated in a standardized format (e.g., that of Bry- 
con by Weitzman 1962) provide perhaps the most 
ready source of information for use in phyletic 
classification. Unfortunately, there is a dearth of 
such accounts. In addition to describing this new 
genus and its two species, therefore, we present 
an account of its osteology. 


MATERIAL AND METHODS 


Material of the two species described herein is 
deposited in the following institutions: Califor- 
nia Academy of Sciences (CAS), Museum of Zo- 
ology of the University of Michigan (UMMZ), 
Smithsonian Institution (USNM), Museum na- 
tional d’Histoire naturelle, Paris (MNHN), Mu- 
seum d’Histoire naturelle de Genéve (MHNG), 
Museum Zoologicum Bogorense, Bogor, Indo- 
nesia (MZB). The osteological account is based 
on a 53.2-mm paratype of 7. smaragdinus and 
a 46.0-mm paratype of 7. tonlesapensis, which 
were cleared and stained 1n alcian blue and aliza- 
rin. Additional osteological observations were 
made on radiographs of the other four type spec- 
imens of 7. smaragdinus and 12 of the largest 
T. tonlesapensis. Although our osteological fig- 
ures are based mainly on 7. smaragdinus, our 
remarks generally relate to the osteology of both 
species. While it is not always so in fishes pre- 
pared with alcian and alizarin, in our material 
of 7. smaragdinus and T. tonlesapensis, except 
for the distal ends of the intermuscular bones, it 
seems that all skeletal elements stained with al- 
cian are true cartilage; all stained with alizarin 
are true bone. In the figures bone is indicated by 
stipple, cartilage by simple diagonal hatching, 
and fenestrae or foramina opening into intracra- 


nial spaces by cross-hatching. In all of the figures 
the scale bar equals 1 mm. : 

We draw attention to a few reservations con- 
cerning the osteology. The parietal laterosensory 
canal bony tubules and supraoccipital crest were 
badly damaged in the 53.2-mm specimen illus- ~ 
trated and were drawn on the basis of reconstruc- 
tion and comparison with whole, unstained spec- 
imens. The shape of the posteriormost portions 
of the prevomer and parasphenoid and their re- 
lationships to other elements were not observed 
with certainty in the specimen illustrated and 
may be in error. In particular, the appearance of 
the prevomer may be due to breakage rather than 
to mere separation. We did not detect an inter- 
calar, an element sometimes absent in Cyprini- 
dae, but are uncertain of its absence in the present 
instance. Otoliths were not removed so that the 
crania could be preserved intact, and thus they 
are not described or figured. 


Thryssocypris, new genus 


Type-species.— Thryssocypris smaragdinus Roberts and Kot- 
telat, new species. 


DiAGNosis.—Small (largest specimen 63.8 
mm), anchovylike cyprinids with highly com- 
pressed head and body; barbels absent; excep- 
tionally large olfactory organ; well-developed 
hyaline eyelid; elongate, moderately upturned, 
and very narrow terminal jaws; acutely pointed 
snout, slightly to strongly projecting anterior to 
upper jaw; low dorsal and anal fins, with falcate 
Margins, originating in posterior half of body; 
abdomen rounded, without keel. 7hryssocypris 
differ from all other known cyprinid genera in 
the following characters: 1) upper jaw entirely 
nonprotrusible, without rostral cap or even ves- 
tigial groove of rostral cap, with posterior half 
of border formed solely by maxilla; 2) cephalic 
laterosensory system with elongate, slitlike su- 
praethmoidal or rostral canal medial to nasal 
canal, supraorbital canals similarly slitlike, not 
enclosed in bony tubules on frontal bone, and 
an elongate dermosphenotic canal enclosed in a 
bony tubule fused for entire length to dorsal sur- 
face of sphenotic bone; 3) ventral portion of gill 
arches highly specialized, ceratobranchials 1-4 
with proximal portions abruptly narrowed and 
ending in elongate cartilaginous extensions, hy- 
pobranchial one minute and hypobranchials 2-3 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 143 


FiGure l. 


absent, and basibranchials extremely slender; 
4) Weberian apparatus with lateral process of 
centrum 2 extremely elongate and strongly curved 
posteriorly dorsal to tripus, extending laterally 
equally as far as fully formed pleural rib of fifth 
vertebra; 5) ischiac process of pelvic girdle with 
elongate and extremely slender anterior and pos- 
terior processes; and 6) posteriormost pterygio- 
phore of dorsal and anal fins with a deeply bi- 
furcate radial projecting considerably beyond base 
of posteriormost fin rays. 

Regarding the diagnostic characters listed 
above: 1) a few other cyprinids have nonprotru- 
sible upper jaws (e.g., the North American Exo- 
glossum and Parexoglossum), but in these and 
all other cyprinids known to us the maxilla is 
entirely or almost entirely excluded from the gape 
by the premaxilla; 2) some other cyprinids ap- 
parently have laterosensory canals in the su- 
praethmoidal region (e.g., Luciosoma), but the 
position and shape of these canals is quite dif- 
ferent from those in 7Jhryssocypris, and they are 
not slitlike. Lekander (1949) reported rostral lat- 
erosensory organs in some European cyprinids, 
but these are more anterior (near snout tip) and 
are joined by a commissure (absent in 7hrys- 


Thryssocypris smaragdinus, 49.4-mm holotype (MZB 3435). 


socypris). Supraorbital canals in Cyprinidae are 
usually enclosed in bony tubules on the dorsal 
surface of the frontal bone. A dermosphenotic 
laterosensory canal is present in most cyprinids, 
but usually occurs in a short segment of bony 
tubule or on a small laminar dermosphenotic 
bone that is completely separate and superficial 
to the sphenotic bone; 3) in all other cyprinids 
we have examined or know about the gill arches 
have ceratobranchials 1-4 uniformly wide and 
hypobranchials one to three present; 4) many 
cyprinids have a very large lateral process on 
centrum two, but in most instances it projects 
directly laterally from the vertebral column, or 
in some instances is strongly curved posteriorly 
but projects ventrally to the tripus (rather than 
dorsally to it as in 7hryssocypris); 5-6) the un- 
usual condition of the ischiac process and of the 
terminal radials in the dorsal and anal fins is 
unlike anything we know of or have seen reported 
in any other cyprinids. 

A more detailed account of the osteology of 
Thryssocypris is given following the species de- 
scriptions. Some additional characters of the ge- 
nus are given here. Lateral line complete, mod- 
erately curved downward anteriorly. Gill rakers 


FiGure 2. 


Thryssocypris tonlesapensis, 50.6-mm holotype (MNHN 1982-1032). 


144 


TABLE 1. 


Character 


Counts 


Gill rakers on first gill arch! 
Pharyngeal teeth (left/right)! 

Dorsal fin rays 

Anal fin rays 

Pectoral fin rays 

Pelvic fin rays 

Procurrent caudal fin rays 

Scales in lateral series 

Scale rows above + below lateral line 
Median predorsal scales 
Circumferential scales 
Circumpeduncular scales 

Abdominal + caudal = total vertebrae 


Proportions (times in standard length) 


T. smaragdinus 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


QUANTITATIVE CHARACTERS IN TAryssocypris. 


T. tonlesapensis 


0+7 0+8 
1+5/5+1 2+5/5+2 
9'(1),10'2(3) 9'%(3), 1029) 
12%2(1),13'2(2),14'2(1) 15%(12),16'2(6) 
11-12 10-12 
7-8 7-8 
9+9 9+8-9 
36-39 43-46 
54+3 64+3 
252-27? 26-31 
18-20 18-21 
14-15 16-17 


214+17 = 38(2),214+18 = 39(3) 


21422 = 43(12),21+23 = 44(1) 


Head 3.8-4.1 4.9-5.3 
Snout 12.0-12.5 15.4-16.9 
Olfactory organ 18.4-22.4 33.6-34.7 
Eye 15.5-17.0 19.2-24.8 
Lower jaw! 7.9 10.7 
Pharyngeal bone! 11.4 20.0 
Body depth 5.1-5.5 6.2-6.8 
Body width 9.9-11.1 12.2-14.3 
Caudal peduncle depth 11.4-12.0 10.6-11.5 
Pectoral fin 5.7-6.1 6.0-7.0 
Pelvic fin 9.0-10.0 8.9-9.7 
Preanal length 1.3-1.4 1.5-1.6 
Other 
Inclination of jaws SP 30-35° 


' From cleared and stained specimens. 


small, short, and somewhat stubby, leading edge 
of lower limb of first gill arch with seven to eight 
rakers, upper limb with none to one. Pharyngeal 
teeth uncinate, in two rows, 1-2 +5/5+ 2-1. Scales 
large, 36—46 in lateral series, approximately cor- 
related in number with vertebrae, which total 
38-44. Quantitative characters of the genus are 
summarized in Table 1. 

Dorsal and ventral profiles of head and body 
anterior to dorsal and anal fins uniformly and 
gently curved (not forming an angle at occiput 
or pectoral fin origin). Dorsal and ventral surface 
of body anterior to median fins rounded from 
side to side. Body moderately tapered caudally 
(markedly tapered in some cheline cyprinids). 
No indication ofa cranial flexure. Morphological 
features related to ability of head to tilt upwards 
in relation to vertebral column (evidently a feed- 
ing adaptation in Asian cyprinids such as Mac- 
rochirichthys, Salmostoma, Oxygaster, and Che- 


la) are absent. Epaxial musculature does not 
invade cranial roof, angle of occiput is relatively 
acute, and parasphenoid relatively horizontal. 
Scales on dorsum, sides (including lateral line 
scale series), caudal peduncle, and abdomen 
morphologically similar, except that scales of lat- 
eral line series have simple tubule for laterosen- 
sory canal, horizontally oriented, originating at 
or near focus and extending for half or more than 
half length of posterior field. Shape modified oval; 
anterior, dorsal, and ventral margins slightly 
convex, posterior margin rounded or even slight- 
ly pointed (especially in median scale rows and 
near bases of paired and median fins), sometimes 
weakly scalloped. Dorsal and ventral fields more 
or less sharply set off from anterior field but grad- 
ing smoothly into posterior field. Radii present 
on all fields, but best developed on anterior and 
posterior fields. Radii on anterior field 8—15, hor- 
izontally oriented (parallel to each other). Radii 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 145 


ik ' 
rn 


FiGuReE 3. 
optics). 


on posterior field 22—30 or more, slightly diver- 
gent from horizontal, more widely separated than 
those on anterior field. Radii on anterior and 
posterior fields originating near focus. Radii on 
dorsal and ventral fields similar, few in number, 
widely divergent, and variable in length, origi- 
nating remote from focus, nearly parallel to up- 
permost and lowermost radii in posterior field, 
with which they seem to form a continuous se- 
ries, and entirely divergent from radii in anterior 
field. Circuli well defined and evenly spaced in 
anterior, dorsal, and ventral fields, but discon- 
tinued or indistinct on posterior field. Circuli of 
anterior field vertical, of dorsal and ventral fields 
horizontal; circuli of dorsal and ventral fields 
meeting at right angles with circuli of anterior 
field at interfield margins, and bisecting growth 


Thryssocypris smaragdinus, 49.7 mm, twenty-third scale in lateral line scale row (Nomarski interference contrast 


lines of posterior field at a sharp angle. Growth 
lines, readily observable in posterior field, much 
more numerous and more nearly circular in ar- 
rangement than circuli. 

Multicellular horny projections, or tubercles 
(also known as breeding tubercles), absent from 
body and fins, and perhaps also from head. Nu- 
merous minute conical projections toward tip of 
snout, on upper lip, and on lacrimal area of head 
(especially near ventrolateral margin of infraor- 
bital 1) appear to be tubercles. Tubercles fre- 
quently occur on dorsal surface of pectoral fins, 
on scales, and on mandible in many cyprinids, 
especially in males, but are absent from these 
places in Thryssocypris. 

EtryMo.Locy.—From the Greek thrissos, a her- 
ring or anchovy, and cypris, a small minnow. 


146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


Ficure 4. 


Thryssocypris, radiographs. a, T. smaragdinus, 49.4 mm (holotype); b, 7. tonlesapensis, 52.0 mm (paratype, CAS 


50946). Note obvious differences between species but also basically identical morphology of skulls. 


Thryssocypris smaragdinus, new species 
(Figures 1, 3a, 4-9, 10b, 11-19) 

Hototyee.—MZB 3435, 49.4 mm, mainstream of upper 
Kapuas River, 6 km w of Putussibau, Kalimantan Barat, In- 
donesia, lat. 0°50.5'N, long. 112°52’r. Seining in late afternoon 
and at dusk, on gently sloping open beach with coarse sand to 
fine gravel bottom and moderate current. 9 Aug. 1976. 

ParatyPes.—CAS 49314, 3: 46.8-54.0 mm, and USNM 
230243, 1: 49.7 mm, collected with holotype. 


DiaGnosis.— Thryssocypris smaragdinus 1s 
most readily distinguished from 7. tonlesapen- 
sis, 1ts only congener, by much larger head, larger 
eye, larger pharyngeal bone, and fewer anal fin 
rays, scales, and vertebrae. These and other 
quantitative characters differing in the two species 
are summarized in Table |. In 7. smaragdinus, 
anal fin origin on a vertical with dorsal fin origin 
(versus anal fin origin far in advance of dorsal 
fin origin in 7. tonlesapensis). Snout tip projects 
strongly beyond upper jaw in four of the five 
specimens of 7. smaragdinus (including the ho- 
lotype), thus differing strikingly from 7. tonle- 
sapensis, in which it projects but very slightly. 
In one paratype of 7. smaragdinus (USNM 
230243, 49.7 mm), however, the snout tip pro- 
jects little more than in 7. tonlesapensis. Some 
additional differences between the two species 
are indicated in the color descriptions below and 
in the osteological account following. 

In life 7. smaragdinus are brilliant emerald 
green on the upper half of the head and body 
and bright silvery below. Preserved specimens 
exhibit a wide longitudinal band, narrowest an- 


teriorly, extending from head to caudal fin, and 
lying entirely in dorsal half of body. Middle of 
caudal peduncle and caudal fin base with dark 
round spot level with longitudinal band. Mela- 
nophores absent from all fins except for a few 
small scattered ones on interradial membranes 
of dorsal and caudal fins. Melanophores almost 
entirely absent on ventral half of body. Dorsal 
half of body with numerous fine melanophores 
or chromatophores in addition to those of lon- 
gitudinal band, but not forming noticeable pat- 
terns such as rows parallel to posterior margin 
of scales. Dorsal midline of body with two or 
three thin longitudinal rows of melanophores. 
Dorsal surface of head, especially dorsal to nasal 
organs, deeply pigmented with numerous large 
melanophores. Inner surface of opercle dusky. 

Gut contents of 53.2-mm paratype comprise 
moderately numerous triturated remains of 
aquatic coleopteran and dipteran larvae, and 
some adult winged dipterans. No other food items 
observed. 

EtyMoLoGy.—From the Greek smaragdinos, 
emerald green. 


Thryssocypris tonlesapensis, new species 
(Figures 2, 3b, 10a) 

Ho.Lotyre.—MNHN 1982-1032, 50.6 mm, Prek Tamen, at 
or near Snoc Trou, Kampuchea, 9 Nov. 1961. 

PARATYPES.—MNHN 1982-1033, 3: 44.7—5S0.3 mm, MHNG 
2119.63-64, 2: 45.5—46.0 mm, CAS 50946, 2: 48.4-52.1 mm, 
collected with holotype; MNHN 1982-1034, 1: 52.3 mm, Prek 
Tasom, at or near Snoc Trou, Kampuchea, June 1961; UMMZ 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 


dermosphenotic 


nasal frontal 


ethmoid 


lateral ethmoid 


Ficure 5. 


210277, 1: 43.4 mm, Mekong River at w end of Con Phong 
(or Con Ho?) Island 2-5 km w of My Tho, Vietnam, 18 July 
1974: UMMZ 210278, 1: 45.7 mm, Mekong River at Con Ho 
Island, My Tho, Vietnam, 19 June 1974; UMMZ 210279, 58: 
20.4—60.1 mm, fish market at Vinh Long, Vietnam, 22 June 
1974: UMMZ 210280, 36: 29.2-63.8 mm, fish market at Can 
Tho, Phong Dinh Province, Vietnam, 23 June 1974. 


DiAGnosis.—Characters distinguishing 7. 
tonlesapensis from T. smaragdinus are given in 
the diagnosis of 7. smaragdinus, in Table 1, in 
the description of coloration and other remarks 
below, and in the osteological account following. 
The species is immediately distinguished from 
T. smaragdinus by its much smaller head, more 
numerous scales (44—46 in lateral line series ver- 
sus 36-39), and more numerous anal fin rays 
(15%-16'2 versus 12'2-14'2). The very striking 
difference in size of the pharyngeal jaws of ton- 
lesapensis (Fig. 4), the linear dimensions of which 
are only about half as great as in 7. smaragdinus, 
is much greater than would be expected if this 
difference were due only to the difference in head 
size, since the head is only about 20% longer in 
T. smaragdinus. 

Coloration of live 7. tonlesapensis has not been 
observed. Specimens observed in fish markets 
by W. J. Rainboth were entirely silvery. Color 
pattern of preserved specimens is similar in its 
basic features to that of 7. smaragdinus, includ- 
ing the longitudinal band and basicaudal spot, 
but differing in some details: longitudinal band 
lower, nearly midlateral in position (confined to 
dorsal half of body in 7. smaragdinus); mela- 


sphenotic 


147 


posttemporal 


ex trascapular : 
P supracleithrum 


exoccipital 


epiotic 
pterotic 


parietal 


Thryssocypris smaragdinus, 53.2 mm, cranium (dorsal view). 


nophores on dorsal half of body tending to form 
rows parallel to posterior margins of scales; and 
inside of opercle clear instead of dusky. 

Guts of numerous specimens are moderately 
to very full of insects, mostly aquatic larvae; no 
other food items observed. 

EtyMoLoGy.— From Tonle Sap, the enormous 
permanent backwater of the lower Mekong, into 
which the Prek Tamen and Prek Tasom flow. 


OSTEOLOGY 


Although the two species of Thryssocypris dif- 
fer strikingly in skull size and there are obvious 
differences in the axial skeleton and median fin 
skeletons related to differences in vertebral num- 
ber and anal fin position and ray number, their 
osteology is very similar in most respects (Fig. 
4). Some osteological differences between the two 
species are noted in the following account, which 
is based on both species, even though the draw- 
ings (except Fig. 10a) are of 7. smaragdinus. 

CRANIUM (Figs. 5-8, 10).—Roof of cranium 
entire, without fontanel, strongly convex trans- 
versely for its entire length. Frontals with nar- 
row, flangelike, nearly horizontal lateral margins, 
and a transverse shallow depression or groove 
overlying tectum cranii or epiphyseal bar. Sphe- 
notic as well as pterotic bones contribute sub- 
stantially to cranial roof, with moderately de- 
veloped sphenotic and pterotic projections or 
spines. Ethmoid (or supraethmoid) very large, 


148 


frontal 


orbitosphenoid 
ethmoid 


preethmoid 


lateral ethmoid 


pterosphenoid 


—l 


FIGuRE 6. 


completely covering mesethmoid so that it does 
not contribute to dorsal surface of ethmoid re- 
gion. Preethmoids cartilaginous. Dilatator fossae 
indistinct. Posttemporal fossae absent. 

In most cyprinids the cranium tends to be dor- 
sally flattened, or even transversely concave. In 
rasborines and bariliines the cranium tends to 
be barrel-shaped (Gosline 1975), with a convex 
dorsum, as in 7hryssocypris. The sphenotic is 
usually entirely or almost entirely excluded from 


optic foramen 


orbitosphenoid 

lateral ethmoid 

ethmoid 
prevomer \ 


pterosphenoid 
sphenotic 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


sphenotic 


dermosphenotic 
parietal 


supraoccipital 


epiotic 


exoccipital 


asteriscus 


parasphenoid pigelic 


basioccipital 


Thryssocypris smaragdinus, 53.2 mm, cranium (lateral view). 


the dorsal roof of the cranium, a noted exception 
occurring in Esomus (Ramaswami 1955), which 
in other respects differs very much from 7hrys- 
socypris. In Cyprinidae the ethmoid is usually 
much shorter than in 7hryssocypris, broader than 
long, and with its anterior margin broadly and 
deeply indented medially to receive the kineth- 
moid (7Thryssocypris has only a very small in- 
dentation anteriorly; see Fig. 5). 

In chelines (Howes 1979) the cranium is more 


prootic —_pterotic 


exoccipital 


basioccipital 


F ee Zz \ “aa a - ‘ 
Wa Lp oe. ve) ticatory plate 
preethmoid W : By 3 mastica 
NWF > a 
mesethmoid = cre a 
nasal frontal Na, 
parasphenoid hyomandibular subtemporal fossa 
= fossa 
Ficure 7. Thryssocypris smaragdinus, 53.2 mm, cranium (ventral view). 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 149 


epiotic 


pterotic 


exoccipital 


basioccipital 


FiGure_ 8. 


or less flat or even concave dorsally, the mes- 
ethmoid forms a shelflike projection extending 
anteriorly to the ethmoid, the anterior half of the 
parasphenoid is oriented at an angle of about 20° 
to 30° from the horizontal, the supraoccipital 
crest tends to be dorsal in position, and the oc- 
ciput is elongated and gently sloping. In all of 
these respects chelines differ from 7hryssocypris. 

Jaws (Figs. 9-11).—Jaws elongate, upturned, 


supraoccipital 


lateral occipital 
fenestra 


foramen magnum 


Cavum Sinus imparis 


aortic canal 


Thryssocypris smaragdinus, 53.2 mm, cranium (occipital view). 


and very narrow. Premaxilla only half as long as 
maxilla, maxilla alone forming margin of pos- 
terior half of upper jaw (maxilla entirely excluded 
or almost entirely excluded from gape in all other 
cyprinids). In 7. smaragdinus maxilla with two 
ascending or ethmoid processes, each with a car- 
tilaginous cap, anterior process moderately elon- 
gate or pedicellate, posterior process short (Fig. 
11); in 7. tonlesapensis maxilla with only a single 


infraorbitals 1-5 


/ 
premaxilla 


maxilla 
dentary 


quadrate 


preopercle 


FIiGuRE 9. 


suprapreopercle 


subopercle 


interopercle 


Thryssocypris smaragdinus, 53.2 mm, jaws and facial bones (lateral view). 


150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


premaxilla 


palatine 


infraorbital 1 


lateral ethmoid 


nasal 


FiGur_e 10. 
53.2 mm, kinethmoid bone (dorsal view). 


ethmoid process, short and broad based (non- 
pedicellate), and with a cartilaginous cap (Fig. 
10). Anterior half of maxilla with a medially di- 
rected shelf arising from its dorsal margin, coex- 
tensive with and largely overlapping lateral sur- 
face of premaxilla. Maxilla with a slender, 
posteriorly curved, broad-based projection aris- 
ing near middle of its dorsal surface and extend- 
ing medially to infraorbital number one, to which 
it is firmly attached by connective tissue, thus 
contributing to immobilization of upper jaw. Only 
posterior half of maxilla free from snout and 
capable of limited movement. Posterior portion 
of maxilla, separated from infraorbitals by a shal- 
low groove, simple, elongate, and flexible; ex- 


dentary 


Meckel’s cartilage 


coronomeckelian 


AE=_==" 


FiGure 11. 


SAE: 


kinethmoid 
prevomer 


preethmoid 


ethmoid 


frontal 


a, Thryssocypris tonlesapensis, 46.0 mm, ethmoid region of skull (dorsal view); b, Thryssocypris smaragdinus, 


tends to proximal end of dentary. Lower jaw 
slightly shorter than upper jaw, elongate and very 
narrow, completely included within gape when 
mouth is closed. Symphysis of lower jaw without 
pronounced dorsal or ventral knobs, although 
vestigial or rudimentary dorsal symphyseal knob 
may be present, especially in 7. tonlesapensis. 
Dorsal margin of dentary with a slight elevation 
just behind symphysis in 7. tonlesapensis; in T. 
smaragdinus elevation absent or barely notice- 
able. Dentary and angular with coronoid pro- 
cesses short, vertically oriented, and separate, 
that of dentary arising very near proximal end 
(coronoid process of dentary frequently very large, 
usually arising near middle of dentary, and not 


maxilla 


coronoid 
process 


retroarticular 


Thryssocypris smaragdinus, 53.2 mm, jaws (medial view). 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 151 


preopercle 


FiGure 12. 


separate from angular in most cyprinids; angular 
without distinct coronoid process in most cyp- 
rinids). Free ventromedian margins of dentary 
nearly straight for entire length, those of opposite 
sides in contact or closely approximated (ven- 
tromedian margin of dentary frequently highly 
modified in shape in cyprinids, as in Danio and 
its relatives, and dentaries of opposite sides usu- 
ally more or less widely separated in cyprinids, 
including genera with comparably elongate and 
narrow lower jaws such as Macrochirichthys, 
Salmostoma, and Securicula). 

The poorly developed condition of the single 
ethmoid process on the maxilla of 7. tonlesa- 
pensis 1s suggestive of the condition in chelines, 
but because the jaws of 7hryssocypris are oth- 
erwise very unlike those of chelines and because 
T. smaragdinus has well-developed ethmoid 
processes, this resemblance is probably due to 
independent reduction (possibly related to small 
size of the head as well as nonprotrusibility of 
the jaws). 

Development of large dorsal symphyseal knobs 
on each dentary (and sometimes almost equally 
prominent ventral symphyseal knobs) is char- 
acteristic of most chelines, and small dorsal sym- 
physeal knobs occur in many cyprinids, includ- 
ing Rasbora and Barilius. In forms with dorsal 
symphyseal knobs the symphysis of the upper 
jaw is frequently indented for their reception; 


symplectic 


hyomandibular 


metapterygoid 


mesopterygoid 


Palatine 


Thryssocypris smaragdinus, 53.2 mm, suspensorium (lateral view). 


there is no comparable indentation in the upper 
jaw of Thryssocypris. 

INFRAORBITAL AND NASAL BONES (Fig. 9).— 
Infraorbital series complete, with five large, la- 
mellar infraorbital bones each bearing a simple 
bony tubule for infraorbital laterosensory canal 
and, together with supraorbital bone, forming an 
almost complete ring around eye. Posterior bor- 
der of supraorbital indented where concave an- 
terior margin of infraorbital 5 fits snugly against 
it; slender, pointed, anteroventral projection of 
supraorbital nearly contacts dorsal margin of in- 
fraorbital 1. Nasal bone moderately elongate, with 
a narrow lamellar portion on either side of bony 
tubule for nasal laterosensory canal. 

Among Asian minnows with elongate, up- 
turned jaws and compressed bodies, a complete 
infraorbital series in which all five infraorbitals 
have well-developed lamina is typical of rasbo- 
rines and bariliines and unusual in chelines (Gos- 
line 1975). In Cyprinidae the primitive number 
of infraorbitals, exclusive of the dermosphenotic 
(sometimes counted as an infraorbital), is five. 
Occasionally a greater number occurs, but this 
is usually (perhaps invariably) due to fragmen- 
tation of more or less tubular infraorbitals in 
which the lamellar component is greatly reduced 
or absent. 

SUSPENSORIUM (Fig. 12).— Palatine arch nearly 
horizontal. Hyomandibular and preopercle ver- 


152 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


urohyal 


dorsal hypohyal 


ceratohyal 


ventral hypohyal 


epihyal 


branchiostegal rays 1-3 


Ficure 13. Thryssocypris smaragdinus, 53.2 mm, hyoid bar and urohyal (lateral view). 


tical. Lateral abductor process of hyomandibular 
dorsally directed, originating just below hyo- 
mandibular articulation with cranium. Meta- 
pterygoid-quadrate fenestra absent or greatly re- 
duced and almost entirely occluded by peculiarly 
expanded distal end of symplectic. Anteroven- 
tral limb of preopercle extremely short. Quadrate 
nearly square, with strong posteroventral exten- 
sion seen in many or most cyprinids, and without 
a foramen immediately posterior to articular fac- 
et for lower jaw. Ectopterygoid apparently absent 
in 7. smaragdinus, present but reduced in T. 
tonlesapensis. Palatine bone with a well-devel- 
oped posteroventral extension underlying lateral 
ethmoid articular facet. An elongate, continuous 
palatine cartilage connects palatine bone to me- 
sopterygoid anteriorly and to metapterygoid and 
quadrate posteriorly. At its anterior end, between 
palatine bone and mesopterygoid, this cartilage 
forms a large, saddlelike facet articulating with 
ventral surface of lateral ethmoid. Opercle gen- 
eralized in shape, deeper than long, its posterior 
margin rounded, dilatator process weakly de- 
veloped. Suprapreopercle short, tubular, fused to 
anterodorsal corner of opercle. 

In chelines the palatine bar is usually upturned 
like the strongly upturned jaws, and is thus 
obliquely oriented. Many cyprinids, including 
some chelines and bariliines, have a well-devel- 
oped metapterygoid-quadrate fenestra, and most 
have the quadrate with a well-developed pos- 
teroventral process, a preopercle with a promi- 
nent anteroventral portion, and a large, broad 


ectopterygoid. In 7. smaragdinus the elongate 
ventral portion of the mesopterygoid extends 
narrowly below the palatine cartilage where the 
ectopterygoid normally occurs, but examination 
with direct and transmitted light failed to reveal 
an ectopterygoid. In 7. tonlesapensis the meso- 
pterygoid does not extend ventral to the palatine 
cartilage, and a very thin, elongate ectopterygoid 
is present. Most chelines (Howes 1979) and Lu- 
clobrama (Howes 1978) have a foramen in the 
quadrate immediately posterior to its articula- 
tion with the lower jaw. Chelines and some other 
cyprinids tend to have a stout, fingerlike dilatator 
process forming the elevated anterodorsal corner 
of the opercle. Some cyprinids have a free su- 
prapreopercle and some lack this element alto- 
gether. Fusion of the suprapreopercle with the 
opercle occurs frequently but is known only in 
the Cyprinidae. The cyprinid suprapreopercle is 
always a simple bony tube enclosing the upper- 
most portion of the preopercular laterosensory 
canal, never with a laminar portion (sometimes 
greatly enlarged) as in characoids. 

Hyorp BAR AND UROHYAL (Fig. 13).—Hyoid 
bar generalized for Cyprinidae, with three bran- 
chiostegal rays as in all members of the family, 
and no unusual features. First branchiostegal ray 
articulated to ventral surface of ceratohyal, which 
has a notch in margin at point of articulation; 
second branchiostegal ray broadly articulated to 
lateral face of ceratohyal; third broadly articu- 
lated to lateral face of epihyal. Basihyal dorso- 
ventrally compressed, its anterior third cartila- 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 153 


basibranchials 1-2 <f/ | 


= 


SS 


Ficure 14. Thryssocypris smaragdinus, 53.2 mm, gill arches and pharyngeal jaws (dorsal view, with upper half of arches 


on left side deflected to reveal ventral surface). 


ginous and with a broadly rounded anterior 
margin (basihyal elongate and narrow for its en- 
tire length in at least some chelines). Interhyal 
tubular and moderately elongate (flattened and 
laterally expanded in some chelines and cul- 
trines). Urohyal with posteroventral process not 
greatly expanded, its distal end simple in 7. 
smaragdinus and with a very small notch in 7. 
tonlesapensis. (Urohyal greatly modified in some 
bottom dwelling cyprinoids including cyprinids; 
posteroventral process very deeply forked in Lu- 
ciobrama and at least some chelines.) 

Gi_L ARCHES AND PHARYNGEAL JAws (Fig. 
14).—Gill arches (main features described above 
under generic diagnosis) basically similar in 7. 
smaragdinus and T tonlesapensis. Narrow prox- 
imal portion of ceratobranchials not quite so long 
as in 7. smaragdinus, but equally strongly nar- 
rowed and with peculiar cartilaginous proximal 
ends as large as in 7. smaragdinus. Basibran- 
chials two in 7. smaragdinus, three in T. ton- 


lesapensis, extremely slender in both species. 
Ceratobranchial 5 (tooth-bearing pharyngeal 
bone) uniformly slender, its length more than 
five times its width, gracefully arched. External 
ala elongate but extremely narrow, its origin 
marked by a small, strongly angular projection 
opposite base of middle tooth in major tooth 
row. Ventral edentulous limb slightly shorter, and 
dorsal edentulous limb slightly longer, than den- 
tigerous portion. Symphyseal half of ventral 
edentulous limb abruptly narrowed and set at 
angle to rest of bone, so its orientation is nearly 
horizontal. Pharyngeal teeth uncinate (conical 
with recurved tips), in two rows; those of inner 
or major row uniformly decreasing in size from 
front to back, those of minor or outer row sub- 
stantially smaller than those of major row. 
Like most cyprinids 7/ryssocypris has but two 
infrapharyngobranchials, here interpreted as in- 
frapharyngobranchials 2-3. They exhibit the 
characteristic two-plus-two relationship with 


154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


np2-3 


~ claustrum 
FON 


scaphium a, eS 
CR 
4 xe <i 


intercalarium 


basioccipital 


Ficure 15. 


tripus 


neural supraneural 
complex 


A neural 
spines 


prezygapophysis 


rib 


Os suspensorium 


Thryssocypris smaragdinus, 53.2 mm, Weberian apparatus (lateral view) (1pc1,2 = lateral processes of centrum 


1 and centrum 2; np2,3 = neural plates of centrum 2 and centrum 3). 


epibranchials 1-2 and 3-4 which seems to be 
characteristic of all cyprinoids. Epibranchials 1- 
2 are united by cartilaginous joints to the distal 
end of infrapharyngobranchial 2, and epibran- 
chials 3-4 are similarly joined to infrapharyn- 
gobranchial 3. Thus the proximal ends of infra- 
pharyngobranchials | and 2 approximate each 
other, and there is a slight but distinct gap be- 
tween epibranchials 1-2 and epibranchials 3-4. 
This peculiar orientation, which seems not to 
have been noted previously, has been observed 
without exception in numerous cyprinoids ex- 
amined by us. Although the orientation is highly 
specialized compared to the primitive one-to- 
one relationship between the infrapharyngobran- 
chials and epibranchials observed in characoids 
and most lower teleosts, it is apparently primi- 
tive for cyprinoids. A curiously similar two-plus- 
two arrangement is present in numerous silu- 
roids examined, including Diplomystes, but the 
relationships of the infrapharyngobranchials to 
the epibranchials appears not to be exactly the 
same. Further investigation should be done to 
determine whether this condition in cyprinoids 
and siluroids indicates a common ancestor for 
these two groups. In any event this specialized 
condition 1s too widespread among cyprinids to 
be of any use in assessing phyletic relationships 
of Thryssocypris. 

In some cyprinoids (mainly bottom feeders, 


such as Labeo and Osteochilus, with inferior, 
suctorial mouths) the basibranchial bones appear 
to be shaped and articulated in such a fashion as 
to permit them to slide over and under each 
other, thus indicating that the ventral half of the 
branchial basket is capable of anteroposterior 
contraction and expansion. In Thryssocypris the 
elongate, slender, and straight basibranchial bones 
are firmly joined end to end, indicating that the 
branchial basket is incapable of such contraction 
and expansion movements. 

WEBERIAN APPARATUS (Fig. 15).—Similar gen- 
erally to Weberian apparatus of Opsariichthys 
(Fink and Fink 1981; Fig. 14). Neural complex 
vertical (not strongly posteriorly sloped as in 
many chelines and cultrines), widely separated 
from occiput. Occiput attached to anterodorsal 
surface of Weberian apparatus by a thin median 
strip of cartilage extending from posterodorsal 
median margins of exoccipitals to claustrum and 
neural process of centrum number two. Lateral 
process of centrum 2 exceptionally elongate and 
strongly curved posteriorly, extending dorsally 
to tripus and laterally as far as major curvature 
of fully formed pleural rib of vertebra 5. Fourth 
pleural rib and parapophysis (bearing os suspen- 
sorium) without a prominent anterodorsally pro- 
jecting lateral process (present in Opsariichthys). 
Intercalarium simple, slender, and elongate, hor- 
izontally oriented, without ascending or articular 


ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS 155 


supracleithrum 


mesocoracoid 
cleithrum 


postcleithrum 


scapula 


coracoid 


Thryssocypris smaragdinus, 53.2 mm, pectoral 


Ficure 16. 
girdle, left half (lateral view). 


processes. Centra 2 and 3 fused to each other, at 
least ventrally (centra 2-3 separate in Opsari- 
ichthys but partially or completely fused in many 
cyprinids). 

Endochondral union of Weberian apparatus 
with cranium is characteristic of cyprinoids and 
is not found in other ostariophysans (Fink and 
Fink 1981). Reduction and loss of ascending and 
articular processes of intercalarium in other os- 
tariophysans, especially siluroids, is reported by 
Fink and Fink (1981:328). In Opsariichthys rel- 
atively large gaps occur between neural arches 
2-3 and the vertebral column, and between neu- 
ral arches 3 and 4; these gaps are greatly reduced 
in Thryssocypris (that between neural arches 3 
and 4 is completely occluded). The element here 
termed neural arch of centrum 2 is the anterior- 
most supraneural according to Fink and Fink 
(1981). 

Fusion of centra 2 and 3 is possibly a primitive 
character for all Cyprinidae. We propose a hy- 
pothesis that this condition evolved only once 
and that the mosaic distribution of cyprinid gen- 


era with fused and with separate centra 2-3 is 
due to repeated secondary failure to fuse and 
reversion to the fused condition, rather than to 
truly independent evolution of fusion between 
these centra in phyletically diverse lines (the hy- 
pothesis seemingly favored by Greenwood et al. 
[1966:385] and Fink and Fink [1981:331]). Cen- 
tra 2-3 are fused in the primitive siluroid Di- 
plomystes, centra 2-4 and sometimes also 5 in 
all other siluroids (Hassur 1970). Whether fusion 
of centra 2-3 evolved independently in siluroids 
or indicates their phyletic relationship to cypri- 
noids has not been resolved satisfactorily (com- 
pare Roberts 1973 with Fink and Fink 1981). 
The centra of the Weberian apparatus are all 
separate in all characoids, gymnotoids, and gon- 
orynchs. In any event this character is unlikely 
to be relevant in considerations of the closer phy- 
letic relationships of Thryssocypris. 

PECTORAL GIRDLE (Fig. 16).—Pectoral girdle 
morphologically generalized, slender, with a sin- 
gle extrascapular (two extrascapulars in some 
cyprinids), dorsal and anteroventral limbs of 
cleithrum slender, postcleithrum moderately 
elongate (slightly more elongate in 7. tonlesa- 
pensis than in T. smaragdinus). Coracoid with a 
strongly developed lateral shelf, but not ventrally 
expanded; coracoid foramen broad. (Coracoid is 
ventrally expanded, sometimes enormously so, 
in most chelines and cultrines. In some, coracoid 
foramen reduced or even occluded.) In chelines 
(Howes 1979), postcleithrum frequently (al- 
ways?) arclike, very slender and elongate, and 
with distal end projecting downwards, quite un- 
like the postcleithrum in Thryssocypris. In 
Thryssocypris the pectoral fin, although slightly 
larger proportionately in 7. tonlesapensis than 
in 7. smaragdinus, is not notably enlarged, nor 
is the simple outer ray thickened or otherwise 
modified. The pectoral axial flap (attached to the 
postcleithrum) is simple, moderately elongate, 
and not overlain by peculiarly shaped scales. In 
all of these respects the pectoral fin of chelines 
and cultrines tends to be modified. 

Petvic GIRDLE (Fig. 17).— Pubic bone deeply 
bifurcate anteriorly (as in most cyprinoids); lat- 
eral pubic projection nearly twice as long as me- 
dial. Ischiac process highly modified, with ex- 
tremely elongate anterior and _ posterior 
projections. In 7. smaragdinus anterior ischiac 
projection about half as long as posterior pro- 
jection; in 7. tonlesapensis anterior projection 


156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


eee 


ischiac process 


Ficure 17. Thryssocypris smaragdinus, 53.2 mm, pelvic girdle (ventral view). 


twice as long as posterior one, extending ante- 
riorly as far as distal end of medial pubic pro- 
jection. Origin of ischiac process opposite distal 
end of pleural rib of vertebra number 9 in 7. 
smaragdinus, and of pleural rib of vertebra 11 
or 12 in T. tonlesapensis. 

AXIAL SKELETON. —Supraneurals 4-6 in num- 
ber, commencing immediately posterior to neur- 
al complex of Weberian apparatus (not greatly 
enlarged or articulated to each other as in some 
chelines). In 7. smaragdinus vertebrae 5-20 and 
in 7. tonlesapensis 5-16 with enlarged prezyga- 
pophyses. 7hryssocypris smaragdinus with well- 
developed dorsal ribs attached to anterior half 
of centrum on vertebrae 11-24; 7. tonlesapensis 
without dorsal ribs (dorsal ribs unreported 1n any 
Cyprinidae until now). Pleural ribs of vertebrae 
5-10 with broad, laminar proximal ends at- 
tached to comparably broad parapophyses; re- 
maining pleural ribs with proximal ends slender, 


proximal radial 


weakly attached to slender parapophyses. In 7. 
smaragdinus proximal portion of pleural ribs on 
vertebrae 6-7 with spurlike, posteriorly directed 
projections (absent in 7. ton/esapensis). 

The counts of abdominal, caudal, and total 
vertebrae in 7hryssocypris (Table 1) are close or 
identical to counts reported in several species of 
Barilius by Howes (1980: table 1). 

DorSAL AND ANAL Fins (Fig. 18).— Dorsal and 
anal fin rays uniformly slender, nonserrate, first 
two dorsal and first three anal fin rays simple. 
Last complete pterygiophore bears two (counted 
as one and a half) fully formed, branched rays; 
bases of ray halves of anterior of these two rays 
overlap and lie externally to ray halves of pos- 
terior ray. 

In the anal fin the bifurcate terminal radial (see 
generic diagnosis) is the medial radial of the last 
pterygiophore, which is complete. In the dorsal 
fin the situation is more complicated, and the 


Ficure 18. 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 157 


homology of the bifurcate terminal radial with 
the proximal, medial, and distal radials is un- 
clear. In 7. smaragdinus 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. —Intermuscular bones 
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 7hryssocypris is not likely to be 
helpful in assessing its relationships to other cyp- 
rinids. 

It should be noted that previous works on os- 


uroneural 


ESSY 


ural centrum 


hypurapophysis 
parhypural 
hemal spine 
—= hypurals 1-6 
FiGure 19. Thryssocypris smaragdinus, 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 7hryssocypris 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 7hryssocypris that 
seem most unusual or highly derived and there- 


158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 11 


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 (Teleoste1). Zool. J. Linn. Soc. 72(4): 
297-353. 

Gos.ine, 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. 

Hassur, 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 Luciobrama macrocephalus (Lacepéde). Bull. 
Brit. Mus. (Nat. Hist.), zool. ser. 34(1):1-64. 

1979. Notes on the anatomy of Macrochirichthys 

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):129-198. 

LEKANDER, B. 1949. The sensory line system and the canal 
bones in the head of some Ostariophysi. Acta Zool. 30:1— 
Si. 

LUNDBERG, J. G., AND J. N. BAskin. 1969. The caudal skel- 
eton of the catfishes, order Siluriformes. Amer. Mus. Nov. 
no. 2399, 49 pp. 

Monon, T. 1968. Lecomplex urophore des téléostéens. Mém. 
Inst. fr. Afr. noire 81:1-705. 

Ramaswaml, 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. 

Roperts, T. R. 1969. Osteology and relationships of chara- 
coid fishes, particularly the genera Hepsetus, 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. 


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PROCEEDINGS 
OF THE JUL 2% 1984 | 
CALIFORNIA ACADEMY OF SCIENCES. 
|; V¥OOGS Hole, hilas i 


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 


Asstract: 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), NV. campbelli (Mount Baker, Whatcom County, Washington), N. 
wallowae (West Fork Wallowa River, Wallowa County, Oregon), N. jeffreyi (South Fork McCoy Creek, 
Harney County, Oregon), NV. haida (Mount Needham, Graham Island, Queen Charlotte Islands, British 
Columbia), N. louiseae (Skedans, Louise Island, Queen Charlotte Islands, British Columbia), N. gebleri 
albimontis (Birch Creek, Mono County, California), N. Jabontei (West Fork Wallowa River, Wallowa County, 
Oregon), N. calva (Mount Baldy, Apache County, Arizona), N. sierrablancae (Sierra Blanca, Lincoln County, 
New Mexico), N. piute sevieri (Parowan Creek, Iron County, Utah), N. steensensis (South Fork McCoy 
Creek, Harney County, Oregon), and WN. 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 = N. 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 = N. trifaria trifaria LeConte. 


INTRODUCTION 


July 12, 1984 


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 13 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] 


160 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 limited to little 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 1136 
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 94118; D. H. Kavanaugh, H. B. Leech. 

CNC—Canadian National Collection of Insects, Biosystem- 
atics Research Institute, Ottawa, Ontario KIA OC6; 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. 

JRLa—J. R. La Bonte, 710 NW 11th, 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—Miichigan State University, East Lansing, Michigan 
48823: R. L. Fischer. 


OSUC-— Ohio State University, Columbus, Ohio 43210; C. A. 
Triplehorn. 

PJJo—P. J. Johnson, 1408 28th Street SE, Auburn, Washing- 
ton 98002. 

QCIM— Queen Charlotte Islands Museum, Skidegate, British 
Columbia VOT 1S0; 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. 

UASM-— University of Alberta, Strickland Museum, Edmon- 
ton, Alberta T6G 2E3; G. E. Ball. 

UMMZ-— University of Michigan, Ann Arbor, Michigan 48104; 
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). 

Ho .orype, 4, 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 label]/““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 2), also deposited in CAS. 

Tyre-LocaLiry.—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 


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. 15) 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). 

Ho totype, 4, 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- 
bellin. sp. det. D. H. Kavanaugh 1983” [red label]/ “California 
Academy of Sciences Type No. 14339.” PARATYPES: two (one 
6 and one 2) deposited in CAS and CNC. 

Type-Locatiry.—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, humeri (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- 


161 


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.] 
@EI@AS): 


Nebria wallowae, new species 
(Figures 4, 17, 24, 31) 

Ho ortypee, 4, 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. 14347.” 
PARATYPES: 87 (36 and 51 2), deposited in CAS, CNC, JRLa, 
UASM, and USNM. 

Type-Loca.ity.— West Fork Wallowa River, 2070-2130 m, 
Wallowa Mountains, Wallowa County, Oregon. 


DIAGNostTIC 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. 31). 

DERIVATION OF TAXON NAME.—This species 
is named for the Wallowa Mountains. 


162 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 


OreEGON: 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 jeffreyi, new species 
(Figures 5, 31) 

Ho .orypee, 6, 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 jeffreyi n. sp. det. D. H. Kavanaugh 1983” [red 
label]/ ‘“‘California Academy of Sciences Type No. 14342.” 
PARATYPES: 35 (11 6 and 24 2), deposited in CAS, JRLa, PJJo, 
UASM, and USNM. 

Type-Loca.ity.—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 
Mountains of Oregon (Fig. 31). 

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 


OreEGon: 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) 

Ho .orype, 6, 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 6 and 83 2), deposited in BCPM, CAS, 
CNC, QCIM, UASM, and USNM). 

Type-Locauity.— 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) 
Ho orypre, ¢, in CAS, labelled: “B. C., Q. C. I. Louise Is., 


KAVANAUGH: NEW NEARCTIC NEBRIA 


Skedans 11.VI.1981 R. A. Cannings’”’/ ‘““Holotype Nebria lou- 
iseae n. sp. det. D. H. Kavanaugh 1983” [red label]/ “California 
Academy of Sciences Type No. 15005.” PARATyPes: four 2, 
deposited in BCPM, CAS, and CNC. 

Type-Locaity.—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 11.0 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) 

Ho ortype, 6, 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 1983” [red label]/ “‘Cal- 
ifornia Academy of Sciences Type No. 14340.” PARATyYPEs: 
four (two 4 and two 2), also deposited in CAS. 

Type-Loca.itry.— 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- 


163 


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. 18) 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” (=a/bus) 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) 

Ho torype, 4, 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 6 and 28 2), deposited in CAS, JRLa, UASM, 
and USNM. 

Type-Loca.ity.— West Fork Wallowa River, 2040-2190 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 12.0 
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. 


164 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) 

Ho .ortyPe, 4, in CNC, labelled: ‘Ariz: Apache Co Mt. Baldy, 
10-11000’ SW of Springerville 13.VII.79, S & J Peck spruce- 
fir forest’’/ ‘Holotype Nebria calva n. sp. det. D. H. Kavanaugh 
1983” [red label]. PARATyPEs: 10 (7 6 and 3 9), deposited in 
CAS and CNC. 

Type-Loca.iry.— Mount Baldy (SW of Springerville), 3050— 
3350 m, Apache County, Arizona. 


DIAGNOSTIC COMBINATION.— Head moderate 
in width and size, dark, with a pair of pale par- 
amedial spots on vertex; pronotum (Fig. 10) with 
midlateral and basolateral setae present; elytra 
with faint but distinct metallic (violet) reflection, 
elytral silhouette subovoid, markedly narrowed 
basally, intervals moderately flat; median lobe 
of male (Fig. 19) with apex straight in ventral 
aspect; bursa copulatrix of female (Fig. 26) with 
very small bursal sclerite; specimen from locality 
in eastcentral Arizona (Fig. 32). 

DERIVATION OF TAXON NAME.—The species 
epithet is formed from the Latin word for “‘bald”’ 
(=calvus), in reference to the type locality, Mount 
Baldy. 

GEOGRAPHICAL DISTRIBUTION.— Figure 32. 
Known only from Mount Baldy in eastern Ari- 
zona. I have studied specimens from the follow- 
ing locality: 


United States of America 


Arizona: Apache County, Mount Baldy (SW of Springerville 
[3050-3350 m}) [July] (11; CNC). 


Nebria sierrablancae, new species 
(Figures 11, 20, 27, 32) 
Nebria trifaria catenata; KAVANAUGH 1978:431 (in part). 
Ho.otyre, 4, in CNC, labelled: ‘““N. M. Lincoln Co. Sierra 
Blanca 10500’ 18.VII.1969 A. Smetana’’/ “Holotype Nebria 
sierrablancae n. sp. det. D. H. Kavanaugh 1983” [red label]. 
ParRATYPEs: 25 (10 6 and 15 2), deposited in CAS, CNC, and 
CUIC. 
Type-Loca.ity.—Sierra Blanca, 3200 m, Lincoln County, 
New Mexico. 


DIAGNOSTIC COMBINATION. — Body color uni- 
formly rufous; head relatively large in relation 


to pronotum, with a pair of pale paramedial spots 
on vertex; pronotum (Fig. 11) with lateral margin 
markedly sinuate basally, slightly to moderately 
angulate at middle, apical angles narrow and 
bluntly pointed, midlateral and basolateral setae 
present; elytra with faint metallic (violet) reflec- 
tion, elytral silhouette subovoid, narrowed ba- 
sally, intervals moderately flat; median lobe of 
male (Fig. 20) with apex deflected left laterally 
in ventral aspect; bursa copulatrix of female as 
in Fig. 27; specimen from locality in Capitan 
Mountains or Sierra Blanca of central New Mex- 
ico (Fig. 32). 

DERIVATION OF TAXON NAME.—This species 
is named for Sierra Blanca, the type locality. 

GEOGRAPHICAL DISTRIBUTION.—Figure 32. 
Known only from Sierra Blanca and the Capitan 
Mountains of central New Mexico. I have stud- 
ied specimens from the following localities: 


United States of America 


New Mexico: Lincoln County, Capitan (1; CUIC), Sierra 
Blanca (Sierra Blanca Ski Area [3200-3510 m]) [July] (25; 
CNC). 


Nebria piute sevieri, new subspecies 
(Figures 12, 21, 28, 32) 


Nebria trifaria trifaria, aactorum—ErRwIN AND BALL 1972:93 
(in part)—KAVANAUGH 1978:430 (in part). 


Ho .otype, 4, in CAS, labelled: ““U.S., Utah, Iron Co., Mar- 
kagunt Plateau, 13.5 mi. s. Parowan, Hwy. 143, Parowan Cr., 
9200’, 21 June 71 DH Kavanaugh & E A Martinko’’/ “D. H. 
Kavanaugh Collection” [orange label]/ ““71-224” [orange la- 
bel]/ “Holotype Nebria piute sevieri n. ssp. det. D. H. Kava- 
naugh 1983” [red label]/ “California Academy of Sciences Type 
No. 14344.” Paratypes: 230 (107 é and 123 2), deposited in 
ANSP, CAS, CNC, KSUC, MCZ, OSUO, UASM, and USNM. 
All specimens studied have been designated as paratypes ex- 
cept for the single (female) specimen from Clay Springs, Navajo 
County, Arizona. At present, I believe that this specimen is 
mislabelled. However, form of the bursa copulatrix in this 
specimen differs markedly from that in other females of N. 
piute sevieri. This may be a teratological example, or, if the 
specimen is correctly labelled, it may represent a distinct form 
not yet adequately sampled. My identification of the specimen 
as belonging to this subspecies is therefore tentative, pending 
additional fieldwork in Arizona. 

Type-Loca.ity. —Parowan Creek (13.5 miles S of Parowan), 
2800 m, Markagunt Plateau, Iron County, Utah. 


DIAGNOSTIC COMBINATION. — Body color uni- 
formly rufopiceous or black; head moderate in 
width and size in relation to pronotum, with a 
pair of pale paramedial spots on vertex; antennal 
scape markedly narrowed basally; pronotum (Fig. 
12) with lateral margin moderately sinuate ba- 
sally, rounded at middle, apical angles relatively 


KAVANAUGH: NEW NEARCTIC NEBRIA 


broad and rounded, midlateral and basolateral 
setae present; elytra without metallic reflection, 
elytral silhouette elongate, subovoid or nearly 
ovoid, narrowed basally, intervals moderately 
flat; median lobe of male (Fig. 21) very thick 
basal to apical orifice, with apex moderate in 
length and broad in lateral aspect, deflected left 
laterally in ventral aspect; bursa copulatrix of 
female (Fig. 28) with bursal sclerite large, narrow 
in dorsal aspect; specimen from locality in south- 
western Utah, east of Tushar Mountains and 
Midget Crest and west of Henry Mountains (Fig. 
32). 

DERIVATION OF TAXON NAME.—This subspe- 
cies is named for the Sevier River and Sevier 
Plateau, important physiographic features of the 
region occupied by members of this species. 

GEOGRAPHICAL DISTRIBUTION.—Figure 32. 
Known at present only from the montane region 
between Salina, Utah, and Cedar Breaks Na- 
tional Monument and east to the Boulder Moun- 
tains. I have studied specimens from the follow- 
ing localities: 


United States of America 


Uran: Garfield County, Cottonwood Peak (38.5 miles SW 
of Antimony at Cottonwood Creek [2440 m]) [July] (72; CAS), 
Mount Dutton (23.4 miles SW of Antimony at North Fork 
Deep Creek [3120 m]) [July] (18; CAS); Iron County, Cedar 
Breaks National Monument ([3200 m]) [June—Aug.] (36; CAS, 
OSUC), Cedar Canyon (Coal Creek [2650 m]) [June] (6; CAS), 
The Mammoth ([3050 m]) [July] (17; ANSP, CAS, CNC, KSUC, 
MCZ, UASM, USNM), Parowan Creek (13.5 miles S of Par- 
owan [2800 m]) [June] (10; CAS); Kane County, Long Valley 
Junction [Aug.] (12; CAS); Sevier County, Monroe Peak (8.6 
[2640 m] and 12.4 [2990 m] miles SE of Monroe) [July] (3: 
CAS), Mount Marvine (0.1 miles N of Johnson Valley Res- 
ervoir at Sevenmile Creek [2590 m]) [Aug.] (10; CAS); Wayne 
County, Bluebell Knoll (31 miles S of Torrey [2440-3050 m]) 
[July] (46; CAS, USNM). 


Doubtful Records: 


Unitep STATES OF AMERICA— Arizona: Navajo County, Clay 
Springs [Sep.] (1; CAS). 


Nebria steensensis, new species 
(Figures 12, 22, 29, 32) 


Ho totype, 6, in CAS, labelled: “U.S.A., Oregon, Harney 
County, Steens Mountains, South Fork McCoy Creek, 2390- 
2560m, 14 July 1982, Stop #82-22 D. H. & J. L. Kavanaugh 
colls.”/ “D. H. Kavanaugh Collection” [orange label]/ ““Ho- 
lotype Nebria steensensis n. sp. det. D. H. Kavanaugh 1983” 
[red label]/ ‘California Academy of Sciences Type No. 14345.” 
ParRAtTyPeEs: 106 (52 6 and 54 9), deposited in CAS, JRLa, PJJo, 
UASM, and USNM. 

Type-Locauity.—South Fork McCoy Creek, 2390-2560 m, 
Steens Mountains, Harney County, Oregon. 


165 


DIAGNOSTIC COMBINATION. — Body color uni- 
formly black; head moderate in width and size 
in relation to pronotum, with a pair of pale par- 
amedial spots on vertex; antennal scape slightly 
narrowed basally; pronotum (Fig. 13) with lateral 
margin very deeply sinuate basally, rounded at 
middle, apical angles relatively broad and round- 
ed, basal angles rectangular, midlateral and ba- 
solateral setae present; elytra without metallic 
reflection, elytral silhouette subovoid, narrowed 
basally, intervals moderately flat; median lobe 
of male (Fig. 22) moderate in thickness basal to 
apical orifice, with distinct, projected ridge on 
right lateral surface, apex deflected left laterally 
in ventral aspect; bursa copulatrix of female (Fig. 
29) with bursal sclerite small; specimen from lo- 
cality in Steens Mountains of Oregon (Fig. 32). 

DERIVATION OF TAXON NAME.—The species is 
named for the Steens Mountains. 

GEOGRAPHICAL DISTRIBUTION. —Figure 32. 
Known only from the Steens Mountains of south- 
central Oregon. I have studied specimens from 
the following localities: 


United States of America 


OreGon: Harney County, Fish Creek Gorge ([2190 m]) [July] 
(1; PJJo), Kiger Headwall ([2680 m]) [Aug.] (1; PJJo), Little 
Blitzen River ({2560 m]) [July] (16; PJJo), Pate Lake ([2260 
m]) [July] (1; PJJo), South Fork McCoy Creek ([2390-2560 
m]) [July] (88; CAS, JRLa). 


Nebria trifaria pasquineli, new subspecies 
(Figures 14, 23, 30, 32) 

Nebria trifaria coloradensis, ERWIN AND BALL 1972:96 (in part). 
Nebria trifaria trifaria, KAVANAUGH 1978:430 (in part). 


Ho torype, 6, in CAS, labelled: “Lefthand Cr., 5 mi. E. Ward, 
Colo. Bould. Co. 20 July 68”/ “D. H. Kavanaugh Collection” 
[orange label]/ ““Holotype Nebria trifaria pasquineli n. ssp. det. 
D. H. Kavanaugh 1983” [red label]/ “California Academy of 
Sciences Type No. 14346.” PAraAtypes: 407 (224 6 and 183 
2), deposited in ANSP, CArm, CAS, CNC, CUB, DEUN, EAMa, 
FMNH, KSUC, LACM, MCZ, MSU, ROM, RTBe, UAFA, 
UASM, UMMZ, USNM, UWEM, and ZMLS. 

Type-Loca.ity.—Lefthand Creek (5 miles E of Ward), Front 
Range, Boulder County, Colorado. 


DIAGNOSTIC COMBINATION. — Body color uni- 
formly black; head moderate in width and size 
in relation to pronotum, with a pair of pale par- 
amedial spots on vertex; antennal scape slightly 
arcuate, only slightly narrowed basally; prono- 
tum (Fig. 14) with lateral margin moderately sin- 
uate basally, rounded at middle, apical angles 
relatively broad and rounded, basal angles rect- 
angular or slightly obtuse, midlateral and baso- 


166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


lateral setae present; elytra without metallic re- 
flection, moderately dull, microsculpture 
moderately impressed, elytral silhouette sub- 
ovoid, elongate, narrowed basally, intervals 
moderately flat, fifth interval interrupted, mod- 
erately or markedly catenate (with 3 to 10 ca- 
tenations, restricted to apical one half or also on 
basal one-half of interval); median lobe of male 
(Fig. 23) slender basal to apical orifice, without 
ridge on right lateral surface, apex moderate in 
thickness and length and bent dorsally in lateral 
aspect, deflected left laterally in ventral aspect, 
apical orifice short, slightly constricted; bursa 
copulatrix of female (Fig. 30) with bursal sclerite 
small, narrow; specimen from locality in eastern 
ranges of Southern Rocky Mountains of central 
or northcentral Colorado or southeastern Wyo- 
ming (Fig. 32). 

DERIVATION OF TAXON NAME.—This subspe- 
cies is named for Pasquinel, the enigmatic, in- 
domitable French trapper in James A. Miche- 
ner’s epic novel, Centennial. The mountains 
traveled and loved by this fictional character are 
within the geographical range of this taxon and, 
in fact, include the type locality. 

GEOGRAPHICAL DISTRIBUTION.—Figure 32. 
Known only from the easternmost ranges of the 
Southern Rocky Mountains, from the Medicine 
Bow and Sierra Madre Ranges of southeastern 
Wyoming south to the Rampart Range of south- 
central Colorado. I have studied specimens from 
the following localities: 


United States of America 


CoLorapbo: (3; ANSP, KSUC, LACM); Boulder County, 
[Aug.] (1; CUB), Arapahoe Pass ([2740-3050 m]) [Aug.] (5; 
RTBe), Jenny Lake ({[3200 m]) [July] (1; CArm), Lake Isabelle 
({3170 m]) [July] (7; CArm), Jasper Lake ((3260 m]) [Aug.] (1; 
CArm), Lefthand Creek (5 miles E of Ward [2530 m]) [July— 
Aug.] (80; CArm, CAS), Little Royal Gorge [Aug.] (1; CUB), 
Long Lake ({3140 m]) [July, Sep.] (6; CArm), Mitchell Lake 
({3290 m]) [July] (7; CArm, CUB), Rainbow Lakes (10 miles 
SW of Ward [3350 m]) [Aug.] (1; CNC), Red Rock Lake ((2900 
m]) [Aug.] (6; CArm), South St. Vrain Creek ([3050 m]) [July] 
(1; CUB), Yankee Doodle Lake ({3140 m]) [July] (1; CArm); 
Clear Creek County, Leavenworth Valley (Argentine Road 
[2740-3350 m], Waldorf Mine [3540 m]) [June-July] (20; ANSP, 
CAS, CNC, DEUN, FMNH, MCZ, ROM, UMMZ, USNM, 
ZMLS), Mount Evans (Echo Lake [3230 m], Summit Lake 
[3960 m]) [July] (3; CAS, CNC), Silver Plume ([2740-3050 
m]) [June] (6; ANSP, CAS, MCZ, UWEM); El Paso County, 
Gold Camp Road (9 miles W of Broadmoor at South Cheyenne 
Creek [2650-2740 m]) [July—Aug.] (50; CAS, EAMa); Gilpin 
County, Rollinsville area [July] (1; CArm); Jackson County, 
Cameron Pass ({3140 m]) [Aug.] (1; UASM); Larimer County, 
Bennett Creek [May-July] (18; RTBe), Browns Lake Trail [Aug.] 
(1; RTBe), Buckhorn Creek [July] (2; RTBe), Cameron Pass 


({3050 m]) [June, Aug.] (6; MSU, RTBe, UASM, USNM), 
Crown Point Road (at Bennett Springs [2290-2350 m], at Crown 
Point Trail [3140-3200 m], 40 miles W of BeHevue [2740 m]) 
[June-Aug.] (20; CAS, RTBe, UASM, USNM), Monument 
Gulch [July] (18; RTBe), North Fork Cache la Poudre River 
[June] (2; RTBe), Zimmerman Lake [Aug.] (5; RTBe), West 
Fork Sheep Creek (16 miles W of Teds Place [3050 m]) [Aug.] 
(3; CNC); Park County, Kenosha Pass [July] (1; CAS); Rocky 
Mountain National Park, Blue Lake [July] (3; RTBe), Chasm 
Lake (stream below) [Aug.] (3; CArm), Endovalley Camp- 
ground [Aug.] (4; RTBe), Fall River ([2620 m]) [Aug.] (1; 
UASM), Hang Lake [Aug.] (1; RTBe), Lake Hiayaha [Aug.] 
(3; UAFA), Longs Peak ([3050-3350 m] and Boulder Field 
(3840 mJ], Larkspur Creek) [July—Aug.] (27; CArm, CAS), 
Thunder Lake ({3080 m] and Thunder Lake Trail [2500-3350 
m]) [June] (2; CArm); Routt County, Walton Creek (above 
Dumont Lake [2900-2960 m]) [Aug.] (3; CAS); Teller County, 
Divide (8 miles S on Cripple Creek Road [2900 m]) [July] (2; 
CNC). WYOMING: Albany County, Brooklyn Lane ([3200 
m]) [July] (18; CAS), Centennial [Aug.] (1; MSU), Douglas 
Creek (1 mile SSE of Keystone [2440 m]) [July] (16; CAS), 
Laramie Peak (south slope at Friend Creek [2440 m]) [July] 
(20; CAS), Little Brooklyn Lake ({[3120 m]) [July] (8; CAS), 
Snowy Range Pass ({3200 m]) [June] (8; UASM); Carbon 
County, South Brush Creek (20 miles SE of Saratoga [2470 
m]) [July] (1; CAS), Hidden Treasure Gulch (11.5 miles WSW 
of Encampment [2870 m]) [July] (2; CAS), Silver Lake ((3170 
m]) [July] (2; USNM), Slaughterhouse Gulch (11 miles SW of 
Encampment [2870 m]) [July] (6; CAS). 


Specimens Without Locality Data: (1; USNM). 


NOMENCLATURAL CHANGES 


Since my last two reports on Nearctic Nebria 
(Kavanaugh 1979 and 1981), additional speci- 
mens and data about geographical and habitat 
distribution have also accumulated for previ- 
ously described taxa. Study of these specimens 
and data, re-examination of some previously 
studied materials, and re-evaluation of species 
and subspecies concepts developed in previous 
reports (Kavanaugh 1978, 1979, and 1981) lead 
me to propose the nomenclatural changes pre- 
sented below. More complete discussions of pro- 
posed synomymies and changes in status of names 
will be presented in a forthcoming monography 
of Nearctic Nebria. 


Nebria intermedia VAN Dyke, 1949:49 [=Nebria 
crassicornis intermedia Van Dyke—NEw 
STATUS]. 


Analysis of the pattern of geographical varia- 
tion shown by samples representing Nebria cras- 
sicornis Van Dyke (1925:121) and N. intermedia 
Van Dyke throughout their ranges suggests that 
these taxa represent allopatric forms that are 
clearly differentiated but not to a degree typical 
of closely related, sympatric species. I suggest 


KAVANAUGH: NEW NEARCTIC NEBRIA 


that they be considered subspecies of a single 
species, N. crassicornis. 


Nebria sonorae KAVANAUGH, 1981:438 [=Ne- 
bria acuta sonorae Kavanaugh— New Status]. 


My description of N. sonorae was based on 
only one male and two female specimens. These 
specimens were only slightly, but consistently, 
different from specimens of Nebria acuta acuta 
Lindroth in several characters of external struc- 
ture. However, form of the aedeagus of the male 
specimen (chosen as holotype) was so different 
from that of N. acuta acuta males that I had no 
doubt concerning specific distinctiveness of the 
two taxa. 

Several additional male specimens of N. so- 
norae recently collected and studied all have ae- 
deagi with form typical of N. acuta acuta males. 
I now conclude that the aedeagus of the holotype 
male of N. sonorae is not typical for that taxon 
and, further, that the two forms are conspecific. 
However, differences in external structural char- 
acters between specimens of N. sonorae and N. 
acuta acuta are both consistent and sufficient to 
support their status as allopatric subspecies of a 
single species. 


Nebria fragilis Casey, 1924:21 [=Nebria arkan- 
sana fragilis Casey—New Status]. 

Nebria arkansana uinta Kavanaugh, 1979:102 
[=Nebria arkansana fragilis Casey —NEw 
SYNONYMYyY]. 

Nebria fragilis teewinot Kavanaugh, 1979:103 
[=Nebria arkansana fragilis Casey —NEw 
SYNONYMYy]. 


Males of Nebria arkansana arkansana Casey 
and N. a. edwardsi Kavanaugh differ distinctly 
from males of “‘Nebria fragilis Casey” in form 
of aedeagus and in several characters of external 
structure. Male specimens from the Uinta and 
northern Wasatch Mountains of northern Utah 
share aedeagal form with males of the two N. 
arkansana subspecies just mentioned, yet differ 
from them slightly in characters of external struc- 
ture. I therefore recognized (Kavanaugh 1979) 
these specimens as representing a distinct sub- 
species of N. arkansana, namely N. a. uinta. Male 
specimens from the Teton, Wind River, and ad- 
jacent Mountains in western Wyoming share ae- 
deagal form with males of N. fragilis; but, again, 
they differ from the latter in several characters 
of external structure and color. Based on these 


167 


similarities and differences, I recognized (Ka- 
vanaugh 1979) the Wyoming specimens as rep- 
resenting a distinct subspecies of N. fragilis, 
namely N. f teewinot. The resulting pattern of 
geographical distribution was one in which the 
range of all subspecies of both N. arkansana and 
N. fragilis were mutually allopatric, although 
ranges of N. arkansana uinta and N. fragilis fra- 
gilis were essentially parapatric in northcentral 
Utah. 

Since 1979 I have studied additional material 
from areas that previously represented gaps be- 
tween the allopatric ranges of described subspe- 
cies, and I re-examined specimens studied ear- 
lier. These studies have shown that samples of 
males from localities in presumed gaps, as well 
as some samples from localities bordering these 
gaps, are mixed in aedeagal form and interme- 
diate in characters of external structure in rela- 
tion to respective allopatric forms. Such findings 
lead me to conclude that N. arkansana and N. 
fragilis are conspecific. Specimens representing 
N. arkansana uinta, N. fragilis fragilis, and N. f. 
teewinot all share a combination of structural 
features which distinguish them from members 
of other N. arkansana subspecies. I therefore 
suggest that these forms (and respective inter- 
mediates between them) together represent a sin- 
gle, distinct subspecies of N. arkansana for which 
the name N. arkansana fragilis has priority. A 
reconstruction of the historical development of 
the complex pattern of geographical variation 
within this subspecies will be presented in the 
monograph now in preparation. 


Nebria trifaria tetonensis ERWIN AND BALL, 1972: 
95 [=Nebria trifaria trifaria LeConte—New 
SYNONYMY]. 


Based on an almost continuous series of sam- 
ples (all collected after 1972) from the area be- 
tween respective type localities for N. trifaria 
trifaria and N. trifaria tetonensis, I suggest that 
these two nominal taxa represent simply the ex- 
tremes of continuous clinal variation in those 
characters previously used to distinguish their 
members. 


Nebria trifaria piute ERWIN AND BALL, 1972:95 
[=Nebria piute piute Erwin and Ball—New 
STATUS]. 


Members of Nebria piute and N. trifaria 
LeConte differ from each other at least as much 


168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


as other closely related, but sympatric, species 
in characters of both external structure and gen- 
italia (of both males and females). I therefore 
suggest that they represent distinct species. 


Nebria trifaria utahensis KAVANAUGH, 1979: 110 
[=Nebria piute utahensis Kavanaugh— New 
STATUS]. 


Based on form of aedeagus of males and bursa 
copulatrix of females, N. utahensis, described as 
a subspecies of N. trifaria LeConte, should in- 
stead be considered a subspecies of N. piute, along 
with N. piute sevieri n.ssp. described above. 


LITERATURE CITED 


Casey, T. L. 1924. Additions to the known Coleoptera of 
North America. Memoirs on the Coleoptera, 11:1-347. 
Erwin, T. L., AND G. E. Batt. 1972. Classification of the 


ovipennis and trifaria groups of Nebria Latreille (Coleoptera: 
Carabidae: Nebriini). Proceedings of the Biological Society 
of Washington, 85:77-108. 

KAVANAUGH, D. H. 1978. The Nearctic species of Nebria 
Latreille (Coleoptera: Carabidae: Nebriini): classification, 
phylogeny, zoogeography, and natural history. Unpublished 
Ph.D. dissertation. Department of Entomology, University 
of Alberta. xlviii + 1041 pp. 

1979. Studies on the Nebriini (Coleoptera: Carabi- 

dae), III. New Nearctic species and subspecies, nomencla- 

tural notes, and lectotype designations. Proceedings of the 

California Academy of Sciences, 42:87-133. 

1981. Studies on the Nebriini (Coleoptera: Carabi- 
dae), 1V. Four new Nebria taxa from western North America. 
Proceedings of the California Academy of Sciences, 42:435- 
442. 

VAN Dyke, E. C. 1925. Studies of western North American 
Carabinae with descriptions of new species. The Pan-Pacific 
Entomologist, 1:111-125. 

1949. New species of North American Coleoptera. 

The Pan-Pacific Entomologist, 25:49-56. 


KAVANAUGH: NEW NEARCTIC NEBRIA 169 


Ficures 1-8. Fig. 1. Right antennal scape, dorsal aspect, Nebria gebleri albimontis n.ssp. (Birch Creek, California); scale 
line = 1.0 mm. Figs. 2-8. Pronotum, dorsal aspect; scale line = 1.0 mm. 2. Nebria altisierrae n.sp. (Olmsted Point, California). 
3. Nebria campbelli n.sp. (Mount Baker, Washington). 4. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 5. Nebria 
Jeffreyi n.sp. (South Fork McCoy Creek, Oregon). 6. Nebria haida n.sp. (Mount Needham, Queen Charlotte Islands, British 
Columbia). 7. Nebria louiseae n.sp. (Skedans, Queen Charlotte Islands, British Columbia). 8. Nebria gebleri albimontis n.ssp. 
(Birch Creek, California). 


170 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


Ficures 9-14. Pronotum, dorsal aspect; scale line = 1. 


10. Nebria calva n.sp. (Mount Baldy, Arizona). 11. Nebri 


sevieri n.ssp. (Parowan Creek, Utah). 13. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon). 


pasquineli n.ssp. (Lefthand Creek, Colorado). 


14 


0 mm. 9. Nebria labontei n.sp. (West Fork Wallowa River, Oregon). 


a sierrablancae n.sp. (Sierra Blanca, New Mexico). 12. Nebria piute 
14. Nebria trifaria 


KAVANAUGH: NEW NEARCTIC NEBRIA 171 


15 


Ficures 15-16. Basal region of left elytron, dorsal aspect; scale line = 1.0 mm. 15. Nebria altisierrae n.sp. (Olmsted Point, 
California). 16. Nebria campbelli n.sp. (Mount Baker, Washington). 


Ficures 17-20. Median lobe of male genitalia, left lateral aspect (or a = left lateral aspect, b = ventral aspect); scale line = 
1.0 mm. 17. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 18. Nebria gebleri albimontis n.ssp. (Birch Creek, 
California). 19. Nebria calva n.sp. (Mount Baldy, Arizona). 20. Nebria sierrablancae n.sp. (Sierra Blanca, New Mexico). 


172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


Ficures 21-23. Median lobe of male genitalia (a = left lateral aspect, b = ventral aspect, c = apical aspect); scale line = 1.0 
mm. 21. Nebria piute sevieri n.ssp. (Parowan Creek, Utah). 22. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon). 
23. Nebria trifaria pasquineli n.ssp. (Lefthand Creek, Colorado). 


KAVANAUGH: NEW NEARCTIC NEBRIA 173 


Ficures 24-26. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect): 
scale line = 1.0 mm. 24. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 25. Nebria gebleri albimontis n.ssp. (Birch 
Creek, California). 26. Nebria calva n.sp. (Mount Baldy, Arizona). 


174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


Ficures 27-28. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect); 
scale line = 1.0 mm. 27. Nebria sierrablancae n.sp. (Sierra Blanca, New Mexico). 28. Nebria piute sevieri n.ssp. (Parowan Creek, 


Utah). 


KAVANAUGH: NEW NEARCTIC NEBRIA 175 


Ficures 29-30. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect); 
scale line = 1.0 mm. 29. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon). 30. Nebria trifaria pasquineli n.ssp. 
(Lefthand Creek, Colorado). 


176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12 


Ficure 31. Map of geographical distributions: Nebria altisierrae n.sp. = solid circle, Nebria campbelli n.sp. = solid diamond; 
Nebria wallowae n.sp. = solid square; Nebria jeffreyi n.sp. = open diamond; Nebria haida n.sp. = solid triangle; Nebria louiseae 
n.sp. = open triangle. 


KAVANAUGH: NEW NEARCTIC NEBRIA 177 


Figure 32. Map of geographical distributions: Nebria gebleri albimontis n.ssp. = solid square; Nebria labontei n.sp. = solid 
triangle; Nebria calva n.sp. = solid diamond; Nebria sierrablancae n.sp. = open triangle; Nebria piute sevieri n.ssp. = inverted 
solid triangle; Nebria steensensis n.sp. = open diamond; Nebria trifaria pasquineli n.ssp. = solid circle. 


ry 
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PROCEEDINGS 


Marine Biolonic 
ole 


fe) 7. 


OF THE 


Vol. 43, No. 13, pp. 179-220, 22 figs., 2 tables. 


July 12, 1984 


SKELETAL ANATOMY AND CLASSIFICATION 
OF THE NEOTENIC ASIAN SALMONIFORM SUPERFAMILY 
SALANGOIDEA (ICEFISHES OR NOODLEFISHES) 


By 


Tyson R. Roberts 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


Apstract: The distinctive, largely cartilaginous skeletal anatomy of the Asian icefishes or noodlefishes is 
described and figured from cleared specimens stained with alcian and alizarin. This information, together 
with examination of types and other material in North American and European museums, leads to the 
following revised classification of these neotenic Salmoniformes: 

Salangoidea new superfamily (coordinate with Osmeroidea, Salmonoidea) 


Salangidae 
Protosalanginae: Protosalanx chinensis 


Salanginae: Salanx (Salanx) ariakensis, S. (S.) cuvieri, S. (Hemisalanx) prognathus, S. (Leucosoma) reevesi 
Salangichthyinae new subfamily: Neosalanx andersoni, N. brevirostris, N. jordani, N. reganius, Salang- 


ichthys ishikawae, S. microdon 


Sundasalangidae: Sundasalanx microps, S. praecox 


The introduction includes a summary of salangoid natural history and a key for their identification. The 
systematic account includes all primary and secondary synonyms of genera and species recognized. New 
information is presented on pectoral girdle morphology in teleosts, relationships of salangoids and other 
salmoniforms, and breeding tubercles, meristic variation, and neoteny in salangoids. 


INTRODUCTION 


The slender, soft-bodied, and transparent or 
translucent salmoniform fishes of the family Sa- 
langidae inhabit the sea coasts, rivers, and lakes 
of East Asia including Japan from Sakhalin, Vla- 
divostok, and the Amur River south to northern 


Vietnam (Tonkin). The greatest concentration of 


genera and species is in China and Korea. Of 11 
species herein recognized, eight occur in China, 
eight or nine in Korea, and four in Japan. Only 
Salangichthys microdon occurs along the outer 


coast of Korea and in Siberia, and only Salanx 


reevesi and Neosalanx brevirostris have been re- 
ported as far south as Tonkin (or Haiphong). 
Members of the Salangidae have almost al- 


ways been referred to in English as icefishes. In 


Japanese, however, they are usually referred to 
as shirauwo (whitefishes) and rarely as hiagio 
(icefishes). In Russian they are usually referred 
to as /apsha-ryba or noodlefish, and an equiva- 
lent name exists in Chinese, mien-tiao-yu 
( #14). They have been referred to as Nudel- 


fische in German, but noodlefishes, a highly ap- 


propriate and distinctive name, seems not to have 
appeared in English except in a translation of a 
Russian work (Berg 1962:480). The flesh is tasty, 
whether cooked as a soup, eaten with vinegar or 
scrambled eggs, or fried (Okada 1955:60). The 
species most commonly eaten in Japan is Sa- 
langichthys microdon, and in China probably 
Neosalanx brevirostris or N. jordani. Protosa- 
lanx and Salanx are also consumed, but I doubt 
that tiny Sundasalanx has ever been dined upon. 


[179] 


| ‘ 

al Lasaratar 
wastit' ‘ 
2t5 Paiury 


180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. i3 


= 


ee ee 
STTUCUI SOCEM CeCe oe ce rece 
a ee ie 


\\ 


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hil lial a eb loll litelltisel 


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OT SALAMA 


SASS 


Axial skeleton. (a) Protosalanx chinensis, CAS-SU 6306, 85.5-mm adult male; (b) Salanx cuvieri, CAS-SU 32454, 


(OE 


FiGure |. 


ee ee eae ae 


JE 


1 1 VI: 


a 


61.7-mm juvenile sex undetermined; (c) Salangichthys ishikawae, CAS 6780, 74-mm adult female; (d) Neosalanx jordani, CAS 
52028, 38.3-mm adult male; (e) Sundasalanx microps, CAS 44220, 17-mm adult sex undetermined. 


Despite their standing as a delicacy—sufficient 
for them to be imported by the Chinese and Jap- 
anese communities of San Francisco and served 
in the city’s sushi bars—relatively little is known 
about the systematics and biology of noodlefish- 
es. An impression of their morphological diver- 
sity can be obtained from Figures | and 2. 

The present study was undertaken in connec- 
tion with the discovery of some minute, scaleless, 
and transparent fishes during my fieldwork in the 


Malay Peninsula (1971, 1973) and on the Kapuas 
River in Kalimantan Barat, Indonesia (1976). 
When first found, although in fresh water, they 
were living close to the sea and were mistaken 
for elopoid leptocephali, which they resemble 
only superficially. In the Kapuas River, however, 
they were living 800 km upriver in the midst of 
a rich riverine fish fauna dominated by Ostar- 
iophysi and with no elopoids. The observation 
that the maxillary bones curved inwards below 


- 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 


TABLE |. 
and pers. obs.). 


181 


REPRODUCTIVE BIOLOGY OF SALANGOIDS (Wakiya and Takahasi 1937, Okada 1960, Senta 1973a, b, Roberts 1981, 


Larg- 
Smallest Larg- est Diam- 
mature est fe- eter 
male male male egg 
(mm) (mm) (mm) (mm) Fecundity Remarks 
Protosalanx chinensis 82 168 146 NSS _ Breeds January-February (Korea) 
Salanx ariakensis = — 147 0.75 _ Breeds October-November (Korea) 
Salanx cuvieri _ — 144 — _ — 
Salanx prognathus 100 111 119 0.85 - Breeds April-May 
Salanx chinensis 130 130 153 — a — 
Neosalanx andersoni 79 100 95 —_ — Breeds April-May (Korea) 
Neosalanx brevirostris _ 64 60 0.7 — _ ; 
Neosalanx jordani 34 56.5 59.5 0.5 - Breeds March—May 
Neosalanx reganius — 56 58 0.9 ~ Breeds February—March 
Salangichthys ishikawae _ 71 74 0.95 _ Breeds April-May 
Salangichthys microdon 65 90 100 0.91-0.99 1300-2700 Breeds March—May 
Sundasalanx microps — — = = Largest specimen (sex unknown) 
19.9 mm 
Sundasalanx praecox 14.9 18.3 17.3. 0.20-0.25 50 Both sexes ripe in June 


the head led to an hypothesis that they are sa- 
langoids, and observations of their skeletal anat- 
omy and particularly the suspensorium con- 
firmed this (Roberts 1981). These fishes differ in 
a number of respects from Salangidae and con- 
stitute a separate family, Sundasalangidae, with 
one genus, and two or more species, one in the 
Malay Peninsula and one or two in the Kapuas 
River (Roberts 1981). Sundasalanx also occur 
in the Mekong basin, as reported herein. This is 
the only truly tropical genus in the entire order 
Salmoniformes. Sundasalanx praecox, with 
males and females sexually ripe at only 14.9 mm, 
is the smallest member of the order, and provides 
a striking example of a minute secondary fresh- 
water fish living in the midst of a rich freshwater 
ichthyofauna dominated by primary freshwater 
Ostariophysi. 

Interest in Sundasalangidae and its relation- 
ships led me to examine other salangoids but my 
observations and drawings quickly became too 
extensive to incorporate in the original descrip- 
tion of the new taxa; hence the present mono- 
graph. 


Food Habits 


All salango:ds, including tiny Sundasalanx, 
appear to be predators. The largest species, Pro- 
tosalanx chinensis and Salanx reevesi, both with 
well-developed teeth on the tongue and jaws, ap- 
parently feed mainly on fishes. Salangichthys 


microdon taken in the Takahashi River had fed 
on larvae of the goby Chaenogobius sp. and on 
the mysid shrimp Neomysis sp. (Senta 1973b). 
Other species of Salanginae and Salangichthy1- 
nae feed mainly on small crustacea (in marine 
environments) or on insects (in fresh water). 
Sundasalanx are known only from fresh water 
and feed on tiny insects (Roberts 1981). 


Reproduction 


While some species are primarily marine or at 
least brackish water inhabitants (e.g., Protosa- 
lanx chinensis), and many spend part of their 
lives in the sea, others are restricted to fresh water 
or have populations which presumably repeat 
their life cycle without leaving fresh water. Basic 
information on salangoid reproductive biology 
is summarized in Table |. Fecundity ranges from 
several thousand eggs in Protosalanginae and 
Salanginae (no precise numbers available) down 
to only about 50 in Sundasalangidae. 

The external egg membrane is adhesive, eggs 
becoming attached to any solid object at the 
spawning site. Wakiya and Takahasi (1937, pl. 
21) published drawings of the basal portion of 
the adhesive strands on the eggs of Protosalanx 
chinensis, Salanx ariakensis and S. prognathus, 
Salangichthys microdon and S. ishikawae, and 
Neosalanx jordani. The eggs illustrated are pre- 
sumably ovarian, since the adhesive strands are 
not detached. For photomicrographs of the 


182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


Ficure 2. Radiographs. (a) Salanx cuvieri, MNHN 9900, 112 mm, holotype; (b) Neosalanx andersoni, NRM 10287, 79 


mm, holotype. 


spawned eggs with detached adhesive threads of 
unfertilized and fertilized eggs of Salangichthys 
microdon see Okada (1960, pl. 18). Spawning 
ecology of this species is described by Senta 
(1973a). According to Wakiya and Takahasi 
(1937:269), after spawning “the body becomes 
very lean and the vertebrae become visible 
through the skin, whence it is generally assumed 
that death then ensues.” I suspect that this is true 
in Salanginae as well as Salangichthyinae but not 
in Protosalanx. 


Sexual Dimorphism 


A notable feature of salangoids is their unique 
sexual dimorphism. In all Salangidae except 
Neosalanx, sexually mature males have the pec- 
toral fins longer and more pointed (falcate) and 
the pelvic fins larger. In all adult male Salangidae 
the anal fin is larger than in females and has 
modified rays. The anterior rays of the anal fin 
are greatly enlarged, the middle rays thin and 
strongly curved, and the posterior rays short and 
widely separated at the base. The morphology of 
the anal fin is very similar in sexually mature 
males of all of the genera and species of Salan- 
gidae. In all Salangidae, mature males have a row 
of large, tightly adherent scales on the body par- 
allel to the anal fin base (Sometimes extending 
posteriorly a short distance beyond the anal fin 
base onto the caudal peduncle). The number of 
anal scales ranges from 14 to 28. Sexual dimor- 


phism has not been observed in Sundasalangi- 
dae. 

Although salangids differ greatly in the size of 
adult males, the morphology of the modified male 
anal fin is remarkably uniform (Fig. la, d). The 
total range of anal fin-rays is 23-32. The first two 
or three rays are simple, the first one or two small 
or minute. The last simple ray and the first four 
to six branched rays are greatly enlarged and 
somewhat thickened; near the base of each of 
these rays is a very large lateral projection. The 
next 12 or so rays are noticeably thinner and are 
deflected backwards near the middle of their 
length, so that their distal portions lie close to- 
gether. In Protosalanx these rays are simple, but 
in other Salangidae they are branched. The pos- 
teriormost rays may be simple or branched, are 
reduced in size and not modified, except that 
their bases tend to be relatively wide apart (much 
more so than the bases of the preceding rays or 
of the corresponding rays in females), especially 
in Salanginae. The proximal pterygiophores, es- 
pecially for the anterior portion of the anal fin, 
are also enlarged in males. In alcian-alizarin 
preparations the anal fin-rays and pterygiophores 
of sexually mature males are deeply stained with 
alizarin, whereas those of females tend to be less 
well stained with alizarin or in some instances 
stained only with alcian. 

Near the middle of the rays in the most mod- 
ified part of the male anal fin, a tough, almost 
tendonlike membrane arises from each ray and 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 183 


extends obliquely and posteroventrally across the 
densely webbed portion of the fin to end in a 
thickened, obliquely oriented non-muscular pad 
of tissue. The distal portion of this oblique pad 
is free from the surface, so it can be readily lifted, 
and even when not lifted forms a sort of groove 
for the length of the pad. This portion of the anal 
fin can be flexed in such a way that it forms a 
slight concavity. The fin may be expanded man- 
ually by pulling on the anteriormost rays; when 
released, it snaps back into a less expanded con- 
dition. Spawning behavior has not been reported 
upon, but presumably the male’s anal fin remains 
in contact with the vent region of the female in 
such a way that it temporarily retains eggs and 
sperm in proximity while fertilization occurs ex- 
ternally. 

In addition to the modified anal fin, sexually 
mature males of all Salangidae bear a row of 
large, cycloid scales on the side of the body above 
and co-extensive with the anal fin or extending 
a short distance beyond it onto the caudal pe- 
duncle. The scales are tightly adherent and 
broadly overlapping (more so anteriorly than 
posteriorly). In addition to the main row of anal 
scales, some specimens exhibit two or three 
smaller scales in a separate row overlying the 
vent. These usually have been overlooked by 
previous authors, and are not included in the 
counts of anal scales in Table 2. 

Breeding tubercles and other forms of tem- 
porary sexual dimorphism have not been re- 
ported previously in salangoids. { have observed 
breeding tubercles in adult males and females, 
apparently in spawning condition, of Protosa- 
lanx chinensis, and in adult males of Salangich- 
thys microdon and Neosalanx jordani. This pre- 
sumably temporary tuberculation is most 
extensive and easily observable in an 120-mm 
male Protosalanx (CAS-SU 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, 11.5 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 ofa 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 Sa/anx 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 exhibit 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 129-132 mm (USNM 120746). 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 NV. andersoni (NRM 10287, 79 
mm). Chyung (1961) shows it well developed in 
a gravid N. andersoni. | 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 (1934a: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 Giinther (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 61 
males and 27 females identified as Hemisalanx 
(=Salanx) progsnathus 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- 
pidae)i eee se: Sie cea Cee ane ll 

1b. 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- 
Leet ae)! ots 12 pearl NEM ne ea em 


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) _. 
SL rani Sh AN ZS 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- 


186 


4a. 


Ab. 
Sa. 


Sb. 


6a. 


6b. 


Ta. 


7b. 


8a. 


8b. 


9a. 


9b. 


10a. 


10b. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


ent; vertebrae 48-65 (Salangichthyinae) 


Tongue with a median row of conical 
teeth (subgenus Leucosoma) 
LA SIRU SRR, Mess DiI) Sate iOas saat ne Salanx reevesi 


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) ne 6 
Head less strongly pointed; lower jaw 
projecting slightly beyond upper jaw; no 
presymphyseal fleshy appendage, bone, 
or teeth; vertebrae 70-73 (subgenus 
Hemisalanx) 0.00000... Salanx prognathus 


Presymphyseal bone usually present in 
specimens over 100 mm standard length, 
relatively elongate and with up to 17 teeth 
on each side; vertebrae usually 77-78 
(rarely sO: One] 9) se Salanx cuvieri 
Presymphyseal bone usually absent, or 
relatively short and with no more than 
6 teeth on each side; vertebrae 72-75 ... 
Li ei Da Sai, Pe eattase rel ole Salanx ariakensis 


Palatal toothplate with minute teeth; 
premaxilla with numerous small or mi- 
nute teeth, snout relatively elongate; ver- 
tebrae 59-65 (Salangichthys) 00... 8 
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 


Pectoralifin-rays 14d=19 1 
Ma oe Raa Meet Re tle Salangichthys microdon 
Rectoralin-rays) 20—28 2a ee 
eG, Teint net FV UehG Salangichthys ishikawae 


Snout relatively short, standard length to 
64 mm, males with 14-21 anal scales, 
VeETLe Dracwe Wels tiannG() sess een eee 10 
Snout relatively elongate, standard length 
to 100 mm, males with 20-28 anal scales, 
vertebrae 63-65 ....... Neosalanx andersoni 


Vertebrae 55-59; standard length to 64 
mm; total rakers on first gill arch 15-19 
(mainland Asia) .... Neosalanx brevirostris 
Vertebrae usually 50-53, rarely 49 or 54; 
standard length usually less than 50 mm; 


total rakers on first gill arch 9-15 (main- 
landtAsia) 22. 3! Neosalanx jordani 
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 


10c. 


lla. 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 _..... 
Benet wt NL KA Sundasalanx microps 
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 
Oi =A Awa Aicaeee Sundasalanx praecox 


1 1b. 


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 (1970) 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 


Ficure 3. Dorsal and ventral view of cranium. Protosalanx chinensis, CAS-SU 6306, 158 mm. 


(his Salanx chinensis and Salangichthys micro- gill arches agree closely with Nelson’s. Rosen 
don). He particularly noted the well-developed (1974; figs. 16g, 26a & b) figured and com- 
fourth hypobranchials, “which so far as known mented briefly upon the caudal skeleton and por- 
are absent from all other adult teleostean fishes.” _ tions of the gill arches of Neosalanx brevirostris 
My own observations and drawings of salangid (his Salangichthys microdon). 


188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


supraethmoid 


frontal 


parasphenoid 


1mm, 


Ficure 4. Dorsal and ventral view of cranium. Salanx cuvieri, CAS-SU 32454, 69.4 mm. 


The advent of a technique for staining whole 
specimens with alcian and alizarin (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 


supraethmoid 


l 2mm 


FiGure 5. 
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. 


frontal | | 7 


parietal 


Dorsal view of cranium and membrane bones on dorsal surface of cranium. Salanx prognathus, CAS 51439, 110 


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 posteriorly recurved jaw teeth 
(especially in Salanginae and Protosalanginae). 


190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


tectum synoticum 


cranial fontanel 


ethmoid plate 


supraethmoid 


taenia marginalis 


pila prooptica 


nasal recess 


trabecula Communis 


hyomandibular fossa 


epiphyseal bar 


auditory capsule 


foramen 


semicircular canals 


basioccipital 


parasphenoid 


\ 1mm ' 


Ficure 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 191 


taenia marginalis 


lamina orbitonasalis 


trabecula Communis 


hypophysial fenestra 


auditory fenestra 


parachordals 


notochordal groove 


1mm 


FIGURE 7. 


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 of ten Neo- 
salanx jordani (39.7-45.7 mm), nine have the 


Dorsal and ventral views of cranium. Sundasalanx microps, CAS 44220, 17 mm. 


cranial fontanel entirely undivided, but one (41.0 
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 


192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


a b 

pe 
_Amm, 1mm , 
Cc cranial fontanel d ethmoid plate lateral fenestra 
i ion lamina 
Plesercoptt . orbitonasalis pila prooptica 
trabecula Communis 14mm 1mm 

Ficure 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) Sundasalanx 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 1s represented by an anterolateral 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 193 


premaxilla 


ethmopalatine 


maxilla 


dentary 
Meckel’s cartilage 


coronoid process 
4 mm 


articular 


lateral 
process 


hyopalatine opercle 


preopercle 


hyomandibular 


quadrate subopercle 


interopercle 


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 
centrum. 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 corner 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 


Ficure 10. Lateral and medial views of jaws, suspensorium, and opercular bones. Salanx cuvieri, 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 Protosalanx 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. 12). 


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 195 


Ficure 11. 


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 | 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 


infraorbital ~~) 


1mm , 


Lateral view of jaws, suspensorium, and opercular bones. Sa/anx prognathus, CAS-SU 51439, 110 mm. 


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 


1mm 


Ficure 12. Lateral view of jaws, suspensorium, and opercular bones. Neosalanx jordani, CAS 52058, 35.1 mm. 


196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


OP 


1mm 


Ficure 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. 


basibranchials 


ceratobranchials 


urohyal 


branchiostegal rays 1mm j 


Ficure 14. Dorsal and ventral views of hyoid and branchial arches. Protosalanx chinensis, CAS-SU 6306, 153 mm. 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 197 


basihyal 


basihyal 
toothplate 


dorsohyal infrapharyngobranchials 


uncinate 
accessory process 


anterohyal Z 
cartilage 


upper pharyngeal 
toothplate 

lower pharyngeal 
toothplate 


4 mm 2mm 
———— 


Figure 15. Dorsal view of hyoid and branchial arches and ventral view of upper pharyngeal elements. Salanx cuvieri, CAS- 
SU 32454, 69.4 mm. 


aN 


Ficure 16. Dorsal view of hyoid and branchial arches; ventral view of infrapharyngobranchial 4 and upper pharyngeal 
toothplate. Neosalanx jordani, CAS 52058, 38.3 mm. 


 imm , 


198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


FiGure 17. 


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 


basibranchial 1+ 2+ 
hypobranchial 1 + 2 


I basibranchial 3 + 


é AK hypobranchial 3 


hypobranchial 4 


basibranchial 4 


1mm 


Dorsal view of hyoid and branchial arches. Sundasalanx microps, CAS 44220, 17 mm. 


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 199 


a posttemporal 


supracleithrum 
 “<. 


medial 
process 


scapulocoracoid 


(e imm 


proximal radials 1-5 


distal radials 


b 


postcleithrum 


FiGure 18. 


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 1s 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. 18a, 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- 


Left half of pectoral girdle. (a) Protosalanx chinensis, CAS-SU 6306, 158 mm (medial view); (b) Salanx cuvieri, 
CAS-SU 32454, 69.4 mm (lateral view); (c) Neosalanx jordani, CAS 52058, 43.1 mm (dorsal view); (d) Salangichthys ishikawae, 
CAS 6780, 74 mm (lateral view). 


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- 
sues. 


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 


Ficure 19. Pectoral girdle. (a) Sundasalanx microps, CAS 44220, 17-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 1s 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. 1b). 

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 


radials 1-4 


- —EE 
———_—— 
—— ———————— 


1mm 


Ficure 20. Ventral view of left half of pelvic girdle. (a) Protosalanx chinensis, 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 | (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 | 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 | is apparently always fused to 
complex hypural centrum. Free radial or ptery- 
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 microdon (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 


a opisthural 
epural 


uroneural 


ural centrum 


hemal spines 
hypurapophysis 
parhypural 


FiGure 21. 


b 


Ez 
i 
Vie 
Oj Ufge= == 


ee a 


WS 
<== 


S=> 


SSS 


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-— 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 Sal/anx 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 1s 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, branchiostegal 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 microdon). 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 
(1971:119) 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- 


204 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 Sundasalanx). 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 1n- 
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, 1902 
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 Protosalanx, 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 
DN) 

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 BASsILewsky, 1855:242. 

Salanx Assott, 1901:490. 

Protosalanx REGAN, 1908b:444 (type-species, by monotypy, 
Salanx hyalocranius Assott, 1901 = Eperlanus chinensis 
BASILEwsky, 1855). 

Paraprotosalanx FANG, 1934a:246 (type-species, by mono- 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 205 


typy, Protosalanx andersoni FANG, 1934a (non RENDAHL, 
1923) = Protosalanx chinensis BASILEWSKY, !855). 


Protosalanx chinensis (Basilewsky, 1855) 


Eperlanus chinensis BASILEwsky, 1855:242 (type-locality “‘in 
sinu Tschiliensis habitat” [not “Pekin” as usually cited)]). 
Salanx hyalocranius Assott, 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 of Salanx 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 1901: 
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 Makao”’ 
has been referred to as Leucosoma or Salanx 
chinensis but never as Eperlanus or Protosalanx 
chinensis, the epithet chinensis is available for a 
species of 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 S. 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 are 
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, 1908b 


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, pl. 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 
7-11 (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, Hoplias, and Acestro- 
rhynchus (Roberts 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 1s 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). 


Salanx Oken, 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, Sa/anx cuvieri VALENCIENNES, 1849). 

Leucosoma Gray, 1831:4 (type-species, by monotypy, Leu- 
cosoma reevesi GRAY, 1831). 

Hemisalanx REGAN, 1908b:444 (type-species, by monotypy, 
Hemisalanx prognathus REGAN, 1908b). 

Parasalanx REGAN, 1908b:444 (type-species, by subsequent 
designation of FANG, 1934a:259, Parasalanx gracillimus 
REGAN, 1908b = ?Salanx cuvieri VALENCIENNES, 1849). 

Reganisalanx FANG, 1934b:509 (type-species, by monotypy, 
Reganisalanx normani FANG, 1934b = Salanx ariakensis 
KisHiNouy_E, 1901). 

Metasalanx WaAKIYA AND TAKAHASI, 1937:293 (type-species, 
by monotypy, Metasalanx coreanus WAKIYA 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 (Sa/anx) ariakensis 

(Kishinouye, 1901) 

Salanx ariakensis KisHtNouyYE, 1901:359 (type-locality Ariake 
Bay, Kiushiu). 

Salanx acuticeps REGAN, 1908a:360 (type-locality Lake Can- 
didius, Formosa). 

Parasalanx acuticeps REGAN, 1908b:446. 

Parasalanx longianalis REGAN, 1908b:446 (type-locality Liao- 
ho, northern China). 

Parasalanx annitae vAN Dam, 1926:342 (type-locality Pei- 
taiho, China). 

Reganisalanx normani FANG, 1934b:509 (type-locality Ichang, 
as herein restricted). 


MATERIAL ExXAMINEP.—AMNH_ 10327, 7:125-147 mm, 
Hunan; BMNH 1888.5.15.1 1-12, 2:141-143 mm, Ichang (lec- 
totype and paralectotype of R. normani); BMNH 1898.2.8.20- 
23, 4:114-123 mm, Liao-ho, northern China (syntypes of P. 
longianalis), BMNH 1904.4.2835-36, 2:116-118 mm, Lake 
Candidius, Formosa (syntypes of S. acuticeps); BMNH 
1927.3.26.3, 125 mm, Nanking; BMNH 1928.6.22.6, 115 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 112.587, 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 S. ari- 
akensis examined have only 26-29 anal fin-rays, 
but two specimens from Ariake Bay have 27 and 
Silk 

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 207 


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 11 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 Sa/lanx) 
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 12, and dentary 
about 10. 


Salanx (Sa/anx) cuvieri Valenciennes, 1849 


Salanx cuvieri VALENCIENNES in CUVIER AND VALENCIENNES, 
1849:360 (type-locality unknown). 

?Parasalanx 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. an- 
gusticeps); BMNH 1891.1.31.20, 111 mm, Shanghai (holotype 
of P. gracillimus); BMNH 1936.10.7.13, 119 mm, Sharp Peak, 
Fukien; CAS 52057, 4:76.5-98.0 mm, Hong Kong (1 alizarin); 
CAS-SU 225732, 112 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, 117 mm, near Pakhoi, 
SW Kwangtung; MNHN 9900, 112 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 Hototype.— 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 
10; diameter of eye (slightly shrunken) approx- 
imately 28. 

NoTEs ON SYNONYMyY.—P. angusticeps 1s 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 | 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 


208 


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ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 


“GSNNILNO)) °7 ATAV 


210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


cillimus as a synonym of P. ariakensis, and per- 
haps they were correct. This matter is of some 
nomenclatural significance, since P. gracillimus 
is type-species of Regan’s Parasalanx. 

The holotype of P. cantonensis has a very elon- 
gate presymphyseal bone with 10 teeth; premax- 
illary teeth 8, maxillary teeth 10, dentary teeth 
13; palatal teeth 11. 

Length of the head (and cranium) is excep- 
tionally variable in all genera of Salangidae, even 
including the relatively short-snouted genus 
Neosalanx, and is particularly variable in the 
long-snouted subfamily Salanginae. Variable 
elongation of the head and particularly the snout 
with its flattened ethmoid plate is evident in the 
large series of Protosalanx chinensis and Neo- 
salanx jordani 1 examined, even though these 
are relatively small specimens. P. angusticeps 1s 
based on a single specimen, the holotype, which 
happens to be the largest referred to S. cuvieri 
that I have examined. In vertebral counts and 
other characters it apparently agrees well with 
other material of S. cuvieri. 


Salanx (Hemisalanx) prognathus (Regan, 1908b) 

new combination 

Hemisalanx prognathus REGAN, 1908b:445 (type-locality 
Shanghai). 

?Salanx brachyrostralis FANG, 1934a:257 (type-locality Nan- 
king). 

?Reganisalanx brachyrostralis FANG, 1934b:509. 

Metasalanx coreanus WAKIYA AND TAKAHASI, 1937 (otherwise 
unpublished manuscript name placed in synonymy of H. 
prognathus by WAKIYA AND TAKAHASI 1937:293). 


MATERIAL EXAMINED.—BMNH_  1873.7.30.69, 111 mm, 
Shanghai (holotype); CAS 51439, 34:97.5-120 mm, Kiangsu 
Prov. (12:97.5-112 mm alcian-alizarin); CAS-SU 33990, 8: 
94-114 mm, Tai Po, New Territory, Hong Kong. 


This species has been recognized as repre- 
senting a monotypic genus since its original de- 
scription, and Wakiya and Takahasi (1937) even 
placed it in a monotypic subfamily (Hemisalan- 
ginae). Apart from its somewhat shorter, blunter 
snout, and slight differences in dentition, how- 
ever, it is very similar to specimens of the sub- 
genus Salanx lacking a presymphyseal bone. The 
magnitude of the differences between Hemisa- 
lanx and Salanx (sensu stricto) is comparable to 
that between the latter and Leucosoma, which I 
also regard as only subgenerically distinct. 

Regarding S. brachyrostralis, Wakiya and Ta- 
kahasi (1937) placed it in the synonymy of S. 
prognathus with a question mark, as is done here. 


Neither Wakiya and Takahasi nor I have ex- 
amined the holotype (“S. 4227, 144 mm total 
length’). 

Wakiya and Takahasi (1937:293) placed Re- 
ganisalanx normani in the synonymy of H. pro- 
gnathus with a question mark, but I have ex- 
amined the type-specimens and have reidentified 
them as S. ariakensis. 


Salanx (Leucosoma) reevesii (Gray, 1831) 


?Albula chinensis OsBecK, 1757 (type-locality West River at 
Canton?; original not consulted). 

Leucosoma reevesi GRAY, 1831:4 (type-locality China). 

Leucosoma chinensis RICHARDSON, 1846:303. 

Salanx reevesii VALENCIENNES if CUVIER AND VALENCIENNES, 
1849:363, Pl. 646. 

Salanx chinensis GUNTHER, 1866:205. 


MATERIAL EXAMINED.—AMNH 10336, 130 mm, Fukien; 
AMNH 11161, 74.5 mm, Fukien; CAS-SU 1511, 4:87.8-140 
mm, Swatow (2:87.8—102 mm alcian-alizarin); CAS-SU 25738, 
127 mm, Canton; CAS-SU 61189, 153 mm, Castle Peak Bay?, 
Hong Kong. 


Salanx reevesi is second only to Protosalanx 
chinensis as the largest and heaviest-bodied sa- 
langoid. It is the only species of Salangidae in 
which males with anal scales have not been re- 
ported previously. A 130-mm specimen from 
Swatow (CAS-SU 1511) is a mature male with 
its anal fin greatly enlarged and 17 anal scales. 
It is the only member of the subgenus Leuco- 
soma, distinguished by a median row of 6-8 teeth 
on the tongue or basihyal bone. 

Albula chinensis Osbeck, 1765, has been iden- 
tified with this species by various ichthyologists 
following Richardson (1846) but, as pointed out 
by Wakiya and Takahasi (1937:291-292), its 
identity cannot be verified. While Albula chi- 
nensis evidently is a member of the Salanginae, 
there apparently are no type-specimen(s) extant 
(Kullander, pers. commun., 1983) and it is im- 
possible to tell from Osbeck’s description which 
species he had. The presence of teeth on the 
tongue, which would positively identify it as S. 
chinensis, is not mentioned. 

In addition to having teeth on the tongue, Leu- 
cosoma has jaw teeth that are somewhat larger 
than those in the subgenera Hemisalanx and Sa- 
lanx. In all other features of skeletal anatomy, 
however, Leucosoma falls in the genus Salanx. 
It has nearly the same vertebral counts as the 
subgenus Hemisalanx, and its cranial shape ap- 
pears to be identical with that in the subgenus 
Salanx. 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 211 


Salangichthyinae, new subfamily 


Cranium only moderately depressed (as in 
Sundasalangidae). Cranial fontanel with poste- 
rior and apparently anterior portions open 
throughout life. Head rounded anteriorly, pre- 
maxillae failing to meet at midline, not project- 
ing significantly beyond snout. Teeth greatly re- 
duced in size, small or minute, those on maxilla 
very numerous (about 15-40). Maxilla expand- 
ed, much larger than premaxilla. Body moder- 
ately elongate. Vertebrae 48-65 (66-79 in all oth- 
er Salangidae). 

This subfamily contains two genera, Neosa- 
lanx and Salangichthys, formerly placed in Pro- 
tosalanginae (Wakiya and Takahasi 1937). 


Neosalanx Wakiya and Takahasi, 1937 


Neosalanx WAKIYA AND TAKAHASI, 1937:282 (type-species, by 
original designation, Neosalanx jordani WAKIYA AND TA- 
KAHASI, 1937). 


The species of this genus are poorly known. In 
museum collections and older literature they are 
usually misidentified as Protosalanx or Salanx. 
Wakiya and Takahasi (1937) recognized four 
species, three of which they described as new. 
Although they recognized that Protosalanx an- 
dersoni Rendahl belonged to Neosalanx, they 
overlooked three previously described taxa, which 
also apparently belong to it: Protosalanx brevi- 
rostris Pellegrin, 1923; Protosalanx tangkahkeii 
Wu, 1931; and Salanx argentea Lin, 1932. All 
three of these were mistakenly placed by Fang 
(1934a:240) in the synonymy of Protosalanx hy- 
alocranius. 1 briefly examined the type-speci- 
mens of Pellegrin’s P. brevirostris in Paris, and 
have since studied radiographs of them, but have 
not seen the types of the taxa described by Wu 
and Lin. 

Wakiya and Takahasi (1937) distinguished four 
species of Neosalanx, mainly on the basis of dif- 
ferences in counts of vertebrae and fin-rays. 
Without knowing more about interpopulational 
meristic and other variation in Neosalanx, it is 
difficult to evaluate the species. Specimens I ex- 
amined tend to fall into groups, based on ver- 
tebral counts, identical to those recognized as 
species by Wakiya and Takahasi, and I have 
identified my material accordingly. 

Wakiya and Takahasi (1937) described Neo- 
salanx as a new genus despite the fact that one 
of its included species, Protosalanx andersoni 


Rendahl, 1923, was proposed by Fang (1934a) 
as type-species for his genus Paraprotosalanx. It 
is obvious from Fang’s account that he mistook 
specimens of Protosalanx chinensis and proba- 
bly either Neosalanx brevirostris or N. jordani as 
Protosalanx andersoni and that his generic di- 
agnosis is based mainly on P. chinensis. At this 
point the nomenclaturally parsimonious solu- 
tion would have been for Wakiya and Takahasi 
to recognize Paraprotosalanx as a valid genus 
with Protosalanx andersoni Rendahl, 1923 (not 
of Fang 1934a), as its type-species. They chose, 
however, to coin a new name and designated as 
type-species the new species N. jordani. Given 
that they did so, and that no publication has 
appeared subsequently in which Paraprotosa- 
lanx is treated except as a synonym of Neosa- 
lanx, I provisionally recognize Neosalanx as val- 
id. It should be noted, however, that judging from 
Article 70 of the International Code of Zoological 
Nomenclature (1964 ed.) this case should be re- 
ferred to the International Commission. 


Neosalanx andersoni (Rendahl, 1923) 


Protosalanx andersoni RENDAHL, 1923:92 (type-locality Chih- 
li, Shan-Hai-Kuan). 
Neosalanx andersoni WAKIYA AND TAKAHASI, 1937:285. 


MATERIAL EXAMINED.—NRM 10287, 79 mm, Chihli, Shan- 
Hai-Kuan (holotype). 


This species differs from all other Neosalanx 
by its relatively large size, to 100 mm (vs. only 
to 64 mm) and more numerous total vertebrae, 
63-65 (vs. 59 or less). Wakiya and Takahasi re- 
ported anal scale counts of 25-28 in N. andersoni 
and only 14-21 in other Neosalanx, but I find 
only 21 or 22 anal scales on the male holotype. 
N. andersoni is known only from rivers in Korea 
and China flowing into the Yellow Sea. 

The jaws of the holotype of Neosalanx ander- 
soni appear to be almost entirely toothless. I de- 
tected a single small conical tooth on the right 
premaxilla, none on the left. The dentaries ap- 
pear to be entirely toothless. The maxillae, al- 
though having irregularities in the margin where 
teeth may have been affixed, are toothless or al- 
most toothless except fora very few minute teeth. 
There seem to be no palatal or glossal teeth. 

The anterior and posterior portions of the cra- 
nial fontanel are widely separated and reduced 
in size (especially the anterior portion). There is 
no indication of a tectum medialis. 


212 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


A small patch of minute, scaly breeding tu- 
bercles occurs near the base of the 9th and 10th 
anal fin-rays, and more extensive areas of tu- 
bercles have evidently been sloughed off. Pelvics 
enlarged but without tubercles; lower jaw very 
strongly projecting. Snout elongate, as in Sa- 
langichthys (snout-tip to anterior margin of 
orbit =4.2 mm). 

Anal scale row continues onto ventral margin 
of caudal peduncle a considerable distance (4 
scales). 

Tip of snout dorsally and tip of lower jaw, 
dorsal surface of cranium over brain, dorsal sur- 
face of body with irregularly distributed small 
melanophores. No melanophores on anal fin, and 
none or almost none on caudal fin, ventral body 
surface, including pelvic and pectoral fin bases. 


Neosalanx brevirostris (Pellegrin, 1923) new 

combination 

Protosalanx brevirostris PELLEGRIN, 1923:351 (type-locality 
Tonkin). 

?Protosalanx tangkahkeii Wu, 1931:219 (type-locality Amoy). 

?Salanx argentea Lin, 1932 (type-localities Sangchang stream; 
Henngchow Bay; Canton). 

Protosalanx brevirostralis FANG, 1934a:232, 236, 240 (mis- 
spelled; referred to synonymy of Protosalanx hyalocranius). 

Neosalanx hubbsi WaktyA AND TAKAHASI, 1937:284 (type- 
locality Tien-tsin). 

Neosalanx tangkahkeii taihuensis CHEN, 1954? (reference not 
seen). 


MATERIAL EXAMINED.—AMNH 10337, 22:43.8-60.7 mm, 
Hunan (2:58.4-60.7 mm alizarin); AMNH 10480, 53.5 mm, 
Tung Ting Lake, Hunan; AMNH 11155, 60.5 mm, Fukien; 
AMNH 37044, 2:49.6-60.8 mm, Foochow, Fukien; AMNH 
51690, 26:48-65 mm, Canton; CAS 52032, 2:46.1—48.3 mm, 
Hong Kong (48.3 mm alizarin); CAS-SU 1540, 3:56.8-57.9 
mm, Swatow; CAS-SU 68888, 3:56.0-58.2 mm, Seoul; MNHN 
1922.184-189, 6:65-70 mm, Tonkin (syntypes of P. breviros- 
tris); USNM 219923, 2: 51.0-57.4 mm, Liang tsi Lake, Hupeh 
(formerly identified as N. tangkahkeii taihuensis). 


Protosalanx brevirostris has not been recog- 
nized as a valid species or even referred to since 
Fang (1934a), without examining the types, er- 
roneously placed it as a junior synonym of Pro- 
tosalanx hyalocranius. 1 briefly examined the 
types, which are in rather poor condition, during 
a visit to Paris in November, 1982, but did not 
have specimens of other Neosalanx on hand for 
comparison. I have since examined radiographs 
of the types. Since their vertebral counts—57(2), 
58(2), 59(1)—correspond only to those reported 
by Wakiya and Takahasi for Neosalanx hubbsi, 
I conclude that they are conspecific. 

Neosalanx brevirostris is known from the 


mainland coast of Asia, from the Yellow Sea 
coast of Korea south to Tonkin. 

In most specimens of Neosalanx the premax- 
illaries are toothless or have relatively few teeth 
(1-6 reported by Wakiya and Takahasi (1937), 
presumably based on examination of numerous 
specimens, and 0-2 in a large number of speci- 
mens I examined). Wakiya and Takahasi (1937) 
reported only 1-2 premaxillary teeth in N. hubb- 
si. Thus it is noteworthy that the 48.3-mm spec- 
imen from Hong Kong (CAS 52032) here re- 
ferred to this species has about 20 minute teeth 
on each premazlla. Tooth counts have not been 
made on the type-specimens of N. brevirostris. 

Despite considerable effort to trace its original 
description, the taxon Neosalanx tangkahkeii 
taihuensis is known to me only from an article 
by Wang et al. (1980), an abstract of which ap- 
peared in Aquatic Sciences and Fisheries Ab- 
stracts. The article reports on its artificial fertil- 
ization and larval development (see Addendum). 


Neosalanx jordani Wakiya and Takahasi, 1937 


Neosalanx jordani WAKIYA AND TAKAHASI, 1937:282 (type- 
locality ““River Rakuto, Corea”). 


MATERIAL EXAMINED.—AMNH 51705, 8:29-33 mm, Can- 
ton; AMNH 51704, 47.5 mm, Foochow, Fukien; CAS 52028, 
177:33.0-45.5 mm, Kiangsu, Chekiang Prov. (14:35.1-44.8 
mm alcian-alizarin); CAS 52029, 1:36.8 mm, Hong Kong; CAS 
52030, 311:22-58 mm, purchased in San Francisco; CAS-SU 
68625, 160:35.3-47.1 mm, and UMMZ 55601, 20:35.5-41.3 
mm, Pei-ho at Tien-Tsin (paratypes of Salanx hyalocranius 
Abbott, 1901) (10:39.7—45.7 mm alcian-alizarin). 


Wakiya and Takahasi (1937) characterize this 
species as having 49-54 vertebrae (average 50.95). 
The range of material I examined is 48-54. The 
species is known from rivers along mainland coast 
of Asia, from the Yellow Sea coast of Korea south 
to Hong Kong. 


Neosalanx reganius Wakiya and Takahasi, 1937 


Neosalanx reganius WAKIYA AND TAKAHASI, 1937:283 (type- 
locality ““Ariake Bay, Kyushu, Japan,” possibly erroneous). 
Neosalanx regani MATSUBARA, 1955:214 (unjustified spelling). 


Wakiya and Takahasi (1937:283) reported 52- 
56 (average 53.75) vertebrae for this species. It 
is known only from the type-specimens, sup- 
posedly collected in Ariake Bay. I have not ex- 
amined any Neosalanx from Japan. 


Salangichthys Bleeker, 1860 


Salangichthys BLEEKER, 1860:101 (type-species, by monotypy, 
Salangichthys microdon BLEEKER, 1860). 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 213 


Salangichthys microdon Bleeker, 1860:101 

Salangichthys microdon BLEEKER, 1860:101 (type-locality 
Yeddo-Tokyo). 

Salangichthys kishinouyei WAKtyYA AND TAKAHASI, 1913:552 
(type-locality Seishin, Korea and Shimane-ken, Japan). 


MATERIAL EXAMINED. —AMNH 13149, 28:50-64 mm, Dao- 
mori-ken, Japan; CAS 52033, 6:74.8-86.7 mm, Kawasaki; CAS 
52034, 9:31.6-36.1 mm, Lake Kituara (alcian-alizarin); CAS- 
SU 134, 10:78-92 mm, Tokyo (5:81-86 mm alcian-alizarin); 
CAS-SU 22637, 61:32-67 mm, Sendai, Matsushima Bay, Ja- 
pan. 


Salangichthys kishinouyei originally was dis- 
tinguished from S. microdon by Wakiya and Ta- 
kahasi (1913) on the basis of its having 1) man- 
dibular, maxillary, and palatal teeth more widely 
spaced; 2) snout shorter; 3) anal fin origin in 
females more anterior; and 4) threads of external 
egg membrane thicker, their ends club-shaped 
and ending freely without fusing to each other to 
form a ring. They later concluded that these dif- 
ferences represent intraspecific variation in pop- 
ulations of S. microdon on the Japan Sea coast 
of Sakhalin, Korea, Honshu, and Kyushu and 
placed S. kishinouyei as a synonym of S. micro- 
don (Wakiya and Takahasi 1937:279-280). 

Matsuoka and Iwai (1983, Fig. 2) illustrated 
an alcian-alizarin preparation of S. microdon (lo- 
cality not indicated) with 65 total vertebrae. The 
highest number of vertebrae otherwise known in 
Salangichthys is 63 (Table 2). 


Salangichthys ishikawae Wakiya and 
Takahasi, 1913 


Salangichthys ishikawae WakIYA AND TAKAHASI, 1913: 552 
(type-locality Miyagi-ken, Japan). 


MATERIAL EXAMINED.—CAS-SU 6780, 9:67.6-74.3 mm, 
Same, Rikuoku (3:70.3-74.2 mm alcian-alizarin); CAS-SU 
68878, 20:48-63 mm, Pacific coast of Aomori-ken, northern 
Honshu; CAS-SU 68889, 70.7 mm, Ariake Bay, Kyushu (lo- 
cality possibly erroneous). 


Salangichthys ishikawae is distinguished from 
S. microdon mainly by having 20-27 pectoral 
fin-rays instead of only 14-19 (see Senta 1973c). 
Differences in counts of vertebrae and anal scales, 
based on few specimens, are of doubtful signif- 
icance (see Table 2). 

According to Wakiya and Takahasi (1937:28 1), 
S. ishikawae “‘lives in water of rather higher sa- 
linity than S. microdon ... and seems to be a 
northern species,” known only on the Pacific coast 
of Honshu. The specimen herein reported from 
Ariake Bay, Kyushu, represents a southerly range 
extension of about 500 km if its locality data are 


correct. It was found with two specimens of Sa- 
lanx ariakensis collected by Kishinouye in Ar- 
iake Bay (CAS-SU 8574). 


Sundasalangidae Roberts, 1981 


Primary pectoral girdle consisting of single 
median element (identical to the condition in 
various larval teleosts, but unknown in adults of 
any other teleost). Pectoral fin rayless, supported 
by a single undivided basal radial. Pelvic fin with 
5 rays. Each half of pelvic girdle with two para- 
pelvic cartilages (unknown in any other teleosts). 
Adipose fin absent. Adult males without anal 
scales or sexually dimorphic anal fins. Neural 
spines much more elongate than in Salangidae. 
Hemal arches of last 12 or so abdominal verte- 
brae with elongate hemal spines (abdominal ver- 
tebrae without hemal spines in all other salan- 
goids). Vertebrae 37—43. Caudal fin skeleton with 
parhypural fused to hypurals 1 and 2 (parhypural 
separate in all other salangoids). 

If specialized is defined as deviation from the 
morphology of any known group of salmoni- 
forms that could possibly serve as the ancestral 
stock of Salangoidea, then Sundasalanx is by far 
the most specialized of all salangoids. 

The samples of Sundasalanx available at the 
time of their original description differed so much 
(in regard to eye size, non-overlapping vertebral 
counts, etc.) that it seemed to me that they could 
not be the same species. Samples examined sub- 
sequently, from Muar and the Mekong, tend to 
be intermediate between the two described 
species. I have tentatively identified the new 
samples with Sundasalanx praecox, but the 
question of the number of species in the genus 
should be reconsidered when more material be- 
comes available. The genus has not been found 
yet in Sumatra but is to be expected there. 


Sundasalanx Roberts, 1981 


Sundasalanx Roperts, 1981:297 (type-species, by original 
designation, Sundasalanx praecox Roserts, 1981). 


Sundasalanx microps Roberts, 1981 


Sundasalanx microps Rosperts, 1981:300 (type-locality Ka- 
puas River at Kampong Nibung, about 100 km NE of Sin- 
tang and 7 km NE of Selimbau). 


MATERIAL EXAMINED. —CAS 44220, paratypes, 34:14.6-19.9, 
Kapuas River at Kampong Nibung (paratypes) (7:13.5—18.0 
mm alcian-alizarin). 


214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


Sundasalanx praecox Roberts, 1981 


Sundasalanx praecox Roserts, 1981:299 (type-locality Khlong 
Falamee, Tale Sap, about 2 km W of Pak Payoon, Kra Isth- 
mus, Southern Thailand). 


MaTERIAL EXAMINED. —CAS 52031, 6:16.4-17.7 mm Khlong 
Falamee, Tale Sap, near Songkhla, Thailand (paratypes; alcian- 
alizarin); UMMZ uncat. and CAS 52036, 65:15.7—22.5 mm, 
Mekong River and tributaries in Thailand from Ban Dan to 
Nakon Phanom (4:18.2-20.3 mm alcian-alizarin, USNM 
229304 and CAS 52037, 121:12.3-18.0 mm, Muar River 7- 
18 miles inland from Bandar Maharani (formerly Muar City) 
(12:16.5-17.8 mm alcian-alizarin). 


When more material becomes available and 
the species of Sundasalanx are re-examined, the 
pigmentation of the Muar specimens (CAS 52037, 
USNM 229304) should be taken into consider- 
ation. In the Sundasalanx examined, each sam- 
ple tends to have a more or less distinctive pig- 
mentation or pattern of melanophore distribution 
shared by all of the specimens. Thus the type- 
series of S. microps is characterized by having a 
series of large midventral melanophores, one per 
myotome, extending from the pectoral fin to the 
anal fin. Midventral melanophores are absent in 
the type-series of S. praecox. In the Muar spec- 
imens here identified as S. praecox, however, a 
series of midventral melanophores extends from 
about pelvic-fin origin to the anal fin, thus re- 
sembling the pigmentation of S. microps. On the 
other hand, each of the Muar specimens has a 
series of large melanohores along the base of 
the anal fin, one melanophore between the base 
of each branched fin-ray. Other samples of Sun- 
dasalanx examined do not exhibit this feature. 

In the original description of S. praecox (Rob- 
erts 1981:299) it is stated that the type-series has 
maxillary teeth about 15-19 vs. about 30 in S. 
microps; the reverse is true. 


DISCUSSION 


Anyone who has looked into recent accounts 
of salmoniform classification realizes that it is in 
disarray. In this group current classifications are 
based largely on skeletal anatomy, and as long 
as the skeletal anatomy of major groups such as 
salangoids remain uninvestigated and others only 
partially investigated, no stable classification can 
be expected. McDowall (1969) briefly examined 
the skeletal anatomy of “Salangichthys micro- 
don” (actually Neosalanx?) stained with alizarin 
and concluded that Salangidae “‘is not part of the 
galaxioid radiation.”’ He also stated ‘“‘a more 


complete study (of salangid skeletal anatomy) 
may be desirable, should a full range of material 
be available, especially if it should reveal some 
species with more complete ossification to enable 
more precise determination of affinities.” A 
species with somewhat more complete ossifica- 
tion actually does exist—Protosalanx chinen- 
sis—but even in this species truly adequate ob- 
servations of skeletal anatomy cannot be based 
on specimens stained solely with alizarin. The 
question is no longer relevant, however, since 
adequate skeletal preparations of salangoids usu- 
ally can be obtained using the alcian-alizarin 
technique. 

All modern accounts of salangid classification 
agree in placing them in the order Salmoni- 
formes. My information on salangoid skeletal 
anatomy, however, has not provided me with 
obvious answers about their relationships to oth- 
er salmoniforms, but has only emphasized their 
distinctness. The presence of two (marginal or 
submarginal) rows of teeth on the basihyal tooth- 
plate of Protosalanx confirms the integrity of the 
order Salmoniformes and the placement of Sa- 
langoidea within this order but nothing more. 
The presence of a taenia medialis in the cranium 
of young Protosalanx and a single specimen of 
Neosalanx suggests a shared derived character 
(synapomorphy) with osmeroids, but this char- 
acter may well prove primitive for salmoni- 
forms, perhaps to be found in many of them. 

Higher classification of Salmoniformes has 
been the subject of considerable interest in the 
last two decades or so, with contributions by 
Gosline (1960), Greenwood et al. (1966), Weitz- 
man (1967), McDowall (1969), Rosen (1974), 
Klyukanov (1975), and Fink and Weitzman 
(1982). A major issue is whether esocoids (pikes, 
northern mud-minnows, and relatives) are Sal- 
moniformes, and should include the southern 
Lepidogalaxias, as advocated by Rosen. This 
view is contested by Fink and Weitzman, who 
exclude esocoids from Salmoniformes and relate 
Lepidogalaxias to Galaxiidae and osmeroids. 
Another major issue is whether relations of the 
southern “‘salmonoids”’ (Galaxiidae, etc.) lie with 
the northern salmonoids (Salmonidae, etc.) or 
with osmeroids. The hypothesis that stomiatoids 
are Salmoniformes (Weitzman 1967) has been 
rejected by Rosen (1974), Klyukanov (1975), and 
Fink and Weitzman (1982). Fink and Weitzman 
also point out that no satisfactory evidence has 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 215 


yet been advanced to support the hypotheses of 
relationships among osmeroids, salmonids, and 
galaxioids. Here I shall comment briefly on re- 
lationships of the salangoids to other Salmoni- 
formes, particularly osmeroids and galaxioids, 
and present some evidence bearing on relation- 
ships of Lepidogalaxias. 

Gosline (1960) suggested a close relationship 
of Osmeridae, Salangidae, and Plecoglossidae, 
distinguishing them from the Southern Hem1- 
sphere Retropinnidae, Aplochitonidae, and Ga- 
laxiidae. Greenwood et al. (1966) placed Salan- 
gidae in a suborder Galaxioidei including 
Retropinnidae, Galaxiidae, and Aplochitonidae 
without explanation. McDowall (1969) conclud- 
ed that Salangidae are a very specialized offshoot 
of the salmonoids and not part of the galaxioid 
radiation. 

In observing the skeletal anatomy of salan- 
goids I have been watchful for specialized char- 
acters indicative of phyletic relationship to other 
Salmoniformes. While my study has revealed 
highly specialized characters (such as the hyopal- 
atine) indicating monophyly of salangoids, it has 
not provided (or at least I have not noticed) char- 
acters that would link salangoids in a monophy- 
letic taxon with Osmeridae or any other group. 
Rosen (1974) pointed to specializations of the 
anal fin and associated scales in males of Lepi- 
dogalaxias and Mallotus, but concluded (p. 304) 
that these do not indicate relationship to Sa- 
langidae, and I agree. The anal scales and anal 
fin modifications of male salangoids appear to 
be a unique specialization, as does the hyopal- 
atine. Salangoids lack some skeletal features 
found in Osmeridae, notably in the ethmoid re- 
gion. Whether this absence is due to loss or re- 
flects a primitive condition is unclear. Devel- 
opment ofa tectum taenia medialis in the cranial 
fontanel, characteristic of the osmeroid chon- 
drocranium, occurs in some salangoids and may 
be indicative of relationship, but further obser- 
vation may reveal that the character is wide- 
spread in Salmoniformes. The peculiar mor- 
phology of the salangoid egg case (Wakiya and 
Takahasi 1937, Okada 1960) may be similar to 
that in osmeroids (compare photographs of Hy- 
pomesus olidus and Salangichthys microdon eggs 
in Chyung 1961, figs. 242-243). Similar spe- 
cializations possibly also occur in Plecoglossus 
but have not been reported in any of the northern 
salmonoids or in galaxioids. 


According to McAllister (1963:6) “the neo- 
tenic Salangidae strongly resemble larval Os- 
meridae but may be distinguished by their re- 
duced pointed head and elongated anterior 
portion of the body, as well as osteological 
characters.” Skeletal preparations of larval Os- 
meridae examined by me differ from salangoids 
in many respects. There are certain similarities 
in appearance of the largely transparent and lightly 
pigmented larvae of osmeroids and salangoids 
but these are mainly such as are to be found in 
larvae of non-related teleosts, and their value in 
assessing relationships is dubious. Skeletal anat- 
omy of larval osmeroids I have examined differs 
from that of salangoids almost as much as does 
skeletal anatomy of adult osmeroids. I have not 
examined skeletal anatomy of Plecoglossidae, but 
this family appears to be very specialized and 
there is no evidence that it is particularly closely 
related to Salangidae. Gosline (1960:346) and 
others have mentioned certain similarities be- 
tween the salangids and the extraordinarily spe- 
cialized galaxoid Lovettia but I believe such re- 
semblance is due to independently acquired 
neotenic characters. 

Considerable interest has centered on the phy- 
logenetic significance of the little salmoniform 
Lepidogalaxias salamandroides recently discov- 
ered in western Australia (Mees 1961). Among 
many peculiar features, it has in sexually mature 
males an extraordinarily modified anal fin par- 
tially covered by a sheath of anal scales sugges- 
tive of the anal scales of male Salangidae. The 
extremely complex modifications of the anal fin- 
rays go far beyond that seen in the anal fin of the 
salangids or any other salmoniform, and I doubt 
that in the relatively simple modifications of the 
anal fin in male salangids any uniquely shared 
specializations (or synapomorphies) with Lepi- 
dogalaxias can be recognized. The presence of a 
sheath of enlarged anal scales, on the other hand, 
demands closer comparison with those of sa- 
langids, which are otherwise unique among sal- 
moniforms (and perhaps all other teleosts). The 
anal scales in Lepidogalaxias are greatly enlarged 
and disposed in two main rows, rather than a 
single row, as in salangids. Two 31-—34-mm spec- 
imens I examined have seven to eight scales in 
the upper row and three in the lower. There also 
appear to be some scales or scalelike structures 
associated with the vent itself. Unlike that of 
salangids, however, the anal sheath covers the 


216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


mesopterygoid metapterygoid 
rae symplectic hyomandibula 
alatine 
: interhyal opercle 


premaxilla 


re he sha 
maxilla 
dentary 
articular 
angular 
quadrate 


1mm preopercle 
see ee ey 


subopercle 


interopercle 


FiGure 22. Lepidogalaxias salamandroides (uncat., 23.5 mm). Jaws, facial ones, and suspensorium (lateral view). 


anal fin, especially the modified portions. In sa- 
langids the anal scales are on the body above the 
anal fin, and the rays themselves are entirely ex- 
posed. Whatever the phyletic significance, it 
would certainly be of interest to know more about 
the functional significance of the anal scales and 
modified anal fins in Lepidogalaxias and in Sa- 
langidae. 

Fink and Weitzman (1982) suggested that a 
single row of mesopterygoid teeth is a synapo- 
morphy indicating monophyly for the osmeroids 
(including Salangidae) and galaxioids (including 
Lepidogalaxias). Although Gosline (1960) stated 
that the mesopterygoid is absent in Salangidae, 
it is actually present in most of them. But while 
most Salangidae have palatal teeth, the salangoid 
mesopterygoid is invariably toothless, even in 
Protosalanx and Salanx chinensis which have 
well-developed basihyal teeth. Thus the concept 
of a “‘tongue-bite” character, based on basihyal 
and mesopterygoid teeth, and uniting galaxioids 
and osmeroids, does not hold for salangoids. 

While my observations of salangoid skeletal 
anatomy have not provided me with the key to 
their higher relationships, I earnestly hope that 
they may do so for future workers who are able 
to make more extensive comparisons. In ex- 
amining Lepidogalaxias I find no characters in- 
dicative of close relationship to Osmeridae or 


Salangidae, but the structure of the jaws and gill 
cover reveals specialized characters linking this 
strange western Australian fish to the southern 
galaxioid radiation. 

Based largely on the assumption that Lepi- 
dogalaxias possesses uniserial mesopterygoid 
teeth, and without having examined skeletal ma- 
terial, Fink and Weitzman (1982) suggested its 
relationships lie with osmeroids and galaxioids. 
I have examined skeletal anatomy of several lar- 
val and juvenile or subadult Lepidogalaxias, the 
largest of these has a well-developed median patch 
of teeth on the prevomer and a pair of well- 
developed tooth patches on the anterior palatine 
area (ectopterygoid? = endopterygoid of Mees 
1960) but the large, well-developed mesoptery- 
goid is toothless. On the other hand, specializa- 
tion of bones in the jaws and gill cover (Fig. 22) 
indicates relationship with galaxioids. The ex- 
traordinary fimbriate condition of the bony oper- 
cle, also of the subopercle, appears to be a highly 
specialized character shared only (i.e., synapo- 
morphic) with galaxioids (see McDowall 1969, 
Fig. 3b-e, and Rosen 1974, Fig. 13). The short 
and laterally directed premaxilla and toothless 
maxilla with its strongly concave margin may 
represent another synapomorphy with galax- 
ioids. My observations suggest that Lepidoga- 
laxias is indeed related to Galaxiidae, as sug- 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 217 


gested by Nelson (1970), but I doubt that it 
represents the “‘primitive sister group of galaxiids 
or galaxioids”’; it is more likely to be a highly 
specialized galaxiid or galaxioid derivative. As 
pointed out by Scott (1966), its osteology should 
be examined carefully and thoroughly; this has 
yet to be done. For the present I would like to 
point out a difference between the branchial 
arches of Lepidogalaxias and Umbridae which 
merits further investigation. In Lepidogalaxias 
and Umbridae the basibranchial plate has five 
basibranchials, and the fifth is cartilaginous. In 
Lepidogalaxias the basibranchial plate is appar- 
ently immobile; that is, the basibranchials ap- 
parently do not move backward and forward in 
relation to each other. In Umbridae, however, 
as exemplified by Novumbra hubbsi, basibran- 
chial 5 is movably articulated to basibranchial 4 
in such a fashion that it can be rocked back and 
forth beneath it; and thus the pair of toothplate- 
bearing fifth ceratobranchials, firmly attached to 
basibranchial 5, are also moved back and forth. 
Whether such basibranchial mobility occurs in 
other Umbridae or in esocoids generally is un- 
known; it has not been observed in galaxioids, 
osmeroids, or salangoids (the latter apparently 
lack basibranchial 5). 

In assessing phylogenetic relationships one 
should not be overly impressed by the presence 
of primitive characters, even in groups in which 
such characters supposedly have been lost for a 
long time. 

Atavism, the expression of ancient characters 
“buried in the genome,” occurs far more often 
than generally recognized. This, rather than a 
Lamarckian interpretation, is doubtless the cor- 
rect explanation for the appearance of breeding 
tubercles on the palms of the midwife toads (A/- 
ytes obstetricans) painstakingly studied by Kam- 
merer (see Koestler 1973). Characters such as 
eyes, teeth, scales, bone, or the pelvic girdle may 
be repeatedly suppressed, and may not be phys- 
ically expressed in any members of quite large 
groups, without ever having been lost from the 
genome. For purposes of phylogenetic analysis, 
I suggest it is parsimonious to assume that prim- 
itive characters are never lost from the genome, 
and that this is really why it is futile to rely on 
them. Thus the fully scaled condition of Lepi- 
dogalaxias does not suggest to me that it rep- 
resents the primitive sister group of the otherwise 
scaleless galaxioids. 


I believe that utilization of the caudal skeleton 
as a guide to phyletic relationships among te- 
leosts, especially those with the primitive teleost 
complement of six separate hypurals and 10+9 
principal caudal fin-rays, has inevitably resulted 
in confusion of primitive with advanced char- 
acteristics in the caudal fin skeleton. While te- 
leosts as a whole exhibit great diversity in their 
caudal skeleton (Monod 1968), the main features 
of the caudal skeleton are remarkably similar in 
many teleosts with forked caudal fins and the 
primitive complement of principal caudal fin- 
rays. Thus the caudal fin skeleton of salangoids 
is strikingly similar in many respects to that of 
Elops, many clupeoids, characoids, and cypri- 
noids as well as of osmeroids and other salmon- 
iforms. There are two possible explanations (or 
hypotheses) for such similarities, both predicated 
upon the assumption that the caudal fin structure 
of Elops and the others is primitive for teleosts. 
The first, and traditional, explanation is that all 
descended from ancestors that never deviated 
from morphologically primitive caudal fins. The 
second is that teleosts with secondarily general- 
ized caudal fins have repeatedly reverted to a 
primitive type of caudal fin skeletal morphology. 

Salangoids often have been referred to as neo- 
tenic. According to Jordan and Snyder (1902: 
592), “the straight alimentary canal, distinct 
muscle segmentation, very thin ventral wall of 
the abdominal cavity, and other characters of 
salangoids suggest a larval stage of develop- 
ment.’’ Among other characters, they were cer- 
tainly thinking of the near perfectly transparent 
state of the fish in life. Thus when the fish die, 
the flesh turns perfectly white, and, in the absence 
of scales, the myotomes stand out very clearly, 
as in many larval fish. Berg (1947), Gosline 
(1960), Weitzman (1967), and McDowall (1969), 
remarked that Salangidae seem to be neotenic 
but did not elaborate. If the term neotenic means 
simply attaining sexual maturity while retaining 
some larval characteristics, salangoids are surely 
neotenic, probably more so than any other sal- 
moniform fishes. The following characteristic 
features of salangoids are among those which 
may be neotenic: 

1) Body almost entirely transparent in life. 

2) Pectoral fin pedunculate, with a broad, fleshy, 
pedestallike base more or less free from the lat- 
eral musculature of the body, and placed high on 


218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13 


the side of the body, just like the pedunculate 
pectoral fin of many lower teleost larvae. 

3) Skeleton in large measure cartilaginous; 
many bones found in adults of other salmoni- 
forms absent. 

4) Body scaleless throughout life except for 
anal scales in sexually mature male salangids. 

5) Ventral body musculature incomplete, ev- 
idently due to an arrested ventral myotomic pro- 
gression, so that the ventral abdominal wall is 
thin and non-muscular. 

6) Left and right halves of pelvic girdle more 
or less widely separated from each other, failing 
to form a cartilaginous union by means of pos- 
teromedian processes as in most other salmon- 
iforms. 

7) Abdomen posterior to pelvic fin with a 
membranous median keel. 

8) Fourth gill arch with well-developed hy- 
pobranchials. 

9) Primary pectoral girdle consisting of a me- 
dian scapulocoracoid in Sundasalangidae (a con- 
dition also present in larvae of E/ops and other 
lower teleosts). 

10) Dorsal and anal fins placed relatively far 
posteriorly (least so in Protosalanx). 

On the other hand, salangoids exhibit consid- 
erable diversification and a number of peculiar 
modifications or specializations which are ob- 
viously not neotenic, including: 

1) Marked sexual dimorphism in Salangidae, 
involving enlarged pectoral and anal fins, mod- 
ified anal fin-rays, and development of the anal 
scales in sexually mature male salangids. 

2) Strongly depressed cranium and _ skull: 
marked ventromedian curvature of maxilla. 

3) Voracious feeding habits and canine den- 
tition, especially in Salanginae. 

4) Vertebral counts of 37-79, the highest counts 
found in Salanginae with excessively elongate 
body form. 

5) Proximal radials of pectoral fin highly mod- 
ified in all Salangidae; pectoral fin-rays very nu- 
merous in all Salangidae except Salanginae. 

6) Fusion of cartilaginous hyomandibular and 
mandibular arches to form a hyopalatine. 

7) Fusion of basibranchials and hypobranchi- 
als in gill arches of Sundasalangidae. 

Thus salangoids resemble larval fish in many 
ways. But the problem arises of distinguishing 
between characters that are truly neotenic and 
characters that represent convergence of adults 
with larvae. Upon first observing the beautifully 


simple condition of the primary pectoral girdle 
in Sundasalanx, and taking into consideration 
the fused condition of ventral elements in the 
branchial arches of Sundasalanx and of the 
pterygoquadrate and hyomandibula in all sa- 
langoids, I was inclined to regard it as due to 
secondarily evolved simplification and/or re- 
duction and fusion of the primitively separate 
left and right halves. But a morphologically iden- 
tical “‘median” pectoral girdle and “‘fused”’ radial 
plate was reported in larval clupeoids by Good- 
rich (1922) and I have found it in larval Elops 
hawaiiensis (Fig. 19b) and anchovies. In all of 
these larvae, as in Sundasalanx, the scapulocor- 
acoid has three clearly divided portions (a trans- 
verse median bar, an ascending process, and a 
posterior projection) and the basal radial or plate 
is perforated by three foramina. In further de- 
velopment, the three foramina of the pectoral 
plate presumably enlarge until the basal plate in 
all of these forms except Sundasalanx divides 
into proximal radials 2-5. (The primitive num- 
ber of proximal radials for all teleosts appears to 
be 5, observed even in many teleosts having 
highly modified pectoral fins.) Swndasalanx is 
the only known fish which retains a median scap- 
ulocoracoid at sexual maturity. The other salan- 
goids presumably have such a median pectoral 
girdle as larvae, but although the scapulocora- 
coid is apparently cartilaginous in all Salangidae 
examined, it is clearly divided into two halves 
in all specimens examined, including Salangich- 
thys microdon of only 39 mm SL. Goodrich 
(1922:508) tentatively concluded that the fusion 
of the endoskeletal pectoral girdle he found in 
young clupeids represents a specialization pe- 
culiar to larval Clupeidae. The present finding 
that morphologically identical girdles occur in at 
least some Elopomorpha and Salmoniformes as 
well suggests that it is indeed a primitive char- 
acteristic of teleosts. The duration of the median 
condition apparently corresponds more or less 
closely with a period when the pedunculate pec- 
toral fins are being used most actively, the pri- 
mary pectoral girdle is still largely or entirely 
cartilaginous, and the mesocoracoid arch has not 
developed. 


ADDENDUM 


The galley proofs of this paper were already 
set when Prof. Xin-Luo Chu of the Kunming 
Institute of Zoology of Academia Sinica visited 
the California Academy of Sciences (January 


ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA 219 


1984) bringing specimens of Neosalanx tangah- 
keii taihuensis for me to examine. These speci- 
mens, CAS 54330, 4: 69.7—76.8 mm, were caught 
in Kunming Lake where the species was intro- 
duced in 1981 from artificially reared stock orig- 
inating in Lake Taihu. The specimens have total 
vertebrae 56(1), 57(2), 59(1) and gill rakers 15(2), 
16(1), 18(1). I therefore conclude that N. tan- 
gahkeii taihuensis is a junior subjective synonym 
of N. brevirostris. N. brevirostris 1s a valuable 
commercial fish and much of the production (es- 
pecially of Lake Taihu) is marketed abroad. In 
the near future it may be widely introduced in 
lakes in China which lie outside the natural range 
of Neosalanx. 

It may be worthwhile for Chinese workers to 
investigate the aquacultural potential of the other 
two species of Neosalanx occurring in China. N. 
andersoni, which may grow slightly larger than 
N. brevirostris, has a relatively restricted north- 
erly distribution, indicating that it is adapted to 
colder waters. N. jordani, the smallest species, 
has a wide range largely overlapping that of N. 
brevirostris, but my observations indicate that 
the two species usually do not occur together in 
nature. N. jordani has fewer gill rakers and this 
together with its smaller size indicates a probable 
difference in feeding habits. It could be of prac- 
tical as well as scientific value to compare the 
ecology and fisheries biology of these three 
species. 


ACKNOWLEDGMENTS 


For providing specimens, information, advice, 
or assistance I wish to thank the following per- 
sons: Tokiharu Abe, Janine Abel, Gerald R. Al- 
len, Vladimir Barsukov, Marie-Louise Bauchot, 
W. H. Butler, Xin-Luo Chu, Alexandra Creigh- 
ton, Oliver Crimmen, Robert Drewes, Norma 
Feinberg, Bo Fernholm, W. I. Follett, Michael 
Hearne, Susan Jewett, Sven Kullander, Robert 
McDowall, Gareth Nelson, Han Nijssen, Nicolai 
Parin, Ingrid Radkey, Walter Rainboth, Donn 
E. Rosen, Lars Wallin, Stanley H. Weitzman, 
and Alwyne Wheeler. 

This study was carried out mainly in the De- 
partment of Ichthyology of the California Acad- 
emy of Sciences, National Marine Fisheries Lab- 
oratory at Tiburon, and the Tiburon Center for 
Environmental Studies (of San Francisco State 
University), and was supported in part by grant 
DEB77-24574 in the Systematic Biology Pro- 
gram of the National Science Foundation. 


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Vol. 43, No. 14, pp. 221-238, 17 figs., 1 table 


PREDATORY BEHAVIOR OF THE WHITE SHARK 
(CARCHARODON CARCHARIAS), WITH 
NOTES ON ITS BIOLOGY 


By 
Timothy C. Tricas 


Department of Zoology, University of Hawaii at Manoa, 
Honolulu, Hawaii 96822 


and 
John E. McCosker 


Steinhart Aquarium, California Academy of Sciences, 
Golden Gate Park, San Francisco, California 94118 


Asstract: The feeding behavior of the white shark (Carcharodon carcharias) was studied at Dangerous 
Reef, South Australia. Cinematographic analyses of shark feeding patterns show that a single bite action is 
comprised of a uniform sequence of jaw and head movements. The components are: 1) snout lift, 2) lower- 
jaw depression, 3) palatoquadrate protrusion, 4) lower-jaw elevation, and 5) a bout-ending snout drop. Du- 
rations for a complete bite action ranged from 0.750 to 1.708 s (x = 0.985 s) for a 3.5 m (TL) subject. Various 
approach behaviors to baits were also documented. 

The stomach contents of nine white sharks captured in northern and central California waters consisted 
entirely of fish prey associated with inshore and pelagic habitats. Records of the stomach contents of 24 
additional sharks were combined and analyzed, and indicated fish to be the most frequent prey items, while 
marine mammals were also common. Analysis of prey type in relation to shark size shows small sharks (<3 
m) feed primarily on fish prey, while larger sharks feed on marine mammals, especially pinnipeds. 

Cursory field experiments and observations indicate sharks detect and are attracted to electric fields. 
Telemetric studies of white shark thermal biology show that they are warm-bodied, approximately 4-5°C 
above ambient water temperature. = 

Length-weight records for 127 sharks were analyzed and found to have the relationship: W = 3.8 x 10-° 
L*+5, where W is weight in kg and L is length in cm. The largest reliable record for a white shark is that of 
a 6.4-m, 3324-kg specimen captured near Cojimar, Cuba, in 1945. 

A hypothesis is proposed to explain the “bite and spit” paradox related to attacks on pinnipeds and humans. 
Comments concerning the risk associated with contemporary surfboard design are included. 


INTRODUCTION Collier 1964: Follet 1974: Ellis 1975: McCosker 


The white shark (Carcharodon carcharias) (Fig. 1981). It is circumglobal in distribution, but most 
1) is the largest piscivorous marine fish in the commonly inhabits the coastal temperate waters 
world and is well known for its aggressive be- of North America, South Africa, and South Aus- 


havior and potential threat to humans (Fast 1955; tralia. 
[221] 


222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


Ficure |. 


In spite of its size and fearsome reputation, 
surprisingly little is known about the natural his- 
tory and behavior of this large fish. Almost all 
published information on the general biology of 
white sharks comes from anecdotal observations 
and notes obtained from commercial fishing or 
whaling operations (Squire 1967), regional species 
lists and range extensions (Bigelow and Schroe- 
der 1948; Day and Fisher 1954; Royce 1963), 
and newspaper articles on captures by fishermen 
or accounts of attacks on humans. 

The predatory behavior and feeding mechan- 
ics involved in prey capture by white sharks has 
remained, until the recent application of scuba 
and high speed photography, essentially un- 
known. Previous studies on the feeding mor- 
phology of other species of sharks were based 
largely on anatomical data where muscle and 
supportive tissue functions were inferred from 
examination of preserved specimens (Luther 
1909; Haller 1926). This approach provided 
functional insight, based largely on articulations 
and spatial arrangements of skeletal tissues and 
head musculature. In some cases, however, the 
inflexibility of preserved materials has led to 
misinterpretations of the true mechanics of jaw 
protrusion and feeding in sharks (see Compagno 


Tagged male white shark swimming near the surface at Dangerous Reef, South Australia. Photo by Al Giddings. 


1977). Whereas examination of fresh pliant spec- 
imens may be more appropriate for functional 
analyses, they still provide only speculative data 
on sequential and temporal relationships of 
structures involved in feeding activity. Moss 
(1972) provided a qualitative analysis of feeding 
mechanisms in living carcharhinid sharks using 
observational, photographic, and electrical mus- 
cle stimulation techniques. Studies on the tem- 
poral and sequential mechanics of feeding be- 
havior in sharks are still lacking, however, when 
compared to the more thorough cinematograph- 
ic studies on teleostean fishes (Osse 1969; Liem 
1978; Lauder 1980). 

Because white sharks are rarely captured, doc- 
umentation of their food habits is scattered 
throughout the literature. Most records come 
from notes on the stomach contents of dead fish 
(Schroeder 1938; Bonham 1942; LeMier 1951; 
Scattergood 1962) or from fortuitous observa- 
tions of feeding in the field (Day and Fisher 1954; 
Pratt et al. 1982). More complete accounts are 
provided on the relationships of white sharks to 
pinnipeds (Ainley et al. 1981; and Le Boeuf et 
al. 1982) and sea otters (Ames and Morejohn 
1980). There still remains, however, the need for 
a comprehensive collation of the prey items tak- 


a 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 223 


en by this predator so that a more complete as- 
sessment of predator-prey relationships can be 
made. 

This paper presents new data obtained during 
a recent expedition to South Australia that relate 
to white shark predatory behavior and general 
biology. We analyze the feeding mechanics of 
white sharks in the field by use of cinemato- 
graphic techniques and provide information on 
their sensory biology and thermal physiology. In 
addition, we have synthesized previously pub- 
lished and unpublished data on the length-weight 
relationships, predator-prey interactions, and 
general behavior of this shark. Based upon what 
is known about the predatory behavior of white 
sharks, we present a new interpretation of the 
curious pattern of non-feeding attacks upon ma- 
rine mammals and humans. 


Stupy AREA AND METHODS 


White sharks were studied in the field during 
a ten-day period in January 1980, in waters near 
Dangerous Reef, South Australia. The reef con- 
sists of two small, low islands approximately 16 
km east of Port Lincoln (Fig. 2). Sharks were 
attracted to the 20-m vessel, Nenad, using tuna 
and meat byproducts as bait. Sharks feeding on 
baits both at and below the surface were pho- 
tographed using Actionmaster 500 cameras and 
7247 Kodak color reversal film exposed at shut- 
ter speeds of 24 and 200 frames per second. 
Frame-by-frame analyses were performed on a 
Movieola 16-mm film editor. 

Stomach content and morphological data from 
nine sharks on record at the California Academy 
of Sciences were analyzed. These data were then 
combined with other published records and fur- 
ther examined. To prevent multiple entries of a 
record into the analyses, only well-documented 
reports that included information on capture lo- 
cality, number of sharks sampled, measured 
lengths and weights (no estimations), and specific 
prey types were used. 

Two types of ultrasonic telemetry packages, 
constructed by the senior author, were used in 
this study to monitor shark body and ambient 
water temperatures. All transmitter circuits con- 
sisted of a crystal-controlled oscillator (carrier 
frequencies = 31.700 or 32.768 kHz) gaited by 
a thermistor-controlled pulse circuit sensitive 
from 10°C to 33°C. The first tag consisted of a 
single transmitter with a thermistor probe 


angerous Ree! 


a. Se \ 
{AUSTRALIA 4 


= ets 
Vv) 


ge 


FiGurReE 2. 
lia. 


The study area, Dangerous Reef, South Austra- 


(embedded on the surface of the transmitter 
housing) that monitored ambient water temper- 
ature around the animal. Its dimensions were 
4.6 x 3.2 x 2.0 cm, and it weighed approxi- 
mately 60 g in air. The second unit consisted of 
a cylindrical package with two transmitters of 
different carrier frequencies. One transmitted 
temperature data from a thermistor in contact 
with the surrounding water, the other from a 
thermistor embedded under the barb of a dart at 
the end of a 31 cm-long wire leader. Total pack- 
age dimensions were length 17 cm x diam. 3.2 
cm, with a weight of approximately 100 g in air. 

The water temperature sensing package was 
applied from underwater using scuba (and a cage). 
A stainless steel dart was attached to an appli- 
cator tip on the end of a speargun shaft, and shot 
3 cm deep into the shark’s mid-lateral muscu- 
lature. The dual-temperature sensor package was 
applied externally to another shark from the 
swimstep of the research vessel via barb and ap- 
plicator pole. Signals were tracked with a tune- 
able ultrasonic receiver and a staff-mounted di- 
rectional hydrophone. Absolute maximum range 
of the transmitter-hydrophone system under ide- 


224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


5000 4 


In W = -12.47 +3.15 (In L) e 
r=98 '. 


n=127 400 fe 

100 e Pr 
see 

" - tat 

2 ee e 5 

= e 

= a fa Gee 

® O e 

= 10 3° 


10 + 1 r 1 r 
100 200 300 400 500 
Total Length (cm) 


5 1 
600 700 


Ficure 3. Length-weight relationship for the white shark. 
Data taken from complete records for 127 sharks. Functional 
(geometric mean) regression equation given on figure (see Rick- 
er 1973 for discussion). Non-transformed power equation for 
relationship is W = 3.8 x 10~¢ L3!5, where W = weight in kg 
and L = total length in cm. 


al conditions was approximately 1500 m. How- 
ever, practical working distances were much less 
due to transmission loss in the shallow waters 
around the reef (20-30 m deep). Ranges were 
estimated by relative audible strength calibrated 
prior to tracking sessions. 

A set of cursory experiments were performed 
to test the sensitivity of sharks to weak electric 
fields. Sharks attracted to the boat by chum were 
presented two pieces of bait, approximately one 
meter apart, suspended on the surface from lines 
attached to the end of 7-m bamboo poles. Two 
saltwater electrodes, similar to those used by 
Kalmijn (1978), were attached to one bait (the 
experimental). Electrodes consisted of one-meter 
lengths of 6.3 mm inside diameter tygon plastic 
tubing filled with a 3% seawater-agar gel. One 
end of each tube was open to the water, while at 
the other end a 32-mm stainless steel pin with 
wire lead was inserted. Lead wires (+ and —) 
were connected to a Grass S-6 stimulator. The 
two saltwater electrodes were attached behind 
the experimental bait and spaced 10 cm apart. 
The control consisted of bait only. The experi- 
mental bait was presented in two different ex- 
periments that used 1) pulsed (2.2 volts at source, 
5 Hz, 1.9 ms duration) current, and 2) constant 
(0.5 and 2.2 volts at source, DC) current elec- 
trical fields. Each test sequence began when a 
shark visually oriented to and approached the 
baits. Once the shark was within 2 m of the baits, 
the stimulator was turned on and choice of bait 
fed upon recorded. Relative positions of the con- 


trol and experimental baits were randomly de- 
termined to control for extrinsic cues. 


RESULTS AND DISCUSSION 


SizeE.— The length-weight relationships of 127 
white sharks, based on records at the California 
Academy of Sciences and those of published and 
contributed sources, are shown in Fig. 3. The 
largest shark record we found was that of a 6.4- 
m-long (21 ft), 3324-kg (7302-Ib) individual cap- 
tured off Cojimar, Cuba, in 1945 (Guitart and 
Milera 1974). The maximum size previously re- 
ported for a white shark originated from an in- 
correct record of an 1 1.1-m individual from Port 
Fairy, Australia, reported by Giinther (1870). 
Randall (1973) re-examined the jaws of this spec- 
imen and concluded that the correct total length 
was approximately 5 m, well within the size dis- 
tribution of sharks currently on record. The pur- 
ported capture of a 9-m (29.5-ft) white shark said 
to be from Vila Franca, Azores, is probably er- 
roneous (see Ellis 1983). The smallest published 
record was a 125-cm (49-in), 20-kg (44-Ib) spec- 
imen reported by Smith (1951). Robert Johnson 
(pers. comm.) has advised us of three juveniles 
captured off Baja California that ranged from 130 
to 135 cm (51 to 53 in) total length and weighed 
less than 18.2 kg (40 Ib). 

Remarkably, we found no well-documented 
records of female white sharks with fetuses or 
pups. Bigelow and Schroeder (1948) reported 
embryos ranging in length from 20 to 61.6 cm, 
but gave no further source information. One fe- 
male taken near Alexandria, Egypt, was reported 
to have nine embryos, each 0.61 m (2 ft) long 
and weighing 49 kg (108 lb). This erroneous 
weight probably represents a total for all nine 
embryos, and translates to a more reasonable 
mean of 5.4 kg (12 Ib) for each fish. We can only 
speculate about this lack of pregnant females in 
the capture record. Females may pup in less fre- 
quently sampled areas, such as remote geograph- 
ic regions, oceanic waters, or deeper pelagic hab- 
itats. Although it is possible that females fast 
while pregnant, this would not completely ac- 
count for the phenomenon, because many of the 
largest females on record were taken by harpoon 
rather than with bait and hook. Perhaps pregnant 
females undergo spontaneous parturition when 
hooked or harpooned and therefore eliminate key 
embryonic evidence before they are landed. More 
critical examination of the reproductive tract of 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 


freshly landed specimens might provide useful 
insight to this enigma. 

FEEDING ETHOLOGY.— The following section is 
based on our observations and the analyses of 
films taken of white sharks feeding on bait at 
Dangerous Reef, Australia. Although baited sit- 
uations can only simulate natural conditions, the 
feeding behaviors observed in these sessions rep- 


Ficure4. Still photographs of white sharks feeding on baits 
near Dangerous Reef, Australia. (a) Shark begins to raise snout 
and depress lower jaw. (b) Mouth opened fully with head and 
snout raised. (c) Palatoquadrate protrusion and lower-jaw el- 
evation. (d) Mouth closéd; head is raised and disassociated 
from upper jaw. (e) Head and snout drop to normal position. 
Photos a, c, d, e by T. Tricas. Photo b by P. Romano. 


resented natural patterns because white sharks 
normally take prey at the surface (Ainley et al. 
1981; personal observations). 

The following descriptions of the structures 
and mechanics involved in biting actions of white 
sharks employ terminology similar to that used 
by Moss (1972, 1977). Cinemaphotographic 
analysis of 36 feeding bouts revealed five basic 


226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


components that constitute a single feeding ac- 
tion. 

1) Snout Lift: This movement involves an up- 
ward lifting of the snout and head, and initiates 
the feeding action (Figs. 4a and b, 5b). The degree 
of snout lift ranged from a slight upward move- 
ment to a pronounced elevation that produced 
an acute angle behind the head (30—40° above 
the longitudinal body axis). The intensity of snout 
lift varied in relation to size of bait, angle of 
approach to the bait, and possibly to level of 
motivation (e.g., hunger). 

2) Lower-Jaw Depression: Like the snout lift, 
lower-jaw depression occurs at the start of a feed- 
ing action. It is characterized by a ventro-pos- 
terior movement of the tip of the lower jaw (Figs. 
4a and b, 5b). This motion, along with the snout 
lift, fully extends the gape. 

3) Palatoquadrate Protrusion: Closure of the 
mouth is marked by disassociation of the upper 
jaw from its original juxtaposition ventral to the 
cranium, and subsequent protrusion out of the 
oral cavity. The upper jaw rotates in an antero- 
ventral direction, while the snout remains at its 
elevated position (Figs. 4c, Sc). During palato- 
quadrate protrusion the teeth become fully ex- 
posed and are directed downward. Eversion of 
the upper jaw was readily visible by exposure of 
the reddish connective tissue on the surface of 
the jaw cartilage. 

4) Lower-Jaw Elevation: Concurrent with the 
initiation of palatoquadrate protrusion, the low- 
er Jaw begins an antero-dorsal (upward) motion 
(Figs. 4c, 5c). These movements collectively pro- 
duce the closing action of the jaws. 

5) Snout Drop: After single-bite feeding bouts 
the snout returns to its normal pre-feeding po- 
sition. This results from a drop of the head and 
snout, and a retraction of the palatoquadrate car- 
tilage to its position immediately ventral to the 
cranium (Figs. 4e, 5d). During multiple-bite bouts, 


— 


Ficure 5. Components ofa feeding action pattern. (A) Shark 
just prior to initiation of feeding action. Snout and lower jaw 
are at normal resting position. (B) Snout lift and lower-jaw 
depression result in maximum gape. (C) Palatoquadrate pro- 
trusion rotates upper jaw forward and downward exposing 
upper teeth. Lower jaw moves forward and upward. These two 
components comprise the actual bite. (D) Snout drop entails 
retraction of palatoquadrate cartilage to its normal juxtapo- 
sition beneath cranium. Snout drop occurs at the end of a 
feeding bout and is not an essential component of the biting 
action. Arrows indicate direction of jaw movements. 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 


to 
tO 
— 


Ae B reo 
5 
Tp) E +—@—4 

B re4 
” 
4) 
o 
& | Maximum 
© ie 
Q | Gape 
Se 
— | 

E -—e—1 
a 
Ole — 
a 
rs B — oo 
w 
Se E -+—e—J 
Qa oo 
° E 
fa 
Wa lz ——_———— e 4 

1 EES EE =! ———) SSE 
10) -l] a. 23 4 ) 6 if 8 9 1.0 1] 1.2 
Time (s) 


Ficure 6. Timing of feeding actions for eleven consecutive bites made by a 3.5 m (TL) white shark. Mean times indicated 
by dots. Horizontal lines show 95% confidence limits. Key: B = begin, D = depression, E = end, Elev = elevation, LJ = lower 
jaw, PQ Prot = palatoquadrate (upper jaw) protrusion, S = snout. 


the snout remains partially elevated prior to the 
next biting action (Fig. 4d). The retention of an 
elevated snout in these cases results in shorter 
time intervals between bites. 

Mean durations for components of eleven 
complete successive feeding events recorded for 
one shark are shown in Figs. 6 and 7. Depression 
of the lower jaw was the fastest component (xX = 
0.140 s), and the snout drop duration the longest 
(x = 0.405 s). Total time for a complete biting 
action, including the snout drop, ranged from 
0.750 s to 1.708 s (X = 0.985 s). Temporal anal- 
yses of film footage showed that the sequence of 
each feeding component fell in a fixed order with 
a non-overlapping range of time limits for each 
individual head and jaw movement. While each 
action showed a range in timing, minimal over- 
lap was detected between events. The four com- 
ponents occurred within a mean time of 0.443 
s, and never was a shark observed to partially 
complete a bite once the snout lift and lower-jaw 
depression actions were initiated. The snout drop, 
however, was not always an integral part of a 


feeding action, except at the termination of a 
feeding bout (as discussed above), and may be 
subject to sensory feedback or motivational 
changes. 

Our observations on the mechanics of jaw pro- 
trusion in the white shark are similar to those of 


S Lift 


LJ Depress 


LJ Elev 
o 
Q | 
5 ° 
Ss x S Drop 
fe) 
=e 
O 2 4 6 8 1.0 
Time (s) 
Ficure7. Range of durations for feeding events. Data from 


same shark as in Fig. 6. 


228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


Alexander (1967) on Squalus. He too found a 
head lift component to precede jaw eversion. This 
action is initiated by contraction of the muscles 
at the posterior region of the head and may lead 
to a pronounced snout lifting prior to feeding in 
other sharks (Backus et al. 1956; Moss 1972). 
Actual jaw protrusion in the white shark begins 
after maximum gape is achieved and the mouth 
begins to close (Figs. 4, 5). In fact, full protrusion 
of the palatoquadrate cartilage is not achieved 
until about midway through the jaw closing ac- 
tion. Once the mouth begins to close, palato- 
quadrate extension is very fast (x = 0.083 s,n = 
11) and represents the actual downward move- 
ment of the fully exposed teeth during a bite. 
The mechanics and function of the protrusible 
jaw in large sharks has played a major role in the 
evolution of their feeding habits (see Moss 1977 
for review). Special hydrodynamic problems ex- 
ist for non-demersal sharks because of their lack 
of a gas-filled swim bladder (but see Bone and 
Roberts 1969) and maneuverable paired fins 
(Alexander 1967). The general streamlined body 
form is considered an evolutionary response to 
this problem (Alexander 1967; Budker 1971; 
Thomson and Simanek 1977). The development 
of a protractile jaw has allowed large lamnid and 
carcharhinid sharks to retain a hydrodynami- 
cally efficient fusiform body and the capacity to 
take clean bites with a subterminal mouth. The 


rounded pattern of bites taken from prey too - 


large to swallow whole comes primarily from the 
upward and forward rotation of the lower jaw 
that secures the mouth to the prey, and the down- 
ward and forward cutting rotation of the upper 
jaw. The detached hyostylic association of the 
upper jaw and chondrocranium also permits the 
upper jaw to close downward much faster than 
it could if it had to pull the head with it as it 
closed. This rapid downward movement of the 
massive unattached upper jaw produces a strong 
resultant force that facilitates the cutting action 
of the serrated teeth. 

PREDATORY BEHAvIOR.—Sharks used various 
capture modes to take baits depending on the 
bait’s size and its position relative to the surface. 
In situations where large pieces of meat were 
suspended or floating at the surface, two com- 
mon approaches were observed. 

1) Underwater Approach: In this behavior, 
sharks swam parallel to and approximately 0.5 
m below the surface until less than 1m away from 


the bait. In situations where bait was freely float- 
ing on the surface, sharks swam at normal swim- 
ming speeds as the prey was engulfed. In cases 
where the bait was suspended by pole and line, 
sharks would typically bite the bait and attempt 
to pull it under by depressing their heads. Sharks 
that did not sever the line would often hang ver- 
tically and repeatedly bite at the bait, displaying 
all components of the bite behavior. Sharks were 
persistent in attempts to take the bait after an 
attack was made. 

2) Surface Charge: The second, less common 
feeding behavior on bait at or near the surface 
was a rapid accelerated rush. Here a shark would 
approach and engulf the bait as it swam by at a 
relatively fast rate. This behavior was most com- 
monly observed on newly arrived sharks in an 
excited state. Unlike the underwater approach, 
a charge was made at the surface. This behavior 
created considerable disturbance well before the 
bait was taken. Charging behavior of a similar 
nature was described for the blue shark (Prionace 
glauca) feeding on dense surface schools of squid 
(inicase!'979): 

White sharks were also observed feeding un- 
derwater, and exhibited different predatory be- 
haviors than when taking prey from the surface. 
Two additional modes are presented. 

3) Normal Underwater Pass: This feeding be- 
havior was observed when a shark approached 
a relatively small submerged bait. Sharks ap- 
proached with the mouth opened wider than dur- 
ing normal swimming and raised the snout slightly 
when approximately | m from the bait. When 
the bait contacted the underside of the snout, the 
lower jaw depressed slightly and the bait was 
taken. In this behavior the snout lift and lower 
jaw movements were present, but not as pro- 
nounced as in surface feeding modes, and there 
was no protrusion of the upper jaw. The under- 
water pass appeared to be first mediated by vi- 
sion prior to contact, and second by tactile sen- 
sory input when the snout touched the prey just 
prior to initiating a feeding action. Additional 
sensory systems (e.g., gustatory or electrorecep- 
tive) may also be involved in normal feeding 
situations at close ranges. 

4) Side-Roll: A similar approach to a normal 
underwater pass occurred where a shark rolled 
onto its side just prior to engulfing the submerged 
prey. Here the shark maintained its horizontal 
approach until approximately 1-2 m away from 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 


the bait. It then rolled aproximately 60° from 
normal, took the prey, and returned to an upright 
swimming attitude. 

These latter two approaches involved no de- 
tectable change in swimming speed and em- 
ployed the characteristic movement sequence of 
head and mouthparts. 

Numerous observations on the variability in 
feeding patterns of sharks in relation to prey type 
and feeding conditions exist. Budker (1971) re- 
ported that in normal feeding situations sharks 
exhibit no body contortions when they consume 
small fish prey which are swimming at their own 
level or slightly below. This appears to be the 
case for white sharks taking small pieces of bait 
in normal underwater passes. He further states 
that there are only two situations that might re- 
quire a different type of approach to a bait. These 
are either when a bait is attached to a hook and 
the shark must turn on its side to avoid the line 
with its snout, or when bait is floating and the 
shark must thrust its snout out of the water to 
get its mouth around the bait. We agree with his 
conclusion in regards to floating baits, with the 
addition that this includes natural feeding situ- 
ations as well. This behavior has been observed 
for white sharks feeding on pinnipeds at the sur- 
face (Ainley et al. 1981; personal observations), 
and for tiger sharks (Galeocerdo cuvier) feeding 
on both surface baits and normal prey (see Moss 
1972; Gilbert 1963 for pictures). In addition, it 
is clear that side-roll behaviors may also occur 
in natural feeding situations, and are not nec- 
essarily responses to obstructions during feeding. 
Tricas (1979) found that blue sharks approached 
small, moving anchovy baits from behind and 
took them in a normal swimming posture, while 
larger whole mackerel baits were taken from be- 
hind by sharks that partially rolled onto their 
sides. This variation was attributed to the size 
of the prey and its position relative to the mouth 
just prior to capture. 

Observations of white shark feeding behavior 
are limited to artificial situations in which sharks 
were attracted to feed on tethered fish or horse- 
meat (this study) and the few instances when 
white sharks were observed feeding on dead ce- 
taceans (e.g., Pratt et al. 1982). Some information 
has been gained from interviews with shark at- 
tack victims, although most of these did not see 
the shark before or during the attack (cf. Miller 
and Collier 1981) and may have made biased 


id 
to 
\o 


observations. White sharks have been kept alive 
for short periods in large aquariums, but none 
have attempted to feed (McCosker 1981). 

On the basis of information discerned from 
white shark attacks on pinnipeds and humans, 
and our observations of their feeding on bait, we 
can best summarize the predatory attack strategy 
as follows. An adult white shark is not agile 
enough to capture a fleeing, darting pinniped;: 
hence, it generally attacks its prey by surprise. 
Bite scars on northern elephant seals (Mirounga 
angustirostris), California sea lions (Zalophus 
californianus), Australian fur seals: (Arctocepha- 
lus doriferus) (Fig. 8), and sea otters (Enhydra 
lutris) (Fig. 9, also see Ames and Morejohn 1980) 
are usually located on the ventral region of the 
body (e.g., haunches and flippers). This indicates 
that attacks were made from behind and beneath 
the prey. A typical attack scenario might entail 
a shark swimming a few meters beneath the sur- 
face, searching for the silhouette of a pinniped 
or sea Otter at the surface. Once a prey is sighted, 
the shark ascends and at close range (approxi- 
mately | m) begins a feeding action as described 
above. After attacking large prey such as an el- 
ephant seal, the shark probably routinely retreats 
a short distance from the injured (and at least 
partially immobilized) prey and swims cautious- 
ly within the area, apparently waiting for the pin- 
niped to bleed to death or lapse into shock. Dur- 
ing the attack, white sharks often roll.their eyes 
posteriorly, which reduces the risk of injury to 
the eye by the teeth or nails of a struggling prey. 
The retreat behavior is also adaptive since it 
eliminates the chance of injury via contact after 
the initial attack is made. This “‘bite and spit” 
strategy might explain why seals that have es- 
caped after attack usually have a single massive 
bite. This might also provide insight into the 
paradox of why humans are rarely consumed 
after being attacked. Since humans rarely dive 
or swim alone, the victim is usually quickly res- 
cued or removed from the attack area by others, 
precluding a second attack. The white shark- 
related human fatalities that have occurred in 
California and Oregon waters since 1926 (5 of 
40 attack victims) have all resulted from trau- 
matic blood loss and did not involve massive 
consumption by the shark. This alternative the- 
ory might be more reasonable than the sugges- 
tion that humans are “distasteful” to white sharks, 


230 


FiGur™e 8. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


TN ee ay 


Wounds inflicted upon pinnipeds by white sharks. It is possible to hypothesize the posture of prey and the attack 


behavior of the shark from bite scars. Elephant seals (Mirounga angustirostris): (A) Subadult survivor at Ano Nuevo Island, 
California. Note lower jaw puncture wounds and tearing caused by upper jaw teeth. Photo by R. Bandar. (B) Adult female 
survivor at Southeast Farallon Island, California. Again note lower and upper teeth wounds. Photo by S. H. Morrell. California 
sea lions (Zalophus californianus): (C) Adult survivor at Ano Nuevo Island. Photo by R. Bandar. (D) Subadult male carcass 
(left) and Richard Ellis (right) at Afio Nuevo Island. Photo by Pam Wing. (E) Subadult male carcass along the central California 
coast. Photo by R. Bandar. Southern fur seal (Arctocephalus doriferus): (F) Large adult male survivor at South Neptune Island, 


South Australia. Photo by J. McCosker. 


particularly when one considers the euryphagic 
diet of the fish. 

PREDATOR-PREY RELATIONSHIPS.— The stom- 
ach contents of nine white sharks (193-511 cm 
total length) captured in northern and central 
California waters are presented in Fig. 10. Seven- 


ty-eight percent of the sharks had recognizable 
food items in their stomachs. The most frequent 
prey was the California bat ray (Myliobatis cali- 
fornica), found in four stomachs; other fish prey 
were less frequent in the diet. Fifty-six percent 
of the sharks examined contained elasmo- 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 231 


eae 


eo 


Ficure 9. Sea otters (Enhydra lutris) from the central California coast. Above, adult in normal feeding or basking posture 
along the edge of a kelp bed in Monterey Bay. Photo by J. McCosker. Below, lacerated carcass from which several white shark 
tooth fragments were removed, suggesting that the animal was bitten at the surface while in a belly up, prone position. Pismo 


Beach. Photo by J. Ames. 


branchs, and 44 percent contained teleost prey 
species. No evidence of predation on marine 
mammals was found in the nine sharks. 
Although the white sharks took prey that nor- 
mally occur in both pelagic and inshore habitats, 
the two most frequent prey are generally asso- 
ciated with demersal inshore communities. The 
California bat ray (M. californica) is common in 
bays and inshore sandy habitats 2-50 m deep, 


where it feeds on benthic sand-dwelling inver- 
tebrates. The spiny dogfish (Squalus acanthias) 
is also demersal, being found in both shallows 
and deeper offshore waters. Other prey species 
that live on the bottom in inshore areas are the 
lingcod (Ophiodon elongatus) and the cabezon 
(Scorpaenichthys marmoratus). These latter two 
species are relatively sedentary, have small home 
ranges, and show cryptic coloration. Limbaugh 


232 
Myliobatis californica (E) KX 11 MW 
Squalus acanthias (E) GG GGJ»™w 
Cetorhinus maximus (E) Wg 
Cynoscion nobilis Gg 
Galeorhinus zyopterus (E) Gg 
Ophiodon elongatus Wg 
Scorpaenichthys 
marmoratus 
Sardinops sagax S39 
Empty MAY 
Ir al) mu T T al 
10) 10 20 30 40 50 


% Occurrence 


Ficure 10. Stomach contents of nine sharks captured in 
northern California waters on record at California Academy 
of Sciences. % occurrence = percentage of the nine shark stom- 
achs that contained that prey item. E = elasmobranch, all oth- 
ers are teleosts. 


(1963) reported cabezon from the stomachs of 
three immature sharks captured at La Jolla, Cal- 
ifornia, and described a number of incidents that 
indicate S. marmoratus is an important prey for 
young white sharks. Earlier researchers ques- 
tioned how sharks could detect and capture such 
inconspicuous and apparently inaccessible prey; 
our studies suggest that weak electric fields might 
be involved in prey detection (see Sensory Bi- 
ology section below). 

The white sea bass (Atractoscion (=Cynoscion) 
nobilis) also occurs in shallow rocky inshore hab- 
itats, and is often found among canopies of giant 
kelp (Macrocystis pyrifera). Unlike the majority 
of the other inshore prey species, however, it 
occasionally swims in the water column as well 
as on the bottom. 

White sharks have been reported to feed on 
the carcasses of captured basking sharks (Fast 
1955), although we know of no published ac- 
counts of predation under natural circumstances. 
However, potential vulnerability of basking 
sharks to large predators was suggested by Lim- 
baugh (1963) in an account of a dead basking 
shark with a large wound probably inflicted by 
killer whales. Basking sharks, which reach lengths 
of more than |1 m, are found seasonally in off- 
shore waters of central and northern California. 
From aerial surveys made over a 2.5-yr period 
near Monterey, California, Squire (1967) found 
that basking sharks were most common from 
September through May, when water tempera- 
tures were generally below 14°C. White shark 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


Fish 


Pinnipeds 


Cetaceans 


£4 Elasmobranchs 


Other prey Actinopterygians 
apa eee ane T | ==) 
ie} 10 20 30 40 50 60 


% Occurrence 


Ficure 11. Stomach contents of 33 white sharks. Data 
combined from this study and other published records. % oc- 
currence = percentage of the 33 sharks that contained the prey 
category. Fish prey subdivided into elasmobranchs and rayed- 
fin fishes (teleosts and sturgeons). Other prey include birds, 
crustaceans, and sea turtles. 


sightings, however, were most common in the 
warmer-water months of May through August, 
when water temperatures neared or exceeded 
14°C. The cause of the seasonal disappearance 
of basking sharks from the coastal waters of Cal- 
ifornia remains unknown. Other prey that in- 
habit pelagic waters include the soupfin shark 
(Galeorhinus zyopterus), the Pacific sardine (Sar- 
dinops sagax), and occasionally bat rays (Myl- 
iobatis californica) (Roedel and Ripley 1950; 
Feder et al. 1974). 

Combined data on the food habits of 33 white 
sharks from this study and other published rec- 
ords are shown in Fig. 11. Here again, fish were 
the most frequent prey items, occurring in over 
half of white sharks in the analysis. Elasmo- 
branchs and rayed-fin fishes (teleosts and stur- 
geons) comprised equal proportions (each oc- 
curred in 30 percent of sharks analyzed) of the 
piscine prey. Pinnipeds were also a major com- 
ponent in the diet of sharks, while cetaceans and 
other prey groups were less common. Bass et al. 
(1975) provided the only other gut content data 
from white sharks useful for comparison. They 
too found both elasmobranchs (40 percent of 
sharks examined) and teleost fishes (25 percent) 
as the most common prey items, although little 
information was given on specific identification. 

Figure 12 shows the distribution of fish and 
mammal prey in relation to shark size. Fish prey 
predominated in the diet of sharks approxi- 
mately 3 m or less (TL), while pinnipeds and 
cetaceans predominated in those of larger sharks. 
This shift in diet may occur for a number of 
reasons. For example, larger sharks are less agile 
and would be less successful in chasing and cap- 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 233 


© Pinnipeds & Cetaceans 


@ Fish 


ooo ° Oo o0c0000 Mo oO 


100 200 300 400 500 600 
Total Length (cm) 


Ficure 12. The relationship between white shark length 
and prey type. Data taken from stomach contents of the 33 
specimens in Fig. 11. 


turing smaller fish prey that dart about when 
pursued. Larger sharks may thus switch to dif- 
ferent prey types and associated new hunting 
modes. In addition, the energetic requirements 
of large, warm-bodied sharks may be better met 
by prey high in fat content (i.e., high-energy— 
density prey). Carey et al. (1982) estimated the 
metabolic rate for a 4.6-m white shark, and con- 
cluded that the animal could survive for ap- 
proximately 1.5 months on 30 kg of whale blub- 
ber (a conservative meal size). They suggest this 
to be adaptive during long intervals between en- 
counters with prey. Although little is known of 
the movements of white sharks, they do show 
seasonal peaks in abundance in California waters 
(Squire 1967; Ainley et al. 1981), which might 
indicate some sort of regional or long-distance 
movement. 

Morphological differences between large and 
small sharks may also account for different pred- 
atory tactics. Fig. 13 shows the relationship be- 
tween tooth shape and shark total length. Smaller 
sharks have a relatively long, narrow tooth shape 
that is better adapted for grasping prey like small 
fishes. This feature is so well developed in small 
white sharks that they are often incorrectly iden- 
tified as mako sharks (Jsurus spp.) (Smith 1951, 
1957). At about 3 m TL, the teeth broaden at 
the base and take on the diagnostic triangular 
serrated form. Unlike the long narrow teeth, this 
shape is well-suited for gouging and cutting pieces 
from prey too large to swallow whole. Le Boeuf 
et al. (1982) found evidence that marine mam- 
mals were the only prey of large white sharks 
they examined from California. Of seven spec- 
imens examined, all but one were approximately 
4 m or longer and had evidence of marine mam- 
mals in their stomachs. The only exception was 
the smallest shark (2.4 m TL), which had only a 
10-cm patch of pinniped pelage in its stomach. 


e 
Hell e 
e e e 
10 4 e 
e? 
we 
© ee 
2 94 e 
e 
SS 
— 
7 e 
S e 
see 85 
ro) 
° 
i 
774 
) 
oe = =r aa | 
1 2 3 4 5 6 


Total Length (m) 


FiGure 13. Therelationship between shark total length and 
tooth shape. Tooth shape expressed as the ratio of width of 
enamel base to medial height of enamel for the first tooth, right 
side, upper jaw of 16 sharks. Low ratio indicates a long narrow 
tooth shape; higher ratio indicates relatively broad triangular 
shape. 


Perhaps this shark’s teeth were too narrow to 
excise a portion of flesh. 

In California waters, elephant seal populations 
at offshore rookeries peak in both the spring and 
winter months (Le Boeuf et al. 1974), but almost 
no predation occurs during the spring peak. Hy- 
pothetical explanations advanced to explain this 
seasonal discrepancy in predation include either: 
1) sharks fasting while breeding; 2) water too cold 
for sharks to feed; or 3) emigrations of sharks 
from the area. Even though sharks occur in Cal- 
ifornia waters during the spring (Miller and Col- 
lier 1981), the decrease in shark attacks is prob- 
ably due to emigrations of large sharks from 
coastal areas (see Squire 1967). Adult male seals 
are more susceptible to shark predation because 
they spend more time in the water near the rook- 
ery during the breeding season than do females 
(Le Boeuf et al. 1982). It is possible that the loss 
of peripheral males to sharks may not adversely 
affect the population because of the polygynous 
mating system of the elephant seal, where rela- 
tively few dominant males do the majority of the 
breeding. 

Although it is clear that white sharks do nor- 


234 


Ficure 14. Underwater photo of a male white shark (ap- 
proximately 3.5 m TL) in a “tail stand” posture with snout 
directly over zinc anode on rudder of study vessel. Photo by 
T. Tricas. 


mally prey upon elephant seals, the significance 
of the interaction is not evident. Ainley et al. 
(1981) reported an increase in the number of 
attacks on elephant seals at the Farallon Islands 
between 1970 and 1979, but their data indicate 
a density-dependent relationship between num- 
ber of attacks and numbers of elephant seals. 
More data are needed on the mortality rates of 
attacked seals and on numbers in the shark pop- 
ulation before any effects of shark predation on 
elephant seal populations can be quantitatively 
assessed. 

SENsoRY BrioLoGy.—Our cursory field exper- 
iments and observations qualitatively indicate 
white sharks are sensitive to electric fields. In the 
pulsed electric field tests, sharks took the exper- 
imental bait 8 times (73 percent) and the control 
3 times (27 percent). In the constant current (DC) 
tests the experimental was taken 4 times (44 per- 
cent) and the control 5 times (56 percent). Al- 
though our sample size was too small to show 
any statistically significant preference for baits 
with either type of electric field, sharks did take 
baits with the pulsed electric field almost three 
times more often than the control. The sharks 
also appeared to be more responsive to pulsed 
fields than to continuous fields. Kalmijn (1971, 
1974) reported that sharks were most responsive 
to weak electrical fields at frequencies from 0 
(DC) to 8 Hz. 

We also observed the behavior of sharks to 
metallic objects attached to the bottom of the 
boat. On three occasions one of us (TCT) watched 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


from underwater a 3.5-m shark approach zinc 
plates attached to the boat’s rudder and assume 
a near vertical “‘tail stand” posture (Fig. 14). The 
shark remained upright for approximately 10— 
20 s as it waved its snout approximately 5-10 
cm above the zinc. Sharks were also observed 
several times to swim back and forth with their 
snouts very near a 10-m-long copper grounding 
strip on the bottom of the boat’s hull. 

We interpret these observations as a response 
by sharks to the galvanic currents produced by 
the electrochemical interaction between the me- 
tallic plates and seawater. White sharks have a 
well-developed system of ampullae of Lorenzini 
(Fig. 15), and although the role of electric detec- 
tion of prey by sharks is well demonstrated (see 
Kalmijn 1978, 1982), the degree of importance 
for such a sensory modality in white sharks re- 
mains unknown. It is noteworthy, however, that 
electric fields produced by large mammals (e.g., 
humans and presumably pinnipeds) in seawater 
are well within the sensory range of elasmo- 
branchs (Kalmijn 1971). Perhaps young white 
sharks are able to detect electrically sedentary 
camouflaged fish prey like the cabezon (Scor- 
paenichthys marmoratus). It also seems reason- 
able that the ampullae would be particularly use- 
ful to detect: 1) the location of a marine mammal 
at the moment Just prior to attack; 2) any change 
in position or escape attempts by the prey; and 
3) any change in the prey’s condition, such as 
bleeding, which might alter the strength or sig- 
nature of the electric field. 

TELEMETRY.— Two sharks were tagged with 
temperature-sensing transmitters during this 
study. The first shark (a 4.5-m male) carried a 
unit that monitored ambient water temperature 
only. After tagging, the shark remained around 
the boat even after all baits were removed from 
the water. The boat was then moved away from 
the area and the shark began to move westward; 
parallel to the north shore of Dangerous Reef. 
Once past the island the shark moved offshore 
in a northwesterly direction. Contact was lost 
with the animal approximately 4 h after initial 
tagging, due to its rapid speed and bad seas that 
created poor tracking conditions. During this time 
the shark swam in waters 20—21°C as indicated 
by the temperature sensor on the transmitter. 

The second shark was tagged on 22 January 
1980. The body temperature probe was placed 
31 cm deep into the lateral musculature, ap- 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 235 


FiGure 15. 
C. J. Slager. 


proximately 25 cm below the first dorsal fin. This 
shark was monitored continuously near the boat 
for approximately 2 hr, until it swam out of range. 
It returned to the anchored boat near midnight, 
and then again departed. Results of the thermal 
data are presented in Table | and Fig. 16. The 
shark swam in water ranging from 20.9° to 21.5°C. 
Mean difference between ambient and body tem- 
perature was 3.7°C, and ranged from 3.2° to 4.3°C. 


TaBLe 1. EpaxiaL MuscLe TEMPERATURES OF A 3.5 M (TL) 
MALE WHitE SHARK MONITORED AT DANGEROUS REEF, SOUTH 
AUSTRALIA ON 22 JANUARY 1980. Mean (AT) = 3.7°C. SD = 
OTe 


Temp (°C) 
Measure- Difference 
ment Water Body (AT) 
1 21.2 DIED 4.0 
2 DNS) 24.7 32 
3 Dj} 24.7 3.5 
4 Die? DS 4.0 
5 Die. WS\f) 4.0 
6 QED. Psyop 4.0 
7 Dey 24.7 3),5) 
8 20.9 Deyo 4.3 
9 20.9 24.7 3.8 
10 20.9 24.2 $53) 
11 20.9 24.2 3)58) 
12 20.9 24.2 33) 


Distribution of the ampullae of Lorenzini on the head of a young female white shark (CAS 37917). Figure by 


Largest and smallest differences were recorded 
when the shark entered water of a different tem- 
perature, before internal temperatures could con- 
form. This time lag to thermal equilibrium and 
variation in muscle temperature indicate that the 
shark did not thermoregulate. Carey et al. (1982) 
found that a 4.6-m white shark had a body tem- 
perature 3—-5°C higher than the surrounding water. 
Their shark swam over deeper waters, and for 
the most part remained in the thermocline. Tem- 
peratures were lower in their study, ranging ap- 
proximately from 5° to 19°C ambient, and 18° 


WHITE SHARK MUSCLE TEMPERATURE 


NIN 


? 


TEMP. White Shark Epaxial Musculature ‘ 


(°C) 23 


Seawater 


Figure 16. Temperature difference between ambient sea- 
water and epaxial musculature of a 3.5 m TL white shark, 
monitored on 22 January 1980 at Dangerous Reef, South Aus- 
tralia. Question marks (?) indicate time interval when shark 
swam away from anchored study vessel and out of telemetry 
range. Figure by K. O’Farrell. 


236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14 


FiGure 17. 


to 23°C muscle temperature. Our study took place 
in relatively shallow waters around Dangerous 
Reef (<30 m), and we found no sign of a marked 
thermocline. The water and shark-muscle tem- 
peratures we recorded were generally higher 
(20.9°-21.2° and 24.2°-25.2°C, respectively), but 
they are consistent with the values for body tem- 
perature elevation over ambient recorded by Ca- 
rey et al. 

One of the primary advantages of being warm- 
bodied is thought to be related to the changes in 
muscle physiology as temperature increases. It 1s 
known that a 10°C increase in temperature may 
result in a three-fold increase in the contraction- 
relaxation rate of frog muscle (Hartree and Hill 
1921). For fish, this may be translated to an in- 
crease in potential tail-beat frequency and a re- 
lated increase in sustained swimming speed. 
Higher speeds may be selectively advantageous 
when chasing prey or fleeing from predators. In 
addition, conservation of heat theoretically al- 
lows for more total energy conversion to work, 
thus enabling an animal to swim longer distances 
on a given meal. Being warm-bodied might also 


(Right) Silhouette of a surfer on a contemporary surfboard. (Left) Silhouette of an adult female (TL = 1.7 m) 
harbor seal (Phoca vitulina). Photo by Al Giddings. 


allow for temporary excursions into colder or 
deeper waters. This thermal inertia (see Neill et 
al. 1976) would not only expand the range of 
environments which the animal could exploit, 
but would also permit increased swimming ef- 
ficiency for predation at otherwise limiting en- 
vironmental temperatures. 

ON WHITE SHARKS AND SURFBOARDS. — In con- 
clusion, we comment on the increasing attacks 
by white sharks upon humans who surf in the 
north Pacific. Since 1972, there have been 11 
recorded white shark attacks upon surfers in Cal- 
ifornia and Oregon (Miller and Collier 1981) and 
one such attack in Hawaii in 1959 (Balazs and 
Kam 1981). The similarity in appearance of the 
silhouette of a prone human on a surfboard or 
“belly board” to a large surface-basking pinniped 
is clear (Fig. 17), and observations of attacks by 
sharks upon surfers fit well with our assessment 
of the feeding strategy of white sharks. Attacks 
have occurred in the vicinity of pinniped rook- 
eries, such as the much-publicized death of Lewis 
Boren on 19 December 1981 at Spanish Bay, 
Monterey, Calfornia. 


TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK 237 


Since the early 1970s, the trend in surfboard 
design has been toward an increase in flotation, 
reduction in board length, multiple posterior- 
fixed rudders (‘‘skegs”), and bifurcated or “V” 
tails. All of these modifications have enhanced 
the similarity between the silhouette of a surfer 
and that of a pinniped, and we suggest that this 
may increase the probability of attack of surfers 
encountered by white sharks. We feel it advisable 
that those who surf be aware of and consider the 
potential risks of surfing in coastal waters known 
to be frequented by white sharks. 


ACKNOWLEDGMENTS 


We are particularly grateful to Al Giddings, 
President of Ocean Images, Ltd., for financial 
assistance, for providing access to his film library 
for cinematographic analyses, and for sharing his 
observations of shark behavior with us. We also 
thank Terry Thompson, Ocean Images, Ltd., for 
his assistance. Additional shark data were pro- 
vided by W. I. Follett (CAS), L. J. V. Compagno 
(CAS), J. Randall (Bernice P. Bishop Museum), 
and the staff of the Department of Ichthyology 
of the CAS. A. Dizon (NMFS, Honolulu) and I. 
Cooke (Bekesy Laboratory, U. of Hawaii) pro- 
vided facilities during construction of our trans- 
mitters. We thank H. Tricas, C. J. Slager, K. 
O’Farrell, S. Middleton, and S. Nakamura for 
assistance with our figures; J. Ames, R. Bandar, 
R. Dunne, A. Giddings, S. Morrell, P. Romano, 
and P. Wing for allowing us to use their pictures; 
and Bob Britcher and Chico Chingwidden, the 
Master and the mate of the Nenad. We give spe- 
cial thanks to Rodney Fox, for his guidance in 
the field in South Australia and for helpful dis- 
cussions concerning shark behavior, and to 
Leighton Taylor, Jr. (Waikiki Aquarium), and 
Phil Motta (Univ. of Montana) for their critical 
reading of this manuscript. 

Senior authorship of this paper was deter- 
mined by the outcome of a pinball match played 
at Port Lincoln, South Australia, in January 1980. 


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RANDALL, J. E. 1973. Size of the great white shark (Car- 
charodon). Science 181:169-170. 

Ricker, W. E. 1973. Linear regression in fishery research. J. 
Fish. Res. Board Can. 30:409-434. 

Roepe, P. M., AND W. E. Riptey. 1950. California sharks 
and rays. Calif. Div. Fish Game, Fish. Bull. 75. 88 pp. 

Royce, W.F. 1963. First record of white shark (Carcharodon 
carcharias) from southeastern Alaska. Copeia 1963:179. 

ScCATTERGOOD, L. W. 1962. White sharks, Carcharodon car- 
charias, in Maine, 1959-1960. Copeia 1962:446-447. 

SCHROEDER, W.C. 1938. Records of Carcharodon carcharias 
(Linnaeus) and Pseudopriacanthus altus (Gill) from the Gulf 
of Maine, summer of 1937. Copeia 1938:46. 

Situ, J. L. B. 1951. A juvenile of the man-eater, Carchar- 
odon carcharias Linn. Ann. Mag. Nat. Hist. 4(12):729-736. 

. 1957. Sharks of the genus /surus Rafinesque, 1810. 
Ichthy. Bull. 6:91-96. 

SPRINGER, S. 1961. Dynamics of the feeding mechanism of 
large galeoid sharks. Am. Zool. 1:183-185. 

Squire, J. L., Jk. 1967. Observations of basking sharks and 
great white sharks in Monterey Bay, 1948-50. Copeia 1967: 
247-250. 

THomson, K. S., AND D. E. SIMANEK. 1977. Body form and 
locomotion in sharks. Amer. Zool. 17:343-354. 

Tricas, T.C. 1979. Relationships of the blue shark, Prionace 
glauca, and its prey species near Santa Catalina Island, Cal- 
ifornia. Fish. Bull. 77:175-182. 


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Vol. 43, No. 15, pp. 239-248, 5 figs., 1 table 


Lights 
| gere sae PROCEEDINGS 


| OF THE 
CALIFORNIA ACADEMY OF SCIENCES 


01 SPOUT WNT A ES 


WV Cc ai IGE 
peer 


September 19, 1984 


DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV. 
(STYELIDAE, ASCIDIACEA) FROM THE PACIFIC COAST OF 
THE UNITED STATES, AND ITS IMPACT ON SOME GONADAL 
CRITERIA OF ITS GENUS AND SUBGENUS 


By 
Andrew Todd Newberry 
Cowell College, University of California, Santa Cruz, California 95064 


Asstract: Dendrodoa (Styelopsis) abbotti, a newly described styelid ascidian from the central and northern 
California coast, the San Juan Islands of Washington, and southwestern Vancouver Island, resembles D. 
carnea but differs in branchial and gonadal traits. Inclusion of D. abbotti in the genus Dendrodoa requires 
modification of the gonadal criteria of the genus to accommodate styelan gonadal resemblances (non-encap- 
sulation of the testis-lobes with the ovary) and styelan or cnemidocarpan spermiducal resemblances (gonad’s 


single vas deferens and spermipore). 


INTRODUCTION 


The tunicate named and described in this pa- 
per, Dendrodoa (Styelopsis) abbotti, is a styelid 
ascidian that has long been collected along the 
central and northern California coast. Donald P. 
Abbott, who first found this ascidian in 1948 
near Point Arena (Mendocino County), included 
it as ““Alloeocarpa sp.” in the urochordate key of 
the second edition of Light’s Manual (Light et 
al. 1954) but, for want of more certain identifi- 
cation, omitted it from that handbook’s third 
edition (Smith and Carlton 1975). 

This ascidian’s aggregative habit does create 
an appearance of budding (Fig. 1A), but adjacent 
zooids’ tests are unfused and easily separated 
from one another; no evidence of budding has 
been found in several hundred zooids from sev- 
eral sites and all seasons. Apparently, then, this 
is a solitary ascidian and cannot be placed in the 
genus Alloeocarpa. It shows Dendrodoa’s restric- 


tion of the single, elongate ovary to the zooid’s 
right side. The ovary’s unbranched shape and 
the pharynx’s simplicity place the species in the 
subgenus Styelopsis of Dendrodoa. The specific 
name, abbotti, honors Professor Donald P. Ab- 
bott, of the Hopkins Marine Station of Stanford 
University, who has shared with his students and 
colleagues a singular keenness of intellect and 
generosity of spirit, and it expresses the esteem 
and affection of his fellow ascidiologists. 


MATERIALS AND METHODS 


This report is based principally on specimens 
collected intertidally at Pigeon Point, San Mateo 
County, California (lat. 37°11'0”N, long. 
122°23'10’”W), at intervals of roughly six weeks 
throughout 1977. I have also drawn on material 
taken over the past two decades from there; from 
Point Pinos, Monterey County, California (lat. 


[239] 


240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15 


Figure 1. A) Living zooids of Dendrodoa abbotti, including one (upper left-center) that has been wounded or severely 
disturbed and is extremely contracted while the others remain relaxed. Zooids are about | cm long. B) Closer view of two slightly 
disturbed zooids, showing siphons in the process of bilabial closure. Brood pouch is detectable in the left zooid by inflated 
aspect of the posterior region of the zooid (to left in photo). Zooids are about | cm long. 


NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV. 24] 


36°38'0’N, long. 121°56’0”W); and subtidally 
from Peavine Pass (lat. 48°35'4”N, long. 
122°45'48”W) in the San Juan Islands, Wash- 
ington. I have used, as well, D. P. Abbott’s un- 
published notes and drawings of specimens from 
northern and central California. 

In all, I have examined approximately 30 spec- 
imens thoroughly. I have examined several doz- 
en more in a cursory way to verify the criteria 
that characterize the species. All specimens were 
relaxed with MgCl, or MgSO,, and menthol, then 
fixed in seawater Bouin’s fluid or 10% formalin, 
and all were preserved in 70% ethanol. The 
Bouin’s-fixed material provided excellent serial 
sections but brittle dissections. Formalin always 

‘fixed adequately for dissections but rarely well 
enough for close scrutiny by serial section (which 
was required, for example, to trace the very fine 
spermiducts). Specimens were dissected in 70% 
ethanol. Dissected specimens usually were 
stained, once opened, with Grenacher’s borax 
carmine; serially sectioned specimens were either 
prestained, often for prior dissection, in Gren- 
acher’s alcoholic borax carmine or stained in sec- 
tion with “standard alum hematoxylin” (Galigh- 
er and Kozloff 1964) and eosin. Prestaining 
proved satisfactory for general examination, but 
staining in section was necessary to reveal finer 
structural details or to take advantage of the bet- 
ter fixation achieved with Bouin’s fluid than with 
formalin. 

CoorDInaTEs.— The endostyle designates the 
anterior-posterior axis and the ventral midline. 
Thus, the dorsal midline extends from the oral 
siphon through and beyond the atrial siphon. By 
these coordinates, the ovary lies against the right 
ventral margin of the zooid, and the loop of the 
gut dominates the left posterior region of the 
zooid (Fig. 2). 


DESCRIPTION OF SPECIES 
Dendrodoa (Styelopsis) abbotti, sp. nov. 


TYPE-SPECIMENS. — Holotype at California Academy of Sci- 
ences, San Francisco, Calif. (CAS #034790). Paratypes at Cal- 
ifornia Academy of Sciences, San Francisco, Calif. (CAS 
#034791). 

TypE-LocaALiry.—North side of Pigeon Point, San Mateo 
County, California (lat. 37°11'0”N, long. 122°23'10”W). 

OTHER REcorpDs.—Intertidal records from Point Pinos and 
Hopkins Marine Reserve (Monterey County), Pigeon Point 
and Moss Beach (San Mateo County), Point Arena (Mendocino 
County), California, and near Sooke, Vancouver Island, British 
Columbia; subtidal records from Peavine Pass (San Juan Coun- 
ty), Washington. 


EXTERNAL APPEARANCE (Fig. 1).—Zooids round 
or oval (lengthened antero-posteriorly) low 
hemispheres; entire sub-endostylar surface ap- 
plied to substrate; attached surface extends be- 
yond ovary on right and gut-loop on left. Spec- 
imens including test reach 8 to 12 mm length, 6 
to 10 mm width, 2 or 3 mm height when relaxed; 
zooids removed from test reach 8 to 10 mm 
length, 6 to 8 mm width, 2 to 3 mm height. Test 
clean, thin, and parchment-like, spreading as a 
thin apron | to 2 mm wide on the substrate 
around the zooid. Ventral test extremely thin. 
Color in life translucent gray tinted with ochre 
or very pale brownish pink, with borders of si- 
phonal apertures sometimes slightly darker. 
Zooids fixed in formalin become plain translu- 
cent white-gray. Alive or fixed, zooid’s branchial 
sac, gut, ovary, and mass of brooded young are 
faintly visible through dorsal and lateral regions 
of test. Oral siphon far anterior; atrial siphon 
placed centrally atop hemispheric zooid; both 
siphons fairly evident in relaxed living animals 
but reduced to obscure slits in contracted ones. 
Relaxed zooids have circular siphonal apertures; 
disturbed zooids close their siphons bilabially 
into transverse slits (Fig. 1B) and flatten them- 
selves against the substrate within a delicately 
crumpled test. Zooids are simple and non-bud- 
ding but often aggregate in pairs or trios (rarely 
groups of more) with young ones often settling 
adjacent to or even on the test “‘apron”’ around 
older zooids (but not on zooidal surfaces them- 
selves). Mature zooids, even when tightly adja- 
cent to one another, attach entirely to the sub- 
strate itself; they do not form clumps of zooids 
growing thickly one upon another. Adjacent 
zooids often are oriented similarly on the sub- 
strate. 

VASCULAR ELEMENTS OF THE TEST.— Test-ves- 
sels not prominent; as revealed by staining, 
branching systems of test-vessels ramify toward 
the margin of the test. Test-vessel ramifications 
connect to zooid by one or more sub-zooidal 
circulatory junctions; tips of all branches of test- 
vessel ramifications end peripherally in slender, 
bulbous vascular ampullae. 

MANTLE.—Thin, lightly muscled mantle ex- 
cept for extensive arrays of fibers radiating from 
each siphon and controlling its bilabial closure; 
fairly conspicuous concentric musculature sur- 
rounding oral siphon, less developed concentric 
musculature around atrial siphon. About a dozen 
endocarps project from the mantle into the atrium 


242 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15 


Ficure 2. Dorsal view of zooid, with pharynx removed to show disposition of ovary and testis-lobes (including far posterior 
group), vasa efferentia and vas deferens (including tiny spermiporal papilla near atrial siphon), mid-ventral endostyle and heart 
beneath it, and gut-tract (stomach, pyloric duct, caecum, intestine). Position of oral and atrial siphons indicated by ovals. 

Ficure 3. Zooid opened by mid-ventral cut to show ovary, testis-lobes (including posterior lobes), larvae in brood pouch, 
several endocarps (stippled), gut-tract, siphons with neural complex between them. Pharynx removed. 


of most zooids; particularly large endocarps usu- 
ally protrude from the atrial mantle anterior to 
the ovary and in the region of the gut-loop. 

ORAL TENTACLES. — 36 to 40 filiform oral ten- 
tacles of three sizes; largest ones most abundant 
(24-30), others about half their size intercalated 
irregularly, a few to many tiny papillae evident 
upon close examination of the band of oral ten- 
tacles. Just distal to this circle of tentacles is a 
siphonal flange that marks the inward limit of 
the test that lines the oral siphon. 

ATRIAL TENTACLES.—40 to 50 tiny filiform 
atrial tentacles in band analogous to that of the 
circle of oral tentacles. Just distal to this incon- 
spicuous circle is the atrial siphonal flange that 
marks the inward limit of the test lining the atrial 
siphon. 

DorsAL TUBERCLE (Fig. 4).—Simple C-shaped 
slit atop a short, stout projection; concavity of 
the C faces posteriorly (toward the dorsal lami- 
na). The dorsal tubercle is set slightly to the right 
of the dorsal midline. 

NEURAL Comp_Lex.—In dorsal or ventral sil- 
houette, whole complex forms a rectangle elon- 


gated antero-posteriorly and extended somewhat 
at each corner. Like the dorsal tubercle, the neu- 
ral complex is set slightly to the right of the dorsal 
midline. 

BRANCHIAL SAC (PHARYNX) (Fig. 5).— Folds 
absent, perhaps represented by internal longi- 
tudinal branchial vessels. In dissection, 4 inter- 
nal longitudinal vessels are evident on each side 
of the pharynx; in transverse serial sections, a 
fifth internal longitudinal vessel is sometimes 
discernible on each side close to the endostyle, 
and in a few specimens even a sixth vessel on 
each side may run only some length of the sac. 
Usually 9 or 10 stigmata lie between these in- 
ternal longitudinal vessels. Ten to 12 transverse 
vessels separate the rows of longitudinally ori- 
ented stigmata, and there are about 10 parastig- 
matic vessels partly or entirely traversing each 
side of the pharynx. Along the ventral midline 
the branchial sac connects with the body wall by 
widely spaced sub-endostylar vascular trabecu- 
lae, not by a continuous sub-endostylar mem- 
brane. Other vascular trabeculae connect the 
branchial sac abundantly in all directions to the 


NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV. 243 


i 


‘NNT tN 
nt 


Ficure 4. Dorsal tubercle in relation to peripharyngeal groove and dorsal lamina. 


FiGcure 5. 


Right side of pharynx, showing several rows of stigmata and the four internal longitudinal branchial vessels of 


the pharynx’s right side. Dorsal lamina at top, endostylar groove at bottom. Drawing based in part on unpublished notes of 


D. P. Abbott, in part on freshly dissected specimens. 


atrial surface of the mantle and to the atrial ep- 
ithelium around the gut. 

DorsAL LAMINA.—Prominent, continuous, 
smooth-bordered dorsal lamina, without lan- 
guets. 

Gut (Figs. 2, 3).—Esophageal aperture far 
dorso-posterior in pharynx; stout esophagus 
bends sharply ventrally into stomach; stomach 
empties anteriorly into fore-intestine, which 
bends to left and passes posteriad on the lateral 
side of the stomach. Hind-intestine then curves 
sharply dorsad and follows the left mantle to the 
anus, which lies slightly to the left-posterior of 
the atrial siphon. Stomach has 16 to 18 mod- 
erately evident external folds corresponding to 
well-developed internal gastric septa. The gastric 
septa are reduced to low ridges in the left pyloric 
region of the stomach, near the pyloric caecum. 
Pyloric caecum is small, sometimes absent. A 
highly vascularized pyloric duct joins the sinus- 
oidal sheath surrounding the stomach with that 
surrounding the fore-intestine. Intestine com- 
prises a fore-intestine with a large typhlosole-like 
longitudinal plication of its wall and a thick si- 
nusoidal jacket between the gut wall and its sheath 


of atrial epithelium, and a hind-intestine of more 
simply tubular section whose atrial sheath is much 
closer to the gut wall. Anus lies dorso-medial or 
slightly to the left, above the stomach; anus is 
cut square to the axis of the rectum; anal margin 
is scalloped into usually 5 lobes that fit together 
when the anus is tightly closed. 

HEART.—Fairly straight within a somewhat 
curved and inflated pericardium; set at about 45° 
obliquely to the endostyle, oriented right-ante- 
rior to left-posterior, centered roughly beneath 
the endostyle in the posterior half of the zooid 
(site and orientation in Fig. 2). 

Ovary (Figs. 2, 3).—Single, unbranched, sau- 
sage-shaped ovary along the nght ventral margin 
of the zooid, extending almost the entire length 
of the zooid, curving sharply dorsad posteriorly 
and following the right mantle to arch halfway 
over the atrium, recurving dorsally to terminate 
in an oviduct directed posteriorly toward the 
brood pouch and away from the atrial siphon. 
Oviduct lies lateral (away from atrium) to main 
mass of ovary, with its lumen penetrating among 
the ripening gametes; lateral surface (away from 
germinal tissue) of oviduct heavily ciliated, other 


244 


oviducal surfaces apparently not ciliated. Ova- 
ries of all specimens examined by dissection or 
serial section show all stages of ovogenesis pres- 
ent, regardless of season. 

Testis (Figs. 2, 3).—A dozen to more than 20 
separate lobate sacs, not encapsulated with the 
ovary but instead lying in the mantle wall ad- 
jacent to but clearly outside the ovary’s delim- 
iting membrane. Most sacs lie medial to the ovary; 
some lie anterior to the ovary; few are lateral; 
many lobes lie partly “beneath” the ovary, in the 
mantle between the ovary and the ventral surface 
of the zooid. In many specimens, but not all, a 
few testis-lobes lie far posteriorly and on the left 
side of the ventral midline, but their spermiducts 
join the vas deferens of the testis-lobes that lie 
beside the ovary. All sacs join by vasa efferentia 
to a single, long vas deferens that lies between 
the ovary and the atrial epithelium. This duct 
follows the ovary to the region of the atrial si- 
phon, and there leaves the ovarian surface to 
project toward the atrial siphon from the dorsal 
roof of the atrium while the ovary bends pos- 
teriad toward its ovipore. The ciliated vasa ef- 
ferentia are extremely thin, visible only in serial 
section; the vas deferens, also scarcely visible 
except in serial section, is a compressed, ciliated 
channel terminating in a tiny, spermipore-bear- 
ing papilla pointing toward the atrial aperture. 
In all specimens examined from all seasons for 
gametic condition, many testis-lobes have tailed 
sperm, but the spermiducts contain only scat- 
tered sperm. 

BROOD CHAMBER AND BROODED YOUNG (Figs. 
1, 3).—The posterior region of the atrium serves 
as a brood chamber, occluded anteriorly by the 
branchial sac, on the left by the gut-loop, on the 
right by the ascending limb of the ovary. All 
specimens examined were brooding young in all 
stages of development from (relatively rarely) 
fertilized eggs and cleavage stages to (usually) 
tadpoles that were still curled (although many of 
these straightened upon removal from the brood 
chamber during dissections). Quantities of 
brooded young vary greatly—fewest (20 to 30) 
in midwinter specimens, most (100 to 200) in 
late spring to midfall specimens. The brood 
chamber often is so swollen with young that it 
is readily apparent in living animals. Young are 
crammed tightly into the chamber; external study 
of zooids divested of test may suggest only a few 
larvae, but dissection then reveals many dozens. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15 


The brood chamber is criss-crossed by many vas- 
cular trabeculae that connect the branchial sac 
and the atrial and gut wall and may keep loose 
young from being swept from the brood chamber 
by atrial water-currents. 

ECOLOGICAL DISTRIBUTION IN CALIFORNIA. — 
Intertidal. At Point Pinos and Pigeon Point, peak 
abundance is at about +0.3 m, and I have found 
no specimens at either site above +0.6 m or 
below 0.0 m. This is an open-coast ascidian in 
California, inhabiting surf-swept rocky habitats 
where the full force of the waves is broken by 
surrounding rocks and reefs. Zooids congregate 
on horizontal undersides of large boulders, usu- 
ally well back from the boulders’ edges. Such 
boulders restrict waterflow underneath, so much 
so that at Point Pinos the rocks that harbor Den- 
drodoa abbotti may lie partially in sand that by 
its odor and color appears to be virtually anoxic. 
At Pigeon Point most rocks with this ascidian 
are slightly propped up by their neighbors, so 
that oxygen remains plentiful in waters perco- 
lating or flowing underneath. Large boulders that 
do not have D. abbotti on them may shelter 
smaller rocks that do. Many rocks that seem ap- 
propriate for this species do not harbor speei- 
mens. This spotty distribution of aggregated in- 
dividuals may indicate a short swimming period 
and quick settlement by brooded larvae, or as 
yet unclear ecological restrictions on the adults. 
At Pigeon Point, other invertebrates found on 
surfaces with Dendrodoa abbotti include the 
anemone Epiactis prolifera, the polyclad Noto- 
plana acticola, the polychaetes Spirorbis and 
Salmacina, the barnacle Balanus glandula (and 
sometimes Chthamalus dalli), porcelain crabs 
such as Petrolisthes, several encrusting bryozo- 
ans such as Eurystomella bilabiata, the asteroid 
Leptasterias pusilla, and the aplousobranch as- 
cidian Aplidium californicum. But none of these 
associated invertebrates seems so severely kept 
back from the margins of boulders, so cryptic in 
its under-rock habitat, as Dendrodoa abbotti. 

ECOLOGICAL DISTRIBUTION IN WASHINGTON. — 
Subtidal. At Peavine Pass, San Juan Islands, 
specimens were dredged from 10 to 12 m. The 
species has been sought elsewhere in rocky areas, 
but only Peavine Pass, which is swept to the 
bottom by strong tidal currents, has proved a 
reliable site for collecting by this method, and 
even there the species is rarely taken. Debris har- 
boring Dendrodoa abbotti contains, as well, Bal- 


NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV. 


TABLE |. 


245 


DENDRODOA CARNEA AND D. ABBOTTI: CONSISTENT DIFFERENCES. 


Feature 


Color in life 


Siphonal apertures 


Dorsal tubercle 


Transverse branchial 
vessels and rows of stig- 
mata 


Internal longitudinal 
branchial vessels and 
folds (DL = dorsal 
lamina, (#) = number 
of vessels in fold, E = 
endostyle) 


Endocarps 


Margin of anus 


Ovary 


Testis 


Dendrodoa carnea 


Bright pink to blood red. 


Bilabial. 


Narrow ovoid slit whose axis is oriented almost 
anterior-posterior. 


17 or more. 


Left: DLO(1)0(1)0(1)0(1)0E as in D. abbotti. 
Right: DLO(4—5)0(1)0(1)0(1)OE; prominent low 
fold carrying at least 4 vessels on right pharyngeal 
wall. 


Many, small, widely scattered over entire atrial 
wall. 


“Reflected but not lobed, often somewhat two- 
lipped” (van Name 1912, p. 587). 


Straight along right ventral margin of body; ovi- 
duct continues so. 


Not clearly encapsulated with ovary, testis-lobes 
extend somewhat into body wall, predominantly 
ventro-lateral to ovary; all testis-lobes close to 


Dendrodoa abbotti 


Gray to ochre, occasionally reddish around si- 
phonal apertures. 


Bilabial, somewhat more pronouncedly so than 
in D. carnea. 


Fairly sharply bent “C” whose long axis is ori- 
ented laterally. 


Canl2: 


Left: DLO(1)0(1)0(1)0(1)OE as in D. carnea. 
Right: DLO(1)0(1)0(1)0(1)OE; no multi-vessel fold 
on right pharyngeal wall. 


Fewer, larger, more (but not entirely) confined to 
ventral atrial surface. 


Scalloped into usually 5 lobes. 


Along right-anterior ventral margin of body, then 
bends sharply into ascending limb, recurvés be- 
hind atrial siphon into dorsal oviduct that pro- 
jects posteriorly. 


Clearly not encapsulated with ovary, testis-lobes 
lie wholly in body wall, predominantly ventro- 
medial to ovary; often one posterior group of 


ovary. 


Spermiduct 


face of ovary (?) 


Brood chamber 
yond oviduct there. 


Brooded young 
brooded at a time. 


(?) as in D. grossularia, many short spermiducts 
converge in multiple spermipores on atrial sur- 


Extensive, including right-posterior region be- 


(From a small sample) only a few dozen embryos 


testis-lobes far from ovary. 


Single, long vas deferens on atrial surface of ovary 
receives vasa efferentia of all testis-lobes, ends 
mid-dorsally in spermipore-bearing papilla 
pointing at atrial siphon. 


More restricted to far posterior part of body. 


Many dozens to more than 100 embryos brooded 
at a time. 


anus nubilis (one of the best indicators that the 
ascidian may be present) and the hydrocoral A/- 
lopora. The ascidian occurs especially around the 
husks of dead barnacles and in crannies in large 
rocks. But dredging of course destroys the set of 
surfaces and actual relationships among mem- 
bers of the fauna at the site, and so no compar- 
ison can yet be made between the subtidal hab- 
itat of Dendrodoa abbotti at Peavine Pass and its 
intertidal circumstances at Pigeon Point. 

The bathymetric contrast between California 
and Washington (San Juan Islands) records of 
Dendrodoa abbotti is striking. The species may 
occur subtidally in California; its inaccessibility, 
beneath large boulders, could account for the cur- 


rent lack of such records by dredging or even by 
diving. But D. abbotti does not occur in the very 
low intertidal zone in California, below about 
mean low-low tidal levels. Thus, if it does occur 
subtidally, there is not a continuous distribution 
of the species from those depths to the low- to 
mid-tidal habitats where it characteristically is 
found. In contrast, in the San Juan Islands, I have 
not found the species at all intertidally in habitats 
that resemble California’s coastal sites—except, 
of course, for the lack of surf in the San Juans. 
Dendrodoa abbotti appears to be only a subtidal 
species in that archipelago. But to the west of the 
San Juan Islands, on the southwest coast of Van- 
couver Island, B.C., Dr. Ivan Goodbody has 


246 


found this species “‘on the underside of boulders 
at extreme low tide ... on the open coast north 
of Sooke.”’ Dr. Goodbody reports (pers. comm.) 
that the site there is “a rough boulder strewn 
shore with many large rounded boulders indi- 
cating heavy wave action.” His record thus ex- 
tends the intertidal range of D. abbotti into those 
Canadian habitats where surf does resemble Cal- 
ifornia’s. Dr. Goodbody’s Canadian specimens 
of D. abbotti are now in the collection of the 
California Academy of Sciences. 


DISCUSSION 


A. Comparison of Dendrodoa abbotti 
with D. carnea 


A comparison of Dendrodoa abbotti with the 
western North Atlantic species D. carnea—the 
styelopsid dendrodoan that most closely resem- 
bles D. abbotti— indicates an array of differences, 
some trivial, some marked, but all consistent. 
This comparison sets a great many specimens of 
D. abbotti against necessarily only a few dissected 
specimens of D. carnea (from the USNM col- 
lection) and others’ reports on D. carnea (see van 
Name 1912, 1945). But differences that emerge 
even in this perhaps unbalanced sampling of these 
species gain force as they become elements in a 
consistent pattern of distinctions between the two 
taxa, and this pattern has become more persua- 
sive with each examination of new specimens. 
Table 1 summarizes the comparison. 

Even arguably minor distinctions (for exam- 
ple, the character of the brood chamber or of the 
endocarps) take on significance in Table 1’s ar- 
ray. The two species are most effectively distin- 
guished, however, by the following criteria: 


1. shape and disposition of the ovary; 

2. testis-ovary relationship, including D. abbot- 
ti’s posterior group of testis-lobes; 

3. structure of the spermiducts, especially of the 
vas deferens; 

4. arrangement and number of internal longi- 
tudinal vessels of the right side of the pharynx; 

5. number of transverse branchial vessels and 
rows of stigmata on both pharyngeal walls; 

6. shape and orientation of the dorsal tubercle; 

7. color in life. 


B. Generic Traits 


By most accounts and diagnoses, in the genus 
Dendrodoa the testis and ovary are “encapsu- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No: 15 


lated’’ within a common sheath (Monniot and 
Monniot 1972), and the testis comprises many 
lobes that do not lie in the body wall but rather 
hug the parietal (away from the atrium) surface 
of the ovary (Huntsman 1913). Most of the go- 
nad of Dendrodoa grossularia shows this con- 
dition of encapsulation and testis-ovary juxta- 
position clearly, although some of the anterior 
testis-lobes do lie more in the body wall than 
wholly against the ovary. Dendrodoa carnea ex- 
hibits a somewhat looser gonadal arrangement: 
the testis-lobes apparently are still encapsulated 
with the ovary and lie largely against its parietal 
surface, but they extend into the adjacent body 
wall much more than do the testis-lobes of D. 
grossularia, especially to the lateral (right) side 
of the ovary. Dendrodoa abbotti carries this loos- 
ening of the testis-ovary bond further still: the 
testis-lobes of D. abbotti lie ““beneath”’ the ovary 
or close by on the medial (left) side of the ovary, 
but they lie in the body wall itself, not against 
the ovarian mass, and there is no sheath enclos- 
ing these gonadal elements into a single structural 
unit of intimately juxtaposed parts. And al- 
though most of the testis-lobes of D. abbotti lie 
very close to the ovary, there is often a group of 
testis-lobes lying in the far posterior atrial floor 
of the zooid, and actually on the left side of the 
zooid, although even this separated and isolated 
group is still connected by a vas deferens to the 
common spermiduct of all the other, “ovary- 
afhiliated”’ testis-lobes. 

Dendrodoa carnea is so much like D. grossu- 
laria (Traustedt’s (1883) type species of his genus 
Styelopsis, now a subgenus of Dendrodoa) that 
Arnbiick (1922) and Hartmeyer (1903) have both 
suggested these could be merely geographic vari- 
ants of a single species—a view not held, how- 
ever, by van Name (1945). The main distinction 
between these two species is their different num- 
ber of internal longitudinal branchial vessels, 
more numerous in D. grossularia than in D. car- 
nea. But the slight gonadal contrast reported here 
also seems to be a consistent one. The difference 
takes on added taxonomic significance when D. 
abbotti joins the comparison, because the genus 
thereby shows a series of testis-ovary juxtapo- 
sitions from a tightly joined one to an appreci- 
ably looser one—from the condition “‘character- 
istic” of the genus Dendrodoa to one rather akin 
to that of the genus Styela. 

Perhaps the perplexing Dendrodoa uniplicata 


NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV. 


Hartmeyer 1903, which Millar (1966) redesig- 
nates Styela uniplicata Bonnevie 1896 because 
“the structure of the gonad agrees better with 
Styela,” extends the grossularia-carnea-abbotti 
series of gonadal arrangements further while re- 
taining dendrodoan features of the pharnyx. Un- 
fortunately, the meager remnants currently 
available of Dendrodoa (or Styela) uniplicata will 
not by themselves resolve this question. 

Another dendrodoan trait from which Den- 
drodoa abbotti diverges involves the spermiduct. 
In the genus Dendrodoa, testis-lobes empty in 
groups into very short vasa deferentia or even 
more cloaca-like pits on the atrial surface of the 
ovary, and there are several such spermiporal 
loci on the ovary (Berrill 1950). The repetition 
of short vasa deferentia, each emptying a group 
of testis-lobes, is not usually as striking in D. 
grossularia as in the somewhat stylized depiction 
of this trait by Lacaze-Duthiers and Delage 
(1892), from which work many accounts of the 
species have been partly drawn. But Riedlinger 
(1902) indicates in his careful study how slight 
or even absent the vasa deferentia may be in that 
species, in place of which spermiporal loci serve 
the converging vasa efferentia of groups of testis- 
lobes. Dendrodoa carnea also appears to have 
multiple spermipores along the atrial surface of 
the ovary (again, though, a condition difficult to 
discern in dissections). In contrast, the gonad of 
D. abbotti has a single, long vas deferens, as in 
Cnemidocarpa and Styela (Fig. 2). All the sper- 
miducts of this species are exceedingly fine, and 
their disposition difficult to trace except in serial 
sections. Such a close scrutiny of D. carnea would 
seem appropriate, to find out if that species is 
intermediate between D. grossularia and D. ab- 
botti in this trait, as it is in testis-ovary juxta- 
positions. 

Dendrodoa (Styelopsis) abbotti is placed in 
Dendrodoa by its possession of a single gonad, 
and in Styelopsis because of its unbranched ovary 
and its simple pharynx, which lacks folds and 
possesses few internal longitudinal vessels. Den- 
drodoa abbotti is so much like D. carnea, which 
in turn is so much like D. grossularia, that this 
placement of the new species seems indisputable. 
But the consequence is to relax and modify long- 
held gonadal criteria of Dendrodoa, recognizing 
that species with styelan gonadal patterns or 
cnemidocarpan spermiducal patterns occur in the 
genus. 


ACKNOWLEDGMENTS 


A grant from the Faculty Research Committee 
of the Academic Senate of the University of Cal- 
ifornia, Santa Cruz has supported much of the 
research reported in this paper. I appreciate the 
assistance of Linda Cole, U.S. National Museum 
of Natural History, who guided me through the 
collection there, with the consequence that Den- 
drodoa carnea came into consideration at a crit- 
ical moment in this study. Professor Ivan Good- 
body has shared with both Professor Abbott and 
me several Canadian specimens of Dendrodoa 
abbotti and ecological information about their 
site; I am grateful for his help and for his read- 
iness to include this important northern inter- 
tidal record in this initial paper about the new 
species. Donald P. Abbott, without realizing at 
the time the nomenclatural consequence of his 
generosity, shared with me his notes and draw- 
ings of many years’ acquaintance with the species 
described in this paper, and I am most grateful 
for these and for many other ways in which he 
has encouraged me. 


LITERATURE CITED 


ARNBACK-CHRISTIE-LINDE, A. 1922. Northern and arctic in- 
vertebrates in the collection of the Swedish State Museum. 
8. Tunicata. 1. Styelidae and Polyzoidae. Jn Kungl. Svenska 
Vetenskapsakad. Handlingar 63(2):1-62, pls. 1-3. 

BeRRILL, N. J. 1950. The Tunicata, with an account of the 
British species. London: Ray Society. 354 pp. 

Bonnevig, K. 1896. Ascidiae simplices og Ascidiae Com- 
positae fra Nordhavs Expeditionen. Jn Norske Nordhavs- 
Expedition 23(2):1-16, pls. 3, 4. 

GALIGHER, A. E. AND E. N. Koztorr. 1964. Essentials of 
practical microtechnique. Philadelphia: Lea & Febiger. 484 
pp. 

HarTMEYER, R. 1903. Die Ascidien der Arktis. Jn Rémer, 
F. and F. Schaudinn, Fauna Arctica 3(2):91—412, pls. 4-14. 

Huntsman, A. G. 1913. The classification of the Styelidae. 
Zool. Anz. 41:482-501. 

LacaAzeE-DuTuierRS, H. DE AND Y. DELAGE. 1892. Etudes sur 
les ascidies des cétes de France. Faune des Cynthiadées de 
Roscoff et des cétes de Bretagne. Mém. Acad. Sci. France 
(ser. 2) 45:1-323. 

Licut, S. F., R. I. Smitu, F. A. Pitecka, D. P. ABBotT AND 
F. M. Weesner. 1954. Intertidal invertebrates of the cen- 
tral California coast. 2nd ed. Berkeley: Univ. Calif. Press. 
446 pp. 

Miiiar, R. H. 1966. Tunicata Ascidiacea. Marine inverte- 
brates of Scandinavia, No. 1. Oslo: Universitetsforlaget. 123 
pp. 

Monniot, C. AND F. Monniot. 1972. Clé mondiale des gen- 
res d’ascidies. Arch. Zool. Exp. Gén. 113:311-367. 

RIEDLINGER, R. 1902. Untersuchungen iiber den Bau von 
Styelopsis grossularia des Ostsee. Nova Acta Akad. Leop.- 
Carol., Halle 81:1-62, pls. 1-6. 


248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No.*15 


Smitu, R. I. AND J. T. CarLTon, eds. 1975. Light’s manual: vAN NAME, W.G. 1912. Simple ascidians of the coasts of 


intertidal invertebrates of the central California coast. 3rd New England and neighboring British provinces. Proc. Bos- 
ed. Berkeley: Univ. Calif. Press. 716 pp. ton Soc. Nat. Hist. 34:439-619, pls. 43-73. 
Traustept, M. P. A. 1883. Vestindiske ascidiae simplices. 1945. The North and South American ascidians. 


2. Molgulidae og Cynthiadae. Vid. Medd. Naturhist. Kjé- Bull. Amer. Mus. Nat. Hist. 84:1-476, pls. 1-31. 
benh., ann. 1882:108—136, pls. 5, 6. 


PROCEEDINGS __ 
tHe JAN 7 


OF 


CALIFORNIA ACADEMY, OF SCIENCES. 


Vol. 43, No. 16, pp. 249-267, 6 figs., 6 tables 


; 
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uy 
16 8 WR eee 


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1985 


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 


Asstract: 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 
resulting 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 Miller, 1834) and Japan 
(E. 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. 
stoutii (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 Giinther’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 Miiller (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) listed a species having 
seven apertures from the Caribbean Sea. Fern- 
holm (1982) has further described it as new. 

In general, we concur with Fernholm 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 
Miiller (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, 
horny 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: Bernice 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). Fernholm and Hubbs 
(1981) reported many of these methods in their 
study of the eastern Atlantic Eptatretus. When 


' Hagfishes are not parasitic; they scavenge dead or mori- 
bund fishes and invertebrates. 


McMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES 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, Na. 16 


FiGur_E 1. 


Po oo © e0000 +” 


A-B: Sketches of an Eptatretus and a Myxine, showing regions of body used in study of myxinids: | to 5, total 


length; | 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 PACIFIC HAGFISHES 


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- 
cilitated 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 mght 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 


253 


Ficure 2. Cusps and basal plates, in excised and spread 
condition, of E. carlhubbsi, paratype USNM 233742, 955 mm 
ales 


embedment in the cartilaginous dental plate is a 
multicusp. 

The two paired sets of cusps (the outer and 
inner rows of Fernholm and Hubbs [1981] and 
Fernholm [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 


254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, Na. 16 


FIGURE 3. 


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 


Ventral view of branchial region of: 1, E. carlhubbsi: 2-3, E. laurahubbsi, showing diversity in afferent branchial 
arteries (ABA) leading off from branches of ventral aorta (VA); 4, E. strahani; 5, E. cirrhatus. 


in marked contrast to the condition shown by 
scanning electron microscopy of E. springeri 
(Fernholm 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 PACIFIC HAGFISHES 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 Paramyxine (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. /aurahubbsi). 

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. 61D) 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 (E. 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 


Eas taraihiamn 


Ficure 4. Sketches (not to scale) of head regions of Ep- 
tatretus cirrhatus and E. strahani showing arrangements of 
sensory canals. The first two pairs of barbels are omitted. 


very small and the surface indications faint, and 
that any apparent erratic appearance might be 
due to the observer. However, on the specimens 
examined by us the canals, when present, were 
readily visible under adequate magnification and 
lighting, and often by the unaided eye. They ap- 
pear as thin lines, about 1-3 mm long, variably 
straight or curved (Fig. 4), often very slightly 
raised above the skin, and sometimes covered 
with a coating of coagulated slime. Histological 
examination was not done, nor have we attempt- 
ed to observe these canals on unpreserved fishes. 

Old, healed scars are often present in areas 
occupied by the sensory canals, and elsewhere 
on the body, mostly anteriorly. These are iden- 
tifiable as shallow depressions, usually wider and 
longer than the sensory canals. Many scars occur 
singly, but often they are in groups of parallel 
lines, the spacing closely resembling that of the 
anterior cusps. Possibly this scarring occurs when 
many hagfishes are feeding in close proximity 
competing for food, or when crowded in a trap. 


256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, Na. 16 


TABLE 1. AVERAGES AND RANGES (IN THOUSANDTHS OF TOTAL LENGTH) OF SELECTED Bopy PROPORTIONS FOR FOUR SPECIES 
OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN. 


N (size range in mm) 


Preocular length 
Prebranchial length 
Branchial length 


Trunk length 
Tail length 


E. carlhubbsi 


9 (813-1160) 


Avg. (range) 


38 (36-54) 
184 (168-197) 
68 (55-77) 
602 (577-623) 
160 (145-176) 


E. laurahubbsi 


8 (240-375) 


Avg. (range) 


50 (44-59) 
193 (184-204) 

55 (52-59) 
561 (545-585) 
198 (181-213) 


E. strahani 


5 (265-520) 


Avg. (range) 


63 (57-68)* 
220 (210-231) 

77 (69-83) 
521 (500-537) 
182 (174-202) 


E. cirrhatus 


8 (481-655) 


Avg. (range) 


60 (52-67) 
225 (214-239) 
76 (69-89) 
550 (525-563) 
154 (135-168) 


Tail depth 97 (89-105) 90 (82-99) 117 (109-125) 83 (77-91) 
Body depth with finfold No finfold 89 (74-97) 111 (101-117) 89 (69-102) 
Body depth without finfold 93 (78-106) 81 (73-91) 98 (94-105) 88 (69-102) 
Body depth at cloaca 73 (65-85) 70 (61-80) 87 (77-94) 67 (57-75) 


* Due to lack of visible eyespots, the preocular length was taken from center of uncovered pupil. 


Waite (1909) placed three adult EF. cirrhatus in 
a bucket of formalin and observed them savagely 
attacking each other. One was bitten at least 15 
times by the other two. 


KEY TO SEVEN-GILLED SPECIES OF Eptatretus 
FROM THE PACIFIC OCEAN 


la. Slime pores of trunk 60-70, low, well be- 
low mid-lateral aspect. Total cusps 61-71. 
JENS OU folteeinth 2 

lb. Slime pores of trunk 45-53, high, near 
mid-lateral aspect. Total cusps 43-53. 
Eyespotsipresent.on absent ee 3 


2a. Ventral finfold absent. Two (rarely three) 
fused cusps on anterior multicusps, three 
on the posterior. Eyespots large, promi- 
Neca nee west ee E. carlhubbsi n.sp. 

2b. Ventral finfold prominent. Two (rarely 
three) fused cusps on each of the four mul- 
ticusps. Eyespots present == 
aE Sit SEER A oe Re E. laurahubbsi n.sp. 


3a. Ventral finfold readily visible. Eyespots 
absent. Ventral margin of tail forming a 
nearly straight line from cloaca to abrupt 
beginning of curve around tail. Anterior 
few gill apertures small, slitlike. No pale 
rings around slime pores or gill apertures. 
Three fused cusps on each of the four mul- 
LICUS) Sees ae ee ee, E. strahani n.sp. 

3b. Ventral finfold vestigial. Eyespots present. 
Tail margin smoothly ovate. All apertures 
rounded. Pale rings around slime pores 
and gill apertures. Three fused cusps on 
each of the multicusps E. cirrhatus 


Eptatretus carlhubbsi new species 


Hotoryrpe.—SIO 68-473, mature female, 961 mm TL, taken 
at 19°18’N, 166°33.5'E, near Wake Island, in a free-vehicle 
trap on bottom at 1574 m, 12-13 Sept. 1968. 

PARATYPES. —SIO 68-473, female, 810 mm TL, taken with 
the holotype; SIO 82-63 (formerly BPBM 27850), female, 1125 
mm TL, taken at Brooks Banks, between French Frigate Shoals 
and Gardner Pinnacles, Leeward Islands, Hawaii, Nov. 1981, 
Mokihana Cruise 81-12, set 35, shrimp trap, depth not given; 
BPBM 27848, male, 1160 mm TL, taken at 12°56'N, 166°22’W, 
French Frigate Shoals, Leeward Islands, Hawaii, 7 Nov. 1981, 
shrimp trap at 684 m; BPBM 27851, male, 830 mm TL, taken 
off the north shore of Molokai Island, Hawaii, 26-27 Dec. 
1981, shrimp trap at 659 m; USNM 227440, male, 900 mm 
TL, taken at 24°48’N, 167°14’W, R/V Cromwell Cruise 80-05, 
Station 57, ina shrimp trap at 835 m; USNM 233742 (formerly 
NMFS P-0289), male, 955 mm TL, taken at 14°59’N, 145°13’E, 
Esmeralda Bank, Guam, 5-6 April 1981, Cruise Typhoon 81- 
01, Station 151, in a shrimp trap at 1061 m; CAS 50705 
(formerly BPBM 27847), male, 1064 mm TL, Leeward Islands, 
Hawaii, Nov.—Dec. 1981, depth and method of capture not 
given; CAS 50706 (formerly BPBM 27849), male, 980 mm 
TL, taken at French Frigate Shoals, East Plateau, north side, 
Leeward Islands, Hawaii, 19 Nov. 1981, in a shrimp trap at 
481 m. 


DiaGcnosis.—A seven-gilled Eptatretus having 
no ventral finfold, very large eyespots, two (rarely 
three) fused cusps on the anterior multicusps and 
three on the posterior. 

DESCRIPTION. — Counts: Those of holotype giv- 
en first (left and right sides), followed by ranges 
for all specimens in parentheses: gill apertures 7, 
7 (all); prebranchial slime pores 15, 16 (12-17); 
branchial pores 7, 7 (6-8); trunk pores 60, 61 
(60-70); cloacal pores 2, 2 (1-3); caudal pores 
11, 11 (11-13); tail pores 13, 13 (12-16); total 
slime pores 95, 97 (93-110). Cusps on anterior 
multicusps 2, 2 (rarely 3); posterior multicusps 
3, 3 (all); anterior unicusps 16, 16 (15-17); pos- 


McMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES 257 


TABLE 2. PREBRANCHIAL, BRANCHIAL, AND TAIL SLIME Pores OF Four SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EP74- 


TRETUS) FROM THE PAciFIC OCEAN. 


Prebranchial slime pores 


12 13 14 15 
E. carlhubbsi l 7 2 5 
E. laurahubbsi 2 3 
E. strahani 1 4 2 
E. cirrhatus 

Branchial slime pores 

6 7 8 
E. carlhubbsi 4 12 2 
E. laurahubbsi 7 6 3 
E. strahani 10 
E. cirrhatus 7 34 2 

Tail slime pores! 

10 11 12 13 
E. carlhubbsi 2 4 
E. laurahubbsi 
E. strahani 3 4 2 
E. cirrhatus 1 6 16 14 


' Tail count is the total of the cloacal and caudal slime pores. 


terior unicusps 12, 13 (11-13); total cusps 68 
(64-71). 

Morphometry: In thousandths of total length; 
values for holotype given first, followed by ranges 
for all specimens: preocular length 38 (36-54); 
prebranchial length 184 (168-197): branchial 
length 68 (55-77); trunk length 602 (577-623); 
tail length 160 (145-176); tail depth 97 (89-105); 
body depth 90 (78-112); depth at cloaca 74 (62- 
86). 

All specimens from Hawaii were frozen ini- 
tially; the body proportions of these may not be 
closely comparable to the other collections, which 
were initially preserved in formalin. It is not 
known what effect freezing may have on subse- 
quent shrinkage, but it is possible that the soft 
tissues of hagfishes are greatly affected by the 


16 17 18 19 20 N 


2 18 
8 3 16 
3 10 
4 17 17 4 1 43 


b 

NO 
—\ 
oO 


lon) 
ae 
Ww 


expansion of cells in freezing. It is known that 
length of time in preservative significantly affects 
the total length; a shrinkage of 10% is not un- 
common. However, to our knowledge no study 
has been done showing the changes in other body 
proportions. Body proportions (Table 1) and 
counts (Tables 2-6) are compared with similar 
data for other seven-gilled Eptatretus from the 
Pacific Ocean. 

Body robust; prebranchial region slightly 
deeper than wide; body increasingly compressed 
laterally to tail, varying in greatest depth from 
8% to 11% of TL. Two to four GP anterior to 
tip of DM, which is somewhat flattened poste- 
riorly. Length of DM 19% (17-21%) of TL, its 
width 15% (9-19%)- of its length, its depth 66% 
(57-87%) of its width. VA short, wide, its width 


TABLE 3. TRUNK SLIME Pores OF Four SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN. 


Trunk slime pores 


45 46 47 48 


E. carlhubbsi 

E. laurahubbsi 

E. strahani D 
E. cirrhatus 


Mm nN 
NN 
Ww 


OG Ie 3 5; 9225 


49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 N 


I 2 3 I ays Des 
3 le 2, 52 [hae 2095) 16 
9 
43 


258 


TaBLe 4. Tota SLIME Pores OF Four SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN. 


Total pores 
82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109110 N 


WS Wi TS TE) SO tit 


18 


N 


iN 


nN 


E. carlhubbsi 
E. laurahubbsi 


E. strahani 
E. cirrhatus 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 


N 


AN 


ioe) 
vr 


39% (35-43%) of its length. Distance between 
DM and branching of VA 17% (12-22%) of DM 
length. GA rather evenly spaced in a line often 
sloping or curving slightly downward posteriorly, 
with the exception of paratype BPBM 27851, 
apparently an aberrant specimen. It has eight GP 
on each side, but with corresponding apertures 
arranged abnormally (Fig. 6); there are eight GA 
on the left side, but only seven on the right, with 
two gills sharing one aperture. 

Head blunt, face sloping at about 30° from the 
vertical. Eyespots large, prominent, ranging from 
nearly round to square or rectangular. Removal 
of overlying integument shows the embedded eyes 
varying from under to well behind center of eye- 
spots. Eyes nearly round, about 5 by 6 mm, slant- 
ing ventrodorsally at about 45°. The first two 
pairs of barbels of nearly equal length on the 
holotype, the second pair slightly shorter than 
the first. Proportions of the barbel pairs on the 
paratypes show great variation, with some of the 
first pairs shorter than the second. The second 
barbel, right side, of paratype SIO 68-473 is bi- 
furcate to the base, both halves of about equal 
length. No sensory canals are present on any of 
the specimens. 

There is no ventral finfold, but an intermittent, 
vague line resembling a median suture extends 
from well behind PCD nearly to cloaca. The thin 
margin of the tail, common on other hagfishes, 
is not readily evident in these large specimens. 
If a caudal finfold is present, it 1s very thick, 
tapering steeply from the tail musculature. This 
may be a species character or merely the result 
of age or size; no juveniles are known. In the 
other three species discussed below (particularly 
the juvenile specimens of FE. /aurahubbsi) the tail 
margins are notably deeper and thinner. The av- 
erage tail length in the nine specimens of E. carl- 
hubbsi is about twice (1.9-2.5) the body depth 
at the origin of the cloaca, but the shapes of the 
tails vary considerably (Fig. 5). The deep notch 
on the dorsal surface of the tail of the holotype 
is apparently an old, entirely healed injury; no 
notch is present on any paratype. The tail depths 
average 58% (49-65%) of the lengths; shape var- 
ies from that of a truncated club (No. 7) with no 
evidence of finfolds to the very deep, expanded 
form of No. 3, which has a notably less thickened 
margin. 

The holotype and three paratypes are tan in 
color; two are from Wake Island, and one each 


McMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES 259 


TABLE 5. UNICUSPS IN THE ANTERIOR AND POSTERIOR SETS OF CusPs OF FouR SPECIES OF SEVEN-GILLED HAGFISHES (GENUS 


EPTATRETUS) FROM THE PACIFIC OCEAN. 


Anterior unicusps 


13 14 15 16 I07/ N 
3) 10 3 18 

l 1 4 7 3 16 
10 

43 


Posterior unicusps 


8 9 10 11 12 
E. carlhubbsi 
E. laurahubbsi 
E. strahani 2 4 4 
E. cirrhatus 7 28 7 l 
7 8 9 10 11 
E. carlhubbsi 
E. laurahubbsi 
E. strahani p 5 3 
E. cirrhatus D 26 15 


from French Frigate Shoals and Molokai, Ha- 
wali. Five paratypes (one from Guam, four from 
Leeward Islands, Hawaii) are a purplish brown; 
the Guam specimen has a more brownish over- 
tone. Scattered, irregularly shaped pale areas of 
varying sizes occur mainly on the anterior por- 
tions of the body and occasionally as small patches 
on the other parts; these light spots are more 
visible on the purplish-brown specimens. Small, 
very dark brown blotches occur randomly on all 
but one of the tan specimens. Pale rings surround 
the GA, but not the slime pores. The color dif- 
ferences are striking, but such variation is known 
in other species wherein blotching, lighter shades, 
and even albinism occur (Dean 1903; Jensen 
1959). 

The color variations discussed above and the 
striking differences in tail shapes and barbel ra- 
tios, as well as wide ranges in other body pro- 
portions and differences, often indicate the pres- 
ence of more than one species, but we find no 
definite correlation between these characters and 
those of counts, sex, other body proportions, or 
geographical areas. Further collections may show 


12 13 14 15 16 N 
8 7 18 
10 4 1 16 
10 
43 


that subspecies or other taxa occur within or be- 
tween the three areas— Hawaii, Wake Island, and 
Guam. 

The first few prebranchial pores curve gently 
downward on the holotype, but on all paratypes 
they are in an essentially straight line (occasion- 
ally the first one or two are slightly depressed). 
There are seven or eight slime pores in the bran- 
chial region, one or two more than expected, 
which is one less than the number of GA. 

Eggs: The holotype contained at least 21 eggs, 
the largest 58.5 by 14.4 mm. These eggs had been 
removed and bottled separately by an earlier in- 
vestigator, and it is not known that all were pre- 
served. None were in the specimen when ex- 
amined by us. Paratype BPBM 27848 contained 
32 eggs, the largest 75.5 by 16.5 mm. Some eggs 
were damaged by freezing, the original preser- 
vative, and were loose in the body; these are now 
bottled separately. Eight eggs, the largest 58.5 by 
14.4 mm, were packaged separately within a larg- 
er pack containing a male and a female (SIO 82- 
63, formerly BPBM 27850). The female con- 
tained only 13 eggs, the largest 63.0 by 12.6 mm. 


TABLE 6. ToTAL Cusps OF Four Species OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN. 


Total cusps 


43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 N 


E. carlhubbsi 

E. laurahubbsi 
E. strahani 1 
E. cirrhatus 1 


eae Ni eae Ba) l 
2 lig) owt 1 


260 
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 


(Ue ees iV 
o) 10 20cm 


McMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES 261 


) R08: 
608: oo: Ene ; 


FiGure6. Sketch ofabberrant arrangement of gill apertures 
of a specimen of EF. carlhubbsi, paratype BPBM 27851, 830 
mm TL. 


Although there was no identifying label with the 
separate eggs, we assume they had been taken 
from the same female. Since the Hawaiian spec- 
imens apparently were all frozen as initial pre- 
servative, these eggs could have been stripped 
prior to immersion in preservative; the specimen 
had not been cut open anywhere on the body. 
Although all were extremely large, none of the 
eggs cited above had the terminal hooks of fully 
ripe eggs (Dean 1899; Jespersen 1975). 

EtyMoLocy.— With great respect and admi- 
ration we dedicate this species of giant hagfish 
to the late Carl L. Hubbs, himself a giant in ich- 
thyology. 


Eptatretus laurahubbsi new species 


Hototype.—SIO 65-643, juvenile female, 375 mm TL, tak- 
en at 33°31'S, 78°5S0’W, near Mas a Tierra, Islas Juan Fernan- 
dez, in a free vehicle trap on bottom at 2400 meters, between 
hours of 2030 and 0830, 12-13 Dec. 1965, Cruise 12 of R/V 
Anton Bruun. 

PARATYPES (remaining material examined).—Seven juve- 
niles (sex questionable) taken with the holotype, are deposited 
as follows: SIO 65-643, two, 369 and 287 mm TL (deposited 
with the holotype); CAS 49125, two, 287 and 358 mm TL; 
USNM 227441, two, 240 and 265 mm TL, Museo Nacional 
de Chile, Santiago, one, 240 mm TL. 


Diacnosis.—A seven-gilled Eptatretus having 
a well-developed finfold and only two (rarely 
three) fused cusps on each of the four multicusps. 

DESCRIPTION. —Counts: Those of the holotype 
given first (left and right sides), followed by ranges 
for all specimens in parentheses: gill apertures 7, 
7 (all); prebranchial slime pores 17, 16 (14-17); 
branchial pores 7, 7 (6-8); trunk pores 67, 66 
(60-67); cloacal pores 2, 3 (2-3); caudal pores 
12, 12 (11-14); tail pores 14, 15 (14-16); total 
slime pores 105, 104 (97-105). Cusps on anterior 


eS 
Ficure 5. 


multicusps 2, 2 (2, 3); posterior multicusps 2, 2 
(2-3); anterior unicusps 15, 16 (13, 17); posterior 
unicusps 12, 12 (11-16); total cusps 63 (61-68). 

Morphometry: Values in thousandths of TL 
given first for the holotype, followed by ranges 
for all specimens (left side): preocular length 53 
(52-59); trunk length 560 (545-585); tail length 
189 (181-213); maximum body depth including 
finfold 89 (74-97), excluding finfold 84 (73-91); 
body depth at cloaca 69 (61-80); tail depth 82 
(82-89). Morphometric data (Table 1) and counts 
(Tables 2-6) are compared with similar data for 
other seven-gilled Eptatretus from the Pacific 
Ocean. 

Body deeper than wide, width at midbody 
about 1.6 in depth. Tail broadly ovate, its depth 
slightly greater than body depth. Ventral finfold 
well developed, originating well behind PCD; a 
broad, thin finfold from posterior margin of clo- 
aca around tail and dorsally until about over 
anterior margin of cloaca (Fig. 5—1). Dorsal pro- 
file of head sloping to a very blunt, nearly straight- 
across rostrum; width of nasopharyngeal 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 


Tail shapes (to scale) and patterns of occurrence of the last four trunk pores and cloacal and tail slime pores of 


four species of seven-gilled Eptatretus from the Pacific Ocean: 1-9 E. carlhubbsii(T = 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, 1160 mm TL; 7—CAS 50706, 908 mm TL; 8—SIO 82-63, 1125 
mm TL; 9—BPBM 27851, 830 mm TL; 10—Holotype, F. /aurahubbsi, SIO 65-643, 375 mm TL; 11—Holotype, E. strahani, 


MNHN 1978-462, 520 mm TL; 12—E. cirrhatus, 655 mm TL. 


262 


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 of EF. /aurahubbsi 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 E. 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 EF. /aurahubbsi, 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 
itis highly probable that adults exceed one meter 
in length. The longest (holotype), although 373 
mm TL, contains minute eggs, seen with difh- 
culty under magnification. In another Eptatretus 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 


(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. 

EtTyMOLoGy.— 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 


Ho.otyre.—MNHN 1978-462, female, 520 mm TL, con- 
taining eggs of about 4.5 mm in length, taken at 14°00’N, 
120°18'.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. 


Diacnosis.—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 PACIFIC 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, 
followed 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 (S00- 
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 
amin)! 

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. 

EtTyMoLoGy.— We are pleased to dedicate this 
new species to Ronald Strahan in acknowledg- 
ment of his important contributions to the study 
of Myxinidae. 


264 


Eptatretus cirrhatus (Bloch and Schneider, 1801) 


Petromyzon cirrhatus BLOCH AND SCHNEIDER, 1801:532 (orig- 
inal description fide Forster ms II:24 [habits: New Zealand)]). 

Homea banksii Fremtnc, 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, 1834:7 (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, 1904:21 (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, 1967:30 (shore 
to 1800 ft; more abundant south of Hawke Bay than else- 
where in New Zealand); WHITLEY, 1968:4 (synonymy); Scott, 
GLOVER, AND SouTHcoTT, 1974:19 (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, 173°41’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, 595 mm TL, 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°51’E, agassiz trawl, 175 fms (320 
m), 27 July 1968; AMS Kapala Station 71-07-03, female, 505 
mm TL, 33°33’—37'S, 152°01'-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°11'S, 152°23’E, otter trawl, 310 fms 
(567 m), 29 April 1971; AMS Kapala Station 71-11-07, two 
females, 491,514 mm TL, 34°40’-35°01'S, 151°10'-07’E, otter 
trawl, 300 fms (549 m), 7 July 1971; AMS Kapala Station 71- 
11-08, 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 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16 


71-11-10, male, 447 mm TL, 35°1 1’-37'S, 150°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. 


DiaGcnosis.—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 Museum, 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 slime pores 86 
(83-90). Three fused cusps on each of the four 
multicusps; 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 
(77-91); 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 PACIFIC HAGFISHES 


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 


265 


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 EF. cirrhatus; and Richard H. Ro- 
senblatt, for the holotype and one paratype of E. 
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, Bernice P. Bishop Museum, Honolulu. 

All specimens of FE. Jaurahubbsi were taken by 
the junior author on Cruise 12, 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 


preciate the efforts of all persons involved. R. H. 
Rosenblatt critically read the manuscript. 


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Scott, T. D., C. J. M. GLover, AND R. V. Sourucott. 1974. 
The marine and freshwater fishes of South Australia (2nd 
ed.). Gov’t. Printer, S. Australia. 1-329. 

STRAHAN, R. 1975. Eptatretus longipinnis, n.sp., a new hag- 
fish (family Eptatretidae) from South Australia, with a key 
to the 5—7 gilled Eptatretidae. Australian Zool. 18(3):137- 
148. 


McMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES 267 


TEMMINCK, C. J., AND H. SCHLEGEL. 1842-1850. Pisces. Pages Government trawling expedition, 1907. Rec. Canterbury Mus. 
1-345 in Fauna Japonica. Siebold, Fr. de. A. Arns et Socios, 1(2):1-26. 
Lugdini. Plates and Suppl. Batavorium (=Leiden). Wuittey, 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 94118 


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Vol. 43, No. 17, pp. 269-282, 5 figs. 


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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- 
CAB POTCS ee an ste eee eee ee Blakea 

Anthers linear-oblong to lanceolate or subu- 
late, the dorsally inclined apical pores ap- 
proximate and often confluent at anthesis 
ie et dass Alan tte Br ete Avtin, 2 Aides Ranh 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; Coclé: Knapp 
5334, CAS; Panama: Sytsma 1154, CAS). These 
additions increase the number of described Pan- 
amanian species to thirteen. 


Blakea elliptica (Gleason) Almeda, comb. nov. 
Topobea elliptica GLEASon, Phytologia 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 
internodes 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, 5-11.5 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—11 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 | 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°21'W, 2200 m, 15 Mar. 1982, Knapp et al. 4233 (MO); end 
of road past Palo Alto NE of Boquete in forest along ridge; 
elev. 6200-6800 ft [1890-2073 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 
verrucose 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 limits 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 of 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- 
florescentia foliorum 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 acuminato basi 
rotundata vel subcordata, chartacea et integra, 
5-7-plinervata. Inflorescentiae pauciflorae in fo- 
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 triangularibus, 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 nec prolongatum nec appendiculatum. 


Stylus 5.5 mm glaber; stigma truncatum; semina 
0.5-1 mm pyriformia. 

Shrubs 1-2 m tall. Older cauline internodes 
glabrate and terete. Distal branchlets, vegetative 
buds, juvenile leaves, and inflorescences copi- 
ously covered with sessile, and stipitate-stellate 
trichomes, the distal cauline internodes 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 apically 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 | 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- 


272 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 


\— 


Ficure 1. Clidemia tetrapetala Almeda. A, habit, x ca. ¥4; B, representative leaf (lower surface), x ca. 34; C, mature berry, x 
ca. 4; D, petal, x ca. 6; E, stamens, % lateral view (left) and dorsal view (right), x9; F, seeds, x ca. 7. (A from Folsom 4871; 
B-F from Folsom 6106.) 


E 


ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 273 


i A sl SE 
STN ay >) 


Than ana} 
mat wpe 


Figure 2. Clidemia trichosantha Almeda. A, habit, x ca. 34; B, representative leaf (lower surface), x ca. 4; C, representative 
flower, buds, bracts and bracteoles, x ca. 4; D, petal, x7; E, stamens, lateral view (left) and ventral view (right), x9; F, seeds, 


x ca. 14. (4-E from the holotype; F from Hammel 965.) 


ry reportedly opaque green but turning translu- 
cent red at maturity, globose, 5-6 mm long to 
the torus, 5-6 mm in diameter. Seeds narrowly 
pyriform, brown, smooth, vernicose, and mostly 
0.5-1.0 mm long. 


Typre.—Panama. Bocas del Toro/Chiriqui Border: Cerro 
Colorado, along intersection of Bocas Road with main ridge 
road, 11.8 km from Chami along path headed into Bocas del 
Toro, elevation 1400-1700 m, 24 Oct. 1977, Folsom 6106 
(holotype: CAS!; isotype: MO). 

ADDITIONAL SPECIMENS EXAMINED.—Panama. Bocas del 


274 


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 1400- 
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 differs most conspicu- 
ously in having ovate-elliptic, 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 1-3(—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; flores 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- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 


go-lanceata. Stamina isomorphica glabra; 
filamenta 2.5 mm longa; antherarum thecae 
1.5 x 0.75 mm oblongae, connectivum nec pro- 
longatum nec appendiculatum. Stylus 4.5 mm 
glaber; stigma punctiforme; semina 0.5 mm cu- 
neata. 

Slender shrubs 1-2.5 m tall. Internodes terete, 
glabrate at maturity, but vegetative buds, young 
leaves, and distal branchlets moderately to 
densely covered with smooth, eglandular, 
spreading trichomes mostly 1—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 axillary, 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, | 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 verruculose angles and a promi- 
nent + translucent lateral raphe, 0.5 mm long. 


Type.— Panama. Coclé: sawmill above El Cope, in forest 
along stream east of sawmill on the Atlantic drainage, elevation 
2300 ft (701 m), 27 July 1978, Hammel 4133 (holotype, CAS!; 
isotype, MO). 

ADDITIONAL SPECIMENS EXAMINED. — Panama. Coclé: above 
El Petroso sawmill at Continental Divide, N of El Cope, 13 
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. 1977, Folsom et al. 7093 
(CAS); El Cope on Pacific side about 2 hour walk from the 
sawmill, 16 Oct. 1979, Antonio 2116 (CAS). Panama: forested 
slope along El Llano-Carti road, 12 km from Panamerican 
Highway, 10 Sept. 1976, D’Arcy 10617 (CAS). San Blas: 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 dichasiai inflorescence of pedunculate ter- 
minal glomerules, the distally constricted fila- 
ments, the cuneate seeds with verruculose angles, 
and the sessile, paired bracts and bracteoles that 
are often fused into amplexicaul collars. 

A fruiting collection from Cerro Habt, San 
Blas (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. 


Graffenrieda 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 acuminato 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. 


Graffenrieda bella Almeda. A, habit, x34; B, representative leaves, lower surface (left) and upper surface (right), 


x ca. %4; 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 et al. 7581.) 


therarum thecae 3—4.5 x 0.5 mm oblongo- 
subulatae; connectivum vix (0.25 mm) prolon- 
gatum, dente dorsali 0.5 mm longo acuto. Stylus 
7.5-9.5 mm glaber; stigma punctiforme; semina 
numerosa 1 mm longa, recta paulo angulata. 
Trees to 5 m tall. Cauline internodes 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 2T7 


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 '%4 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. 


Tyre.— Panama. Chiriqui: along road between Gualaca and 
the Fortuna Dam site at 10.1 m; NW of Los Planes de Hornito, 
elev. 1300 m, 8 Apr. 1980, Antonio 4078 (holotype: CAS!; 
isotype: MO). 

ADDITIONAL SPECIMENS EXAMINED.—Panama. Chiriqui: 
Cerro Hornitos, 40 km NW of Gualaca in cloud forest dom- 
inated by Quercus spp., Podocarpus, and Drimys, 26 Jul. 1975, 
Mori & Bolten 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). 


DiIsTRIBUTION.— Known only from cloud for- 
ests on Cerro Tute in Veraguas province and 
Cerro Hornitos 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 (13-20 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 1000 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, inflorescence 
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 relies 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 


278 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 


Ficure 4. Miconia arboricola Almeda. A, habit, x ca. 3; B, representative leaf (lower surface) with enlargement (left), x 
ca. 2; 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 x 3.5-7.1 cm late ovata vel cordata, apice 
breviter acuminato basi subcordata vel cordata, 
5—7-nervata, firme 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 279 


antherarum thecae 1.5 x 0.5 mm oblongo- 
cuneatae; connectivum nec prolongatum nec ap- 
pendiculatum. Stylus 4 mm glaber; stigma cap- 
itatum; semina 0.5 mm galeiformia. 

Pendent woody vines or hemiepiphytes ad- 
hering to bark of host tree by adventitious roots. 
Older cauline internodes 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 vernicose 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 rounded 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 | 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. 3 to 4 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, 3-4 mm long and 3-4 mm in 
diameter. Seeds galeiform, white, papillate to 
costate-papillate, 0.5 mm long. 


Type.— Panama. Chiriqui: slope NW of confluence of Rio 
Hornito 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 
Urén, Cordillera Talamanca, | Sept. 1898, Pittier 12683 (US). 
Panama. Coclé: hills N of El 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 El 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. 

Miconia arboricola is quite unlike all other 
species of Miconia 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. mollicula Triana and M. cordata Triana 
in section Miconia. In foliar shape, internodal 
pubescence, and inflorescence structure, M. ar- 


280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17 


Pach 


Wi ; 
fii 


\ i: 

Whiaeeaaier \) 
pie CNH a 
\Eae fs ve 


Nes wy ye ) 


eB D 


Ficure 5. Topobea calophylla Almeda. A, habit, x2; B, stamens, ventral view, x1; C, hypanthium with floral bracts 
removed, x ca. 1%; D, cauline trichomes, x ca. 4; E, petal, x ca. 1%; F, outer floral bract (outer surface), x ca. 1; G, inner 
floral bract (inner surface), x ca. 1. (A & D from Nee 9873; B, C & E from Mori & Kallunki 5353; F-G from the holotype.) 


ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA 281 


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 7opobea to be described from Panama since 
the appearance of Gleason’s (1958) summary. 
The following description of 7. calophylla, the 
transfer of T. elliptica to Blakea (also proposed 
herein) and the discovery of 7. 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. (1-)3—9 mm longis in- 
duti. Petioli 2.2-5 cm longi; lamina 14.5-37.5 x 
8.6-17.8 cm, elliptico-ovata, apice abrupte 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 
in foliorum superiorem axillis oppositis plerum- 
que 2-4; bracteae liberae; bracteae exteriores 1.7- 
2.3 x 1.5-1.7 cm late ovatae; bracteae interiores 
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, poris 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- 
ternodes + 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-S5 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, 11-14 mm longand | 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, 7. 
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 7. calophylla can become an epi- 
phyte of massive proportions comparable to Jo- 
pobea durandiana 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 TJopobea 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 7. 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 


AtMEDA, F. 1974. A new epiphytic Blakea (Melastomata- 
ceae) from Panama. Brittonia 26:393-97. 

. 1980. Central American novelties in the genus Blak- 
ea (Melastomataceae). Rhodora 82:609-15. 

CoGntAux, C. A. 1891. Melastomaceae. Pages 1-1256. In 
A. and C. de Candolle, eds., Monographiae phanerogama- 
rum, vol. 7. G. Masson, Paris. 

Gieason, H. A. 1958. Melastomataceae. Jn R. E. Woodson, 
Jr. and R. W. Schery, Flora of Panama. Ann. Missouri Bot. 
Gard. 45:203-304. 

WurbDACcK, J. J. 1973. Melastomataceae. Jn 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. 


5 


Biereniber 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 


Asstract: 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 (Ginther, 
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 


Ogooué 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 


garita. Our studies, however, indicate that it is 
a valid species, albeit a close relative of D. mar- 
garita and the previously undescribed D. mar- 
garitella. Dasyatis garouaensis 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 fiir Seefischerei, Hamburg 
(ISH); Muséum National d’Histoire Naturelle, 
Paris (MNHN); Musée 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 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


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- 
born 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 DAsyATipAE 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 


a 


Ficure 1. 


b 


£® & 8 


285 


Disc shape in West African Dasyatidae. (a) Dasyatis violacea (trapezoidal); (b) Dasyatis centroura (diamond- 


shaped); (c) Taeniura grabata (circular); (d) Urogymnus asperrimus (oval). 


la. 


1b. 


2a. 


2D. 


3a. 


Sb: 
4a. 


4b. 


Sa. 


Sb: 


Key TO DASYATIDAE OF WEST AFRICA 


Disc oval (Fig. 1d); tail without dermal 
folds; sting invariably absent 
Lee 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) 00... 2 


Ventral tail fold extending to end of tail; 
dise circular (Fig) lc) 22 2 Taeniura 
grabata (E. Geoffroy Saint-Hilaire, 1817) 
Ventral tail fold ending far anterior to 
ena Ontall(Dasyalis) 3 


Disc trapezoidal or diamond-shaped (Fig. 
la—b) 


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 
RNase D. violacea (Bonaparte, 1832) 
Disc diamond-shaped, anterior margin 
angular, snout projecting as an angular 
lobe from disc (Fig. 1b); lower surface of 
GNIS Calo Mn (serene rs Me oes ee tS S 


Upper surface of disc with a dark retic- 
ular pattern; ventral tailfold very short, 
about twice length of sting 0. 
Parent D. marmorata (Steindachner, 1892) 
Upper surface of disc plain; ventral tail- 
fold long, much more than twice sting 
lene Ghee Rites reste ns FRE co 6 


6a. 


6b. 


Ta. 


7b. 


8a. 


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 (Giinther, 1870) 


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 


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 
fear tip) Of pectoral fins eee eens 
pare tenors eevee. D. pastinaca (Linnaeus, 1758) 


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 


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) 


286 


8b. 


9a. 


9b. 


10a. 


10b. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


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. centroura (Mitchill, 1815) 


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 
Sa ess D. ukpam (Smith, 1863) 
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: ARs. ae EG Vee EC) 10 


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- 
born specimens) and 27-32% of disc 
width; disc very flat, thickness at scap- 
ulocoracoid 8-11% (usually less than 
11%) of disc width; lateral prepelvic pro- 
cesses of pelvic girdle greatly expanded 
asl G oe leathers iS ee rs aor ee, D. garouaensis 
(Stauch and Blanc, 1962) 

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 19-26% of disc width; 
disc thicker, 11-15% of disc width over 
scapulocoracoid; lateral prepelvic pro- 
cesses slightly expanded) = 32s 11 


lla. 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 
(i00 sere D. margarita (Giinther, 1870) 
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 
i oA eles D. margaritella new species 


Iti: 


Dasyatis Rafinesque, 1810 
Dasyatis RAFINESQUE, 1810:16 (type-species Dasyatis ujo Ra- 
finesque, 1810 [=Raja pastinaca 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 Miller and 
Henle, 1837 in general morphology, heavy den- 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 


287 


FiGure 2. 
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 Miiller 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. 


(a) Dasyatis margarita, lectotype, 200-mm immature female, West Africa (BMNH 1865.7.4:1); (b) Dasyatis 


Dasyatis margarita (Giinther, 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 D. margarita; 
designated lectotype below); USNM 222589, 130-mm late fetal 
or newborn male, Sierra Leone; BMNH 1930.3.24:3, 212-mm 
immature male, Accra, Ghana; BMNH 1936.8.20:2-3, 216- 
mm and 315-mm immature females, Lagos, Nigeria, BMNH 
1899.2.20:35, 206-mm immature female, Banana, Congo Riv- 
er mouth, Zaire; AMNH 40408, 235-mm female, Angola. 

LectotyPeE DESIGNATION. —In the original description of D. 
margarita Giinther (1870:479) listed two specimens from West 
Africa without indicating either as holotype: 


a. Disk 81 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 


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COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 289 


CONTINUED 


TABLE 1. 


D. ukpam 


D. margaritella 


D. margarita 


Ogooué R. 


D. garouaensis 


Non-types 
27:109-262 


Types 


3:170-191 


Non-types 
5:130-315 


Zaire R. 


Old Calabar 3:499-650 


Lagos 


Cross R. 


Benue R. 
8:68-340 mm 


Lectotype 
200 mm 


266 mm mm 360 mm 


255 mm 


342 mm 


mm 


mm 


64 
22 


65-70 
17-19 


65 
59-64 


3p) 


56 
18 


51 


50-51 49-50 53-56 


55-56 


17-21 
59-60 


58 


18 


Propterygial radials 


20 


17 
a2 
23 


16-20 
49-52 


15-22 
51-56 
20 


16-19 
50-54 
22-24 


Mesopterygial radials 


58 


61 


59 


Metapterygial radials 


23 


Pelvic radials male 


Det 


28-34 


29 
149 


29 
157-162 


27 
128-131 


28 


Pelvic radials female 


155(1) 


122 


162 


139-151 


162 


Total vertebral segments 
Total separate centra 


108-120 


126 


120 


119-128 


116-124 


130 


38 


40 


31 


33-35 


34 


Ist synarcual segments 


2-5 


3-7 


0-2 
20-26 
37-42 


83-88 


2-4 


Ist synarcual free centra 
Intersynarcual centra 


113} 
41 


18 
40 


2nd synarcual centra 


39 


44-50 
57-65 


45 


40-42 


42-44 
63-66 


45 


2nd synarcual + monosp. 


Diplospondylous 
Post-sting centra 


72 


81 


78 


61-77 


V1 


11-17 


6-11 


9-13 


spine; specimen b apparently is BMNH 1852.8.30:23, a 116- 
mm fetal or newborn male with two relatively small oval pearl 
spines. Giinther’s description clearly is based mainly on spec- 
imen a, which we therefore designate as lectotype. Specimen 
b belongs to D. margaritella. 


DiAGcnosis.—A large marine and estuarine 
Dasyatis, 65 cm or more, most closely related to 
the much smaller marine and estuarine species 
D. margaritella and to the extremely flat fresh- 
water species D. garouaensis. Pearl spines always 
present and very large (much smaller in D. mar- 
garitella and absent or represented by 2-3 low 
denticles in D. garouaensis). Denticles on dorsal 
surface of disc smooth, restricted to middle por- 
tion. 

Disc oval, moderately flat, its medial lobe 
broad-based and moderately exserted, its an- 
terolateral margin somewhat concave; disc depth 
13.0-14.5% of disc width, disc width 0.9-1.1 
times disc length; snout moderately long, preoral 
length 22-24% of disc width; dorsal surface of 
disc naked in young (except for pearl spine), but 
in older individuals (larger than about 200 mm) 
middle third of disc covered with small, circular, 
flat denticles and tail covered with small prickles: 
no enlarged thorns on disc or tail; a massive, 
usually circular, button-shaped pearl spine on 
back, 4.8-5.5 mm long; usually a single large 
slender sting on tail, its length 22% of disc width 
in newborn; eyes moderately large and somewhat 
elevated, eyeball length 1.4-2.3 times in inter- 
orbital space, interorbital space 1.5-2.0 in preor- 
bital length; spiracles moderately large and flat- 
tened; floor of mouth with 5 elongate papillae; 
total tooth rows 26-30/31-34; pelvic fins short, 
anterior margins 18-21% of disc width; tail evenly 
tapering to a slender whiplash behind sting, but 
broad opposite and in front of it, its length when 
intact about 1.8—2.5 times disc width; base of tail 
horizontally oval and depressed in section; ven- 
tral tailfold moderately high, dorsal tailfold re- 
duced to a low keel behind sting; disc and pelvic 
fins gray-brown above, without spots or prom- 
inent markings, uniformly pale or whitish below; 
intestinal valve turns 12-14; total pectoral ra- 
dials 133-135; total vertebral centra 130, and 
total vertebral segments 162: vertebral centra ex- 
tending behind origin of sting; a moderately large 
stingray, exceeding 65 cm. 

Proportional measurements (as percent of disc 
width) and counts are given in Table 1. Preoral 
length 2.5-3.0 times internarial width and about 
1.0 times width between first gill slits. Snout fair- 


290 


ly broad, angle in front of eyes 123°. Spiracle 
length 1.0-1.3 times eyeball length, 0.8-0.9 times 
internarial width, and 1.8—2.0 times in distance 
between fifth gill slits. Nasal curtain with a fringed, 
weakly trilobate posterior margin. Mouth weakly 
arched, midline of lower jaw with a prominent 
indentation; a shallow to deep, curved groove 
extending posteriorly from posterior nasal flap 
around corners of mouth. Floor of mouth with 
5 papillae, including a transverse row of one small 
medial and two large elongate paramedial pa- 
pillae, and two smaller lateral papillae near ends 
of dental bands. 

Teeth in quincunx pavement, with crowns 
closely adjacent to one another. Teeth similar in 
upper and lower jaws, upper and lower jaw teeth 
about equally large at symphysis, varying con- 
tinuously lateral to synphysis, largest and most 
elongate at or near symphysis and smallest and 
least elongate at mouth corners. Upper dental 
band with a prominent knob of considerably en- 
larged teeth at symphysis, separated from simi- 
lar, very large knobs at either side by depressions 
with smaller teeth; lower dental band with a well- 
marked symphyseal depression into which sym- 
physeal knob of upper jaw fits, and a pair of 
prominent lateral knobs fitting into depressions 
in upper dental band. 

Dorsal surface naked except for pearl spine in 
young below about 200 mm, but in larger spec- 
imens moderately large, heart-shaped or circular 
denticles form a middorsal belt of denticles on 
disc; lateral parts of disc naked. Larger individ- 
uals with small conical prickles on dorsal surface 
of tail behind sting. 

Neurocranium, observed in radiographs, sim- 
ilar to that of Himantura signifer as described 
by Compagno and Roberts (1982) but with long- 
er, more ovate nasal capsules. 

Pelvic girdle (Fig. 10e) broadly arched, semi- 
circular, and relatively narrow, with a medial 
angle but without a medial prepubic process. Lat- 
eral prepubic processes low, rounded, lobate, and 
not greatly expanded; iliac processes well devel- 
oped; ischial processes short. Four obturator fo- 
ramina present. 

Claspers not studied in detail, but morpholog- 
ically similar to those of D. garouaensis. 


Dasyatis margaritella, new species 
(Figures 2b, 3) 


Trygon margarita GUNTHER, 1870:479 (in part). 
Dasyatis margarita BLACHE ET AL., 1970:53, fig. 116. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


Ho.otyre.—CAS-SU 68915, 190-mm male, Mbode, Cam- 
eroun, A. I. Good, 7 December 1940. 

ParatyPes. —CAS-SU 68916, 169-mm male and 180-mm 
female, same collection data as holotype. 

ADDITIONAL MATERIAL EXAMINED But Not INCLUDED IN 
Type-SERIES.—BMNH 1852.8.30.23, 116-mm immature male, 
West Africa, apparent syntype of 7rygon margarita Giinther, 
1870; USNM 222590, 4 immature males, 109-139 mm, 2 
females, 132-164 mm, and USNM 222593, 3 immature males, 
129-141 mm, 146 mm female, both lots from Guinea-Bissau; 
ISH 141-142/62, 3 immature males, 135-196 mm, 201-mm 
adolescent male, 228-mm adult male, 3 females, 149-222 mm, 
Guinea; ISH 183/63, 226-mm adult male and 248-mm female, 
Conakry, Guinea; ISH 295/59, 167-mm immature male, Du- 
breka, Guinea; USNM 222591, 145-mm female, Liberia; 
BMNH 1920.8.12.1, 262-mm female, Sierra Leone; USNM 
222591, 149-mm female, Liberia; BMNH 1914.11.2.75, 133- 
mm female, Lagos, Nigeria; BMNH 1937.4.19.4, 208-mm adult 
male, Lagos Lagoon, Nigeria, AMNH 41515, 204-mm female, 
Congo River mouth. 


DERIVATION OF NAME.—margaritella, dimin- 
utive of Latin margarita, f. “pearl”; for the smaller 
size of this stingray and of its pearl spine, as 
compared to D. margarita. 

DiaGnosis.—D. margaritella is a marine 
species and the smallest dasyatid in West Africa. 
Males sexually mature at 208 mm or less; largest 
known specimen a 262-mm female. Most similar 
to the marine species D. margarita, which attains 
over 600 mm, has a larger pearl spine, and more 
numerous pectoral radials (133-135 vs. 116-127). 
It is also similar to D. garouaensis, a freshwater 
species with a much flatter disc and a longer 
snout. 

Disc oval, moderately flat. Medial lobe broad- 
based and exserted, its anterior margin concave. 
Disc depth 11.0-15.5% of disc width, disc width 
0.9-1.1 times disc length; snout moderately long, 
preoral length 25-27% of disc width; dorsal sur- 
face of disc naked in young (except for pearl spine), 
but above about 130-140 mm middle third of 
disc covered with small heart-shaped or circular 
flat denticles and tail posterior to sting with small 
prickles; no enlarged thorns on disc or tail; a 
moderately large, usually longitudinally oval pearl 
spine on back, 2.4—4.1 mm long; usually a single 
large slender sting on tail, 24-32% of disc width; 
eyes moderately large and somewhat elevated, 
eyeball length 1.2—1.9 times in interorbital space, 
interorbital space 1.7—2.4 in preorbital length; 
spiracles moderately large and flattened; floor of 
mouth with 5 elongated papillae; total tooth rows 
35—43/38-50; pelvic fins short, anterior margins 
13-22% of disc width; tail evenly tapering to a 
slender whiplash behind sting, but broad oppo- 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 291 


FiGureE 3. 


site and in front of it, its length from vent to tip 
when intact about 2.5—3.4 times disc width; base 
of tail horizontally oval and depressed in section; 
a moderately high ventral tailfold present but 
with only a low keel on the dorsal surface of the 
tail behind the sting; disc and pelvic fins gray- 
brown above, without spots or prominent mark- 
ings, white below and without a marginal dark 
band; intestinal valve turns 13-14; total pectoral 
radials 116-127; total vertebral centra 116-128, 
and total vertebral segments 139-151; vertebral 
centra extending behind origin of sting. A small 
stingray, probably not exceeding 30 cm. 
Proportional measurements (as percent of disc 
width) and counts are given in Table 1. Preoral 
length 2.5-3.3 times internarial width and 1.1- 
1.3 times width between first gill openings. Snout 
fairly narrow, angle in front of eyes 113-119°. 
Spiracle length 0.7—1.1 times eyeball length, 0.6- 
0.9 times internarial width, and 1.8—2.5 times in 
distance between fifth gill openings. Nasal cur- 
tain with a fringed, nearly straight or weakly tri- 
lobate posterior margin. Mouth weakly arched, 
midline of lower jaw with a prominent inden- 
tation; a shallow-to-deep, curved groove extend- 
ing posteriorly from posterior nasal flap around 


Dasyatis margaritella, holotype, 190-mm immature male, Mbode, Cameroun (CAS-SU 68915). 


corners of mouth. Skin on ventral surface of low- 
er jaw more or less corrugated and papillate. Pal- 
ate behind fringed maxillary valve with three 
strong, short ridges, a medial longitudinal ridge 
and a pair of diagonal lateral ridges. Floor of 
mouth with 5 oral papillae, including a trans- 
verse row of one smaller medial and two larger 
elongated paramedial papillae at midline of 
mouth, and two smaller lateral papillae near ends 
of dental bands. 

Upper jaw with about 6, lower jaw with about 
8 functional series of teeth. Teeth in quincunx 
pavement, close-set, with crowns closely adja- 
cent to one another. Teeth similar in upper and 
lower jaws; uppers about as large as lowers at 
symphysis, varying continuously lateral to sym- 
physis; teeth largest and longest relative to width 
at or near symphysis and smallest and shortest 
at mouth corners. Upper dental band with a 
prominent knob of slightly enlarged teeth at sym- 
physis, separated from similar knobs at either 
side by depressions with smaller teeth; lower 
dental band with a well-marked symphyseal 
depression into which symphyseal knob of upper 
jaw fits, and a pair of prominent lateral knobs 
that fit into depressions in upper dental band. 


292 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


Ficure 4. Dasyatis garouaensis, 340-mm mature male, Benue River at Makurdi, Nigeria. 


Teeth of both jaws with low, broad-tipped, cusp- 
less, rhomboidal crowns that have a transverse 
blunt keel or carina, reticulating ridges on their 
labial sides, a strong basal ledge, and concave, 
inset basal groove separating crown and root. 
Roots moderately large, fairly low, pedicellate, 
bilobed basally, with a transverse groove and 
nutrient foramen. Teeth of adult males some- 
what longer than in females, but without elon- 
gated cusps. 

Dorsal surface naked in young below about 
130-140 mm, except for pearl spine, but in larger 
specimens moderately large, heart-shaped or cir- 
cular denticles form a middorsal belt of denticles 
on disc; lateral parts of disc naked. Also, small 
conical prickles on dorsal surface of tail behind 
sting in larger individuals. 

Neurocranium, observed on radiographs, sim- 
ilar to that of Himantura signifer as described in 
Compagno and Roberts (1982), but with longer, 
more ovate nasal capsules. 

Pelvic girdle (Fig. 10f) broadly arched, semi- 
circular, relatively narrow, with a medial angle 
but without a medial prepubic process. Lateral 
prepubic processes low, rounded, lobate, not 


greatly expanded; iliac processes well developed; 
ischial processes short. Four obturator foramina 
present. 

Claspers not studied in detail, but similar mor- 
phologically to those of Dasyatis garouaensis. 


Dasyatis garouaensis (Stauch and Blanc, 1962) 
(Figures 4-5) 


Potamotrygon garouaensis STAUCH AND BLANC, 1962:166, fig. 
1-4 (type-locality Benue River at Malape, Cameroun); Da- 
GET AND STAuUCH, 1963:85-107 (reference); CAsTEXx, 1967: 
167-176 (discussion). 

Dasyatis sp. KreFFr, 1968:70, pl. 6 (““Rafin Kunama, Neben- 
fluss des Nigers etwa 300 Meilen oberhalb der Miindung”’). 

Dasyatis garouaensis CASTELLO, 1973:67 (placed in Dasyatis); 
THORSON AND Watson, 1975:701—712 (placed in Dasyatis, 
size, range, physiology, additional specimens reported from 
Niger and Benue rivers of Nigeria and Cameroun); REID AND 
SYDENHAM, 1979:46, 54-55 (possibly synonymy with D. 
margarita, range in Benue River system); COMPAGNO AND 
Roserts, 1982:321 (reference). 


MATERIAL EXAMINED.—MNHN 1962-303, holotype, 202- 
mm female; MNHN 1962-304, 68-mm male and 78-mm fe- 
male fetuses, MNHN 1967-441, 263-mm female; MNHN 1967- 
440, 300-mm adult male; MNHN 1967-439, 340-mm adult 
male; all from upper Benue River, Cameroun; CAS 49147, 
342-mm adult female, Cross River, 5—10 km downstream from 
Mamfe, Cameroun; CAS 53108, 311-mm female, 340-mm 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 293 


adult male; Thorson collection, uncat., 238-mm female (ra- 
diograph only); Benue River at Makurdi, Nigeria; BMNH 
1949.10.24.1, 255-mm adult male, Lagos, Nigeria. 


DiaGnosis.—A moderately large and thin- 
bodied freshwater dasyatid. Largest known spec- 
imen a 342-mm adult female. Disc flatter than 
in any other West African dasyatid, its depth 
only 8.7—11.0% of disc width (vs. 11.0% or more 
in other species). Denticles on dorsal surface of 
disc highly variable, sometimes absent, always 
restricted to central portion. Pearl spine absent 
or represented by 2-3 low denticles. Most similar 
to the small marine species D. margaritella and 
the larger marine species D. margarita, both of 
which have well-developed pearl spines and 
shorter snouts. 

Disc oval, very flat. Medial lobe fairly broad- 
based and elongated, its anterior margin straight 
or concave; disc width 0.9-1.0 times disc length; 
snout moderately long, preoral length 29-32% of 
disc width; dorsal surface of disc either entirely 
naked or with scattered, small, heart-shaped or 
circular flat denticles on its middle, no pearl spines 
(a few slightly enlarged, flattened, heart-shaped 
denticles occasionally present in midscapular 
area), and no enlarged conical denticles on disc 
and tail; a single, large, slender sting present on 
tail; sting 22.6-26.3% of disc width in adults; 
eyes small and hardly elevated, eyeball lengths 
1.3-2.0 times in interorbital space; interorbital 
space, 2.4-3.2 times in preorbital length; spira- 
cles moderately large and flattened; floor of mouth 
with 5 elongated papillae; total tooth rows 32- 
40/37-45; pelvic fins short, their anterior mar- 
gins 20-25% of disc width; tail rapidly tapering 
to a slender whiplash behind sting but broad op- 
posite and in front of it, its length from vent 2.5— 
3.5 times disc width when intact; base of tail 
horizontally oval and depressed in section; a 
moderately high ventral tailfold present but with 
only a low keel on dorsal surface of tail behind 
sting; disc and pelvic fins medium gray or gray- 
brown above, without spots or prominent mark- 
ings, white below and without dark margins; tail 
darker or blackish, mottled, lighter below, un- 
derside of base white; intestinal valve turns 10; 
total pectoral radials 122-125; total vertebral 
centra 120-131 and total segments about 149- 
162; vertebral centra extending posterior to sting 
origin. 

Proportional measurements (as percent of disc 
width) and counts are given in Table 1. Preoral 


length 4.0-6.0 times internarial width and 1.2- 
1.9 times width between first gill slits. Snout 
moderately broad, angle in front of eyes 110- 
116°. Spiracle length 0.9-1.2 times eyeball length, 
0.8—1.2 times internarial width and 1.8-2.1 times 
in distance between fifth gill slits. Internal gill 
openings with close-set transverse ridges on gill 
arches, apparently serving as gill rakers. Nasal 
curtain with a fringed, concave posterior margin. 
Mouth nearly straight, midline of lower jaw 
slightly concave. A groove extending posteriorly 
from posterior nasal flap around. mouth corner. 
Skin on anteroventral surface of lower jaw cor- 
rugated and papillate. Palate behind heavily 
fringed maxillary valve with three strong short 
ridges, a medial longitudinal ridge and a pair of 
diagonal lateral ridges; two short ridges also pres- 
ent behind the three palatine ridges. Floor of 
mouth with a transverse row of three elongated 
oral papillae, a larger medial papilla and a small- 
er lateral one behind each end of dental band; a 
pair of additional large papillae just behind and 
lateral to medial papilla. Upper jaw with 5-7 and 
lower jaw with 7-9 functional tooth series. Teeth 
in quincunx pavement, close-set, with crowns 
adjacent to one another. Teeth similar in upper 
and lower jaws, uppers slightly larger than low- 
ers; teeth varying continuously lateral to sym- 
physis, largest and longest relative to width at or 
near symphysis, smallest and shortest near mouth 
corners. Upper dental band with a small knob 
of slightly enlarged teeth at symphysis, separated 
from similar weak knobs on either side by 
depressions with smaller teeth; lower dental band 
with a corresponding symphyseal depression into 
which upper symphyseal knob fits, and a pair of 
very low lateral knobs which fit into upper par- 
asymphyseal depressions. Teeth of both jaws with 
moderately high (females) to very high (males), 
peaked, broad-tipped (females) or acutely point- 
ed, cuspidate (males) crowns shaped like mush- 
room caps, with a strong, sharp cutting edge 
(males) or a broad, blunt carina or transverse 
keel (females), a strong basal ledge, and a con- 
cave, inset basal groove separating crown from 
root. Root small, moderately high, pedicellate, 
bilobed basally, with a transverse groove and 
nutrient foramen. Dentition sexually dimorphic; 
teeth of males with triangular, concave, cuspi- 
date crowns; females with low, rhomboidal, 
truncated, cuspless crowns. 

Dorsal surface of disc either completely naked, 
without denticles (2 fetuses, the holotype, and a 


294 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


FiGur_e 5. 


340-mm adult male), or with a variable number 
of small, heart-shaped or circular denticles bur- 
ied in skin on middorsal area (remainder of spec- 
imens examined). Two large specimens (includ- 
ing 342-mm male from Mamfe) have a few 
scattered denticles along midline of back; two 
others have a small rectangular area of flat den- 
ticles centered on midscapular region but not 
extending outwards to cover middle third of disc 
as in D. ukpam, D. margarita, and D. marga- 
ritella. Five large specimens had 3 or 4 large, 
flattened, wedge-shaped denticles in midscapular 
region, but these not formed as domed, rounded 
pearl spines (although probably homologous to 
pearl spines). 

Neurocranium, examined from radiographs, 
generally similar to that of Himantura signifer 
as described in Compagno and Roberts (1982), 
except frontoparietal fontanelle shorter and 
broader, and posterior margin of nasal capsules 
more transverse; width of nasal capsules about 
75% of nasobasal cranial length. 

Pelvic girdle (Fig. 10c—d) less arched than that 
of D. ukpam, convex anteriorly and without a 
medial prepubic process or a median angle. Pel- 


Dasyatis garouaensis, 342-mm mature female, Cross River near Mamfe, Cameroun (CAS 49147). 


vic girdle of D. garouaensis differs from that of 
D. ukpam as well as D. margarita and D. mar- 
garitella in having large, laterally expanded lat- 
eral prepelvic processes; these were prominent 
on all D. garouaensis radiographed, including 
specimens from Mamfe, Lagos, and Benue River 
in Nigeria and Cameroun. Pelvic girdle also with 
short ischial processes, long, slender iliac pro- 
cesses, and 4 obturator foramina. 

Claspers of adult male short and stout, length 
of outer margin 10.2—11.6% of disc width, oval 
in cross section and somewhat depressed; height 
about % of width at midlength. Dorsal surface of 
clasper slightly flattened, ventral surface broadly 
convex, lateral edge convexly arched, medial edge 
undulated, and tip bluntly pointed. Apopyle on 
anterodorsal surface, connected to hypopyle by 
an open, posteriorly curved clasper groove. 
Clasper glans simple, dorsal lobe supported by 
dorsal marginal and terminal cartilages, ventral 
lobe supported by ventral marginal and terminal 
cartilages as well as ventral covering piece. No 
structures inside hypopyle. A large pseudopera 
laterally situated on ventral lobe below hypopyle; 
a small pseudosiphon on dorsomedial surface of 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 295 


dorsal lobe, its cavity lying entirely below flange 
of dorsal marginal cartilage. Ventral lobe without 
clasper spur or associated terminal (T, cartilage). 
Clasper skeleton simple, with two cylindrical 
basal segments connecting pelvic basipterygium 
to axial cartilage. Axial cartilage cylindrical, 
nearly straight, with rodlike tip reaching ends of 
terminal cartilages. Beta cartilage present, a long, 
slender, separate, flattened plate running along 
lateral surface of clasper skeleton from posterior 
end of basipterygium to anterior end of dorsal 
marginal cartilage. Dorsal marginal cartilage 
broad and subquadratetriangular, with a diago- 
nally truncate posterior edge, broad medial flange, 
and narrow lateral flange that forms roof of clasp- 
er groove. Ventral marginal cartilage a narrow, 
laterally expanded plate on axial cartilage, with 
a straight lateral margin forming floor of clasper 
groove. Dorsal terminal cartilage large, broad, 
wedge-shaped, and axially convex, with a broad 
anterior base articulating with posteroventral edge 
of dorsal terminal, a narrow posterior tip op- 
posite tip of axial cartilage, and medial edge ar- 
ticulating with axial cartilage. Ventral terminal 
cartilage large, complex, oval, and scoop-shaped, 
with a broad, arched, lateral flange forming roof 
of pseudopera and a recurved, expanded pos- 
teroventral tip forming a partial floor for it along 
with ventral covering piece. Ventral covering 
piece large, elongate-oval, broadly convex ven- 
trally, and scoop-shaped, enclosing terminal car- 
tilages and rear tips of marginals ventrally. 


Dasyatis ukpam (Smith, 1863) 

(Figures 6-9) 

Hemitrygon ukpam Smitn, 1863:69 (type-locality Old Calabar 
River, Nigeria); FowLer, 1936:126 (placed in synonymy of 
D. centroura),; Fowxer, 1969:186 (in synonymy of D. cen- 
troura). 

Trygon ukpam GUNTHER, 1870:480 (description after Smith, 
1863, placed in Trygon = Dasyatis). 

Dasyatis margarita LousBens, 1964:11 (freshwater lakes south 
of Lambarene district, Ogooué basin; presumably no spec- 
imens preserved). 

Dasyatis ukpam STEHMANN, 198 1:4 (in key to marine Dasyatis 
of West Africa); COMPAGNO AND Ropserts, 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, Ogooué River sys- 
tem, Gabon; CAS 42761, 650-mm immature female, Lake 
Ezanga near Nzame-Akesile village, Ogooué River system, Ga- 
bon; MNHN 1979-244, 499-mm immature female, Booué, 
Ogooué River, Gabon; MRAC 55778, 361-mm immature fe- 
male, Binda, Congo (Zaire) River, Zaire. 


DiaGnosis.—D. 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, witha 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, 
ora single small, possibly vestigial one, only 6.5% 
of disc width in 520-mm specimen; eyes small 
and strongly elevated, eyeball 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 ongin. 

Proportional measurements (as percent of disc 
width) and counts are given in Table 1. Preoral 
length 2.3—2.6 times internarial width and 0.9- 
1.0 times width between first gill openings. Snout 
broad, angle in front of eyes 123-132°. Spiracle 


296 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


Ficure 6. Dasyatis ukpam, 266-mm fetal male syntype, Old Calabar, Nigeria (BMNH 1874.5.23:1). 


Ficure 7. Dasyatis ukpam, 650-mm immature female, Lake Ezanga, Ogooué basin, Gabon (CAS 42761). 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 297 


Ficure 8. Dasyatis ukpam, 520-mm immature female, Lake Ezanga, Ogooué basin, Gabon (USNM 219780). 


length 1.5—2.0 times eyeball length, 0.8—1.0 times 
internarial 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 corners of mouth. 
Skin on ventral surface of lower jaw more or less 


Ficure 9. Dasyatis ukpam, 361-mm immature female, lower Zaire River at Binda, Zaire (MRAC 55778). 


298 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


Ficure 10. 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 corners. 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- 
males. 

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 (361-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. 10a—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 IItis 1965; Blache et al. 1970; 
Stehmann 1981). 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 | kg 
while the other attains easily 15-20 kg and per- 
haps much more (quoted in Daget and IItis 1965: 
15). The small species is D. margaritella, the 


COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS 299 


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 crystalline 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 of 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 
lives as much as 150-miles upriver (Smith 1863), 
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 Ogooué basin was first brought to our atten- 
tion by an ichthyological colleague, Dr. Jaques 
Géry, 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 1964. The Ivindo flows into the Ogooué 


300 


near Booué, 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 100 km upriver 
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 AI- 
wyne Wheeler, Mandy Holloway, and Oliver 
Crimmen, British Museum (Natural History); D. 
F. Thys van den Audenaerde, Musée Royale de 
l’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. Fishes of the 
western North Atlantic. Sears Found. Mar. Res., Mem. No. 
1(2). 


BLACHE, J., J. CADENAT, AND A. StAucH. 1970. Clés de dé- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18 


termination des poissons de mer signalés dans |’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. Rosperts. 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):321-339, 
12 figs. 

Dacet, J., AND A. ILtis. 1965. Poissons de Céte d’Ivoire 
(eaux douces et saumatres). Mém. Inst. franc. Afr. noire no. 
74. 

DaGcET, J., AND A. StAucH. 1963. Poissons de la partie Ca- 
merounaise du bassin de la Benoué. Mém. Inst. franc. Af- 
rique noire 68:85-107. 

Fow ter, H.W. 1936. The marine fishes of West Africa. Bull. 
Amer. Mus. Nat. Hist. 70(1):1-606. 

. 1969. Acatalog of world fishes (XI). Quart. J. Taiwan 
Mus. 22(3-4):125-190. 

GUNTHER, A. W. 1870. Catalogue of the fishes in the British 
Museum. 8. 

KreFrt, G. 1968. Knorpelfische (Chondrichthyes) aus dem 
tropischen Ostatlantik. Atlantide Rep. 10:33-76, pls. 3-6. 
Lousens, G. 1964. Travaux en vue de développement de la 
péche dans le bassin inferieur de l’Ogooué. Publ. Cent. Tech. 

Forest. Trop., Nogent-sur-Marne (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 pls. 

Reip, 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. 

Staucnu, A., AND M. BLANnc. 1962. Description d’un sélacien 
rajiforme des eaux douces du Nord-Cameroun, Potamotry- 
gon garouaensis n. sp. Bull. Mus. Nat. Hist. Natr. (Paris) 
34(2):166-171. 

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. 

TuHorson, 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 94118 


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Vol. 43, No. 19, pp. 301-315, 12 figs., 1 table. 


PROCEEDINGS 
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CALIFORNIA ACADEMY OF SCIENCES 


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 of Fish and Game, 
Long Beach, California 90802 


Asstract: Four species of catalufas inhabit eastern Pacific waters: Cookeolus boops (Schneider, 1801), 
Heteropriacanthus cruentatus (Lacépéde, 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 cruentatus, 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. 


December 11, 1984 


INTRODUCTION 


For years, fishermen aboard long-range sport- 
fishing boats that operated out of San Diego 
sought only such large game species as yellowfin 
and bigeye tuna (Thunnus albacares and T. obe- 
sus), wahoo (Acanthocybium solanderi), yellow- 
tail (Seriola lalandi), giant sea bass (Stereolepis 
gigas) and several kinds of large serranids (Epi- 
nephelus spp. and Mycteroperca spp.). During 
1978, however, at the urging of California De- 


partment of Fish and Game (DFG) biologists, 
skippers and crew members of these vessels com- 
menced fishing for and saving miscellaneous 
small fishes caught at the Revillagigedo Islands, 
Alijos Rocks, and other fishing spots off southern 
Baja California. 

Among the first of these incidentally caught 
species turned over to DFG personnel were a 
half-dozen catalufas that appeared to represent 
three species of Priacanthus. A literature search 


[301] 


302 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


Imus 
a 


Loy 

XY » 
Armies 
macropame 


a RE RE RN NR a ae ba) 


Figure 1. Type specimen of Anthias macrophthalmus Bloch, 1792 (ZMB 8156). Photo courtesy of Hans-J. Paepke, Museum 


fiir 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 fiir 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 Starnes (pers. comm.), 
to whom we sent a copy of the photograph, has 
confirmed that Bloch’s Anthias macrophthalmus 
is conspecific with Sciaena hamrur Forsskal, 
ld: 


Key TO EASTERN PAcIFIC PRIACANTHIDAE 


la. Pored scales in lateral line 35 to 40; dorsal 
soft rays 11 (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 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 


1b. 


2a. 


2b. 


Ficure 2. Pseudopriacanthus serrula. Photo by Jack W. Schott. 


under anal fin base; ctenii on scales in par- 
allel species, presenting a corrugated or 


12 (rarely) or more dorsal soft rays; 12 or 
more anal rays; dorsal and ventral profiles 
tapering gradually to caudal peduncle; 
ctenii on scales not as above 
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 
caudal fins yellow, edged with black; ctenii 
thick and bristlelike, in rows 


ea _........ Cookeolus boops 


Eight to 10 scale rows between 8th dorsal 
spine and lateral line; fewer than 15 scale 
rows on dorsum of caudal peduncle; anal 
rays 14; peduncle depth fits about 14 times 
into horizontal eye diameter; dorsal, anal 


3a. 


3b. 


and caudal fins never with yellow or tipped 
with black; ctenii thin and filamentous ... 
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- 
opercie completely, scaled] eas 


pete mers EBA be 9 Wy s) Uiee 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- 


303 


304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


amentous, only on scale margin, sparse; 
posterior margin of preopercle without 
SCALES) 22 ace ak it ede eee 


SPECIES ACCOUNTS 


Pseudopriacanthus serrula (Gilbert, 1891) 
(Figure 2) 


DIAGNOSTIC CHARACTERS.—35 to 40 pored 
scales in lateral line; 10 to 12 (typically 11) dorsal 
soft rays; 10 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,10-12; A. THI,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 11 dorsal soft rays, one had 10, 
and one had 12. 

MAximMumM SizeE.—The largest individual we 
observed was a female 274 mm SL (353 mm TL) 
that weighed 1300 g. This was one of 24 indi- 
viduals caught in gill nets in “30-40 fm” (55- 
73 m) off Magadalena Bay, Baja California, 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 13 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. 11) 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 Californian 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. — 18 to 20 scale rows 
between eighth dorsal spine and lateral line; 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. III,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 SizeE.—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- 
ern 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 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 


Ficure 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 (1.9 by 2.4); d) Pseudopriacanthus 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 


Ficure 4. Cookeolus boops. Photo by Lee Stockland. 


PERCENT OF PELVIC FIN LENGTH 


STANDARD LENGTH 


STANDARD LENGTH (MM) 


160 180 200 220 240 260 280 300 320 340 360 380 400 


Ficure 5. Pelvic fin length (as percent of SL) plotted against SL for 32 specimens of Cookeolus boops. 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 307 


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 Priacanthus 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 Ist 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. 
2 See Figure 5. 


308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


FIGURE 7. 


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, 1904 
(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- 


Priacanthus alalaua. Photo by Lee Stockland. 


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). 

Me_ristic Data.—D. X,13; A. III,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 19 elements in the pectoral fin; the rest 
had 18. 

MAXIMUM Size.—The longest individual we 
observed was a female 261 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 14 


‘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- 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 309 


Ficure 8. Otoliths (sagittae) of eastern Pacific Priacanthidae (otolith length and fish SL, in mm, in parens): a) Pseudo- 
priacanthus serrula (7.2 and 2.9); b) Heteropriacanthus cruentatus (5.3 and 195); c and d) Cookeolus boops (7.6 and 257; 5.5 
and 180); e) Priacanthus alalaua (3.6 and 221). Photos a through d by Jack W. Schott; SEM photo e by Brian White. 


casion, it has been taken mostly atnightindepths ‘‘A. III,13”) were reported by Fitch and Schultz 
of “5 to 25 fm or more” (9-46 m). (1978) along with data from seven cruentatus 

REMARKS. — Although three individuals of P. taken at the same time and place. Fortunately, 
alalaua were taken in gill nets set overnight just their unique otoliths had been removed and 
upcoast from Braithwaite Bay, Socorro Island, saved, and counts and measurements were made 
in April 1955, they were misidentified as P. before they were skeletonized, so subsequent 
cruentatus, and meristic data from them (e.g., identification as P. alalaua was easily verified. 


310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


Ficure 9. Gas bladder (124 mm total length) from Pria- 
canthus alalaua 220 mm SL. Drawing by Mary Butler. 


We examined the entire collection (worldwide) 
of priacanthids at Los Angeles County Museum 
of Natural History and found that arenatus, 
hamrur, macracanthus, meeki, and tayenus pos- 
sessed similar peculiar gas bladders and otoliths. 
Therefore, we believe these species should be 
assigned to Priacanthus. 


Heteropriacanthus new genus 
Type-spEcies. — Priacanthus cruentatus (Lacépéde, 1801). 


DiaGnosis.—Second dorsal, anal and caudal 
fins with rust-colored spotting (dusky in forma- 
lin-preserved specimens); pelvic fin membranes 
lightly pigmented to clear; gas bladder thin- 
walled, contained entirely within the body cav- 
ity; otoliths (sagittae) oval in outline with normal 
anteroventral rostrum (Fig. 8b); ctenii filamen- 
tous, sparse, and only on scale margin (Fig. 3c); 
posterior margin of preopercle scaleless. 

EtyMoLoGy.— From Greek heteros (different), 
alluding to its being different from other Pria- 
canthus. 


Heteropriacanthus cruentatus (Lacépéde, 1801) 
(Figure 10) 

DIAGNOsTIC CHARACTERS. —As for the genus. 

Meristic Data.—D. X,12-13; A. HI,14—-15; 
P. 17-19; GR 5-6 + 17-20 = 22-25; pored lat- 
eral line scales 57-65; vertebrae 10 + 13 = 23. 
Only two of the 61 specimens we examined for 
meristic data had 12 dorsal soft rays; the rest had 
13. One specimen had an anal count of V,13 
(LACM 32283), obviously a freak, so it was not 
included. Of the remaining 60 specimens, two 
had counts of III,15 and the rest III,14. All but 
eight specimens had 18 total elements in their 
pectoral fins; three had 17 and five had 19. 

Maximum SizeE.—The largest individual we 
observed (sex undetermined) from the eastern 
Pacific was 247 mm SL (315 mm TL). This fish 
(SIO 70-136) was from the Gulf of Chinqui, Pan- 
ama. We did not obtain a weight for it or for any 
other large H. cruentatus. 

RANGE. — Worldwide in tropical and subtrop- 


ical seas. In the eastern Pacific, H. cruentatus 
ranges from Guadalupe Island, Baja California 
(SIO 60-18) to the Galapagos Islands, Ecuador 
(numerous West Coast collections) in depths of 
*2 to 15 fm” (3.6—27 m) at least. Within this 
range, it has been collected at all of the Revil- 
lagigedo Islands, Hurricane Bank, the Tres Mari- 
as Islands (Mexico), Cocos Island (Costa Rica), 
and many islands off Panama. Its mainland dis- 
tribution is not so extensive, ranging from Cape 
San Lucas, Baja California (UCLA-W52-259) to 
Panama (SIO 70-140). 

REMARKS.— Typically an overall reddish or 
crimson when alive, H. cruentatus often will de- 
velop silvery marbling or blotching on the sides 
and back. Thomson et al. (1979) suggest that this 
color pattern results from stress. 

Although we examined specimens of H. cruen- 
tatus from several localities far removed from 
the eastern Pacific, we were unable to find any 
differences that could be considered of specific, 
or even subspecific, magnitude. 


FossiL RECORD 


Fossil priacanthids have been reported only 
from Europe and only from Eocene deposits. 
These fossil remains have consisted of skeletal 
bones and impressions from Italy (Pristigenys 
substriata: see Fritzsche and Johnson 1981), oto- 
liths from Belgium and France (Pristigenys rutoti 
and P. caduca: Stinton and Nolf 1970; Nolf 1973), 
and otoliths and dorsal fin spines from England 
(P. bella, P. spectabilis, and P. dentifer: Stinton 
1980). 

In identifying fossil fishes, an ideal situation 
would be to have a three-dimensional specimen 
with all bony elements and conventional char- 
acters present (e.g., viscera, gas bladder, scales, 
otoliths, etc.). Unfortunately, this does not hap- 
pen. Soft parts can only be inferred. Rarely are 
three-dimensional fossil fishes found, and even 
two-dimensional skeletal impressions are not all 
that common compared with isolated teeth, 
scales, otoliths, and bones (Schafer 1972). Ob- 
viously, the more complete the fossil specimen 
or specimens, the greater the likelihood of mak- 
ing a correct identification. 

The skeletal impressions of P. substriata from 
Italy generally have been in excellent condition 
and have permitted direct comparison of many 
salient features with the same features on extant 
priacanthids. None of these two-dimensional 
skeletal impressions has contained otoliths, how- 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 311 


Ficure 10. Heteropriacanthus cruentatus. Photo by Paul Gregory. 


ever; to associate isolated otoliths with the genus 
Pristigenys is pure speculation. 

In using otoliths (sagittae) as a taxonomic char- 
acter, the most important feature for determining 
family and/or generic relationships (including 
ancestry) is the sulcus or groove on the inner face: 
its position, configuration, proportions, angle of 
curvature, and other features. If features of the 
sulcus do not match exactly, the otolith cannot 
belong to the same genus as the one to which it 
is being compared. Overall otolith shape, ratio 
of height into length, marginal ornamentation 
and similar surface features are important pri- 
marily at species level. 


Fosstt ACCOUNTS 


Pristigenys substriata (Blainville, 1818) 
(Figure 11) 


White (1936) presents an excellent account of 
the taxonomy of this species as follows: 


Among the numerous fossil fishes described by Volta (1796 
...) from the lower Lutetian of ‘Monte Bolca’ was a small 
imperfect specimen which he identified with the living species 
Chaetodon Striatus, . . . illustrating his description with a figure 
that is unusually obscure; this specimen was later referred by 
de Blainville (1818 . . .) toa new species C. substriatus. Agassiz 
(1835 ...) recognizing that the fossil was not a Chaetodon, 


renamed it completely, Pristigenys macrophthalmus, and 
sketched a few of its more obvious characters, which led him 
to suppose that this was ‘un genre voisin de Beryx’; and under 
Agassiz’ name it was listed among the Berycidae by A. S. 
Woodward (1901 .. .); finally, Eastman (1905 . . .) added de- 
tails omitted by previous authors, and re-figured the unique 
original specimen. Eastman rightly named the species Pristi- 
genys substriatus (for while Agassiz’ genus is good, so is de 
Blainville’s species), and retained it in the Berycidae. 


White went on to report that the fossil was not 
a berycoid but a priacanthid, which, in his opin- 
ion, was identical with the extant Pseudopria- 
canthus Bleeker, 1869, and that this generic name, 
by reason of its later publication date, must be 
replaced by Pristigenys. 

White’s report appears to have been over- 
looked by subsequent authors until Myers (1958) 
called it to the attention of contemporary ich- 
thyologists. Subsequently, as discussed by 
Fritzsche and Johnson (1981), there has been 
considerable controversy as to the validity of 
placing Pseudopriacanthus in the synonymy of 
Pristigenys. In concurring with such synonymy, 
Fritzsche and Johnson point out that a “‘predor- 
sal bone” is a character shared only by Pristi- 
genys and Pseudopriacanthus among all priacan- 
thid genera. Interestingly, they support their 
contention of close relationship by presenting four 


312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


TMT 


Pristigenys substriata (BMNH P.16370) 155 mm SL. Photo courtesy of Colin Patterson, British Museum (Natural 


Ficure 11. 
History). 


sets of morphometric data and two sets of me- 
ristic characters (from Pristigenys substriata and 
Pseudopriacanthus altus), while one paragraph 
later they state that ‘““morphometric characters 
are not generally considered to be valid indica- 
tors of relationship” within the perciforms. 

After carefully examining all available char- 
acters for the fossil Pristigenys substriata and for 
three species of the extant Pseudopriacanthus (i.e., 
altus, niphonius, and serrula), itis our contention 
that both Pristigenys and Pseudopriacanthus are 
valid genera. While we agree that these two gen- 
era share at least one character that appears to 
be of generic magnitude (i.e., the ““‘predorsal bone” 
of Fritzsche and Johnson), there are other salient 
characters of equal magnitude that are shared 
with other priacanthid genera or are distinctive 
within their own genus. 

Based upon the associated Monte Bolca fish 
fauna, Pristigenys substriata was living in a pe- 
lagic environment. The pelagic environment and 


extremely long pelvic fins (Fig. 5) are shared with 
juvenile Cookeolus, and apparently all known 
specimens of P. substriata represent juveniles as 
none exceeds 155 mm SL. The scales of Pseu- 
dopriacanthus (Fig. 3d) are unique among pria- 
canthids for the shape and arrangement of ctenii; 
scales of Pristigenys appear to resemble those of 
Priacanthus or Heteropriacanthus (Colin Patter- 
son, pers. comm.). Other features of Pristigenys 
(e.g., scale size, body shape, dorsal fin spine, and 
ray lengths) are intermediate to the same features 
as found on Cookeolus and Pseudopriacanthus. 
There is other less salient evidence to support 
retention of both Pristigenys and Pseudopria- 
canthus as valid genera. 


Pristigenys rutoti (Leriche, 1905) 
(Figure 12b) 
Otoliths of this species were described and fig- 


ured from Belgium by Leriche as ““Sparidarum 
rutoti.”” Subsequently, Schubert (1916) reported 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 313 


Ficure 12. Otoliths (sagittae) of: a) Pseudopriacanthus ni- 
phonius (type species of Pseudopriacanthus) 6.5 mm long; b) 


their occurrence in England (refuted by Stinton 
1980) and mentioned their similarity to ‘‘Oto- 
lithus (Sparidarum) rutoti Leriche [sic],” but then 
erroneously referred them to Koken’s (1891) 
Otolithus (Sparidarum) gregarius using the com- 
bination “Otolithus (Pagellus?) gregarius.” Ad- 
ditional errors in their taxonomy appeared in 
later years by other authors, climaxed by their 
being placed in ‘“‘family Pseudopriacanthidae”’ 
(genus Pseudopriacanthus) by Stinton and Nolf 
(1970). Because Leriche used the genitive plural 
in giving these a generic name, he cannot be 
considered the authority for the species, nor can 
the 1905 publication date be considered valid. 
Schubert might possibly qualify as the author, 
but depending upon interpretation of the Inter- 
national Rules of Zoological Nomenclature, this 
also is questionable. It is entirely possible that 
Stinton and Nolf are the authorities for “‘Pristi- 
genys rutoti,’ with a publication date of 1970, 
but the matter is irrelevant to priacanthid tax- 
onomy because features of the sulcus of this fossil 
otolith do not permit its placement in family 
Priacanthidae. In sagittae of extant priacanthids 
(except Priacanthus) the upper and lower rims 
of the ostium (anterior part of the sulcus) con- 
verge posteriorly to appear slightly ovoid and 
funnel-shaped (Fig. 7a—d). 

In the fossil otolith, the dorsal rim of the sulcus 
sweeps posteriorly in a continuous, gentle sig- 
moid curve, making it difficult to distinguish os- 
tium from cauda without reference to the ventral 
rim, which sweeps abruptly dorsad at its juncture 
with the cauda. The ostium comprises less than 
37% of total otolith length in three species of 
Pseudopriacanthus, whereas in Pristigenys rutoti, 
ostium length exceeds 41% of otolith length. Fi- 
nally, when priacanthid otoliths (except Pria- 
canthus) are placed with the outer face down on 
a flat surface, the sulcal side is highly arched 
(convex). Sagittae of P. rutoti lie almost flat when 
placed in this position. 


Pristigenys caduca Nolf, 1973 
(Figure 12c) 


Features of the sulcus of this otolith do not 
permit a placement in the family Priacanthidae. 


_— 


Pristigenys rutoti 2.7 mm long; and c) Pristigenys caduca 2.7 
mm long. Photo of Pseudopriancanthus by Jack W. Schott; 
SEM photos of Pristigenys by Brian White. Sulcus of fossils 
highlighted with broken inked line to show configuration. 


314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19 


Again, the dorsal rim of the sulcus forms a con- 
tinuous, gentle sigmoid curve, while the lower 
rim of the ostium sweeps abruptly upward at its 
posterior terminus—a beryciformlike ostial 
character, but also found in such unrelated fishes 
as Lactarius, Epigonus, Ambassis, Glaucosoma, 
some pempherids, and others. 


Pristigenys bella Stunton, 1980 


We have not handled otoliths from this species, 
but from Stinton’s excellent photographs, it ap- 
pears that the holotype and paratype shown in 
his text figure 34 are valid priacanthid sagittae. 
We cannot vouch for their generic placement, 
however, as otoliths do not accompany the skel- 
etal impressions of Pristigenys substriata, the type 
of the genus. Again, because of ostial configu- 
ration, we question the assignment of the otoliths 
(presumably P. bella) shown in Stinton’s Plate 
13 (figs. 27 and 28) to family Priacanthidae. Stin- 
ton states that these otoliths “are from immature 
fish and demonstrate the ontogenetic changes 
[found in otoliths of these fishes].’’ Except for 
Cookeolus, which goes from a pelagic existence 
as juveniles to a benthic existence as adults, we 
have not observed ontogenetic changes in pria- 
canthid sagittae, and the changes in Cookeolus 
otoliths are not the same as those attributed by 
Stinton to his fossil P. bella. 


Pristigenys spectabilis Stinton, 1980 


We have examined a dozen otoliths from this 
species, and judged by sulcal characters, it should 
not be assigned to family Priacanthidae. Stin- 
ton’s excellent photographs of type material also 
show the non-priacanthid ostial configuration that 
precludes their being priacanthids. 


Pristigenys dentifer Stinton, 1980 


Based upon Stinton’s photographs of the oto- 
liths of P. dentifer, we believe that more than 
one species is involved. Some of his illustrated 
sagittae are very similar to priacanthid otoliths 
in sulcal characters, while others do not appear 
to be. One of his paratypes was found associated 
with ‘“‘a few bones and spines” inside the cavity 
of a gastropod, Clavilithes macrospira. Accord- 
ing to Stinton “some [of the] peculiarly prickly 
fin spines . . . were independently identified as a 
species of Pristigenys.” 

We were loaned one of these fin spines by Colin 
Patterson (BMNH), who informed us (pers. 


comm.) that these prickly spines “‘are certainly 
different from those of Pristigenys substriata, 
which are smooth apart from longitudinal ribs.” 
We compared the fossil spine with fin spines of 
all extant genera of priacanthids and could find 
no agreement with any of them. Camm Swift 
(LACM) examined the spine and suggested it 
bore resemblance to some beryciform fin spines, 
but was unable to suggest a family or generic 
afhliation. 

Otoliths (especially sagittae) generally are ex- 
cellent taxonomic tools, but when working with 
fossils, factors other than those observed on the 
otoliths themselves must be considered. Zooge- 
ography, environment and habitat preferences 
are especially important. If zoogeography had 
been considered, it is doubtful that embiotocid 
perch (presently restricted entirely to the North 
Pacific Ocean) and Leuresthes, Atherinops, and 
other New World atherinids would have been 
reported from the tropical and subtropical Eocene 
of Europe. 

Except for the early pelagic stage of Cookeolus 
boops, all extant priacanthids inhabit areas of 
high relief. Fishes living in these kinds of habitats 
frequently fossilize, but their remains rarely con- 
tain otoliths. Fossil deposits that contain otoliths 
almost invariably represent faunas that inhabit 
flat relief, or pelagic and mesopelagic realism. 
The occasional otolith from an inhabitant of 
rocky, high-relief habitat found in a fossil deposit 
generally represents a prey item or a straggler 
into the flat-relief area, a not uncommon phe- 
nomenon today. Such otoliths are rare, however, 
so the abundance of Eocene sagittae assigned to 
family Priacanthidae (Pristigenys spp.) by Eu- 
ropean paleontologists indicates that habitat 
preference was not a consideration. As already 
pointed out, features of the sulcus also were over- 
looked when making such assignments. Regard- 
less, otoliths are excellent taxonomic tools, ex- 
tremely abundant in the fossil record, and if 
properly used, can furnish an insight into the past 
that can not be gained any other way. 


ACKNOWLEDGMENTS 


As with any such project, our investigation 
could not have been completed without our bor- 
rowing or making use of specimens, library ma- 
terials, work space, ideas, and the special talents 
of others. Our sincere thanks are extended to 
Lillian J. Dempster and W. I. Follett (CAS); Paul 


FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS 315 


A. Gregory, Jack W. Schott and Donald L. 
Schultze (DFG); Mary Butler, Robert J. Laven- 
berg, Camm C. Swift and Bran N. White 
(LACM); Richard H. Rosenblatt (SIO); Boyd W. 
Walker (UCLA); William A. Bussing (Univ. Cos- 
ta Rica); John E. Randall (Bishop Museum, Ha- 
wail); Wayne J. Baldwin (Hawaii Institute of Ma- 
rine Biology); Bruce B. Collette (National Marine 
Fisheries Service, Washington, D.C.); Frederick 
H. Berry (NMFS, Miami); Richard W. Hud- 
dleston (Chevron Oil Field Research Co.); James 
B. Shaklee (now CSIRO, Australia); Peter Forey 
and Colin Patterson (British Museum Natural 
History); Hans-J. Paepke (East Germany); Chris- 
tine Karrer (West Germany); Dirk Nolf (Bel- 
gium); F. C. Stinton (England); George Coates 
(Australia); and Lee Stockland (Tustin, Calif.). 

Wayne C. Starnes (USNM) and Ronald A. 
Fritzsche (Humboldt State Univ.), who also are 
conducting research on priacanthids, were ex- 
tremely helpful. 

Finally we wish to thank the skippers and crews 
of the Qualifier 105, Red Rooster and Royal Po- 
laris and in particular George Cargal and Nor- 
man Kagawa. 


LITERATURE CITED 


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figurés dans !’Ittiolitologia Veronese. Neues Jahrb. Miner., 
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ANDERSON, W. D., Jr., D. K. CALDWELL, J. F. MCKINNEY, AND 
C. H. FARMER. 1972. Morphological and ecological data 
on the priacanthid fish Cookeolus boops in the western north 
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BLAINVILLE, H. M. D. 1818. Poissons fossiles (Jn Nouveau 
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27, Paris). (Not seen, ref. in White 1936.) 

BLEEKER, P. 1869. Neuviéme notice sur la faune ichthyolo- 
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Berlin Vol. 6, iv + 126 p. 

EASTMAN, R. 1905. Les types des Poissons fossiles du Monte 
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Fitcu, J. E., AND R. J. LAVENBERG. 1975. Tidepool and near- 
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Fitcn, J.E., ANDS. A. ScHuttz. 1978. Some rare and unusual 
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Fish Game 64(2):74-92. 

ForsskAL, P. 1775. Descriptiones animalium avium, am- 


phibiorum, piscium, insectorum, vermium; quae in itinere 
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FritzscHe, R. A. 1978. The first eastern Pacific records of 
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FRITZSCHE, R. A., AND G. D. JoHNson. 1981. Pseudopria- 
canthus Bleeker, a synonym of the priacanthid genus Pris- 
tigenys Agassiz. Copeia 1981(2):490-492. 

GrLBerT, C.H. 1891. A supplementary list of fishes collected 
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one new genus and three new species. U.S. Natl. Mus., Proc. 
13:449-455. 

Gos.ing, W. A., AND V. E. Brock. 1960. Handbook of 
Hawaiian fishes. Univ. Hawaii Press, Honolulu. ix + 372 p. 

JORDAN, D. S., AND B. W. EvERMANN. 1904. Descriptions of 
new genera and species of fishes from the Hawaiian Islands. 
U.S. Fish. Comm., Bull. 22:161-208. 

Koken, E. 1891. Neue Untersuchungen an tertidren Fisch- 
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LacEPEDE. 1801. Histoire naturelle des poissons, Paris. Vol. 
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LerIcHE, M. 1905. Les Poissons éocénes de la Belgique. Mu- 
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Myers, G. S. 1958. The priacanthid fish genus Pristigenys. 
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Notr, D. 1973. Deuxiéme note sur les téléostéens des sables 
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Oxen, L. 1817. Cuviers und Okens Zoologien neben einander 
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ScHAFER, W. 1972. Ecology and palaeoecology of marine 
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SCHNEIDER, J.G. 1801. M.E. Blochii. Systema ichthyologiae 
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ScHuBERT, R. J. 1916. Obereoc&ne Otolithen vom Barton 
Cliff bei Christchurch (Hampshire). Kaiserlich-kéniglichen 
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Stinton, F.C. 1980. Fish otoliths from the English Eocene. 
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Stinton, F. C., AND D. Noir. 1970. A teleost otolith fauna 
from the Sands of Lede, Belgium. Bull. Soc. belge Géol., 
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TuHomson, D. A., L. T. FINDLEY, AND A. N. Kerstitcu. 1979. 
Reef fishes of the Sea of Cortez. John Wiley & Sons, New 
York. XV + 302 p. 


Vota, G. S. 1796. Ittiolitologia veronese del museo Boz- 
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1936.) 

Wire, E. I. 1936. On certain Eocene percoid fishes. Ann. 


Mag. Nat. Hist., ser 10, 18(103):43-54. 

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(Not seen, ref. in White 1936.) 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, CA 94118 


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20, pp. 317-321, 3 figs. 


Vol. 43, No 


3" PROCEEDINGS 
. OF THE 


| CALIFORNIA ACADEMY OF SCIENCES 


December 11, 1984 


AMAZONSPRATTUS SCINTILLA, NEW GENUS AND SPECIES 
FROM THE RIO NEGRO, BRAZIL, THE SMALLEST 
KNOWN CLUPEOMORPH FISH 


By 


Tyson R. Roberts 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


ABSTRACT: Amazonsprattus scintilla new genus and species, inhabiting the Rio Negro and its tributaries in 
the Amazon basin of Brazil and feeding on minute aquatic Diptera and planktonic Cladocera, is the smallest 
known clupeomorph fish. The largest specimen is 19.5 mm standard length, but males and females are 
sexually ripe as small as 14-16 mm. Superficially it looks very much like a herring and particularly resembles 
the tropical western Atlantic genus Jenkinsia (Clupeidae). On the other hand, further investigation might 
indicate that it is closely related to some small and poorly known Amazonian anchovies currently placed in 


Anchoviella (Engraulididae). 


INTRODUCTION 


Among the secondary freshwater fishes sharing 
the Amazon basin with 1500 or so species of 
primary freshwater Ostariophysi are somewhat 
more than a dozen species of the clupeomorph 
families Clupeidae and Engraulididae. Amazo- 
nian Clupeidae, or herrings, include several 
species of J/isha or Pellona, some piscivorous 
and attaining nearly one meter; Pristigaster cay- 
ana Cuvier, 1829, with its greatly expanded tho- 
rax, attaining perhaps 250 mm; and Rhinosar- 
dinia amazonica (Steindachner, 1880), typically 
40-60 mm long. The Engraulididae, or ancho- 
vies, include Cetengraulis juruensis Boulenger, 
1898, attaining perhaps 200 mm, and a number 
of species currently placed in Anchoviella, some 
as small as 30-40 mm. Recently, while searching 
for comparative material of larvae, I was thus 
surprised to find some sexually ripe Amazonian 
fishes, which looked like clupeomorphs, less than 


20 mm in standard length. Study of cleared and 
stained preparations confirmed that these spec- 
imens are not Ostariophysi and that they rep- 
resent an undescribed genus and species of 
clupeomorph. 


Amazonsprattus, new genus 


TYPpe-SPECIES. —Amazonsprattus scintilla, new species. 


DiaGnosis.— Minute, slender, scaleless clu- 
peomorphs without pre- or post-pelvic abdom- 
inal scutes. Ventral myotomic progression in- 
complete. Premaxilla absent or minute and 
toothless. Maxilla with 16-20 very small conical 
teeth. Two supramaxillae. Dentary and palate 
usually toothless (one tooth observed on dentary 
in one specimen). Branchiostegal rays 4-5. Dor- 
sal fin with 12-13 rays, its origin in posterior half 
of body; anal fin with 14-16 rays, its origin below 
anterior third of dorsal fin. Pectoral fin rays 7- 
9. Pelvic fin rays 6. 


[317] 


318 


FiGure |. 


Amazonsprattus scintilla, new species 
(Figures 1-3) 

Ho.otype.—CAS 52175, 17.0 mm (sex undetermined), Rio 
Jufari between Castanheiro Grande and Santa Fé, collected by 
Martin Brittan, 21 April 1964. 

ParATyPes.—CAS 52176, 18: 14.3-19.0 mm (five males, 
eight females, six sex undetermined), collected with holotype 
(5: 16.5—19.0 mm cleared and stained with alcian and alizarin); 
CAS-SU 68891, 19.5 mm (sex undetermined), Rio Negro at 
Santa Isabel, collected by Carl Ternetz, 17 January 1925. 


DESCRIPTION.— Head compressed and mod- 
erately elongate, its length almost four times in 
standard length. Eyes moderately large and 
strongly compressed or flattened laterally. Eye 
diameter about four times in head length. Entire 
medial surface of eyes closely approximate (so 
that forebrain is confined to a small space dorsal 
to eyes), and medial surface of eyes just as flat 
as lateral surface. Ventral surface of eyeball with 
a prominent choroid fissure. Hyaline eyelid well 
developed. Snout moderately elongate, its length 
about equal to eye diameter. Nasal organ mod- 
erately large, with rather small anterior and pos- 
terior nostrils. Mouth terminal. Lower jaw elon- 


palatine 


supramaxillae 


lower jaw 


metapterygoid 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 20 


aoe a — — = 


> pSsiass 


Amazonsprattus scintilla, 17.0 mm (holotype, CAS 52175). 


gate, extending posteriorly to below posterior 
margin of eye, but gape much restricted and en- 
tirely anterior to eye. Maxilla slender and mod- 
erately elongate, extending posteriorly to below 
anterior margin of eye. Anterior and posterior 
portions of cranial fontanel open, posterior por- 
tion entirely bridged by a narrow, cartilaginous 
taenia medialis. Circumorbital bones apparently 
five (very fragile, hence easily broken, and stain- 
ing very weakly with alcian). Hyosymplectic en- 
tirely cartilaginous, without large, axe-shaped 
anterodorsal process usually present in clupeids. 
Opercle with strongly concave dorsoanterior and 
dorsoposterior margins; dilatator process of 
opercle present but weakly developed. First gill 
arch with a total of 18-19 large, elongate gill 
rakers on its leading edge. Rakers on trailing edge 
of first gill arch and on successive arches slightly 
less numerous and considerably smaller. Upper 
and lower pharyngeal toothplates small, with 8- 
10 conical teeth (branchial and hyoid arches oth- 
erwise edentulous). Hyoid and branchial arches 
entirely cartilaginous except for branchiostegal 
rays, gill rakers, and pharyngeal toothplates. 


hyosymplectic 


opercle 


te ee ie at 


subopercle 
interopercle 


Ficure 2. Amazonsprattus scintilla, 18.0 mm, CAS 52176. Lateral view of jaws, suspensorium, and opercular bones (hy- 


osymplectic and palatine cartilaginous). 


ROBERTS: MINUTE AMAZONIAN CLUPEOMORPH 


319 


Figure 3. Amazonsprattus scintilla, 18.8 mm, CAS 52176. Axial skeleton. Total vertebrae 17 + 21 = 38; pectoral girdle 
abnormally incomplete; a series of 10-11 small, irregularly triradiate intermuscular bones lateral to vertebrae 2-13 has been 


omitted for clarity. 


Body compressed, slender, its greatest depth 
about 6-8 in standard length. Abdomen round- 
ed. Myotomes well defined, those fully formed 
33-34, or about four fewer than total vertebrae. 
Dorsal and anal fin rays low set, with moderately 
falcate margins, dorsal fin rays about 1.5 times 
longer than anal. Pectoral and pelvic fins rela- 
tively small, pelvic smaller than pectoral. Pelvic 
scute variably developed, absent or failing to stain 
in some specimens, weakly to strongly stained 
with alcian or alizarin in others. In specimens 
with relatively well developed scutes there is an 
elongate anteromedian process and an elongate 
lateral ascending process on each side. Pectoral 
girdle with bony posttemporal, supracleithrum, 
and cleithrum (postcleithra absent), cartilaginous 
scapulocoracoid, and three rows of radials (prox- 
imal, medial, and distal, with five, five, and eight 
radials respectively). Caudal fin moderately 
deeply forked, upper and lower lobes about equal 
and with rounded margins. Upper and lower lobes 
overlapping slightly when adducted. Principal 
rays 4-5 of upper lobe and 3-4 of lower lobe 
with delicate alar flaps (not illustrated). In an 
18.8-mm specimen the large alar flap on ray 5 
consisted of four overlapping scalelike laminae 
(lightly stained with alcian) that may actually be 
modified scales. Caudal fin with 10 + 9 principal 
rays, 8—9 upper and 8 lower procurrent rays. Cau- 
dal fin skeleton with a parhypural, six separate 
hypurals, and a single epural. Hypural 2 fused to 
complex ural centrum (as in many clupeoids). 

Total vertebrae 37(4) or 38(1), 16-17 abdom- 
inal plus 20-21 caudal. All vertebrae with a sim- 
ple, slender neural spine. Supraneurals 6-8. Or- 
igin of pelvic fin below vertebrae 13-14, of dorsal 
fin above vertebra 18, and of anal fin below ver- 
tebrae 20-21. All abdominal vertebrae except 
first two with fully developed ribs. First two ver- 


tebrae usually without ribs, sometimes second 
vertebrae with incompletely developed ribs (Fig. 
3). Distal ends of most abdominal ribs deflected 
posteriorly. First two caudal vertebrae with re- 
duced ribs. Intermuscular bones well developed 
anteriorly and posteriorly. Anteriorly two dis- 
similar and morphologically complex sets of in- 
termuscular bones. A series of about 13 epipleu- 
ral intermuscular bones parallel to abdominal 
vertebrae 3-16. Anteriormost eight epipleurals 
with a well developed anteromedial process 
proximally; this process is absent from last five 
epipleurals, which become progressively smaller. 
Distal ends of epipleurals, except reduced pos- 
teriormost one, closely approximated to distal 
half of ribs (Fig. 3). In addition to epipleurals, a 
series of 10-11 small, irregularly triradiate in- 
termuscular bones lies just dorsal to epipleurals 
and directly lateral to centra of abdominal ver- 
tebrae 2-12 or 13 (not illustrated). Posterior two 
processes of these triradiate elements lie quite 
near body surface, but anterior process lies much 
deeper. Posteriorly, two similar series of 8-10 
simple dorsal and ventral intermuscular bones 
extend laterally just above and below caudal ver- 
tebrae 9-19 (Fig. 3). 

Alimentary canal with a well-defined stomach. 
Pyloric caeca in two or three groups: a dorsal 
group with one or two elongate caeca; a ventral 
group with about four or five elongate caeca; and 
sometimes a second ventral group of about four 
short, poorly defined or only partially separate 
caeca. Intestine straight. Gut contents of four 
specimens were as follows: specimen 1) two dip- 
teran pupae; specimen 2) a single dipteran pupa; 
specimen 3) numerous small Cladocera of two 
size classes, 270 x 150 umand 72 x 55 um; and 
specimen 4) a single dipteran pupa, several small 
dipteran larvae, and moderately numerous 


320 


cladocerans of a single kind. My sketches of the 
cladocerans in this fourth specimen, 424 x 255 
um, with a pair of curved, divergent, strongly 
deflected horns 184 um long projecting from the 
rostrum, were tentatively identified as Bosmi- 
niopsis deitersi Richard, 1895, by Thomas Zaret. 

Gonads readily identifiable in most of the type- 
specimens. Eight, 15.9-18.2 mm, have creamy 
or pale orangish ovaries with eggs in more or less 
good condition observable through body wall with 
transmitted light. One of these, 17.3 mm, con- 
tained 20 eggs 0.2 mm in diameter. Five, 14.3- 
16.2 mm, have milk-white testes. In two of these 
the testes are particularly well developed and ex- 
hibit numerous “segments” or laminae, about 
five per myotome, comparable to laminae ob- 
served in testes of other minute teleosts (e.g., 
Sundasalanx, Roberts 1981, fig. 1a). In both sexes 
the gonads appear to be single and occupy only 
the posterior half of the body cavity, from about 
the origin of the pelvic fin to the vent. In seven 
specimens, 16.0-19.5 mm, gonads not observed 
and sex undetermined. 

Judging from preserved material live Ama- 
zonsprattus probably are translucent or even 
transparent. Only a few melanophores on head, 
largest and most obvious a group of about six 
superficial to cleithrum and clearly visible through 
gill cover, and two large ones on either side of 
dorsoposterior margin of hindbrain (Fig. 1). A 
number of large, deep-lying melanophores as- 
sociated with posteroventral portion of cranium 
(not illustrated). Dorsal surface of cranium oth- 
erwise usually devoid of pigment. Tip of snout 
and lower jaw, and side of head just below eye 
and midway between eye and end of gill cover 
sometimes with a few small melanophores. Body 
with relatively few melanophores, mostly on 
ventral half, including base of anal fin and caudal 
peduncle. A row of about 10 melanophores, one 
per segment, along each ventral myotomic bor- 
der, and a midventral row of about five near 
pelvic fins. Two rows of segmental melanophores 
near anal fin base: one row at ventral end of 
myotomes and between pterygial muscles, 
another on bases of anal-fin rays. A row of small 
melanophores, more than one per segment, on 
ventral portion of caudal peduncle, and some 
small melanophores near base of caudal fin. A 
nearly straight row of small segmental melano- 
phores on side of body just above midline (with- 
out evident anatomical relationship to any un- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 20 


derlying structures). In holotype and most 
paratypes this row commences posterior to dor- 
sal fin origin (Fig. 1), but in some paratypes it 
extends nearly entire length of body. Dorsal sur- 
face of body devoid of pigmentation. A few me- 
lanophores on basal portion of anterior dorsal 
fin rays, but dorsal fin without basal melano- 
phores like those of anal fin. Caudal fin relatively 
densely pigmented, with large melanophores 
more or less regularly distributed on upper and 
lower lobes (Fig. 1); an area near middle of caudal 
fin devoid of melanophores. Sexual differences 
in pigmentation not observed. 

Note ON Type-Loca.ities. — The Rio Jufari is 
a low-gradient, swampy tributary with an enor- 
mous mouth-bay at its confluence with the Rio 
Negro, about 20 km upriver from the relatively 
narrow mouth of the much more important Rio 
Branco. Maps I have seen do not show Castan- 
heiro Grande or Santa Fé;saccording to the col- 
lector (pers. commun. M. Brittan, March 1983) 
several days of slow boat travel up the Jufari were 
required to reach the collecting sight. Santa Is- 
abel is an old name for the modern town of Ta- 
purucuara. This portion of the Amazon basin lies 
within equatorial rain forest; here the waters of 
the Rio Negro and its tributaries (excepting the 
Rio Branco) are generally darkly tinted, have a 
pH of 4-5, and are extremely low in mineral 
conduct. 

ETyMoLoGcy.—Amazon; and sprattus (Latin, 
masc.), a herring or herringlike fish; scintilla (Lat- 
in, masc.) a spark, hence the smallest trace or 
particle (employed as a noun in apposition). 


DISCUSSION 


Few collectors have preserved specimens of 
the smallest Amazonian fish species, and it may 
well be that the smallest species of various other 
groups have yet to be discovered. In the Amazon, 
where community structure of freshwater fishes 
may be more complex than anywhere else, mi- 
nute body size seems to have arisen primarily in 
response to biotic factors. This, in my opinion, 
is the general reason why so many of the smallest 
Amazonian fishes belong to secondary freshwa- 
ter fish groups which in terms of relative numbers 
of species represent an insignificant fraction of 
the fauna. Amazonsprattus provides an excellent 
example of the survival of a group of secondary 
fishes in the midst of the Amazonian ichthy- 
ofauna by evolution of minute body size and an 


ROBERTS: MINUTE AMAZONIAN CLUPEOMORPH 


entirely freshwater life history, possibly involv- 
ing year-round continuous reproduction of mi- 
nute young. Other particularly striking examples 
of the phenomenon include the two minute 
species of Poecilia (or Pamphorichthys) and the 
two or more minute species of Microphilypnus. 
These are the only members of the families Poe- 
ciliidae and Eleotrididae inhabiting the interior 
of the Amazon basin. For further discussion of 
these and other examples see Roberts (1972). 
Relationships of Amazonsprattus are unclear, 
and for the moment it is perhaps best left un- 
assigned to family. Having examined its skeletal 
anatomy and made comparisons with a number 
of herrings and anchovies, I was inclined to place 
the genus in Clupeidae, largely on account of its 
jaw structure, which is unlike anything I have 
observed in Engraulididae. But my colleague 
Gareth Nelson, who is studying Amazonian An- 
choviella and has examined some small unde- 


321 


scribed species I have not seen, is inclined to 
believe that it may belong to this group of En- 
graulididae. 


ACKNOWLEDGMENTS 


For information, comments, or other assis- 
tance I wish to thank Martin Brittan, Norma 
Chirichigno, Dusty Chivers, Lillian Dempster, 
William N. Eschmeyer, Karsten Hartel, Michael 
Hearne, Albertina Kameya, Vincent Lee, Sarah 
Ward, Thomas Zaret, and George Zorzi. The 
manuscript was reviewed by Gareth M. Nelson 
and Peter J. Whitehead. 


LITERATURE CITED 


Roserts, T. R. 1972. Ecology of fishes in the Amazon and 
Congo basins. Bull. Comp. Zool. 143(2):117-147. 

. 1981. Sundasalangidae, a new family of minute fresh- 

water salmoniform fishes from Southeast Asia. Proc. Cali- 

fornia Acad. Sci. 42(9):295-302, 6 figs. 


CALIFORNIA ACADEMY OF SCIENCES 
Golden Gate Park 
San Francisco, California 94118 


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