sa I tad re sha eve bea et viet eek vase gare Poe nbn i Em ena pene! latetsiteTacmerte 20 aca eit arm uF ty - ) a , is - +! ty PROCEEDINGS OF THE California Academy of Sciences Volume 44 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1985-1986 » 3 PROCEEDINGS OF THE California Academy of Sciences Volume 44 SEP 2 4 1999 LIBRARIES SAN FRANCISCO PUBLISHED BY THE ACADEMY 1985-1986 PUBLICATIONS COMMITTEE Daphne G. Fautin, Scientific Editor Sheridan Warrick, Managing Editor Frank Almeda Luis F. Baptista Wojciech J. Pulawski Frank 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 (Compiled by Tomio Iwamoto) New names Acanthogilia 111-125 Asarcenchelys 12 Asarcenchelys longimanus 12-15 Cosmochilus cardinalis 3-7 Emoia campbelli 49-51 Emoia trossula 47-49 Gastrocopta cordillerae 244-245 Gastrocopta (Albinula) montana 241-243 INDEX TO VOLUME 44 Gastrocopta (Albinula) sagittaria 244 Helminthoglypta bozemanensis 255-256 Mixomyrophis 10 Mixomyrophis pusillipinna 10-11 Phenacostethus trewavasae 226-235 Pupoides (Ischnopupoides ) tephrodes 246-247 Radiocentrum taylori 249-251 Radiocentrum \aevidomus 25 1—252 New names in boldface type Acanthogilia | 1 1-126 gloriosa 113-125 Adephaga 67-68, 73, 81, 83-92, 94-97 Agabus cordatus 98 Agonidae 17, 160 Agonopsis chiloensis 17 vulsa 19-22 Agonus 17, 31 cataphractus 17, 31 Ahlia 9 egmontis 15 Allophyllum 117 glutinosum 124 Ammonitella 252, 264 lunata 264 Ammonitellidae 238, 252, 264 Amphizoa 68-72, 78, 81, 83, 98-100, 102 carinata 67-68, 75-78 davidi 67-68, 70-73, 79-81, 98-102, 107 davidis 70 insolens 68, 70-75, 79-80, 82, 98-101, 103-106 josephi 70, 72 kashmirensis 68 lecontei 67-68, 70-80, 98-100, 103-104, 106-107 planata 75 striata 70-71, 74-75, 78-80, 82, 98-100, 103-106 Amphizoidae 67-69, 81, 84, 89, 100 Anableps 233 Anachis 278 Anachis? sp. indet. 278 Anchistoma parvulum 253 Anguispira 262 holroydensis 262 russelli 262 Anomia peruviana 59 Archostemata 83-84, 86-87, 89, 91, 94-97 Arionidae 262 Artediini 172 Artedius 157-163, 165-168, 170-175, 177, 182, 185, 188-189, 191, 194, 198-199, 213, 217, 221 corallinus 159, 161, 171-172, 188 creaseri 157-159, 161, 163, 166-168, 170-175, 177, 213-217, 221 fenestralis 157, 159, 161-168, 170-182, 184, 188 harringtoni 157-159, 161-162, 164-168, 170-175, 179, 181-185, 188 lateralis 157-164, 166-168, 170-175, 178-179, 182, 184-188 meanyi 157-159, 161, 163, 166-168, 170-175, 177, 216-221 notospilotus 159, 161, 171-172, 188 type 2 158 type 3 157, 161-164, 168, 170, 172-175, 179, 182, 184, 188-191 Asarcenchelys 12 longimanus 9, 11-15 Aspidophoroides 38-39 bartoni 18-20, 22, 37 monopterygius 17 olriki 18-20, 22, 37-38 Asterotheca 18 Atherinomorpha 225 Avicula 60-61 Balanidae 60 Balaninae 60 Balanoidea 60 Balanus 60 calidus 55-56 poecilus 55-56, 59-61, 65 sp. cf. B. calidus 55-56, 59-60, 65 [285] 286 tintinnabulum galapaganus 61 trigonus 55-56, 59-60, 65 Bathyagonus 38 alascanus 18-20, 22, 37 infraspinatus 18-20, 22, 37-39 nigripinnis 18-20, 22, 37-38 pentacanthus 18-20, 22, 37 Binneya 263 antiqua 263 Biomphalaria 258 glabrata 258 pseudammontius 239, 258 sp. cf. B. pseudammonius 258 Bonplandia 117, 120-121 geminiflora 117, 123 Bostryx (Peronaeus ) 265 Bothragonus 38-39 swant 18-20, 22, 37 Brachinus 88 Buccinacea 277 Buccinidae 277 Bulimulidae 262 Bulimulus 262 sp. 262 Bulimus nitidulus 245 Bursa 276 caelata 275-276 Bursidae 276 Camaenidae 263, 265 Cantharus 277 (Gemophos) 277 (Gemophos) sanguinolentus 277 Janellii 277 sanguinolentus 275, 277 Cantua 111, 116-118, 120-121, 125 buxifolia 118-121, 125 candelilla 121-123, 125 pyrifolia 118, 123 quercifolia 121, 125 Cantueae 118, 125 Carabidae 68, 71, 85 Carabinae 89 Carabini 87 Carabus 86 Caracolus 263 aquilonaris 263 Carbinae 83 Cardiacea 275 Cardiidae 275 Carlhubbsia kidderi 234 Carychium 261 sp. 276 Ceratostethus bicornis 228, 234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Charopa 262 Charopidae 262 Chelonibia 56, 58 testudinaria 55-58, 64 Chelonibiinae 56 Chitonotus 170, 173 Cicindelini 87 Clausiliidae 262 Clinocottus 157-160, 162-168, 170-175, 177, 199, 201, 204, 209, 213, 217, 221 acuticeps 157-162, 164-168, 170-172, 174-175, 199-204, 205, 207 analis 157, 159, 162, 164, 168, 170-171, 174— 175, 203, 205, 209, 212-213 embryum 157, 159, 162, 164, 166-168, 170-171, 174-175, 203-208, 210 globiceps 157, 159-162, 164, 166-168, 170-172, 174-175, 203, 205, 208-212 recalvus 157-159, 164, 168, 170-171, 174-175, 205, 209 Cobaea 117, 120-121, 125 biaurita 123 Codakia 65 Coleoptera 67, 85, 87, 92, 95 Collomia 117, 121 Collumbellidae 277 Columbella 277 castanea 275, 278 cf. C. strombiformis 277 lanceolata 278 major 278 pyrostoma 277 strombiformis 277 Colymbetinae 88 Conacea 279 Concavus 55-56, 61 (Arossia) 61 (Arossia) panamensis 56 (Arossia) panamensis eyerdami 55 (Arossia) sp. cf. C. panamensis eyerdami 55, 61- 62, 65 aquila 61 henryae 61 panamensis eyerdami 61 panamensis panamensis 61 Conidae 279 Conus 279 (Asprella) 279 (Asprella) arcuatus 275, 279 (Chelyconus) 279 (Chelyconus) orion 279 (Cylindrus) 279 (Cylindrus) lucidus 275, 279 (Lithoconus) arcuatus 279 arcuatus 279 loomisi 279 lucidus 279 orion 279 vittatus 279 INDEX Coptoclava 96 Coptoclavidae 83 Coronula 58 diadema 55, 57-58, 61, 64 Coronulidae 56 Coronulinae 58 Coronuloidea 56 Coscinodiscus 128 eccentricus 137 lacustris 128, 144 lineatus 131 Cosmochilus 1-7 cardinalis |—7 falcifer 2-3, 6-7 harmandi 2-3, 6-7 Cottidae 157-158, 160, 173, 199, 204, 208 Cottus 167 Craterarion 263 pachyostracon 263 Cybister 88 Cyclocheilichthys 7 Cyclophoridae 261 Cyclopteridae 160, 168 Cymatiacea 276 Daedalochila 263 Dentatherina 225 mercerl 226 Discidae 262 Dysmathes sahlbergii 72 Dytiscidae 68, 83-84, 89, 98, 101 Dytiscus 88 Ellobiidae 261 Emoa samoensis 45 Emoia 41-44, 46, 48, 50 aneityumensis 42, 48 caeruleocauda 41 campbelli 41-42, 49-51, 52 concolor 41-43, 45-47, 49, 51-52 cyanogaster 41 cyanogaster tongana 46 cyanura 41 murphyi 41-43, 52 nigra 41-42, 49, 51 nigromarginata 42-43, 51 parkeri 41-43, 52 physicae group 42 resplendens 42, 45 samoense 43, 45, 47 samoensis 41-47, 49, 51-52 Samoensis group 41-43, 48, 50-51 sanfordi 42, 49, 51 speiseri 42 trossula 41-44, 46-49, 51-52 Engina 277 pyrostoma 277 Enophrys bison 160, 168 Eodromeinae 83, 89, 97 Eohipptychia eohippina 271 Eretes 88 Eriastrum 117-118, 125 Eucalodium eophilum 260 Eumeces 44 samoensis 43, 47 Euprepes concolor 45 resplendens 45 samoensis 45 Fasciolariidae 278 Fasciolariinae 278 Fraginae 275 Fusinus centrifugus 278 Garra 7 Gastrocopta 237, 240-241, 244, 258-259 (Albinula) 241, 243-244, 256-257, 259 (Albinula) contracta 241, 243, 256 (Albinula) dupuyi 244 (Albinula) falcis 241 (Albinula) holzingeri 241, 243, 256 (Albinula) montana 238, 241-244 (Albinula) proarmifera 241 (Al/binula) sagittaria 238, 242-244, 257 (Albinula) sp. a 238, 242-244 (Albinula) tridentata 241 (Gastrocopta) 241 (Gastrocopta) cristata 243 (Gastrocopta) sp. 261 armifera 241-242 contracta 244, 256-257 cordillerae 238, 242, 244-245, 258 Sagittaria 258 sp. cf. G. montana 242, 258 Gastrodonta coryphodontis 263 imperforata 263 Gastropoda 237-238, 240, 275 Geadephaga 68, 81 Gehringia 88 Gilia 111, 116-117, 121-123, 125 gloriosa 111, 115, 117 incisa 121-122 insignis 121 leptomeria 122 rigidula 121-122, 125 ripleyi 121-122, 125 sect. Giliastrum 111, 118, 121-122, 125 sect. Leptodactylon 111 tricolor 122 288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Gilieae 125 Hodopoeus 265 Glypterpes crassus 263 rotundatus 264 hesperarche 263 veternus 264 Holospira 262, 265 Gonyodiscus 248 adventicia 262 Gymnosteris 117, 121 dyeri 262 Gyrinidae 68, 84, 88, 101 grangeri 262 leidyi 262 Haliotis 270 sp. 262 Haliplidae 89 Horaichthys Haplochromis setnai 234 elegans 229 Humboldtiana 256 Helicina palmeri 256 cokevillensis 261 Huthia 117-118, 120-121, 125 cretacea 261 Hydaticus 88 vokesi 261 Hydnophytum 50 Helicinidae 261 Hydradephaga 67-68, 73, 81, 85-87, 91, 95-98, 101 Helix Hydronebrius 68, 98 adapis 264-265 Hygrobiidae 68, 83, 88-89, 101 nacimientensis 248, 264-265 Hypsagonus 17, 30 polygyrella 252 quadricornis 17, 19-22, 30 riparia 256 span Ose a0: Icelidae 158 tudiculata 253 Teelinae 158 maaan TSS Icelinus 157, 161-163, 166-167, 170-174, 177, 217, Helminthoglypta 237, 253-259, 264 224 aes sp. 160, 168 bop 2°? Icelus 166, 173 bozemanensis 238, 254-256, 258-259, 264 Ipomopsis 111, 116-118, 123, 125 californiensis 254-255, 259 gloriosa 11 1 115: 117 fieldi 259 ape Isomeria 265 hertleini 254 mailliardi 254, 259 nickliniana 255 Kanabohelix nickliniana awania 259 kanabensis 263 obtusa 254 reederi 254 Labeo 7 sp. 264 Labeoinae 7 sp. cf. H. bozemanensis 255 Laccophilinae 88 stocki 254 Langloisia 117 traskii 254 Latirus 278 tudiculata 254 centrifugus 275, 278 Helminthoglyptidae 238, 249, 255, 264 Leistus 86 Hemilepidotus 166-167, 173 Leptarionta 256 hemilepidotus 168 Leptocottus 166 spinosus 160 armatus 160, 168 Hemitrochus 255-256, 264 Leptodactylon 111, 117-118, 121, 123, 125 hoemastomus 255 gloriosa 115, 117 sp. 264 gloriosum 111 Hendersonia 261 pungens 124 evanstonensis 261 Liadytidae 83, 89 oregona 261 Linanthus 117 Heterodonta 275 dianthiflorus 124 Hexagrammidae 160 grandiflorus 124 Hexagrammos Loeselia 111, 116-118, 120, 123 sp. 160, 168 amplectens 124 Hiletus 86 gloriosa 111, 115 INDEX gregell 124 mexicana 118 purpusil 118 Lucidella 261 buttsi 261 Lygosoma cyanogaster 45 cyanogaster tongana 45 samoense 43, 45, 47 Lymnaea 238 Lysinoe 256, 264 breedlovei 264 Malacocottus 166 Megabalaninae 61 Megabalanus 61 clippertonensis 61 galapaganus 55-56, 61-65 sp. indet. 62-64 tanagrae 61 Menesiniella 61 Mesogastropoda 275 sagensis 264 Metaphos laevigatus 277 Microsteris 117, 121 gracilis 123 Mirophallus bikolanus 225-226, 232, 234-235 Mixomyrophis 10-11 pusillipinna 9, 10-11, 14 Monadenia 256, 260, 264 (Shastelix) marginicola 264 antecedens 264 dubiosa 264 Morulius chrysophekadion 7 Muraenichthys 9 puhioilo 11 Myoxocephalus 162-163, 172-174, 221 sp. 160, 168 Myrophinae 9 Myrophini 9 Myrophis 9-10, 15 vafer 14 Nassariidae 278 Nassarius 278 caelolineatus 278 nodicinctus 278 Natica uber 276 Naticacea 276 Naticidae 276 Navarretia 117, 121 fossalis 122 mitracarpa 122 Nebria 86 Nebriini 83, 86-87 Necronectulus 84, 89-90, 94-96 Neenchelys 11, 14-15 Neostethidae 225 Neostethinae 225-226, 229, 231 Neostethus 229, 233 Noteridae 83, 89 Notiokasiini 83, 86 Notiophilini 87 Ocella dodecaedron 19-22 Odontopyxis 38-39 trispinosa 18-20, 22, 37-38 Oleacinidae 263 Oligocottinae 158, 171-172 289 Oligocottus 157-160, 162-168, 170-175, 177, 199, 208, 2133.21; 2211 maculosus 157-164, 167-168, 170, 172-175, 191- 195, 197 rimensis 159, 163, 172 rubellio 159, 163, 172 snyderi 157, 159-164, 167-168, 170, 172-175, 194-199, 221 Omma 88 Omphalina laminarum 263 oreodontis 263 Ophichthidae 9 Opisthiini 83, 87 Oreoconus Jepseni 262 planispira 262 spp. 262 Oreohelicidae 238, 247-249, 264 Oreohelix 248-249, 251-252, 259-260 angulifera 248 chiricahuana 247 Steini 263 thurstoni 248 Orthonopias 173 Orthurethra 240 Osteochilus 7 Paleocyclotus sp. 271 Pallasina barbata 19-22 Parahygrobiidae 83 Paricelinus 173 Pelecypoda 275 Pelophila 86 Percis japonicus 19-22 Phallostethidae 225, 233 Phallostethinae 225-226 290 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Phallostethus 225, 229, 231-233 dunckeri 225-226, 228-229, 232 Phenacostethus 226, 229, 231-232 posthon 225-228, 232-235 smithi 225-230, 232-233 thai 226 trewavasae 225-235 Phlox 117, 121 andicola 123 Phos 277 (Metaphos) 277 (Metaphos) laevigatus 277 chelonia 277 laevigatus 275, 277 Physa 239 Planorbis amplexus 253 Platymantis 41 Pleurodonte (Dentellaria) 263 (Dentellaria) sp. 263 (Pleurodonte) 263 (Pleurodonte) wilsoni 263 Pleurotomaria 272 Podothecus 17, 31 acipenserinus 19-22, 31 gilberti 31 thompsoni 31 veternus 31 Polemoniaceae 111, 116-118, 121-123, 125 Polemonium 116, 121 Polinices 276 (Polinices) 276 (Polinices) uber 276 intemeratus 276 uber 275-276 unimaculatus 276 Polygyra 263 expansa 263 martini 263 petrochlora 263 sp. 263 venerabilis 253 veternior 263 Polygyrella 237, 252-253, 257-259, 264 amplexa 264 amplexus 253 parvula 253, 264 polygyrella 251-253, 257, 264 sp. 264 sp. cf. P. polygyrella 238, 251, 253, 257, 264 venerabilis 264 Polygyridae 263, 265 Polymita 264 texana 264 Polyphaga 97 Prosobranchia 261 Protorabinae 83, 89 Protornatellina isoclina 261 Pseudarinia convexa 261 pupilla 261 uniplica 261 Pseudocolumna 265 haydeniana 262 spitzia 262 spp. 262 vermicula 262 Pseudomyrophis 11, 14-15 micropinna 14 Pteria beilana 61 peruviana 61 sterna 60 Pulmonata 237-238, 240, 261 Pupa acarus 240 contracta 241 hordacea 245 incolata 245 Pupillidae 238, 240-241, 243-244, 246-247, 261 Pupoides 237, 245-246 Pupoides (Ischnopupoides) 237, 258 ([schnopupoides) 245-247 ([schnopupoides) hordaceus 245-246, 261 ([schnopupoides) inornatus 245 (Uschnopupoides) sp. 245, 247, 261 ([schnopupoides) sp. cf. P. (I. ) hordaceus 238, 246— 247, 261 ([schnopupoides) tephrodes 238, 245-247, 261 (Pupoides) albilabris 246 hordaceus 246-247, 258-259 inornatus 247, 259 tephrodes 246-247, 258 Purpura Sanguinolentus 277 Radiocentrum 237, 247-249, 251-252, 256-258, 260 anguliferum 250, 252, 264 avalonense 257, 259 chiricahuanum 250, 252 chiricahuanum chiricahuanum 250 chiricahuanum obsoletum 249-250 grangeri 248, 264 hachetanum 251-252 hendersoni 248, 250-251, 264 laevidomus 238, 250-252, 257-258, 264 taylori 238, 249-252, 258, 264 thurstoni 250, 252, 264 Radulinus 166 asprellus 160, 168 Ranella caelata 276 INDEX Rhiostoma 261 americana 261 Ruscariops 173 Salvia mellifera 257 Sarritor Srenatus 19-22 Schizophoridae 83, 89 Scincidae 41 Scorpaenichthys 166 marmoratus 160, 168 Scorpaenidae 160 Scorpaeniformes 157 Sebastes 163 flavidus 160, 168 Shastelix 264 Sigmurethra 247 Sinilabeo 7 Spanglerogyrus 84, 89-91, 94-95 Stellerina 163 xyosterna 168 Stellerinna xvosterna 160 Strobilops 261 sp. 262 Strobilopsidae 261 Stombina 278 (Strombina) 278 (Strombina) lanceolata 278 gibberula 278 lanceolata 275, 278 recurva 278 Subulinidae 262 Systolosoma 87, 94, 96 Terebra 279 armillata 279 plicata 279 Terebridae 279 Tetraclita 55-56, 58 milleporosa 55-61, 64-65 panamensis 58 porosa var. communis 58 rubescens 55, 60 rubescens rubescens 55, 58 sp. indet. 55, 58-60, 64 squamosa milleporosa 58 stalactifera 58-59 stalactifera confinis 58 stalactifera stalactifera 58 Tetraclitidae 58 Tetraclitinae 58 Thalassiosira 127-128, 130-132, 137-138, 142, 146, 151, 153-154 angstil 128 anguste-lineata 128, 146, 151, 153 291 decipiens 127-128, 137-138, 142, 144, 152-154 eccentrica 128, 137-138, 142, 153-154 endoseriata 128, 146, 151, 153 hendeyi 127-128, 130-132, 150, 153-154 incerta 128, 132, 146, 151, 153-154 lacustris 128, 146, 151, 153 leptopus 131 lundiana 128, 137, 142, 151, 153 minuscula 128, 132, 137-138, 153 nodulolineata 127, 130-132, 150-151, 153-154 nordenskioeldti 127-128, 130, 138, 153 oestrupii var. venrickae 130, 137, 153 pacifica 127, 130, 138, 142, 153 punctigera 128, 130, 137-138, 142, 151, 153 rotula 127-128, 130, 146, 151, 153 simonsenti 130-132, 153 stellaris 130, 146, 151, 153 tenera 130-132, 153 visurgis 127, 130, 138, 142, 146, 151-153 wongli 127, 130, 132, 136, 152-154 Tornatellinidae 261 Tozerpina douglasi 261 mokowanensis 261 rutherfordi 261 Trachypachidae 82, 85, 87 Trachypachinae 83, 89, 97 Trachypachus 94 Triadogyrus 90, 94 Triaplidae 89-90 Triglops 173 Trigonocardia 275 biangulata 275 Trigonocardia? sp. 274-275 Triodopsis 263 spp. 263 Turridae? 280 Turritella 275-276 broderipiana 275-276 broderipiana marmorata 274-276 gonostoma 275-276 marmorata 275 nodulosa 276 rubescens 274, 276 Turritellacea 275 Turritellidae 275 Turritellinae 275 Urocoptidae 262 Veneroida 275 Ventridens 263, 265 lens 263 sp. 263 Vespericola 263 dalli 263 292 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Xeneretminae 18 alascanus 17 Xeneretmus 17-18, 30-36, 38 latifrons 17, 32 (Xeneretmus) 18, 36 pentacanthus 17 (Xeneretmus) triacanthus 17-18, 32, 36-38 triacanthus 17, 32, 36 (Xenopyxis) 17-18, 32, 36, 38-39 Xenodexia (Xenopyxis) latifrons 17-18, 32-35 ctenolepis 234 (Xenopyxis) leiops 17-18, 32, 35-36 Xenopyxis 32 (Xenopyxis) ritteri 17-18, 32, 36 latifrons 32 infraspinatus 17 leiops 35 latifrons 18-21, 31-35, 37 ritteri 36 leiops 17-21, 32-34, 36-37 Xerarionta 256, 264 ritteri 17-21, 30, 32-34, 36-38 waltmilleri 264 triacanthus 18-38 Xenochirus 17, 31-32 Zonitidae 263 CONTENTS OF VOLUME 44 X1n-Luo Cuu, AND Tyson R. Roserts. Cosmochilus cardinalis, a new cyprinid fish from the Lancang-Jiang or Mekong River in Yunnan Province, China. Published August 29, 1985 McCosker, JOHN E. Two new genera and two new species of deepwater western Atlantic worm eels (Pisces: Ophichthidae). Published August 29, 19850000. LerrpertTz, STEVEN L. A review of the fishes of the agonid genus Yeneretmus GilbertabublishedyAucust:29 alo 8 Sine ee ee BRowNn, WALTER C., AND JOHN R. H. GipBons. Species of the Emoia samoensis group of lizards (Scincidae) in the Fiji Islands, with descriptions of two new SPEGlESMRUDIISHeE Cm) Amu allay aes ew lio S.C peseeeeee ater ere ae ee rs ZULLO, Victor A. Quaternary barnacles from the Galapagos Islands. Published | REY op Wee 7 Tle CSS ee KAVANAUGH, Davin H. A systematic review of amphizoid beetles (Amphizoi- dae: Coleoptera) and their phylogenetic relationships to other Adephaga. Pub- lished February 7, 1986 Day, Ava G., AND Retp Moran. Acanthogilia, a new genus of Polemoniaceae from Baja California, Mexico. Published February 7, 19860 Manoop, A. D., G. A. FRYXELL, AND M. McMILLAN. The diatom genus Thalassiosira: Species from the San Francisco Bay system. Published May 6, INSEE sek lle cle a A a a8 OR Sg AL meee ee, Ota oer ace ear WASHINGTON, Betsy B. Systematic relationships and ontogeny of the sculpins Artedius, Clinocottus, and Oligocottus (Cottidae: Scorpaeniformes). Published IVE eayid OSI S 6 rere cee era eee PR SE ar a ca err en ee PARENTI, LYNNE R. Bilateral asymmetry in phallostethid fishes (Atherino- morpha) with description of a new species from Sarawak. Published May 6, 1986 Rotn, Barry. Land mollusks (Gastropoda: Pulmonata) from early Tertiary Bozeman Group, Montana. Published May 6, 1986.0 Pitt, WILLIAM D., MATTHEW J. JAMES, CAROLE S. HICKMAN, JERE H. Lipps, AND Lois J. Pirr. Late Cenozoic marine mollusks from Tuff Cones in the Galapagos Islands4BublishedsMay6s 1:93 6 2a et ee I GYG CESS {iC WYO) ADT 0 OY cer eee eee ta cer ac OE Ree ae a Pages 41-53 55-66 67-109 111-126 127-156 157-223 225-236 237-267 269-282 285-292 i ii - gg rt s 7 _ - (aw ; Ln ie Naa 0rn PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 1, pp. 1-7, 5 figs. August 29, 1985 COSMOGHILUS CARDINALIS, A NEW CYPRINID FISH FROM THE LANCANG-JIANG OR MEKONG RIVER IN LIBRARIES 5 and By Xin-Luo Chu nming Institute of Zoology of Academia Sinica, gfhing, Yunnan Province, The People’s Republic of China YUNNAN PROVINCE, CHINA Tyson R. Roberts California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Asstract: Cosmochilus cardinalis is a large, deep-bodied cyprinid fish recently discovered in southern Yunnan Province in the mainstream of the Lancang-jiang or Mekong River. It is distinguished from all other cyprinids in China and Southeast Asia in having all of its fins bright red; it is further distinguished from the two previously known species of Cosmochilus by having longer barbels, reduced labial papillae, a concave nape, more numerous scales (also more numerous vertebrae?) and nine instead of eight branched dorsal fin rays. INTRODUCTION In the present paper we describe a distinctive new species of large cyprinid fish from the Me- kong River or Lancang-jiang of China. The type- specimens were collected during an ongoing, long- term ichthyological survey of Yunnan Province undertaken by the Kunming Institute of Zoology of Academia Sinica. The Lancang-jiang or Mekong is the largest river in Southeast Asia and probably has the rich- est ichthyofauna of any river in Asia. It is some 4300 km long and arises at an elevation of 5090 m below an enormous glacier on the northern slopes of the Dza-Nag-Lung-Mung or Tanglha Range of the Tibetan highlands of China’s Tsing- hai Province. In Tibet it is known as the Lan- cang-jiang or Dza Chu. It leaves Tibet and enters Yunnan Province at an elevation of about 2800 m, assuming a generally southerly course of near- ly 900 km through mountainous and hilly coun- try of Yunnan. In its upper reaches in Tibet and Yunnan it flows through canyonlike gorges be- tween and parallel to the Salween and Yangtze. In lower Yunnan, where the new species of cyp- rinid was collected, the elevation is only about 500 m. Here the river has a relatively gentle gra- dient and moderate current and is about 150 m wide during the dry season. The bottom is rocky in places but predominantly muddy. The local fishermen know the new species as bia lang or hong chi (“red-finned fish”’). We have identified it as an undescribed Cosmochilus. 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 1 90° 100° \ \ ES ; * i <) ~ * Cardinalis IN a falcifer e harmandi 1 i} ! FiGure |. Cosmochilus Sauvage, 1878 Cosmochilus SAuvAGE, 1878:240 (type-species Cosmochilus harmandt SAUVAGE, by monotypy). DiaGnosis.— Large, deep-bodied and laterally compressed labeoine cyprinids; dorsal fin large and falcate, with 4 simple and 8-9 branched rays; last simple dorsal fin ray greatly enlarged, its pos- terior border more or less strongly serrated for its entire length; anal fin relatively small, with 3 simple and 5-6 branched rays; head relatively small, compressed; snout truncate, without en- larged tubercles or pores; mouth small and in- ferior, its opening transverse; rostral and max- illary barbels large and relatively elongate; lips moderately thick, entirely covered with large, contiguous papillae; horny jaw sheaths trans- verse, moderately thick but with relatively weak cutting edge; gill rakers fleshy, relatively unspe- 110° 120° 130° \ \ \ —20° \ 2 y, DS —10° —0° 0 a g eee S SS —10° | ! | Geographical distribution of the species of Cosmochilus. cialized, 15-18 on first gill arch; pharyngeal teeth triserial, morphologically generalized for Cy- prinidae, usually 1,3,5/5,3,1 or 2,3,5/5,3,2; lat- eral line almost perfectly straight; each lateral line tubule with a short ventroposterior branch terminating in a small pore on exposed portion of posterior shield; scales in lateral line series 35— 48; circumpeduncular scales 16-18; scales ob- long, with relatively huge posterior shields; radi of posterior shield strongly convergent; radii of anterior shield frequently conjoined or bifurcate; vertebrate 35-43. In addition to C. harmandi from the Chao Phrya and Mekong rivers, the genus includes C. Jalcifer Regan, 1906 from the Baram and Rejang rivers in Sarawak and Kapuas in Kalimantan Barat (western Borneo). The distribution of the species is shown in Figure 1. Cosmochilus har- mandi is known to undertake lengthy spawning CHU AND ROBERTS: COSMOCHILUS CARDINALIS, A NEW CYPRINID FISH 3 FiGcure 2 migrations. This is suspected in C. falcifer, and thus may also be characteristic of C. cardinalis. Mekong localities for C. harmandi are from Rainboth et al. 1976. Cosmochilus cardinalis new species (Figures 2-5) Ho.otyre.—KIZ (Kunming Institute of Zoology) 735113, 177 mm (standard length), mainstream of Lancang-jiang near Jinghong, southern Yunnan Province, lat. 21°50’N, long 100°55’E, gill net, May 1973 Paratypes.—KIZ 734079-80, 734082, 735107, 735109-112, 735025, 735030, 735075, 735160, 12: 165-326 mm, same locality and collecting method as holotype, April-May 1973.! DiaGnosis.—A large, deep-bodied, and later- ally compressed Cosmochilus, attaining at least 400 mm standard length. Rostral and maxillary barbels relatively long and thick, dorsal fin high with a falcate margin and elevated base; dorsal fin rays iv9-1/2, last simple ray an elongate stiff spine strongly serrate posteriorly; anal fin rays 1116-1/2. It differs from all or almost all other cyprinids in China and Southeast Asia in having all of the fins including the pectoral and pelvic ' KIZ 735111, 251 mm, has been transferred to the Ichthy- ology Department collection of the California Academy of Sci- ences and is now CAS 55592. Cosmochilus cardinalis, holotype (KIZ 735113, 177 mm). and both caudal lobes entirely bright cardinal red in life. Body dusky dorsally, silvery or whitish on sides and abdomen. Opercle golden. Scales in lateral series 46-48, between dorsal fin and lat- eral line 9-10, between lateral line and pelvic fin origin 5, circumpeduncular 16-18. Vertebrae 24 + 19 = 43 (holotype). Head relatively small and laterally com- pressed, its length 4.0—4.5; dorsal profile of head to nape moderately sloped, then abruptly steeper at nape until dorsal fin origin. Snout 2.7—3.2 in head, eye 4.1—5.3 in head, interorbital width 2.4— 2.7 in head. Eye with a narrow gelatinous rim or hyaline eyelid. Barbels thick and relatively long; anterior or rostral barbel extending posteriorly almost to below posterior border of eye, its length 1.8-2.2 in head; posterior or maxillary barbel somewhat longer, extending posteriorly almost to pectoral fin, its length 1.4—1.7 in head. Mouth subinferior and moderately wide, extending pos- teriorly to directly below anterior margin of eye or slightly farther. Rostral cap well developed with deeply incised rostral groove complete be- tween rostral barbels of either side. Sublacrimal groove also well developed and deeply incised, extending from rostral barbel to posterior end of jaws. Lips moderately well developed; upper lip FiGure 3. well defined, entirely separate from rostral cap and upper horny jaw sheath; lower lip with oral margin separated from horny jaw sheath only by a shallow groove, and with posterior margin well defined laterally but entirely interrupted for transverse portion of lower jaw; upper and lower horny jaw sheaths well developed, with a broadly rounded surface and very weakly developed transverse grooves or sulci. Anterior margin of lower jaw truncate. Rostral cap and horny jaw sheaths relatively smooth; lips and oral epithelium (including gular flap) covered with large, close-set or contiguous but low-lying papillae (probably unculiferous) comparable in distribution and basic morphol- ogy to the greatly enlarged and elevated contig- uous papillae characteristic of the other two species of Cosmochilus. Gill openings relatively broad, isthmus nar- row; upper portion of gill cover with posterior margin very slightly concave. First gill arch (Fig. 3) with 5 + 11 or 12 = 16 or 17 gill rakers on PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 1 Cosmochilus cardinalis, holotype. First gill arch (lateral view; bony supports of gill rakers in black). anterolateral margin and 0 + 11 gill rakers on posterodorsal margin in holotype; paratypes with 17-19 anterolateral gill rakers on first gill arch. Gill rakers all relatively short and fleshy, those on lower limb of gill arches more or less trian- gular in shape with broad bases; posterior gill rakers similar in shape to anterior rakers but relatively somewhat larger. Pharyngeal bones (Fig. 4) strongly arched; dorsal edentulous limb and dorsal half of toothbearing portion thick and evenly curved; external ala moderately broad; edentulous lower limb and ventral half of tooth- bearing portion below external ala with concave lateral margin. Entire medial surface of lower limbs of either pharyngeal bone forming a broad symphysis. Pharyngeal teeth basically uncinate, 2,3,5/4,3,2 in holotype, those of principal row compressed. Scales (Fig. 5) large, slightly longer than high, and relatively numerous. Anterior margin rela- tively straight but with a well defined convex median projection; posterior margin broadly CHU AND ROBERTS: COSMOCHILUS CARDINALIS, A NEW CYPRINID FISH 5 \ 5 mm | Ficure 4. rounded; radii most numerous and longest on posterior field, least numerous and shortest on lateral fields; radii near central portion of pos- terior field strongly convergent. Lateral line com- plete and nearly straight. Scales of lateral line with straight primary tubules and a single small pore arising from a short ventroposteriorly di- rected secondary tubule originating from ante- rior half of primary tubule. Scales in lateral line series 46-48, 9-10 above and 5 below lateral line; predorsal scales 24-26, circumpeduncular 16-18. Two or three scale rows extend on caudal fin base posterior to last pored scale of lateral line series. Scales between vent and anal fin or- igin 2. Circumferential scales 35-37 (18-20 dor- sal + 2 lateral line + 15 ventral). Dorsal fin origin near middle of body, some- what closer to snout tip than to caudal fin base. Anal fin origin distinctly posterior to vertical through base of last dorsal fin ray. Pelvic fin or- Cosmochilus cardinalis, holotype. Pharyngeal jaws (dorsal view). igin well in advance of vertical through dorsal fin origin. Base of dorsal fin with well developed, strongly convex scale sheath comprising three somewhat irregular rows of large scales (of which the middle row is somewhat smaller); anal fin scale sheath only slightly convex, with one or two rows of scales. Pelvic fin with two moder- ately elongate axillary scales. Dorsal fin spine length 3.0-3.7, with 20 serrae in holotype. Anal fin much smaller than dorsal fin, with last simple and first two branched rays slightly prolonged to form a lobe. Pectoral and pelvic fins almost equal in shape and size, pectoral fin extending poste- riorly almost to pelvic fin origin and pelvic fin reaching or almost reaching vent; pectoral fin length 3.8—4.3, pelvic 4.0—4.3. Pectoral fin rays usually 15, pelvic 9. Caudal fin deeply forked, upper and lower lobes nearly equal in length and shape, with slightly pointed tips. Abdomen anterior to and between pelvic fins 5 mm FiGure 5. Cosmochilus cardinalis, holotype. Scales. Below, 30th scale in lateral line scale series. Above, scale two scale rows about 30th scale in lateral line scale series. Dashed lines indicate areas of overlap by scales neighboring exposed portion of posterior shield. somewhat flattened from side to side; abdomen posterior to pelvic fins rounded or very slightly (almost imperceptibly) carinate. Body depth 2.4. Caudal peduncle laterally compressed and mod- erately deep, its length 6.1-7.9 and depth 6.4—- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 1 7.2. Dorsal surface of head and lateral surface of lacrimal region more or less uniformly covered with fine, widely spaced granular tubercles; stronger tuberculation or tubercles on fins not observed in specimens of either sex. Swim blad- der with two chambers. Intestine about three times as long as body. COMPARISON WITH OTHER SPECIES OF COSMO- cHILUS.—The basically similar morphology of the scales, scaly fin sheaths, fleshy gill rakers, pharyngeal teeth, and papillose lips of C. car- dinalis, C. harmandi, and, so far as known, C. falcifer, leads us to conclude that the three are correctly placed together within Cosmochilus. Our new species differs from C. harmandi and C. falcifer in having more numerous scales (44— 48 vs. 36-39 in lateral series), three instead of two scale rows in scaly sheath on dorsal fin base, 9 instead of 8 branched dorsal fin rays, dorsal profile strongly concave at nape (vs. relatively evenly sloped), more elongate barbels, labial pa- pillae much reduced in size, and all fins bright red. It additionally differs from C. harmandi in having 6 instead of 5 branched anal fin rays, a less pointed snout viewed from above or from the side, and 43 instead of only 35 vertebrae (number of vertebrae unknown in C. falcifer). Cosmochilus cardinalis also differs from C. fal- cifer in having the fourth simple dorsal fin ray relatively less elongate, very straight, and with large, strong serrae on its posterior margin. In C. falcifer this ray is exceptionally elongate (some- times extending posteriorly to caudal fin base when adpressed), very strongly curved, and with very weak serrae on its posterior margin. Life color of C. harmandi is recorded as back rich pale blue, dorsal and caudal fins black-edged; in some specimens anal fin with black tip (Smith 1945:132). We note that the dorsal fin may also be black-tipped. Life color of C. falcifer has not been reported previously. A fresh 316-mm spec- imen caught in the Kapuas and photographed in the market at Sintang by the junior author had overall color white or milk white, especially ven- trolaterally and ventrally; dorsolaterally and dor- sally distinctly brownish or violaceous brown; posterior margins of scales, especially on upper parts of body, with broad dark margins; dorsal surface of head faintly yellowish and entire gill cover distinctly yellow; iris and ventral portion of head milk white; entire dorsal fin rosy pink or faintly orangish except black at tip; pectoral fin CHU AND ROBERTS: COSMOCHILUS CARDINALIS, A NEW CYPRINID FISH 7 white, pelvic white or pinkish; anal and caudal fins dusky, caudal very dark, its posterior margin almost black. In conclusion, C. harmandiand C. falcifer seem to be much more similar to each other than either of them is to C. cardinalis. RELATIONSHIPS OF COSMOCHILUS Sauvage (1878:240) stated that Cosmochilus is related to Labeo but did not discuss this assess- ment. The pattern of strongly convergent radii on the posterior shield and conjoined or bifur- cating radii on the anterior shield of the scales of Cosmochilus is seen in relatively few Chinese cyprinids (Chu 1935). Those in which this pat- tern is most clearly present are species of Osteo- chilus, Sinilabeo, and Garra (Chu 1935, pls. 15- 16). These genera are referable to the cyprinid subfamily Labeoinae. We have observed scales with a similar pattern of radii in various non- Chinese Labeoinae including Morulius chryso- phekadion and various African and Asian species currently assigned to Labeo, but not in genera belonging to other subfamilies. This suggests that Cosmochilus might indeed belong in Labeoinae. On the other hand, the general morphology of Cosmochilus, including the relatively simple mouth parts; deep, laterally compressed body; and elongate, serrate dorsal fin spine suggest re- lationship to a group of Southeast Asian barbels including Cyclocheilichthys. A serrate dorsal fin spine 1s unknown in the Labeoinae. ACKNOWLEDGMENTS We wish to express our gratitude to Walter J. Rainboth, who first noted the similarity of the new species to Cosmochilus, Susan Middleton for photographing the holotype, George Zorzi and David Catania for preparation of radio- graphs, and Reeve M. Bailey and Douglas W. Nelson for loaning specimens of C. harmandi. The manuscript was typed by Francis Bertetta. LITERATURE CITED Cuu, Y. T. 1935. Comparative studies on the scales and on the pharyngeals and their teeth in Chinese cyprinids, with particular reference to taxonomy and evolution. Biol. Bull. St. John’s Univ. 2, x + 226 pp., 30 pls. RainsotH, W. J., K. F. LAGLerR, AND S. Sontirat. 1976. Maps of freshwater fish distribution in the lower Mekong basin. Mekong Secretariat Working Document 31, xv + 406 pp. Sauvace, H. E. 1878. Note sur quelques poissons d’espéces nouvelles provenant des eaux douces de I’Indo-Chine. Bull. Soc. philomath. Paris 7(2):233-242. Smitu, H. M. 1945. The freshwater fishes of Siam or Thai- land. Bull. U.S. Nat. Mus. 188, xii + 622 pp., 9 pls CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 CASES PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENC Vol. 44, No. 2, pp. 9-15, 6 figs. August 29, 1985 TWO NEW GENERA AND TWO NEW SPECIES OF DEEPWATER WESTERN ATLANTIC WORM EELS (PISCES: OPHICHTHIDAE) By John E. McCosker California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Apstract: Two new genera and two new species of Atlantic worm eels, family Ophichthidae, subfamily Myrophinae, tribe Myrophini, are described and illustrated. Mixomyrophis pusillipinna, gen. nov., sp. nov., trawled from deepwater off the Lesser Antilles, is an elongate species with uniserial conical teeth, a labial posterior nostril, a minute pectoral fin, and 178 vertebrae. Asarcenchelys longimanus, gen. nov., sp. nov., collected off Belem Brazil, is an elongate species with biserial conical teeth, a labial posterior nostril, well- developed pectoral fins, and 149 vertebrae. Osteological characteristics of the new genera are described from radiographs and compared with those of related myrophines. INTRODUCTION The ophichthid worm eels of the subfamily Myrophinae (sensu McCosker 1977) occupy a variety of sand and mud habitats as well as the midwater environment, ranging from the shal- low intertidal to depths of 400 fathoms or more. The shallow-water species of the genera Myro- phis, Muraenichthys, and Ahlia are common in collections and are probably abundant within their milieu. The deeper water species are rare, being difficult to trawl or dredge, and are often known from but a single specimen. While preparing the ophichthid eel section of The Fishes of the Western North Atlantic (FWNA), the late James E. B6hlke discovered a single specimen of a new species of myrophine collected by trawl off Anguilla, Lesser Antilles. In taking over the completion of the FWNA proj- ect, I discovered another new species of deep- water myrophine, from Brazil, which is also ge- nerically distinct. I intend to make these [9] taxonomic names available for the FWNA vol- ume and to describe the significant osteological characters that are visible by radiographic ex- amination. It is my hope that subsequent spec- imens will be discovered, which will allow a more thorough osteological examination and compar- ison with other myrophine genera. MATERIALS AND 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 calipers (all other measurements) and recorded to the nearest 0.1 mm. Body length comprises head and trunk lengths. Head length was mea- sured from the snout tip to the posterodorsal margin of the gill opening; trunk length was taken from the end of the head to mid-anus; maximum body depth did not include the median fins. Ver- tebral counts (which included the hypural) were 10 oe PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 2 % LAS is Ficure 1. of Mixomyrophis pusillipinna McCosker, sp. nov. taken from radiographs. Stained and cleared gill arches were prepared using the Taylor (1967) trypsin technique. Institutional abbreviations of material examined are explained in the acknowl- edgments section of this paper. Mixomyrophis McCosker, gen. nov. Tyre species. — Mixomyrophis pusillipinna McCosker, sp. nov. DiaGcnosis.—An elongate myrophine, tribe Myrophini, with tail longer than head and trunk, laterally compressed, particularly posteriorly; snout subconical, broad from above, not grooved ventrally; anterior nostril tubular, posterior nos- tril on outer edge of lip and covered by a flap; dorsal fin origin in mid-trunk; pectoral fin a mi- nute flap in posterodorsal corner of upper gill opening; eye large, behind middle of jaw; third preopercular pore present; head and lips smooth, without cirri or lappets; teeth of jaws and vomer small, conical, uniserial, and close set, with slightly retrorse tips; gill arches well developed for a myrophine, first basibranchial ossified, up- per pharyngeal toothplates fused; neurocranium stout, slightly sloping posteriorly; suspensonum anteriorly inclined; pterygoid stout, not bracing maxilla; maxillae elongate, tapering posteriorly; opercular series apparently moderately devel- oped; pectoral girdle reduced to a slender clei- Holotype of Mixomyrophis pusillipinna McCosker, sp. nov., ANSP 152305, 407 mm TL. Inset: Head of holotype thrum and supracleithrum; epipleural ribs on all precaudal vertebrae; caudal transverse processes apparently absent; caudal vertebrae more nu- merous than precaudal. Other characteristics those of single species. EtyMoLoGcy.—From the Greek és, mixis, a mixing, and Myrophis (masculine), a genus of ophichthid eel. Named in reference to the com- bination of myrophine characters that this eel possesses. Mixomyrophis pusillipinna McCosker, sp. nov. (Figures 1, 2, 6b) Hototype.—ANSP 152305 (originally UMML 30290), 407 mm, a female with ripening ovaries, captured off Anguilla, Lesser Antilles (18°26.4'N, 63°12.6’W to 18°28'N, 63°11.1'W), by 10-m otter trawl, between 393-451 m depth, by the RV Pillsbury, sta. 984, on 22 July 1969. COUNTS AND MEASUREMENTS (IN MM).—Total length 407; head length 36.5; trunk length 115.5; tail length 255; body depth at gill openings 9.0; body width at gill openings 7.1; body depth at anus 8.8; body width at anus 6.0; gill opening 1.5; snout tip to origin of dorsal fin 94; left pectoral fin length 1.0; snout length 7.9; upper jaw length 12.1; eye diameter 2.9; fleshy interorbital distance 4.2 Total vertebrae 178; predorsal ver- tebrae 33; preanal vertebrae 57. DESCRIPTION. — Body elongate, its depth 45 in total length (TL), laterally compressed in tail re- gion. Head and trunk 2.7 and head 11.2 in TL. Snout subconical, broad as seen from above; lower jaw included, its tip reaches the anterior McCOSKER: TWO NEW SPECIES OF ATLANTIC WORM EELS 11 nostril bases. Anterior nostrils tubular, directed ventrally, their anterior edge looped upward; posterior nostril at outer edge of lip, covered by a flap. Eye large, its anterior edge behind midpoint of upper jaw. Gill opening mid-lateral, a constricted open- ing. Median fins low, lying partially within a groove, but elevated above last 20 vertebrae, meeting each other and extending beyond caudal tip. Dorsal fin arises above posterior trunk region. Pectoral fin minute. Head pores developed. Single temporal and interorbital pores. Six pores along left mandible; 5 along right. Two pores between anterior and posterior nostrils. Four supraorbital pores. Three preopercular pores. Left lateral line pores ca. 149; 9 above branchial basket; 57 before anus. Teeth small, conical, nearly uniform in size. An intermaxillary chevron of 8 teeth, followed by 2 on each side, closely adjoining ca. 20 uni- serial vomerine teeth and 25 maxillary teeth. Ap- proximately 30 uniserial mandibular teeth, with a secondary pair at symphysis. Gill arches removed, stained and cleared. Ba- sibranchial | ossified, basibranchials 2—4 absent. Hypobranchials 1 and 2 ossified; hypobranchial 3 cartilaginous. Ceratobranchials 1-4 ossified; ceratobranchial 5 absent. Infrapharyngobran- chials 2 and 3 ossified. Lower tooth plate small, with 2 rows of conical teeth, medial row largest. Upper pharyngeal tooth plate fused, subrectangular, with 4-5 rows of conical teeth, medial row largest. Body color in isopropyl alcohol yellow on head, chin, tail, and dorsal surface of trunk. Throat and belly whitish. Finely peppered throughout body and tail with small brown specks. Peritoneum black. EtyMoLoGy.—From the Latin pusillus, puny or insignificant, and pinna, fin, to be treated as a noun in apposition. ReEMARKS.— Mixomyrophis is separable from all other myrophines by the combination of its minute pectoral fin, elongate body, and posterior nostril located within the outer lip. Mixomyro- phis appears most similar to the elongate species of Pseudomyrophis, which differ by having the posterior nostril before the eye, more extreme body elongation, a reduced and rounded neu- rocranium (cf. Fig. 6b and 6d), and reduced gill fi Ficure 2. Dentition of holotype of Mixomyrophis pusil- lipinna McCosker, sp. nov., ANSP 152305. arch components (Bohl ke 1960; McCosker 1977). The nostril condition of Pseudomyrophis appears to be a slight posterodorsal translocation of the opening from its location along the lip (although it is difficult to interpret which state might be the primitive condition), and the other characters seem to be advanced specializations. The nearly bulbous snout, large eye, and body elongation of Mixomyrophis are conditions shared by other deepwater myrophines such as the species of Pseudomyrophis, Neenchelys, Asarcenchelys lon- gimanus, and Muraenichthys puhioilo McCosker (1979). 12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 2 Ficure 3. mm TL. The actual specimen is intact, but badly torn in the anterior trunk region. Asarcenchelys McCosker, gen. nov. Tye species. — Asarcenchelys longimanus McCosker, sp. nov. DiaGnosis.—A very elongate myrophine, tribe Myrophini, with tail longer than head and trunk, laterally compressed throughout trunk and tail; snout subconical, tumid, not grooved ventrally; anterior nostril tubular; posterior nostril on outer edge of lip and covered by a flap that is incised posteriorly; dorsal fin origin in anterior trunk region; anal fin elevated; pectoral fin lanceolate, well developed, slightly longer than snout; eye large, behind middle of jaw; third preopercular pore present; head and lips smooth, without cirri or lappets; teeth of jaws and vomer large, not close set, conical and slightly recurved; teeth bi- serial anteriorly in jaws and vomer, outer row smaller; gill arches appear to be well developed for a myrophin; neurocranium stout, truncate posteriorly; supraoccipital crest developed; sus- pensorium posteriorly inclined; maxillae taper posteriorly; pectoral girdle reduced to stout Reconstructed appearance of holotype of Asarcenchelys longimanus McCosker, sp. nov., MNHN 1968-215, 277 cleithrum and thin supracleithrum; epipleural ribs present only on anterior trunk vertebrae; caudal temporal processes apparently absent; caudal vertebrae more numerous than precaudal. Other characteristics those of single species. EtyMoLoGcy.—From the Greek acapxos, asar- kos, lean, and evxedvs, enchelys, eel (treated as feminine according to Opinion 915 of the Bul- letin of Zoological Nomenclature, 1970), in ref- erence to its emaciated appearance. Asarcenchelys longimanus McCosker, sp. nov. (Figures 3-5, 6c) Hototyrpe.—MNHN 1968-215, 277 mm, sex undeter- mined, captured near Belém, Brazil, at 5S m depth by P. Four- manoir, September 1966. ParATyPeE.—MNHN B. 2994, 147 mm, sex undetermined, collected with the holotype. COUNTS AND MEASUREMENTS (IN MM). — Data for the paratype parenthetically follow those of the holotype. Total length 277 (147); head length 27 (18); trunk length 78 (46); tail length 172 (83); body depth behind gill openings 3.8 (~2); body width behind gill openings 3.6 (1.4); body depth at anus ~1.5 (~2); Ficure 4. Head of holotype of Asarcenchelys longimanus McCosker, sp. nov., MNHN 1968-215. McCOSKER: TWO NEW SPECIES OF ATLANTIC WORM EELS 13 body width at anus 1.5 (1.2); gill opening 1.4 (0.9); snout tip to dorsal fin origin 55 (34); left pectoral fin length 4.0 (3.3); snout length 3.8 (3.7); upper jaw length 6.6 (5.8); eye diameter 1.4 (1.3); fleshy interorbital distance 1.6 (1.3). Total vertebrae 148 (131, tail incomplete); predorsal vertebrae 27 (27); preanal vertebrae 53 (55). DEscrIPTION.— Body very elongate, its depth 72.9-73.5 in TL, laterally compressed behind head. Head and trunk 2.3-2.6, and head 8.2- 10.3 in TL. Snout subconical, bulbous; lower jaw includ- ed, its tip reaches to front of anterior nostril bas- es, leaving several intermaxillary teeth exposed. Anterior nostrils tubular, directed ventrally; pos- terior nostril at outer edge of lip, covered by a flap whose posterior edge is incised. Eye large, anterior edge of orbit above middle of upper jaw. Gill openings mid-lateral, not as constricted as those of most myrophines, about equal in length to isthmus. Dorsal fin low, arising in anterior trunk region. Anal fin elevated. Median fins expanded in pos- terior tail region, extended beyond caudal tip. Pectoral fin lanceolate, broad based, well devel- oped for a myrophine. Head pores developed, much more apparent than those of lateral line. Single temporal and interorbital pores. Five pores along mandible, widely spaced posteriorly. Two pores between anterior and posterior nostrils. Four supraorbital pores. Three preopercular pores. Lateral line pores difficult to discern; 14 above branchial basket. Teeth conical, fairly large for a myrophine, not close set, nearly uniform in size, recurved. An intermaxillary chevron of 6 teeth, visible when mouth is closed, followed by closely abutting vo- merine dentition consisting of 3—4 pairs of teeth and a uniserial row of 13 teeth. Maxillary teeth biserial anteriorly, with an inner row of 6 teeth and an outer row of 22-24 smaller teeth. Lower jaw biserial anteriorly, with an inner row of 5 teeth and an outer row of 23-25 smaller teeth. Gill arches, as viewed from radiograph, appear to be myrophin-like and not reduced. First ba- sibranchial is ossified, others appear to be car- tilaginous or absent. Hypobranchials | and 2 os- sified; hypobranchial 3 appears cartilaginous or absent. Ceratobranchials 1-4 ossified; cerato- branchial 5 not apparent. Upper and lower tooth plates appear to have 2-3 rows of conical teeth; upper plate appears to be fused. Ficure 5. Dentition of holotype of Asarcenchelys longi- manus McCosker, sp. nov., MNHN 1968-215. Body coloration in isopropyl alcohol cream to white, numerous fine, chocolate-brown spots overlying snout, dorsal surface, and area behind eye. All fins transparent. Peritoneum light col- ored. EtyMoLoGy.—From the Latin /ongus, long, and manus, hand, to be treated as a noun in apposition. Named with reference to the elongate pectoral fins. REMARKS.— This new myrophine is separable from all related ophichthids by a combination of internal and external morphological charac- 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 2 Ficure 6. Enlarged radiographs of neurocrania of selected myrophins: A) Myrophis vafer, CAS 17823, 220 mm TL. B) Mixomyrophis pusillipinna, ANSP 152305, 407 mm TL. Gill arches have been removed. C) Asarcenchelys longimanus, MNHN 1968-215, 277 mm TL. D) Pseudomyrophis micropinna, CAS 50978, 109 mm TL. ters. Particularly significant are its well-devel- oped pectoral fins, posterior nostril located on the edge of the lip, elongate body and tail, and elongate dentition. The body elongation, anal fin development, nostril location, and snout shape are not unlike those of certain species of Neenchelys and Pseudomyrophis (cf. McCosker 1982: Smith and Bohlke 1983), and are probably McCOSKER: TWO NEW SPECIES OF ATLANTIC WORM EELS 15 adaptive for living in deepwater soft benthic hab- itats. Its close affinities, however, lie with the species of Myrophis, which share with it the de- rived character state of having lost epipleural ribs beyond the fifteenth vertebra, a condition shared as well with Ahlia egmontis. Asarcenche- lys longimanus also shares with the species of Myrophis the primitive states of neurocranial shape (Fig. 6a, 6c), gill arch condition, and pec- toral fin development. Species of Pseudomyro- phis and Neenchelys are further specialized and separable from the “Myrophis group” in having a much-reduced neurocranium, and posterior nostrils before the eye and lacking a flap (Mc- Cosker 1977, 1982). It should be noted that both specimens of 4. longimanus are damaged and thereby the total length measurement of each specimen may be in error by a few percent. The paratype is intact, but the radiograph indicates that the tail has probably been severed and regrown. The speci- men has 17 fewer vertebrae than the holotype. During capture the holotype was broken behind vertebra 15 and is twisted in preservative. The head remains attached by the skin to the trunk region and is sufficiently intact to allow precise measurements to be taken and characters to be analyzed. ACKNOWLEDGMENTS The curators of several institutions generously supplied information and specimens used in this study. I thank the ichthyological staffs of: Acad- emy of Natural Sciences of Philadelphia (ANSP), British Museum (Natural History) (BMNH), California Academy of Sciences (CAS), Paris Museum of Natural History (MNHN), Scripps Institution of Oceanography (SIO), Uni- versity of Miami Marine Laboratory (UMML), and the United States National Museum of Nat- ural History (USNM). Special assistance was supplied by Eugenia B. B6hlke (ANSP), Marie- Louise Bauchot (MNHN), and C. Richard Ro- bins (UMML). | also wish to thank: Mary Fuges and Amy Pertschuk for the preparation of illus- trations, Susan Middleton for photographic as- sistance, Lillian J. Dempster and W. I. Follett for nomenclatural assistance, and Eric Anderson for critically reading a draft of this manuscript. LITERATURE CITED BOu ke, J. E. 1960. A new ophichthid eel of the genus Pseu- domyrophis from the Gulf of Mexico. Notul. Nat., No. 329: 1-8. McCosker, J. E. 1977. The osteology, classification and re- lationships of the eel family Ophichthidae. Proc. Calif. Acad. Sci., Ser. 4, 41:1-123. 1979. The snake eels (Pisces, Ophichthidae) of the Hawaiian Islands, with the description of two new species. Proc. Calif. Acad. Sci. 42(2):57-67. . 1982. A new genus and two new species of remark- able Pacific worm eels (Ophichthidae, subfamily Myrophi- nae). Proc. Calif. Acad. Sci. 43(5):59-66. Situ, D. G., AND J. E. BOHLKE. 1983. Neenchelys retropin- na: a new worm eel (Pisces: Ophichthidae) from the Indian Ocean. Proc. Acad. Nat. Sci. Philadelphia 135:80-84. Taytor, W. R. 1967. An enzyme method of clearing and staining small vertebrates. Proc. U.S. Natl. Mus. 122(3596): 1-17. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 oy 2) Sot ironeii. te Rader pi hind Ww ae = vite Tabi | wit nr CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 3, pp. 17-40, 18 figs., 6 tables. August 29, 1985 A REVIEW OF THE FISHES OF THE AGONID GENUS XENERETMUS GILBERT By Steven L. Leipertz School of Fisheries WH-10, University of Washington, Seattle, Washington 98195 Asstract: The agonid genus Xeneretmus is reviewed and found to be composed of two subgenera: Xeno- pyxis, containing X. latifrons, X. leiops, and X. ritteri; and Xeneretmus, containing only X. triacanthus. The osteology of the type species of the genus, X. triacanthus, is described, illustrated, and compared with the other members of the genus, as well as to members of the agonid genera Agonus, Hypsagonus, and Podothecus. On the basis of a comparison with 15 other agonid taxa, the subgenera Xeneretmus and Xenopyxis are demonstrated to be monophyletic. A key is provided, along with synonymies, diagnoses, and descriptions for the genus, the subgenera, and species. Lectotypes are designated for X. triacanthus and X. latifrons. INTRODUCTION The family Agonidae is composed of typically small, benthic, scorpaeniform fishes that are al- most totally encased in rows of overlapping der- mal plates; the centers of these plates often bear spines or protuberances. The majority of the species are found in the North Pacific Ocean and Bering Sea; only 3 of the approximately 50 rec- ognized species (distributed among some 20 cur- rently recognized genera) are restricted to other regions: two in the North Atlantic Ocean (As- pidophoroides monopterygius and Agonus cata- phractus), and one off southern South America (Agonopsis chiloensis). Only two major reviews of the family have been written. The first is that of Jordan and Ev- ermann (1898). While this work was primarily concerned with American species, all known agonids were considered; little osteology was dis- cussed, and only a very few specimens of each species were examined. The second review of the family is that of Freeman (1951), a widely known, but unpublished doctoral dissertation written at (17] Stanford University. Once again, little osteology was examined, and few specimens were used. Only two notable osteological investigations of agonids have been published: Rendahl’s (1934) work on Hypsagonus quadricornis and Ilina’s (1978) work on the genera Podothecus and Agon- us. Both were limited to aspects of cranial os- teology. The taxonomic history of the genus Xeneret- mus began with Gilbert’s (1890) erection of Xen- ochirus, established to contain three species: Y. triacanthus, X. pentacanthus, and X. latifrons. Five years later, Gilbert (1895) described a fourth species, Xenochirus alascanus. In 1903 Gilbert (in Jordan 1903) became aware of the prior use of the name Xenochirus by Gloger (1842) for a genus of marsupial mammals, and therefore of- fered Xeneretmus as a replacement name. In the following year, a fifth member of the genus, Y. infraspinatus, was described by Gilbert (1904). In his final paper on this genus, Gilbert (1915) described two additional species, XY. ritteri and X. leiops, moved X. alascanus, X. infraspinatus, 18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 and X. pentacanthus into a new genus, Astero- theca, and created two subgenera within Yeneret- mus: Xenopyxis containing X. latifrons, X. leiops, and X. ritteri; and Xeneretmus containing only X. triacanthus. Jordan et al. (1930), without ex- planation, raised the subgenus Xenopyxis to ge- neric status. Although a few later workers (Barn- hart 1936; Clemens and Wilby 1961) followed Jordan et al. (1930), the majority of recent work- ers (Freeman 1951; Peden and Gruchy 1971; Miller and Lea 1972; Hart 1973; Barraclough and Peden 1976; Robins 1980; Eschmeyer et al. 1983) retained Gilbert’s (1915) classification. Since Gilbert (1915), only minor publications on the genus have appeared. Bolin (1937), while noting that Gilbert’s (1915) labels on the illus- trations of X. ritteri and X. leiops were switched, extended the geographic range of X. /eiops north from Santa Catalina Island to Monterey Bay. Pe- den and Gruchy (1971) expanded the range of X. triacanthus into British Columbian waters. Ginn and Bond (1973) extended the range of ¥. leiops north to the Columbia River. Three years later, Barraclough and Peden (1976) extended the range of XY. /eiops further north to southern British Columbia and noted, as did Bolin (1937), the switching of Gilbert’s (1915) labels. The purposes of this study are to provide a complete osteological description of the genus Xeneretmus, to describe variation in a number of systematically important characters, to des- ignate type material for the species where it is in question, and to investigate the phyletic nature of the genus, subgenera, and closely related taxa. MATERIALS AND METHODS All measurements were taken from the right side of the fish. Standard length (SL), used throughout, was measured from the tip of the snout to the posteroventral corner of the last su- pralateral plate. Other measurements were made as follows: Anal, first dorsal, and second dorsal lengths. — from the tip of the snout to the insertion of the first ray of the respective fin Caudal peduncle length.—from the insertion of the posteriormost anal ray to the posteroven- tral corner of the last supralateral plate Vent length.—from the tip of the snout to the anterior margin of the anal opening Ventral head length.—from the tip of the snout to the posterior margin of the isthmus Depth at first and second dorsal.—the shortest distance from the first ray of that fin to the ventral contour of the body Head length.—from the tip of the snout to the posteriormost margin of the opercular mem- brane Supraoccipital pore to snout.—from the tip of the snout to the anterior edge of the acoustico- lateralis pore located dorsal to the supraoccipi- tal bone Snout length.—from the tip of the snout to the anterior margin of the orbit Upper jaw length. — from the anteriormost extent of the premaxilla to the posteriormost margin of the maxilla Length of orbit.—greatest distance between the rims of the orbit Interorbital width.—least distance between the lateral margins of the frontals Length of pectoral and pelvic fins.—from the base of the longest ray to its tip Caudal depth. —least depth of the caudal pedun- cle Pectoral width.—the greatest width measured between the pectoral bases Ural centra are included in vertebral counts ob- tained from radiographs. Nomenclature for, and the method of enu- meration of, dermal plates follow the system out- lined by Gruchy (1969) with the following ad- ditions: the number of plates anterior or posterior to a fin is counted to or from, but not including, the plate on which the first or last ray of the fin is inserted; the ventrolateral series of plates is considered to start at the pelvic fin base. Cladistic analysis was performed using Joseph Felsenstein’s (Department of Genetics, Univer- sity of Washington) Package for Inferring Phy- logenies. Summary statistics were calculated us- ing SPSS and SCSS (Nie et al. 1975, 1980). All programs were run on a Digital Electronics Cor- poration VAX, under the VMS operating system. For the phylogenetic analysis, characters from the following in-group were recorded: Aspido- phoroides bartoni, A. olriki, Bathyagonus alas- canus, B. infraspinatus, B. nigripinnis, B. pen- tacanthus, Bothragonus swani, Odontopyxis trispinosa, Xeneretmus latifrons, X. leiops, X. rit- teri, and X. triacanthus. A close relationship of these taxa was hypothesized by Freeman (1951); all are members of his subfamily Xeneretminae. A set of morphological characters (Table 1) was WD) LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS Y¥ENERETMUS GILBERT 1yatys 2s00] so ways jussqe Sok Wyss juasqe ou ou asoo]| SoA ou juasqe sok yonur =. yuasqe ou yonul 9800] Sod yonu 2s00] sod yonw = yuasqe ou yonw = quasqe ou yonur =. juasqe ou 1yars yan sok yonuw 1430 SoA yonuw 9S00] Soh ou as00] sak yonuw 1yan SoA yonuw 1y3n Soh ou aso] SoA ou 1y3n Sod skel saiejd sun esoised = yaoyd—s -1Nqe. [e1U9A jo saiejd jouon juow jseaig -oafoid -aduReLy oytiasuy poua “yoryy jo quour -dojaaog quasoid juasoid juasaid quasqe juasoid quasoid quasaid quasoid quasaid quasoid quasoid juasaid quasqe quasoid quasaid juasaid quasoid quasqe juasqe soutds aieid Apoq jeuwog quou juasqe ouou juasqe auou juasqe auou juasqe auou quasqe guou juasqe auou quasqe deiy) = Juasaid quo =. }uasaud €-I uasaid ouo =.» }uasoid auo =. "1}uasaid guou = juasaid aay = yuasaid aay = yuasaid dag. quasaid aay: yuasaid auou =: juasaid auou = juasaid souids aed ajejd = [eso jeusoy pasodxq "YXWT GINOOY G3aLI9IT9S WOW SHAILOVUYYVH’) AO NOLLNAIYLSICT Inoj Ol= ouo0 ouou OM) 9u0 2a1y) om} OM) 9u0 ouo0 92u0 9u0 OM} OM} OMI OM} auo ouo sjaqueq AreyIXe quasaid juasqe quasaid quasaid quasoid quasoid quasaid quasqe quasoid quasoid quasoid juasqe juasqe juasqe juasqe quasoid juasqe quasaid quasoid snuryist jo PlOJ-9914 -nosedo OM} ouo OM) OM) OM} OM) OM) OM) auo auo auo quasqe juasqe OM} OM} OM) OM} quasqe quasqe souids Ie] -31d quasaid quasoid juasqe juasqe juasqe juasqe juasoid quasqe juasqe quasqe juasqe juasqe juasqe juasqe quasqe juasqe juasqe juasqe juasqe souids prowye -eidns posodxq OM) OM] auo 9u0 ouo 9uo0 OM} 9u0 9uo0 ouo 9u0 9u0 9u0 ouo 9uo0 ouo0 9u0 9uo0 9u0 souids yesen quasoid juasoid juasaid quasqe quasqe quasaid quasaid juasaid juasoid juasaid quasaid juasoid quasqe quasoid quasaid quasoid quasoid quasqe juasqe souids [e1u01y SNIDUIA AOPUIDS: snulasuadiop snaayjopod snouodvl s1240g pDq4vq DUISD]]0d uospav2apop DIJ2IO siusooluponb snuospsda psjna sisdouos py SNYIUDIVIAT snudjasaUay Malytd SNULJOIIUAY sdo1a] snujasaUay suodfijD] SNULJALIUIX psouldst} sixddojuopo 1UDMS snuosp1ylog snyjuvopjuad snuospdylog siuuidusiu snuospAylog snjouidsp.fur snuospAyog snuDosp]D snuospAylng 1y14]0 sapiosoydopidsy 1u0]40q sapiosoydopidsp saisadg quasoid = yuasaid quasqe = juasaid = juasoid quosoid = =juasoid = juasaid = juasoid = juasaud juasqe quasqe quasqe = juasoid = juasoid juasqe quasqe juasqe juasqe juosqe juasqe quasqe quasqe quasqe quasqe quasoid juasqe quosqe juasoid juasqe quasoid = juasaid = juasoid = juasaid = juasoid juasaid = juasaid juasqe = juasaid quasqe quasoid = juasaid juasaid juasaid juasqe quasaid = yuasaid quasqe = juasoid juasqe quasaid =. juasaid quasqe juasoid quasqe juasqe quasqe. quasqe quasqe = uasaid juasqe quasqe quasqe juasqe quasqe quasaid = juasaid = juasaid juasaid = juasaid quasaid = juosoid = juasoid = juasoid juasqe quasoid = yuasaid quasqe juasoid juasoid qugsoid = uasoid quasqe juasaid juasoid juasqe juasqe juosqe juasqe quasqe quasqe quasqe quasqe quasqe juasqe soutds souids sauids sauids ¢ sourds | eloueg je1od = NON ~=s«RVIQUO —-[RIIqUO -W911s0g -wndy = -WNdJID ‘| aavy Binary CoDING OF THE CHARACTERS LISTED IN TABLE 1. TABLE 2. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 sAei [e10190d oytiosuy jo quowidojaaaq sajejd yooyo jo JuswodueLy suninge sajeid iseoig saieid Apoq jewuap uo sauids aiejd yensoi uo aulds auQ aed yessos uo soutds 9014], aieyd jensor uo saulds daly aired [el1so1 uo souldg aed yenso1 pasodxq sjaqueg AreyIxey Ploy-20y snus] ouids re[nosadoa1g duids prlowya -eidns posodxq soutds yeseNy soutds [equol4 auids [eloueg ouids jesodwiay1s0g auids 9101914 souids ¢ [ewqiownsitD sautds | [eiqsounoit Aspidophoroides bartoni Aspidophoroides olriki Bathyagonus alascanus Bathyagonus infraspinatus Bathyagonus nigripinnis 1 Bathyagonus pentacanthus Bothragonus swani Odontopyxis trispinosa Xeneretmus latifrons Xeneretmus leiops Xeneretmus ritteri Xeneretmus triacanthus Agonopsis vulsa Hypsagonus quadricornis Ocella dodecaedron Pallasina barbata Percis japonicus Podothecus acipenserinus Sarritor frenatus Estimated ancestral character states LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS XENERETMUS GILBERT 21 collected and converted to binary characters (Ta- ble 2) by additive binary coding (Sokal and Sneath 1963; Kluge and Farris 1969). The ancestral state of each binary character was established by ex- amining an out-group, that is, a group of taxa considered not to be members of the smallest monophyletic unit that contains all members of the in-group. The most frequent character state found among the species of the out-group was considered to be the primitive character state for the taxa of the in-group. The out-group was com- posed of Agonopsis vulsa, Hypsagonus quadri- cornis, Ocella dodecadron, Pallasina barbata, Percis japonicus, Podothecus acipenserinus, and Sarritor frenatus, seven agonid species consid- ered to be related to but not members of the in- group (Jordan and Evermann 1898; Freeman 1951). Several methods of estimating phylogenies from binary data have been proposed (Felsen- stein 1982, and references cited therein). Of these methods, Wagner analysis (Farris 1970; Farris et al. 1970a, 1970b) has been the most widely em- ployed (Baird and Eckhardt 1972; Simon 1979: Presch 1980; Jensen and Barbour 1981; Miya- moto 1983) and has been examined in detail (Colless 1981; Felsenstein, 1973, 1978, 1979: Mickevich 1978, 1980, Mickevich and Farris 1981; Schuh and Farris 1981; Schuh and Pol- hemus 1980; Sokal and Rohlf 1981). Felsenstein (1973, 1978, 1979) has shown some assumptions of the method: 1. Characters evolved independently. 2. Changes of character states through time are a priori improbable. 3. Polymorphisms of character states for a species are exceedingly unlikely. 4. Inequality of lengths of segments of the tree is not so extreme that two changes of states along a long segment is more probable than one change along a short segment. 5. Different lineages evolved independently. While these assumptions do not exactly express how the world is believed to work, without as- sumptions no explicit model could be advanced; with them at least we know the assumptions upon which the hypothesis rests. All systematists make assumptions when trying to work out a phylog- eny, but their assumptions are not as open to examination as are those of a model. The Wagner method searches for the cladogram that requires the fewest number of steps for all the characters. Some hypothesized monophyletic sets may not be supported by uniquely derived characters. Osteological material was cleared and stained with alizarin red S following the method of Tay- lor (1967). Osteological drawings were prepared with the aid of a Wild MS stereomicroscope and camera lucida. Osteological terminology follows Weitzman (1974). The following cleared and stained specimens were examined: Xeneretmus latifrons: UW 18216, 3 (148-171 mm); X. leiops: OSU 7309, 2 (186, 191 mm); X. ritteri: SIO 59-92, 1 (141 mm); X. triacanthus: UW 20948, 3 (146-167 mm). The following abbreviations are used in the osteological illustrations: angular articular process mesopterygoid metapterygoid ASP ascending process MVP mid-ventral plate BPT basipterygium N nasal BR __ branchiostegal ray NZ __ neural zygapophysis BSB _basibranchial OP _ opercle BSO __basioccipital PC __postcleithrum CBR ceratobranchial PHY parahypural CHY ceratohyal PLT palatine CL cleithrum PM _premaxilla CO _ circumorbital POP preopercle COR coracoid PPH_ parapophysis DH _ dorsal hypohyal PRO prootic DLP dorsolateral plate PRT parietal DN dentary PSP parasphenoid ECT ectopterygoid PTG pterygiophore EPB_ epibranchial PTO pterotic EPH epihyal PTS _pterosphenoid EPO epiotic PTT posttemporal EPU _ epural Q quadrate ER _ epiplural nb R radial EXO exoccipital RAT retroarticular F frontal RP _ rostral plate FHA first haemal arch SBO _ subopercle HPP hypural plate SCL | supracleithrum HYB hypobranchial SCP scapula HYM hyomandibular SET supraethmoid IHY interhyal SLP supralateral plate ILP _ infralateral plate SOC _ supraoccipital INT intercalar SPH _ sphenotic IOP interopercle SPN spine IPB infrapharyngo bran- SYM _ symplectic chial at tabular LC _ lacrimal UC _ ural centrum LE lateral ethmoid URN uroneural LLS _lateral-line scale Vv vomer M maxilla VH _ ventral hypohyal MDP mid-dorsal plate VLP ventrolateral plate MIS medial interopercular socket Material examined is deposited at the follow- ing institutions: California Academy of Sciences, tN tN San Francisco (CAS); Natural History Museum of Los Angeles County (LACM); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ); National Museum of Canada, Ottawa (NMC); Oregon State University, Corvallis (OSU); Scripps Institution of Oceanography, La Jolla, California (SIO); Stanford University (SU), material now housed at CAS; University of Alberta, Museum of Zo- ology, Edmonton, Alberta (UAMZ); United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM); and School of Fisheries, University of Washington, Seattle (UW). COMPARATIVE MATERIAL EXAMINED Agonopsis vulsa: UW 4798 (27), UW 5359 (40). Aspidophoroides bartoni: CAS 10842 (2), CAS 15508 (1), CAS 15509 (1), CAS 22355 (2), CAS 26764 (3), CAS 26773 (1), MCZ 28323 (1), MCZ 32463 (1), SU 20421 (1), SU 26136 (5), SU 31699 (3), USNM 125584 (5), USNM 149047 (7), UW 20940 (1), UW 20941 (1), UW 20942 (1), UW 20943 (1), UW 20944 (4), UW 20945 (2), UW 20946 (2), UW 20947 (3). Aspidophoroides olriki: NMC 77-1537 (26), USNM 177610 (1), UW 20935 (3), UW 20936 (2), UW 20937 (1), UW 20938 (2). Bathyagonus alascanus: NMC 650219 (3), NMC 65-319 (3), NMC 66-16 (1), SIO 69-138 (3), SU 3088 (13), UAMZ 1985 (5), UAMZ 2774 (4), USNM 48741 (1), USNM 53582 (1), USNM 53583 (3), USNM 53586 (2), USNM 53589 (5), USNM 53592 (1), USNM 60484 (1), USNM 208391 (2), UW 1422 (5), UW 14392 (11). Bathyagonus infraspinatus: CAS 14911 (2), NMC 65-259 (1), SIO 63-203 (2), SIO 69-110 (2), SIO 72-230 (1), SIO 72- 239 (1), SU 24967 (3), USNM 53593 (1), USNM 53595 (4), USNM 53596 (1), USNM 53597 (2), USNM 53598 (1), USNM 60416 (1), USNM 104676 (15), USNM 207968 (1), USNM 207990 (1), USNM 208117 (1), UW 1660 (1), UW 2886 (5), UW 5006 (1), UW 7583 (1). Bathyagonus nigripinnis: CAS 45524 (1), CAS 45525 (1), CAS 45526 (1), SIO 63-205 (8), SIO 69-140 (3), USNM 46613 (3), UW 7333 (4), UW 18147 (8), UW 20931 (1), UW 20932 (7), UW 20933 (1). Bathyagonus pentacanthus: CAS 15130 (4), NMC 65-397 (1), NMC 65-423 (6), NMC 71-693 (1), SIO 75-355 (3), SIO 80-9 (1), USNM 46612 (3), USNM 63444 (1), UW 18145 (15), UW 18472 (6), UW 19140 (3), UW 20934 (1). Bothragonus swani: UW 14155 (2), UW 17971 (1), UW 20929 (1), UW 20930 (2). Hypsagonus quadricornis: UW 11721 (32). Ocella dodecaedron: UW 20999 (5). Odontopyxis trispinosa: UW 1752 (4), UW 4375 (5). Pallasina barbata: UW 4206 (3). Percis japonicus: UW 21000 (1), UW 21001 (1). Podothecus acipenserinus: UW 3977 (125), UW 7340 (12). Sarritor frenatus: UW 20998 (5). OSTEOLOGY OF XENERETMUS TRIACANTHUS CRANIUM (Figs. 1, 9).—Rostral plate unpaired, situated anterodorsal to nasals, most anterior os- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 teological element and bears a single, dorsally directed spine; on either side, a laterally directed spine. Nasals in contact anteriorly, but separated posteromedially by anterior third of supraeth- moid. Each, bordered laterally by respective lac- rimal (Fig. 9), bears a strong, posterodorsally di- rected nasal spine. Lateral ethmoids lie posterior to nasals. Each comes into contact with frontal medially, lacri- mal laterally (Fig. 9), and parasphenoid ventro- laterally. Frontals are in contact with each other on mid- line for most of their length, separated by su- praethmoid anteriorly. Each frontal bordered posteriorly by sphenotic, pterotic, and parietal; posteroventrally by pterosphenoid. A sharp spine is present on the dorsal surface of each frontal, just posterodorsal to the orbit. Parietals meet on dorsal midline. Bordered along lateral margin by pterotic, tabular, and posttemporal. Each bears two posterodorsally di- rected spines: anteriormost spine knoblike, pos- teriormost strong and sharp. Most of the anterodorsal surface of supraoc- cipital, covered by parietals, comes into contact with exoccipitals along posterior margin. Pterot- ic meets sphenotic anteriorly, tabular posterior- ly, and prootic, exoccipital, intercalar, and post- temporal ventrally. Tabular dorsal to the epiotic; posttemporal bears a spine at posterior margin. Supracleith- rum articulates with posteroventral surface of posttemporal. Epiotic attaches at posterolateral corner of cranium, situated ventral to tabular and post- temporal. Exoccipital forms lateral and dorsal borders of foramen magnum; anteriorly, it forms posterolateral portion of otic capsule. A condyle at its posteroventral corner contacts the lateral process of the anteriormost vertebral centrum (preural centrum 41). Basioccipital, broad anteriorly, narrowing pos- teriorly, forms ventral margin of foramen mag- num. A single large condyle situated posteriorly, abuts against the anteriormost vertebral cen- trum. A posterior projection of parasphenoid overlaps anterior midline of basioccipital ven- trally; its anterolateral corner forms postero- medial portion of otic capsule. Parasphenoid runs from vomer anteriorly to basioccipital posteriorly, forms ventral margin of cranium; anteriorly receives shaft of vomer. N CE F SPH PTS PTO Ficure 1. Dorsal, left lateral, and ventral views of cranium of Yeneretmus triacanthus, UW 20948, 158 mm SL. Dotted lines portray canals of the acoustico-lateralis system. 23 24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 ASP ee a go PM Ficure 2. Left lateral view of upper jaw of Xeneretmus triacanthus, UW 20948, 158 mm SL. Lateral ethmoids border on its anterolateral sur- face and its dorsolateral projections abut on pter- osphenoids dorsally and form posterior margin of orbits. Vomer is “tear’’-shaped; teeth borne M along its anteroventral surface arranged in semi- circular pattern. Prootic forms anterior portion of otic capsule; does not reach posterior margin of orbit. DN ANG RAT Ficure 3. Left lateral view of lower jaw of Yeneretmus triacanthus, UW 20948, 158 mm SL. Dotted lines portray acoustico- lateralis canals. LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS YENERETMUS GILBERT nN wn HYM op |OP Ficure 4. Lateral view of suspensorium and opercular apparatus of Yeneretmus triacanthus, UW 20948, 158 mm SL, right side reversed. Dotted lines portray acoustico-lateralis canals. Intercalar approximately circular; dorsally bordered by exoccipital and pterotic. Antero- medially directed projection of posttemporal overlies its posteroventral face. Upper JAw (Fig. 2).— Premaxilla toothed along op POP | \ f)) \ \ F | line \ kK | at piiowa- 4 ; | = Ci | > SSeS | SBO MIS |OP Ficure 5. Left medial view of opercular apparatus of Yene- retmus triacanthus, UW 20948, 158 mm SL. entire ventral surface in a broad band. The max- illa forked anteriorly to receive ascending process of premaxilla and widens abruptly posteriorly. Lower JAw (Figs. 3, 4).—Anterodorsal three- fourths of dentary toothed. Angular bears socket DH CHY EPH Ihe VH Noy BR Ficure 6. Left lateral view of hyoid apparatus of Yene- retmus triacanthus, UW 20948, 158 mm SL. 26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 FiGure 7. on posteromedial surface to receive a process of quadrate (Fig. 4). Retroarticular attaches to the posteroventral corner of angular. SUSPENSORIUM (Figs. 1, 4-6, 9).—Palatine toothed for a third of its length, teeth centered about midpoint. It articulates posteriorly with mesopterygoid and ectopterygoid. Anterodorsal surface of palatine articulates with ventral sur- face of lateral ethmoid (Fig. 1). A lateral process of palatine articulates with medial surface of lac- rimal. Mesopterygoid borders ectopterygoid ventrally, quadrate posteriorly; does not contact metapter- ygoid. The ectopterygoid lies between palatine, mesopterygoid, and quadrate. Posteroventral surface of quadrate contacts preopercle. Metap- terygoid thin and flat; borders symplectic an- teroventrally and hyomandibular posteroven- trally. Hyomandibular has three dorsal articulating facets: anteriormost, articulating with sphenotic and prootic; medial articulating with pterotic; and posteriormost articulating with anterodorsal corner of the opercle (Fig. 1). Preopercle crescent-shaped with two spines along posterior margin. It is dorsally overlain by circumorbital 3 (Fig. 9). Elongate interopercle bears medial socket that fits onto posterior cor- Dorsal view of branchial basket of Yeneretmus triacanthus, UW 20948, 158 mm SL. ner of epihyal (Figs. 5, 6). Opercle triangular and slightly striated; a socket on anterodorsomedial face receives posterior facet of hyomandibular. Subopercle V-shaped, with crotch of the V lying dorsal to opercle (Fig. 5); posterior arm long and thin, lying on medial face of opercle for majority of its length; anterior arm short, its most dorsal point reaching only half the height of opercle. Hyoip Arcu (Figs. 5, 6).—Dorsal hypohyal anterodorsal to ceratohyal. Ventral hypohyal forms an anterior cap over ceratohyal. Cerato- hyal has four branchiostegal rays connected to it: anterior two ventrally attached, posterior two ventrolaterally. Epihyal connected to the re- maining two basibranchials ventrolaterally. Pos- terodorsally, interhyal connects epihyal to hyo- mandibular. Posterior corner of epihyal fits into medial socket of interopercle (Fig. 5). BRANCHIAL ARCHES (Fig. 7). —Hypobranchials 1-3 broad and flat. Hypobranchial 2 two-thirds length of hypobranchial 1; hypobranchial 3 tear- shaped and two-thirds length of hypobranchial 2. All four ceratobranchials have anterior rows of gillrakers that bear toothlike structures; cer- atobranchials 1-3 also possess posterior rows of similar, “‘tooth’’-bearing gillrakers. Ceratobran- chials 3 and 4 articulate with hypobranchial 3. Ceratobranchial 5 oval, completely toothed. Epi- LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS YENERETMUS GILBERT 27 —— = ee we a Ficure 8. Dorsal view of urohyal of Yeneretmus triacan- thus, UW 20948, 158 mm SL. branchials 1-4 all articulate dorsally with a sin- gle, large, well-toothed infrapharyngobranchial. Epibranchial | forked dorsally in some speci- mens. Uronyvat (Fig. 8).—Urohyal triangular, with a dorsomedial ridge rising posteriorly. CIRCUMORBITAL SERIES (Figs. 1, 9).—Lacrimal forms majority of dorsal surface area of snout and connects with lateral ethmoid and nasal (Fig. 1). Circumorbital 2 is tubelike. Circumorbital 3 has a single centrally located and posteriorly di- rected spine on lateral surface. Circumorbital 4 also tubelike, forming posterior margin of orbit. PecTORAL GIRDLE (Fig. 10).—Three rectan- gular radials and two postcleithra present. Scap- ula crescent-shaped and attached to posterior margin of cleithrum by two arms. Coracoid L-shaped, its anterior arm in contact medially with ventrolateral face of cleithrum; the dorsal arm with ventral margin of scapula and anterior borders of two ventralmost radials. Cleithrum the largest element of pectoral girdle; dorsally attaches to supracleithrum. PeLvic GirDLE (Fig. 1 1).—Basipterygia paired and connected medially to each other along pos- terior tenth of their length. A ventral ridge runs anteroposteriorly. One spine and two rays pres- ent; lateralmost ray tightly bound to medial sur- face of spine. VERTEBRAL COLUMN (Fig. 12).—There are 41 preural centra in all the specimens of Yeneretmus triacanthus dissected. The 41st preural centrum has three anterior concave facets (a large central facet and two smaller lateral ones) that articulate with posterior surface of cranium; neurapophy- ses not dorsally ankylosed; 41st through 31st preural centra bear epiplural ribs. Neural zyg- apophyses become more pronounced posterior- ly. Haemal spines on 29th through first preural tc co2 c Ficure 9. Left lateral view of circumorbital bones of Xene- retmus triacanthus, UW 20948, 158 mm SL. Dotted lines por- tray acoustico-lateralis canals. centra. Haemal spines posterior to posteriormost anal fin pterygiophore lie ventral to adjacent pos- terior centrum (this tendency increases poste- riorly). Posterior two rays of anal and second dorsal fin articulate with last pterygiophore of ventral and dorsal series, respectively. Four pterygio- phores lie between first and second dorsal fins. No ray articulates with these pterygiophores. An- terior two pterygiophores of ventral series do not articulate with any rays. Neural spines on 40th through first preural centra; centra posterior to last dorsal pterygio- phore have broad neural spines that lie between neural zygapophyses of adjacent posterior ural centrum. One parahypural fused to ventral margin of hypural plate. One uroneural tightly bound to dorsal margin of hypural plate. Epurals absent. DERMAL P ates (Fig. 13).—All dermal plates of supralateral, dorsolateral, and mid-dorsal se- ries bear posteriorly directed spines. All infra- lateral plates bear similar spines as well, except those medial to pectoral fin. Ventrolateral plates spineless, except for those behind pelvic fin in- sertion to approximately four plates anterior to insertion of first anal fin ray. Mid-ventral plates spineless. Anterior supralateral plates overlain poste- riorly by next supralateral plate for approxi- mately half their length; those more posterior in position overlain by as little as 20% of their length. Supralateral plates overlain by dorsolateral and mid-dorsal plates dorsally, and lateral-line scales ventrally. Anterior infralateral plates have half their length overlain by adjacent posterior infralateral plate; length covered reduced to 20% posteriorly. Infralaterals overlain by lateral-line scales dor- 28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 ScP PC | PC2 COR Ficure 10. Left lateral view of pectoral girdle of Xeneretmus triacanthus, UW 20948, 158 mm SL. sally and ventrolateral and mid-ventral plates __teriorly by next ventrolateral plate. Medially, ventrally. ventrolateral plates slightly overlap each other Ventrolateral plates bordering anal fin have on midline. medial projections that meet on midline such Each mid-ventral plate posteriorly overlain by that fin rays surrounded by plates. A third of the immediately posterior mid-ventral plate; length length of each ventrolateral plate overlain pos- covered approximately 15%. Figure 11. Ventral view of pelvic girdle of Xeneretmus triacanthus, UW 20948, 158 mm SL. LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS XENERETMUS GILBERT 29 URN uc Ficure 12. Left lateral view of vertebral centra of Yeneretmus triacanthus, UW 20948, 158 mm SL. 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 SLP MDP DLP Speen = oe SLP eS ILP — ‘ee IEP, MVP VLP FiGure 13. Dermal body plates of Xeneretmus triacanthus, UW 20948, 158 mm SL: (A) Dorsal view of plates immediately posterior to insertion of posteriormost ray of second dorsal fin. (B) Left lateral view of plates immediately posterior to pectoral fin. (C) Ventral view of plates immediately posterior to inser- tion of posteriormost anal fin ray. Each dorsolateral plate overlain posteriorly by next dorsolateral plate for approximately 35% of length. Like ventrolaterals, dorsolateral plates bordering first and second dorsal fins have me- dial projections that meet on dorsal midline such that only a small break in plates exists where dorsal rays insert. Each mid-dorsal plate overlain posteriorly for 20% of length by adjacent posterior mid-dorsal plate. ACOUSTICO-LATERALIS SYSTEM (Figs. 1, 3, 4, 9).—Acoustico-lateralis system of cranium passes posteriorly through nasals. Canal enters frontals through an anterior pore and extends along me- dial border to a medial pore where it turns lat- erally, continuing to posterolateral pore of fron- tal where it branches (Fig. 1). Anteriorly directed branch of acoustico-lateralis system passes through entire circumorbital series (Fig. 9). Pos- teriorly directed branch passes through pterotic and branches again beneath tabular. Medially directed branch passes beneath posterior spine of parietal to a medial pore dorsal to supraoc- cipital where it meets its counterpart from the other side. Posteriorly directed branch passes through posttemporal and supracleithrum, con- tinues posterolaterally along entire length of fish (Fig. 1). A second acoustico-lateralis canal passes posteriorly through dentary, angular, and pre- opercle (Figs. 3, 4). COMPARATIVE OSTEOLOGY The osteology of the other species of the genus is almost identical with that of X. triacanthus described above; very few differences were found between species that exceeded variation within species. Xeneretmus triacanthus has two spines on the posterior margin of the preopercle where- as its congeners possess only one. The rostral plate of the other members of the genus do not have lateral spines (except in some individuals of X. leiops). Xeneretmus ritteri has two spines on circumorbital 3, whereas the other species of Xeneretmus have only one. Finally, the pterotic of X. ritteri bears two spines while in the other members of the genus only a ridge may be dis- cerned. The following comparisons can be made with Rendahl’s (1934) work on the agonid species Hypsagonus quadricornis. The rostral plate, di- agnostic for Yeneretmus (Fig. 1), does not occur in Hypsagonus (Rendahl 1934, figs. 1-3). The vomer of Hypsagonus is toothless (Rendahl 1934, fig. 2). The frontal spine of Hypsagonus is much larger than that of Xeneretmus (Fig. 1; Rendahl 1934, figs. 1, 3). In Hypsagonus, Rendahl (1934, fig. 1) depicted the supraoccipital as lying along the entire medial border of the parietal, in such a configuration that the parietals are not in con- tact along their medial edges; the parietals meet along their entire medial edges in Yeneretmus (Fig. 1). Rendahl (1934, figs. 28A, 28B) depicted the retroarticular as lying only on the medial face of the angular whereas in Xeneretmus the re- troarticular is visible from both a medial and lateral view (Fig. 3). Rendahl (1934, fig. 24A) showed the mesopterygoid and metapterygoid of Hypsagonus to be in contact with each other; in Xeneretmus the quadrate is between these two bones such that they do not meet (Fig. 4). Finally, the posterior arm of the subopercle of H. quad- ricornis is considerably shorter than is the case for Xeneretmus (Figs. 4, 5; Rendahl 1934, figs. 24A, 24B). LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS YENERETMUS GILBERT 31 The following comparisons can be made with the work of Ilina (1978). Neither of the genera Podothecus and Agonus possesses a rostral plate. Ilina (1978, figs. 2-4, 6) portrayed the supraoc- cipitals of Podothecus acipenserinus, P. veternus, P. gilberti, and P. thompsonias lying between the parietals, such that they do not meet along their medial edges, as they do in Xeneretmus (Fig. 1). The posttemporal of Podothecus acipenserinus, P. veternus, and Agonus cataphractus apparently makes no contact with the intercalar (Ilina 1978, figs. 2, 3, 7); in Xeneretmus the anteriorly di- rected projection of the posttemporal touches the intercalar (Fig. 1). SYSTEMATICS Genus Xeneretmus Gilbert Xenochirus GiBert, 1890:90 (type-species Yenochirus tria- canthus GiLBerRT, 1890, by original designation; preoccupied by Xenochirus GioGer, 1842, a genus of marsupial mam- mal). Xeneretmus GiLBertT, in Jordan 1903:360 (substitute name for Xenochirus GitpBert, 1890 [preoccupied, therefore taking the same type-species Xeneretmus triacanthus)]). DiaGnosis.— The genus Xeneretmus is distin- guished from all other agonid genera by the ab- sence of a supraoccipital pit and by the presence of an exposed rostral plate bearing a single dor- sally directed spine. DEscrIPTION. — Body tapering uniformly from pectoral girdle to caudal fin; anterior cross sec- tions octagonal, cross section through caudal peduncle hexagonal; completely encased in over- lapping dermal plates. All dorsolateral, mid-dor- sal, and supralateral plates bearing posteriorly directed spines; all but those plates medial to pectoral fin of infralateral series bearing poste- riorly directed spines; ventrolateral plates be- tween pelvic fin insertion and insertion of first anal fin ray bearing posteriorly directed spines; no spines present on mid-ventral plates; dorso- lateral plates 22-24; mid-dorsal plates 12-19; supralateral plates 39-45; infralateral plates 35- 42; ventrolateral plates 21-23; mid-ventral plates 14-20. In comparison with the other genera of Agonidae, Xeneretmus has long spines on slightly flexible angular dermal plates. Cephalic spines. One nasal spine; one frontal spine dorsal to posterior edge of orbit; two pa- rietal spines; one posttemporal spine; one or two spines on circumorbital 3; one or two spines on posterior margin of preopercle. Fin rays. All simple; four ventralmost pectoral Ficure 14. Right lateral view of head of two species of Xeneretmus: (A) X. triacanthus, (B) X. latifrons. Arrows in- dicate dermal plates of cheek region. rays thickened (in comparison to the dorsalmost pectoral rays), and projecting fingerlike from the fin membrane; dorsal two thickened rays longest rays of pectoral fin. First dorsal, 5-8; second dor- sal, 6-8; anal, 5-8; pectoral, 12-16; pelvic I, 2; branchiostegal rays, 6. Mouth. Both jaws of equal length, mouth ter- minal; teeth present on premaxilla, dentary, pal- atine, and vomer. Barbels present along ventral margin of dentary at edges of acoustico-lateralis pores and at posterior corner of maxilla. Measurements. The following ranges for pro- portions of all species of the genus are expressed in thousandths of standard length (number of specimens measured in parentheses): anal length, 305-510 (201); vent length, 210-335 (197); cau- dal peduncle length, 370-479 (199); second dor- sal length, 443-592 (201); depth at second dorsal, 47-72 (197); first dorsal length, 258-386 (203); depth at first dorsal, 65-132 (171); pectoral length, 123-217 (189); pelvic length, 51-105 (197); pec- toral width, 92-157 (189); head length, 166-241 (203); ventral head length, 80-155 (200); length from supraoccipital pore to snout, 147-189 (203). The following proportions, associated with characteristics of the head, are expressed in thou- sandths of head length (number of specimens measured in parentheses): orbit length, 249-476 (207); upper jaw length, 229-363 (141); snout Ww is) TABLE 3. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 Rance, MEAN, STANDARD DEVIATION, AND SAMPLE SIzE FOR MERISTIC CHARACTERS OF SPECIES OF Yeneretmus. Character X. latifrons X. leiops X. ritteri X. triacanthus First dorsal spines 6-8 x= 6.9 6-7 X= 6.7 6-7 X= 6.4 5-7 X= 6.0 SD =0.50 n=110 SD =0.45 n= 30 SD = 0.54 n=7 SD = 0.28 n=63 Second dorsal rays 6-8 x=7.0 7-8 = 7.4 6-7 X= 6.9 6-7 X= 6.6 SD = 0.43 n= 110 SD = 0.49 n= 30 SD = 0.38 n=7 SD =0.50 n= 63 Anal fin rays 6-8 X=7.2 6-8 X= 6.9 6-7 X= 6.9 5-7 xX=6.1 SD=0.15 n= 110 SD = 0.57 = 30 SD = 0.38 n=7 SD = 0.49 n= 63 Pectoral fin rays 13-15 X= 14.1 13-15 X= 14.0 16 X= 16 12-14 X= 13.0 SD =0.40 n= 109 SD = 0.33 n= 29 SD=0.00 n=7 SD =0.22 n=61 Eyeball plates 3-6 X= 4.2 0 x= 0.0 3-6 X= 5.0 2-6 X= 3.9 SD =0.62 n=110 SD = 0.00 n= 30 SD=1.0 n= SD = 0.76 n= 63 Supralateral plates 39-42 x= 40.9 43-45 X=43.9 40-41 X= 40.7 41-43 X= 42.0 SD = 0.65 n= 109 SD =0.80 n= 28 SD=0.49 n= SD = 0.46 n=58 Infralateral plates 35-40 X = 37.6 39-42 X= 40.3 36-38 x = 36.9 38-40 xX = 39.0 SD =0.92 n= 108 SD = 0.98 n= 28 SD =0.69 n=7 SD = 0.49 n=58 Mid-dorsal plates 12-16 X= 14.7 16-19 X= 17.7 14-15 x= 14.7 15-17 X= 16.2 SD = 0.63 n= 108 SD=0.72 n=29 SD = 0.49 n=7 SD = 0.54 n=58 Dorsolateral plates be- 3-5 x=4.1 45 x= 4.4 45 X= 4.6 46 X=5.1 tween first and sec- SD =0.53 n=110 SD = 0.50 n= 30 SD=0.54 n=7 SD = 0.33 n= 64 ond dorsal fins Mid-ventral plates 14-17 x= 15.4 16-20 X= 18.2 14-16 x= 14.9 16-18 X= 16.7 SD = 0.66 n= 109 SD = 0.83 n= 28 SD = 0.69 n= SD =0.50 n=59 Cheek plates O-l X=0.0 0 x= 0.0 0 x= 0.0 1-4 X= 2.75 SD=0.10 n=110 SD = 0.00 n= 30 SD=0.00 n= SD = 0.69 n= 64 Vertebrae 40-42 X=40.6 43-45 X=43.7 4041 x=40.7 42 X= 42.0 SD =0.79 n= SD = 0.82 n=6 SD=0.52 n=6 SD=0.00 n=6 length, 157-330 (206); interorbital length, 58- 123 (107). KEY TO THE SPECIES OF THE GENUS XENERETMUS Cheek plates present, filling area between circumorbitals and elements of the lower jaw (Fig. 14) .. = ; Wenereimin fe Nenereinia) iriacantHs. p. 36 . Cheek plates absent, or rarely represented by a single plate that does not fill the area between the circumorbitals and elements of the lower jaw (Fig. 14) ... Ee = ~ Subgenus Xenopyxis 2 2 ; Two o or more dermal plates on eyeball _. 3 . No dermal plates on eyeball ae Xeneretmus (Xenopyxis) leiops, p. 35 . Two or more barbels present at posterior corner of maxilla, 16 pectoral rays _ Xeneretmus (Xenopyxis) ritteri, p. 36 One barbel present at posterior corner of maxilla, 13-15 pectoral rays . : Xeneretmus (Xenopyxis) latifrons, p D. 32 3b. Subgenus Xenopyxis Gilbert Xenopyxis GILBERT, 1915:345 (type-species Yeneretmus (Xen- opyxis) latifrons Gilbert 1890, by original designation). Jor- dan et al. 1930:396 (elevated to generic level). DraGnosis. — Distinguished from the subgenus Xeneretmus by the absence of dermal plates in the cheek region (Fig. 14B), and the failure of the breast plates to abut against each other. It further differs in having higher average counts for rays in the first and second dorsal, anal, and pectoral fins (Table 3); having larger eyes (Table 4); and in general being more robust. Xeneretmus (Xenopyxis) latifrons (Gilbert) [Blacktip Poacher] (Figure 15) Xenochirus latifrons GitBert, 1890:92 (original description, lectotype USNM 43091). Xeneretmus latifrons GicBert, in Jordan 1903:360 (new com- bination, Yenochirus preoccupied). Xeneretmus (Xenopyxis) latifrons GiLBerT, 1915:345 (descrip- tion, key). Xenopyxis latifrons JORDAN ET AL., 1930:396 (checklist). MATERIAL EXAMINED.—One hundred and eighty-two spec- imens, 62 to 173 mm. LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS YENERETMUS GILBERT 33 TABLE 4. RANGE, MEAN, AND SAMPLE SIZE FOR Bopy ProporTIONs OF Species OF Xeneretmus. Character X. latifrons Anal length/SL 305-486 X= 462 n= 109 Vent length/SL 220-335 X= 253 n= 109 Caudal length/SL 370-447 X= 414 n= 106 Second dorsal length/SL 443-592 X= 487 n= 108 Depth at second dorsal/SL 48-72 X=59 n= 106 First dorsal length/SL 258-386 X = 307 n= 108 Depth at first dorsal/SL 73-132 X=91 n=88 Pectoral length/SL 123-217 X= 167 n= 104 Pelvic length/SL 57-105 X=84 n=108 Pectoral width/SL 97-157 X= 113 n= 102 Head length/SL 166-241 X= 212 n= 109 Ventral head length/SL 80-146 X= 121 n= 105 Supraoccipital pore 147-186 to snout length/SL X= 166 n= 109 Orbit length/head length 249-476 X= 374 n= 109 Upper jaw length/head length 255-363 X= 293 n= 53 Snout length/head length 157-330 X= 258 n= 109 Interorbital length/head length 58-123 X= 100 n= 107 Caudal depth/caudal length 37-68 X=47 n=95 Lecrotyre.—USNM 43091, 131 mm, Albatross station 2935, San Diego, California, 32°45'N, 117°23’W, 227 m. PARALECTOTYPES.—CAS 5072, 3 (108-110 mm), Albatross station 2973, Point Conception, California, 34°20'N, 119°44’W, 124m; USNM 46602, 8 (72-136 mm), Albatross station 2935, San Diego, California, 32°45’N, 117°23’W, 227 m; USNM 46605, 2 (110-112 mm), Albatross station 3059, Lincoln City, Oregon, 44°56'N, 124°13’W, 141 m; USNM 46608, 120 mm, Albatross station 2972, Santa Barbara, California, 34°19’N, 119°41'W, 112 m; USNM 46611, 111 mm, Albatross station 2948, Santa Cruz Island, California, 33°56’N, 119°42'W; UW 1416, 2 (109-110 mm), Albatross station 2973, Santa Barbara, California, 34°20'N, 119°44’W, 124 m. ADDITIONAL Non-Type MATERIAL.—CAS 12572, 141 mm, Farallones, California, 37°43'N, 123°03'W; CAS 14282, 2(128, 131 mm), San Pedro, California, 33°45’N, 118°11’'W; CAS 26404, 2 (114, 134 mm), Port Heuneme, California, 34°09'N, X. leiops X. ritteri X. triacanthus 434-493 484-508 448-510 X= 457 n=28 X=498 n=7 X= 477 n=S7 224-288 253-280 210-255 X= 244 n=24 X= 265 n=7 X= 230 n=57 384-479 380-409 384-439 X= 434 n= 30 X= 393 n=7 X= 410 n= 56 446-498 490-516 459-500 X= 464 n=29 X=501 n=7 X= 483 n= 57 48-64 58-64 47-67 X=57 n=29 X=61 n=7 X=58 n=S55 269-319 315-335 287-317 X= 291 n= 30 X= 323 n=7 X = 302 n= 58 72-98 86-117 65-109 X=85 n=22 X=95 n=7 X=82 n=54 139-205 163-188 151-202 X=171 n=29 X= 173 n=7 X=175 n=49 56-96 72-99 51-94 X=77 n=25 X=83 n=7 X=77 n=57 92-151 107-127 95-125 X= 106 n= 23 X=119 n=7 X= 109 n= 57 191-227 231-248 190-215 X= 204 n=25 X= 238 n=7 X= 201 n=57 99-155 133-151 110-143 X= 113 n= 30 X= 142 n=7 X= 125 n=58 150-181 167-189 149-171 X= 163 n= 28 X= 180 n=7 X= 160 n=57 335-419 334-372 299-358 X= 374 n= 30 X= 358 n=7 X= 326 n=61 234-313 278-300 229-283 X= 280 n= 28 X= 286 n=7 X=251 n= 53 230-316 249-271 258-301 X = 284 n= 30 X= 258 n=7 X = 282 n= 60 66-96 84-125 71-112 =81 n=30 X=95 n=7 xX=89 n=60 35-48 49-60 39-58 Xx=40 n=28 X=57 n=7 X=45 n=53 119°12'W; CAS 26447, 116 mm, Gaviota, California, CAS 26554, 3 (112-113 mm), Goleta Point, California, 34°27'N, 119°50'W; CAS 26560, 4 (97-112 mm), Santa Barbara Point, California, 34°30'N, 120°00'W; CAS 26563, 4 (118-130 mm), Santa Barbara Channel, California, 34°15’N, 119°55'W; CAS 26596, 120 mm, Santa Monica, California, 33°50'N, 118°38'W; CAS 26630, 19 (62-103 mm), Point Dume, California, 34°00'N, 118°50'W; CAS 37497, 149 mm, Half Moon Bay, California, 37°10'N, 122°42’W; CAS 40310, 136 mm, Santa Cruz, Cali- fornia, 34°07'N, 119°42’'W; CAS 47107, 4 (99-110 mm), Go- leta, California, 34°27’N, 119°50'W; CAS 47110, 8 (91-128 mm), Santa Monica Bay, California, 33°58'N, 118°38’W; CAS 47111, 6 (66-108 mm), Morro Bay, California, 35°30'N, 121°15'W; CAS 47112, 156 mm, Marin County, California. NMC 65-0259, 5 (121-153 mm), Kwatna Inlet, British Co- lumbia, 52°07'N, 127°38'W. SU 16903, 2 (77, 95 mm), Santa Barbara Channel, Califor- 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 en A : bee A Ficure 15. Xeneretmus latifrons, 142 mm SL. Courtesy of R. H. Gibbs, Jr., and the Fish Division, National Museum of Natural History. nia, 30°26'N, 120°14'W; SU 39779, 118 mm, Santa Barbara Channel, California, 34°25'N, 120°18'W. USNM 61176, 152 mm, Albatross station 3671, Santa Cruz, California, 37°00'N, 122°20’'W; USNM 63435, 3 (70-114 mm), Point Soma, California, 32°41'N, 117°14’W; USNM 63437, 3 (82-123 mm), Point Soma, California, 32°41'N, 117°14'W. UW 1415, 131 mm, Albatross station 3174, Bodega Bay, California, 38°16’N, 123°14'W; UW 2943, 10 (116-150 mm), Camano Island, Washington, 47°59’N, 122°13’W; UW 3151, 123 mm, Burrard Inlet, British Columbia, 49°10'N, 123°00'W; UW 3168, 4 (93-138 mm), Elliot Bay, Washington, 47°36'N, 122°22'W; UW 3907, 43 (66-142 mm), Hoodsport, Washing- ton, 47°30'N, 123°10'W; UW 4224, 12 (76-130 mm), Hood Canal, Washington, 47°17'N, 122°42'W; UW 4308, 129 mm, Hood Canal, Washington, 47°30'N, 123°10'W; UW 5780, 2 (137, 144 mm), Hoodsport, Washington, 47°30'N, 123°10'W; UW 5861, 120 mm, Golden Gardens, Washington, 47°40'N, 122°24’'W; UW 5872, 124 mm, Tulalip Bay, Washington; UW 5960, 132 mm, Meadow Point, Washington, 47°36’N, 122°22'W; UW 7347, 4 (104-133 mm), Puget Sound, Wash- ington; UW 8016, 142 mm, Ballard, Washington, 47°40'N, 122°25'W; UW 18216, 5 (158-163 mm), Columbia River, 46°N, 124°W; UW 18297, 162 mm, Columbia River, 46°N, 124°W; UW 18507, 3 (162-173 mm), 46°N, 124°W; UW 20939, 146 mm, Bainbridge Island, Washington, 47°37'N, 122°33'W. DiaGcnosis. — Distinguished from other mem- bers of subgenus by following combination of characters: three to six spine-bearing dermal plates on each eyeball; one barbel at posterior corner of maxilla; 13-15 pectoral rays (Table 5). DescriPTION.—Posterior free-fold of bran- Tasie 5. CHARACTERS USED IN DISCRIMINATING AMONG THE Species OF Xeneretmus. Characters Maxil- Eye- lary ball — bar- Cheek Pectoral Taxa plates bels plates rays Xeneretmus latifrons 3-6 ~— 1 Oorsmall 13-15 Xeneretmus leiops 0 1 0 13-15 Xeneretmus rittert 36 2 0 16 Xeneretmus triacanthus 2-6 2-3 1-4 12-14 chiostegal membrane narrow; two barbels on ventral surface of dentary, one at each posterior margin of two anteriormost acoustico-lateralis pores; breast plates surrounded by skin, and hav- ing slightly raised centers; first dorsal fin with black distal margin; second dorsal fin membrane lightly pigmented along rays, clear between rays; counts and proportions are given in Tables 3 and 4. DistTRIBUTIONS. —Gilbert (1890) described YX. latifrons from specimens obtained from Alba- tross stations situated off the coasts of California and Oregon, ranging between approximately 33° and 45°N latitude. Material examined in this study ranged from Ensenada, California to Kwat- na Inlet, British Columbia (Fig. 16). Gilbert (1890, 1915) reported YX. /atifrons occurred in depths from 35 to 399 m. All the lots examined for this study fell within that depth range. Comments.—In Gilbert’s (1890) original de- scription of the species, no type-specimen was designated. When the single specimen in a lot now registered as USNM 43091 was transferred to the United States National Museum by Gil- bert and his associates, it was indicated in an accompanying letter that this specimen was soon to be described as the type of the species (Susan Jewett, USNM, personal communication, 7 June 1982). After examination of this specimen and the other members of the syntypic series avail- able to me (CAS 5072, USNM 46602, USNM 46605, USNM 46608, USNM 46611 and UW 1416), USNM 43091 is hereby designated as the lectotype. This decision was reached for the fol- lowing reasons: It appears to have been Gilbert’s intention to designate this specimen as the type for the species, it is very close to the average for the species in the majority of characters, and its condition is as good as, if not better than, that of any other member of the syntypic series. In comparison to its congeners, X. /atifrons has LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS YENERETMUS GILBERT 35 50 30 @ ee 30 ) 20 M 20 130 125 120 115 110 105 Ficure 16. Distribution of Yeneretmus latifrons (circles) and X. ritteri (stars). low counts for numbers of mid-ventral plates, mid-dorsal plates, infralateral plates, supralater- al plates, and vertebrae; and high counts for numbers of unpaired fin rays (Table 3). Its snout is shorter, orbits longer, and interorbital distance greater than those of the other Yeneretmus (Table 4). Xeneretmus (Xenopyxis) leiops (Gilbert) [Smootheye Poacher] Xeneretmus (Xenopyxis) leiops Girpert, 1915:348 (original description, key, illustration, holotype USNM 75813). Xenopyxis leiops JORDAN ET AL., 1930:396 (checklist). MATERIAL EXAMINED. — Forty-four specimens, 67-211 mm. Hovotyre.—USNM 75813, 163 mm, Albatross station 4410, Catalina Island, California, 323-357 m. ParatyPe.—SU 22988, 2 (108, 136 mm) Albatross station 4410, Catalina Island, California. ADDITIONAL Non-TyPe MATERIAL.—LACM 93744, 5 (83- 160 mm), Catalina Basin, California, 32°N, 118°W. NMC 67-0348, 2 (147, 163 mm), Rennell Sound, British Columbia, 53°21’N, 133°04’W; NMC 72-061 3 (192-211 mm), Barkley Sound, British Columbia, 48°30'N, 126°10'W. 45 xd ee 2a 4? 7 \ 25 i) me 25 be vv \ 130 125 120 115 110 105 Ficure 17. Distribution of Yeneretmus triacanthus (cir- cles) and_X. /eiops (stars). OSU 7305, 67 mm, Newport, Oregon, 44°40'N, 124°10'W; OSU 7309, 15 (134-191 mm), Columbia River, 46°10'N, 124°05'W. SIO 72-81, 206 mm, Neah Bay, Washington, 48°22'N, 126°10'W. SU 3623, 7 (132-172 mm), Central California Coast, 34°45’N, 121°29'W; SU 16711, 2 (153, 165 mm), Monterey Bay, Cal- ifornia, 36°49’N, 122°30'W; SU 26420, 3 (161-182 mm), Mon- terey Bay, California, 36°49’N, 122°30'W. UW 18123, 198 mm, Columbia River, 46°N, 124°W; UW 18473, 2 (171, 179 mm), Columbia River, 46°N, 124°W. DiAGnosis.— Distinguished from the other members of the subgenus by the following com- bination of characters: absence of dermal plates on eyeball; one barbel at posterior corner of max- illa; 13-15 pectoral rays (Table 5). DescriPTION.—Posterior free-fold of bran- chiostegal membrane wide; breast plates thin and completely surrounded by skin; one to three bar- bels at posterior margin of anteriormost acous- tico-lateralis pore of dentary, and one or none at posterior margin of middle acoustico-lateralis 36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 pore of dentary; first dorsal whitish at base, black at distal margin, black pigmentation nearly reaching origin of fin, retreating distally poste- riorly; second dorsal black at distal margin. DisTRIBUTION.—Gilbert (1915) described X. leiops from specimens captured off Santa Cata- lina Island, southern California (Albatross sta- tion 4410). Yeneretmus leiops has a geographic distribution that ranges from Santa Catalina Is- land north to the Queen Charlotte Islands (Fig. 17). Specimens examined during this study were captured from depths between 183 and 357 m. Comments.—Relative to its congeners, X. leiops has high counts for second dorsal rays, supralateral plates, infralateral plates, mid-dor- sal plates, mid-ventral plates, and vertebrae (Ta- ble 3). It also has a shorter precaudal region, longer caudal peduncle and orbits, and a smaller interorbital distance than the other members of the genus (Table 4). Xeneretmus (Xenopyxis) ritteri (Gilbert) {Stripefin Poacher] Xeneretmus (Xenopyxis) ritteri GILBERT, 1915:350 (original de- scription, key, illustration, holotype USNM 75814). Xenopyxis ritteri JORDAN ET AL., 1930:396 (listed). MATERIAL EXAMINED. — Nine specimens, 106-141 mm. Ho.otyre.—USNM 75814, 123 mm, Albatross station 4366, Point Loma, California, 320-331 m. ParatyPe.—SU 22980, 106 mm, Albatross station 4322, San Diego, California, 353-415 m. AppITIONAL Non-Type MATERIAL.—LACM 88182 2 (111, 137 mm) Gulf of California, Mexico, 29°N, 112°W. SIO 59-92, 4 (121-141 mm), Cedros Island, Mexico, 28°23'N, 115°21’W; SIO H50-245B, 126 mm, Torrey Pines, California, 32°10'N, 117°10'W. DiaGcnosis.— Distinguished from the other members of the subgenus by the following com- bination of characters: three to six spine-bearing plates on eyeball, two barbels at posterior corner of maxilla, 16 pectoral rays (Table 5). DescripTION.—Posterior free-fold of bran- chiostegal membrane narrow; two barbels on ventral margin of dentary, one at posterior mar- gin of each of two anteriormost acoustico-la- teralis pores; breast plates with bony prickles at centers, each surrounded by skin (such that they do not contact each other at their edges); dorsal fins with black bars along base and distal margin. DIsTRIBUTION.— Gilbert (1915) described YX. ritteri from specimens captured near San Diego (Albatross stations 4366 and 4322). Since that time YX. ritteri has been obtained from Cedros Island, Baja California, north to Malibu, Cali- fornia, and in the northern section of the Gulf of California (Fig. 16). Specimens examined for this study were captured at depths from 274 to 415 m. Comments.—In comparison to its congeners, X. ritteri has low counts for supralateral plates, infralateral plates, mid-dorsal plates, mid-ven- tral plates, and vertebrae (Table 3). It also has a larger head, a longer precaudal region, and a shorter caudal peduncle than other species of Xeneretmus (Table 4). Its spines and ridges are more strongly developed than those of its con- geners. Subgenus Xeneretmus Gilbert Xeneretmus (Gilbert) 1915:345 (Type species Yeneretmus (Xeneretmus) triacanthus Gilbert, 1890, by original designation.] Diacnosis. — Distinguished from the subgenus Xenopyxis by the presence of one to four dermal plates in the cheek region leaving little or no skin exposed in the cheek region (Fig. 14), and the tight arrangement of the breast plates. It further differs in having lower average counts for fin rays in the first and second dorsal, anal, and pectoral fins (Table 3); having larger eyes (Table 4); and in general being slender in comparison. Xeneretmus (Xeneretmus) triacanthus (Gilbert) [Bluespotted Poacher] Xenochirus triacanthus GiLBerT, 1890:91 (original description, lectotype USNM 43089). Xeneretmus triacanthus Gicpert, in Jordan, 1903:360 (New combination, Yenochirus preoccupied). Xeneretmus (Xeneretmus) triacanthus GiLBerT, 1915:345 (de- scription, key). MATERIAL EXAMINED. —Seventy-six specimens, 74-167 mm. LectotyPe.—USNM 43089, 151 mm, Albatross station 2893, Santa Barbara Channel, California, 34°13'N, 120°33’W, 265 m. PARALECTOTYPES.—USNM 46601, 2 (124, 154 mm), Al- batross station 2893, Santa Barbara Channel, California, 34°13/N, 120°33'W, 265 m; USNM 46606, 117 mm, Albatross station 3059, Lincoln City, Oregon, 44°56’N, 124°13'W, 141 m:; USNM 125577, 5 (137-152 mm), Albatross station 2973, Point Conception, California, 34°20'N, 119°44’W, 124 m. AppitioNAL Non-Tyre MATERIAL.—CAS 13100, 2 (78, 90 mm), San Pedro, California, 33°43’N, 118°23’W; CAS 14270, 4 (132-144 mm), Monterey Bay, California, 36°48’N, 122°07’W; CAS 26405, 134 mm, Port Hueneme, California, 34°10'N, 119°10’'W; CAS 26441, 138 mm, Gaviota, California, 34°05'N, 119°02’W; CAS 47113, 2 (142, 147 mm), Point Baja, Califor- nia, 30°05'N, 115°58’W. LACM 320303, 141 mm, Bahia San Quintin, Mexico, LEIPERTZ: A REVIEW OF THE FISHES OF THE AGONID GENUS XENERETMUS GILBERT 37 Bathyagonus infraspinatus Bathyagonus alascanus Bathyagonus pentacanthus Bathyagonus nigripinnis Xeneretmus triacanthus Xeneretmus ritteri w ® ® o AS) Ae] Ge ee Lees) c 9 5 [= E aS fo) is 2 -— wo = a o a aes ® c o o 2 4 fs) . “ 6 . ow - £ =e 5 9 6 oo oa sim 5 sow eel OR es (Ss eC ‘oO oOo ® r= {9} ou fat, fe o = Ch 32) FS o : Ce ea Ce As) oe x = orf OQ SG atsy csi 5) Ficure 18. Cladogram for selected agonid taxa based on a Wagner analysis of data presented in Tables | and 2. 30°18’N, 115°53’W; LACM 322463, 5 (120-155 mm), Santa Monica Bay, California, 33°54’N, 118°25’W. NMC 65-258, 147 mm, Kwatna Inlet, British Columbia, 52°25'N, 127°34'W. SIO 51-255-56, 7 (88-151 mm), Channel Islands, California, 34°01'N, 119°24'W; SIO 6047156, 154 mm, Baja California Norte, Mexico, 31°18'N, 116°38’W; SIO 63104256, 8 (75-154 mm), Point Arguello, California, 34°10'N, 120°00’W. SU 16712, 3 (141-142 mm), Monterey Bay, California, 36°44’N, 121°58’W; SU 19172, 104 mm, Santa Barbara Chan- nel, California, 34°25'N, 120°906’'W; SU 21363, 4 (127-148 mm), Point Pinos, California; 36°37’N, 121°55'W; SU 39780, 131 mm, Santa Barbara Island, California, 33°37'N, 119°05'W; SU 39781, 6 (105-134 mm), Santa Barbara Island, California, 34°25'N, 120°18'W. USNM 59370, 74 mm, Albatross station 3171, Russian Riv- er, California, 38°21'N, 123°20'W; USNM 63422, 129 mm, Point Soma, California, 32°41'N, 117°16'W; USNM 63423, 122 mm, Point Pinos, California, 36°38'N, 121°56’'W; USNM 63427, damaged, Santa Cruz, California, 36°58’N, 122°01'W; USNM 103719, 136 mm, Mukilteo, Washington, 47°57'N, 122°18'W. UW 4175, 117 mm, Saratoga Passage, Washington, 47°50'N, 122°30'W; UW 4725, 4 (106-159 mm), Richmond Beach, Washington, 47°50'N, 122°30’'W; UW 20948, 9 (129-167 mm), Ballard, Washington, 47°30'N, 122°30'W. DEscRIPTION.— Two to six spine-bearing plates on eyeball; two, rarely three, barbels at posterior corner of maxilla; 12-14 pectoral rays (Table 5): branchiostegal membrane without a posterior free-fold; three barbels on ventral surface of den- tary, one at each posterior margin of three acous- tico-lateralis pores; dorsal fins unpigmented, blue spots present on head. DIsTRIBUTION.—Gilbert (1890) described X. triacanthus from specimens captured at Alba- tross stations located off the coasts of California and Oregon, between approximately 34° and 45° N latitude. Material examined in this study ranged from Point Baja, Baja California, north to Kwat- na Inlet, British Columbia (Fig. 17). Gilbert (1915) reported X. triacanthus occurred in depths from 73 to 364 m; all lots examined for this study fell within that depth range. Comments.—As was the case for_X. /atifrons, Gilbert (1890) did not designate a type-species for X. triacanthus. When the single specimen of 38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 3 TaABLe 6. CHARACTER STATES, NUMBER OF EVOLUTIONARY Steps, AND THE LOCATION OF THE EVOLUTIONARY STEPS FOR THE DATA OF TABLES | AND 2 ON THE CLADOGRAM ILLUSTRATED IN Fic. 18. Num- ber of steps Branches where Character states the steps take place Spination Circumorbital | spines Present, absent 3 6, 9, Bathyagonus nigripinnis Circumorbital 3 spines Present, absent 1 10 Pterotic spines Present, absent 2 2, XYeneretmus rittert Posttemporal spines Present, absent 1 7 Parietal spines Present, absent 1 10 Frontal spines Present, absent 1 9 Nasal spines One, two (0) - Exposed mesethmoid spines Present, absent 0 —- Preopercular spines Present, absent 1 10 2<, 22 2 5; 10 Spines on rostral plate Present, absent 2 YRoul 5, #5 1 3 3, #3 1 Xeneretmus triacanthus 5, Odontopyxis Woes 2 trispinosa Spines on dermal body plates Present, absent 1 9 Free-fold of isthmus Present, absent 4 5, 8:11, Bathyagonus infraspinatus Maxillary barbels 7 : ’ iA - _ , . j 4 7) - - ifs tes See ay Ty 7 = = a ue eee > ie : ‘7 _ oes _ ’ a 7 7 = a ae es - ieee, & “Gray tem, a e Say anton : ao : a iT ooo . aa —— rs Te a ; ae te NS Tia Je yo ae _ Toe Sine sv soars neaal OPy eset ers oes 7a oh a 7 =a) - _ ' way =! - 7 — = ’ a ~ ir ot FAVq=o5 ~~ ee 5 a ie =e : we i (ih a OE Fe i —s eon _ ‘ce v~ 7 7 ' . a) : i a, ke _ : a ~ A — -— = i 7 a Je . A 7 7 co — = > =) "we a U , : en = : » i : 7 - ‘ 7 a : p a6 : » Mite, ” ‘ ‘oon a na) 7 4 i , : ' Laas - >| = ov ° _ aa : To . ion 7 ust 7 7 _ 7 7 »e2 : is -* oe - - ‘ 7 - _ ares i eS a > Ge i 7 aa Ty ¥ = — : i - 7 U _ ‘oi = ae oe | Di we iY # i] + ae) , a a », = » an a -_ oO Leary _— - i va H ; pave 7 1) - it ’ a ie a a : ‘ n) - ' AZ P- - _ - ; =z, 7 re ae ' | Soe ie 7) , nL " i} =o : - > eee. ’ ) rr wr: i in by atl Ns a + ‘i « fa be 4 "yA : “i wing Bah 7 = ’ nhs oe AVS) | ek rae ; a oe ra ie i ae } : ha oir 242 hae Pe ae ie ec - : - ean mn _ - io =_ 7 es @ er pry Lehi Ti. 4 ; alr a i Ss 7 ea ae eee See ae ae _ ii C253X NH PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 5, pp. 55-66, 37 figs., 1 table. February 7, 1986 QUATERNARY BARNACLES FROM THE GALAPAGOS ISLANDS By Victor A. Zullo Department of Earth Sciences, University of North Carolina at Wilmington, North Carolina 28403 ABSTRACT: Quaternary marine deposits on six islands in the Galapagos Archipelago have yielded at least nine species of balanomorph barnacles, seven of which are present in the extant Galapagan fauna. The mid- intertidal species Tetraclita milleporosa Pilsbry and the lower intertidal to subtidal species Megabalanus galapaganus (Pilsbry) and Balanus trigonus Darwin are most common. The whale barnacle Coronula diadema (Linnaeus) and the turtle barnacle Chelonibia testudinaria (Linnaeus) are represented by unique specimens at separate localities. Balanus poecilus Darwin and a shell tentatively identified with B. calidus Pilsbry were found at one locality. A species of Concavus that might represent C. (Arossia) panamensis eyerdami (Henry), and a Tetraclita shell bearing marked similarity to that of 7. rubescens rubescens Darwin were present at single localities. Neither Concavus nor Tetraclita rubescens is known from the extant Galapagan fauna. INTRODUCTION As a participant in the 1964 Galapagos Inter- national Scientific Project, I had an opportunity to study the extant cirriped fauna of the Gala- pagos Archipelago, and to collect a few fossil barnacles from Cerro Colorado on Isla Santa Cruz. Some aspects of the extant fauna were pub- lished (Zullo 1966; Zullo and Beach 1973). The lack of adequate fossil material has prevented any serious speculation on the antiquity of the extant fauna and, indirectly, on the antiquity of intertidal and shallow-water habitats in the Ga- lapagos Archipelago. During February 1982, Carole Hickman, Mat- thew James, Jere Lipps, and Lois and William Pitt made an extensive survey of fossiliferous marine deposits in the Galapagos. Of the 84 sam- ples taken on seven islands, 12 localities on six islands contained barnacle remains (Fig. 36, 37). Lipps and Hickman (1982) argued that none of the Galapagan fossil localities is older than two million years, and that some types of deposit are only a few hundred years old. This conclusion is contrary to previous Miocene or Pliocene age estimates for several of these localities (e.g., Dall and Ochsner 1928; Durham 1964), but the com- pletely modern aspect of the fossil barnacle fauna would appear to support a Quaternary age as- signment. PALEONTOLOGY All of the species represented by fossils are either found today in the intertidal zone or at depths less than 20 m. The mid-intertidal species Tetraclita milleporosa Pilsbry, and the low in- tertidal zone and shelf species Megabalanus galapaganus (Pilsbry) and Balanus trigonus Dar- win are the most abundant fossils. The remaining species, including Balanus sp., cf. B. calidus Pils- bry, B. poecilus Darwin, Concavus (Arossia) sp., cf. C. (A.) panamensis eyerdami (Henry), Tetra- clita sp. indet., Chelonibia testudinaria (Lin- naeus), and Coronula diadema (Linnaeus) are represented by one or a few specimens from single localities. Quaternary distribution of barnacles mirrors modern distribution patterns. Tetraclita [55] 56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 milleporosa and, particularly, Megabalanus gal- apaganus prefer high energy environments and are most abundant and grow to greatest size on windward sides of islands in areas of consider- able wave action. Balanus trigonus, on the other hand, prefers low energy environments on the leeward sides of islands or in protected areas below wave base. Balanus calidus and B. poe- cilus are most common in current-swept areas below wave base. Table | indicates the relationship between fos- sil barnacle occurrences and orientation of lo- calities with respect to prevailing wind direction. The majority of localities yielding specimens of Tetraclita milleporosa and Megabalanus gala- paganus are on southeast-facing shores that pres- ently bear the brunt of wave energies generated by the southeast trade winds. Balanus trigonus, on the other hand, is found predominantly at localities on west-facing, or present leeward shores. Only at CASG locality 61392 does B. trigonus occur with Megabalanus galapaganus. This apparent contradiction can be explained by the presence of Balanus sp., cf. B. calidus and B. poecilus, both subtidal species, and the small size of the Megabalanus galapaganus specimens, typical of subtidal populations. Locality 61392 probably represents a depositional environment below wave base with substantial current action. The two leeward Tetraclita localities are notable in that neither has yielded B. trigonus, suggesting that local wave energies were sufficient to main- tain Tetraclita populations, but too high to per- mit establishment of B. trigonus. ORIGIN OF THE GALAPAGAN BARNACLE FAUNA The major objective sought in this study, but not completely attained, was a clue to the time of origin of the Galapagan barnacle fauna. Clear- ly, the modern Galapagan fauna was already es- tablished in the Pleistocene, and its origins must be looked for in Neogene deposits, if such de- posits exist. This conclusion is supported by studies of north temperate and tropical eastern Pacific Cenozoic barnacle faunas. The major fau- nal break occurs at the Tertiary-Quaternary boundary, with the barnacles of the Pleistocene being essentially of modern aspect, whereas those of the Pliocene are primarily of extinct species- groups that evolved at the end of the Oligocene. The presence of Concavus cannot be adequate- ly explained. The two subspecies of Concavus (Arossia) panamensis (Rogers) range throughout much of the Panamic faunal province (Newman 1982). It is possible that the species has been overlooked in the extant fauna, or was elimi- nated from the fauna in the recent past. SYSTEMATICS Superfamily CoRONULOIDEA Newman and Ross Family CoRONULIDAE Leach Subfamily CHELONIBIINAE Pilsbry Genus Chelonibia Leach Chelonibia testudinaria (Linnaeus, 1767) Figures 3, 4 MarTeRIAL. —One lateral compartment, CASG locality 61281. Discussion.—The single lateral plate in the collection is 31 mm high and has a basal width of 28 mm. The presence of deep cavities between basal septa and well-developed oblique grooves and ridges on the radial and alar edges of the paries readily identify this specimen with C. tes- tudinaria. This common and widely distributed turtle barnacle has been reported from the Pacific loggerhead, green, hawksbill, and ridley turtles. Fossils are known from Miocene and younger TasLe 1. DistRIBUTION OF ENVIRONMENTALLY SENSITIVE GALAPAGAN Fossit BARNACLES WITH Respect TO LocaLity ORI- ENTATION. Orientation of localities* E-facing SE-facing (windward) W-facing (leeward) Species 392 387 281 282 285 286 386 388 389 390 391 Tetraclita milleporosa x Xx Xx Xx x Megabalanus galapaganus >, 4 D4 ? Xx Balanus trigonus »4 xX x >, < Balanus sp., cf. B. calidus Xx Balanus poecilus x * Locality numbers in table are last three digits of CASG numbers (e.g., 6/392). ZULLO: GALAPAGOS QUATERNARY BARNACLES 57 Ficures 1-11. Fig. 1, 2. Coronula diadema (Linnaeus, 1758), basal and side views of shell, hypotype CASG 61364, CASG locality 61229; x 1.3. Fig. 3, 4. Chelonibia testudinaria (Linnaeus, 1767), lateral and interior views of lateral plate, hypotype CASG 61365, CASG locality 61281; x 1.6. Fig. 5-11. Tetraclita milleporosa Pilsbry, 1916. Fig. 5, 6. Exterior and basal views of shell, hypotype CASG 61366, CASG locality 61387; x 1.6. Fig. 7, 8. Basal and exterior views of shell, hypotype CASG 61367, CASG locality 61386; x 1.6. Fig. 9. Interior of scutum, hypotype CASG 61368, CASG locality 61286; x 2.7. Fig. 10. Interior of scutum, hypotype CASG 61369, CASG locality 61286; x 2.7. Fig. 11. Exterior of shell rasped by fish, hypotype CASG 61370, CASG locality 61282; «1.6. 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 deposits in Mediterranean, Atlantic, and Carib- bean regions (Zullo 1982). To my knowledge, this is the first reported fossil occurrence of Che- lonibia from the Pacific basin. Subfamily CORONULINAE Leach Genus Coronula Lamarck Coronula diadema (Linnaeus, 1767) Figures 1, 2 MateRIAL.—One complete specimen, CASG locality 61229. Discussion.—The single specimen is 27 mm in height and its greatest diameter is 31 mm. The well-developed transverse corrugations on the external surfaces of the transverse flanges suggest the ornamentation seen in the Pliocene species C. barbara Darwin, but the absence of similar corrugations on the inner surfaces of the flanges and the lack of infilling between radii and the alar plates indicate that the Galapagos Coronula is merely a highly corrugated specimen of C. diadema. Coronula diadema, with a modern cosmopol- itan distribution on humpback, fin, blue, and sperm whales (Newman and Ross 1976), has been reported from numerous Pliocene and Pleisto- cene localities in the Pacific basin region. Family TETRACLITIDAE Gruvel Subfamily TeTRACLITINAE Gruvel Genus Tetraclita Schumacher Tetraclita milleporosa Pilsbry, 1916 Figures 5-11 Tetraclita porosa var. communis Darwin, 1854:329 (in part). Tetraclita squamosa milleporosa Pilsbry, 1916:257, pl. 60, fig. 1-1d; Newman and Ross 1976:48. MarteriAL.—One shell, CASG locality 61386; one shell, CASG locality 61387; one shell, CASG locality 61391; one shell, CASG locality 61281; 21 shells, two partial shells, CASG locality 61282; 28 shells, four compartmental plates, six scuta, and one partial tergum, CASG locality 61286. DiscussiIon.—The extant tropical American taxa T. milleporosa, T. panamensis Pilsbry, T. stalactifera stalactifera (Lamarck), T. stalactifera confinis Pilsbry, and T. stalactifera floridana Pilsbry form a group within the genus Tetraclita characterized by similarities in shell coloration and opercular plate morphology that readily dis- tinguish them from other Tetraclita species and suggest close phylogenetic relationships. It is as- sumed that 7. milleporosa, known only from the Galapages Archipelago, was derived from a mainland 7. sta/actifera stock. In the eastern Pa- cific, subspecies of 7. stalactifera are restricted to Panamic faunal province mainland localities. Tetraclita panamensis occurs along the Central American Pacific coast, but is also found on Bay of Panama islands. The intertidal Tetraclita of Cocos Island off the coast of Costa Rica appears to be conspecific with 7. panamensis, but may represent a distinct subspecies. The opercular plates of 7. milleporosa are sim- ilar to those of T. stalactifera, but the shell of the Galapagos species differs in being thicker and having much smaller and more numerous pari- etal tubes. Tetraclita milleporosa approaches T. panamensis in thickness and density of small pores, but differs particularly in opercular plate morphology. None of the fossils in the present collections shows any deviation from morphologies exhib- ited by extant 7. milleporosa populations. The shells (Fig. 5-8) are typical of 7. milleporosa in being peltate, and in having tiny orifices, obscure sutures, eroded external surfaces exposing in- filled parietal tubes, and thickened walls with very small and numerous parietal tubes. A few shells show evidence of rasping by fish (Fig. 11). The well-preserved scuta from CASG locality 61286 (Fig. 9, 10) are typical as well, being about as high as wide, with small, closely set denticulae on the inflected occludent margin, and a rela- tively short adductor ridge that nearly merges with the lower part of the articular ridge, being separated by only a shallow groove. The tergum is too worn to be of aid in identification. Tetraclita sp. indet. Figures 12-14 MateriAL.—One shell without opercular plates, CASG lo- cality 61286. Discussion.—A single shell associated with numerous specimens of 7. milleporosa, from CASG locality 61286, represents a second species of Tetraclita. The shell is high conic, with a rel- atively thin shell wall and correspondingly fewer rows of parietal tubes formed of larger individual tubes. The radii are narrow, but well developed and conspicuous, and the exposed filling of the upper parts of the parietal tubes is red. This shell is remarkably similar to that of T. rubescens ru- bescens which presently ranges between San Francisco, California and Cabo San Lucas, Baja California. The combination of shell features, ZULLO: GALAPAGOS QUATERNARY BARNACLES 59 Ficures 12-21. Fig. 12-14. Tetraclita sp. indet., lateral and basal views of shell, hypotype CASG 61371, CASG locality 61286; x 2.5. Fig. 15, 16. Balanus sp., cf. B. calidus Pilsbry, 1916, top and lateral views of shell, hypotype CASG 61372, CASG locality 61392; x 2.5. Fig. 17-19. Balanus poecilus Darwin, 1854, CASG locality 61392; x 2.5. Fig. 17, 18. Top and side views of shells, hypotype lot CASG 61373. Fig. 19. Lateral view of shell, hypotype CASG 61374. Fig. 20, 21. Balanus trigonus Darwin, 1854, CASG locality 61388. Fig. 20. Top view of shells, hypotype lot CASG 61375; = 2.5. Fig. 21. Shells on Anomia peruviana Orbigny, hypotype lot CASG 61376; x 1.6. particularly the color of the internal filling of the species. The Panamic species, related to or con- parietal tubes, is unlike that of 7. milleporosa or — specific with 7. stalactifera, range in color from any of the known Panamic faunal province gray to purple-black, usually lack well-defined 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 radii, and usually have a conic shell with a small orifice. If this unique specimen 1s indeed repre- sentative of T. rubescens, 1am ata loss to explain its presence in the Pleistocene Galapagan fauna. Superfamily BALANoIDEA (Darwin) Newman and Ross Family BALANIDAE Darwin Subfamily BALANINAE Darwin Genus Balanus Da Costa Balanus trigonus Darwin, 1854 Figures 20, 21 MAaArTeRIAL.—Ten shells, CASG locality 61388; two shells, CASG locality 61389; one shell, CASG locality 61390; six shells, CASG locality 61392. Discussion. — Although shells characteristic of B. trigonus were collected at the four localities listed above, no opercular plates were present in the collections. Balanus trigonus is found in most of the warm-water regions of the world from the lower intertidal zone to the edge of the shelf, but is most common in those parts of the immediate subtidal and inner-shelf zones that are protected from wave shock. In the Galapagos Archipelago, extant B. trigonus is abundant on the leeward sides of islands in the lower intertidal and im- mediate subtidal. Considering the widespread distribution and abundance of this species in modern shallow seas, few verifiable reports of fossil B. trigonus exist. The species is fairly common in Pleistocene de- posits of the Gulf of California region (Ross 1962) and has been identified by William A. Newman (personal communication 1982) from the Pleis- tocene of Hawaii. To my knowledge, fossil B. trigonus has not been reported from the western margins of the Pacific basin. Western Atlantic reports of B. trigonus include those of Withers (1953) from the (?)Miocene of Cuba, and of Ross (1965) from the Pliocene Tamiami Formation of Florida. Ross’s (1964) report of this species from the Pliocene Yorktown Formation of Vir- ginia was later stated to be in error (Ross 1965). Western Tethyan reports include those of Ko- losvary (1957) from the Tortonian (Miocene) of Hungary, and Davadie (1963) from the Pliocene of Italy, the Red Sea, and the Coralline Crag of England. These fossil occurrences, coupled with the modern distribution of the species, would suggest that B. trigonus is an old Tethyan element that has managed to survive to the present. There are two problems, however, that cause me to ques- tion this conclusion. First, many of the afore- mentioned reports are based on species lists without substantiating descriptions or illustra- tions. Their validity is placed in question par- ticularly in the knowledge that other students of barnacles who have monographed the faunas of the same regions (e.g., Darwin 1854; Alessandri 1906; Menesini 1966) did not uncover B. tri- gonus. Secondly, although both extinct and ex- tant representatives of the B. trigonus complex are common in Neogene and Pleistocene depos- its of southern California and the southeastern United States that I have examined (e.g., Zullo 1979), B. trigonus is absent. This 1s particularly odd, because the faunas of these units indicate that hydroclimates were substantially the same or warmer than those in the same regions today, and that depositional environments were fully within the present bathymetric range of B. tri- gonus. The origin and historical biogeography of B. trigonus remain in doubt, and their resolution will, in part, be dependent on a thorough eval- uation of previously reported occurrences. Balanus sp., cf. B. calidus Pilsbry, 1916 Figures 15, 16 MarerIAL.—One complete shell, CASG locality 61392. Discussion. —A single shell, lacking opercular plates, is tentatively identified with B. calidus based on its coarsely ribbed, volcaniform shell and small orifice. Only a few extant specimens of B. calidus were obtained during the 1964 ex- pedition, and all came from shallow, subtidal depths. Off the East Coast of the United States, B. calidus is found on the shelf at depths below significant wave action. Balanus poecilus Darwin, 1854 Figures 17-19 Balanus poecilus Darwin, 1854:246, pl. 5, fig. 3a, b; Henry 1960:142, pl. 2, fig. a, c, d, pl. 5, fig. b-d. MArTEeRIAL.—Eight shells without opercular plates, CASG locality 61392. Discussion. — The “west coast of South Amer- ica, Mus. Cuming; attached to an Avicula” was cited by Darwin (1854) as the type and only lo- cality in his original description of B. poecilus. The species went unreported until Henry (1960) obtained some individuals of Pteria sterna (Gould) from the vicinity of Guaymas in the Gulf ZULLO: GALAPAGOS QUATERNARY BARNACLES of California. Because of the unusually broad distribution indicated by the recorded occur- rences, and because of the ambiguity of the type locality, I requested the aid of J. P. Harding, British Museum (Natural History) in attempting to refine these data through identification of the “‘Avicula” to which the types are attached. Dr. Harding kindly located the type-lot and for- warded the following information provided by S. P. Dance (personal communication August 5, 1965): The shell to which the type specimens of Ba/anus poe- cilus Darwin are attached closely resembles a recently described species, Preria beilana Olsson. The type lo- cality for this species is Venado Beach, Canal Zone, Pan- ama. Pteria peruviana Reeve may be an earlier name for this taxon but there is not enough material in the British Museum (Natural History) collections to decide this. Whichever name is used for it there can be little doubt that the shell to which the Darwinian barnacles are at- tached is a member of the Panamic-Pacific faunal prov- ince. Dr. Harding also reported that the specimens bear the label ““West coast of America,” rather than South America, and as it is known that Hugh Cuming made extensive collections on the west coast of Central America, and especially in Panama during the period 1832-1856 (Keen 1958:2), it seems likely that the types of B. poe- cilus are from the same region. Based on collections made during the 1964 Galapagos expedition, and previously unre- ported specimens in the collections of the Allan Hancock Foundation and the California Acad- emy of Sciences, Balanus poecilus is found to range throughout the Panamic faunal province. The Allan Hancock Foundation collection in- cludes specimens from off San Pedro Nolasco Island in the Gulf of California, off Jicarita Island and Bahia Honda, Panama, off Gorgona Island, Colombia, and off La Libertad, Ecuador, as well as from the Galapagos off Gardiner Island, near Espanola. Genus Concavus Newman Subgenus Arossia Newman Concavus (Arossia) sp., cf. C. (A.) panamensis eyerdami (Henry, 1960) Figures 22-24 MATERIAL.—Two shells without opercular plates, CASG locality 61390. Discussion.—The genus Concavus is not known to be represented in the extant Galapagan 61 fauna. According to Newman (1982), modern representatives of this Tethyan Tertiary genus are restricted to the eastern Pacific, ranging from San Francisco, California to Valparaiso, Chile. Newman (1982) established two subgenera for extant species: Menesiniella for C. aquila (Pils- bry) and C. regalis (Pilsbry); and Arossia for C. henryae Newman, C. panamensis panamensis (Rogers), and C. panamensis eyerdami (Henry). The Galapagan fossils, with their plicate, but not regularly or strongly ribbed parietes, appear to be assignable to Arossia in the absence of the more definitive features of the opercular plates. Within Arossia, these fossil shells most closely approach those of C. panamensis eyverdami in having a high conic shell with the rostrum higher than wide, a straight carina, and no evidence of beaded growth lines. The preserved reddish-pur- ple coloration of the shell and the closely spaced transverse septa appear to distinguish the fossils from C. henryae, the sole representative of Con- cavus in the Peruvian faunal province. Subfamily MEGABALANINAE Newman Genus Megabalanus Hoek Megabalanus galapaganus Pilsbry, 1916 Figures 28-35 Balanus tintinnabulum galapaganus Pilsbry, 1916:70, pl. 12, fig. 1-1b. MArTerIAL.—Eight shells, 5 shell fragments, CASG locality 61281; 2 partial shells, CASG locality 61282; 24 shells, 26 compartmental plates, two bases, four scuta, and one tergum, CASG locality 61286; 2 shells, CASG locality 61392. Discussion.—After Tetraclita milleporosa, shells of Megabalanus galapaganus are the most abundant barnacle fossils obtained during the 1982 expedition. The specimens range from re- cently settled juveniles to mature individuals over 5 cm in height and 4 cm in greatest diameter. Many individuals retain the parietal color or col- or striping, and the parietal spines characteristic of extant populations. The opercular plates, al- though somewhat worn, are typical for 7. gala- paganus. The scutum is flat, bears a well-defined adductor ridge, and lacks a definite lateral de- pressor muscle pit. The tergum has a longer and narrower spur than the closely related species M. clippertonensis (Zullo) from Clipperton Island and M. tanagrae (Pilsbry) from the Hawaiian Islands (Zullo 1969). Extant M. galapaganus 1s relatively abundant in low intertidal rocky areas subject to heavy 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 ¥ re cel be Cal, ge NO ol FiGures 22-27. Fig. 22-25. Concavus sp., cf. C. (Arossia) panamensis eyerdami (Henry, 1960), CASG locality 61390; x 1.6. Fig. 22, 23. Lateral views of shell, hypotype CASG 61377. Fig. 24. Basal view of shell, hypotype CASG 61378. Fig. 25. Broken radial sutural edge showing tubes, hypotype CASG 61378. Fig. 26, 27. Megabalanus sp. indet., lateral views of shell, hypotype CASG 61379, CASG locality 61285; x 2.5. wave shock. It is in this region that the species reaches its maximum size. At subtidal depths specimens are locally abundant on lobster car- apaces, gastropod shells, and coral heads, but rarely attain more than 2 cm basal diameter and are usually less than | cm high. Pilsbry (1916) based this species on specimens from the inter- tidal of Espanola Island. Collections made during the 1964 expedition and augmented by collec- tions from the Allan Hancock Foundation and the California Academy of Sciences extend the range of M. galapaganus not only through most of the Galapagos Archipelago but to Cocos Is- land (Costa Rica) to the north and Port Utria, Colombia on the South American mainland. Megabalanus sp. indet. Figures 26, 27 MatTerRIAL.—One shell without opercular plates, CASG lo- cality 61285. ZULLO: GALAPAGOS QUATERNARY BARNACLES 63 x. i) Ficures 28-35. Megabalanus galapaganus (Pilsbry, 1916). Fig. 28, 29. Exterior and interior of tergum, hypotype CASG 61380, CASG locality 61286; x 1.3. Fig. 30, 31. Exterior and interior of scutum, hypotype CASG 61381, CASG locality 61286; x 1.3. Fig. 32, 33. Interior and exterior of scutum, hypotype CASG 61382, CASG locality 61286; «1.3. Fig. 34. Lateral view of shells, hypotype lot CASG 61383, CASG locality 61286; = 1.0. Fig. 35. Top view of shell clump, hypotype CASG 61384, CASG locality 61282; «1.0. Discussion.—The single barnacle specimen _ to question its identification. The shell is 15 mm from CASG locality 61285 differs sufficiently high, 22 mm in carinorostral diameter, and is from the typical growth form of M. galapaganus low conic, rather than cylindric to subglobose in 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 |. Pinta / GALAPAGOS ISLANDS a4 G @® 7 |. Genovesa |. Marchena I. San Salvador 61387 61388- o | Rabida 61391 mI. Baltra wa. 61392 |. Santa Cruz | ace yt I. ; dues see Fig.37 San Cristobal I. Santa Fe 61229 E11" “9 I. ’ Santa Maria IL. Espanola Ficure 36. Generalized map of CASG Galapagos localities containing fossil barnacles (map provided by J. H. Lipps). 4 shape. The sub-diamond-shaped orifice is, re- February 1982. Chelonibia testudinaria, Tetraclita sultingly, rather small, and the radii are corre- milleporosa, Megabalanus galapaganus. spondingly narrow. The parietes show no evi- ©1282 Isla Santa Fe. Fossils from top of sedimentary sequence shia overlain by basalt. Red, tuffaceous, crossbedded sand- dence of color or striping, and bear low, rounded, stone with stratified fossils at top, about 8 m above sea irregular ribs. In the absence of opercular plates, level. Same horizon as CASG locality 61281, but 30 however, there is no way to determine its iden- m farther seaward. Collected 14 February 1982. Tet- tity. raclita milleporosa, Megabalanus galapaganus. 61285 Isla Santa Fe. Terrace deposits of boulders, cobbles, pebbles, and sand containing molluscs and barnacles Locatity DESCRIPTIONS at top of cliff in small cove at landing site; 3.5—-4 m above sea level. Locality is north of CASG locality All barnacle specimens are in the collection of 61282. Collected 15 February 1982. Megabalanus sp. the Department of Geology, California Academy net of Sciences, San Francisco (CASG). Locations of 61286 Isla Santa Fe. Terrace deposit about 100 m from shore collection sites are shown in Figures 36 and 37. near eastern end of south coast. Loose, white to tan, medium- to coarse-grained sand containing many bar- 61229 Isla Isabella. White to tan, loose, silty sand containing nacles. Collected 15 February 1982. Tetraclita mille- abundant shells at site of airport at Villamil. Collected porosa, Tetraclita sp. indet., Megabalanus galapaga- 3 February 1982. Coronula diadema. nus. 61281 Isla Santa Fe. Beach deposit about 8 m above sea level 61386 Isla San Salvador, James Bay. Shelly, basaltic sand in on southeast shore north of Punta Miedo. Calcareous, line of trees north of mining camp. Collected 8 Feb- sometimes stratified sand up to 2.5 m thick and inter- ruary 1982. Tetraclita milleporosa. mixed with basalt boulders and cobbles. Collected 14 61387 Isla Rabida. Storm-tossed shell and bone in small, cliff- ZULLO: GALAPAGOS QUATERNARY BARNACLES SA Sa Ip ( ron ST \ Cie) : mI 0°50'S + “lea, 90°05’W ZT Mea Sng Fe CAS 61285* 2a CAS 61283 ML S Las CAS 61284 300 CAS 61282* O Po freas 61281* CAS 61286 * ay ISLA SANTA FE (@) km 4 EE contour interval in feet from U.S. Naval Hydrographic chart 5939 Ficure 37. backed cove on south side of island. Collected 9 Feb- ruary 1982. Tetraclita milleporosa. 61388 Isla Baltra. Bedded to crossbedded, reddish-brown, silty sandstone with abundant Codakia shells at basal con- tact (Unit 4). South shore of Caleta Aeolian, directly south of Punta Noboa. Collected 10 February 1982. Balanus trigonus. 61389 Isla Baltra. Crossbedded, white sandstone containing shell debris and abundant pectinids (Unit 1). Locality about 30 m east of CASG locality 61388 along a 100-m stretch of exposure. Collected 10 February 1982. Bal- anus trigonus. 61390 Isla Baltra. Basal 0.5—1.5-m-thick boulder and cobble bed containing abundant coralline algae and casts and molds of molluscs. Same area as CASG locality 61389. Collected 10 February 1982. Balanus trigonus, Con- cavus (Arossia) sp., cf. C. (A.) panamensis everdami. 61391 Isla Baltra. Bulldozed pit (old anti-aircraft gun em- placement) about 170 m back of sea cliff. Collected 10 February 1982. Tetraclita milleporosa. 61392 Isla Santa Cruz, Cerro Colorado. Fossils from top of limestone shelf on north side of Cerro Colorado. Col- lected 17 February 1982. Balanus trigonus, Balanus sp., cf. B. calidus, B. poecilus, Megabalanus galapaga- nus. ACKNOWLEDGMENTS I thank Jere H. Lipps, Department of Geology, University of California, Davis, and William D. Collecting sites on Isla Santa Fe. CASG localities with asterisks yielded barnacles (map provided by J. H. Lipps). Pitt, Sacramento, California for providing the specimens, maps, and locality data used in this study, and for their advice during manuscript preparation. Funding for this study was provided by the Marine Sciences Research Program of the University of North Carolina at Wilmington. This paper 1s contribution number 372 of the Charles Darwin Foundation. LITERATURE CITED ALESSANDRI, G. DE. 1906. Studi monografici sui Cirripedi fossili d'Italia. Palaeontogr. Italica 12:207-324. Dati, W. H. AND W. H. Ocusner. 1928. Tertiary and Pleis- tocene Mollusca from the Galapagos Islands. Proc. Calif. Acad. Sci., ser. 4, 17:89-139. Darwin, C. 1854. A monograph on the sub-class Cirripedia, the Balanidae, the Verrucidae. Ray Society, London. 684 pp. Davapigz, C. 1963. Systématique et structure des Balanes d'Europe et d’Afrique. Editions Centre Natl. Recherche Scient. 146 pp. Durnam, J. W. 1964. The Galapagos Islands expedition of 1964. Am. Malacol. Union, Inc. Annu. Rep. 1964:53. Henry, D. P. 1960. Thoracic Cirripedia of the Gulf of Cal- ifornia. Univ. Washington Publ. Oceanogr. 4(4):135-158. Keen, M.A. 1958. Seashells of tropical west America. Stan- ford Univ. Press, Calif. 624 pp. 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 5 KotosvAry, G. 1957. Enumeration des Balanides fossiles de la Hongne. Bull. Soc. Linn. Lyon 26(2):30-32. Linnaeus, C. 1758. Systema naturae. Holmiae, Editio De- cima, Reformata, Vol. |. 824 pp. . 1767. Systema naturae per regna tria naturae—editio duodecima, reformata. Holmiae 1(2):533-1327. Lipps, J. H. AnD C. S. HickMAN. 1982. Paleontology and geologic history of the Galapagos Islands. Abs. with Pro- grams, Geol. Soc. Am. 14(7):548. Menesini, E. 1966. I balani Miocenici delle “Arenarie di Posano” (Volterra, Provincia di Pisa). Palaeontogr. Italica 60:99-129. Newman, W. A. 1982. A review of extant taxa of the “Group of Balanus concavus” (Cirripedia, Thoracica) and a proposal for genus-group ranks. Crustaceana 43:25-36. Newman, W. A. AND A. Ross. 1976. Revision of the ba- lanomorph barnacles; including a catalog of the species. San Diego Soc. Nat. Hist. Mem. 9:1-108. Pitspry, H. A. 1916. The sessile barnacles (Cirripedia) in the collections of the U.S. National Museum; including a mono- graph of the American species. U.S. Natl. Mus. Bull. 93:1- 366. Ross, A. 1962. Results of the Puritan-American Museum of Natural History expedition to western Mexico, 15. The lit- toral balanomorph Cirripedia. Am. Mus. Novitates 2084: 1-44. 1964. Cirnpedia from the Yorktown Formation (Miocene) of Virginia. J. Paleontol. 38:483-491. 1965. A new barnacle from the Tamiami Miocene. J. Fla. Acad. Sci. 27(4):273-277. Witners, T. H. 1953. Catalogue of fossil Cirripedia in the Department of Geology, Vol. III. Tertiary. British Mus. (Nat. Hist.), London. 396 pp. Zutto, V. A. 1966. Zoogeographic affinities of the Balano- morpha (Cirripedia: Thoracica) of the eastern Pacific. Pp. 139-144 in The Galapagos, R. I. Bowman, ed. Univ. Cal- ifornia Press, Berkeley. 1969. A new subspecies of Balanus tintinnabulum (Linnaeus, 1758) (Cirripedia, Thoracica) from Clipperton Island, eastern Pacific. Proc. Calif. Acad. Sci., ser. 4, 36(16): 501-510. 1979. Thoracican Cirripedia of the lower Pliocene Pancho Rico Formation, Salinas Valley, Monterey County, California. Contrib. Sci., Nat. Hist. Mus. Los Angeles Co. 303:1-13. . 1982. Anew species of the turtle barnacle Chelonibia Leach, 1817, (Cirripedia, Thoracica) from the Oligocene Mint Spring and Byram formations of Mississippi. Mississippi Geol. 2(3):1-6. Zu.io, V. A. AND D. B. Beacu. 1973. New species of Mem- branobalanus Hoek and Hexacreusia Zullo (Cirripedia, Ba- lanidae) from the Galapagos Archipelago. Contnb. Sci., Nat. Hist. Mus. Los Angeles Co. 249:1-16. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 11 C253Xx NH PROCEEDINGS OF THE CALIFORNIA ACADEMY OF CTENGES Vol. 44, No. 6, pp. 67-109, 28 figs., 2 tables. SMITHSON ar 2 6 1085 A SYSTEMATIC REVIEW OF AMPHIZOID BEETLES RELATIONSHIPS TO OTHER ADEPHAGA By David H. Kavanaugh Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Asstract: Rediscovery of type material for Amphizoa davidi Lucas, 1882, the only known Palaearctic amphizoid, is reported, with a lectotype designated, and the type area emended (from Tibet to Szechwan Province, China). A key is provided for identification of adults of the four known amphizoid species. Form and structure, geographical and habitat distributions, and geographical relations with other taxa are described and illustrated for each species. Amphizoa carinata Edwards, 1951, is recognized as a junior synonym of A. lecontei Matthews, 1872. Through cladistic analysis, using out-group and character correlation criteria, and a review of known Mesozoic fossil material, a hypothesis of phylogenetic relationships among extinct and extant Adephaga is developed, discussed, and related to geologic time. A semi-aquatic, rather than terrestrial, common ancestor is proposed for Adephaga. Amphizoids diverged from their sister-group, which includes all Hydradephaga except haliplids, in Triassic time. All extant amphizoid species had differentiated by late Pliocene time, in response to a series of vicariant events. Quaternary climatic and geologic events resulted in changes in geographical distributions of these species and structural, physiological, and behavioral adapta- SE eee ee tions of their members. TABLE OF CONTENTS Evidence for relationship between amphizoids and trachypachids A hypothesis of adephagan PAGE phylogeny Introduction 68 Phylogenetic relationships Materials and Methods... ees Sees 5p) of amphizoid species. Systematics of Amphizoidae...... 69 Zoogeography and Evolution Introduction... eae Dees re rete 69 Present pattern of amphizoid A Key for Identification of distnibutionte2= : Amphizoa Adults... 70 Mesozoic events and the Amphizoa davidi Lucas... 70 origin of amphizoids...... Amphizoa insolens LeConte................ eZ. Tertiary events and Amphizoa striata Van Dyke... 75 amphizoid radiation........ Amphizoa lecontei Matthews... = 75 Quaternary history and development Phylogeny... ms 78 of the present amphizoid fauna Bayorenene Pnnonehine of Prospectus for Future Research... amphizoids .... 81 Acknowled amen ts sess se seer eee Evidence from ine Focal ees al 84 Literature Cited... : (AMPHIZOIDAE: COLEOPTERA) AND THEIR PHYLOGENETIC 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 INTRODUCTION For several decades, the location of type-ma- terial for Amphizoa davidi Lucas, 1882, de- scribed from Tibet, remained a mystery (Ed- wards 1951; Kavanaugh 1980; Kavanaugh and Roughley 1981). Although material Lucas stud- ied was known to have been deposited in the Muséum National d’Histoire Naturelle in Paris, several independent efforts to locate specimens of A. davidi in appropriate parts of that collection had failed. Equally perplexing was the fact that no additional specimens representing this taxon had been found since its original description. Amphizoa davidi is an especially important taxon for two reasons. First, it is a member of the small family Amphizoidae, which 1s consid- ered by many workers to represent an interme- diate evolutionary grade between the so-called Geadephaga, or terrestrial Adephaga (1.e., Ca- rabidae, in the broadest sense), and the remain- ing Hydradephaga, or aquatic Adephaga (i.e., Dytiscidae, Hygrobiidae, Gyrinidae, etc.). Amphizoa davidi Lucas: lectotype male, dorsal Ficure | aspect, total length = 11.4 mm. Knowledge of amphizoids is seen as a major key to understanding adephagan evolution and phy- logeny; and knowledge of 4. davidi in particular is critical for understanding Amphizoidae. Sec- ond, Amphizoa kashmirensis Vazirani, 1964:145, described from the Himalaya of India, has re- cently been shown to be a dytiscid, referable to genus Hydronebrius Jakovlev, rather than an am- phizoid (Kavanaugh and Roughley 1981). As a result, 4. davidi is the only known Palaearctic amphizoid; and because no specimens of this species had ever been seen by current workers, doubts had arisen with regard to its familial af- finities (Kavanaugh 1980; Kavanaugh and Roughley 1981). What are the phylogenetic re- lationships between A. davidi and the Nearctic species, and what are the zoogeographic impli- cations of this phylogeny and the disjunct dis- tribution of genus Amphizoa? Answers to these questions might shed new light on the origins and history of the Holarctic fauna in general and of certain relict, taxonomically isolated taxa in particular. In early 1983, Terry L. Erwin (U.S. National Museum, Washington, D.C.) discovered several amphizoids pinned in one corner of a Schmidt box labelled “‘Australian Carabidae” at the Mu- séum National d’Histoire Naturelle in Paris. In- cluded were a few specimens of Amphizoa in- solens LeConte from western North America and one specimen (Fig. 1) from Mou-pin, Tibet, the type-locality for A. davidi. Suspecting that he had found the long-sought type of 4. davidi, Erwin arranged for shipment of the specimen to me on loan. Jean Menier, curator at the museum in Paris, provided photocopies of relevant entries in the museum’s catalog, specifically for the accession of material from Mou-pin, Tibet, re- ceived from Armand David and upon which Lu- cas’s description was based. Subsequently, I have determined that the specimen is the type-speci- men of Amphizoa davidi Lucas through a study of the specimen itself and the labels it bears (in- cluding one with the proper catalog number). The purposes of this paper are: (1) to report on the rediscovery of type-material for 4. davidi Lucas; (2) to designate a lectotype for same; (3) to redescribe this material in comparison with Nearctic forms, and illustrate certain character- istics of form and structure for the first time; (4) to update distributional records that have ac- cumulated since Edward’s (1951) revision of the family; (5) to propose one new synonymy; (6) to KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES provide a revised key to species that reflects new findings; and (7) to initiate consideration of the phylogenetic relationships among extant Am- phizoa species and the zoogeographic implica- tions of these relationships. A cladistic analysis of relationships among major extant and known extinct adephagan groups is presented as a basis for the intrageneric analysis of Amphizoa. MATERIALS AND METHODS Descriptions of form and structure, taxonomic conclusions, geographical distributions, and oth- er findings reported here are based on exami- nation of more than 1,250 adult specimens of Amphizoa and more than 300 specimens rep- resenting other extant adephagan taxa. The fol- lowing acronyms are used in the text to refer to collections from which specimens were received for study and/or in which specimens are depos- ited. Curators and collecuuon managers respon- sible for these collections are also listed, and I thank them sincerely for their help in providing specimens on loan for study. BMNH British Museum (Natural History), London SW7 5BD, England; M. E. Bacchus. BYUM Brigham Young University Museum of Natural His- tory, Provo, Utah 84602; R. W. Baumann. CAS — California Academy of Sciences, San Francisco, Cal- ifornia 94118; D. H. Kavanaugh. CNC Canadian National Collection of Insects, Biosys- tematics Research Institute, Ottawa, Ontario K1A 0C6; A. Smetana. DMad_D. Maddison, University of Alberta, Edmonton, Al- berta T6G 2E3. GCha_ G. Challet, Orange County Vector Control District, Garden Grove, California 92643. GLPa_ G. L. Parsons, Oregon State University, Corvallis, Oregon 97331. GLPe_ G.L. Peters, Oregon State University, Corvallis, Or- egon 97331. LGBe _L. G. Bezark, California State Department of Food and Agriculture, Sacramento, California 95814. MCZ Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, Massachusetts 02138; A. F. Newton, Jr. MNHP Muséum National d’Histoire Naturelle, Paris, 75005 France; J. Menier. Nevada State Department of Agriculture, Reno, Ne- vada 89504; R. C. Bechtel. OSUO_ Oregon State University, Corvallis, Oregon 97331; J. Lattin, G. L. Peters. NSDA PUCA Pacific Union College, Angwin, California 94508; L. E. Eighme. RERo_ R. E. Roughley, University of Manitoba, Winnipeg R3T 2N2. SJSU San Jose State University, San Jose, California 95114; J. G. Edwards. UASM University of Alberta, Strickland Museum, Edmon- ton, Alberta T6G 2E3; G. E. Ball. 69 UCB University of California, Essig Museum of Entomol- ogy, Berkeley, California 94720; J. A. Chemsak and G. Ullrich. UCD University of California, Davis, California 95616; R. O. Schuster. USNM United States National Museum, Smithsonian Insti- tution, Washington, D.C. 20560; P. J. Spangler, T. L. Erwin. UZMH Universitetets Zoologiska Museum, Entomologiska Avdelningen, SF-00100 Helsingfors 10, Finland; H. Silfverberg. Methods, including techniques for dissection of male and female genitalia and criteria for rank- ing taxa, are discussed by Kavanaugh (1979). The only measurement used in this paper, stan- dardized body length (SBL), is the sum of three measurements: length of head along midline from apical margin of labrum to a point opposite pos- terior margin of left eye; length of pronotum along midline from anterior to posterior margin; and length of elytron along midline from apex of scu- tellum to a point opposite apex of longer elytron. Line drawings were made with the aid of a camera lucida attached to a Wild Model M-5 stereoscopic dissecting microscope. The scan- ning electron micrograph (Fig. 2) was obtained using a Hitachi model S-520 SEM (with accel- erating voltage = 5 kV and specimen uncoated). Cladistic analyses were carried out using man- ual methods (see Phylogeny below for further discussion); but results were compared with those generated using “WAGNER” and “SOKAL” programs from the “Phylogenetic Inference Package” (PH YLIP) for microcomputers created by J. Felsenstein (University of Washington, Se- attle), as modified by T. K. Wilson (Miami Uni- versity, Oxford, Ohio). In general, cladograms obtained using manual and computer-assisted methods were similar. However, placement of individual taxa in cladograms generated by the PHYLIP programs varied markedly, subject to changes in the order in which taxa were listed in the database (and, therefore, compared by com- puter). SYSTEMATICS OF AMPHIZOIDAE Introduction Edwards’s (1951) monograph of Amphizoidae stands as the definitive systematic treatment of this group. His extensive review of the literature and detailed descriptions, comparative studies, and discussions of form and structure serve as a sound basis for all subsequent work on amphi- 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 zoids, as well as for comparisons of members of this group with those of other adephagan taxa. Because of more liberal institutional lending policies, I was able to borrow type-material that was unavailable to Edwards and designate lec- totypes for Amphizoa insolens LeConte, A. jo- sephi Matthews, and A. /econtei Matthews (Ka- vanaugh 1980). Moreover, a great deal of new material has been collected during the past 30 years. Through loans and my own fieldwork, I have had access to almost five times as many specimens as Edwards studied, many of these from areas in which amphizoids were previously unknown. These new distributional records have important taxonomic and zoogeographic impli- cations. In order to make this report minimally redundant with respect to Edwards’s paper, I have limited my descriptive presentations to brief list- ings and discussions of distinguishing character- istics, except where my findings depart from Ed- wards’s. The reader should consult Edwards (1951) for more detailed descriptive and com- parative information on amphizoids, as well as comprehensive coverage of the literature prior to that date. The format used for presentation of Amphizoa species below is as follows: (1) a synonymy (in- cluding for each name the author, date, and page citation for original description; status, sex, and depository for holotype or lectotype; type-local- ity; and literature citations that were not listed by Edwards [1951]); (2) additional comments on nomenclature, type-specimens, and/or type- locality as needed; (3) a brief listing of distinguish- ing characteristics of adults, with additional dis- cussion of form and structure as needed; (4) habitat distribution; (5) geographical distribu- tion, including distributional summary state- ment, map illustrating known localities, and for- mal listing of localities for specimens studied (with area and month[s] of collection, number of spec- imens studied, and depository[ies] for same); and discussions of (6) geographical variation; and (7) geographical relationships with other Amphizoa species. A Key for Identification of Amphizoa Adults Ne Elytron (Fig. 6b) with blunt but distinct carina on fifth interval, area medial to carina elevated, flat, area lateral to ca- rina slightly concave uu... Amphizoa lecontei Matthews Me Elytron (Fig. 3b, 4b, 5b) evenly convex or slightly concave paralaterally, with- out carina . fy 2. . Prosternal intercoxal process (Fig. 1) short, round; body form narrower (Fig. 3a); specimen from southwestern China (Fig. 17) . Amphizoa davidi Lucas De Prosternal intercoxal process (Fig. 12) long, spatulate; body form (Fig. 4a, 5a) relatively broader; specimen from west- ern North America ...... mS) . Elytral silhouette (Fig. 5a) broad basally and distinctly narrowed subapically, elytral surface only faintly rugose in lat- eral one-half; pronotum (Fig. 9) broad- est at base, with lateral margins not or only slightly crenulate a Amphizoa striata Van Dyke BN Elytral ‘silhouette (Fig. 4a) subovoid, slightly narrowed basally, slightly broader subapically, elytral surface moderately or coarsely rugose in lateral one-half; pronotum (Fig. 8) at least as broad at middle as at base, with lateral margins markedly crenulate .. Amphizoa insolens LeConte Amphizoa davidi Lucas (Figures 1-3, 7, 11, 13, 17) Amphizoa davidis Lucas, 1882:157 [incorrect spelling]. Lec- totype (here designated), a male, in MNHP, labelled: ““Mu- seum Paris, Mou-pin, A. David 1870°/ “398"/ “774 70” [yellow-backed disk]/ ““Amphizoa davidis, Lucas” [label double-pierced by pin, hence vertical on pin]; “Type” [red label]/““Muséum Paris”/ ‘‘Lectotype Amphizoa davidi Lu- cas designated by D. H. Kavanaugh 1983” [red label]. Type- Locality.— Pao-hsing, Szechwan Province, People’s Repub- lic of China. Edwards 1951:322. Kavanaugh 1980:289. Amphizoa davidi Lucas [justified emendation]. Edwards 1951: 322. Kavanaugh 1980:289. Kavanaugh and Roughley 1981: 269. Leech and Chandler 1956:301. Notes ON NOMENCLATURE AND TYPE- SPECIMEN.—Mou-pin, Tibet, the area originally cited as type-locality, is now called ““Pao-hsing”’ (30°22'N, 102°50’E). This region is no longer part of Tibet, but rather the western part of Szechwan Province, People’s Republic of China. DISTINGUISHING CHARACTERISTICS. —Size small, SBL male = 11.4 mm; body form (Fig. 1, 3a) narrow; body color piceous, with antennae, maxillary and labial palpi, and tarsi rufopiceous; head (Fig. 2) finely and densely punctate; prono- tum (Fig. 2) coarsely and densely punctate, with KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES areas between punctures convex, granulate in ap- pearance; elytra finely and densely punctate, slightly rugose at base and in lateral one-fourth; pronotum (Fig. 7) broadest at base, with lateral margins arcuate at middle, markedly sinuate an- terior to basal angles, not crenulate, median lon- gitudinal impression present but faintly im- pressed; prosternal intercoxal process (Fig. 11) short, round; posterolateral angle of proepister- num and posteromedial angle of proepipleuron abut evenly to form smooth prothoracic margin (see Edwards 1951:321, “Plate 4°’); elytral sil- houette (Fig. 3a) moderate in width basally and distinctly narrowed subapically, elytra (Fig. 3b) evenly convex, without carinae: elytral striae complete but faintly impressed and finely punc- tate; front tibia with posterodorsal groove pres- ent on apical three-fourths, with fringe setae in groove very short and restricted to apical one- half; male median lobe (Fig. 13) with shaft slen- der at middle, evenly arcuate ventrally, apex slightly deflected ventrally, left paramere narrow basally, with vestiture restricted to apical one- fourth; female unknown; specimen from south- western China (Fig. 17). Edwards’s description of A. davidi (1951:322) was an English translation of the original de- scription in French (Lucas 1882). Based on my examination of the type-specimen, additional comments and certain amendments to the orig- inal description seem appropriate. Lucas de- scribed the type of 4. davidi as “noir mat ... avec les palpes ... d’un brun teinté de ferrugi- neux. Les antennes ... d’un brun ferrugineux brilliant” (i.e., dull black, with reddish-brown antennae and palpi). In my view, the specimen is as dull as adults of A. insolens and A. striata but less dull than adults of 4. lecontei. Its body color is piceous, not black as in A. insolens adults: and its antennae and palpi are rufopiceous, not reddish-brown. The median longitudinal impression (median furrow), which was de- scribed as “ne presente pas” (i.e., absent), is pres- ent and as deeply impressed as in A. striata adults, less so than in A. /econtei and A. insolens mem- bers. Lucas described the scutellum as “‘tres fine- ment chagriné” (very finely granulate); but be- cause this character state is shared with adults of the other Amphizoa species, it is of no taxo- nomic use. According to the original description, the elytral striae are “‘les parcourent obsoléte- ment accusées et non ponctuées” (obsolete and ———— —— — — rs 71 FiGure 2. Amphizoa davidi Lucas: scanning electron mi- crograph of head and pronotum, dorsal aspect, magnification = 25 (specimen uncoated). impunctate); but they appear to be complete and clearly (although very shallowly) impressed. Due to dense punctation of the entire elytral surface, it is difficult to distinguish the fine punctures which are found in the striae. Close examination of the elytra of the lectotype of A. davidi has revealed a previously unrecorded feature. Due to the relatively faint development of macrosculpture on the elytral surface of this specimen, I found small but distinctly foveate punctures on the third, fifth, seventh, and ninth intervals. No setae appear to be associated with these punctures. Identical punctures were sub- sequently found in adults of all three Nearctic Amphizoa species, although they are much less obvious in Nearctic specimens, least so in 4. striata adults. Similar, but seta-bearing, punc- tures are found among adults ofa broad spectrum of tribes and genera of Carabidae. The presence of such setiferous punctures on odd-numbered elytral intervals (except the first) in amphizoids suggests that this may be an ancestral (plesiotyp- ic) adephagan trait. Absence of setae from the punctures may be an apotypic trait associated with development of an aquatic lifestyle. Ab- sence of the punctures themselves (such as is seen in adults of the other hydradephagan groups) may represent a more highly evolved trait. However, a majority of carabid groups also lack some or all of these punctures; so the evolution of this character has been complex and homoplastic, no matter how the polarity of its states is interpret- ed. HasitAt DistriBuTION.— Unknown. 72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 GEOGRAPHICAL DISTRIBUTION.—This species is known only from the type-locality in south- western China (Fig. 17), in the Min River drain- age, an upper tributary of the Yangtze River. This watershed flows first south, then east more than 2,000 km to the Pacific Ocean, at 30°N latitude, and has no Himalayan drainage com- ponent. Past mislocation of the type-locality (i.e., Tibet rather than Szechwan, China) has apparently led collectors astray. Several workers, all with knowledge of the habits and habitat preferences of Nearctic amphizoids, have collected in var- ious parts of the Himalayas (e.g., India, Nepal, Tibet, Sikkim) in recent years without finding representatives of this species. This suggests that the range of Amphizoa in the Palaearctic Region may not extend west to include the main Hi- malayan ranges. Furthermore, the People’s Re- public of China has been closed to most western collectors for decades (and until very recently); and this may account for the lack of additional specimens in European or North American col- lections during this century. GEOGRAPHICAL RELATIONSHIPS WITH OTHER Species.—The known range of this species is al- lopatric with respect to ranges of all other known species of Amphizoa. Amphizoa insolens LeConte (Figures 4, 8, 14, 18, 28) Amphizoa insolens LeConte, 1853:227. Lectotype (designated by Kavanaugh 1980) male in MCZ. Type-Locality.—Sac- ramento, California. Edwards 1951:323, 1954:19. Hatch 1953:194. Kavanaugh 1980:290. Leech and Chandler 1956: 301. Dysmathes sahlbergii Mannerheim, 1853:265. Location of type- specimen unknown. Type-Locality.—Sitka, Alaska. Ed- wards 1951:323. Kavanaugh 1980:291. Synonymized by Sallé Amphizoa josephi Matthews, 1872:119. Lectotype (designated by Kavanaugh 1980) male in BMNH. Type-Locality.— Van- couver Island, British Columbia. Edwards 1951:323. Hatch 1953:194. Kavanaugh 1980:290. Synonymized by Horn 1873: WT: Notes ON NOMENCLATURE AND TYPE- SPECIMENS.—The problem with location of the holotype of Dysmathes sahlbergii Mannerheim was discussed by Kavanaugh (1980). DISTINGUISHING CHARACTERISTICS. — Size var- ied (small, medium, or large), SBL male = 10.9— 13.6 mm, female 11.1-15.0 mm; body form moderately broad (Fig. 4a); body black, with an- tennae, maxillary and labial palpi, and tarsi black or rufopiceous; head finely and densely punctate; pronotum medially with coarse, sparse punc- tures with areas between punctures flat, laterally with punctures coarser, denser, confluent, sur- face markedly gnarled; elytra finely and densely punctate, markedly rugose at base and in lateral one-half; pronotum (Fig. 8) as broad (or broader) at middle as (than) at base, with lateral margins arcuate at middle, moderately or markedly sin- uate anterior to basal angles, markedly crenulate, median longitudinal impression deeply im- pressed; prosternal intercoxal process (Fig. 12) slightly elongate, spatulate; posterolateral angle of proepisternum and posteromedial angle of proepipleuron abut evenly to form smooth pro- thoracic margin (see Edwards 1951:321, “Plate 4’’); elytral silhouette (Fig. 4a) subovoid, slightly narrowed basally, less narrowed subapically, ely- tra (Fig. 4b) evenly convex, without carinae; ely- tral striae complete but faintly impressed (diffhi- cult to define laterally because of macrosculpture) and finely punctate; front tibia with posterodor- sal groove restricted to apical one-half or two- thirds, with fringe setae in groove restricted to apical one-third or one-half; male median lobe (Fig. 14) with shaft slightly thickened at middle, evenly arcuate ventrally, apex slightly deflected ventrally and extended apicodorsally, left par- amere narrow basally, with vestiture restricted to apical one-third; female coxostylus (““coxite” of Edwards 1951:321, see his “Plate 4”) with stylar region short, with vestiture of only a few scattered, minute setae; specimen from western North America (Fig. 18). Among specimens studied, I observed greater variation in pronotal shape than that reported by Edwards (1951). Although many adults of A. insolens have pronota broadest at the middle, in most they are equally broad at middle and base. There is also notable variation among individ- uals with respect to tibial grooves and associated fringe setae (“hairs” of Edwards 1951). These structures are discussed more fully below in my treatment of 4. lecontei. In A. insolens adults, the posterodorsal groove on the front tibiae is varied in length, either restricted to the apical one-half of the tibia or extended basally to oc- cupy the apical two-thirds of the tibia. Fringe setae in this groove are restricted to the apical one-third of the tibia in most individuals, but several adults were seen with fringe setae also at the middle of the tibia or even on the apical part of the proximal one-half. As noted by Edwards KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES (1951:324), A. insolens adults have the least completely developed complement of fringe se- tae among extant amphizoids (see further dis- cussion in section on Zoogeography and Evolu- tion). Hasitat DistRIBUTION.— Members of this species are most often found at the edges of cold, swift-flowing streams, under rocks or in coarse gravel at shoreline, clinging to exposed roots be- neath undercut banks, or in floating debris that has collected in backwater eddies. Adults are often found in greatest numbers at the bases of water- falls, which represent a first stretch of quiet water after a steep drop downstream. The occasional occurrence of these beetles in ponds or lakes, where they are almost always found near the in- lets of torrential streams, probably results from their being washed downstream and does not represent permanent residence in such standing bodies of water. GEOGRAPHICAL DISTRIBUTION.—The known range of this species (Fig. 18) extends from south- ern Yukon Territory and southeastern Alaska south to the San Bernardino, San Gabriel, and San Jacinto mountains of southern California, and from the Pacific Coast, including the Queen Charlotte Islands and Vancouver Island, east across the Columbia Plateau and Great Basin to western Alberta, central Montana, western Wy- oming, central Idaho, and eastern Nevada. I have examined 398 males and 360 females from the following localities: CANADA Alberta: Banff National Park, Banff [May] (1; CAS). British Columbia: Yoho National Park, Kicking Horse River (20.9 km W of Field) [June] (2; USNM); other localities, Ainsworth Hot Springs [July] (1; USNM), Fernie [Aug.] (1; CAS), Haines High- way (km 92.1) [July] (1; UASM), Inverness [July—Aug.] (2; USNM), Kaslo [June] (1; USNM), Kay Falls [July] (1; CAS), Kuskanook (Kootenay Lake [530 m]) [Oct.] (1; RERo), Kwi- nitza (Telegraph Point) [June] (2; UASM), Nicomen Ridge [July] (1; CAS), North Bend [June] (1; USNM), Prince Rupert (north slope of Mount Hayes near base [1 20 m]) [July] (1; CAS), Revelstoke (25.7 km W) [July] (1; CAS), Seltat Creek (Haines Highway km 78.5) [June] (1; UASM), Skagit (40 km E of Hope) [July] (1; RERo), Stanley [June] (1; CAS), Stawamus River (2 km S of Squamish on Highway 99) [July] (1; CAS), Wynndel [Aug.] (10; CAS, OSUO); Queen Charlotte Islands, Graham Island (Ghost Creek drainage 7.3 km NW of Rennell Sound Road [240 m], Juskatla area, Nebria Peak at Lower Nebria Lake [620 m)}) [July] (11; CAS), Moresby Island (3 km NE of Jedway [6-50 m], Mount Moresby at High Goose Lake [640 m]) [July-Aug.] (2; CAS); Vancouver Island, Tyee (4.9 km NW) {June] (2; UASM). Yukon Territory: Upper Frances River (at Route 10) [June] (3; DMad). 73 UNITED STATES OF AMERICA Alaska: Juneau [June] (1; CAS), Lituya Bay (9.7 km N [240- 590 mJ) [Aug.] (21; CAS). California: El Dorado County, Pino Grande ({1,370 m]) [July] (3; UCD), Pollock Pines [July] (1; UCD), Riverton [July] (2; CAS, UCD), Whitehall area [June] (1; CAS); Fresno County, Barton Flat ({1,580 m]) [May] (1; UCB), Huckleberry Meadow [May] (1; CAS); Inyo County, Lone Pine (12.9 km N) [June] (1; CAS); Kings Canyon National Park, Bubbs Creek Canyon ({3,200 m]) [July] (1; CAS); Lake County, Bartlett Springs [June] (1; CAS); Lassen County, Susan River (12.9 km N of Susanville on Highway 36) [Aug.] (1; CAS), Los Angeles County, Coldwater Canyon [Aug.] (2; CAS), Little Jimmy Creek [June] (1; GCha), Los Angeles area (1; USNM), San Antonio Creek [June] (6; GLPe); Madera County, Boggy Meadows ([1,830 m]) [July] (10; CAS, NSDA, SJSU); Mariposa County, Sweetwater Creek ({1,220 m]) [July] (2; CAS); Mono County, Twin Lakes [Aug.] (4; USNM); Nevada County [Aug.] (1; CAS), Sagehen Creek (near Hobart Mills) [Aug.] (9; UCD), Truckee [Aug.] (1; CAS); Placer County, Emigrant Gap ({1,620 m]) [June] (1; UCD), Shirttail Creek (below Yellow Pine Reservoir) [May] (1; BYUM); Plumas County, North Fork Feather River ({910 m]) [Apr.] (1; CAS); Riverside Coun- ty, San Jacinto Mountains (Idlewild) [July] (6; CAS); San Ber- nardino County, Camp Baldy [July, Sep.] (9; CAS, UCD), Cie- nega Seco (6.4 km E of Barton Flats on Highway 38) [Aug.] (1; GCha), Mill Creek (0.16 km E of Forest Falls [1,650 m]) {May] (4; CAS), San Gorgonio Mountain ({2,130 m]) [Sep.] (22; CAS); San Mateo County, Tunitas Creek [Aug.] (1; UASM); Santa Clara County, Corte Madera Creek [Apr.] (1; CAS), Los Gatos [June] (1; CAS), San Francisquito Creek (Stanford Uni- versity Campus) [July] (1; USNM), San Jose [Sep.] (1; CAS), Uvas Creek (Sveadal, Uvas County Park, Uvas Meadows) (Mar.—May, July—Aug.] (11; LGBe, SJSU); Santa Cruz County, Boulder Creek [Apr.] (1; SJSU), Castle Rock State Park [May] (1; LGBe); Sequoia National Park ({610-910 m]) [May-June] (7; CAS, UCD), Ash Mountain (Kaweah Powerhouse #3) [June-— July, Sep.] (22; UCB, UCD), Cahoon Meadow (({2,290 m]) {Aug.] (1; CAS), Giant Forest [Aug.—Sep.] (1; CAS), Paradise Valley ({910-2,130 m]) [May, July] (2; CAS), Potwisha ([610- 1,520 m]) [May, July] (5; CAS, UCD, USNM), Wolverton ({2,130-2,740 m]) [June] (1; CAS); Shasta County [July] (1; USNM), Castle Crags [July] (4; CAS), Lost Creek (at Twin Bridges Road [1,450 m]) [Aug.] (3; CAS), McArthur-Burney Falls State Park ([910 m]) [June—-Sep.] (36; CAS, OSUO, SJSU), Old Station ([1,220 m]) [Sep.] (2; CAS), Viola ([1,370 m] and 6.4 km W) [June] (3; CAS, NSDA); Sierra County, Sierraville (8 km S [1,830 m]) [Aug.] (1; CAS); Siskiyou County [July] (7; CAS), Big Flat Campground [Aug.] (6; CAS), Cement Creek (S of Callahan [1,220 m]) [Aug.] (1; CAS), East Fork of South Fork Salmon River (headwaters at Cecilville/Callahan Road [1,830 m]) [July] (1; CAS), McCloud [June] (4; CAS, USNM), Mount Shasta (Panther Creek [2,440 m]) [July] (1; CAS), Shasta Springs (Shasta Retreat [730 m]) [July] (14; CAS), Taylor Lake Road ({1,750 m]) [Aug.] (1; CAS), Yreka area (1; USNM); Tehama County, Soap Creek ({610 m]) [July] (1; CAS); Trinity County, Boulder Creek (at Goldfield Campground [1,070 m]) [July] (5; CAS), Doe Gulch (1.6 km W of Altoona Mine on Ramshorn/Castella Road [1,230 m]) [Aug.] (1; CAS), Emerald Lake ({1,680 m]) [Aug.] (1; CAS), Rarick Gulch Creek (8 km S of Dedrick [640 m]) [Aug.] (1; CAS), Swift Creek ({1,520 m]) {May] (12; PUCA); Tulare County, California Hot Springs [July] (7; LGBe), Franklin Creek ([2,500-2,990 m]) [July] (1; CAS), Kaweah [June—Aug.] (10; CAS), Mineral King [Aug.] (2; CAS), South Fork Kaweah River [July] (2; USNM); Tuolumne 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 County, Herring Creek ({1,980 m]) [Aug.] (1; CAS); Yosemite National Park, Yosemite Valley (Lower Merced River) [June] (3; CAS); county unknown, Alpine Lake (1; CAS). Idaho: Blaine County, Petit Lake Creek (4.8 km WSW of Highway 93 on Twin Lakes Trail [2,130-2,440 m]) [Aug.] (1; CAS); Boise County, Rocky Bar ({1,830 m]) [June] (1; CAS); Elmore Coun- ty, South Fork Boise River (4.8 km N of Pine at Dog Creek {1.460 m]) [Aug.] (1; CAS); Nez Perce County, Waha [June] (1; CAS); Shoshone County, Wardner [July] (6, CAS, OSUO); county unknown, Twin Creek Forest Camp ([1,520 m]) [July] (2; OSUO). Montana: Cascade County, Belt Creek (27.4 km S of Monarch on Highway 89 [2,100 m]) [July] (4; CAS); Glacier National Park [July, Sep.] (5; CAS, SJSU, UCD), Howe Creek {July] (4; SJSU), St. Mary Lake [July] (3; CAS), Swiftcurrent Creek (at Many Glacier Ranger Station) [Aug.] (3; SJSU), Two Medicine Lake [July] (7; CAS); Sweetgrass County, Big Timber Creek (at Half Moon Campground [2,230-2,290 m]) [July] (4; CAS). Nevada: Elko County, Lamoille Creek (near headwaters) {June] (1; BYUM), Thomas Creek (12.9 km SE of Lamoille at Thomas Creek Campground [2,320-2,380 m]) [Aug.] (1; CAS); Lander County, Hilltop [Aug.] (1; NSDA), Skull Creek [Sep.] (3; NSDA); Washoe County, Galena Creek (17.7 km W of Highway 395 on Highway 27 [2,290 m]) [July] (3; CAS), Third Creek (at Highway 28 [2,210 m]) [July] (22; CAS), Whites Creek (near Reno) [Oct.] (1; NSDA); White Pine County, Taft Creek ([{2,130-2,440 m]) [July] (3; CAS). Oregon: Baker Coun- ty, Pine Creek (16.1 km W of Baker [1,220 m]) [June—July, Sep.] (17; CAS, OSUO, USNM); Benton County, Marys Peak (Parker Creek at Road 1245 and Road 1296) [June] (9; GLPe, OSUO, SJSU), Yew Creek (14.5 km E of Alsea) [May] (1; OSUO), Clackamas County, Brightwood [May] (1; OSUO); Deschutes County, Indian Ford Creek (8 and 9.7 km NW of Sisters) [May—July, Sep.—Oct.] (81; GLPe, OSUO, SJSU), Squaw Creek (Highway 20 at Sisters [980 m]) [Aug.] (1; CAS); Hood River County, Mount Hood (Sand Creek) [July] (4; CAS); Jef- ferson County, Camp Sherman [Aug.] (2; UCD), Metolius (9; OSUO), Metolius River [June] (2; OSUO);, Klamath County, Deming Creek (17.7 km NE of Bly) [June] (4; GLPa); Lane County, McKenzie River (8.4 km W of McKenzie Bridge [350 m]) [May] (2; CAS), South Fork McKenzie River [Sep.] (1; OSUO); Linn County, H. J. Andrews Forest (Mack Creek at Road 1553 [810 m]) [May] (1; CAS), North Santiam River (near Idanha) [May] (1; OSUO); Multnomah County, Bonne- ville [July] (1; BYUM), Horsetail Falls ([120 m]) [May, July] (8; CAS, GLPe, OSUO), Multnomah Falls [July] (5; CAS, OSUO); Wallowa County, Lostine River ([{1,310 m]) [Aug.] (1; OSUO), Wallowa Lake ({1,830 m]) [June-July] (9; CAS, OSUO, USNM), West Fork Wallowa River (at Sixmile Meadow [1,830 m]) [July] (1; CAS); Wasco County, Bear Springs (40 km W of Maupin [980 m]) [May] (1; OSUO). Washington (1; OSUO): Chelan County, Buck Creek [Aug.] (1; SJSU); Clallam County, Soleduck River [Sep.] (3; CAS, SJSU); King County, Fall City [July] (1; OSUO), Greenwater River (1; OSUO), North Bend {July] (1; CAS), Seattle [July] (1; OSUO), Snoqualmie (1; OSUO), Snoqualmie Pass [Sep.] (2; OSUO), Tokul Creek (at Tokul) [July] (4, CAS, GLPe, OSUO, UCD), Wellington [July] (4; CAS, USNM), White River (8 km W of Greenwater on High- way 410 [490 m]) [Aug.] (35; CAS); Kitsap County, Seabeck [Aug.] (1; OSUO), Kittitas County, Iron Creek Pass ({1,520 m]) [Aug.] (2; OSUO); Lewis County, Horse Creek (near Long- mire) [July] (1; CAS); Mason County [June] (2; OSUO), Pebble Ford Creek [June] (1; OSUO), Skokomish River [May] (1; OSUO); Mount Rainier National Park, Longmire [July] (2; CAS), Narada Falls ({1,370 m]) [July] (1; CAS), Paradise River ({1,490 m]) [June] (1; USNM); Olympic National Park, De- ception Creek (at Dosewallips Trail [960 m]) [July] (1; CAS), Olympic Hot Springs ({760 m]) [June-July] (9; CAS, OSUO), Pass Creek (at Dosewallips Trail [S60 m]) [July] (1; CAS), Sol Duc Hot Springs [Aug.] (2; USNM), Upper Twin Creek (at Dosewallips Trail [670 m]) [July] (2; CAS); Pierce County, Goat Creek (6.4 km E of Ashford on Highway 706 at Nisqually River [900-910 m]) [July] (4; CAS), Poch Creek (Carbon River Canyon) [Aug.] (1; OSUO); Whatcom County, Mount Baker (3.3 km E of Picture Lake on Highway 542 at Bagley Creek [670 m]) [Aug.] (1; CAS); Yakima County, Glenwood (2.7 km N [700 m]) [May] (2; CAS), Mount Adams (Bird Creek [1,370- 2,130 m]) [July] (44; CAS, OSUO, USNM), Naches River (6.9 km SE of Cliffdell on Highway 410 [740 m)) [July] (1; CAS), White Pass [June] (2; SJSU); county unknown, Mount Adams ({1,830-2,440 m]) [July] (3; CAS). Wyoming: Yellowstone Na- tional Park, Gardiner River (at Mammoth Hot Springs) [Aug.] (1; OSUO). Locality unknown: (2; CAS, USNM). GEOGRAPHICAL VARIATION.—Although con- siderable intrapopulational variation is evident for several characters, the only character in which I observed variation associated with distribution is body size. Adults from southern California, at the southern range limit for 4. insolens, are the largest specimens I have seen. The smallest adults are from coastal Alaska, at the extreme northern range limit of the species. Adults from inter- mediate areas are intermediate in size, but the pattern is not strictly clinal. For example, adults from the area around Portland, Oregon, are larg- er than those from the Mount Rainier, Wash- ington, area; and adults from interior localities (e.g., Alberta, Montana, and Wyoming) are al- most as small as Alaskan specimens and clearly smaller than specimens from west coast localities at equivalent latitudes. Hence, the pattern is one of decreasing size from south to north and from west to east, with minor exceptions to the pattern in a few areas (such as Portland). GEOGRAPHICAL RELATIONSHIPS WITH OTHER Species. — The known range of A. striata (Fig. 19) is completely within the range of A. insolens (Fig. 18). Nevertheless, the two taxa may not be mi- crosympatric. The only record of their co- occurrence (perhaps in different streams) is at North Bend, King County, Washington. The geographical ranges of 4. insolens and A. lecontei (Fig. 20) overlap broadly: from south- central and southeastern British Columbia, east to southwestern Alberta (Banff National Park) and northwestern Montana (Glacier National Park), and south to northeastern Oregon (Wal- lowa Mountains) and central Idaho (Sawtooth Mountain system). Ranges of these species ap- pear to overlap also in northwestern British Co- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES lumbia/southern Yukon Territory. Adults of both species have been found together in several lo- calities (see respective locality lists). Amphizoa striata Van Dyke (Figures 5, 9, 15, 19, 28) Amphizoa striata Van Dyke, 1927b:197. Holotype male in CAS. Type-Locality.— North Bend, King County, Washing- ton. Edwards 1951:324. Hatch 1953:194. Kavanaugh 1980: 291. Leech and Chandler 1956:301. DISTINGUISHING CHARACTERISTICS. — Size large, SBL male = 13.1-14.2 mm, female 13.2-14.9 mm; body form very broad (Fig. 5a); body dark brown or piceous, with antennae, maxillary and labial palpi, and tarsi piceous or rufopiceous; head very finely and densely punctate; pronotum coarsely, moderately densely punctate over en- tire surface, with areas between punctures flat; elytra finely and densely punctate, slightly rugose in lateral one-third; pronotum (Fig. 9) broadest at base in most individuals (as broad at middle as at base in a few individuals), with lateral mar- gins slightly or moderately arcuate at middle, not sinuate or slightly sinuate anterior to basal an- gles, slightly or moderately crenulate, median longitudinal impression present but faintly im- pressed; prosternal intercoxal process moderate- ly elongate, spatulate; posterolateral angle of proepisternum and posteromedial angle of pro- epipleuron abut evenly to form smooth protho- racic margin (see Edwards 1951:321, “Plate 4’’); elytral silhouette (Fig. 5a) very broad basally, markedly narrowed subapically, elytra (Fig. 5b) convex, except slightly concave in lateral one- half posterior to humeral area, without carinae; elytral striae complete but faintly impressed, coarsely punctate; front tibia with posterodorsal groove extended along entire length, with fringe setae in groove restricted to apical two-thirds or four-fifths of tibia; male median lobe (Fig. 15) with shaft distinctly thickened at middle, slightly bulged ventrally, apex slightly deflected ventral- ly, not extended apicodorsally, left paramere broad basally, with vestiture restricted to apical one-fourth; female coxostylus (“‘coxite” of Ed- wards 1951:321, see his “Plate 4”) with stylar region medium in length, with dense vestiture of minute setae; specimen from western North America (Fig. 19). HasitAt DistRIBUTION.— Members of this species have been found in cool (but not cold), slow-flowing streams (Edwards, pers. comm.) and 75 in roadside ditches. Their distribution in such streams is similar to that of members of A. in- solens. GEOGRAPHICAL DiIsTRIBUTION.—The known range of this species (Fig. 19) extends from south- ern Vancouver Island and the Olympic Penin- sula and Cascade Range of northern Washington, south to southwestern Oregon, and east to Yak- ima County, Washington, and Wasco County, Oregon (both east of the Cascade Range). I have examined 73 males and 63 females from the following localities: CANADA British Columbia: Vancouver Island, Duncan (Koksilah Creek) [Aug.] (5; MCZ, OSUO, USNM), Little Qualicum Falls Provincial Park (Little Qualicum River) [Aug.] (4; CAS, OSUO). Unitep STATES OF AMERICA Oregon: Benton County, Sulphur Springs (9.7 km NW of Corvallis) [May] (2; GLPe); Clackamas County, Colton [Aug.] (1; CAS); Jackson County, Little Applegate River (7.2 km S of Ruch [520 m]) [May] (1; CAS); Lincoln County, Deer Creek (12.9 km S of Toledo) [June] (1; OSUO); Wasco County, Tygh Valley [June] (1; OSUO). Washington: Clallam County, La Push [July] (1; OSUO); King County, Bothell (North Creek, Swamp Creek) [May-July] (12; CAS, GLPe, OSUO, SJSU, UCD), North Bend [July] (3; CAS), Seattle (Swamp Creek) (July, Sep.] (10; BYUM, CAS, OSUO, SJSU, UCD), Swamp Creek [May—Aug.] (71; GLPe, OSUO); Kitsap County, Brem- erton [Apr.] (1; NDSA); Kittitas County, Parke Creek (near Kittitas) [Aug.] (1; LGBe); Mason County, South Fork Sno- homish River [July] (2; OSUO); Snohomish County, Hazel (Stillaguamish Club) [May] (1; OSUO), Swamp Creek [Sep.] (5; SJSU); Yakima County, Satus Creek (near Toppenish [610 m]) [Aug.] (7, CAS, UCD, USNM); county unknown, Pack Forest [Aug.] (1; OSUO), ‘*Pisht R.” [July] (1; CAS). GEOGRAPHICAL VARIATION.— Although intra- populational variation is evident in body size, pronotal shape, and several other characters, I found no characters in which variation is asso- ciated with distribution. GEOGRAPHICAL RELATIONSHIPS WITH OTHER Species.— Refer to discussions under this head- ing for 4. insolens and A. lecontei. Amphizoa lecontei Matthews (Figures 6, 10, 12, 16, 20, 28) Amphizoa lecontei Matthews, 1872:121. Lectotype (designated by Kavanaugh 1980) male in BMNH. Type-Locality.— Van- couver Island, British Columbia [doubtful record, see com- ments below]. Edwards 1951:327, 1954:19. Hatch 1953:195, Kavanaugh 1980:290. Leech and Chandler 1956:301. Amphizoa planata Van Dyke, 1927a:98. Holotype female in CAS. Type-Locality.— Beaver Creek, Alberta. Edwards 1951: 327. Hatch 1953:195. Kavanaugh 1980:291. Synonymized by Van Dyke 1927b:197. Amphizoa carinata Edwards, 1951:326. Holotype male in CAS. 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Type-Locality.—Conejos River near Menkhaven, Conejos County, Colorado. Kavanaugh 1980:289. Leech and Chan- dler 1956:301. New Synonymy. Notes ON NOMENCLATURE AND Types.—The lectotype of A. /econtei is supposed to have been collected on Vancouver Island, British Colum- bia, as noted both in Matthews’s original de- scription and on labels affixed to the specimen. However, I have not seen any other specimens from the island nor from the adjacent coastal mainland. Because the present type-locality ap- pears to be well outside the known geographical range of A. /econtei (see Fig. 20 and text below), it is probable that the lectotype is mislabeled and that the type-locality should be emended. How- ever, I choose not to do so at this time, pending further field efforts on Vancouver Island. DISTINGUISHING CHARACTERISTICS. — Size me- dium, SBL male = 11.7-12.7 mm, female 12.2- 14.0 mm; body form moderately broad (Fig. 6a); body dark brown or piceous (specimens from Arizona almost black), with antennae, maxillary and labial palpi, and tarsi piceous or rufopiceous; head very finely and densely punctate; pronotum medially with coarse, sparse punctures, with areas between punctures flat, laterally with punctures coarser, denser, more or less confluent, surface unevenly rugose in appearance; elytra finely and densely punctate, with punctures confluent over large areas, moderately rugose in lateral one-half; pronotum (Fig. 10) broadest at base in most in- dividuals (as broad at middle as at base in a few individuals), with lateral margins slightly or moderately arcuate at middle, not or slightly sin- uate anterior to basal angles, slightly or moder- ately crenulate, median longitudinal impression faintly or deeply impressed; prosternal intercoxal process moderately elongate, spatulate or sub- lanceolate; posterolateral angle of proepisternum and posteromedial angle of proepipleuron either abut evenly to form continuous posterior pro- thoracic margin or proepipleuron is distinctly shorter than proepisternum and the two do not abut evenly, posterior prothoracic margin there- fore with distinct jog (see Edwards 1951:321, “Plate 4’); elytral silhouette (Fig. 6a) broad ba- sally, markedly narrowed subapically, elytron (Fig. 6b) with blunt but distinct carina on fifth interval, area medial to carina elevated, flat, area lateral to carina slightly concave; elytral striae complete but faintly impressed, coarsely punc- tate; front tibia with posterodorsal groove ex- tended along entire length or restricted to apical four-fifths, with fringe setae in groove restricted to apical one-half or two-thirds of tibia; male median lobe (Fig. 16) with shaft distinctly thick- ened at middle, slightly bulged ventrally, apex slightly deflected ventrally, not extended apico- dorsally, left paramere broad basally, with ves- titure restricted to apical one-fourth; female cox- ostylus (“coxite” of Edwards 1951:321, see his “Plate 4°’) with stylar region long and slender, with dense vestiture of minute setae; specimen from western North America (Fig. 20). There is considerable intrapopulational vari- ation in the development of tibial grooves and associated fringe setae among adults of all Am- phizoa species; for this reason, I have experi- enced considerable difficulty in trying to interpret tibial characters that Edwards used to distinguish A. lecontei and A. carinata adults. I have found no differences between specimens Edwards iden- tified as A. carinata and specimens of A. /econtei from various localities throughout its range in development of tibial grooves or in length or distribution of fringe setae, except such as can be attributed to intrapopulational variation. Edwards also described and illustrated differ- ences in shape of valvifers and paraprocts be- tween A. /econtei and A. carinata females. Among my own dissections of females from within the range of 4. carinata and from other localities for A. lecontei, 1 found only the /econtei form illus- trated by Edwards (1951:321, ‘Plate 4°’). I have also examined material dissected by Edwards, including the specimen that he illustrated for A. carinata. Although his drawing is a true repre- sentation of form of the valvifers and paraprocts of the latter specimen, other specimens from the same series differ from it in form and are, in fact, similar to other females of A. /econtei. It appears, therefore, that 4. /econtei and A. carinata females are similar in form of valvifers and paraprocts, and that the A. carinata specimen illustrated by Edwards is atypical in this regard. Hasitat DistriBuTION.— Members of this species are found in cool or cold, slow- or fast- flowing streams, in the same microhabitats as those described above for 4. insolens members. However, they are more common in stretches of slow-moving water and streams that drop less steeply than are members of the latter species. GEOGRAPHICAL DISTRIBUTION.—The known range of this species (Fig. 20) extends from south- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES ern Yukon Territory south along the Rocky Mountain system to the Chuska Mountains of northeastern Arizona and Sangre de Cristo Range of northern New Mexico, and from the Wallowa Mountains of northeastern Oregon and Indepen- dence Mountains of northeastern Nevada east to the Bighorn Mountains of northcentral Wyo- ming and Front Range of central Colorado. I have examined 190 males and 202 females from the following localities: CANADA Alberta: Banff National Park, Banff (and at Cascade River) [May, July—Aug.] (5; CAS, USNM); other localities, Beaver Creek [May] (4; CAS, USNM), Blairmore [Aug.] (1; CAS), Edmonton (1; OSUO), Happy Valley [Aug.] (2; CAS, USNM), Lundbreck [Aug.] (1; CAS), Mill Creek (72.4 km W of Fort Macleod) [Aug.] (2; USNM), Whitecourt (21 km SE on High- way 43) [May-June] (2; RERo). British Columbia: Creston (Goat River) [July—Aug.] (16; CAS, GLPe, OSUO), Fernie (Liz- ard Creek) [July] (1; CAS), Golden [Apr.] (1; CAS), Midday Valley (near Merritt) [July—Aug.] (4; CAS), Stanley [June] (1; CAS), Vernon (1; USNM). Yukon Territory: Haunka Creek (Highway 8 N of Atlin, British Columbia) [July] (1; UASM). Unitep STATES OF AMERICA Arizona: Apache County, Lukachukai Creek (8 km NE of Lukachukai at Wagon Wheel Campground [2,250-2,260 m]) [May, July—Aug.] (21; BMNH, CAS, MCZ, UASM, USNM). Colorado (2; MCZ): Archuleta County, Pagosa Springs area ({2,440-2,740 m]) [Aug.] (2; MCZ), Upper San Juan Valley ({2,130-3,200 m]) [Aug.] (6; MCZ, USNM); Boulder County, Coal Creek (3.2 km E of Wondervu) [May] (1; CAS), Lefthand Creek (9.7 km WSW of Highway 36 [2,010 m]) [Aug.] (12; CAS); Conejos County, Menkhaven (Conejos River) [June] (2; CAS); Jackson County, Cameron Pass ({2,740—2,930 m]) [Aug.] (5; CAS, SJSU), Gould (Michigan River near Cameron Pass) {Aug.] (2; BYUM); Larimer County, Virginia Dale [June] (1; USNM); Pueblo County, Beulah [Aug.] (4; MCZ); San Miguel County, South Fork San Miguel River ([2,590 m]) [July] (12; MCZ). Idaho: Adams County, New Meadows [June] (2; CAS, OSUO); Bear Lake County, Bloomington Creek (11.1 km SW of Bloomington [2,130 m]) [Aug.] (2; CAS); Camas County, Carrie Creek (57.9 km ESE of Ketchum [2,100 m]) [Aug.] (13; CAS), Little Snake Creek [Sep.] (1; GLPe), South Fork Boise River (22.5 km E of Featherville at Skeleton Creek [1,550 m]) [Aug.] (2; CAS); Cassia County, Goose Creek [July] (2; GLPe), Magic Mountain (Rock Creek at Ranger Station [1,890 m]) (July] (5; OSUO); Clark County, Birch Creek [July] (1; GLPe); Elmore County, South Fork Boise River (4.8 km N of Pine at Dog Creek [1,460 m]) [Aug.] (3; CAS), Wood Creek (1.6 km S of Pine [1,370 m]) [Aug.] (16; CAS); Valley County, Bear Valley [July] (1; GLPa). Montana: Cascade County, Dry Fork Belt Creek (at Henn Gulch [1,620 m]) [July] (9; CAS); Chouteau County [Aug.] (1; OSUO); Glacier National Park [July—Sep.] (10; CAS, SJSU), Kintla Lake [June] (1; CAS), Swiftcurrent Creek (at Many Glacier Ranger Station [1,460 m]) [June—Aug.] (32; CAS, SJSU). Nevada: Elko County, North Fork Humboldt River [Oct.] (1; BYUM). New Mexico: Taos County, Red Riv- er (6.6 km W of Red River [2,580 m]) [June] (1; CAS). Oregon: Baker County, Cornucopia (14.5 km NW of Halfway) [July] (1; GLPe), Richland area ([{1,220 m]) [June] (1; CAS); Grant 77 County, Clear Creek (3.2 km W of Granite) [Aug.] (1; GLPe); Wallowa County, Bear Creek (at Boundary Camp) [Sep.] (1; USNM), Lostine River (16.1 km S of Lostine [1,310 m]) [July— Aug.] (7; CAS, OSUO, UCD, USNM). Utah: Box Elder County Clear Creek (at Clear Creek Campground) [Mar.] (1; BYUM), George Creek Campground [Apr.] (1; BYUM); Emery County, Huntington Creek (at Stuart Ranger Station) [July] (1; BYUM); Garfield County, Steep Creek [Aug.] (1; BY UM); Kane County, East Fork Virgin River (7.9 km NE of Glendale [1,860 m]) [June] (2; CAS); Salt Lake County, City Creek [June-July] (14; USNM); Piute County, Beaver Creek (below national forest boundary) [May] (1; BYUM); Sevier County, Mount Marvine (0.2 km N of Johnson Valley Reservoir at Sevenmile Creek [2,590 m]) [Aug.] (15; CAS); Summit County, Tryol Lake (1; BYUM); Utah County, Hobble Creek ({1,830 m]) [July-Aug.] (29; BYUM, CAS, NSDA, SJSU), Provo ([1,490 m]) (1; CAS); Wasatch County, Little South Fork Provo River [July] (1; BYUM), Lost Lake Campground ([2,990 m]) [Aug.] (1; CAS), Upper Provo River (5.5 km E of Hailstone Junction on High- way 89A/150 [1,890 m]) [Aug.] (35; CAS), West Fork Du- schesne River [Aug.] (1; BY UM); Weber County, Ogden [July] (2; USNM), Weber River (Highway 30 at Mountain Green [1,510 m]) [Aug.] (3; CAS); county unknown, Uinta Moun- tains [June] (2; BYUM), “Wasatch” [June] (1; USNM). Wash- ington: Pend Oreille County, Sullivan Lake [Aug.] (2; CAS, OSUO); Stevens County, Crystal Falls [Aug.] (1; CAS). Wy- oming: Big Horn County, Granite Creek (12.9 km SW of Gran- ite Pass on Highway 14 [2,380 m]) [July] (2; CAS); Converse County, LaPrele Creek (61.2 km SW of Douglas on Highway 91 at Camel Creek Campground [2,530 m]) [July] (7; CAS); Grand Teton National Park, Colter Bay [Aug.] (1; SJSU), Delta Lake ({2,730 m]) [July] (1; SJSU); Johnson County, South Fork Clear Creek (25.7 km W of Buffalo on Highway 16 [2,350 m]) [July] (7; CAS), Tie Hack Camp [Aug.] (2; SJSU); Sheridan County, Little Tongue River (20.9 km WSW of Dayton on Highway 14 [2,380 m]) [July] (5; CAS); Sublette County, Ho- back River (3.2 km NW of Bondurant [2,100 m]) [Aug.] (28; CAS); Teton County, Jackson (1; USNM); Washakie County, Tensleep Creek (17.7 km NE of Tensleep on Highway 16 [1,890 m]) [July] (1; CAS); Yellowstone National Park, Grand Canyon of the Yellowstone (above Tower Falls) [Aug.] (1; MCZ), Indian River Campground [Aug.] (1; USNM), Spirea Creek [Aug.] (2; SJSU). GEOGRAPHICAL VARIATION.—In his original description of A. carinata, Edwards (1951:327) suggested that this form might represent “merely a geographical subspecies” of 4. /econtei, but added that “it seems probable that no intergra- dation occurs between these populations.” How- ever, subsequent collections from geographically intermediate areas demonstrate intergradation, and the incongruence found among geographical variation patterns of different characters has led me to treat 4. /econtei and A. carinata as con- specific. Nonetheless, the pattern of variation in A. lecontei merits description. Mature (i.e., non-teneral) adult specimens from northeastern Arizona are black whereas mature specimens from other parts of the species range are piceous or dark brown. 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Characters of pronotal shape, including shape of lateral margins, of apical and basal angles, and relative width at base versus at middle, are all highly varied among adults of 4. /econtei. All character states cited by Edwards as unique for A. carinata adults fall within the range of vari- ation seen among 4. /econtei adults from other geographical areas. Edwards described the me- dian longitudinal impression as deep in A. car- inata adults but shallow and indistinct in A. lecontei adults. Specimens with pronotal char- acteristics of the carinata form predominate in the region from southcentral Wyoming, south through Colorado and northern New Mexico, east through northeastern Arizona, and north through south and central Utah. Adults with the typical lecontei form predominate in all other areas. Specimens with prominent excavations of the prosternum anterior to the front coxal cavities, described by Edwards as a feature unique to A. carinata adults, are found in localities through- out the range of A. /econtei, although always in lower numbers than specimens from which these excavations are lacking. Furthermore, not all specimens exhibiting other features characteris- tic of A. carinata have these excavations (e.g., most specimens from Arizona). Similarly, the relationship between the proepisternum and proepipleuron described and illustrated by Ed- wards (1951:321, “Plate 4’’) does not hold up as a distinguishing feature of 4. carinata adults. Samples from localities in southcentral Wyo- ming, northern Colorado, northern New Mexico, northeastern Arizona, and southern and east- central Utah include specimens exhibiting both states of this character, as well as intermediates between these extremes. In most adults from northeastern Arizona, northern New Mexico, and Colorado, the pro- sternal intercoxal process is slender, elongate, and sublanceolate, whereas it is slightly broader, shorter, and spatulate in adults from other areas. Several of the characters noted above are use- ful for describing the carinata form. Its geograph- ical range is centered at the southern extreme of the range of A. /econtei, in northeastern Arizona, and extends northwestward (through Utah) and northeastward (through New Mexico, Colorado, and southcentral Wyoming). In successively more northern populations within this range, the car- inata form is represented by a lower percentage of individuals. However, adults demonstrating one or more A. carinata traits are found in low numbers throughout the range of 4. /econtei; adults that are intermediate between the A. car- inata and typical /econtei forms (for one or more characters) are abundant in northern parts of the range of the 4. carinata form and present in low numbers throughout that range. Given this pat- tern, there appears to be insufficient reason for retaining the name 4. carinata even at subspe- cific rank. GEOGRAPHICAL RELATIONSHIPS WITH OTHER Species. — The geographical ranges of 4. lecontei and 4. insolens overlap extensively over a broad north/south area (see above under this heading for A. insolens),; and adults have been collected together in several localities (e.g., at Swiftcurrent Creek, Glacier National Park, Montana; see also respective locality lists). Based on material I have examined, the ranges of A. lecontei and A. striata are allopatric. If, however, A. /econtei is represented on Vancou- ver Island, the original type locality for the species, then these two species are at least macrosym- patric in that area. Phylogeny Prerequisite to understanding the evolutionary and distributional histories of the species of Am- phizoa 1s formulation of a hypothesis of phylo- genetic relationships among them. Cladistic analysis is the best available technique for elu- cidation of these relationships (Hennig 1966; Ka- vanaugh 1972, 1978a). Briefly, the analytical procedure is as follows. (1) For each character, the direction of its evolution (i.e., the so-called “polarity” of the transformation of its different character states) is determined, from most prim- itive (plesiotypic) to most derived (apotypic) state or states. (2) Taxa are then grouped together, solely on the basis of shared derived (synapo- typic) character states, into successively more in- clusive groups. (3) Because synapotypy is ac- cepted as evidence for common ancestry, and because degree of phylogenetic (cladistic) rela- tionship 1s equivalent to relative recency of com- mon ancestry, the hypothetical branching pat- tern of phylogenetic relationships inferred is simply the grouping sequence read in reversed order (i.e., from most to least inclusive). The crucial step in cladistic analysis is deter- mination of the polarity of transformations of character states for each character. Several cri- teria have been proposed and/or used (Ball 1975; KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES 79 a a 3 4 Cs b ee gs» 4 cceeenes (UN OR Q NZ b Ficures 3-6. Body form (a = dorsal aspect, right elytron omitted; b = cross-sectional dorsal silhouette of left elytron at point one-third of elytral length from base); scale line = 1.0 mm. Figure 3. Amphizoa davidi Lucas (Pao-hsing, China). Figure 4. Amphizoa insolens LeConte (Indian Ford Creek, Oregon). Figure 5. Amphizoa striata Van Dyke (Swamp Creek, Washington). Figure 6. Amphizoa lecontei Matthews (Upper Provo Canyon, Utah). Ross 1974; Ekis 1977; Kavanaugh 1978b; Crisci and Stuessy 1980; Watrous and Wheeler 1981; and references therein) to determine which states are relatively plesiotypic and which are relatively apotypic. Of these, only two have intrinsic merit. First, and most important, is the so-called “‘out- group” criterion, which can be stated as follows: for a given character with two or more states within a group, the state occurring in related groups is assumed to be the plesiomorphic [=ple- siotypic] state (Watrous and Wheeler 1981). This criterion is relatively straightforward and easy to apply, except when an appropriate out-group is difficult to recognize or when more than one character state is represented in the out-group. Recently, Maddison, Donoghue, and Maddison (1984) proposed a practical method for out-group analysis using parsimony criteria which should prove useful even when phylogenetic relation- ships among out-group components are inade- quately known. The second criterion, “character correlation” (Ekis 1977; Hennig 1966; Kavanaugh 1978)), can be stated as follows: characters for which the polarities of transformation series have been de- termined with confidence can be used to infer polarities in transformations of other characters in which evolutionary sequence is less easily in- ferred. This is the criterion of choice only when the out-group criterion cannot be applied on its 80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Ficures 7-10. Pronotum, dorsal aspect; scale line = 1.0 mm. Figure 7. Amphizoa davidi Lucas (Pao-hsing, China). Figure 8. Amphizoa insolens LeConte (Indian Ford Creek, Oregon). Figure 9. Amphizoa striata Van Dyke (Swamp Creek, Washington). Figure 10. Amphizoa lecontei Matthews (Upper Provo Canyon, Utah). own. Without formally recognizing it, Watrous and Wheeler (1981) invoked the character cor- relation criterion in order to recognize functional in-groups and functional out-groups where con- ventional (i.e., their so-called “‘taxonomic’’) in- groups and out-groups proved useless. In prac- tice, a tentative phylogenetic tree (cladogram) is constructed based on one or more characters for which character-state polarities are well estab- lished. Depending on the structure of the clado- gram derived, it may be possible to recognize a functional out-group (e.g., the most basal diver- gent lineage in the cladogram), which can be used in analysis of other characters. The distribution of states of another character, polarity of which cannot be determined by reference to the out- 12 he Ficures !1, 12. pect; scale line = 1.0 mm. Figure 11. Prosternal intercoxal process, ventral as- Amphizoa davidi Lucas (Pao-hsing, China). Figure 12. Amphizoa lecontei Matthews (Lukachukai Creek, Arizona). I Ficures 13-16. Median lobe and left paramere of male genitalia, left lateral aspect; scale line = 1.0 mm. Figure 13. Amphizoa davidi Lucas (Pao-hsing, China). Figure 14. Am- phizoa insolens LeConte (Indian Ford Creek, Oregon). Figure 15. Amphizoa striata Van Dyke (Swamp Creek, Washington). Figure 16. Amphizoa lecontei Matthews (Swiftcurrent Creek, Montana). \ KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES 81 Ficure 17. group criterion using the conventional (taxo- nomic) out-group, is then determined over the tentative cladogram. A derivative out-group cor- relation may then be possible, making use of the functional out-group recognized using other characters. The choice of a suitable out-group for Am- phizoa is not a simple one. Although amphizoids have long been considered to represent an evo- lutionary grade between the Geadephaga and the more specialized groups of Hydradephaga (LeConte 1853; Edwards 1951), their phyloge- netic relationships with other adephagan groups are not clearly understood. Among the Adephaga are both terrestrial and aquatic groups, each with generalists, specialists, and hyperspecialists (Er- win 1979) in their ranks. Structural, functional, and behavioral diversity within the suborder is great, and independent evolutionary trends, some in opposite directions, are numerous. Character state distributions of many important characters are so complex within the suborder, at least in Map of geographical distribution of Amphizoa davidi Lucas. our present understanding of them, that the log- ical out-group for amphizoids, the Adephaga- minus-Amphizoidae, is not a particularly useful group for cladistic analysis. I have, therefore, tried to limit the scope of the out-group to some subgroup of Adephaga to maximize the useful- ness of the out-group criterion as a tool in anal- ysis. Phylogenetic relationships of amphizoids To many workers (Edwards 1951, and refer- ences therein), amphizoids appear to represent a primitive grade of dytiscoid evolution. Amphi- zoids lack structural adaptations of highly spe- cialized swimmers (Hlavac 1975; Evans 1982), such as dytiscids. In fact, they are much more efficient as runners on land than as swimmers in water. Characteristics of thoracic and male and female genitalic structure and numerous other features approximate what could be expected in a suitable common dytiscoid ancestor. Among 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Ficures 18-20. Map of geographical distribution. Figure 18. Amphizoa insolens LeConte. Figure 19. Amphizoa striata Van Dyke. Figure 20. Amphizoa lecontei Matthews. extant forms, no other hydradephagan adults ap- pear to represent the early Mesozoic grade of dytiscoid evolution (Ponomarenko 1977) as well as amphizoids. But are amphizoids related only to the dytiscoids, and ifso, to which most closely? Several more particular affinities have been proposed for amphizoids. Horn (1881) anda few later authors have suggested close relationship with hygrobiids. Might the latter alone serve as a suitable out-group? Probably not. Evidence linking hygrobiids with amphizoids is minimal and likely based on symplesiotypic traits (Ham- mond 1979). Bell (1966) suggested that amphizoids, living in habitats where swimming is hazardous, may have evolved from more advanced dytiscids, with their apparent plesiotypic characteristics repre- senting secondary loss or reduction of swimming adaptations. However, characteristics of pro- thoracic (Hlavac 1975) and pterothoracic struc- ture (Ponomarenko 1977; Evans 1982, also in press) support a phylogenetically more remote (basal) relationship between amphizoids and dy- tiscids. Although a suitable out-group for am- phizoids must include dytiscids, extant forms of the latter represent a more highly specialized grade of adephagan evolution and are probably not suf- ficient as an out-group. Several lines of evidence suggest that Trachy- pachidae form a monophyletic group with the dytiscoid families, including all the hydradepha- gan groups except, perhaps, haliplids and gy- rinids (Bell 1966, 1967, 1982; Crowson 1981; Evans 1977, 1982; Hammond 1979; Forsythe 1981; Roughley 1981). This relationship is es- tablished on the basis of numerous supposed syn- apotypic features (Hammond 1979; Roughley 1981; Evans 1982) involving characters of an- tennal pubescence, locomotory function and structure (of legs, wings, and associated struc- tures), male and female genitalic structure, and female reproductive system. If trachypachids are KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES closely related to amphizoids and other dytis- coids, then they should be included in any suit- able out-group for analysis. Determination of apotypic states of at least several characters linking trachypachids with dy- tiscoids is based on the assumption, either stated or implied, that the common adephagan ancestor was terrestrial rather than aquatic in habits (Crowson 1955, 1981; Bell 1966, 1967, 1982; Evans 1977, 1982, also in press; Hammond 1979; Forsythe 1981; Roughley 1981). However, this contention is not universally accepted. Erwin (1979) suggested that the Adephaga arose from an aquatic neuropteroid ancestor similar to ex- tant amphizoids, at least in habits. Based on re- view of both Palaeozoic and Mesozoic fossil bee- tles, Ponomarenko (1977) proposed an aquatic origin of Adephaga, probably in late Permian time, from aquatic schizophoroid Archostemata. In fact, his separation of fossil specimens of Adephaga from those of Schizophoridae of that age was admittedly somewhat arbitrary (Po- nomarenko 1977). Perhaps this distinction is one of grade rather than clade. Crowson (1981) and Ponomarenko (1977) agree on both the time (Late Permian) and source group (Archostemata: Schizophoridae) for the probable origin of Adephaga. They differ, how- ever, in their views on the ancestral adephagan habitat (whether terrestrial or aquatic), which by extension, could have been inherited from either terrestrial or aquatic schizophoroid ancestors, both of which are known from Permo-Triassic time. Amphizoids are only semiaquatic in habits. Adults are able to carry on most if not all life functions (e.g., feeding, locomotion, oviposition) at least as well out of water as in it and based on my fieldwork, do so routinely in nature. At least under laboratory conditions, eggs and larvae also thrive out of water, and pupation occurs on land. These habits are reflected by structure. Adults lack special adaptations for fast swimming and are barely able to move freely underwater except by clinging to substrata. Their most effective mode of locomotion in water is passively drifting with stream currents. Here I use the term ‘‘semi- aquatic” to refer to the combination of amphib- ious habits, structure that is relatively unspe- cialized for aquatic life, and ineffective swimming capability that is characteristic of extant amphi- zoids. 83 If the ancestral adephagan was a terrestrial or- ganism, then amphizoids may represent, at least structurally, a first stage in adephagan adaptation to aquatic life. Primitive (plesiotypic) character states for Hydradephaga, including amphizoids, should be represented among their near terres- trial (i.e., geadephagan) relatives, including the Mesozoic Eodromeinae and Protorabinae (Po- nomarenko 1977), both living and extinct Trachypachinae (Bell 1966, 1982; Evans 1977, 1982; Ponomarenko 1977; Roughley 1981), and living basal-grade Carabinae, such as Notioka- suni, Nebriini, and Opisthiini. It might, there- fore, be a waste of effort to include other extant hydradephagan groups in the out-group for cla- distic analysis because their members may dem- onstrate only relatively apotypic character states associated with more advanced stages of spe- cialization to aquatic life. Alternatively, if the common adephagan an- cestor were aquatic, then plesiotypic character states should be associated with aquatic rather than terrestrial organisms. Any suitable out-group for cladistic analysis of amphizoids would have to include extinct aquatic groups, such as the Mesozoic Parahygrobiidae, Coptoclavidae, and Liadytidae (Ponomarenko 1977), as well as other living dytiscoids (Hygrobiidae, Dytiscidae, and Noteridae). If, however, the common adephagan ancestor were only semiaquatic, similar in both habits (Erwin 1979) and structure to living am- phizoids, then extant dytiscoids might again be too specialized to be useful in out-group com- parisons. Composition of a suitable out-group for anal- ysis of extant Amphizoa species depends, at least in part, on the evolutionary hypothesis proposed to account for the origin and initial radiation of Adephaga—whether from a terrestrial, aquatic, or semiaquatic common ancestor. Faced with a choice from among five conflicting hypotheses (none of which he could reject with available evidence) to explain the relationships of trachy- pachids with other Adephaga, Bell (1982) called for additional efforts to discover new evidence bearing on the question. Perhaps a similar call for additional data is most appropriate here as well. However, even a preliminary cladistic anal- ysis of Amphizoa species at this time requires selection of an out-group for comparative pur- poses; such a selection requires a choice among alternative hypotheses for adephagan origin. In 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 my view, evidence favors the origin of Adephaga from a semiaquatic common ancestor for reasons outlined below. Evidence from the fossil record Thanks to Ponomarenko’s (1977, and other papers cited therein) outstanding work on late Palaeozoic and Mesozoic beetle fossils, infor- mation about the early stages of adephagan evo- lution is now available. It is evident, for example, that a significant aquatic radiation of schizoph- orid Archostemata, presumptive ancestors of Adephaga, had occurred by Permo-Triassic time (Ponomarenko 1977). By early Mesozoic time, the adephagan radiation was already diverse. Forms that, structurally, could have given rise to all major extant adephagan groups—gy- rinoids, haliploids, dytiscoids, and caraboids— were represented in the Jurassic fauna of Asia. However, the aquatic adephagan component was clearly more diverse and more advanced (i.e., more similar to extant forms) than the terrestrial component of that time. The carabid fauna, for example, did not take on a modern aspect (i.e., one in which middle- and higher-grade carabids are evident) until mid- to late-Cretaceous time (Ponomarenko 1977). This suggests that the aquatic radiation of Adephaga preceded that of terrestrial groups. Much can also be learned about plesiotypic (primitive) versus apotypic (derived) character states for Adephaga from study of the diverse and beautifully preserved Mesozoic fossil ma- terial illustrated by Ponomarenko (1977). For example, it is clear, from review of these fossil specimens and out-group comparisons with schi- zophorid fossil material, that contribution to the lateral wall of the mesocoxal cavity by the met- episternum is plesiotypic in Adephaga. This trait was widespread among extinct (and extant) Ar- chostemata as well as the extinct eodromeine trachypachids, protorabine carabids, triaplids, and some (but not all) Mesozoic dytiscoid groups. Among extant forms it is restricted to Amphi- zoidae, some Dytiscidae, and members of genus Spanglerogyrus among Gyrinidae. Similarly, the form of hind coxae seen among extant trachy- pachids, dytiscids, amphizoids, hygrobiids, gy- rinids, and haliplids—in which the lateral coxal wing extends laterally to the elytral epipleuron, completely separating thoracic from abdominal sclerites (i.e., the “incomplete” form of Bell 1967, or “interrupted” form of Roughley 1981)—ap- pears to be plesiotypic, based on out-group com- parisons with schizophorids and Mesozoic fossil adephagans. In form and structure of hind coxae, relation- ships of mesepimera and metepisterna to me- socoxal cavities, and every other structural detail that can be observed in the fossil material, am- phizoids appear to demonstrate the character state that can be interpreted as plesiotypic in relation to a semiaquatic ancestor and divergent lines of more specialized forms. Liadytids (Ponoma- renko 1977), which probably represent a basal grade of Mesozoic dytiscoids, have hind coxae more specialized (hence, apotypic) for rapid swimming than amphizoids, and coptoclavids (Ponomarenko 1977) have metepisterna exclud- ed from lateral walls of mesocoxal cavities by anterolateral extensions of the metasternum. Amphizoids are very similar in appearance and structure to Mesozoic eodromeine trachy- pachids, except that their metacoxae are slightly larger and more closely contiguous medially than the latter. Presumably, eodromeines were ter- restrial beetles, not aquatic or semiaquatic. Perhaps the only known form more similar to eodromeines than amphizoids is Necronectulus (Ponomarenko 1977), described from a single, legless specimen of Early Jurassic age from Asia. Its metacoxae were typical of those in eodro- meines, but nothing is known of its distal leg structure. Based on body structure and form of antennae, Ponomarenko suggested that it could have been either terrestrial or aquatic in habits, but he favored the latter view. Possibly, it rep- resents the first stage of adaptation to purely ter- restrial life among Adephaga, although the ear- liest known eodromeines predate the only known occurrence of Necronectulus in the fossil record. In summary, I suggest that a review of Me- sozoic fossil material provides two insights. First, character states demonstrated by extant amphi- zoid adults can, in almost every instance, be in- terpreted as plesiotypic in relation to respective character states in known Mesozoic and extant aquatic Adephaga, as well as extant trachy- pachids and carabids. Second, there is little with which to distinguish amphizoids and eodro- meine trachypachids, except their habitats. If this similarity is based on synapotypic features, then adephagan relationships could be as illustrated in Figure 21a or 21b. If it is based on symple- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES TRACHYPACHIDAE HYDRADEPHAGA 21 TRACHYPACHIDAE 85 CARABIDAE HYDRADEPHAGA CARABIDAE TRACHYPACHIDAE HYDRADEPHAGA CARABIDAE Ficure 21. carabids. siotypic features, then adephagan groups could be related as in Figure 21c. Evidence for relationship between amphizoids and trachypachids As noted above, several workers (Bell 1966, 1982; Evans 1977, 1982; Hammond 1979; and Roughley 1981) have provided evidence in sup- port of close relationship between trachypachids and the dytiscoids, including amphizoids, hy- grobiids, noterids, and dytiscids. All of these au- thors presumed a terrestrial origin for Adephaga. Hammond (1979) listed 7 and Roughley (1981) 10 (for a total of 14 different) proposed syn- apotypies uniting these groups. Each should be considered separately, in light of all available data about extant and fossil forms. Illustrations of alternative hypotheses of phylogenetic relationships among Hydradephaga, trachypachids, and 1. Antennal pubescence. Both Hammond and Roughley considered the glabrous antennae of adult hydradephagans to be apotypic, with the plesiotypic state—antennae pubescent—associ- ated with carabids (i.e., terrestrial forms). The condition in trachypachids—glabrous, except for an apical whorl of setae on each antennomere and fine pubescence on antennomere 11 only— was considered synapotypic with the condition found among Hydradephaga. Most Coleoptera have flagellar antennomeres covered with a dense coat of short sensory setae, as do adults of most other insect orders; and presence of such pubes- cence would appear to be plesiotypic for Adeph- aga. If we assume an aquatic or subaquatic an- cestry for the suborder, however, presence of pubescence could be interpreted as apotypic in 86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 carabids; pubescence on antennae of trachy- pachids, albeit greatly restricted, could be syn- apotypic for trachypachids and carabids. 2. Open procoxal cavities with postcoxal bridge. This combination of two characters (1.e., (a) procoxal cavities open or closed and (b) post- coxal bridge absent or present) is difficult to in- terpret. Most workers agree that open procoxal cavities represent the plesiotypic state of the first character (a). However, presence or absence of a postcoxal bridge (b) is more difficult to interpret. A bridge has been reported from trachypachids and dytiscoids and cited as a synapotypy for these groups. However, Hlavac (1975) and Hammond (1979) noted the presence of a bridge in adults of Carabus, Hiletus, and the nebriine genus Leis- tus, whereas no bridge is evident in members of related carabid groups, including other nebriine genera (i.e., Nebria and Pelophila). Presence or absence of a postcoxal bridge does not appear to be a reliable character for demonstrating phy- logenetic relationships among Adephaga. 3. Prosternal process. Roughley (1981) pro- posed a similarity (synapotypy) among trachy- pachids and Hydradephaga in length and shape of the prosternal intercoxal process. I disagree with this contention. The process in both trachy- pachids and amphizoids (Fig. 11, 12) is very similar to that in Nebriini, Notiokasiini, and oth- er basal-grade carabids in size and shape and unlike more specialized aquatic adephagans such as dytiscids and hygrobiids. I regard this char- acter as symplesiotypic in trachypachids, am- phizoids, and carabids, apotypic in higher dytis- coids. 4. Prosternal-metasternal contact. Roughley (1981) suggested that contact between the pro- sternal intercoxal process and the anteriormost portion of the metasternum was possible in trachypachids as in most dytiscoids and, further, that this represented a synapotypic feature. As with character 3 (prosternal process) above I dis- agree with this interpretation. Contact between prosternum and metasternum is no greater in either trachypachids or amphizoids than in ne- briines or other basal-grade carabids. The con- dition in amphizoids, trachypachids, and basal- grade carabids is surely symplesiotypic. 5. Coadaptation of posterior border of prono- tum and anteriorly truncate elytra. Hammond (1979) proposed that coadaptation of the pos- terior pronotal margin and elytral base among trachypachids and hydradephagans represents a synapotypic feature. It is true that pronota and elytra are very closely juxtaposed in trachy- pachids and most dytiscoids, but no more so than in a number of carabid groups (e.g., omophron- ines, amarines, and some migadopines, bembi- diines, pterostichines, and harpalines). Close, relatively inflexible association of prothoraces and pterothoraces and a continuous, evenly arcuate, uninterrupted lateral silhouette (also including the head in many instances) is broadly charac- teristic of aquatic beetles. Assuming an aquatic origin for Adephaga, this form may represent the plesiotypic condition. Apparent support for this interpretation is provided by Ponomarenko’s (1977) numerous illustrations of Mesozoic Adephaga. Among beetles illustrated, including eodromeine trachypachids and protorabine ca- rabids, a form typical of extant trachypachids predominates. This evidence suggests that early carabids were more similar in form to extant trachypachids than to a majority of extant cara- bids. Perhaps the relatively narrow-waisted, flexibly-joined carabids are apotypic rather than plesiotypic in this regard, with members of groups such as omophronines and amarines having ac- quired a trachypachid-like form secondarily, as an adaptation to particular, specialized biotopes. 6. Metacoxal cavities interrupted (“incom- plete,” Bell 1966). As discussed above, meta- coxae of trachypachids, amphizoids, and other Hydradephaga (both extinct and extant) are sim- ilar in form and lateral extent to those seen in Archostemata, including presumptive schizo- phoroid ancestors of Adephaga. The main dif- ference between these adephagan metacoxae and archostematan metacoxae is that the former, unlike the latter, are countersunk into the body wall (i.e., into the basal abdominal sterna), so that they appear to divide the first (basal) visible sternum into two lateral parts. Continuity of this sternum internal to the metacoxa (i.e., dorsally) can be confirmed by dissection. I agree with Po- nomarenko (1977) that this form of metacoxae is plesiotypic in Adephaga, rather than apotypic as suggested by most recent workers (Bell 1966, 1967, 1982: Evans 1977, 1982: Hammond 1979; Forsythe 1981). Evans (1977, 1982), Forsythe (1981), Hammond (1979), and others have con- structed and/or reviewed various hypotheses to explain why trachypachids should have meta- coxae preadapted for aquatic life, and why car- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES abids should have even partially immobilized coxae for rapid running. Again, these workers assumed a terrestrial origin of Adephaga. In light of both out-group comparisons with Archoste- mata and form and structure of known Mesozoic Adephaga, it seems simplest to suggest that trachypachids have legs adapted for aquatic life because their ancestors were aquatic, and cara- bids have immobilized coxae because, like trachypachids, their ancestors lived in the water and such coxae are advantageous there. Carabid leg structure is adapted to terrestrial life, but it still reflects the constraints of ancestry. 7. Metacoxal fusion. Roughley (1981) stated that “in trachypachids and Hydradephaga the metacoxae are fused medially, the fusion being marked by a single internal intercoxal septum continuous with the metafurca and the median sternal ridge.”” My own dissections do not sub- stantiate the extent of fusion Roughley reported. In both trachypachids and amphizoids, the me- dial walls of the metacoxae are not fully fused to form a single septum, but are merely very closely approximated, slightly more so in amphizoids than in trachypachids. Roughley is correct in not- ing the close association between the metacoxae and the metafurca and median sternal ridge in adults of these groups. In the carabids examined, the metafurca is positioned far forward in rela- tion to the metacoxal base medially. It is unclear, however, which of these conditions (states) is apotypic. Presumably the state seen in trachy- pachids and Hydradephaga is an adaptation to ‘aquatic existence” (Roughley 1981:276). Again, assuming an aquatic ancestry, the state seen in carabids could be apotypic, rather than plesio- typic as Roughley and others have suggested. 8. Similarities in wing venation and folding. According to Hammond (1979), hindwings of trachypachids and dytiscoids share numerous features (e.g., wing folding pattern, position of oblongum cell in relation to apical and posterior wing margins). Adephaga are characterized by having an exceptionally strong spring mecha- nism for wing folding, which is aided, in a ma- jority of groups, by one or another kind of ab- dominal movement. Almost complete reliance on the spring mechanism alone is seen among the related, basal-grade carabid tribes Nebriini, Opisthiini, Notiophilini, Carabini, and Cicin- delini. Hammond (1979) interpreted this latter condition as (probably) plesiotypic for Adepha- 87 ga; but he noted that this hypothesis requires that increased reliance on abdominal movements, and development of special structures associated with same, occurred independently in several adeph- agan lineages. Again, without information from direct out-group comparisons with other Co- leoptera, especially Archostemata, it is difficult to recognize the most plesiotypic condition with any confidence. Although they may represent only a basal grade of carabid evolution, nebriines, opisthiines, and the other groups listed above may also form a monophyletic assemblage that diverged from other carabids at an early evolu- tionary stage, members of which are character- ized by sole reliance on the spring mechanism for wing folding. 9. Subcubital binding patch of hindwing. Both Hammond (1979) and Roughley (1981) cited presence of this binding patch, posteriorly near the apex of the hindwing, as a synapotypic fea- ture uniting trachypachids with dytiscoids. Ab- sence of such a binding patch from hindwings of carabids, haliplids, and gyrinids was seen as a plesiotypic condition. Not all dytiscoids, how- ever, have the binding patch; in all Systolosoma (Trachypachidae) adults that I examined, the patch was nonpigmented and very poorly de- fined, if it could be claimed as present at all. A subcubital binding patch is absent from hygro- biid wings and from wings of members of some bidessine, hydrovatine, and hyphydrine dytiscid genera. Hammond (1979) noted a marked as- sociation between small body size and absence of the binding patch in the dytiscid groups cited above. He suggested functional reasons why the subcubital binding patch might not be necessary in small beetles and proposed that its absence represented a secondary loss in the above dytis- cid groups. Size considerations do not, however, account for the absence of the patch from hy- grobiid hindwings nor its reduction or absence from Systolosoma adults. Although I see no rea- son to doubt that presence of the subcubital bind- ing patch is an apotypic feature in Adephaga, I suggest that its absence from hygrobiid and car- abid hindwings may also represent secondary losses. If this is correct, presence of the binding patch may, in fact, be synapotypic for Adephaga, with its secondary loss having evolved indepen- dently in some or all members of the dytiscoid, trachypachid, carabid, and haliplid lineages. 10. Male genitalia with long, apically nar- 88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 rowed parameres. Parameres of trachypachid males and of at least some dytiscoid group males are very similar in length, shape, and vestiture. Hammond (1979) suggested that the long, api- cally narrowed form seen in males of these groups represented a synapotypic feature. However, males of certain carabid groups, including car- abines, cychrines, pamborines, and cicindelines also have parameres resembling those of trachy- pachids in form. It seems simpler to suggest that this condition represents a plesiotypic rather than apotypic condition, with the great diversity of forms seen among extant carabids having evolved through several independent, apotypic trends di- verging from the basic form. 11. Size and armature of internal sac. Rough- ley (1981) and other workers have assumed that presence of a well-developed internal sac found in the median lobe of the aedeagus, such as in carabid males, represents the plesiotypic condi- tion among Adephaga; he further suggested that presence of an armature of setae and spines, on or in the sac, 1s also plesiotypic. Without knowl- edge of these characteristics in proposed schi- zophoroid common ancestors of Adephaga, nor even in extant archostematan males (such as in Omma species), it is difficult to interpret differ- ences in size and development of the internal sac among extant Adephaga in a cladistic sense. As Roughley suggested, it is also possible that the small, slightly developed internal sac of trachy- pachids and dytiscoids represents the plesiotypic adephagan condition. Male gyrinids, which ap- pear to be only distantly related to other Adeph- aga, based on many other characters (Evans 1982; Ponomarenko 1977), also have a slightly devel- oped internal sac. This suggests that the large, well-developed internal sac of carabids repre- sents an apotypic, rather than plesiotypic, con- dition. Some basal-grade carabid males (e.g., ne- briines, notiokasiines, and opisthiines) lack ev- ident armature on the internal sac. Although as- sociated spines and/or setae are found in males of some basal-grade carabids and are widespread among those of middle- and high-grade carabid groups, I see no reason to suggest that their oc- currence represents a plesiotypic condition among Adephaga, and I do not consider their absence to be synapotypic for trachypachids and dytis- coids. 12. Dilator muscle of vagina. The presence of this muscle in a majority of dytiscoids examined (Burmeister 1976) led Roughley (1981) to suggest that its occurrence represents a synapotypic fea- ture among dytiscoids (including amphizoids) and trachypachids. Its absence from carabid fe- males was considered plesiotypic. The source of Roughley’s data for trachypachids and amphi- zoids (Roughley 1981, table 1) is unclear; but I assume that these data are from his own dissec- tions because Burmeister (1976) did not present data for these groups (see his table 1, p. 216). Assuming that Roughley is correct, and this mus- cle is present in trachypachids and amphizoids as well as in haliplids, gyrinids, hygrobiids, and most dytiscids, but not in carabids (Burmeister 1976), it would seem simpler to suggest that its presence is plesiotypic, and its absence (in car- abids and a few dytiscids) apotypic among Adephaga. As with the previous character, it will be useful to examine extant archostematans as a possible out-group test of alternative hypotheses. 13. Giardina bodies. Roughley (1981) suggest- ed that the nature of so-called “Giardina bod- ies,” which contain extrachromosomal DNA and appear in oogonia at the preoocyte stage of 0o- genesis, might represent a synapotypic feature for dytiscoids and trachypachids. He noted that these bodies ‘“‘appear to be of a different type in Dy- tiscoidea than in other insects.” They have been found in female representatives of Gyrinidae, Hygrobiidae, and some Dytiscidae studied. For example, they occur in Colymbetinae, Lacco- philinae, and some (e.g., Hydaticus, Dytiscus), but not all (e.g., Eretes, Cybister), dytiscines, and are absent from the few hydroporines studied. More significantly, however, their presence (or absence) remains unknown for noterids, hali- plids, amphizoids, trachypachids, and carabids. Roughley’s primary intent was to initiate a sur- vey of the occurrence of Giardina bodies among Adephaga—to introduce a new character into adephagan systematics. Available data cannot support the hypothesis that presence of a partic- ular type of Giardina body is synapotypic for dytiscoids and trachypachids. 14. Ligula absent from labium of larva. Ham- mond (1979) cited this character state as a pos- sible synapotypic feature uniting trachypachids and dytiscoids; but he noted that a ligula is absent from larvae of various carabid groups (e.g., Bra- chinus, Gehringia, and lebiines) as well. Distri- bution of this characteristic among extant Adephaga is not yet fully known, nor have de- tailed out-group comparisons with archostema- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES Permian Triassic Jurassic WWa,21a,27a 13a Cretaceous 89 Paleogene Neogene SCHIZOPHORIDAE (ARCHOSTEMATA) (8a) A GYRINIDAE la®,3a,4a®,9a, 10a, 11c,12a®, 160, 19a®,22a® ? — |) B Spanglerogyrus (la*,3a,12a®, 160, 17a, 18a, 19a®) C COPTOCLAVIDAE PARAHYGROBIIDAE E HYGROBIIDAE F NOTERIDAE 28a & (6b, 18a) ae 8a,11b,13a,25a 220,238 12a®, 16a" (8a,11b,13a,22a) H Ja, 17a®,20a G DYTISCIDAE LIADYTIDAE I AMPHIZOIDAE 12a,16a,17a J Necronectulus 114,264 1) 4a,7a,20b (18a) 22 Na®,15a,22a,23a,24a 1a,2a,6b,8a,11a®, 13a, 18a®,19b,20a, 22a, 23a, EODROMEINAE TRACHYPACHINAE PROTORABINAE (11a®,25a,27a®) N CARABINAE 25a ' 5a,6a,18a P TRIAPLIDAE FiGure 22. QO HALIPLIDAE Reconstructed phylogeny of Adephaga, including both extinct and extant groups. Time is represented by the horizontal axis; but neither position nor gap width on the vertical axis is intended to reflect divergence considerations. Thickened portions of tree branches indicate known temporal occurrence in the fossil record. Number and letter symbols placed adjacent to solid dots refer to synapotypic features presented in Table | and discussed in the text. Symbols in parentheses refer to apotypic features found in some, but not all, members of the lineage directly below them. tan larvae been made. Therefore, significance of the co-occurrence of this feature among dytis- coids and trachypachids cannot be properly eval- uated. It may represent another symplesiotypy for Adephaga. In summary, there is little, if any, unequivocal evidence to support strict monophyly of a group including dytiscids, hygrobiids, amphizoids, and trachypachids but excluding carabids. This view is based on a re-evaluation of character polarities proposed and/or supported by Hammond (1979), Roughley (1981), and several other workers (e.g., Bell, Crowson, Evans, and Forsythe, as previ- ously cited). These workers may be correct in their interpretations. Nonetheless, I offer an al- ternative interpretation of data and relationships as perceived from my studies of nebriines and other basal-grade carabids over the past few years; I hope that these interpretations and conclusions will be rigorously tested by current and future colleagues. A hypothesis of adephagan phylogeny The hypothesis of adephagan relationships that I have used below as a basis for out-group com- parison in cladistic analysis of amphizoids is il- lustrated in Figure 22. Some relationships pro- posed are highly speculative in relation to available data, and relatively few characters have been adequately studied and applied to a cladis- tic analysis of Adephaga. Consequently, the monophyly of certain groups proposed is not substantiated, or is inadequately substantiated, 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 by synapotypic features at present. Nevertheless, I hope that others will be encouraged to challenge proposed relationships through a search for ad- ditional apotypic features that support or refute the phylogenetic hypothesis. In this regard, a comprehensive comparative study of larval structure, including what can be gleaned from review of fossil material, will undoubtedly pro- vide extremely valuable data. In both comparisons made and conclusions drawn, I have accepted adephagan family limits as presently defined. Some of the family-group taxa so delimited may not represent strictly monophyletic groups, and better understanding the phylogenetic relationships among some of these so-called “families” (e.g., dytiscids and no- terids) will require further cladistic analyses of member subgroups and relationships among them. Familial status of certain taxa known only from fossils (e.g., liadytids, parahygrobiids, and coptoclavids) is unclear, but I have accepted pro- posed familial ranking for each herein to facili- tate comparisons with extant taxa of familial rank. Based mainly on Ponomarenko’s (1977) re- view of Mesozoic fossil material, I have also tried to relate the branching sequence of the proposed cladogram to geologic time (but not specifically to events in Earth history). Among possible sources of error in establishing timing of diver- gent events in adephagan phyletic history are: (1) that fossil occurrence of a group provides only a minimum estimate of the time of its origin, and disappearance or absence of a group from the known fossil record does not rule out its existence at a particular time or place; and (2) that the geographical distribution of currently available material which represents the Mesozoic (and ear- ly Cenozoic) adephagan fauna is highly biased. Almost all useful specimens are from Asia, and some groups, such as amphizoids, hygrobiids, and haliplids, may well have evolved in other areas and much earlier than the known fossil record suggests. Of the five hypotheses of adephagan relation- ships reviewed by Bell (1982), that of Ponoma- renko (1977) is most similar to the one proposed here. Ponomarenko suggested that the common adephagan ancestor gave rise to three major, in- dependent lineages, which Bell (1982) termed the “haliplomorph,” ‘“‘dytiscomorph,” and “cara- bomorph” ancestral lineages, respectively. Ac- cording to Ponomarenko, extant haliplids may be descendants of the Triassic haliplomorph group, Triaplidae; gyrinids diverged, probably in Lower Triassic time, from the common ancestor of other dytiscoids (including amphizoids, hy- grobiids, dytiscids, and a number of extinct Me- sozoic forms); and extant carabids and trachy- pachids are descendants of a common, terrestrial carabomorph ancestor, which also evolved in, or just before, the Triassic. The only major difference between Ponoma- renko’s hypothesis and that illustrated in Figure 22 involves the relationship of haliplids to other Adephaga. I doubt that any close phylogenetic relationship exists between haliplids and tria- plids. Evidence cited by Ponomarenko as linking these two groups more likely represents conver- gence. Instead, several synapotypic features link haliplids with caraboids, and triaplids probably have no extant descendants or near relatives. Several other relationships proposed here are noteworthy. The recently discovered gyrinid ge- nus Spanglerogyrus (Folkerts 1979) appears to be a relict form less closely related to other extant gyrinids than is the Upper Triassic form, 7ri- adogyrus (Ponomarenko 1977) (see details be- low). Nothing is known about external structure of parahygrobiid adults, and placement of this group in the cladogram is problematic at present. Some evidence exists to link hygrobiids with the extinct coptoclavids rather than with other ex- tant dytiscoids. Both coptoclavids and hygro- biids appear to be more closely related to gyrinids than to dytiscids and amphizoids. The Lower Jurassic form, Necronectulus (Ponomarenko 1977), known from only a single specimen with- out legs, shares apotypic features with no known adephagan lineage. I have, therefore, indicated its derivation from an unresolved trichotomy with the dytiscomorph and carabomorph lineages. It may be related to either of these lines, but evi- dence for one or the other affinity is currently lacking. Evidence in support of relationships proposed in Figure 22 is presented in Table 1. Code letters used for taxa in the table are the same as those used in Figure 22. Coding of character states, both in Figure 22 and Table 1, is as follows: (1) each character is represented by a unique, Arabic number; (2) the plesiotypic state of each char- acter is represented by the letter 0; (3) indepen- dently derived apotypic states are represented by different letters (a, b, etc.), where states a and b KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES evolved independently from state 0; (4) sequen- tially derived apotypic states are represented by a letter (a) or a letter plus asterisk (a*), where state a evolved from state o and state a* evolved from a; and (5) apotypic states that include a combination of independently and sequentially derived conditions are represented by letters (a, b, etc.) and letters with different symbols (a*, a#, etc.), where states a and ) evolved independently from state o, and both a* and a# evolved inde- pendently from state a. Polarities of transfor- mation for 25 of the 29 characters used for cla- distic analysis were determined by means of the out-group criterion. The character correlation criterion was used to determine polarities for characters 4 (antennal pubescence), 27 (gonostyli of female ovipositor), 28 (thoracic defense glands), and 29 (pygidial defense gland cells). Implica- tions of the distributions of states of the char- acters presented in Table | in relation to the cladogram in Figure 22 are as follows. Character 1. General habitat. If a semiaquatic lifestyle, similar to that of extant amphizoids, is accepted as plesiotypic for Adephaga, then a fully aquatic lifestyle may have evolved only twice: in a lineage including all Hydradephaga except amphizoids and haliplids, and in haliplids. A more highly evolved lifestyle, one specializing in water surface activity apparently evolved twice— once in gyrinids, and again in some coptoclavids (see Ponomarenko 1977). Haliplids and amphi- zoids swim with an alternating (walking) leg mo- tion. In the former group, this trait may reflect a semiaquatic (or even terrestrial) ancestry and independent adaptation to fully aquatic life. Ad- aptation to passive drifting in streams shown by amphizoids is no doubt an apotypic feature. Character 2. Food habits/feeding. Ponoma- renko (1977) suggested that triaplids and hali- plids shared herbivorous feeding habits, but he noted also that this trait could have been plesi- otypic in triaplids. If the relationship of haliplids to caraboids proposed here is correct, then algal feeding must be apotypic in haliplids. Character 3. Compound eyes. Both gyrinids and a majority of known coptoclavids have com- pound eyes divided into dorsal and ventral por- tions. Based on other characteristics, this co- occurrence appears to be convergent in the two groups. In all extant gyrinids, except Spanglero- gyrus adults, the dorsal and ventral eye portions are moderately or broadly separated by an an- 91 terior extension of the gena. In Spanglerogyrus adults, the eye portions are broadly contiguous, with their division marked by only a thin sep- tum. This feature, in combination with others listed below, suggests a very ancient divergence of this monobasic group from the main line of gyrinid evolution. Character 4. Antennal pubescence. As noted above, this character is problematic. Other au- thors (e.g., Roughley 1981) have suggested that absence of antennal pubescence is apotypic, a trait evolved in association with the change to an aquatic lifestyle. Yet terrestrial trachypachids lack antennal pubescence (except on antenno- mere 11) and aquatic gyrinids have pubescence (but of a peculiar form and distribution). If, as I suggest here, presence of antennal pubescence is an apotypic feature where it occurs in Adephaga, then this trait may have evolved only twice: once in the lineage including trachypachids and car- abids, and again in gyrinids. The minimal pu- bescence seen in extant trachypachids can be in- terpreted as a first step in a transformation series leading to the condition found in a majority of carabids. Character 5. Orientation of mouthparts. Po- nomarenko (1977) suggested that apparent opis- thognathy seen in triaplid fossil specimens may reflect a grazing style of feeding, characteristic of a variety of herbivorous beetle groups. The known occurrence of opisthognathy among schizopho- roid Archostematan an Adephagan is such that it must be apotypic for triaplids. Character 6. Prosternal intercoxal process. Most extant and extinct Archostemata and Adephaga have a well-developed prosternal in- tercoxal process. Known triaplids appear to have lacked such a process, at least externally. This probably represents an apotypic feature. In re- lation to those of other groups, haliplids, omo- phronine carabids, and some noterids have intercoxal processes markedly expanded and strikingly similar in form and degree of contact with the mesothorax. However, adult haliplids and noterids have open procoxal cavities, where- as omophronines have them closed. While this difference may be significant, Bell (1967) pointed out that the type of procoxal closure found in omophronines was apparently unique to them. Hence, it is likely that the immediate ancestor of omophronines had open procoxal cavities. Shape of the prosternal intercoxal process is just 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Taste |. DistTRiBUTION OF STATES OF SELECTED CHARACTERS AMONG MEMBERS OF CERTAIN SUPRA-SPECIFIC TAXA OF ADEPH- AGA (CoLeopTerRA) (See Fig. 22 for Code Letters for Taxa and Text for Discussion of Characters). Taxa and character state distributions Character ee eee eons a ee Character state A Ba Ca Doerr ak: GP eke gc MN C6 ie & 1. General habitat a* a® aval a> sia ia a 236-7 tr b 1) ala. Semiaquatic, 0 Aquatic, a Aquatic, surface, a* Terrestrial, b Semiaquatic, passive drifter, c i) Food habits/feeding (0) o Oo o © 0 to) 0| 0. 50 <0 fo) 0 40 ava, Predaceous, 0 Herbivorous (on algae), a 3. Compound eyes a aoa? 00 (e) o Oo 0 © to) (ome) oe) Undivided, o Dorsoventrally divided, a 4. Antennal pubescence a* ane 2 0)8 70, to) Tea ON Beam a wae ‘oe? Without pubescence, o Only antennomere |1 pubescent, a Pubescence widespread, a* 5. Orientation of mouthparts ° ene) 27> 0) 0) to) oO 0 0 te) (omens) oa Prognathous, o Opisthognathous, a 6. Prosternal intercoxal process to) oOo ? 0; “o;\bi to 0) {01 “or “oO oO one) ba Narrow, o Absent, a Broad, b 7. Protibial antenna cleaner te) o Oo y 10 oO to) 0. O10 7) ca a aa Oo 7? Absent, 0 Present, a 8. Scutellum oa 0 0 2 Oa Oia oO OG oOo (0) 0 -‘o ao Visible externally, 0 Concealed, a 9. Mesothoracic length a ao To O- 10 [e) O1/.0') O70 (o) (ouene) (ome) Short, 0 Long, a 10. Mesocoxal shape a a o 2 O10 fe) 0 © 10; (0 ie) (oe) (one) Round, o Laterally expanded, a 11. Ventral mesocoxal articulation c Ca dmecoyin slo} ‘ob ev) b a 7 laylak Hahn Absent, 0 Coxal lobe, sternal stop, a Coxal peg, sternal socket, a* Coxal groove, sternal ridge, b Coxae otherwise immobilized, c 12. Metasternal transverse ridge a* at anar “ial kas a* a) al ore a oo © 0 Present, laterally extended, o Present, laterally reduced, a Absent, a* 13. Relationship of metepisternum to mesocoxal cavity a o a Pa ta 6,4 0 © (0 0 a oa a 0 Forms part of lateral wall, o Excluded from lateral wall, a 14. Metacoxal position a aoa TeiFacesa a ‘ao. cauikals ca a aa aoa Free of abdomen, o Countersunk into abdomen, a 15. Metacoxal width to) o Oo i Manns rice) (0) QO: 02.0) +0 oO aa o Oo Wide, o Narrow, a KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES Taste 1. CONTINUED. Taxa and character state distributions Character Character state A Bec 16. Metacoxal length to) Oo O,a Short, 0 Medium, slightly expanded anteriorly, a Long, markedly expanded anteriorly, a* 17. Metacoxal fusion a* ak@ana’: Not fused medially, o Partially fused medially, a Extensively fused medially, a* 18. Metacoxal femoral plates te) 0 0O,a Absent or small, 0 Present, moderately large, a Present, very large, a* 19. Legs, distal modifications for swimming a* a* a,a* Absent, 0 Slight modifications, a Extensive modifications, a* Femoral modifications only, b 20. Legs, fringe setae a aoa Present, slightly developed, o Present, well developed, a Absent, b 21. Hindwing apex in repose a Ey Bt Spirally rolled, o Folded, a . Hindwing, subcubital binding patch a* Present, 0 Absent, a Absent, suboblongum patch present, a* i) to 23. Oblongum cell position a a ? Posteroapical, 0 Near center of wing, a 24. Male median lobe, internal sac ° o ? Short, slightly developed, o Large, better developed, a 25. Male genitalia, parameres (a) Or Symmetrical in length and shape, 0 Asymmetrical in length and shape, a 26. Male genitalia, ring sclerite to) o ? Split posterodorsally, o Complete posterodorsally, a 27. Female ovipositor, gonostylus a Distinct, o Fused with gonocoxite, a Apparently distinct, a* 28. Thoracic defense glands te) o ? Absent, o Present, a 29. Pygidial defense gland cells fo) Type I cells absent, o Type I cells present, a E. oF Ga si hake E apa a* anna OO ° ans at a* aaal sour Oo fo) 0) (0;!a" 0 (On .O% 103) 103.2" 0) aa a (oy fo iP = Xo) fe) aa a a e071 a7, 1b, b aoa a eh a heb are a o a OpB iin ona” Cree aah fo) a o (0) fg Win ate leg ° oe) to) Dole 2) (7 ° oa o Dt eae? een te) ,a oo to) een One wee a aoa a Te aA 8. a ae ao a ee Pa te) o Oo a i eto Ape 87: te) a* 93 94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 one of several similarities (see below) among no- terids, haliplids, and omophronines that appear to represent convergences, based on data from other characters. Character 7. Protibial antennal cleaner. Sev- eral authors (e.g., Hammond 1979) have sug- gested that absence of a protibial cleaning organ may be an apotypic feature among hydradepha- gans. Also among carabid groups (e.g., paussines) in which specialized antennal structure precludes grooming by means ofa protibial cleaning organ, such an organ is absent. I see no evidence, how- ever, to suggest that presence ofa protibial clean- ing organ is plesiotypic among Adephaga, and I view its occurrence as an apotypic feature linking trachypachids and carabids. Character 8. Scutellum. A scutellum is visible externally in extant and extinct Archostemata and in Adephaga, except noterids, haliplids, omophronine carabids, some gyrinids, and some dytiscids. Because the distributions of apotypic states of several other characters are incongruous with the distribution of a concealed scutellum among Adephaga (1.e., character correlation cri- terion), there is little evidence to suggest that this trait is synapotypic for any two or more of the exceptional taxa. It probably evolved indepen- dently in each, although its co-occurrence among noterids and certain dytiscids may reflect close phylogenetic relationship. Character 9. Mesothoracic length. A signifi- cant increase in length of the mesothorax is seen in extant and fossil gyrinids, including adults of Spanglerogyrus and the inadequately known Triassic fossil form, Triadogyrus. This feature appears to be autapotypic (i.e., uniquely derived) in gyrinids. Character 10. Mesocoxal shape. Distribution of states of this character is identical with that in character 9. No doubt, the two characters are closely correlated. Among known Adephaga, only gyrinids have laterally expanded mesocoxae. Even coptoclavids, which share several other features with gyrinids, had round mesocoxae typical of the remainder of the suborder. Character 11. Ventral mesocoxal articulation. Evans (1977) reviewed various structural means found among Adephaga for ventral articulation of mesocoxae with the metasternum. He noted that amphizoids, hygrobiids, and some dytiscids evidently lack special structural means of ventral articulation. Carabids and extant trachypachids (i.e., adults of both Trachypachus and Systolo- soma species) have a coxal lobe/sternal stop mechanism, but noterids and those dytiscids with evident ventral articulations have a sternal ridge/ coxal groove arrangement. Gyrinids have me- socoxae that are practically immobilized by a unique structural arrangement which is probably independently derived. It seems that the absence of ventral articular structure is plesiotypic in Adephaga, and therefore, that articular struc- tures evolved independently in (1) noterids and some dytiscids, and (2) the lineage including trachypachids and carabids. A special coxal peg/ sternal socket arrangement is found in haliplids and omophronines (Evans 1977), although po- sition of the socket 1s different in members of the two groups. This is yet another similarity be- tween these groups, but it probably evolved in- dependently in each from the coxal lobe/sternal stop arrangement seen in other caraboids. Character 12. Metasternal transverse ridge. Evans (1977) discussed this structure (also known as the ““metasternal suture’’), its functional sig- nificance, and its distribution among Adephaga. Its presence appears to be plesiotypic and its loss or lateral reduction apotypic within the suborder. The single known Necronectulus specimen has a well-developed transverse ridge. Presence of the laterally reduced ridge in amphizoids and hygro- biids suggests that, if the cladogram is correct, its loss has occurred three times independently: in dytiscids and noterids, in gyrinids, and in some (but not all) coptoclavids. Eodromeines appar- ently had well-developed, laterally extended transverse ridges, like extant carabids, and so the laterally reduced ridge found in extant trachy- pachids probably represents reduction conver- gent with that in the dytiscomorph lineage. Ste1- ner and Anderson (1981) reported presence of a metasternal ridge in adults of Spanglerogyrus. In my own examination of representatives of this genus, I found the structure in question to be wholly part of the metacoxa rather than the metasternum. I suggest that the suture (or ridge) at the base of the metacoxa in Spanglerogyrus adults is autapotypic among them and not ho- mologous with the metasternal ridge found among Adephaga as listed in Table 1. Character 13. Relationship of metepisternum to mesocoxal cavity. As noted earlier, the plesio- typic condition among Adephaga is that in which the metepisternum contributes to the lateral wall of the mesocoxal cavity. Among extant forms, this condition is found only in amphizoids, some KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES dytiscids, and adults of the gyrinid genus, Span- glerogyrus. Although this condition may have been achieved secondarily in members of Span- glerogyrus, it is more likely that it represents yet another feature suggesting ancient ancestry for this unique genus. Exclusion of the metepister- num from the mesocoxal cavity appears to have evolved at least seven times: in (1) gyrinids (after divergence of Spanglerogyrus from the main lin- eage), (2) the lineage including hygrobiids and coptoclavids, (3) noterids, (4) some dytiscids, (5) trachypachines [this is the feature that distin- guishes them from eodromeines], (6) carabids [again, this feature distinguishes extant carabids from protorabines], and (7) haliplids. This struc- tural change must be highly advantageous me- chanically for it to have become fixed in so many different lineages making use of both terrestrial and aquatic habitats. Character 14. Metacoxal position. In all known Adephaga, the metacoxae are countersunk into the base of the abdomen so that they divide the first visible abdominal sternum externally into two triangular lateral portions. This feature dis- tinguishes Adephaga from other Coleoptera, in- cluding Archostemata. It is, no doubt, a syn- apotypic feature. Character 15. Metacoxal width. The narrowed metacoxae found in protorabines and all extant carabids (except gehringiines) are clearly apotyp- ic. The condition found in rhysodines is not equivalent to the plesiotypic state, because the metacoxae extend laterally only to the postero- lateral corner of the metasternum, just as in car- abids. The metepisterna are hidden posteriorly under the elytral epipleura, but they are com- pletely laterad of the lateral margins of the meta- coxae. Evans (1977) noted that, unlike those in dy- tiscids, gyrinids, and other Hydradephaga, meta- coxae of haliplids have a lateral coxal condyle, as do carabid metacoxae. I agree this feature in- dicates close affinity with a presumed terrestrial ancestor, namely carabids. However, a coxal condyle is also present, although not as well de- veloped, in extant trachypachines and amphi- zoids, but not in hygrobiids and other dytiscoids. Presence of a coxal condyle may represent the plesiotypic condition among Adephaga. The lat- erally extended metacoxae of haliplids may rep- resent either the plesiotypic adephagan condition or secondary acquisition of a similar condition as part of an adaptation of metacoxae for a new 95 function (see further discussion under character 18). Character 16. Metacoxal length. Slight to moderate expansion of metacoxae anteriorly, and attendant reduction in size of the metasternum, is seen in amphizoids, liadytids, hygrobiids, and some coptoclavids. In dytiscids and noterids, metacoxae are greatly expanded anteriorly. Based on distributions of states of other characters, it is likely that the trend for anterior expansion, which was initiated in the common dytiscoid ancestor, has been reversed at least twice inde- pendently: in gyrinids and in some coptoclavids. Character 17. Metacoxal fusion. This char- acter was discussed above in consideration of the relationship between amphizoids and _ trachy- pachids. Based on character states represented in extinct and extant Archostemata and Mesozoic Adephaga, it is clear that the unfused metacoxae are plesiotypic. As noted by Evans (1977) and others, the metacoxae of haliplids are not fused medially. It is therefore likely that at least partial medial fusion of metacoxae represents a syn- apotypic feature for Hydradephaga exclusive of haliplids (and Necronectulus, if its members were aquatic). A trend for more extensive medial fu- sion of metacoxae may have evolved only once— in the common ancestor of all dytiscomorphs except amphizoids. If so, then this trend was also reversed at least once, in the ancestor of some (but not all) coptoclavids. Eodromeine trachy- pachids appear to have had more widely sepa- rated metacoxae than extant trachypachids. Hence, a trend toward increased medial conti- guity, if not fusion as suggested by Evans (1977) and Roughley (1981) for extant trachypachids, probably represents a development independent of that in Hydradephaga. Character 18. Metacoxal femoral plates. Po- nomarenko (1977) described posteroventral ex- tensions of metacoxae, which he termed femoral plates, in triaplids, some coptoclavids, and some eodromeines among Mesozoic fossil forms. He noted that such plates could be rather easily bro- ken offand, therefore, that their distribution may have been taxonomically more extensive than present fossil material illustrates. He also sug- gested that presence of metacoxal plates may be plesiotypic for Adephaga. Such structures are ap- parently unknown among schizophoroid Ar- chostemata, however, and among extant forms, femoral plates are found only in noterids and haliplids. In my view, it is simplest to consider 96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 presence of femoral plates as apotypic where they occur among Adephaga. There is little or no evi- dence to suggest that this feature is synapotypic for any two or more of the extinct and/or extant groups whose members are known to possess them. Haliplid metacoxal femoral plates are much larger, both posteriorly and laterally expanded, than those of noterids and the extinct groups listed above, including triaplids. The role of these plates in haliplid respiration has been well doc- umented (Hickman 1931). It is unlikely that they served this highly specialized function in triap- lids, coptoclavids, or eodromeines, and no respi- ratory role has been suggested for them among noterids. Character 19. Legs, distal modifications for swimming. Based on comparisons with Archo- stemata and Mesozoic Adephaga, it appears that distal leg structure in amphizoids represents the plesiotypic state among Adephaga. Special struc- tural modifications of femora, tibiae, and/or tarsi as adaptations for rapid swimming are consid- ered apotypic. Relatively slight modifications of this kind have apparently evolved twice inde- pendently: in the common ancestors of (1) no- terids and dytiscids and (2) hygrobiids, copto- clavids, and gyrinids. In each of these groups, structure of distal leg parts is quite distinctive in detail. Gyrinids and members of the extinct ge- nus Coptoclava (Ponomarenko 1977) are similar in that their middle and hind legs are (or were) markedly flattened and expanded. This feature is probably apotypic relative to more conserva- tive leg modification, but distributions of states of other characters suggest that it is not syn- apotypic for these two groups. Middle and hind legs of haliplids show no spe- cial structural adaptations for swimming. Hali- plid hind femora are unique in that they are markedly narrowed basally—a feature probably evolved to facilitate leg movement within the narrow space between abdominal venter and metacoxal femoral plates. Character 20. Legs, fringe setae. Unfortu- nately, some of the most important extinct Me- sozoic groups are known only from specimens without distal leg parts. Among these are Tri- aplidae and genus Necronectulus. Hence, it is dif- ficult to know whether or not ancestral Adephaga had legs bearing fringe setae (or so-called “‘swim- ming hairs’’). Assuming a semiaquatic ancestry, the condition found in extant amphizoids, in which fringe setae are present but short and lim- ited in distribution, could be considered the ple- siotypic condition. Absence of fringe setae would then be synapotypic for trachypachids and car- abids. If the cladogram in Figure 22 is correct, then more extensive development of fringe setae would also be apotypic. But this feature would have had to have evolved at least twice inde- pendently: in (1) the common ancestor of all Hy- dradephaga except amphizoids, and (2) haliplids. Fringe setae are longer and more extensively dis- tributed in haliplids than in amphizoids—this is perhaps associated with a slightly better devel- oped aquatic lifestyle. Character 21. Hindwing apex in repose. Mem- bers of all Adephaga groups examined have the hindwing apex folded, rather than spirally rolled as in Archostemata. This feature is probably syn- apotypic for the suborder Adephaga. Character 22. Hindwing, subcubital binding patch. If presence of the subcubital binding patch is synapotypic for suborder Adephaga (hence, plesiotypic within Adephaga, see above), then loss of the patch has evolved at least five times: in (1) the common ancestor of carabines and (probably) protorabines, (2) haliplids, (3) trachy- pachids of genus Systolosoma, (4) some dytis- cids, and (5) the common ancestor of gyrinids, hygrobiids, and (probably) coptoclavids. Gyrinid specimens examined have a narrow patch of short setae or long microtrichia along the posterior margin of the oblongum cell that may aid in wing folding as an alternative to or replacement for the subcubital patch. Character 23. Oblongum cell position. Ham- mond (1979) noted that the oblongum cell is positioned closer to the posterior margin of the wing apex in trachypachids, amphizoids, noter- ids, and dytiscids than in other Adephaga and considered this to represent a synapotypic fea- ture for the groups noted. Position of the oblon- gum cell in archostematan hindwings, however, is also close to the posterior margin of the apex, just as in amphizoids and other taxa noted by Hammond. I conclude that this feature is ple- siotypic, and further, that a more anterior and basal placement of the cell is apotypic. If this view is correct, then the apotypic state could have evolved as few as three times: in (1) cara- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES bids, (2) haliplids, and (3) the common ancestor of hygrobiids, coptoclavids, and gyrinids. Character 24. Male median lobe, internal sac. As noted above, it is likely that a large, well- developed internal sac, such as is found in most carabid males, is apotypic among Adephaga. Males of basal-grade rhysodid lineages have larg- er internal sacs than those of more highly evolved lineages, but this trend appears to reverse that seen among carabids in general. Character 25. Male genitalia, parameres. Based on comparisons with genitalia of extant Archostemata, it appears that the plesiotypic form of parameres among Adephaga demonstrates symmetry in both length and shape. Asymmet- rical parameres are found in noterids, haliplids, and most, but not all, carabids. Based on the distribution of this feature in relation to char- acter-state distributions of other characters, it is likely that asymmetry of parameres evolved in each of these groups independently. Character 26. Male, ring sclerite. The ring sclerite (Kavanaugh 1978+) and associated struc- tures probably represents the sclerotized remains of the ninth abdominal segment (the genital seg- ment, or urite X of Jeannel 1941), and it serves as arim for attachment of muscles from the base of the median lobe. In all Hydradephaga ex- amined, except haliplids, the ring is split postero- dorsally in the midline, into what might be termed ““hemitergites,” but is continuous anteroventral- ly (see Edwards 1951, “Plate 2”’). This condition is shared with Archostemata males examined. In trachypachids, carabids, and haliplids, however, the ring is complete posterodorsally as well as anteroventrally—a feature that is probably syn- apotypic for these three groups. Character 27. Female ovipositor, gonostylus. Bell (1982) and others have suggested that the apparent absence of a gonostylus (or stylomere two) from ovipositors of female trachypachids, isochaetous carabids, and hydradephagans may represent a synapotypic feature uniting these groups. In fact, a majority of basal-grade carabid groups (e.g., opisthiines, notiokasiines, nebri- inies, and notiophilines) also have females in which a gonostylus is either absent from the ovi- positor or fused with the gonocoxite (stylomere one) so as to appear absent. I agree with Bell that this feature is apotypic, but suggest that it is syn- apotypic for the suborder Adephaga rather than 97 just for a subgroup of that taxon. The structures that have been called gonostyli (or second sty- lomeres) in female carabines, cychrines, cicinde- lines, and a majority of intermediate- and advanced-grade carabids are probably not ho- mologous with the gonostyli of female Archo- stemata and Polyphaga. Character 28. Thoracic defense glands. For- syth (1968, 1970) noted that, among Adephaga, only hygrobiids and dytiscids possess thoracic defense glands in addition to the pygidial defense glands common to all Adephaga. Presence of such thoracic glands is no doubt apotypic in hygro- biids and dytiscids, but based on the character correlation criterion, I agree with Forsyth that this similarity represents convergence rather than common ancestry. Character 29. Pygidial gland cells. In a series of papers describing the structure of pygidial and other defense glands among Adephaga, Forsyth (1968, 1970, 1972) provided numerous excellent characters, while making detailed comparisons among members of included taxa, but he did not consider states of these characters from a cladis- tic perspective. The relationships he suggested were based on simple similarity, rather than on synapotypy, and I have been unable to recognize patterns of synapotypy among the mass of data he provided for included adephagan taxa. Forsyth (1968) recognized two types of secre- tory cells (Type Land Type II cells) in the pygidial glands of dytiscids. Apparently, only Type II cells are found in these glands in other Adephaga, and presence of Type I cells in dytiscid pygidial glands must be autapotypic. Summary of phylogenetic reconstruction. Sev- eral final points should be made in reference to the proposed cladogram and data provided in Table 1. First, the monophyly of a lineage in- cluding all Adephaga except triaplids is unsup- ported at present by evidence in the form of syn- apotypic features. We know too little about triaplid structure and lifestyle to recognize fea- tures in which their proposed sister-group may be considered specialized (i.e., apotypic). I also failed to discover any synapotypic feature uniting Trachypachinae with Eodromeinae. However, eodromeines probably represent the ancestral stock from which both trachypachines and car- abids evolved. A group including trachypachines and eodromeines but excluding carabids would 98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 therefore be paraphyletic, which may explain why synapotypic features for such a group are lacking. Monophyly of a group including both extinct and extant trachypachids, carabids, and haliplids is supported by fewer and less compelling syn- apotypic features than might be desirable. The only proposed synapotypies for this group are the following: (1) mesocoxal ventral articulation by means of a coxal lobe and sternal stop or derivative of this arrangement, and (2) male gen- italia with ring sclerite complete posterodorsally. Nonetheless, available evidence supports a clos- er phylogenetic relationship between haliplids and carabids than between the former and other Hy- dradephaga. As can be seen in Table 1, noterids share apo- typic features (e.g., see characters 8, 11, and 13) with some, but not all, dytiscids. This suggests a close relationship between noterids and only some dytiscids. It is therefore possible that if Dytis- cidae (in the broad sense) is a monophyletic tax- on (and there is considerable doubt in this regard; Roughley, pers. comm.) then it would be a para- phyletic taxon if noterids were excluded and/or recognized as a separate family. On the other hand, dytiscids possess thoracic defense glands and Type I secretory cells in their pygidial de- fense glands, whereas noterids studied to date have neither of these features. Available evi- dence is therefore equivocal with regard to the question of relationship between noterids and dytiscids. However, I suggest that noterids and dytiscids should be taken together as a mono- phyletic unit of greater inclusiveness, whether at the familial or some higher taxonomic level, to assure that appropriate comparisons are made in future studies. The proposed relationship between hygrobiids and coptoclavids also requires further comment. Among characters used in this study, I found no apotypic states that distinguish all members of either group from all members of the other. Some coptoclavids have apotypic features not shared with hygrobiids, but the reverse does not apply, except perhaps for the presence of thoracic de- fense glands in hygrobiids (but coptoclavids may also have had such glands). Hygrobiids are most similar to certain members of Necronectinae (Ponomarenko 1977). Together, these groups ap- pear to represent a basal grade of coptoclavid evolution, and I predict that future studies will indicate that hygrobiids and coptoclavids should be included in a single family. Phylogenetic relationships of amphizoid species Based on assumed adephagan phylogenetic re- lationships as illustrated in Figure 22, a cladistic analysis was conducted to ascertain relationships among extant amphizoid species. A total of 14 selected characters was used. For each, the out- group criterion was used to establish polarity (from plesiotypic to most apotypic) of character- state transformation. Characters and character- state distributions among amphizoid species are presented in Table 2, and the cladogram that results from analysis of these data is illustrated in Figure 23. Format and coding for characters and character states used in Table 2 and Figure 23 are as explained above for Table | and Figure 2? If the hypothesis of phylogenetic relationship proposed—namely that Amphizoa davidi is the sister-group of the other three species, and that A. insolens is the sister of the group including 4. striata and A. lecontei—is correct, then the fol- lowing comments are appropriate. Character-state distributions of all characters analyzed are compatible with each other over the cladogram, except for character 3 (sinuation of the lateral margin of the pronotum). Develop- ment of a deep sinuation basolaterally is evident in adults of 4. davidi and A. insolens. Although nothing is presently known about habitat re- quirements and/or tolerances of A. davidi mem- bers, those of 4. insolens are often found in swift- er-flowing, more precipitous streams than are members of A. striata or A. lecontei. A deep sub- basal sinuation of the lateral pronotal margin is also found in certain dytiscids (e.g., members of genus Hydronebrius Jakovlev and of the cordatus group of Agabus), which also live in fast-flowing streams. This suggests that the apotypic state of this character (1.e., lateral margin deeply sinuate sub-basally) may be associated with adaptation to life in swift-flowing streams, and distributions of states of other characters suggest that this fea- ture evolved independently in 4. davidi and A. insolens. An alternative, equally parsimonious interpretation of the distribution of character states is that a deep, sub-basal sinuation evolved among members of the common ancestor of ex- tant Amphizoa species and is therefore synapo- typic for the genus. An evolutionary reversal then occurred in members of the common ancestor of A. striata and A. /econtei. If this interpretation KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES Oe TaBLe 2. DisTRIBUTIONS OF STATES OF SELECTED CHARACTERS AMONG Members OF Amphizoa Species (See Text for Discussion of Character Coding). Taxa and character state distributions Character Character state davidi Insolens striata lecontei 1. Macrosculpture, elytra to) a fo) b Not rugose or slightly rugose basally, punctures distinct, 0 Markedly rugose basally, punctures distinct, a Slightly rugose basally, punctures confluent, b i) . Pronotum, shape fe) a oO (o) Widest at base, 0 Width at middle and base equal, a 3. Pronotum, sinuation of lateral margin a a to) fs) Absent or shallow, 0 Deep, a 4. Pronotum, lateral margin o a* a a Not crenulate, 0 Slightly crenulate, a Markedly crenulate, a* 5. Prosternal intercoxal process, shape a fo) to) (o) Elongate, spatulate, o Short, circular, a 6. Elytra, silhouette (dorsal aspect) fe) a b b Moderately broad basally, narrowed subapically, o Subovoid, slightly narrowed basally, less narrowed subapically, a Very broad basally, narrowed subapically, b 7. Elytra, silhouette (cross-sectional aspect) te) fo) a a* Evenly convex, 0 Convex medially, slightly concave laterally, a Carinate, flat medially, concave laterally, a* 8. Male median lobe, shaft thickness oO a a* a* Slender at middle, o Slightly thickened at middle, a Markedly thickened at middle, a* 9. Male median lobe, ventral margin fo) oO a a Evenly arcuate, 0 Slightly bulged, a 10. Male median lobe, shape apex fo) a to) to) Slightly deflected ventrally, o Extended apicodorsally, a 11. Male left paramere, shape fo) fo) a a Narrow basally, o Broad basally, a 12. Male parameres, vestiture fo) a oO te) Restricted to apical one-fourth, 0 Restricted to apical one-third, a 13. Female ovipositor, length of coxostylus 2, fo) a a* Short, 0 Medium, a Long, a* 14. Female ovipositor, vestiture of coxostylus ? a to) fe) Dense, evenly distributed setae, 0 Sparse, scattered setae, a is correct, then absence of a deep sinuation from _ others used for descriptive purposes only), no adults of the last two species mentioned repre- _apotypic feature was found to unite all members sents yet another synapotypy for these taxa. of A. striata, although six (or seven, see above) Among the characters used in this analysis(and — synapotypic features support a sister-species re- 100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 striata davidi 23 Ficure 23. 6b,7a,8a*®,9a, 11a, 13a lecontei insolens 1b,7a®,13a*® la, 2a,3a,4a®,6a, 10a, 12a, 14a Reconstructed phylogeny of species of Amphizoa. Number and letter symbols placed adjacent to solid dots refer to synapotypic features presented in Table 2 and discussed in the text. lationship for A. striata and A. lecontei. Adults of A. striata are distinctly larger than most mem- bers of other amphizoid species; but it was not possible, using the out-group criterion, to affirm that this represents an apotypic feature. Zoogeography and Evolution In this section, I briefly review the present geo- graphical and habitat distributions of amphi- zoids and then discuss what can now be inferred about the zoogeographic and evolutionary his- tory of this group. Present pattern of amphizoid distribution The present pattern of geographical distribu- tion of Amphizoidae is disjunct across the north- ern Pacific Basin, with three species (Fig. 18- 20) restricted to western North America and one (Fig. 17) to central China. This pattern reflects a vicariance relationship, with the Palaearctic species recognized as the sister-group of the three Nearctic forms. Among North American species, the distri- bution of A. insolens (Fig. 18) is mainly coastal (1.e., east to the Sierra Nevada and Cascade Range), with range extensions east into mountain ranges of the Great Basin in Nevada, Oregon, and Idaho, and to the Northern Rocky Moun- tains of Idaho, Montana, Alberta, British Colum- bia, and Yukon Territory. The sister-group of this species includes A. /econtei, restricted to the Rocky Mountain region (Fig. 20), and 4. striata, restricted to western Oregon, western and central Washington, and Vancouver Island, British Co- lumbia (Fig. 19). A vicariance relationship is ap- parent between A. /econtei and A. striata across the northern Great Basin and Columbia Plateau. However, because the ranges of A. striata and A. insolens overlap extensively, 4. insolens and its sister-group are not strictly vicariant at present. The habitat distribution of extant amphizoids is apparently quite limited. Members of all three Nearctic species are confined to cool or cold streams. Members of 4. striata are found in slow- flowing, relatively warm streams, those of A. /e- contei in cooler or cold, moderate- to fast-flowing streams, and those of 4. insolens in cold, fast- flowing or cascading streams. Habitat is un- known for A. davidi members; however, the type- locality of this species is in a region occupied by vegetation types that Wolfe (1979) called “‘no- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES tophyllous broad-leaved evergreen forest” and “mixed broad-leaved evergreen and coniferous forest.” Western North American vegetation types with apparently equivalent temperature re- quirements include “mixed coniferous forest,” “‘mixed evergreen forest,” and “California wood- lands” (in part) (Griffin and Critchfield 1972). These vegetation types are almost completely re- stricted to areas in California at present, and members of A. insolens are found in streams as- sociated with such forests. I suggest that the hab- itat of A. davidi members will be found to be similar to that for 4. insolens members, although the former may prefer slightly warmer and slow- er-flowing streams than the latter. Mesozoic events and the origin of amphizoids According to the hypothesis of adephagan phylogenetic relationships proposed above and illustrated in Figure 22, amphizoids are the sis- ter-group of all other Hydradephaga, except hal- iplids. If this is correct, then divergence of these sister-groups probably occurred at about the Per- mo-Triassic boundary, and certainly no later than Upper Triassic time. Although there are no am- phizoid fossil specimens known from that time, fossils representing a diverse array of other hy- dradephagan taxa document a relatively exten- sive radiation of the structurally more advanced sister-group of amphizoids by Upper Triassic and Jurassic time. In the Triassic, the supercontinent of Pangaea was still intact (Smith, Briden, and Drewry 1977), and climate was apparently warm and equable over the entire landmass (Hallam 1981). Local climatic anomalies, associated with physiogra- phy and/or relative proximity to the ocean, may have provided some diversity of habitats, but there is no evidence for broad, latitudinally lim- ited climatic zones such as occur on continents at present. Both early and late Palaeozoic glacia- tions have been recognized (Tarling 1978), with most of these associated with high latitude po- sitions of the continents affected. No major gla- ciations appear to have occurred during all of the Mesozoic, however, probably because the con- tinents were all positioned at relatively low lat- itudes. At present, we have no information from which to infer the geographical and habitat distribu- tions of amphizoids during early Mesozoic time. Again, these beetles are not known from the fossil 101 record; their sister-group includes both extinct groups, presently known only from Mesozoic Asia, and extant groups with widely disjunct (e.g., Hygrobiidae) or worldwide (e.g., Dytiscidae and Gyrinidae) distributions. Because (1) there is ex- tensive sympatry at the familial level and (2) comprehensive hypotheses of phylogenetic re- lationships within families have not yet been for- mulated, it is currently impossible to recognize vicariance relationships between amphizoids and their sister-group. Hence, amphizoids could have been either widely distributed in Pangaea or geo- graphically restricted to some unknown part of that supercontinent. Structurally, extant amphizoids appear to have diverged little, if at all, from the hypothetical common ancestor of all Hydradephaga (exclud- ing haliplids). It is their sister-group, whose ex- tant descendants include hygrobiids, dytiscoids, and gyrinoids, that evolved rapidly away from the presumed ancestral form and lifestyle in adapting to a more fully aquatic existence. How then were amphizoids able to survive presumed early competition with members of their ad- vanced sister-group, whereas other lineages with- in their sister-group (e.g., liadytids and most cop- toclavids) appear to have been replaced by more highly evolved forms? Amphizoids may have persisted in geographical isolation from their sis- ter-group for an extended period. Eventually, a shift of habitat— namely to faster-flowing water— may have reduced the potential for competition with other, more rapidly diversifying Mesozoic hydradephagan groups. Even to the present, dy- tiscoids and their allies have exploited lotic hab- itats in only a limited manner, especially in geo- graphical areas where amphizoids now occur. There is no reason to suggest that amphizoids also became adapted to cool- or cold-water hab- itats so early in their history. Such habitats may have been available locally, but as noted above, climate was generally warm and equable throughout Pangaea (Hallam 1981) at that time. Cool- or cold-water specialization would seem to have been a risky adaptive strategy at that time—one that could well have led to extinction during or before early Cenozoic time (see below). At present, there is no way to infer what (if any) effect Mesozoic plate-tectonic processes, re- sulting in fragmentation of Pangaea, may have had on the Mesozoic amphizoid fauna. Of po- tentially greater impact, however, were eustatic changes in Jurassic and Cretaceous time that re- 102 sulted in formation of epicontinental seas in Eur- asia (the so-called “‘Turgai Sea,” late Middle Ju- rassic through Oligocene) and North America (mid-to-late through latest Cretaceous) (Hallam 1981). Because there appear to have been con- tinental connections between eastern North America and western Europe on one hand and western North America and Asia on the other, two new land masses were formed, which Cox (1974) called ““Euramerica”’ and ‘‘Asiamerica,” respectively. Fossil evidence suggests that biotas subsequently evolved independently on each landmass (Cox 1974; Hallam 1981), resulting in increased endemism in each area by the end of the Mesozoic. The geographical range of extant amphizoids is confined to land area derivatives of Asiamerica, and it is tempting to suggest that amphizoids were at least present on that land- mass, if not also restricted to it, during Creta- ceous time. Tertiary events and amphizoid radiation As just noted, there is no evidence to suggest that the late Mesozoic distribution of amphi- zoids extended outside an area including eastern Asia and western North America (Fig. 24), al- though a more extensive distribution was cer- tainly possible. The first direct land connection between these areas occurred well before the end of the Cretaceous, as a consequence of spreading of the North Atlantic (Hallam 1981), and per- sisted continuously until late Miocene time (Hopkins 1967). Then, between 10 and 12 mil- lion years before present (mybp), a_trans- Beringian seaway developed, which linked the North Pacific and Arctic basins but interrupted the exchange of terrestrial and freshwater (aquat- ic) biota between North America and Asia. A land connection was re-formed in Pliocene time and permitted renewed biotic exchange until about 3.5—4.0 mybp, at which time the trans- Beringian seaway opened again (Hopkins 1967). In Quaternary time, the Beringian land connec- tion was re-established during several, if not each, of the major glaciations, and further biotic ex- change is known to have occurred during this period (Repenning 1967). Finally, the seaway opened for the last time more than 11,000 years ago, and it has remained a substantial barrier to east-west biotic movement since that time. Palaeobotanical and other evidence indicates that early Cenozoic climates were as warm and equable as those of the Mesozoic. Then, in late PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Eocene time, an abrupt cooling occurred in the northern hemisphere. This cooling trend leveled off in Oligocene time; but cool conditions have persisted, with both major and minor fluctua- tions (e.g., the various Pleistocene glaciations), to the present. Another set of events that had a profound effect on climate, especially in western North America, were the episodes of orogenic and volcanic activity in Miocene and Pliocene times. This activity produced topographic relief that resulted in local and regional rain-shadow effects, increased diversity of microclimates, and increased seasonality. Geographical regions of Asia and North Amer- ica that are now occupied by extant Amphizoa species appear to have shared closely related flo- ras in early Tertiary time. These floras were of the evergreen sclerophyllous broad-leaved and mixed mesophytic forest types (Leopold and MacGinitie 1972). Floral affinities between Asia and western North America were very close in Paleocene and early Eocene time. However, by middle-to-late Eocene time, floras of the Rocky Mountain region were quite distinctive. Leopold and MacGinitie (1972) suggested that edaphic conditions associated with local volcanic activity may have stimulated selection for xeric-adapted vegetation. Although affinities between floras of southeastern Asia and the Pacific coast of North America decreased more gradually, they were nonetheless very slight indeed by late Miocene time (Wolfe and Leopold 1967). Differentiation of the North American floras appears to have been closely related to general cooling begun in late Eocene time and to middle through late Ter- tary orogenic activity in the Pacific Northwest region. Two features that seem to characterize devel- opment of the North American floras more than contemporary floras of southeastern Asia include wholesale selective elimination of broad-leaved evergreen elements, and recruitment of subtrop- ical and temperate elements from Neotropical floras (Wolfe 1978). The first feature is no doubt related to decreasing temperatures and/or in- creased seasonality in the region; the second may simply indicate that derivative Neotropical ele- ments were already well suited to life in arid regions and could readily move into such habi- tats as they appeared and expanded. The historical factors that resulted in the vi- cariance relationship observed between A. davidi and the three Nearctic species may be the same KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES Ficure 24. factors that led to initial isolation of ancestral stocks and thereby permitted differentiation to proceed. These factors may include: (1) the gen- eral cooling trend that began abruptly in late Eocene time, which resulted in the elimination of subtropical and warm temperate vegetation types and their biotic associates from the Be- ringian region by late Miocene time; (2) Miocene orogenic and volcanic activity, particularly in western North America, which resulted in lati- tudinal and altitudinal climatic zonation, in- creased climatic and habitat diversity, and de- velopment of physiographic and local climatic barriers to north-south and east-west continuity of biotic distribution; and (3) opening of the trans- Beringian seaway in late Miocene time, which effectively severed faunal continuity between Asia and North America for about two million years. Any of these factors, either singly or in combi- nation, could have effected a division of the geo- graphical range of the common ancestor of extant amphizoids into Asian and North American iso- lates, and all three point to a middle-to-late Mio- cene age for the vicariance event in question. Hypothetical distribution of ancestral amphizoid stock, late Cretaceous to middle Eocene time. Because extant Nearctic and Palaearctic am- phizoids all appear to be cool-adapted, it is likely that their common ancestor was also cool-adapt- ed rather than that such an adaptation was ac- quired independently in the two lines. If this is correct, then it is another indication that isola- tion of respective ancestral stocks occurred after initiation of the late Eocene cooling trend, hence in Miocene time. Because there do not appear to have been any extensive areas of cool-temperate climate in the northern Pacific region prior to late Eocene or Miocene time, it is unlikely that amphizoids had specialized at an earlier time for life in a cool climate. As noted above, a vicariance relationship is not readily apparent between A. insolens and its sister-group, including A. /econtei and A. striata, due to rather extensive sympatry. However, present distribution patterns of these species are at least suggestive of an initial split of the an- cestral Nearctic stock into eastern and western vicars, the latter represented at present by 4. insolens, the former by its sister-group (Fig. 25). 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Ficures 25, 26. The present range of A. insolens is primarily cen- tered in and west of the Sierra Nevada and Cas- cade Range. Present populations in mountain ranges of the Great Basin, in the northern Rocky Mountains, and in Yukon Territory, could be viewed as representing more recent dispersal eastward from areas along the Pacific Coast. Development of the Cascade Range and Sierra Nevada was a gradual process (King 1977) that apparently had little effect on Pacific Northwest biota before late Miocene time. At that time, differences between floras east and west of the divide first became apparent (Wolfe 1969). Flo- ras east of the divide began to include elements adapted to drier summers and increased season- ality, while composition of the western flora con- tinued to reflect a more humid, somewhat less seasonal climate. From late Miocene time to the present, topographic relief has continued to in- crease, resulting in greater seasonality and aridity in the east, and increasingly greater differences between trans-montane climates and associated biotas. Hypothetical distributions of amphizoid ancestral stocks. Figure 25. Late Miocene to early Pliocene time; ancestral stocks of 4. insolens (stippled areas) and A. striata and A. lecontei (cross-hatched area). Figure 26. Middle Pliocene to end of Tertiary; ancestral stocks of 4. insolens (stippled area), 4. /econtei (cross-hatched area), and A. striata (obliquely hatched area). Based on proposed phylogenetic relationships among extant Nearctic amphizoid taxa and re- spective habits of their members at present, it seems likely that Nearctic amphizoids were adapted for life in cool (but not cold), slow- to only moderately fast-flowing, lowland or lower- montane streams during late Miocene time. Con- sequently, development of the extensive north- south trending Sierra-Cascade mountain system served as a barrier that effectively isolated the ancestors of A. insolens west of the divide and the common ancestors of A. /econtei and A. stria- ta east of it (Fig. 25). Based on inferred associations of amphizoids with particular early and mid-Tertiary vegeta- tion types and the known distributions of the latter and/or their descendant vegetation types during mid-Tertiary time (Leopold and Mac- Ginitie 1972; Wolfe 1969, 1978), I suggest that the common ancestor of 4. striata and A. lecontei occupied a broad geographical range—one that extended from the eastern flank of the Sierra- Cascade divide eastward to include at least parts KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES of the Rocky Mountain region—during late Mio- cene and/or early Pliocene time (Fig. 25). The northern extent of this range was probably lim- ited by development of a much cooler, conti- nental climate east of the Coast Mountain system in British Columbia. Pliocene fossil assemblages from areas east of the Cascades reflect increasing aridity, probably due to the enhanced rain-shadow effect of the rising Cascade Range, and increased seasonality in the region (Wolfe 1969). Eventually, this trend resulted in isolation of the last (relict) broad- leaved deciduous remnants of early Tertiary flo- ras on opposite sides of the Columbia Plateau and northern Great Basin (i.e., just east of the Cascades in central Oregon [Wolfe 1969] and on the western fringe of the Rocky Mountain system in central Idaho [Leopold and MacGinitie 1972]). This climatic change may have been the histor- ical event that isolated respective ancestral stocks of A. striata (in the west) and A. /econtei (in the east) (Fig. 26) and led to their divergence and, ultimately, speciation. A vicariance relationship between these taxa is still apparent at present. Quaternary history and development of the present amphizoid fauna If the sequence and timing of vicariance and speciation events suggested above is correct, then extant amphizoid diversity was achieved prior to Quaternary time (Fig. 26). Pleistocene and Recent events appear to have played a relatively minor role in the evolution of the present am- phizoid fauna. Nonetheless, available evidence suggests that important changes in geographical (Fig. 27) and habitat ranges of the Nearctic species and in structural, physiological, and behavioral characteristics of their members occurred during Quaternary time. Geologic, climatic, and biotic events of the Quaternary are relatively well known, and the reader is referred to Black, Goldthwait, and William (1973), Heusser (1960), Wright and Frey (1965), and references therein for pertinent information on the period. Amphizoa insolens LeConte. The ancestral stock of this species appears to have been isolated in the area west of the Cascade-Sierra divide in late Miocene time (Fig. 25). Subsequently, and probably in response to profound cooling (as- sociated with local and regional glaciation) and the continued rise of the Cascade-Sierra and Coastal mountain systems during early Pleisto- 105 cene time, members of this species acquired sev- eral adaptations for life in cold, fast-flowing montane streams. Adult structural changes apparently associated with adaptation to such streams included (1) modification in pronotal and elytral shape, which actually appears to have reduced streamlining, and (2) reduction in the size and extent of fringe setae on legs. Both of these changes may have accompanied a shift in locomotory behavior among members of this species from limited use of both swimming movements and passive trans- port with stream current to almost complete reliance on the latter locomotory mode. This lo- comotory strategy, common to all extant am- phizoids, is most highly developed in A. insolens adults. Reliance on passive transport with stream cur- rent in montane areas presents amphizoids with a high risk of drifting downstream into lowland areas of warmer climate where they cannot sur- vive. To counteract downstream displacement, they may resort to either crawling back upstream on the substratum (in the water against the cur- rent, or out of water along stream banks) or flight. I have observed the former activity repeatedly, but this must result in very slow progress. Am- phizoids have very large, thick-veined hind- wings, and they appear to be capable of strong flight. The only record for amphizoid flight to date, however, is that of Darlington (1929). Finally, increased cold-tolerance is also evi- dent among 4. insolens members, and this trait probably accounts for the success of this species in extending its geographical range so remarkably (Fig. 18). Eastward range expansion across the Great Basin and more northern Columbia and Central plateaus probably occurred during a ma- jor glacial (or pluvial in this area) period (Fig. 27). A general lack of evident differentiation among members of widely isolated populations over a large part of the Great Basin and western Rocky Mountain flank suggests that the present extent of range was achieved relatively recently, perhaps during the Wisconsinan. Similarly, members of populations in coastal Alaska and British Columbia are undifferentiated from those in populations to the south. Hence, occurrence of these populations in formerly glaciated areas probably represents postglacial range extension through dispersal from the south (Fig. 28). Amphizoa striata Van Dyke. Although they share several apotypic features with members of 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Ficures 27, 28. A. lecontei and no doubt represent the sister- group of that species, 4. striata adults are sur- prisingly similar to the hypothetical common ancestor of Nearctic amphizoids in general form, structure, and habits. Their occurrence in rela- tively warm, slow-flowing streams is unique among extant amphizoids, but such streams probably represent the ancestral (plesiotypic) habitat. The present geographical distribution of this species (Fig. 19) suggests that its members are only marginally adapted to a continental cli- mate. I have proposed that the ancestral stock of this species first became isolated and differentiated on the eastern flank of the Cascade-Sierra divide, at the western limit of the Columbia Plateau and northern Great Basin, during Pliocene time (Fig. 26). This hypothesis requires that the present distribution pattern (Fig. 19) resulted from sub- sequent westward range extension over, around, or through the divide in Pliocene or Quaternary time. Several present lowland routes through or around the divide (e.g., through the lower Fraser and Columbia River valleys or across the low Figure 27. Hypothetical distribution of ancestral amphizoid stocks, mid-Pleistocene glacial period. Figure 28. Present distributions of Nearctic Amphizoa species. Limits of geographical distribution: 4. insolens = solid line; A. striata = dotted line; 4. /econtei = dashed line. area north and east of the Pit River in northern- most California) probably also existed through at least part of Pliocene and Pleistocene time. Populations of A. striata, members of which were marginally adapted to the regional climate of the Great Basin and Columbia Plateau, were appar- ently able to disperse westward along lowland routes and subsequently expand their range through the Willamette and Puget lowlands and into adjacent low mountains. Because potential dispersal routes were probably either filled with, or greatly restricted by, montane glaciers during major glaciations (Fig. 27), it is more likely that westward range extension coincided with some interglacial period. Nevertheless, an early post- glacial origin for the present pattern cannot be ruled out (Fig. 28). Amphizoa lecontei Matthews. Adaptation to a continental climate was probably well under way among western Rocky Mountain populations of the common ancestor of A. /econtei and A. striata (Fig. 25) even before the complete isolation of eastern and western descendant stocks (Fig. 26). The present geographical distribution of A. /e- KAVANAUGH: SYSTEMATICS OF AMPHIZOID BEETLES contei (Fig. 20) suggests that members of this species now require such a climatic regime for survival. At present, Amphizoa lecontei is widely dis- tributed in the Rocky Mountain region. Many extant populations, especially at the southern limits of distribution, occupy mountain ranges that are now widely separated by warm, arid low- lands. There is considerable geographical vari- ation in characters of form and structure among members of these disjunct populations, but the pattern of variation is highly discordant (see above). This suggests that the ancestral stock of this species became widely distributed through- out the central and southern Rocky Mountain regions during a major glacial period (probably the Illinoian) (Fig. 27). During a subsequent in- terglacial (e.g., the Sangamon), the formerly con- tinuous geographical range became fragmented, and isolated populations differentiated to a lim- ited degree. During one or more subsequent gla- cial periods (probably the Wisconsinan glacia- tions), ranges of previously isolated populations came in contact, and secondary intergradation occurred among several differentiated forms. Ex- tant populations achieved their present geo- graphical relationships (Fig. 20, 28), as disjunct isolates, in response to postglacial warming; the present pattern of discordance in geographical variation reflects a history of repeated episodes of isolation and dispersal among several evolving populations or groups of same. Adults of A. /econtei are similar to those of A. insolens in their physiological and behavioral ad- aptations for life in cooler, relatively faster-flow- ing streams. Perhaps the most striking features of A. /econtei adults are the broad elytral carinae. The functional significance of these carinae is yet unknown, but their dorsal position suggests that they may somehow contribute to stability during passive transport in stream currents. PROSPECTUS FOR FUTURE RESEARCH Clearly, much remains to be learned about ex- tant amphizoids and their evolutionary history. More information is needed about Amphizoa davidi—its geographical and habitat ranges, adult locomotory habits, and the form and structure of females. Because amphizoids are often diffi- cult to find, even in areas where they are known to occur, it is yet uncertain whether or not other species occur in eastern Asia. Concerted field- 107 work in this region, carried out by individuals familiar with the habits of Nearctic amphizoids, is required to resolve this question. Comparative morphological study of amphi- zoid larvae and those of other adephagan groups should provide valuable new data that can be used in tests of hypotheses of phylogenetic re- lationship among both amphizoid species and adephagan families. This potential source of data has gone largely untapped and much basic de- scriptive work on larvae is still lacking. In order to learn more about the historical development of amphizoids in space and time, search must continue among fossil materials of Mesozoic as well as Cenozoic age. To the best of my knowledge, amphizoids are not represented anywhere in the known fossil record, even during Quaternary time. Organisms living in lotic en- vironments are much less likely to be preserved as fossils than are their lentic equivalents, and this punctuates the notion that absence from the fossil record at any particular time does not pre- clude occurrence at that time. Clearly the search for additional fossil assemblages of appropriate age must be continued. ACKNOWLEDGMENTS On different occasions, George E. Ball, Terry L. Erwin, Jean Menier, and Rob E. Roughley each tried to locate type-material for 4. davidi Lucas on my behalf in MNHP and/or in other European collections, and I thank them for their efforts in this regard. Both J. Gordon Edwards and Rob Roughley provided helpful comments and insights, based on their extensive personal experience with amphizoid beetles, during var- ious phases of my study. The photograph of Am- phizoa davidi was taken by Susan Middleton, and Mary Anne Tenorio took the scanning electron micrograph. Access to the PHYLIP program package was made possible through J. Russo (Of- fice of Information Management, Smithsonian Institution). Alan Leviton provided access to the computer facility in the Department of Herpe- tology (CAS) and assisted me with use of the PHYLIP programs. LITERATURE CITED Batt, G. E. 1975. Pericaline Lebiini: notes on classification, a synopsis of the New World genera, and a revision of the genus Phloeoxena Chaudoir (Coleoptera: Carabidae). Quaest. Entomol. 1 1:143-242. 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 6 Bett, R. T. 1966. Trachypachus and the origin of the Hy- dradephaga. Coleopt. Bull. 20:107-112. 1967. 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U.S. Geol. Surv. Prof. Pap. 1106. iti + 37 pp. Wo re, J. A. AND E. B. LEopotp. 1967. Neogene and Early Quaternary vegetation of northwestern North America and northeastern Asia. Pp. 193-206 in The Bering land bridge, D. M. Hopkins, ed. Stanford University Press, Stanford. xill + 495 pp. Wricut, H. E. anp D. G. Frey, eps. 1965. The Quaternary of the United States. Princeton University Press, Princeton. x + 922 pp. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 cl — = _~_ i 253X NH PROCEEDINGS OF THE Vol. 44, No. 7, pp. 111-126, 14 figs., 1 table. CALIFORNIA ACADEMY OF SCIB ACANTHOGILIA, A NEW GENUS OF POLEMONIACEAE FROM BAJA CALIFORNIA, MEXICO By Alva G. Day and Reid Moran Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ApsTract: We propose the genus Acanthogilia for the spiny desert shrub first named Gilia gloriosa Bran- degee. The gametic chromosome number is nine, also the basic number for nine other genera and for the family. Acanthogilia is unique among Polemoniaceae in its extreme leaf dimorphism, its persistent woody-spinose primary leaves, and its coarsely verrucate zonocolporate pollen grains. Though formerly placed in Gilia, Ipomopsis, Leptodactylon, and Loeselia, it differs further from all these genera in its persistent secondary leaf bases with deciduous blades, its numerous closely spaced corolla veins connected at several levels, and its winged seeds. It does share several unusual characters, such as the superficially adnate filaments, with species of Gilia sect. Giliastrum. Acanthogilia seems closest to the Andean genus Cantua. Cantua, like Acanthogilia, is shrubby, with leaves dimorphic, on long shoots and axillary short shoots, with persistent leaf bases, with corolla veins connected at several levels, with winged seeds, with superficially adnate filaments (in some species), and with coarsely verrucate pollen grains (in one other species). Cantua differs in having the leaves broad and herbaceous, only weakly dimorphic, and neither woody-spinose nor with deciduous linear blades, the calyx entirely herbaceous, the pollen pantoporate, and the chromosome number hexaploid. INTRODUCTION Gilia gloriosa, of T. S. Brandegee (1889), is a spiny but truly gloriose desert shrub of rather local occurrence on the Pacific drainage of north- central Baja California (Fig. 1-3). This plant is seldom seen and little known, and its best generic position has remained uncertain. Brand (1907) placed it in Gilia sect. Leptodactylon, and Wher- ry (1945) called it Leptodactylon gloriosum. Johnston (1924) informally listed it as Loeselia gloriosa. Current floras (Wiggins 1964, 1980) treat it as Ipomopsis gloriosa, following Alva Grant (in V. Grant 1956). New information on the chromosome num- ber, pollen grain type, and some other aspects shows that Gilia gloriosa differs from all de- scribed genera of the family and has some unique characters. We therefore propose for it the fol- lowing new monotypic genus. SYSTEMATIC TREATMENT Acanthogilia Day et Moran, genus novum mexicanum Polemoniacearum, ob folia valde di- morpha, primariis rigide spinosis persistentibus, granaque pollinis zonocolporata supraverrucata bene distinctum; Cantuae Juss. fortasse proxi- mum, quae autem calyce toto herbaceo, aetate non rumpenti, pollinis granis pantoporatis, chro- mosomatumque numero polyploideo differt. Si vis descriptionem latine recipere, involucrum praeinscriptum praesolutumque mitte. Stiff spiny shrub with dimorphic leaves, the (111) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 DAY AND MORAN: ACANTHOGILIA, NEW GENUS OF POLEMONIACEAE 2mm Ficure 2. Acanthogilia gloriosa in flower and fruit. A. Flowering and fruiting branches on an older branch, with dehisced capsules from previous season; B. Green, submature seeds, contents of a single locule; C. Dry seeds from dehisced capsule; D. Calyx with mature, undehisced capsule; E. Calyx with dehisced capsule from previous season; F. Segment of branch with spinose primary leaves and fascicled herbaceous secondary leaves. a Ficure 1. Inflorescence of Acanthogilia gloriosa (Brandg.) Day and Moran, El Colosal, Baja California, Mexico, 13 June 1976. 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 | 30° *El Rosario e San Fernando Santa as Catarinae 4 * Cajiloab “San. Luis Gonzaga *Laguna Punta’ Moe + Chapala Gonads

>,, 5 pm ~@ 10 pm Ficure 4. Chromosomes of Acanthogilia gloriosa. Left, mitosis; right, meiosis. Traced from micrographs. Pollen grains 5-6 colporate (zonocolporate), suboblate to spheroidal; diameter 55-64 um x 61-71 um. Colpi short, only slightly longer than pores are broad. Pores lalongate to circular; diameter 5-7 um = 7-10 um. Exine 2.4-2.9 um thick; nexine 0.8-1.2 um thick, thickened up to 1.7 um in pore area, finely perreticulate. Lumina vari- able in shape and size; diameter less than 0.5 um to | um; muri supported by simple bacula densely spaced, COMPARISON OF THE GENERA OF POLEMONIACEAE. I. 9, 8 = intrageneric aneuploidy; 9/8 and 8/7 = dibasic polyploidy. II. Pollen groups 1-4 are the alliances of Taylor and Levin (1975), with Acanthogilia added. III. Pin = leaves pinnately veined, dissected, or lobed; PinC = leaves pinnately compound; Palm = leaves palmately lobed; * = true foliage leaves lacking. IV. N = seeds not winged; NW = seeds very narrowly winged; W = seeds broadly winged. V. A = filaments superficially adnate, M = filaments merged with corolla; I = filaments intermediate: merged below. VI. M = calyx membranous below sinuses; H = calyx herbaceous throughout. VII. A = veins connected at base of lobe and in upper lobe; B = veins connected only at base; C = veins connected only well above base; D = veins free; * = venation too simplified to classify. I Basic II VI Vil chromosome Pollen Ill IV Vv Calyx Venation of number group Leaf form Seed type Filaments type corolla lobe Acanthogilia 9 1 Pin Ww A M A Cantua 9 1 Pin A, M H A,B (2n = 27n) Huthia ? 1 Pin WwW I B Cobaea 9/8 2 PinC WwW A H A (2n = 26) Phlox 7 2 Pin N M M A,B Microsteris if 2 Pin N M M u Gymunosteris 6 2 a N M H * Polemonium 9 3 PinC N M, I H A,B Bonplandia 8/7? 3 Pin N, NW M H A (Qn = 15y)t Gilia 9,8 4 Pin N A, M, I M A, B, D Collomia 8 4 Pin N M H B,C Eriastrum il 4 Pin N M, I M B Navarretia 9 4 Pin N M M Bye Ipomopsis 7 4 Pin N M, I M B,D Langloisia 7 4 Pin N M, I M BC Allophyllum 9,8 4 Pin N M M Cc Loeselia 9 4 Pin N, NW M M GD Leptodactylon 9 4 Palm N M M D Linanthus 9 4 Palm N M M D + Bonplandia geminiflora chromosome number, 27 = 15 bivalents, determined from the following collections: Sinaloa, Mexico, D. E. Breedlove 44637, Chiapas, Mexico, D. E. Breedlove 56256. Vouchers deposited at CAS. Counted by A. Day. 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 sometimes merged 2-3 together. Diameter of bacula ca. 0.5 um. Reticulum supraverrucate. Verrucae on surface of exine very variable in shape and size, from very small and flat with diameter ca. 1 um to circular or oval with diameter to 10 um, in young pollen grains very densely spaced. Surface of verrucae ultra-finely striate or rugu- late, as seen with scanning electron microscope at 7000 x magnification. This diagnosis is based on a single collection (Day and Moran 76133, CAS). Other collections show that the grains may be 7-8-colporate (Mo- ran 7498, CAS) or the colpi may be longer (Fig. 6; Wiggins 5731, DS). Variation in the number and distribution of verrucae 1s seen by comparing Figures 5 and 6. The most distinctive feature of Acanthogilia pollen grains is the coarsely verrucate exine (Fig. 5-8). Among other Polemoniaceae with zono- colporate grains, only Eriastrum and Gilia sect. Giliastrum have the exine verrucate, but there the verrucae are minute. Dr. Stuchlik (pers. comm. 1980) remarked that Acanthogilia has probably a new pollen type for the family. Pollen grains with large verrucae do occur, however, in Cantua. In C. buxifolia Juss. ex Lam. (Fig. 9, 10) the exine appears much as in Acan- thogilia. In both C. buxifolia and Acanthogilia the verrucae are diverse in size and shape, the larger ones supported by groups of bacula. Viewed with SEM (Fig. 9, 10), the verrucae of C. bux- ifolia differ from those of Acanthogilia only in being somewhat broader and flatter. In the Cantueae (Cantua and Huthia) the exine is semitectate and, as illustrated (SEM) by Taylor and Levin (1975), generally consists of large, closely spaced areoles (Taylor and Levin’s term) or insulae (Stuchlik 1967). However, Cantua buxifolia is exceptional in having areoles of such small diameter that they have been described as large verrucae (Erdtman 1952). This exine pat- tern may have evolved through reduction of larg- er areoles. Despite the similarity in exine, the pollen grains of Cantua buxifolia differ from those of Acan- thogilia in aperture type; for, as in other Can- tueae, they are pantoporate, not zonocolporate. Since, however, both zonocolporate and panto- porate grains can occur within a single genus else- where (Collomia, Loeblich 1964, Chuang et al. 1978; Gilia, Stuchlik 1967), this difference be- tween Acanthogilia and Cantua is not necessarily fundamental. In view of other notable shared characters (Table 1), we interpret the similarity in exine as a mark of relationship. On the basis of pollen morphology, Taylor and Levin (1975: fig. 1) grouped the genera of Pole- moniaceae into four unnamed alliances (pollen groups 1-4 of our Table 1). One alliance included only Cantua and Huthia, but we would add Acanthogilia. Leaves.—In Acanthogilia the leaves of long shoots and axillary short shoots are markedly different, with no intermediates (Fig. 2F). The leaves of long shoots are woody-spinose and per- sistent, as in no other Polemoniaceae. Base and blade are scarcely delimited, and the blade is pinnately divided, with terete rachis and lobes. On the contrary, the fascicled axillary leaves are each clearly divided by a constriction into a per- sistent base and a deciduous blade (Fig. 11A). The broadened bases remain indefinitely in a compact spiral on the short shoot, but the blades fall at one time throughout the plant with drying of the season. These are smaller blades than those of the primary leaves, mostly simple, linear but flattened, herbaceous, and greener. Cantua, Huthia, Leptodactylon, Loeselia, and some species of Jpomopsis also have leaves on long shoots and in axillary fascicles; but although the fascicled leaves may be smaller, all leaves are nearly alike. Cantua is somewhat exceptional: the primary leaves are large and more or less lobed and fall early, whereas the secondary, ax- illary leaves are more persistent and in most species are smaller and have entire margins (In- fantes Vera 1962; Gibson 1967). In C. buxifolia grown in San Francisco, we note that, except for young shoots, leafy stems bear only the smaller secondary leaves. In various other Polemonia- ceae, especially annuals, leaves are gradually dif- ferent from base to apex, grading into bracts above. Only Acanthogilia, however, has mark- edly dimorphic leaves. In most perennial Polemoniaceae the leaves wither persistent, though they may finally erode away. In several evergreen shrubs (Cantua, Hu- thia, Loeselia mexicana (Lam.) Brand, L. pur- pusii Brandegee), however, leaf blades finally fall, leaving the persistent bases conspicuous (Fig. 11B). In Cantua buxifolia and C. pyrifolia Juss. ex Lam., both of which produce fascicled leaves, the short shoots and crowded leaf bases, with blades gone, somewhat resemble those of Acan- thogilia (Fig. 11A, B). DAY AND MORAN: ACANTHOGILIA, NEW GENUS OF POLEMONIACEAE 119 Ficures 5-10. Pollen grains of Acanthogilia gloriosa and Cantua buxifolia Juss. ex Lam.; Fig. 5, 6. (light microscope) Acanthogilia; Fig. 5. Day and Moran 76-133 (CAS); Fig. 6. Wiggins 5731 (DS); Fig. 7, 8. (SEM) Acanthogilia, Wiggins 5731 (DS); Fig. 9, 10. (SEM) Cantua buxifolia, cultivated, McClintock s.n., 15 Mar 1976 (CAS). Ss te Eee DOr bir a t Baar a 3 Sate eae eae 2 Oe oe ot RD Sone Ones FAP MOL or: x} i W988 5 VAS hie PHU. cf 2. 4s (@) Ficure 11. have fallen, the leaf bases from previous seasons compacted below, primary leaf mostly eroded away; B. Short shoot of Cantua buxifolia with crowded leaf bases, growing out into long shoot above; C. Calyx of Acanthogilia gloriosa at anthesis; D. Part of calyx, ventral side, showing venation in rib; E. Calyx of Cantua buxifolia at anthesis. Seeps.—The seed of Acanthogilia is flat and is bordered by a membranous wing 1-3 mm wide (Fig. 2C). In most Polemoniaceae, seeds are wingless, though in Bonplandia and Loeselia they PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 I\C JAN YA it Wy S/F HH WV, Wa 1% f ‘ i AE YM { [4 ¢ | WATE Efe fEoPEL Doe Hhodiy thee! Wot TE uy N AY et eta Lg a Nab bay q WU i i | mili i fy oe 5 Wf ZA & mm “*s, Ly fife ty Frapaeas” ra eo 28 Short shoots and calyces. A. Short shoot of Acanthogilia gloriosa with persistent leaf bases after most blades are sometimes very narrowly winged. Only in Cantua, Cobaea, and Huthia are the seeds like- wise flat and broadly winged. In these genera, however, both seeds and wings are considerably DAY AND MORAN: ACANTHOGILIA, NEW GENUS OF POLEMONIACEAE 121 wider; and the wings, though thin, are opaque rather than membranous. Acanthogilia has only 1-6 seeds per locule, whereas Cantua, Cobaea, and Huthia have many, in platelike layers. However, the ovary of 4can- thogilia has 14-24 ovules per locule, suggesting that the ancestral capsule may have had many more seeds. A hint of layering in the capsules suggests that if more seeds were present they might form layers as in the other genera. STAMENS.—The filaments of Acanthogilia are attached to the corolla tube about midway but are only superficially adnate below; they are well formed, with free margins, and are distinguish- able to the base. Likewise in Cohaea, Cantua (C. candelilla Brand and C. quercifolia Juss. but not C. buxifolia), and Gilia sect. Giliastrum (G. ri- gidula Benth. and G. ripleyi Barneby but not G. insignis (Brand) Cory and Parks or G. incisa Benth.) the filaments are superficially adnate. This appears to be a rare and primitive condition in the family. In our sampling of other genera, the filaments are so mérged with the corolla, at least basally and commonly to the point of insertion, that they are not distinguishable. CaLyx.—The calyx of Acanthogilia is narrow- ly membranous and veinless below the sinuses and has many veins crowded in the herbaceous ribs (Fig. 11C, D); it ruptures between ribs as the capsule grows. In eleven other genera (Table 1), including Gilia and most allies as well as Phlox and Microsteris, similarly, the calyx is narrowly to broadly membranous below the sinuses, with veins again confined to the ribs; it may or may not rupture in fruit. In all examples seen, veins are fewer and less crowded than in Acanthogilia. In all these genera, including Acanthogilia, lat- eral veins of adjacent ribs are connected only near the base of the calyx. On the other hand, in Bonplandia, Cantua (Fig. 11E), Cobaea, Collomia, Gymnosteris, Huthia, and Polemonium the calyx is not alternately ribbed and membranous but is herbaceous or somewhat chartaceous throughout, and it en- larges without rupturing as the capsule grows. Venation is various but generally is spread out more than in the genera with membranous calyx. Lateral veins of adjacent sectors may be con- nected just below the sinuses (Fig. 1 1E) or much lower. They are connected in Cantua quercifolia near the base of the calyx but in C. buxifolia at various levels, even in the same calyx (Fig. 1 1E). The herbaceous calyx type, found also in re- lated families, presumably is primitive in the Polemoniaceae, the membranous calyx perhaps arising independently in more than one line in arid habitats. A division of the family by calyx types then would separate some related genera. Thus Collomia (herbaceous calyx) belongs with the Gilia group (otherwise membranous), and Phlox and Microsteris (membranous calyx) seem related to genera with herbaceous calyx (Table 1). Similarly, Acanthogilia appears related to Cantua despite the difference in calyx (Table 1). CoROLLA VENATION.—Surveying corolla ve- nation in the family, Day has found patterns to link Acanthogilia with some genera and to sep- arate it from others. Generally in the family, each sector of the corolla has a median vein and two laterals more or less parallel in the tube, with branches in the lobes and commonly with con- nections; positions of vein connections are char- acteristic for many taxa. Figures 12-14 show ex- amples from 12 out of 19 genera—all traced from photographs of dissected corollas stained with safranin. The staminal veins, alternating with the corolla veins in the tube, are omitted. Venation patterns fall mainly into four types (identified in Table 1 by letters A-D): A. Cantua type—veins connected near corolla orifice, curv- ing and connected once or twice above in the lobe (Fig. 12G-H, 13K-O); B. Gilia type—veins connected near orifice but straighter above and without other connections (Fig. 1 2B—D); C. Loe- selia type—veins often connected near middle or apex of lobe but not near orifice (Fig. 14P-R); and D. Leptodactylon type—veins free, not con- nected in orifice or lobes, even in large corollas with many veins; often each sector with only a single vein in basal half of tube (Fig. 12E-F, 14S-U). Some genera show only one venation type, some two, and one three (Table 1). In very small corollas (Microsteris, Fig. 13J; many Na- varretia species, Fig. 12A; Gymnosteris; and oc- casional species in other genera) venation may be so simplified that it tells little of relationship. In Acanthogilia the corolla veins are connected at several levels in the lobes (Fig. 12H), much as in Cantua, Cobaea, and Phlox (Fig. 13). Acan- thogilia differs from them in having more closely spaced veins that are nearly parallel and less curved. Its pattern is perhaps most closely ap- co t nae bar=I mm PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 Ficure 12. Corolla venation patterns in Polemoniaceae, each showing a sector from base of tube to apex of lobe. Dashed lines show where corolla tube cut. Stamens and staminal veins not shown. A. Navarretia fossalis Moran; B. N. mitracarpa Greene; C. Gilia tricolor Benth.; D. G. leptomeria Gray; E. G. incisa Benth.; F. G. rigidula Benth.; G. G. ripleyi Barneby; H', H”. Acanthogilia gloriosa Day and Moran. proached in Cantua candelilla (Fig. 13M). On the other hand, despite more numerous veins with connections at several levels, the pattern of Acanthogilia resembles that of Gilia and allies (Fig. 12B-—D) in its straighter and closer-spaced veins. And although most species of Gilia sect. Giliastrum have free veins (Fig. 12E-F), the anomalous G. rip/eyi (Fig. 12G) has connections DAY AND MORAN: ACANTHOGILIA, NEW GENUS OF POLEMONIACEAE 123 bar=I!mm Ficure 13. Corolla venation patterns, cont. J. Microsteris gracilis (Hook.) Greene; K. Cobaea biaurita Standl.; L. Cantua pyrifolia Juss. ex Lam.; M’, M”. C. candelilla Brand; N. Phlox andicola Nutt. ex Gray, O. Bonplandia geminiflora Cav. at several levels, thus somewhat approaching Acanthogilia. Although the species gloriosa has been placed in Gilia, Ipomopsis, Leptodactylon, and Loese- lia, each of these genera has a venation pattern different from that of Acanthogilia. The distinc- tive patterns of Leptodactylon and Loeselia es- pecially seem to make close relationship with Acanthogilia unlikely. Woop ANATOMyY.—Carlquist et al. (1984) studied the wood anatomy of the Polemoniaceae, comparing the relatively few woody species. Be- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 Ficure 14. Corolla venation patterns, cont. P. Loeselia greggii S. Wats., Q. L. amplectens (Hook. and Arn.) Benth.; R. Allophyllum glutinosum (Benth.) A. and V. Grant; S. Linanthus dianthiflorus (Benth.) Greene; T. L. grandiflorus (Benth.) Greene; U. Leptodactylon pungens (Torr.) Rydb. DAY AND MORAN: ACANTHOGILIA, NEW GENUS OF POLEMONIACEAE 125 sides Acanthogilia, these species fall in Cantua and Huthia (Cantueae), Cobaea (Cobaeeae), and Eriastrum, Ipomopsis, and Leptodactylon (Gi- lieae). In general, they thought the wood anato- my probably more closely correlated with growth form as related to habitat, than with systematic relationships. The authors noted that Acanthogilia and a species of Jpomopsis are alike in having banded axile parenchyma—unusual in the family but oc- curring also, in its most incipient form, in Can- tua. Likewise, in the imperforate tracheary ele- ments and in the vascular rays, Acanthogilia is similar to Jpomopsis on the one hand and to the Cantueae on the other. The coincidence in these characters among Acanthogilia, the Gilieae, and the Cantueae somewhat parallels other similar- ities we report here (Table 1) and would seem to be due to relationships rather than to environ- mental factors alone. RELATIONSHIPS Acanthogilia is unique among Polemoniaceae in its extreme leaf dimorphism, its persistent woody-spinose primary leaves, and its coarsely verrucate zonocolporate pollen grains. Although A. gloriosa, the sole species, has been placed in Gilia, Ipomopsis, Leptodactylon, and Loeselia, it differs further from all these genera in its persis- tent secondary leaf bases with deciduous blades, its numerous closely spaced corolla veins with interconnections at several levels, and its winged seeds, and from all these except for two species of Gilia in its superficially adnate filaments. It differs still further from Gilia and Ipomopsis in its large shrubby habit, from Jpomopsis in its basic chromosome number of nine, and from Leptodactylon in its pinnate leaves and its three corolla veins instead of one in each sector of the lower tube. Among North American Polemoniaceae, Acanthogilia seems to have most in common with Gilia and allies, and especially with species of Gilia sect. Giliastrum. As in Acanthogilia, all Gilia species have the calyx membranous below the sinuses, and most, including sect. Giliastrum, have zonocolporate pollen and have the primi- tive x = 9. In this polymorphic genus of five sec- tions, usually the pollen is blue and the exine reticulate to striate and not verrucate. In sect. Giliastrum, however, the pollen is yellow as in Acanthogilia, and the exine is somewhat similar, being pertectate and minutely verrucate whereas in Acanthogilia it is perreticulate and coarsely verrucate. Although in most species of Gilia the filaments merge with the corolla below, in G. ripleyi and G. rigidula, of sect. Giliastrum, the filaments are superficially adnate, as in Acantho- gilia. Most species of Gilia are annual and none are truly woody, but G. rip/eyi is a suffrutescent perennial. Finally, although most species of sect. Giliastrum have free corolla veins, G. ripleyi is unique in Gil/ia and further resembles Acantho- gilia in having the veins connected at several levels. Acanthogilia is perhaps most closely related to the Andean genus Cantua. Cantua, like Acan- thogilia, is shrubby, with leaves dimorphic, borne on long shoots and axillary short shoots, with crowded leaf bases remaining on the short shoots after the blades have fallen, with corolla veins connected at more than one level, with seeds flattened and broadly winged, with a basic chro- mosome number of nine, with superficially ad- nate filaments in C. candelilla and C. quercifolia, and with coarsely verrucate pollen in C. buxi- folia. The lower branches of C. buxifolia (grown in San Francisco) take root, as do those of Acan- thogilia. Cantua differs in that the leaves are broadly herbaceous and only slightly dimorphic, with primary leaves deciduous, not at all woody- persistent, and secondary leaves more persistent; the calyx herbaceous, not membranous below the sinuses, and not rupturing in age; the pollen pan- toporate, not zonocolporate; the chromosome number hexaploid, not diploid. We suggest that Acanthogilia may be a specialized desert descen- dent of a diploid line also ancestral to Cantua. Since Cantua is hexaploid, however, and prob- ably amphiploid, such divergent characters as the herbaceous calyx may perhaps derive from some other line. Grant (1959) divided the Polemoniaceae into five tribes. Acanthogilia probably belongs to the Cantueae but apparently has some relationship also with the Gilieae. Much new evidence bear- ing on generic relationships has accumulated, es- pecially from pollen studies, since Grant’s clas- sification, and the time seems ripe fora new tribal arrangement. ACKNOWLEDGMENTS We are grateful to all who have helped with this paper, and especially the following: Dr. Leon 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 7 Stuchlik of the Botanical Institute, Cracow, Po- land, for his pollen diagnosis of Acanthogilia and observations concerning it; Jerry Morgan of Uni- versity of California, San Francisco, for the SEM micrographs; the Gift Fund, Dudley Herbarium of Stanford University for support for the SEM work; Terry Bell for line drawings of Acantho- gilia; Colleen Sudekum for the corolla venation tracings; Barbara Stewart for drawing the distri- bution map; and Dave and Dru Binney for send- ing two points for the map, from far, out-of-the- way places. We appreciate suggestions given by Dr. Robert Patterson on the pollen morphology section, by John Thomas Howell on the Latin diagnosis, and by Dr. Dennis Breedlove and Dr. Leslie Landrum on various points along the way. An early preview of the wood-anatomy manu- script was kindly offered by the senior author, Dr. Sherwin Carlquist. We thank Dr. Frank Al- meda, Dr. Christopher Davidson, Dr. Verne Grant, and two unsigned reviewers for reading this manuscript and making many helpful com- ments. We appreciate the courtesies extended at the following herbaria: California Academy of Sciences (CAS), Dudley Herbarium of Stanford University (DS), San Diego Museum of Natural History (SD), and University of California, Berkeley (UC). Lastly, for making the color plate possible, we are most grateful to seventy-two col- leagues and friends enlisted by Annetta Carter and Lincoln Constance as Los Amigos de Acan- thogilia. LITERATURE CITED Branp, A. 1907. Polemoniaceae. A. Engler, Das Pflanzen- reich 4(250). Branpecee, T. S. 1889. A collection of plants from Baja California, 1889. Proc. Calif. Acad. Sci. Ser. 2, 2:117-216. Car.ouist, S., V. M. ECKHART, AND D. C. MICHENER. 1984. Wood anatomy of Polemoniaceae. Aliso 10(4):547-572. CHUANG, T., W. C. HstEH, AND D. H. WiLKEN. 1978. Con- tribution of pollen morphology to systematics in Collomia (Polemoniaceae). Am. J. Bot. 65:450-458. ErptMAN, G. 1952. Pollen morphology and plant taxonomy: angiosperms. The Chronica Botanica Co., Waltham, Mas- sachusetts. 539 pp. Gipson, D. N. 1967. Polemoniaceae. /n Flora of Peru, Mac- bride, ed. Field Mus. Nat. Hist., Bot. Ser. 13(5a:2):112-131. Grant, V. 1956. A synopsis of Jpomopsis. Aliso 3:351-362. 1959. Natural history of the Phlox family: 1. Sys- tematic botany. Martinus Nijhoff, The Hague, Netherlands. Grant, V. AND K. A. Grant. 1965. Flower pollination in the Phlox family. Columbia Univ. Press, New York. INFANTES VERA, J. G. 1962. Revisién del género Cantua (Polemoniaceae). Lilloa 31:75-107. Jounston, I.M. 1924. Expedition of the California Academy of Sciences to the Gulf of California in 1921: the Botany (the vascular plants). Proc. Calif. Acad. Sci. Ser. 4, 12:951- 1218. Loesuicn, A. R. 1964. The pollen grain morphology of Col- lomia as a taxonomic tool. Madrono 17:205-216. Martinez, M. 1937. Catalogo de nombres vulgares y cien- tificos de plantas Mexicanas. Ediciones Botas, México. Moran, R. 1952. The Mexican itineraries of T. S. Brande- gee. Madrono 11:253-262. Stucuiik, L. 1967. Pollen morphology and taxonomy in the Polemoniaceae. Grana Palynol. 7:146-240. Taytor, T. N. AND D. A. Levin. 1975. Pollen morphology of Polemoniaceae in relation to systematics and pollination systems: scanning electron microscopy. Grana 15:91-112. Wuerry, E. T. 1945. Two Linanthoid genera. Am. Midl. Naturalist 34:38 1-387. Wiaains, I. L. 1964. Flora of the Sonoran Desert. Pp. 189- 1740 in Vegetation and flora of the Sonoran Desert, Shreve and Wiggins, eds., 2 vols. Stanford Univ. Press, Stanford, California. . 1980. Flora of Baja California. Stanford Univ. Press, Stanford, California. 1025 pp. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 @ ii = a $ PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 8, pp. 127-156, 106 figs., 2 tables. May 6, 1986 THE DIATOM GENUS THALASSIOSIRA: SPECIES FROM, THE SAN FRANCISCO BAY SYSTEDEN 7 f By | A. D. Mahood Department of Geology, California Academy of Sciences, Golden Gate Park, San Fragcisco, Ci alifornia, RES = % ace, ae G. A. Fryxell Department of Oceanography, Texas A & M University, College Station, Texas 77843 M. McMillan Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602 ABSTRACT: Twenty species of the diatom genus Thalassiosira, including one previously undescribed species, were collected from diverse habitats of San Francisco Bay, California, extending from nearly freshwater in Suisun Bay to marine salinity near the Golden Gate. In this paper the morphology of these 20 species is elucidated by scanning electron microscopy (SEM) and light microscopy (LM). The species range from large taxa with linear areolae to small, lightly silicified forms with eccentrically arranged areolae. The basic form of the genus is seen as a moderately small diatom with rectangular outline in girdle view and a simple process pattern of one central and one marginal ring of strutted processes and a single labiate process. The distribution of species is influenced by salinity. Major species show limited distributions: 7. visurgis Hustedt, fresh to brackish water; 7. decipiens (Grunow) Jorgensen, brackish water; 7. nodulolineata (Hendey) Hasle and G. Fryxell, tidal marine to brackish; 7. hendeyi Hasle and G. Fryxell, and 7. wongii Mahood sp. nov., tidal marine water; 7. nordenskioeldii Cleve and T. pacifica Gran and Angst, coastal marine. These distributions demonstrate the value of species as indicators of salinity patterns within the bay ecosystem. INTRODUCTION oe : : ‘ their interactions begun at the University of Cal- The San Francisco Bay system has been stud- ied by many investigators since 1816, when the Russian ship Rurik anchored in the bay (Hedg- peth 1979). The bay estuarine system extends from the mouth of the Guadalupe River in the south to the lower reaches of the Sacramento- San Joaquin delta near the city of Pittsburg (Fig. 1) in the north. Early studies of the bay concen- trated on hydrology, fisheries, and physical pa- rameters of the system. Not until the early 1920s was a serious effort to study the bay’s biota and ifornia, Berkeley (Hedgepeth 1979). The phy- toplankton flora was not examined until 1939, when F. W. Whedon, using a Sedgewick-Rafter chamber, made a limited study of San Francisco Bay phytoplankton and presented a brief species list, including Thalassiosira rotula Meunier. The flora of San Francisco Bay remained large- ly unstudied until 1958, when the Sanitary En- gineering Research Laboratory (SERL) of the University of California, Berkeley, began a mul- tidisciplinary study of the bay (Harris et al. 1961; [127] 128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 Taste 1. Key To CoLLection Sites, DATES OF COLLECTION, AND COLLECTORS. Date Collector Location 3 September 1980 R. L. J. Wong 1 16 October 1980 R. L. J. Wong 2 2 May 1971 A. D. Mahood 3 6 July 1981 G. A. Fryxell/ 3 A. D. Mahood 6 June 1971 A. D. Mahood 4 20 February 1972 Cc. A. McNeil 4 6 July 1981 A. D. Mahood 5 24 April 1971 A. Hauser 6 June 1972 P. Wares 6 1 November 1979 L. L. Lack 7 3 September 1980 R. L. J. Wong 8 13 November 1980 R. L. J. Wong 9 12 September 1979 A. D. Mahood 10 7 July 1981 A. D. Mahood 10 Storrs et al. 1966). Although the species list gen- erated by SERL was established from raw ma- terial (i.e., not cleaned of organic material so that the siliceous parts can be clearly seen) and was thus of limited systematic value, it is one of the few comprehensive references to the San Fran- cisco Bay diatoms available. Hasle (1978a, b) published the first careful taxonomic papers on Thalassiosira from the bay system. The most recent inventory of bay diatom species (Wong and Cloern 1981), though still in progress, offers a basis for future diatom floristics and phyto- plankton ecology studies of the bay. Thalassiosira Cleve, one of the dominant dia- tom genera in the bay’s phytoplankton, has been frequently mentioned in species lists: T. decip- iens (Grunow) Jorgensen; T. eccentrica (Ehren- berg) Cleve; 7. /acustris (Grunow) Hasle (=Cos- cinodiscus lacustris Hustedt); T. nordenskioeldii Cleve; T. punctigera (Castracane) Hasle (=T. angstii (Gran) Makarova); and 7. rotula Meu- nier. The diversity of species of the genus in the bay system is not surprising because the Tha- lassiosira probably evolved in a similar estuarine and coastal environment (Round and Sims 1981). The species diversity of diatoms is limited by salinity variations within the bay system (pers. comm. Wong, United States Geological Survey, 1983). The use of the genus Thalassiosira as an indicator of marine influence in the bay system has been hindered by several conditions: the con- fusion with other centric genera (e.g., Coscino- discus), difficulty in identification (especially in water mounts), problems connected with sample preparation (including the need for mounting in a medium with a high refractive index for light microscopy), and widely scattered pertinent taxonomic literature. In this paper we report on the distribution of Thalassiosira species in the bay and discuss mor- phology and its application to systematic prob- lems within the bay. METHODS AND MATERIALS Samples were collected with a Kemmerer water bottle and net over a broad range of the navigable section of the San Francisco Bay estuarine system (Fig. 1). Net hauls are particularly useful in con- centrating material for identification before quantitative work is attempted on water sam- ples. Fixed samples were examined in Utermohl settling chambers using an Olympus IMT in- verted microscope. Most often positive identi- fication must be made from cleaned material, mounted in medium of high refractive index such as Hyrax (Hanna 1930). Material was cleaned using the Van der Werff (1955) hydrogen per- oxide method. Strewn and arranged slides were prepared for light microscope (LM) from cleaned materials and mounted in Hyrax. Similar strewn and also arranged material (mounted by the se- nior author) was prepared on scanning electron microscope (SEM) stubs for examination and photography using Jeolco JSM-35, Texas A&M University Electron Microscopy Center. Texas A&M Department of Oceanography phyto- plankton cultures F190, F206, F209, and F226 were isolated by T. P. Watkins from collections made about a mile north of the Golden Gate Bridge, 6 July 1981. They were maintained in the Department of Oceanography phytoplankton culture collection: F/2 medium, 30% salinity, 16 hours light and 8 hours dark cycle, in 16°C growth chambers. Terminology follows that of the work- ing Party on Diatom Terminology (Anonymous 1975; Ross et al. 1979). RESULTS AND DISCUSSION Twenty species of Thalassiosira were re- covered from 33 samples from San Francisco Bay: T. anguste-lineata (A. Schmidt) G. Fryxell and Hasle; T. decipiens (Grunow) Jorgensen; T. eccentrica (Ehrenberg) Cleve; T. endoseriata Ha- sle and G. Fryxell; T. hendeyi Hasle and G. Fryxell; 7. incerta Makarova; T. lacustris (Gru- now) Hasle; 7. Jundiana G. Fryxell; T. minuscula MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA 129 tenes l «| F 122°30' ih aS 122°15' if \22°00° \ (\ y Z 2 . ~ SG VA : A NN \ ) No alee San Pablo Bay ® Carquinez x@) Strait 5 38°00'— ee ° 5 10 20 MILES © Central Bay ! ee ® ° 5 10 20 30 KILOMETERS 3 Golden alle Ys Gate V B 37°45'— Pacific : Ocean NORTH South Bay 37°30: — pam aos \ — saute River ie eS Ficure 1. San Francisco Bay System. 130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 Krasske; T. nodulolineata (Hendey) Hasle and G. Fryxell; T. nordenskioeldii Cleve; T. oestrupii var. venrickae G. Fryxell and Hasle; T. pacifica Gran and Angst; 7. punctigera (Castracane) Ha- sle; T. rotula Meunier; T. simonsenii Hasle and G. Fryxell; 7. stellaris Hasle and Guillard; T-. tenera Proschkina-Lavrenko; 7. visurgis Hus- tedt; and 7. wongii Mahood sp. nov. An understanding of the individual species, their similarities and differences, the habitats in which they occur, and their distributions within the bay system is essential to understanding the phytoplankton dynamics of the bay and the role of the Thalassiosira species in the bay ecosystem. The genus is characterized by a number of morphological features: one to a few labiate pro- cesses (Fig. 16); many strutted processes (Fig. 11); none to many occluded processes (Fig. 38); eccentric, linear, or fasciculated patterns of the rows of areolae, in a basically radial pattern; and the placement of the areola cribrum on the in- ternal side of the loculate areolae and the fora- men on the external side (Fig. 14, 17). For this paper and to assist the light microscopist, we have stressed the characters particularly visible in LM: number and location of labiate processes; number and arrangement of the areolae; and lo- cation and number of strutted processes. Several species can be identified under LM, whereas oth- ers require SEM in order to resolve definitive characters. Because our primary purpose includes gaining a greater understanding of San Francisco Bay, an essential goal is to distinguish between marker species with similar morphological characteris- tics but differing distributions, species which have been confused in earlier literature. For such com- parisons, we have separated the Thalassiosira species in this paper into five morphological groups: 1) species with basic linear areola pat- terns, 2) species with eccentric areola patterns, 3) species with one central strutted process and one marginal ring of strutted processes, 4) species with no central strutted processes and modified rings of strutted processes, and 5) two otherwise dissimilar species that have radial areola patterns and multiple central strutted processes. These are form groupings only; they are not placed together to imply close taxonomic relationships. Group 1.—Thalassiosira species with linear ar- eola patterns. Thalassiosira simonsenii Hasle and G. Fryxell, 1977 (Figures 2-5) DetAILeD Description. Hasle and Fryxell (1977). —Cell di- ameter 30-57 um; areolae 45.5 in 10 um across the valve, 8- 10 in 10 um at mantle edge (Fig. 2, 4); one small central strutted process (Fig. 5); two rows of alternating strutted processes on margin, five to six in 10 um (Fig. 3); two opposing labiate processes (Fig. 2); distinctive marginal ribs, eight in 10 um; large tubular occluded processes, one in 10 um on margin above strutted processes (Fig. 4). Marginal ribs distinctive in SEM, but not always clear in LM. DisTRIBUTION. — Marine, found only in central San Francisco Bay in association with other ma- rine diatoms. Observed previously at 28°00'N, 112°17.5'W, Pacific (Hasle and Fryxell 1977). Thalassiosira hendeyi Hasle and G. Fryxell, 1977 (Figures 6-11, 86) DetatLep Description. Hasle and Fryxell (1977).—Cell di- ameter 38-120 um; areolae, regularly linear, five to six 10 um; prominent central strutted process (Fig. 6, 8) set to one side of central areola; two closely adjacent rings of marginal strutted processes (Fig. 9) alternating in orientation (Fig. 11, internal view), not easily resolved with LM (Fig. 86); wavy mantle ridge (Fig. 9, 86); two labiate processes (Fig. 6, external [arrow]; Fig. 10, internal); labiate process with two adjacent strutted pro- cesses (Fig. 7); valve slightly concentrically undulated (Fig. 6). DisTRIBUTION.—Common but never abun- dant from San Pablo Bay to south San Francisco Bay. Previously found in warm coastal waters of West Africa (Hasle and Fryxell 1977), Uruguay and Brazil (Muller-Melchers 1953). Thalassiosira nodulolineata (Hendey) Hasle and G. Fryxell, 1977 (Figures 12-17, 87, 90) DetarLep Description. Hasle and Fryxell (1977).—Cell di- ameter 26-58 um; areolae linear, regular, 3.5 (one with 9.0) in 10 um (Fig. 12); four strutted processes in 10 um with spines (Fig. 12, 15) along the margin; five to six strutted processes inside the central areola (Fig. 14, external; Fig. 17, internal; Fig. 90); single labiate process at valve margin (Fig. 12, 16). In our samples, central areolae surrounded by six symmetrical areolae (Fig. 90), although central areolae surrounded by five areolae have been reported (Hasle and Fryxell 1977). DisTRIBUTION.—Central San Francisco Bay, San Pablo Bay, and Suisun Slough; common but never in large numbers. Thalassiosira tenera Proschkina-Lavrenko, 1961 (Figures 18-23, 103-104) DeraiLepD Description. Hasle and Fryxell (1977).—Cell di- ameter 10-29 um; areolae 9-16 in 10 um; marginal strutted MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA Ficures 2-5. processes three to five in 10 um, one central strutted process (Fig. 18-20, 23); one labiate process (Fig. 20, 22, arrow); ar- rangement of areolae linear (Fig. 23, 103, 104) or fasciculated; marginal strutted processes often with siliceous overgrowths and flattened (Fig. 18, 21, 23). DisTRIBUTION.— Restricted to coastal waters (Hasle and Fryxell 1977). Material examined from San Francisco Bay entirely prepared from 131 Thalassiosira simonsenii Hasle and G. Fryxell. SEM. Figure 2. Scale = 5 um, external view of valve, two labiate processes across valve from each other (arrow). Figure 3. Scale = 2 um, two rows of alternating strutted processes on margin. Figure 4. Scale = 1 um, large tubular occluded process on margin above strutted processes. Figure 5. Scale = 1 um, view of central strutted process. cultures from samples taken just north of the Golden Gate (6 July 1981). Discussion.—T. hendeyi, T. simonsenii, and T. nodulolineata have been confused in bay stud- ies and erroneously reported as Coscinodiscus lineatus Ehrenberg (=T. /eptopus [Grunow] Ha- sle and G. Fryxell). In our samples, 7. nodulo- lineata specimens were most easily distinguished 132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 from the other two by having a single labiate process and six symmetrical areolae surrounding the central areola (Fig. 14, 17). The five to six strutted processes inside the central areola fur- ther confirm the identification. Although 7. fe- nera and T. incerta Makarova (to be discussed in Group 5) have a similar appearance to 7. nod- ulolineata, the coarser areolae of T. nodulolineata distinguish it from the two with finer areolae. In addition, the marginal strutted processes of T. tenera often are in heavily silicified areas of the margin and are markedly distinct in LM, in con- trast to other species. Distinguishing specimens of 7. hendeyi and T. simonsenii is extremely difficult under LM. Ex- amination of SEM micrographs of the margin of T. hendeyi (Fig. 6) and the central area (Fig. 8) will assist the light microscopist in differentiating these two species. Both species have two labiate processes and similar central areas. For identi- fication under the LM (Fig. 86), careful exami- nation of the margin is necessary. Under the SEM (Fig. 6, 9), the margin of 7. hendeyi is irregular and wavy, but with the light microscope a clean break can be noticed when focusing from the margin to the valve face. The valve face forms a rather sharp angle with the margin in 7. hen- deyi, while in T. simonsenii (Fig. 4) the transition from valve face to margin is smooth. Although the marginal strutted processes of 7. simonsenii are more prominent than those of 7. hendeyi, this characteristic is not easily resolved under LM. In our samples the salinity ranges for T. hen- deyi and T. nodulolineata overlap, and both are found from south San Francisco Bay to Suisun Bay, indicating a more brackish environment. Thalassiosira simonsenii, much less common than the others, was confined to the central San Francisco Bay and was associated with other ma- rine diatoms. Thalassiosira tenera was collected near the Golden Gate Bridge in a marine habitat. Thalassiosira tenera is the only one of this group with strutted processes in the middle of the cen- tral areola. Group 2.—Thalassiosira species with eccentric areola patterns. Thalassiosira wongii Mahood, sp. nov. (Figures 24-29, 99-101) DetalLeD Description. Mahood.— Diameter 27-51 «um; ar- eolae radial, fasciculated, 9-11 in 10 um with areolar rows parallel to central areolar row of fascicle; strutted processes at margin three to five in 10 um (Fig. 24-26, 29, 99-101); four to five strutted processes observed around central areola (Fig. 25, 26); strutted processes usually forming two irregular rings between central areola and margin (Fig. 26, 99, 100); one mar- ginal ring of strutted processes near margin, four to five in 10 um (Fig. 29); one labiate process set slightly inside marginal ring (Fig. 27, internal and external, Fig. 28, 99, 100), three spines in 10 um around margin in same ring as labiate process (Fig. 24); small spines often above each strutted process on outside of valve (Fig. 27, internal and external; Fig. 29). Diameter 27-51 um; areolae 9-11 in 10 um; fultoportulae ad marginem, 3-5 in 10 um; 4-5 fultoportulae circum areolam centralem visae; fultoportulae plus minusve annulos duos ir- regulares formantes inter areolam centralem et marginem; an- nulus marginalis unicus fultoportularum prope marginem 4-5 Ficures 6-11. > Thalassiosira hendeyi Hasle and G. Fryxell. SEM. Figure 6. Scale = 10 um; external view of valve; two labiate processes (arrow), valve slightly concentrically undulated, distinctive marginal ridge, linear areolae. Figure 7. Scale = 1 um, labiate process with two adjacent processes. Figure 8. Scale = 1 um, prominent central process. Figure 9. Scale = 1 um; wavy marginal ridge, two closely adjacent rows of strutted processes. Figure 10. Scale = 10 um, internal view of valve, alternating marginal strutted processes. Figure 11. Scale = 1 um, internal view of valve, alternating marginal strutted processes. Ficures 12-17. Thalassiosira nodulolineata (Hendey) Hasle and G. Fryxell. SEM. Figure 12. Scale = 5 um; external view of valve; linear areolae, marginal strutted processes (sp), marginal spines (ms), single labiate process (Ip). Figure 13. Scale = 1 um, single labiate process, bands. Figure 14. Scale = 1 um; strutted processes in the central areola, six symmetrical areolae surrounding central areola with radial threads. Figure 15. Scale = 0.5 um, external marginal spine, strutted processes. Figure 16. Scale = 1 um, internal view of valve, internal labia with external labiate process (arrow). Figure 17. Scale = 1 um, internal view of central strutted processes. Ficures 18-23. Thalassiosira tenera Proschkina-Lavrenko. SEM. Figure 18. Scale = 2 um; external view of valve; marginal strutted processes, one central strutted process. Figure 19. Scale = 1 um, distinctive central strutted process. Figure 20. Scale = 2 um, internal view of valve, one central strutted process. Figure 21. Scale = 1 um, external view of marginal strutted process with siliceous overgrowths. Figure 22. Scale = 0.5 um, internal view of labia (arrow). Figure 23. Scale = 2 um, external view of valve; single external labiate process (arrow), marginal strutted processes with siliceous overgrowths. MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 x eS a 9 a wy = J aay S MAHOOD, FRYXELL, AND McMILLAN: PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 | Ficures 24-29. Thalassiosira wongii Mahood, sp. nov. SEM. Figure 24. Scale = 10 um; external view of valve; fasciculated areolae, small marginal spines, single labiate process (arrow). Figure 25. Scale = 1 um, external view of irregularly arranged processes surrounding the central areola. Figure 26. Scale = 10 um; internal view of valve; single labia, two irregular rings of strutted processes, marginal ring of strutted processes. Figure 27. Scale = 2 um; internal view of labia and external labiate MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA in 10 wm; rimportula unica annuli marginalis parum penitus posita, spinis 3 in 10 um circum margineum in eodem annulo cum rimoportula; spinae parvae saepe supra fultoportulam omnem extus valvarum. DIsTRIBUTION.—South and central San Fran- cisco Bay, associated with marine diatoms. Ho.otyre.—Slide, deposited at California Academy of Sciences, CAS 61243. Thalassiosira oestrupii var. venrickae G. Fryxell and Hasle, 1980 (Not figured) DetarLeD Description. Fryxell and Hasle (1980).—Cell di- ameter 5.5—39 um; areolae 6-9 in 10 um in central area, 7-11 in 10 wm toward margin; one labiate process, usually located three areolae from center; one central strutted process, one to two in 10 um, on margin. May be confused with 7. decipiens, but marginal strutted processes internal rather than external. Dominant characteristics: labiate process away from margin and marginal strutted processes with internal projection seen in same plane of focus (Fryxell and Hasle 1980, fig. 15A, B, 17). DisTRIBUTION. —Coastal, temperate waters. Central San Francisco Bay, rare. Thalassiosira eccentrica (Ehrenberg) Cleve, 1904 (Figures 30-35, 102) DerAILeD DescriPTION. Fryxell and Hasle (1972).—Cell di- ameter 12-101; areolae 5—8 in 10 um in central area, 7-10 in 10 um toward margin (Fig. 102); scattered strutted processes across valve face (Fig. 32); irregular spines around margin, three to four in 10 um; central areola surrounded by seven areolae with single strutted process next to the central areola (Fig. 30, 33); two to three rings of strutted processes in 10 um near margin (Fig. 31; Hasle 1979); one prominent labiate pro- cess (Fig. 34, internal; Fig. 35). Valve face relatively flat. DisTRIBUTION. — Possibly brackish to definite- ly saline waters, common in central San Fran- cisco Bay. Discussion. — The confusion within this group stems in part from the areola patterns of T. ec- centrica (Fryxell and Hasle 1972), which varies from the classic eccentric pattern to a fascicu- lated arrangement. For example Cupp (1943), a primary reference source for Bay studies, does not clearly distinguish 7. decipiens (to be dis- cussed in Group 3) and 7. eccentrica (=Cosci- nodiscus eccentricus). The overall eccentric pat- tern is representative of three species in this group. 137 Other morphological characteristics may be used to facilitate identification. Thalassiosira oestru- pli var. venrickae lacks external labiate process tubes. Only 7. wongii has multiple central pro- cesses, fasciculated valves, and spines above each strutted process. A single row of marginal strut- ted processes further distinguishes 7. wongii from T. eccentrica. The spines above each strutted process of 7. wongii are only seen under SEM. Group 3.—Thalassiosira species with one cen- tral and one marginal ring of strutted processes and one labiate process near the margin. Thalassiosira minuscula Krasske, 1941 (Figures 36, 37) DetaiLeD Description. Hasle (1976).—Cell diameter 10- 20 um; areolae small, 30 in 10 um in rows parallel to radial row; one large radially oriented labiate process located in from margin (Fig. 36); strutted processes in ring on margin four to five in 10 um near margin, one strutted process in center (Fig. 36), plus one adjacent to labiate process (Fig. 37). Distin- guished from other fasciculated species of Tha/assiosira in San Francisco Bay by the unique arrangement of a strutted process adjacent to the labiate process and by the finer areolae. DisTRIBUTION. — Originally described from coastal plankton of Chile (Krasske 1941). Central San Francisco Bay, rare. Thalassiosira lundiana G. Fryxell, 1975a (Figures 36-41, 89) DerarLep Description. Fryxell (1975a).—Cell diameter 7— 43 um; areolae 24-30 in 10 um, fasciculated; marginal striae; strutted processes in ring inside margin (Fig. 38), approxi- mately 10 in 10 um; often irregularly arranged large occluded processes in ring farther from margin (Fig. 38, 39); one central strutted process (Fig. 40, arrow; Fig. 41, csp); one labiate pro- cess inside marginal ring (Fig. 41, arrow) in same ring as oc- cluded processes; weakly silicified. DistRiBuTION.—Inshore euryhaline (Fryxell 1975a); found in our samples from central San Francisco Bay to Suisun Bay, indicating a broad- er freshwater range than previously proposed. Thalassiosira punctigera (Castracane) Hasle, 1983 (=T. angstii (Gran) Makarova) (Figures 42-48, 92) DetaiLep Description. Gran and Angst (1931), G. Fryxell (1978).—Cell diameter 43-145 um; areolae across valve face _ process, small spines above marginal strutted processes. Figure 28. Scale = 2 um; external view of margin at the single labiate process, valve strutted processes (arrow). Figure 29. Scale = 2 um, marginal strutted processes with short spines, large spines back from edge of margin. 138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 15 in 10 um; areolae fasciculated with areolae arranged parallel to center of fascicle (Fig. 45); strutted processes four to five in 10 um along margin (Fig. 44, 46, 92); single central process (Fig. 43, arrow); one labiate process (Fig. 46; Fig. 47, internal); large occluded processes irregularly arranged around margin (Fig. 42), although some specimens lack large occluded pro- cesses entirely (Fig. 48). DisTRIBUTION. —Coastal, large population of T. punctigera reported in Richardson Bay (A. Hauser, pers. comm., 1976), which is constantly flushed by waters from the Golden Gate. Thalassiosira nordenskioeldii Cleve, 1873 (Figure 106) DETAILED Description. Hasle (19784).—Cell diameter 10- 50 um; areolae 14-18 in 10 um across valve face; marginal strutted processes three in 10 um; separated from margin by ca. 6-8 areolae. Single strutted process in center of valve (Fig. 106); one labiate process in same ring as marginal strutted processes but not in a constant position relative to a strutted process; marginal striae 18-20 in 10 um in mantle rim. DisTRIBUTION.— Marine, cold water (Hasle 19785); only in samples near the Golden Gate Bridge. Thalassiosira pacific Gran and Angst, 1931 (Figures 49-55, 105) DetaiLeD Description. Hasle (1978b).—Cell diameter 7— 46 um; areolae 10-18 in 10 um in central area, 20 in 10 um at margin (Fig. 49); one labiate process (Fig. 54); pronounced, regular marginal strutted processes, four to seven in 10 um (Fig. 52); single central strutted process (Fig. 51, internal; Fig. 53, external; Fig. 105) adjacent to central areolae. Areolae usually in fasciculated rows with areolae parallel to radius. DisTRIBUTION.— Marine, from central San Francisco Bay through Golden Gate to Gulf of Farallons. Thalassiosira visurgis Hustedt, 1957 (Figures 56-61, 95, 96) DetaiLeD Description. Hasle (1978a).—Cell diameter 9- 18 um; areolae in central area 13-14 in 10 um, 18 in 10 um at the margin (Fig. 58); one central strutted process (Fig. 58, external; Fig. 60, internal [arrow]), and four to five strutted processes in 10 um in ring near margin (Fig. 56, external; Fig. 57); two labiate processes on opposing sides, each found be- tween marginal strutted processes and slightly inside the ring of strutted processes (Fig. 56, 58-60, 95, 96). In our prepa- ration, valves often convex or concave, suggesting heteroval- vate cells with one convex and one concave valve. Granules on valve often extending onto processes (Fig. 57). DisTRIBUTION.— Usually found surrounded with silt and clay, fresh to brackish water. Tha- lassiosira visurgis occasionally the dominant Thalassiosira in Suisun Slough, a brackish en- vironment. Thalassiosira decipiens (Grunow) Jorg., 1905 (Figures 62-67, 97, 98) DetaiLep Description. Hasle (1979).—Cell diameter 9-29 um; areolae across valve face 8-12 in 10 um, much smaller on mantle (Fig. 62); single ring of marginal strutted processes four to six in 10 um (Fig. 62, 64); one central strutted process (Fig. 63, arrow); labiate process located between two marginal strut- ted processes (Fig. 62, arrow; Fig. 97, arrow; Fig. 98), closer = Ficures 30-35. Thalassiosira eccentrica (Ehrenb.) Cleve. SEM. Figure 30. Scale = 20 um; external view of valve; eccentric areolar pattern, irregular spines. Figure 31. Scale = 2 um, large marginal spines, two rings of marginal strutted processes. Figure 32. Scale = 20 um, internal view of valve, scattered processes across valve. Figure 33. Scale = 3 um; internal view of central area, seven areolae with cribra surrounding central areola (ca), central strutted process (csp) just off center. Figure 34. Scale = 2 um, internal view of labia, scattered strutted processes. Figure 35. Scale = 3 um; external view of valve; single labiate process (Ip), scattered strutted processes, marginal strutted processes. Ficures 36, 37. Thalassiosira minuscula Krasske. SEM. Figure 36. Scale = 10 um; internal view of valve, single labia set back from margin, single row of marginal strutted processes, central strutted process. Figure 37. Scale = 2 um; same valve, internal view of marginal strutted processes, labia set back from margin. Ficures 38-41. Thalassiosira lundiana G. Fryxell. SEM. Figure 38. Scale = 10 um; external view of valve; irregular strutted processes ring valve, irregular occluded processes (arrow). Figure 39. Scale = 2 um; large occluded process, marginal strutted processes. Figure 40. Scale = 2 um; external view of central area (arrow), fasciculation. Figure 41. Scale = 10 um; internal view of valve; scattered valve strutted processes, single labiate process (arrow), single central strutted process (csp). Ficures 42-48. Thalassiosira punctigera (Castracane) Hasle. SEM. Figure 42. Scale = 20 um; external view of valve; marginal strutted processes, irregular large occluded processes. Figure 43. Scale = 2 um, single strutted process in central area. Figure 44. Scale = 2 um, marginal strutted processes. Figure 45. Scale = 20 um, internal view of fasciculated areolae, marginal strutted processes. Figure 46. Scale = 2 um, external view of labiate process, marginal strutted processes. Figure 47. Scale = 2 um, internal view of labia and external labiate process. Figure 48. Scale = 20 um, external view of valve lacking occluded processes. MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 140 MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA 141 142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 to one than the other. Often found coated with sediment (Fig. 67). DisTRIBUTION. —San Joaquin estuary, a brack- ish environment (Hasle 1979). Dominant Tha- Jassiosira in the Suisun Bay area (Arthur and Ball 1980; Wong and Cloern 1981). Discussion.—Confusion of the species 7. punctigera and T. lundiana may lead to misin- terpretation of environmental conditions. Tha- lassiosira punctigera is the larger of the two species, although there is a slight overlap between the smaller diameters of 7. punctigera and the larger diameters of 7. /undiana. The larger di- ameter of 7. punctigera, the more heavily silic- ified valve, and the prominent marginal strutted processes are the most important distinguishing features (Fig. 44, 46, 48, 92), with only a single strutted process in the center of the valve. Tha- lassiosira lundiana is fragile and weakly silicified making the fine areolar pattern more difficult to resolve, but strutted processes are scattered over the valve face (Fig. 89). The differentiation of 7. decipiens and T. vi- surgis has proven both difficult and interesting. In our samples these species commonly appear in large numbers in Suisun Slough, Suisun Bay, and the delta of the Sacramento and San Joaquin rivers. The presence of the two labiate processes clearly distinguishes 7. visurgis (Fig. 56, 60, 95, 96) from T. decipiens. Confusion of the species is possible if one process is obscured by detritus (Fig. 67). The best distinguishing character in both cleaned and uncleaned specimens is the ar- rangement of the areolae. In 7. decipiens the number of areolae in 10 wm is constant across the valve face (Fig. 97, 98). On these small valves, the cell diameter, number of areolae, and the presence of the second labiate process may not be clear enough under the light microscope to differentiate the two species. A peculiar charac- teristic of T. visurgis is seen as the light micro- scope focuses through the valve: the central area areolae display a winking effect, optically sepa- rating the central area from the margin (Fig. 95, 96). Unless the material is cleaned and mounted in Hyrax or other suitable medium, however, we cannot differentiate these two species. The overall eccentric pattern, overlap of cell diameter, and overlap of number of areolae in 10 wm make it difficult to differentiate 7. eccen- trica and T. decipiens in the 30 um diameter range without reference to more recent works. Distinctive characteristics that aid their differ- entiation include pronounced, regular strutted processes seen on 7. decipiens versus the irreg- ular spines of 7. eccentrica and the concave or convex valves of 7. decipiens versus the rela- tively flat valve of 7. eccentrica. Thalassiosira eccentrica is a coastal marine species, while vi- able T. decipiens cells in the bay system are usu- ally restricted to the Suisun Bay—Delta, a brack- ish environment; non-viable cells may be flushed from Suisun Bay by tidal action. It is possible that 7. eccentrica reported in eco- = Ficures 49-55. Thalassiosira pacifica Gran and Angst. SEM. Figure 49. Scale = 10 um; external view of valve; fasciculated areolae, single central strutted process, marginal strutted processes. Figure 50. Scale = 2 um, single labiate process (arrow) on margin between strutted processes. Figure 51. Scale = 1 um, internal view of single strutted process. Figure 52. Scale = 10 um, regular marginal strutted processes. Figure 53. Scale = 1 um, external details of central strutted process. Figure 54. Scale = 1 um, internal view of labia and external labiate process. Figure 55. Scale = 10 um; internal view of valve; regular marginal strutted processes, single central strutted process, internal labia. Ficures 56-61. Thalassiosira visurgis Hustedt. SEM. Figure 56. Scale = 2 um; external view of valve; strutted processes (sp) at margin, two labiate processes (Ip), single central strutted process (csp). Figure 57. Scale = 2 um, labiate process between two strutted processes on margin. Figure 58. Scale = 2 um, central areolae distinct from those toward margin. Figure 59. Scale = 2 um, internal view of convex valve, regular marginal strutted processes. Figure 60. Scale = 2 um, internal view of concave valve, two opposing labia. Figure 61. Scale = 2 um, external view of concave valve. Ficures 62-67. Thalassiosira decipiens (Grunow) Jorgensen. SEM. Figure 62. Scale = 10 um; external view of valve; single labiate process (arrow), regular marginal strutted processes. Figure 63. Scale = 1 um; single central process (arrow), fine siliceous granulations on external side of valve. Figure 64. Scale = 10 um; internal view of valve; single labia (arrow), regular marginal strutted processes. Figure 65. Scale = 10 um; external view of valve; consistent areola pattern across valve, regular arrangement of marginal strutted processes. Figure 66. Scale = 1 um, labiate process between two strutted processes. Figure 67. Scale = 10 um, detritus accumulation surrounding cell. MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 S x A n ie) > iF) 7) = J SS = Zz xt a S = iS) = Q Z < =) 4 w ~ z fa) ie) fe) eo < = 146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 logical studies of San Francisco Bay (Arthur and Ball 1980) is in reality a complex of 7. decipiens and 7. visurgis, two species usually found in the entrapment zone in Suisun Bay. Freshwater from the Sacramento and San Joaquin rivers flows westward to meet the tidal wedge moving east- ward in Suisun Bay. This mixing area produces vertical and horizontal circulation that tends to concentrate populations of phytoplankton (Ar- thur and Ball 1980). The average salinity is low (0.1-5%o, Arthur and Ball 1980; Sitts and Knight 1979) and well within the range established for T. decipiens, T. visurgis, and T. incerta. Group 4.—Thalassiosira species with no central strutted process but a modified ring of processes on the face of the valve. Thalassiosira stellaris Hasle and Guillard, 1977 (Not figured) DetaiLeD Description. Fryxell and Hasle (1977).—Cell di- ameter 6-20 yum; areolae elongate, fasciculated, 30 in 10 um; marginal strutted processes three to five (sometimes six) in 10 um; a ring of two to seven strutted processes 2 the distance between the center and the margin. DisTRIBUTION.— Marine (Fryxell and Hasle 1977), San Francisco Bay samples described from cultures (F226) developed at Texas A&M Uni- versity and maintained at ca. 30% salinity. Thalassiosira lacustris (Grunow) Hasle and Fryxell, 1977 (Figures 68-73, 88) DetaiLeD Description. As Coscinodiscus lacustris, Hustedt Ficures 68-73. undulated, marginal strutted processes. Figure 69. Scale = (1930).—Cell diameter 20-75 um; areolae 10-14 in 10 um in central area; valve face tangentially undulated (Fig. 68, 88); areolae fine (Fig. 71), arranged in dichotomous branching ra- diating rows (Fig. 71); five to seven strutted processes in ring ca. 4 radius from center of valve (Fig. 68, external; Fig. 69, internal); also ring on edge of mantle (Fig. 69). Labiate process large, with slit parallel to the margin (Fig. 72). This species marked by tangentially undulated structure of the valve face (Fig. 68, 69) and areolar pattern. Other tangentially undulated centric species usually in San Francisco Bay include members of the genus Cyclotella. DIsTRIBUTION.—Specimens in our samples from Suisun Slough indicate a brackish prefer- ence. Thalassiosira endoseriata Hasle and G. Fryxell, 1977 (Figure 91) DetaILeD Description. Hasle and Fryxell (1977).—Cell di- ameter 20-60 um; areolae 1 1-18 in 10 um; one labiate process, located 4 distance from margin to center; marginal strutted process projecting internally 5-6 in 10 wm; central irregular ring of 4-14 strutted processes. Areolae fasciculated with rows of areolae parallel to the radius (Fig. 91). DisTRIBUTION.—The number of specimens from samples was too small to draw positive conclusion concerning the distribution of this species, but it apparently lives at ca. 20%o salin- ity. Thalassiosira anguste-lineata (A. Schmidt) G. Fryxell and Hasle, 1977 (Figures 74-79, 93) DertaiLepD Description. Fryxell and Hasle (1977).—Cell di- ameter 17-78 um; areolae fasciculated 8-18 in 10 um; one = Thalassiosira lacustris (Grunow) Hasle. SEM. Figure 68. Scale = 10 um; external view of valve; tangentially 1 um; internal view of valve; ring of valve strutted processes, pronounced tangential undulation. Figure 70. Scale = 1 um; external view of margin, labiate process (arrow) between marginal strutted processes. Figure 71 (arrow). Scale = 1 um; internal view of valve strutted process, areolae in dichotomous branching rows. Figure 72. Scale = 1 um, internal view of labia and marginal strutted process. Figure 73. Scale = 10 wm; internal view of marginal strutted processes, dichotomous branching rows of areolae. Ficures 74-79. Thalassiosira anguste-lineata (A. Schmidt) G. Fryxell and Hasle. SEM. Figure 74. Scale = 20 um; external valve view; areolae fasciculated, regular marginal strutted processes, arc of valve strutted processes in each fascicle (arrow). Figure 75. Scale = 1 um, external view of valve strutted processes. Figure 76. Scale = 2 um, marginal strutted processes. Figure 77. Scale = 2 um; single labiate process (arrow) between two marginal strutted processes, small processes above each strutted process. Figure 78. Scale = 2 um, internal view of labia. Figure 79. Scale = 2 um, internal view of valve strutted processes. Ficures 80-85. Thalassiosira rotula Meunier. SEM. Figure 80. Scale = 2 um, external view of central area strutted processes. Figure 81. Scale = 2 um, internal view of central area strutted processes and scattered strutted processes on valve. Figure 82. Scale = 10 um; external view of center and margin; weakly silicified valve, many marginal and valve strutted processes. Figure 83. Scale = 10 um, internal view of valve, central and valve strutted processes. Figure 84. Scale = 10 um, external view of valve, single labiate process (arrow), radial arrangement of areolae. Figure 85. Scale = 2 um, external labiate process (arrow). . MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 = ge > nH is) 4 n 7) = J w & MAHOOD, FRYXELL, AND McMILLAN: 150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 Ficures 86-94. LM. Scale = 10 um. Figure 86. Thalassiosira hendeyi Hasle and G. Fryxell. Linear arranged areolae, two labiate processes (arrow), wavy margin. Figure 87. T. nodulolineata (Hendey) Hasle and G. Fryxell. Linear arranged areolae, six symmetrical areolae surrounding central area, marginal strutted processes, single labiate process (arrow). Figure 88. 7. MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA 151 labiate process (Fig. 77, arrow; Fig. 78); ring of strutted pro- cesses in small arcs (Fig. 74, 75, 79) in each fascicle located ca. 2 the distance between the margin and the central areolae (Fig. 74). Characteristic arc-shaped grouping of strutted pro- cesses regularly arranged in each fascicle diagnostic for this species. DIsTRIBUTION. — Coastal, prefers cold to tem- perate waters, usually associated with other ma- rine forms. Central San Francisco Bay to Golden Gate. Discussion. — Thalassiosira stellaris, a lightly silicified species, was found in culture from a sample taken north of Golden Gate Bridge. Tha- lassiosira endoseriata was found only once. As previously mentioned, Thalassiosira lacustris displays a tangentially undulated valve face (Fig. 68), an aid in identification. The largest concen- tration of viable cells of 7. /acustris was found in the brackish waters of Suisun Slough (Mahood 1981). Thalassiosira anguste-lineata, with its fasciculated areolar pattern and distinguishing arrangement of strutted processes in each fascicle (Fig. 74, 79, 93) is easily distinguished from other fasciculated species seen within the Bay in cleaned material in permanent mounts. However, the strutted processes on the valve face are not al- ways easily seen in an uncleaned sample, and lack of resolution may result in some confusion. When cells are united in chains, however, several threads can be seen to extend from one cell to the next, one from each cluster of strutted pro- cesses instead of the single central thread com- monly seen in the genus. Group 5.—Thalassiosira, otherwise dissimilar species that have radial areolae patterns and mul- tiple central processes. Thalassiosira rotula Meunier, 1910 (Figures 80-85, 94) DETAILED DescriPTION. Fryxell (19754); Syvertsen (1977).— Cell diameter 8-61 um (40-61 um, Gran and Angst 1931); areolae very fine, only clearly seen in the central area (Fig. 84); cluster of strutted processes in center (Fig. 80, external; Fig. 81, internal), with scattered processes across entire valve face (Fig. 83, 84); single labiate process (Fig. 84, arrow; Fig. 85, arrow); valve weakly silicified (Fig. 82). Usually observed in chains (Fig. 94). DisTRIBUTION.—In our samples 7. rotula re- stricted to the more saline portion of the central bay and Golden Gate with other marine forms. Thalassiosira incerta Makarova, 1961 (Not figured) DertaILep Description. Hasle (1978a).—Cell diameter |3- 38 um; areolae 8-16 in 10 um; three to four strutted processes in 10 um at margin; four to six strutted processes surrounding central areolae. DisTRIBUTION. —Fresh to brackish water, rare in Our samples. Discussion.— Both brackish and marine members of this group show distinctive char- acteristics that clearly differentiate them from other Thalassiosira species of the bay. Thalas- siosira rotula, first mentioned by Whedon (1939), is usually found in the central bay (Wong and Cloern 1981). This species has been reported from similar estuarine environments (Marshal 1980). Our experience has been to associate 7. rotula with other coastal or marine forms. Its charac- teristic chain formation with coin-shaped cells connected by a thick central thread, narrow gir- dle bands, delicate structure, and scattered strut- ted processes on the valve face aid in the iden- tification. Thalassiosira incerta, T. decipiens, and T. vi- surgis are all characterized by small size, eccen- tric to linear areolar patterns, and an accumula- tion of detritus around the margin of the living cell. This group has proven to be the most dif- ficult to differentiate by light microscopy. All three species have similar diameters and general char- acteristics (Table 2) and can only be differen- tiated after the sample has been cleaned of or- ganic material. In cleaned material 7. incerta possesses characteristics that distinguish it from _— lacustris (Grunow) Hasle. Tangential undulations, radial arrangement of areolae. Figure 89. 7. /undiana G. Fryxell. Weakly silicified, large irregular occluded processes at margin, irregularly arranged strutted processes on valve. Figure 90. T. nodulolineata (Hendey) Hasle and G. Fryxell. Detail of central area, strutted processes in central areola. Figure 91. T. endoseriata Hasle and G. Fryxell. Fasciculated areolae, ring of strutted processes near center of valve. Figure 92. 7. punctigera (Castracane) Hasle. Regular marginal strutted processes, weakly silicified. Figure 93. 7. anguste-lineata (A. Schmidt) G. Fryxell and Hasle. Fasci- culated, arc of strutted processes in each fascicle. Figure 94. T. rotula Meunier. Central strutted process, random valve strutted processes, forming chains. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 Ficures 95-106. LM. Scale = 10 um. Figure 95. Thalassiosira visurgis Hustedt. Focused on center, two labiate processes (arrows), radial pattern of areolae. Figure 96. T. visurgis Hustedt. Same valve as Figure 95, focused on margin. Figure 97. T. decipiens (Grunow) Jorgensen. Focused on center, regular marginal strutted processes, one labiate process (arrow). Figure 98. T. decipiens (Grunow) Jorgensen. Same valve as Figure 97, focused on margin, uniform areolae across valve. Figures 99-101. T. wongii Mahood sp. nov. Fasciculated, two circles of strutted processes on valve, one labiate process (arrow), raised central MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA 153 TABLE 2. THALASSIOSIRA SPECIES FROM SAN FRANCISCO BAy AREA. Measurements from the literature and our observations. Marginal Labiate strutted Central Diameter Areolae pro- processes strutted Distinctive Species (um) in 10 um cesses in 10 um ___ processes characteristics Group | —Species with a linear areolar pattern T. hendeyi 38-120 5-6 2 5-6 1 wavy margin T. nodulolineata 26-58 3.5-7.0(9.0) 1 3-5 5-6 strutted processes in central areola T. stmonsenii 30-57 4.0-5.5 2 5-6 1 two marginal rings of strutted processes T. tenera 10-29 9-16 1 3-5 1 flattened marginal processes Group 2—Species with eccentric patterns T. eccentrica 12-101 5-10 1 irregular 1 seven areolae around central 10-20 areola T. oestrupii var. 5.5-39.0 6-9 1 1-2 1 strutted processes project in- venrickae ward T. wongil 27-51 9-11 1 3-5 4-5 fasciculated central ring of strutted processes Group 3—Species with | central process and | marginal ring of strutted processes T. lundiana 7-43 24-30 1 5-10 1 fasciculated, weakly silicified T. punctigera 43-145 15 1 4-5 1 fasciculated, regular marginal strutted processes T. nordenskioeldii 10-50 14-18 1 3 1 radial, marginal strutted pro- cesses back from margin T. pacifica 7-46 10-18 1 47 1 fasciculated, raised central process T. minuscula 10-20 30 1 45 labiate process away from margin T. visurgis 9-18 13-14 2 4-5 1 irregular radial pattern T. decipiens 9-29 8-12 1 4-6 1 eccentric radial pattern Group 4—Species with no central strutted processes but a modified ring of strutted processes T. stellaris 6-20 30 1 3-5 (6) 2-7 processes in ring, fascicu- lated T. lacustris 20-75 10-14 1 5-7 tangentially undulated T. endoseriata 20-60 11-18 1 5-6 4-14 processes in ring, fasci- culated T. anguste-lineata 17-78 8-18 1 3-4 ring of arcs on face of valve, fasciculated Group 5—Dissimilar species with radial patterns T. rotula 8-61 20 1 47 cluster cluster of central strutted pro- cesses T. incerta 13-38 8-16 1 3-4 4-6 central processes around cen- tral areola radial pattern T. decipiens and T. visurgis. The central areola _u/olineata, the central strutted processes in 7. of T. incerta is surrounded by five to six strutted incerta are extremely small and are just visible processes similar to the arrangement seen in 7. _ under the light microscope. Thalassiosira incerta nodulolineata (Fig. 14). Unlike those in JT. nod- was rare in our collections and was only found — strutted process, pronounced and regular marginal strutted processes. Figure 102. T. eccentrica (Ehrenb.) Cleve. Eccentric areolar pattern, irregular marginal spines. Figures 103, 104. 7. tenera Proschkina-Lavrenko. Linear arrangement of areolae, robust flattened marginal strutted processes, central area raised around the central areola. Figure 105. 7. pacifica Gran and Angst. Fasciculated pronounced regular marginal strutted processes, distinctive central process. Figure 106. 7. nordenskioeldii Cleve. Large marginal strutted processes set back from margin, raised central process. 154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 8 from the Suisun Bay material. Hasle (1978a) has also reported 7. incerta from waters of the San Joaquin delta. CONCLUSIONS The 20 species presented require additional study to delineate more clearly their ecological variables. Other than salinity, little is known of their habitat requirements. Of the species pre- sented, 7. decipiens, T. nodulolineata, T. eccen- trica, T. hendeyi, and T. wongii appear to be best suited as indicators of salinity. ACKNOWLEDGMENTS Samples were collected by R. L. J. Wong, C. A. McNeil, P. Wares, and L. L. Lack. T. P. Wat- kins isolated clonal cultures from the study area and provided technical assistance. P. A. Fryxell kindly provided the Latin description of Tha- lassiosira wongii. Expert assistance was provided by the Texas A&M University Electron Micros- copy Center. M. M. Hanna and the California Academy of Sciences were helpful with literature resources. We would like to express our thanks to M. A. Hoban of the California Academy of Sciences for his careful and constructive editing. G. R. Hasle and L. K. Medlin participated in helpful discussions, and the Texas A&M Uni- versity Department of Oceanography provided partial support for maintenance of phytoplank- ton cultures. Partial support for this work was provided by National Science Foundation Grants DEB 79-23159 and DEB 77-15908 and the Cal- ifornia Academy of Sciences, G Dallas Hanna Memorial Publication Fund, and is gratefully ac- knowledged. Figure 1 was prepared by J. F. DeMouthe. LITERATURE CITED Anonymous. 1975. Proposal for a standardization of diatom terminology and diagnoses. Nova Hedwigia, Beih. 53:323- 354. ArTHUuR, J. F. AND M. D. Batt. 1980. The significance of the entrapment zone location to the phytoplankton standing crop in the S.F. Bay-Delta Estuary. U.S. Department of Interior, Water and Power Resources Service, p. 89. Cteve, P. T. 1873. On diatoms from the Arctic Sea. Bih. K. Svenska Vetensk.— Akad. Handl. 1(13):1-28. 1904. Plankton table for the North Sea. Bull. Cons. Explor. Mer. 1903-1904:216. Cupp, E. E. 1943. Marine plankton diatoms of the west coast of North America. Bull. Scripps Inst. Oceanogr. Tech. Ser. 5(1):1-238. Fryxe.t, G. A. 1975a. Three new species of Thalassiosira: with observations on the occluded process, a newly observed structure of diatom valves. Nova Hedwigia, Beih. 53:57-75. 1975b. Morphology, taxonomy, and distribution of selected species of the diatom genus Thalassiosira Cleve in the Gulf of Mexico and antarctic waters. Ph.D. Dissertation, Texas A&M University, College Station, Texas. 1978. The diatom genus Thalassiosira: T. licea sp. nov. and 7. angstii (Gran) Makarova, species with occluded processes. Bot. Mar. 21:131-141. FryxeLi, G. A. AND G. R. Haste. 1972. Thalassiosira ec- centrica (Ehrenb.) Cleve, 7. symmetrica sp. nov. J. Phycol. 8:297-317. 1977. The genus Thalassiosira: some species with a modified ring of central strutted processes. Nova Hedwigia, Beih. 54:67-98. 1980. The marine diatom Thalassiosira oestrupii: structure, taxonomy, and distribution. Am. J. Bot. 67(5): 804-814. Gran, H. H. ano E. E. AnNcst. 1931. Plankton diatoms of Puget Sound. Puget Sound Mar. (Biol.) Stn. 1929-31 7:417- 519. Hanna, G D. 1930. Hyrax, a new mounting medium for diatoms. J. Microsc. (Oxf.), Series 3, 50(4):424426. Harris, H. S., D. L. Feurstern, AND E. H. PEARSON. 1961. A pilot study of physical, chemical, and biological charac- teristics of waters and sediments of San Francisco Bay, SERL. College of Engineering and Public Health, University of Cal- ifornia Berkeley, pp. 9-14. Haste, G. R. 1976. Examination of diatom type material: Nitzschia delicatissima Cleve, Thalassiosira minuscula Krasske, and Cyclotella nana Hustedt. Br. Phycol. J. 11: 101-110. 1978a. Some freshwater and brackish water species of the diatom genus Thalassiosira Cleve. Phycologia, 17(3): 263-292. 1978b. Some Thalassiosira species with one central process (Bacillariophyceae). Norw. J. Bot. 25:77-110. 1979. Thalassiosira decipiens (Grun.) Jorg. (Bacil- lariophyceae). Bacillaria 2:85-108. 1983. Thalassiosira punctigera (Castr.) comb. nov., a widely distributed marine planktonic diatom. Nord. J. Bot. 3:593-608. Haste, G. R. AND G. A. Fryxett. 1977. The genus Thalas- siosira: some species with a linear array. Nova Hedwigia, Beih. 54:15-66. HepcGpetn, J. W. 1979. San Francisco Bay: the unsuspected estuary. San Francisco Bay, the urbanized estuary. Pacific Division of the American Association for the Advancement of Science. California Academy of Sciences, Golden Gate Park, San Francisco, California. P. 493. Hustept, F. 1930. Die Kieselalgen Deutschlands, osterreichs und der Schweiz unter Berucksichtigung der ubrigen Lander Europas sowie der angrenzenden Meeresgebiete in Krypto- gamen-Flora von Deutschland, Osterreich und der Schweiz. (ed.) L. Rabenhorst. Pp. 609-920. 1957. Die Diatomeen Flora des Fluss-systems der Weser im Gebiet Hansestadt Bremen. Abh. naturw. ver. Bremen, 34(3). JorGENSEN, E. 1905. Protist plankton. Diatoms in bottom samples. Pp. 1-254 in Hydrographical and biological inves- tigations in Norwegian fiords, O. Nordgaard, ed. Bergens Museum Strift, 1905. John Grieg, Bergen. MAHOOD, FRYXELL, AND McMILLAN: THALASSIOSIRA 155 KrasskE, G. 1941. Die Kieselalgen des chilenischen Kus- tenplankton. Arch. Hydrobiol. 38:260-286. Mauoop, A. D. 1981. Phytoplankton analysis of Suisun Slough, Fairfield-Suisun Sewer District, Fairfield, California, Unpublished report to the Fairfield-Suisun Sewer District. Makarova, I. V. 1961. Diatomaceae novae familiae Cos- cinodiscaceae e Mari Caspico borealis. Notulae Systematicae e Sectione Cryptogamica Instituti Botanici Nomine V. L. Komarovii Academiae Scientiarum U.S.S.R. 14:49-52. MarsHat, H.G. 1980. Seasonal phytoplankton composition in Lower Chesapeake Bay and Old Plantation Creek, Cape Charles, VA. Estuaries 3(3):207-216. Meunier, A. 1910. Microplancton des Mers de Barents et de Kava. Duc d’Orleans, Campagne Archique 1907. Murier-Meccuers, F, C. 1953. New and little known dia- toms from Uruguay and South Atlantic coast. Comun. Bot. Mus. Hist. Nat. Montev. 3(30):1-11. PROSCHKINA-LAVRENKO, A.I. 1961. Diatomeae novae e Mari Nigro (Ponto Exino) et Azoviano (Maeotico). Notulae Sys- tematicae e Sectione Cryptogamica Instituti Botanici Nom- ine V. L. Komarovii Academiae Scientiarum U.S.S.R. 14: 33-39. Ross, R., E. J. Cox, N. I. KArAyeva, D. G. Mann, T. B. B. Pappock, R. SIMONSEN, AND P. A. Sims. 1979. An am- mended terminology for the siliceous components of the diatom cell. Nova Hedwigia, Beih. 64:513-533. Rounp, F. E. anp P. A. Sims. 1981. The distribution of diatom genera in marine and freshwater environments and some evolutionary considerations. Proceedings of the sixth symposium on recent and fossil diatoms. Budapest Sept. 1- 5, 1980. Otto Koeltz, Konenstein, Germany:30 1-320. Sitts, R. M. AnD A. W. Knicut. 1979. Plankton ecology in the Sacramento-San Joaquin estuary. Water Science and Engineering No. 4509. University of California, Davis. P. 145. Storrs, P. M., E. A. PEARSON, AND F. F. Secteck. 1966. A comprehensive study of San Francisco Bay. Final Report, V. Summary of physical, chemical, and biological water and sediment data. University of California, Berkeley SERL, 67(2): 1-140. SyveERTSEN, E. E. 1977. Thalassiosira rotula and T. gravida: ecology and morphology. Nova Hedwigia, Beih. 54:99-112. VAN Der Werrr, A. 1955. A method of concentrating and cleaning diatoms and other organisms. Int. Assoc. Theor. App. Limno. Proc. 12:276-277. Wuepon, W. R. 1939. A three year survey of the phyto- plankton in the region of San Francisco, California. Int. Rev. Hydrobiol. Pp. 459-476. Wona, R. L. J. AND J. E. CLoern. 1981. Plankton studies in San Francisco Bay. II Phytoplankton and Species com- position. July 1977-December 1979. U.S. Geological Sur- vey. Open File Report 81-214:103. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 \ a det C255xX NH PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 9, pp. 157-224, 33 figs., 7 tables. May 6, 1986 SYSTEMATIC RELATIONSHIPS AND ONTOGENY OF THE SCULPINS ARTEDIUS, CLINOCOTTUS, AND OLIGOCOTTUS (COTTIDAE: SCORPAENIFORMES) Se eo i iSO iis a By Fogo os cine athe Ih ee Betsy B. Washington! hay 15 1880 \ Gulf Coast Research Laboratory, East Beach Drive, ics F Ocean Springs, Mississippi 64 yi Apstract: Using the methods of phylogenetic analysis proposed by Hennig (1966), characters of the larvae of 13 species of Artedius, Clinocottus, Oligocottus are examined in terms of synapomorphic states. Number and pattern of preopercular spines, gut diverticula, body shape, and a bubble of skin at the nape are identified as synapomorphic characters useful in systematic analysis of this group. The synapomorphic character, multiple preopercular spines, provides strong evidence that Clinocottus acuticeps, C. analis, C. embryum, C. globiceps, C. recalvus, Oligocottus maculosus, O. snyderi, Artedius fenestralis, A. harringtoni, A. lateralis, and A. Type 3 form a monophyletic group within the Cottidae. Within this group, the species of Clinocottus and Oligocottus are very closely related; each genus, however, appears to be monophyletic. Larvae of all species of Clinocottus possess the synapomorphy, auxiliary preopercular spines. Larval Oligocottus maculosus and O. snyderi share two derived characters, dorsal gut bumps and a bubble of skin at the nape. Artedius fenestralis, A. harringtoni, A. lateralis, and A. Type 3 also form a monophyletic group closely related to Clinocottus and Oligocottus on the basis of a unique multiple preoper- cular spine pattern. Synapomorphic characters of the larvae provide strong evidence that A. creaseri and A. meanyi are more closely related to Icelinus than to species of Clinocottus, Oligocottus maculosus, O. snyderi, Artedius fenes- tralis, A. harringtoni, A. lateralis, and A. Type 3. Characters of the larvae strongly indicate that the genus Artedius as defined by Bolin (1934, 1947) is not monophyletic and that A. creaseri and A. meanyi should be placed separately from the other species of Artedius. Clarification of the exact position of these two species in relation to the Artedius-Clinocottus-Oligocottus group and other cottids must await reexamination of characters of adults. Complete, identified, developmental series of larval cottids Artedius fenestralis, A. creaseri, A. meanyi, Oligocottus snyderi, Clinocottus embryum, and C. globiceps are described for the first time. Partial devel- opmental series of two species, Artedius Type 3 and Clinocottus analis, are also described and illustrated for the first time. In addition, four species, Artedius harringtoni, A. lateralis, Oligocottus maculosus, and Clino- cottus acuticeps are redescribed providing new and comparative information on larval development. INTRODUCTION era and 300 species (Nelson 1976). Most of these species are marine and generally distributed in coastal waters of all oceans except the Indian Ocean. Cottids are most speciose in the North Pacific where 90 species distributed in 40 genera ‘Current address: National Marine Fisheries Service, Sys- ate reported to occur between Baja, Califormia tematics Laboratory, National Museum of Natural History, and the Aleutian Islands, Alaska. Sixteen of these Washington, D.C. 20560. species belonging to the genera Artedius, Clino- (157] The Cottidae are a large, morphologically di- verse family of fishes composed of nearly 67 gen- 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 cottus, and Oligocottus are common intertidal and subtidal inhabitants of the northeast Pacific coast (Table 1). Despite their abundance the sys- tematic status and early life history of members of Artedius, Clinocottus, and Oligocottus have received little study. The few systematic studies that have ad- dressed these genera have yielded contradictory results. Most workers have placed members of Artedius, Clinocottus, and Oligocottus in the family Cottidae (Jordan and Evermann 1898; Regan 1913; Bolin 1934, 1944, 1947; Berg 1940; Taranets 1941; Howe and Richardson 1978): Jordan (1923), however, placed the genus Arte- dius in the family Icelidae. Bolin (1934, 1947), in a review of marine cottids of California, pro- posed that Artedius, Clinocottus, and Oligocottus were closely related genera that evolved from an evolutionary line of cottids tending towards a reduction of gills, pelvic fin rays, preopercular spines, and squamation. In contrast, Taranets (1941) separated members of these genera into two subfamilies. He placed Clinocottus, Oligo- cottus, and five species of Artedius in the subfam- ily Oligocottinae, and placed Artedius creaseri and A. meanyi in the Icelinae. Howe and Rich- ardson (1978) suggested that Artedius, Clinocot- tus, and Oligocottus form a closely related group, Oligocottus and Clinocottus being most closely related. However, they did not discuss characters that led to their proposal. Much of the confusion in the systematic treat- ment of these genera is due to the use of primitive and reductive characters in classifications, and the failure of past workers to define the genera on the basis of unique, derived characters. Char- acters used in past studies have not been exclu- sive to this group and are present in other cottid genera. The usefulness of characters of larvae and ju- veniles in elucidating systematic relationships has been demonstrated in several groups (Bertelsen 1951; Moser and Ahlstrom 1970, 1972, 1974; Johnson 1974; Okiyama 1974; Kendall 1979; Richardson 1981; and Moser etal. 1984). Results of these studies have indicated that ontogenetic characters provide an independent set of char- acters with which to evaluate phylogenetic re- lationships. Characters of larvae have been par- ticularly helpful in groups in which characters of adults have been reductive or generalized (Moser and Ahlstrom 1972, 1974). Although larvae of most species of Artedius, Clinocottus, and Oligocottus are frequently col- lected in nearshore plankton samples in the northeast Pacific, until recently larvae of few species have been described. Of the 16 nominal species, developmental series of identified larvae have only been described for A. harringtoni, A. lateralis, C. acuticeps, C. recalvus, and O. mac- ulosus. Other forms that belong to this group based on larval morphology, but not identified to species, were described by Richardson and Washington (1980) as Artedius Type 2, Cottidae Type 1, Type 2, and Type 3. Previous descrip- tions of A. /ateralis and O. maculosus are inad- equate for specific identification. Descriptions by Richardson and Washington (1980) were based on incomplete developmental series and too few specimens for specific and/or generic identifi- cation. The objectives of this study were: 1) to eval- uate phylogenetic relationships of Artedius, Cli- nocottus, and Oligocottus within the Cottidae following the phylogenetic methodology of Hen- nig (1966) and 2) to describe the ontogeny of larvae and juveniles of as many species of 4r- tedius, Clinocottus, and Oligocottus as possible. METHODS AND MATERIALS Systematic Procedures The investigation of systematic relationships in this study follows the phylogenetic approach first put forth by Hennig (1966) and modified and debated by Brundin (1966, 1968), Cracraft (1974), Sneath and Sokal (1973), Ashlock (1974), Mayr (1974), and others. This methodology is considered best suited to the objectives of this study because the meth- odology is a phylogenetic approach, and is well defined in regard to character evolution. Other approaches currently used are not as well suited to the purpose of this study. Numerical taxon- omy, best described by Sneath and Sokal (1973), is a phenetic approach in which taxa are clustered by overall similarity. Evolutionary systematics described by Mayr (1969) and Simpson (1961) combines both phyletic and phenetic informa- tion; however, a well-defined, repeatable meth- odology has not been incorporated into this ap- proach. The basic tenet of Hennig’s approach is that the shared possession of derived character states is the only valid criterion for establishing phy- logenetic relationships. Hennig (1966) defines a 159 WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS “APNAS SITY) UL Paquosap aie IeAIE] YOTYM JO} satoadg , ‘(1S61) SHOW Aq paquosap seAIPT , “uoreuR]dxa Joy 1x9} 9a “snjojIdsojou “F 10 SNUI]]D4OI “P JOYA 1B IBAIL] E AGA SHIPAI PY ¢ "yeandAy sapn[ouy ; (8.61) UOspseyory pue aMoy JO uONe]IdwWos WOY pur s}UNOD [eUIsLO WOL , 9 (SE ‘pE) LEE 9+9 € (pI) SI-ZI (pI) SI-ZI (61) O@-LI (IILA) XI-IIA MapaUus SN1OI081O x 9 (€€) SE-ZE 9+9 € (pI) SI-€1 (€1) bI-ZI (91) LI-ST (IIIA) XI-IILA o1jjaqna sni10908110 9 (9€) LE-vE 9+9 € (pI) SI-€I (pI) SI-€1 (81) 61-91 (XI) X-IIIA SISUBULA SNIIOI08YO 9 (€€) pE-€E 9+9 € (pI) SI-TI (€1) bI-ZI (L1) 8I-SI (IIIA) XI-TIIA SNSOJNIDU SN110908110 9 (€€) €€-ZE 9+9 € (pI) SI-€1 (ZI) €1-6 (91) 91-41 (XD XI-IIIA SNA]DIAA SNNOIOUY Dy (9) L-9 (pE “€€) PE-ZE 9+9 € (pI) SI-€T (11) ZI-II (91) LI-€I (XT) X-IIIA $d991qO]3 SNIOIOU 9 (pe “€€) SE-EE 9+9 € (pI) SI-TZI (OI) TI-6 ($1) LI-FI1 (XD X-IIIA Uindquia SNIOIOUND 9 (Z€) SE-1€ 9+9 € (S1) SI-FI (pI ‘Z1) PI-II (LI ‘91) 8I-F1 (XD X-XI SIJDUD SNHOIOUN 9 (€€ ‘TE) E€-1E 9+9 € (pI) SI-€1 (Z1) €1-6 (91 ‘S1) LI-€I (IILA) XIILA Sd9I1INID SNIJOIOU! ) 9 (ZE) pE-ZE 9+9 € (91) LI-+1 (ZI) €I-11 (S1) 9I-F1 (XD X-XI snjopidsojou snipaly, 9 (p€) SE-€€ 9+9 (Z) €-Z (ST) 9I-F1 (ZI) ZI-01 (91) LIVI (X) X-XI 1AUDIUL SNIPAM 9 (€€) pE-ZE 9+9 € ($1) 9I-F1 (€1) PI-ZI (91) LI-SI (XD X-IIIA S1]D19]D] SNIPAUY » L (pe ‘€£) PE-ZE 9+9 € (pI) SI-€I (€1) pI-0l (L1) 8I-SI (XD X-IIIA MMOJBULDY SNIPAUY » 9 (S€) S€-ZE 9+9 € (S1) 9I-F1 (€1 ‘ZI bI-ZI (L1) 81-91 (XD XI-IIIA S1]DAISAUAY SNIP 9 (1€) 1€-0€ 9+9 € (91) LI-SI (Ol) 11-6 (€1) PI-ZI (X) X-XI MaSDad) SNIPAIF 9 (€€ ‘TE) CE-1E 9+9 £ (91) 9I-F1 (€1) €I-ZI (91) 9I-SI (XD) XI-IIIA SNUI]]D1OI SHIPAUY sAei [eda1s z@PIQIUIA [RIO], sel [epnes shel skel skel skel souids -orgourlg jediouug uy SIA[ag uy [e10109g uy jeuy uy [esioqg uy [esioqd (‘sasayjuared ut pareoiput st apoy)) ,SOLSTAa] AIH], GNV SA.LLOIO9ITE ANY ‘SALLODONIT ‘SAIdaLUy AO SALAS “] FTAV | 160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 monophyletic group as a group in which all members are descended from a single stem. The common possession of one or more derived char- acters is the only conclusive evidence that a group is monophyletic. Shared, plesiomorphic char- acters are not used because primitive character states inherited from an ancestral taxon may re- main unchanged in various divergent lineages and may not be evidence of close relationship. Monophyletic groups that arose from a common stem by the same splitting process are called sis- ter groups. Every monophyletic group, together with its sister group, constitutes a monophyletic group of higher taxonomic rank. Since only derived character states are used in determining phylogenetic relationships, the po- larity of character states was determined through outgroup comparisons. The outgroup taxa ex- amined in this study included larvae of seven different cottid genera: Scorpaenichthys mar- moratus, Hemilepidotus spinosus, Leptocottus armatus, Enophrys bison, Myoxocephalus sp., Icelinus sp., and Radulinus asprellus. Members of these genera are quite varied and represent several divergent lineages within the Cottidae (Bolin 1934, 1947; Taranets 1941; Howe and Richardson 1978). Larvae of several other scor- paeniform families also were examined for out- group comparisons. These taxa included: Se- bastes flavidus (Scorpaenidae); Hexagrammos sp. (Hexagrammidae), Cyclopteridae Type | (Cy- clopteridae), and Stellerina xyosterna (Agoni- dae). SELECTION OF CHARACTERS FOR ANALYSES. — A variety of characters were examined in Arte- dius, Clinocottus, and Oligocottus larvae includ- ing meristics, morphology, pigmentation, spi- nation, and developmental osteology. However, of the 5O characters initially examined, many were deleted from final analysis. The criteria used in deleting characters are as follows: (1) Characters that exhibit a large amount of variability were deleted from analysis. High- ly variable characters are poor indicators of phylogenetic relationships (Bolin 1947; Simpson 1961; Mayr 1969). Examples of variable characters are head pigmentation and number of posttemporal-supracleithral spines. (2) Derived character states found in only one species were deleted. Again characters of this nature are of no value in determining intra- group relationships. Hindgut diverticula of Clinocottus acuticeps are an example of a specific character. (3) Characters in which the sequence of change or the primitive and derived states could not be identified were deleted from the analysis. Many of the morphometric and pigmenta- tion characters fell into this category. Taxonomic Procedures Larval descriptions are based on both labo- ratory-reared larvae and field-collected speci- mens. Egg masses were spawned from ripe C/i- nocottus globiceps, Oligocottus maculosus, and O. snyderi collected from tidepools along the central Oregon coast during winter-spring 1979 and 1980. Larvae were maintained at 12—13°C. In addition to reared specimens, developmental series were put together with larvae obtained with 70 cm bongo nets and neuston nets off the coast of Oregon between 1969 and 1978. Samples were taken in all months of the year from an area concentrated along an east-west transect off Newport, Oregon (lat. 44°39.1'N). Specimens were also obtained from estuarine and coastal collections of the Southwest Fisheries Center, La Jolla Laboratory, National Marine Fisheries Ser- vice; Scripps Institution of Oceanography; Los Angeles County Museum of Natural History; Marine Ecological Consultants; California Acad- emy of Sciences; Humboldt State University; Northwest Fisheries Center, Seattle Laboratory, National Marine Fisheries Service; and Univer- sity of Washington. Reared specimens of Arte- dius lateralis from College of Fisheries, Univer- sity of British Columbia and Oligocottus maculosus and Clinocottus acuticeps from Van- couver Public Aquarium were also utilized. Transforming and juvenile specimens were col- lected monthly from 1977 to 1980 from tide- pools along the central Oregon coast. All specimens were preserved in 5 or 10% buff- ered formalin and some material was subse- quently transferred to 36 or 40% isopropyl al- cohol. Developmental series of larvae and juveniles were assembled for 12 of the 16 species of Ar- tedius, Clinocottus, and Oligocottus. The num- ber of specimens examined in each series varies from 11-38 according to availability of material. Developmental series were formed utilizing field- caught larvae, except as noted below, because of WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 161 the large amounts of variation in morphology and pigmentation in laboratory-reared larvae. Newly hatched, reared larvae are included in se- ries of Artedius fenestralis, A. lateralis, Clino- cottus acuticeps, C. globiceps, Oligocottus mac- ulosus, and O. snyderi. In addition, reared larvae were used to supplement incomplete develop- mental series of field-caught larvae of Clinocottus globiceps and Oligocottus maculosus. Marked differences between developmental series based on field specimens and laboratory-reared speci- mens are noted in the descriptions. Developmental stages follow the terminology of Ahlstrom et al. (1976), except that the tran- sitional period between the larval and juvenile stages is marked by an increase in pigmentation particularly over the head and in saddles along the dorsum, a reduction in the size and number of preopercular spines, an ossification of the pel- vic fin spine and rays, and the formation of scales. Specimens are referred to as juveniles when they settle from the plankton and assume a benthic existence. MorpHometrics.— Measurements of selected body parts were made to the nearest 0.1 or 0.01 mm using an ocular micrometer in a stereo- microscope. Measurements were made following the definitions of Richardson and Laroche (1979) except as follows: body depth at anus = vertical distance from the dorsal to ventral body margin at the anus, snout to pelvic fin origin = horizon- tal distance from the tip of the snout to a vertical through the origin of the pelvic fin, and origin of pelvic fin to anus = horizontal distance from a vertical through the origin of the pelvic fin to the anus. Head length is abbreviated as HL. Detailed tables documenting the development of meristic elements for larvae of Artedius, Clinocottus, and Oligocottus are presented by Washington (1981). All body lengths given in this study refer to either notochord length (NL), which is defined as snout tip to notochord tip preceding devel- opment of the caudal fin; standard length (SL), which is defined as snout tip to the posterior margin of the hypural plates; or total length (TL), which is defined as snout tip to the posteriormost margin of the caudal fin. Unless otherwise in- dicated, all lengths given are standard length. Meristics.— Following the methods of Din- gerkus and Uhler (1977), several larvae were cleared and stained with Alcian Blue and Ali- zarin Red S for each species when specimens were available in sufficient numbers. Counts were a EE Lao a a =—=-—— TS made of dorsal fin spines and rays, anal fin rays, pelvic fin spines and rays, principal caudal rays, branchiostegal rays, preopercular spines, and vertebrae. Vertebral counts always included the urostyle. All meristic elements were counted if they absorbed Alizarin stain. Principal caudal rays are defined as the number of caudal fin rays that articulate with the upper and lower hypural plates. Counts of meristic elements were also made on unstained larvae from the developmental se- ries used in the morphometric examination. All fin rays and spines, branchiostegal rays, pre- opercular spines, and myomeres were counted when visible under magnification. In this study, all fin rays and spines were counted, regardless of whether they arose from the same pterygio- phore. (For detailed meristic and spination tables of developmental series of known Artedius, Cli- nocottus, and Oligocottus larvae see Washington 1981). SPINATION.— Spine terminology generally fol- lows Richardson and Washington (1980) in which spines are named for the bones from which they originate. TAXONOMIC TERMINOLOGY.—Results of this study do not agree with previously recognized limits of the genus Artedius. In order to avoid confusion, species in this group are designated as Artedius Group A including A. fenestralis, A. harringtoni, A. lateralis, and A. Type 3 (either A. corallinus or A. notospilotus) or Artedius Group B including A. creaseri and A. meanyi. Larvae referred to as Artedius Type 3 are either A. cor- allinus or A. notospilotus. Positive identification is not possible at this time because of the lack of late-stage larval specimens. (See descriptions for discussion of identification.) RESULTS Description of Characters Considered PREOPERCULAR SPINATION.—The number of preopercular spines is a relatively stable, con- servative character in larval cottids. Most cottid larvae (22 of 28 known genera) possess four ap- proximately equal-sized spines situated along the posterior margin of the preopercle. Generally the dorsalmost spine increases in size with devel- opment while the lower three spines are reduced or lost. A modification of this basic preopercular pat- tern is found in larvae of several species of Ice- 162 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 2 2 ZY BX Ficure 1. Multiple preopercular spines in larval Artedius, Clinocottus, and Oligocottus. A) Artedius harringtoni, B) A. fenestralis, C) A. lateralis, D) A. Type 3, E) Clinocottus acuticeps, F) C. embryum, G) C. globiceps, H) C. analis, 1) Oligocottus snyderi, J) O. maculosus. linus and Myoxocephalus. These larvae possess an additional small, auxiliary spine situated on the inner shelf of the preopercle anterior to the bases of the four principal preopercular spines. A third pattern of preopercular spination is found in larvae of Clinocottus, Oligocottus, and Artedius Group A (Fig. 1). These larvae possess 5-24 small spines situated along the posterior margin of the preopercle. Two basic patterns of multiple preopercular spines occur in larvae of this group. In four species of Artedius (Group A), the dorsalmost, middle, and ventralmost spines become enlarged relative to the other preoper- cular spines. During transformation, the dorsal- most spine continues to increase in size, while the middle spines (7-9), midventral spines (1 1- 14), and ventralmost spines each fuse together, forming three large bumps on the preopercular margin. The other preopercular spines gradually disappear. In larvae of Clinocottus and Oligocottus the dorsalmost spine increases in size relative to the other preopercular spines. During transforma- tion the lower spines regress and disappear while the dorsalmost spine remains prominent. Outgroup comparisons with larvae of closely related Scorpaeniformes indicate the presence of five or fewer approximately equal-sized pre- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 163 Figure 2. Preopercular spines of larval Artedius meanyi and A. creaseri. opercular spines. Sebastes and Stellerina larvae possess five and four spines, respectively; hexa- grammid larvae possess five to six, three, or no spines, and cyclopterid larvae have lost all pre- opercular spines. The cottid taxa that possess four equal-sized preopercular spines also tend to have many other primitive character states. The presence of four equal-sized spines is considered the plesiomorphic state for preopercular spines in cottid larvae. The modified pattern of spines found in larval Icelinus and Myoxocephalus could easily be de- rived from the basic pattern of four preopercular spines. In fact, larvae of several species of Ice- linus and Myoxocephalus possess only four pre- opercular spines. The presence of an auxiliary spine on the preopercle probably represents an intermediate character state leading toward mul- tiple preopercular spines. The multiple preopercular spines of larvae of Artedius (Group A), Clinocottus, and Oligocottus are unique to this group. Multiple preopercular spines are not present in any other known cottid or scorpaeniform larvae. Multiple preopercular spines are derived character states indicative of the monophyletic origin of this group. BASAL PREOPERCULAR SPINE.—Larvae of Ar- tedius meanyi and creaseri, larvae of at least two species of Icelinus, and larvae of Myoxocephalus possess small projections or spines on the base ofeach of the four main preopercular spines (Fig. 2). These basal spines project out at 90° angles to the axis of the main preopercular spines. The basal spines are most pronounced in early post- flexion larvae. With development, four bony ridges form on the inner shelf of the preopercle, parallel to each basal spine. These bony ridges grow toward the basal spines and gradually fuse with them, forming bony arches over the forming lateral line canal of the preopercle. These basal spines are not present in other cottid or scor- paeniform larvae examined and probably are a derived character state. INNER SHELF PREOPERCULAR SPINES.— Larval Clinocottus possess one or two tiny spines on the inner shelf margin of the preopercle. Clinocottus acuticeps larvae have only one inner shelf spine. All other Clinocottus larvae examined have two. These spines are transient features which form in postflexion larvae and are lost before trans- formation. They appear to be unique to this group and, as such, to be derived character states. Nape Bussie.—Larvae of Oligocottus macu- losus and O. snyderi possess a distinctive bubble of skin in the nape region just anterior to the origin of the dorsal fin (Fig. 3). This bubble is present at hatching and persists for two or three weeks (to about the beginning of flexion of the notochord). No other known cottid larvae pos- sess a bubble of skin at the nape; accordingly, this bubble is probably a derived character unique to these two species. (Larvae of O. rimensis and O. rubellio are unidentified and it is not known if they also possess this character.) Gut DiverticuLA.—Long protrusions or di- verticula extend dorsolaterally from either side of the abdominal cavity in larvae of Artedius fenestralis, A. lateralis, and A. Type 3 (Fig. 4). These diverticula are present at hatching and per- sist throughout larval development. Newly 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 3. hatched larvae of Oligocottus maculosus and O. snyderi possess similar but less pronounced bumps or protrusions on either side of the dorsal surface of the abdominal cavity. These bumps are present at hatching, but they disappear after two to three weeks at about the onset of noto- chord flexion. The diverticula of Artedius fenes- tralis, lateralis, and Type 3 and the smaller pro- trusions of Oligocottus appear to be homologous structures, with the smaller bumps constituting an intermediate form. These diverticula are unique, derived characters not known in other cottid larvae. Larvae of C. acuticeps also possess long di- verticula which extend posteriorly on either side of the anus. Larvae of C. globiceps, C. embryum, and C. analis have bulges on either side of the anus. Although these bulges appear to form an intermediate state in the evolution of hindgut diverticula, histological sections of the guts of larval Clinocottus yielded inconclusive results. Larval C. analis, C. embryum, and C. globiceps possess an enlarged coelom on either side of the hindgut, but no distinct diverticula. Hindgut di- verticula appear to be unique to C. acuticeps. PARIETAL AND NuCHAL SpINeEs.— Most larval cottids develop two spines, a large anterior pa- rietal and a smaller posterior nuchal spine at the posterior edge of the parietal bones. The anterior parietal spine develops first, followed by a small- er nuchal spine that forms just posterior to it. These spines are generally transient structures that form late in larval development and are re- duced or lost during transformation. In many Nape bubble of larval Oligocottus snyderi. species of cottids, the spines appear to fuse to- gether enclosing a small canal between the bases of the two spines. This canal eventually becomes part of the cranial lateral line system. In other species, the spines decrease in size without fusing together. Concurrently, sheets of bone extend an- teriorly and posteriorly from the spines and eventually fuse together, forming an incipient cranial arch. Similar parietal and nuchal spines occur in lar- vae of most other scorpaeniform families and appear to be homologous to those of cottid lar- vae. The presence of a parietal and nuchal spine is probably the primitive or ancestral condition in the Cottidae. Parietal and nuchal spines have undergone modification and elaboration in larvae of most of the species of Clinocottus and Oligocottus, and in Artedius fenestralis, A. harringtoni, A. later- alis, and A. Type 3. Two species (Artedius har- ringtoni and Clinocottus acuticeps) have lost these spines completely. Three species (Clinocottus analis, C. embryum, and C. recalvus) have re- tained the primitive condition of possessing a parietal and nuchal spine. The remaining species (A. fenestralis, A. lateralis, O. maculosus, O. sny- deri, and C. globiceps) have tended toward an elaboration and increase in number of parietal spines. Generally, these larvae develop a cluster of three to six spines, which are situated in two transverse rows at the posterior margin of the parietal region. During transformation, these clusters of spines decrease in size and disappear. At the same time, sheets of bone extend ante- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 165 Be SOND ie Ficure 4. Dorsal gut diverticula in larval Artedius fenestralis. riorly and posteriorly from the bases of the two rows of spines and eventually fuse together. This bony arch becomes a part of the cranial lateral line system in juveniles. The absence of parietal spines in A. harringtoni and C. acuticeps is probably a secondary loss and, as such, represents a derived condition. The elaboration of spines into clusters is apparently unique to larvae of Artedius, Clinocottus, and Oligocottus and is also a derived state. PIGMENTATION. — Melanistic pigmentation varies greatly among cottid larvae ranging from relatively unpigmented forms to heavily pig- mented ones. Larvae of Artedius, Clinocottus, and 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Oligocottus are all lightly pigmented and possess numerous, intense melanophores over the dor- solateral surface of the gut and in a row posterior to the anus. The shape and number of midline melanophores varies between species. Larvae of all but one species, A. creaseri, possess several melanophores in the nape region. The presence of head pigment and anteroventral gut melano- phores varies among the species of Artedius, Cli- nocottus, and Oligocottus. Although pigment patterns are diagnostic at the specific level, they are difficult to evaluate for use in systematic analysis. Many fish larvae in distantly related families, and even orders, are similarly pigmented. Several cottid genera other than Artedius, Clinocottus, and Oligocottus also possess similar pigment patterns. Hence, it is dif- ficult, if not impossible, to determine which pig- ment patterns are primitive and which are de- rived. Nevertheless, trends in certain areas of pig- mentation can be discerned. Among known cot- tid larvae, a discrete nape pigment patch is found only in members of Artedius, Clinocottus, Oli- gocottus, Enophrys, Myoxocephalus, and Gym- nocanthus. Nape pigment is probably derived in cottid larvae. The number of ventral midline me- lanophores situated posterior to the anus ranges from 2 to 33 in larvae of Artedius, Clinocottus, and Oligocottus. In addition, the shape and spac- ing of these melanophores varies from small dots to long slashes extending onto the ventral finfold to large pigment blotches. Artedius creaseri and A. meanyi both possess irregularly shaped blotches of pigment along the ventral midline. Artedius fenestralis, A. harringtoni, and A. lat- eralis all possess distinctive pigment slashes. Oli- gocottus and Clinocottus larvae possess small, round melanophores. Although these pigment patterns bind certain species together, it is dif- ficult to determine ancestral versus derived states. Both pigment blotches and distinctive midline slashes are found in larval Jcelinus. Pigment, as well as other characters, indicates that Jcelinus shares close affinities with Artedius; however, the direction of the evolution of pigment patterns cannot be determined. MorpHometrics. —Cottid larvae exhibit a di- versity of body forms. Body shape ranges from short and stubby (Artedius [Group A] and Eno- phrys) to long and slender (Radulinus and Icelus) to globose (Malacocottus). Larval Artedius (Group A), Clinocottus, and Oligocottus have short, stubby bodies with blunt, rounded snouts. Gut length is moderately long and the posterior por- tion of the hindgut trails well below the rest of the body. Measurements of body parts frequently over- lap in larval Artedius (Group A), Clinocottus, and Oligocottus because of their similarity in body shape. These similarities make it difficult to de- termine discrete character states or transforma- tion series. In addition, many body parts change markedly during larval development, frequently exhibiting allometry. Because of the extreme di- versity of forms found in larval cottids, it is dif- ficult to evaluate morphometric characters for systematic analysis. Trends can be observed in only a few body parts. Larvae of Artedius creaseri and A. meanyi have long, pointed snouts. They develop relatively long ascending processes on the premaxillary. The un- derlying ethmoid cartilage is relatively large, causing a pointed, humped appearance of the snout. Larval Jcelinus exhibit pointed snouts similar to that of A. meanyi and A. creaseri. In contrast, all other larval Artedius, Clino- cottus, and Oligocottus have blunt, rounded snouts. The ascending processes of their pre- maxillaries are relatively short, and the ethmoid cartilage forms late in development. Snout length is variable in the outgroup taxa. Sebastes larvae have a somewhat long, pointed snout, but the hexagrammid, cyclopterid, and agonid larvae examined have shorter, rounded snouts. Within the cottids, Enophrys, Leptocot- tus, Hemilepidotus, and Scorpaenichthys larvae all have blunt, rounded snouts. This condition is probably the primitive condition relative to larvae of Artedius, Clinocottus, and Oligocottus because it is widespread in several divergent gen- era of cottids and scorpaeniforms. The pointed snout appears to be a derived condition. Although gut length varies greatly among cot- tid larvae (Richardson and Washington 1980), larval Artedius (Group A), Clinocottus, and Oli- gocottus possess distinctive guts with the hindgut coiled very loosely and extending posteriorly. The tip of the hindgut extends ventrally well below the rest of the body and posteriorly to the origin of the anal fin. This condition is most pro- nounced in Clinocottus larvae. Larval C. acuti- ceps and C. embryum have especially long, trail- ing hindguts which extend posteroventrally past nn WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 167 the anal fin origin. A trailing gut is unique to larvae of Artedius (excepting A. meanyi and A. creaseri), Clinocottus, and Oligocottus, and is as- sumed to be a derived condition. Petvic Fin Rays.—The number of pelvic fin rays in cottids ranges from one spine and five rays to no spines or rays. Pelvic fin rays are generally considered to be undergoing reduction in the cottids. The primi- tive state is I,5 fin rays as in other Scorpaeni- formes. Reduction in number of rays is a derived state. Larvae of Artedius, Clinocottus, and Oligocot- tus all possess I,3 pelvic fin rays, except for A. meanyi. Artedius meanyi usually possesses I,2 pelvic fin rays. The outermost of these fin rays is markedly long and thickened, and the tips of this ray are separated. Jcelinus also possesses I,2 pelvic fin rays; however, both rays are relatively short and fine. The thickened outer ray of A. meanyi may have evolved through the fusion of two fin rays. If so, this condition may constitute an intermediate state between the three pelvic fin rays of Artedius, Clinocottus, and Oligocottus and the two pelvic fin rays of Icelinus. BRANCHIOSTEGAL RAys.—The scorpaenids, considered to be the most generalized scorpae- niform (Bolin 1947; Quast 1965), possess seven branchiostegals. The hexagrammids and the zan- iolepids, which occupy an intermediate position between the scorpaenids and cottids (Quast 1965), both possess six branchiostegals. Most cottids possess six branchiostegal rays; however, the psychrolutids and some freshwater Cottus species have seven branchiostegals. The psychrolutids are a distinct group which possess many derived characters and have apparently diverged from other cottids. Similarly, members of Cottus also possess many derived characters that apparently reflect adaptation to a freshwater habitat. Cottids that are generally considered to be primitive be- cause they retain many primitive features all pos- sess six branchiostegal rays. Although the possession of seven branchioste- gal rays is probably the primitive condition in the scorpaeniforms, possession of six branchi- ostegals appears to be the primitive state within the cottids. Cottid genera such as Jcelinus and Hemilepidotus, which Bolin (1947) considered to have evolved from the evolutionary line lead- ing to Artedius, Clinocottus, and Oligocottus, all possess six branchiostegal rays. Six branchioste- gals is probably also the primitive state relative to Artedius, Clinocottus, and Oligocottus. Be- cause those cottids generally considered to be from the same evolutionary lineage as Artedius (Bolin 1934, 1947) all have six branchiostegals, it is assumed that the seven branchiostegals found in Artedius harringtoni and in some Clinocottus globiceps are secondarily derived. POSTTEMPORAL-SUPRACLEITHRAL SPINES. — Larvae of most known scorpaeniforms (includ- ing most known cottids) develop three spines in the posttemporal-supracleithral region of the head. Generally, two spines form first on the ven- tral portion of the posttemporal bone, and one spine forms midway along the posterior margin of the supracleithrum. These spines persist dur- ing transformation at which time the surround- ing portions of the posttemporal and supracleith- ral bones undergo modification and canals form in these bones between the spines. This entire complex then develops into the junction point of the cephalic lateral line system and the lateral line system. This pattern of spines probably rep- resents the plesiomorphic condition in larval cot- tids. In larvae of Artedius, Clinocottus, and Oligo- cottus the posttemporal-supracleithral spines are frequently modified. The modifications appear to be correlated with those of the parietal and nuchal spines. Larvae that have lost parietal spines do not develop posttemporal-supracleith- ral spines, and larvae that have evolved complex clusters of parietal spines also develop clusters of posttemporal-supracleithral spines. Neither 4. harringtoni nor C. acuticeps larvae develop any spines in the posttemporal-supracleithral region. Larvae of A. creaseri, A. meanyi, C. embryum, and C. recalvus possess two posttemporal and one supracleithral spine. Artedius fenestralis, A. lateralis, O. maculosus, O. snyderi, and C. glo- biceps all develop more than three posttemporal- supracleithral spines. As described for parietal and nuchal spines, the posttemporal-supracleithral spines vary among species of Artedius, Clinocottus, and Oli- gocottus. This variability among closely related species suggests that these spines may be undergoing rapid modification in this group and loss of spines or possession of clusters of spines may represent convergent or parallel evolution. As with the parietal and nuchal spines, the absence of posttemporal-supracleithral spines in 168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Tasie 2. CHARACTER STATES USED IN SYSTEMATIC ANALYSIS AND THEIR DisTRIBUTION AMONG LARVAL ARTEDIUS, CLINOCOTTUS, AND OLIGOCOTTUS AND THE OUTGROUP TAXA. Preopercular spine pattern Basal No. of Boral preopercular Bubble of Dorsal mid, & ; ae A preopercular : spine Auxiliary pre- skin at nape spines SS Nee ae opercular spines > =a Equal- larg- spine Pres: \7 22 Se eee Pres- <5 = sized est largest Absent ent Absent One Two’ Absent ent Artedius fenestralis Xx x > 4 xX x Artedius harringtoni >, « x ».4 Xx Xx Artedius lateralis »« x x x »« Artedius Type 3 », 4 Xx >, 4 Xx x Oligocottus maculosus xX > 4 > 4 Xx 4 Oligocottus snyderi X x xX x x Clinocottus aculiceps xX Xx », 4 »¢ xX Clinocottus analis Xx Xx Xx Xx Xx Clinocottus embryum >, 4 x Xx Xx > 4 Clinocottus globiceps », 4 »« Xx 4 Xx Clinocottus recalvus » 4 Xx Xx x Xx Artedius creaseri x », 4 Xx X Xx Artedius meanyi Xx x »4 Xx X Scorpaenichthys marmoratus X >, 4 xX », 4 X Hemilepidotus hemilepidotus XxX Xx 4 », & X Leptocottus armatus X xX xX », 4 Xx Enophrys bison x x xX >, 4 xX Myoxocephalus sp. Xx x x X >, « Icelinus sp. x x Xx x Xx Radulinus asprellus x x X xX Sebastes flavidus x Xx Hexagrammos sp. z >, * Xx », « Cyclopteridae Type 1 > bd * X X Stellerina xyosterna Xx > 4 x >, ¢ ».¢ * Character absent. A. harringtoni and C. acuticeps is probably asec- criteria for character selection listed in the meth- ondary loss and hence a derived state. The trend ods. toward an elaboration of these spines is found Character 1: Number of preopercular spines. only in members of Artedius, Clinocottus, and A) 5 Oligocottus and is also considered a derived state. B) 0 CHARACTERS SELECTED FOR SYSTEMATIC C) 4 ANALYsIS.—Of the 50 characters examined, 10 D) >5 characters were selected for use in the phyloge- Character 2: Preopercular spine pattern. netic analysis (Table 2). These 10 best fit the A) preopercular spines equal-sized WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 169 TABLE 2. CONTINUED. Panetal Hindgut shape Dorsal gut spines Trails diverticula Modi- No. pelvic fin rays Snout shape ‘ites iPatih Absent Small Large Two fied >3 3 7) Rounded Pointed Compact ly greatly i x x x me x x ae x xX X x x x x X Xx Ke xX 5,8 x x x x X XG x x x x EG x x x x > x x x x og x xi x X x: xg de x x: De as x xe Xs axe X x xe x x Ke x x x x x x Xx xX x x Xe x x xX x x x x xX Xi x x x x x xX x x x x x x xX x x Ke x x K xi x x x x x x x xe x x * x x xX x x x xX B) dorsalmost spine largest ©C)2 C) dorsal, middle, and ventral spines larg- Character 5: Bubble of skin at nape. est A) absent Character 3: Basal preopercular spines. B) present A) absent Character 6: Diverticula on dorsal surface of B) present gut. Character 4: Auxiliary preopercular spines. A) absent A) absent B) small bumps B) 1 C) long diverticula 170 7) >| z| | a | TON! ERI RE AL MEANYI HA R FENESTR PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 A TYPE 3 GLOBICEP A & ANALIS CE Ficure 5. cladogram indicate synapomorphies. Character 7: Parietal spines. A) 2 B) modified: clusters of spines C) modified: absent Character 8: Number of pelvic fin rays. A) >3 Byes ©)r2 Character 9: Snout shape. A) rounded B) pointed Character 10: Hindgut shape. A) compact; not trailing below body; end- ing anterior to anal fin origin B) hindgut trailing slightly below body; ex- tending to origin of anal fin C) hindgut trailing well below body; ex- tending posterior to anal fin origin PHYLOGENETIC RELATIONSHIPS A hypothesis of evolutionary relationships among species in the cottid genera Artedius, Cli- nocottus, and Oligocottus is presented in Figure 5. Two main lineages or sister groups of Artedius, Clinocottus, and Oligocottus larvae are repre- multiple preopercular spines 2a. “Artedius” spine pattern 2b. dorsal spine longest 3 basal preopercular spine 40. auxiliary spine (one) 4b. auxiliary spine (two) 5S. nape bubble 6a. dorsal gut diverticula 6b. dorsal gut bumps 8. 2 pelvic fin rays 9 pointed snout 10a. trailing gut 10b. greatly trailing gut Cladogram of systematic relationships between Artedius, Clinocottus, and Oligocottus. Characters numbered on sented in the resulting cladogram based on shared derived characters of the larvae. Species with larvae possessing the synapomorphic characters, multiple preopercular spines and trailing guts, form one major group. In addition to the pos- session of shared, derived characters, larvae of this group are extremely similar in pigmentation and body shape. The second major evolutionary line consists of species sharing two derived char- acters, basal preopercular spines and pointed snouts. This line includes Artedius creaseri, A. meanyi, and Icelinus. The two main evolutionary lines or groups of Artedius, Clinocottus, Oligocottus, and Icelinus correspond to Taranets’s (1941) classification of these species. Taranets (1941) placed Artedius meanyi and A. creaseri in the subfamily Icelinae, along with members of Jcelinus and Chitonotus. He based this decision on the following char- acters: the upper preopercular spine larger than the lower spines; two rows of bony plates on the body—one along the lateral line, the other at the base of the dorsal fins; and scales usually present on other parts of the body. All of these characters are undergoing reduction and are present in sev- eral different genera of cottids. As such, they have WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 171 low systematic value. Taranets (1941) combined Clinocottus, Oligocottus, Artedius corallinus, A. fenestralis, A. harringtoni, A. lateralis, and A. notospilotus in the subfamily Oligocottinae. This subfamily was characterized by the absence of spines or ridges projecting through the skin on the head, weakly developed preopercular spines, body naked or with bony plates reduced, and **.. other characters.” Unfortunately, Taranets did not mention which other characters he ex- amined. In contrast, other investigators have placed Ar- tedius meanyi and A. creaseri in the genus Ar- tedius along with the five other species of Arte- dius discussed above. Howe and Richardson (1978) and Bolin (1947) proposed that although Artedius shared close affinities with Jcelinus, it was more closely related to Clinocottus and Oli- gocottus. Jordan (1923), however, stated that Ar- tedius was most closely related to Icelinus be- cause of the common possession of bony plates on either side of the dorsal fins. The evolutionary lineage (Fig. 5) containing larvae with multiple preopercular spines in- cludes three distinct groups of species. One of these groups includes all the species of the genus Clinocottus. Larvae of this group share one syn- apomorphic character, inner-shelf preopercular spines. Larvae of Clinocottus possess one or two auxiliary spines on the inner preopercular shelf. These spines appear to be a derived character unique to members of this genus and provide evidence that the genus is monophyletic. This genus has been previously recognized and de- fined on the basis of adult characters, e.g., loss of scales, an advanced anus, and possession of a heavy, blunt penis (Hubbs 1926; Bolin 1934, 1944, 1947; Taranets 1941; Howe and Richard- son 1978). None of these characters is unique to members of Clinocottus. Several different lin- eages of cottids exhibit reduction in squamation, advanced anuses, and penes. Within the genus Clinocottus, C. embryum and C. acuticeps are grouped together on the basis of a synapomorphic character, a trailing hindgut which extends posterior to the origin of the anal fin. Clinocottus acuticeps larvae are unique in their possession of the autapomorphy, distinct hindgut diverticula. Clinocottus embryum and C. acuticeps larvae also are similar in possessing moderately pointed snouts relative to the blunt, rounded snouts of other Clinocottus larvae, a loose bubble of skin in the head region, and light pigmentation. Synapomorphic characters for clarifying inter- specific relationships among C. analis, C. recal- vus, and C. globiceps were not identified. Never- theless, several pigmentation and morphological characters, for which direction of evolution is not known, do suggest possible relationships among these species. Both Clinocottus globiceps and C. recalvus larvae have intense pigmentation over the snout, head, and nape. Both have very blunt, globose heads, large bulging guts, and relatively deep bodies. Juveniles of the two species are nearly inseparable based on external morphol- ogy. These characters suggest that C. globiceps and C. recalvus may be a very closely related species pair. Postflexion larvae of C. analis differ from all other postflexion Clinocottus larvae in possessing an intense band of melanistic pigment over the lateral surface of the body. Unfortu- nately, the polarity (direction of evolution) of many of the transformation series of morpho- metric and pigment characters could not be de- termined; hence, these characters could not be used in the phylogenetic analysis. Pigmentation and morphometric characters have been useful in several systematic studies based on larvae and have frequently been correlated with other de- rived characters (Johnson 1974; Moser and Ahl- strom 1974; Okiyama 1974; Richardson 1981). Clarification of relationships among the species of Clinocottus must await identification of de- rived characters or a better understanding of the evolution of pigmentation and body shape with- in cottid larvae. The relationships among species of Clinocottus postulated by Bolin (1947) are in close agreement with relationships suggested by larval characters. Bolin placed C. acuticeps in its own subgenus because of its unique possession of a modified penis with a tri-lobed tip, and a membrane con- necting the innermost pelvic fin ray with the ab- domen. Bolin also placed C. analis in its own subgenus because of the retention of minute prickles covering the body. All other members of the genus are scaleless. Clinocottus embryum, C. globiceps, and C. recalvus were placed in the same subgenus because of their large, rounded heads and the retention of a pore behind the last gill. The latter character is plesiomorphic and, hence, of little value for evaluating relationships. Bolin (1947:163) described C. recalvus and C. 172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 globiceps as ‘‘two extremely closely related species” because of their hemispherical head shape, an increased number of cirri on the head, and a pair of lateral knobs near the tip of the penis. Another group of species within the lineage having multiple preopercular spines consists of Oligocottus maculosus and O. snyderi. They share two synapomorphies—a bubble of skin at the nape, and dorsal gut bumps. Larval O. rimensis and O. rubellio are not yet identified and, there- fore, it is not known if these larvae also possess the synapomorphic characters binding O. snyderi and O. maculosus together. Bolin (1934, 1947) defined the genus Oligo- cottus on the basis of the following adult char- acters: absence of scales; presence ofa long, slen- der, simple penis; and modification of the anterior anal fin rays in males. Only the last character appears to be unique to members of this genus. As mentioned above, evolution of penes and loss of scales have occurred in several diverse cottid genera. Bolin placed O. maculosus, O. snyderi, and O. rubellio in the same subgenus because of the greatly modified anal fin in males, a per- manently external penis, and loss ofall but lateral line scales. He further speculated that O. mac- ulosus was the least specialized member of the subgenus and O. rubellio was the most special- ized. Larval O. snyderi possess the derived, auta- pomorphic characters of multiple prickles cov- ering the parietal and posttemporal regions of the head. In addition, larval O. snyderi possess an accessory spine at the anterior base of most of the main spines on the posterior margin of the preopercle. Both of these conditions are unique specializations of larval O. snyderi. Clarification of relationships of other Oligocottus species must await identification of larval O. rimensis and O. rubellio. Larval characters indicate that while Oligo- cottus and Clinocottus are each a monophyletic group, they are closely related. Larvae of both genera are linked together into a higher-level monophyletic unit by the possession of a dis- tinctive preopercular spine pattern. Taranets (1941) also concluded that Oligocot- tus and Clinocottus are closely related. He placed both genera in the supragenus Oligocottini be- cause of the presence of a penis and the absence of bony plates in both groups. Artedius fenestralis, A. harringtoni, A. later- alis, and A. Type 3 form the third group of larvae with multiple preopercular spines. These larvae share one synapomorphy, an Artedius-type pre- opercular spine pattern. In addition, larvae of this group possess distinctive pigment slashes on the ventral midline posterior to the anus and a strongly humped appearance in the nape region. Known larvae of Oligocottus and Clinocottus do not possess either of these characters; however, larvae of several species of Icelinus and Myox- ocephalus do possess similar pigment slashes on the ventral midline. Although these characters are not unique to Artedius species, they provide additional support for the cohesiveness of this group. Within the Artedius group, A. fenestralis, A. lateralis, and A. Type 3 form a distinct subgroup. Larvae of these species share one synapomorphic character, dorsal gut diverticula. Characters identified in this study do not define relation- ships among these three species. Artedius har- ringtoni is probably less specialized than the three species possessing gut diverticula. Artedius har- ringtoni is further distinguished from other Ar- tedius larvae by the possession of seven bran- chiostegal rays. All other cottid larvae examined in this study possess six branchiostegal rays ex- cept several laboratory-reared Clinocottus glo- biceps. Although seven rays are probably a prim- itive character in scorpaeniforms, A. harringtoni appears to have secondarily derived this condi- tion, since none of the outgroup cottids have seven branchiostegals. This grouping of Artedius larvae with multiple preopercular spines corresponds to Taranets’s (1941) classification. He placed Artedius coral- linus, A. fenestralis, A. harringtoni, A. lateralis, and A. notospilotus together in the supragenus Artediini in the subfamily Oligocottinae. Other workers have placed all species of Ar- tedius in the same subfamily and genus (Hubbs 1926; Bolin 1934, 1947; Rosenblatt and Wilkie 1963; Howe and Richardson 1978). Bolin (1947: 161) included A. creaseri in Artedius because of the retention of hemilepidotid-like scales ‘in various degrees of reduction,” large head, an un- advanced anus, and “‘normal structure of the pel- vic fins.” Artedius meanyi was not reported to occur off California at the time of Bolin’s work. Hence, he did not include this species in his classification. WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 173 Rosenblatt and Wilkie (1963) described 4. meanyi as being extremely similar to A. creaseri and placed it in Bolin’s subgenus Ruscariops along with A. creaseri. Reduction in squamation and number of pel- vic fin rays is found in many different cottid genera and appears to have evolved separately several times. Icelinus, Chitonotus, and Ortho- nopias also possess hemilepidotid-like scales in various degrees of reduction. Several species of Icelinus, Orthonopias, and Chitonotus possess large heads and unadvanced anuses. Characters of the larvae indicate that Artedius creaseri and A. meanyi form a distinct grouping separate from the other species of Artedius. Two synapomorphic characters, a pointed snout and basal preopercular spines provide strong evi- dence that A. creaseri, A. meanyi, and Icelinus form a monophyletic group. In addition, 4. meanyi, A. creaseri, and Icelinus larvae are very similar in other pigmentation, morphometric, and spination characters, giving further support for the cohesiveness of this group. In a phenetic study of larval cottids, Richardson (1981) also placed A, meanyi in a group with Chitonotus, Parice- linus, Triglops, and Icelus. (A. meanyi was mis- identified as Icelinus spp. in Richardson’s study. See literature section of A. meanyi description.) Although her study was based on similarities of the larvae and not synapomorphies, it supports the grouping of 4. meanyi and A. creaseri with Icelinus. Both phenetic and synapomorphic characters of the larvae provide strong evidence that the genus 4rtedius (as defined by Bolin 1934, 1947) is not monophyletic and that A. creaseri and A. meanyi should be placed separately. Artedius meanyi and A. creaseri appear to be more closely related to species of Jcelinus and other members of Richardson’s Group 2 than to other species of Artedius. Clarification of relationships among A. meanyi and A. creaseri must await a reex- amination of characters of adult Artedius. Although larvae of the A. meanyi-creaseri group and the Artedius-Clinocottus-Oligocottus group are distinct from one another, they share certain similarities in comparison to other cottid larvae. Both groups have similar pigment pat- terns, morphology, and meristics, suggesting that species of these two groups share a common ancestor. Bolin (1947:159) also speculated that Icelinus, Chitonotus, Artedius, Clinocottus, and Oligocottus constitute a single evolutionary line within the Cottidae. He suggested that “certain details of the more primitive members, partic- ularly the scales, indicate that while these forms undoubtedly did not spring from the modern ge- nus Hemilepidotus, they shared a common and not particularly remote ancestor with the fishes of that genus.” Although characters of the larvae do not exclude the possibility of a hemilepidotid- type ancestor, they do indicate that it would be a relatively distant ancestor. Larval Hemilepi- dotus differ markedly from larvae of Artedius, Clinocottus, Oligocottus, and Icelinus in many characters including meristics, morphometrics, osteology, spination, and pigmentation. It is much more likely that the ancestor of this group pos- sessed characteristics similar to both the Arte- dius-Icelinus group and the Artedius-Clinocot- tus-Oligocottus group. Larvae of at least one species of Jcelinus and several species of Myox- ocephalus possess a fifth or sixth accessory pre- opercular spine. Larvae of Myoxocephalus also possess two distinct patterns of pigment: one type is lightly pigmented similar to the two Artedius groups, whereas the other has intense bands of lateral pigmentation. An ancestor similar to Jce- linus or Myoxocephalus may well have given rise to Artedius, Clinocottus, Oligocottus, and Iceli- nus. This hypothesis is supported by the presence of one or two accessory preopercular spines in Myoxocephalus larvae. This preopercular spine condition appears to be intermediate between the primitive pattern of four preopercular spines and the derived pattern of multiple preopercular spines. Hence, larvae of the ancestor of Artedius, Clinocottus, and Oligocottus were probably rel- atively lightly pigmented with melanophores present on the head, nape, dorsal surface of the gut, and along the ventral midline posterior to the anus. In addition, the ancestral larvae prob- ably possessed four large preopercular spines with one accessory spine on the inner preopercular shelf, two parietal spines, and three posttempor- al-supracleithral spines. In summary, the hypotheses of relationships between Artedius, Clinocottus, and Oligocottus based on larvae characters is in general agree- ment with previous classifications based on adult characters. Synapomorphic characters of the lar- vae provide strong evidence that Clinocottus, Oligocottus maculosus, O. snyderi, A. fenestralis, A. harringtoni, A. lateralis, and A. Type 3 form 174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 a monophyletic group within the cottids. Within this group, the genera Clinocottus and Oligocot- tus are very closely related; however, each genus appears to be monophyletic. Artedius fenestralis, A. harringtoni, A. lateralis, and A. Type 3 also form a monophyletic species group closely re- lated to Clinocottus and Oligocottus. However, synapomorphic characters of the larvae provide strong evidence that A. creaseri and A. meanyi are more closely related to Jcelinus than to species of Clinocottus, Oligocottus, and other Artedius. The genus Artedius as defined by Bolin (1934, 1947) does not appear to be monophyletic; A. meanyi and A. creaseri should be placed sepa- rately. Clarification of the exact position of these two species in relation to Jcelinus and Myoxo- cephalus and the Artedius-Clinocottus-Oligocot- tus group must await identification and exami- nation of larvae of additional species of cottids and reexamination of adult characters. TAXONOMIC DESCRIPTIONS Larvae of Artedius, Clinocottus, and Oligocot- tus have been difficult to identify at both the specific and generic levels because of their strik- ing similarities. Previous descriptions of larvae of this group have been inadequate to separate larvae at both the specific and generic levels be- cause of inaccuracies or insufficient detail. Many of the diagnostic characters useful in separating these larvae are transient features which are pres- ent during only a part of larval development (e.g., head spines, nape bubble). Hence, frequently a combination of several characters is necessary for identification of the larvae. Therefore, to fa- cilitate identification, larval descriptions are ar- ranged in species groups formed by the shared presence of diagnostic characters (Table 3). This matrix table is based on a set of characters that will allow identification of the early life history stages of 13 species of Artedius, Clinocottus, and Oligocottus. Larvae of Artedius fenestralis, A. harringtoni, A. lateralis, A. Type 3, Oligocottus maculosus, O. snyderi, Clinocottus acuticeps, C. analis, C. embryum, C. globiceps, and C. recalvus (Groups A, B, and C) are very similar in morphology, pigmentation, and spination. They are all rela- tively lightly pigmented with melanophores pres- ent on the nape, dorsolateral surface of the gut, and in a series on the ventral midline of the tail. Presence and amount of head pigmentation var- ies within the group. All of these larvae possess blunt, rounded snouts, stubby bodies, and a bulg- ing gut which trails somewhat below the rest of the body. These larvae are readily distinguished from all other known cottid larvae by the pres- ence of multiple preopercular spines (> 5). Larvae in Group A, Artedius fenestralis, A. harringtoni, A. lateralis, and A. Type 3, all have a distinctively stubby shape, a rounded snout, and a humped appearance in the nape region. They are further distinguished by a series of ven- tral midline melanophores posterior to the anus that extend onto the ventral finfold as charac- teristic pigment slashes in flexion and postflexion larvae. These Artedius larvae possess distinctive preopercular spination; postflexion larvae have a relatively high number (=14) of preopercular spines. The dorsalmost, middle, and ventralmost spines are larger than the other spines creating the “Artedius” spine pattern unique to larvae of this group. Characters such as number of pre- opercular spines, number of ventral midline me- lanophores, size at formation of head pigmen- tation, presence of gut diverticula, and number of branchiostegal rays distinguish larvae of each species of Artedius. Larvae in Group B are Oligocottus maculosus and O. snyderi. These larvae can be distin- guished by the presence of a distinctive bubble of skin situated just anterior to the origin of the dorsal finfold in preflexion and early flexion lar- vae. Larvae of both species are more slender than larvae in Groups A and C and have a relatively short, compact gut. In contrast to larvae of Group A, the dorsalmost preopercular spine becomes larger than other spines in flexion and postflexion larvae. Characters useful in distinguishing larvae of the two species of Oligocottus are number and position of ventral midline melanophores, num- ber of preopercular spines, number of parietal spines or prickles, and presence of melanophores on the nape bubble. Group C includes Clinocottus acuticeps, C. an- alis, C. embryum, and C. globiceps. Larvae of C. recalvus, described by Morris (1951), also belong to Group C based on morphology. This is the least cohesive group in that larvae vary more in morphology and pigmentation than in the other groups. In general, larvae have a long gut, the posteriormost portion of which trails below the rest of the body. Larvae of all species except C. 175 WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS “BAIT UOTXayIsod uy # “TS61 SIO UO AT[eIUed paseg || ‘soroydourjaw! 9¢-97 ary eIQUIN]OD Ysnug Woy seAJE] pores ‘soroydourjau Q7—p] 2APY UOZIICO puke eIWIOJITED WO 9BAIR] Pareay § ‘ysadie] auids 1ejnosedoaid 1add¢ + ‘ysadie] sautds sejnoiadoaid Jamo] pue ‘ajpprur ‘saddyq | ‘quawdojaaap jo 1Ys1ay 1B JAQUINN y a BE a Re = ae a + fe = = = = 9 €I-8 = = P iMuvau snipapy = = = + + — — - 9 W=L — _ v MASDIAI SNIPAV “ = = + + + + = = = 9 7-1 a = GIss \|snaypoaa snnosound - - + + + + - - - L109 8 + = 61-91 sdanigojs snyroz0urD - - + + = = = = = 9 17ST a = PI-€l wnduquia snjjoz0uN + = a ar 4h iP = = = 9 CC-91 + = 11-6 SyouD snijoooul|) - = aD 4p + 4F 4b = = 9 OI-t + ae eI-Il Sdaaynap snyosoury) “D - - + =r =r = = + = 9 L-€ it = 77-81 uapdus snj10908110 = — fe ne + = = at = 9 §9€-91 + ~_ 11-6 SNSOJNIDU SNOIOBYO “F . ~ - . - # 9 £1-6 = a b7-T7Z € adh] smipaup = = + + qP = = = te 9 1€-@ = + 9I-F1 S1]D4ajD] SNIpaLy - ~ + - - - - = + 9 6I-€1 = 4 7-81 MopBULDY SNIpay - - + ~ - - - - = L €7-1Z = +r 7-81 sypaisauaf snipaupy “WY ee a a ae eee #iusWdId yuo quouwl uolxey uOIxe4 uoIKxeay = RN = Bqqnq) = ae[NNN syedoqs soioyd fuia {ur} ssoutds exe] [esa -aid -aid -180q “Nq -OIMZAIP aden -I0AIP = -o1ourig -ourjsu -jed -jed aejnd eT Ploy aden juswaid pesy in3puryy inp OUI[prur auids ouids -1ado0a1g “uly ; yen1ua A, «SNN0D snip -Ouly),, -aldV,, ——— a a ee ee ee ee ee ee “SHALOVUAVHD OLLSONOVIG NIVLYAD NO Gasvg AVAUVT] SALLOIOIITE ANY ‘SALLODONIT) ‘SAIGTLYp 40 SONIGNOULD) “¢ ATAV] 176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 6. Larvae of Artedius fenestralis: A) 3.0 mm NL, B) 3.0 mm NL, C) 4.7 mm NL, D) 6.0 mm NL (from Richardson and Washington 1980). WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 177 WN BZ Za EB FS ss = \\ Ficure 7. 1980). embryum have melanistic pigmentation on the head and nape. The dorsalmost preopercular spine is larger than other preopercular spines in postflexion larvae. Characters such as number of preopercular spines, number and spacing of ven- tral midline melanophores posterior to the anus, and presence of hindgut diverticula or bulges are useful in separating larvae of each of these Cli- nocottus species. Group D consists of Artedius creaseri and A. meanyi. These larvae differ from all other larvae of Artedius, Clinocottus, and Oligocottus species listed above in morphology, pigmentation, and spination. They have pointed snouts and large Larvae of Artedius fenestralis: A) 7.2 mm SL, B) 9.9 mm SL, C) 11.8 mm SL (from Richardson and Washington heads, light pigmentation, and four preopercular spines. These characters bind them more closely with Jcelinus larvae. In addition, A. creaseri and A. meanyi larvae are further distinguished by large blotch-like melanophores situated along the ventral midline posterior to the anus. Snout to anus length, meristics, finfold pigmentation, and nape pigmentation are useful characters in sep- arating larvae of the two species. Artedius fenestralis (Figures 6-8; Table 4) LITERATURE. — Blackburn (1973) illustrated an 8.5 mm SL larva similar to Artedius fenestralis, 178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 8. Juvenile of Artedius fenestralis, 19.1 mm SL. which he described as Cottid 4. Eldridge (1970) and White (1977) briefly described and illus- trated 3.2 mm and 3.9 mm larvae, respectively, which are similar to A. fenestralis. These illus- trations also closely resemble A. /ateralis larvae. Richardson and Pearcy (1977) listed these larvae as Artedius sp. 2. Richardson and Washington (1980) described and illustrated specimens 3.0, 4.7, 6.0, 7.2, 9.9, and 11.8 mm long as Artedius Type 2. IDENTIFICATION.—Juveniles and adults were identified by the following combination of char- acters: high dorsal fin ray counts, absence of nasal and preorbital cirri, and the presence of scales on the head under the entire orbit and in a dense patch on the caudal peduncle. The develop- mental series was linked together primarily by pigmentation, body shape, gut diverticula, and preopercular and parietal spination. Identifica- tion of larvae was further confirmed through comparison with larvae reared from known eggs. Postflexion and transforming larvae were linked with juveniles using pigmentation, cirri patterns, spination, and meristics. DISTINGUISHING FEATURES.—A combination of characters is useful in distinguishing preflexion A, fenestralis larvae including prominent gut di- verticula protruding from the dorsal surface of the abdominal cavity, melanistic nape pigmen- tation, lack of head melanophores, and a series of 13-19 ventral midline melanophores poste- rior to the anus. Late flexion and postflexion larvae are further distinguished by the presence of 18-22 preoper- cular spines with the dorsalmost, middle, and ventralmost spines being larger than the others. Postflexion larvae also have a cluster of 5 or 6 spines situated on the posterior margin of each parietal bone. Juveniles of A. fenestralis are distinguished by meristics, dark pigmentation over the dorsolat- eral surface of the body, and 13-16 ventral mid- line melanophores posterior to the anus. Other useful characters include the absence of a nasal and preorbital cirrus, the presence of one or two small cirri on the eyeball, and two frontoparietal cirri. PIGMENTATION. — Newly hatched larval Arte- dius fenestralis reared in the laboratory have no melanistic pigmentation on the head or nape. Intense melanophores are scattered over the dor- solateral surface of the gut. These lateral gut melanophores are frequently faded and difficult to see in field-collected larvae. Posterior to the anus, a series of 13-19 melanophores originates under the third or fourth postanal myomere and extends posteriorly along the ventral body mid- line. An additional 1 or 2 melanophores extend onto the ventral finfold near the notochord tip. These ventral midline melanophores are evenly spaced approximately one every other myomere. During larval development, the head region remains unpigmented. Two to four melano- phores are added on the nape in larvae 3.4 mm long and become embedded in musculature over the notochord by ~7 mm. By that size the pos- terior half of the series of ventral midline me- lanophores appear as distinctive slashes that ex- tend onto the ventral finfold. During transformation (planktonic specimens ~ 12-14 mm long) juvenile pigmentation begins to develop. Melanophores are added on the dor- sal surface of the head, on the tip of the lower jaw, and on the pectoral fin base. Gradually, me- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 179 TasLe 4. Bopy Proportions OF LARVAE AND JUVENILES OF ARTEDIUS FENESTRALIS, A. HARRINGTONI, A, LATERALIS, AND A. Tyre 3. Values given are percent standard length (SL) or head length (HL) including mean, standard deviation, and range in parentheses. Item Artedius fenestralis Artedius harringtoni Artedius lateralis Artedius Type 3 Head length/SL: Preflexion 22.2 + 1.41 (21.1-25.0) 20.9 + 2.30(18.7-22.3) 20.2 + 2.68 (16.4-22.9) 21.8 + 0.92 (20.7-23.2) Flexion 22.1 + 1.22 (20.3-23.3) 23.1 + 3.86 (24.9-27.0) 23.6 + 1.92(19.9-26.1) 24.6 + 1.71 (23.3-27.1) Postflexion 26.1 + 2.85 (20.8-29.4) 29.0 + 0.50 (22.4-34.1) 26.9 + 1.89 (24.3-30.8) 29.0* Juvenile 33.9 + 2.08 (32.0-34.9) 33.6 + 3.46 (30.4-36.5) 22.7 + 3.21 (19.1-25.3) - Snout length/HL: Preflexion 15.1 + 3.77 (10.5-20.0) 18.9 + 5.52 (9.9-25.1) 28.0 + 1.83 (25.8-30.1) 19.8 + 3.85 (15.4-24.6) Flexion 20.4 + 5.96 (12.3-26.6) 23.7 + 6.14(18.3-19.1) 25.2 + 1.91 (22.9-29.4) 23.2 + 3.51 (19.2-26.7) Postflexion 21.8 + 3.57 (17.5-24.5) 24.2 + 4.19 (15.8-31.4) 24.7 + 3.77 (19.4-30.6) 29.4* Juvenile 21.6 + 2.08 (19.8-23.2) 23.1 + 1.00 (22.3-24.3) 23.0 + 1.73 (18.9-24.7) - Eye diameter/HL: Preflexion 39.7 + 4.32 (33.7-46.9) 44.7 + 4.57 (39.7-50.1) 44.7 + 2.87 (40.8-47.1) 46.7 + 6.53 (39.0-57.6) Flexion 41.2 + 3.42 (37.4-46.5) 40.4 + 7.33 (38.4-43.2) 40.4 + 2.88 (37.4-45.5) 37.1 + 2.16 (35.3-40.2) Postflexion 36.9 + 7.10 (27.3-52.1) 32.6 + 4.09 (23.2-39.8) 36.6 + 4.06 (37.9-44.6) | 37.9* Juvenile 29.1 + 1.00 (24.6-27.3) 31.2 + 5.77 (29.8-31.2) 35.3 + 5.03 (22.1-24.8) ~ Snout to anus length/SL: Preflexion 45.3 + 3.59 (41.2-50.8) 42.0 + 3.21 (38.2-48.2) 40.4 + 4.83 (33.3-45.6) 44.8 + 2.76 (41.4-49.8) Flexion 45.9 + 2.49 (44.5-50.1) 47.3 + 8.30 (36.5-55.9) 42.7 + 3.65 (38.2-47.9) 45.1 + 2.63 (43.2-48.1) Postflexion 48.4 + 2.26 (42.7-51.4) 49.9 + 2.67 (45.9-55.4) 48.4 + 2.18 (44.4-50.9) 50.2* Juvenile 49.1 + 2.00 (47.0-51.5) 47.4 + 3.21 (43.1-49.2) 48.7 + 4.04 (44.1-51.2) - Snout to pelvic fin origin/SL: Preflexion - - - - Flexion 25.6 + 3.21 (20.2-29.8) - - 25.3 + 4.24 (21.9-27.6) Postflexion 26.1 + 3.21 (20.2-29.8) 27.8 + 1.72 (24.5-30.8) 28.1 + 3.11 (24.9-34.1) 29.3* Juvenile 30.4 + 3.51 (26.5-30.1) 27.4 + 2.65 (24.1-29.4) 27.0 + 3.61 (23.8-31.0) - Pelvic fin origin to anus/SL: Preflexion - - - - Flexion 22.2 + 5.35 (16.1-27.0) - - 23.1 + 4.95 (19.1-26.1) Postflexion 26.1 + 3.21 (20.2-29.8) 22.8 + 4.68 (17.2-34.8) 20.1 + 4.20(14.4-26.7) 20.9* Juvenile 18.9 + 1.15 (18.2-20.2) 19.9 + 1.15 (18.9-21.3) 22.3 + 2.08 (20.3-24.1) - Body depth at pectoral fin base/SL: Preflexion 21.7 + 1.81 (19.0-23.8) 23.7 + 2.99 (20.9-29.7) 23.0 + 3.16 (17.9-26.3) 25.9 + 1.77 (23.4-28.2) Flexion 28.5 + 2.70 (24.9-30.1) 28.1 + 2.45 (28.0-31.2) 26.6 + 2.33 (24.2-27.1) 28.2 + 1.76 (26.3-30.1) Postflexion 28.2 + 1.82 (24.2-32.1) 30.5 + 2.74 (23.4-34.1) 28.1 + 2.29 (24.1-32.2) 30.0* Juvenile 25.8 + 1.53 (24.4-26.8) 22.3 + 4.15 (24.6-25.2) 19.7 + 3.06 (21.9-25.0) = Body depth at anus/SL: Preflexion 19.0 + 2.49 (14.8-22.2) 27.8 + 2.45 (17.2-27.6) 20.4 + 2.51 (15.8-21.8) 22.9 + 3.28 (18.2-27.2) Flexion 24.1 + 2.59 (23.6-27.1) 30.4 + 3.32 (28.1-34.7) 26.3 + 2.29 (24.5-31.3) 29.1 + 2.06 (26.1-31.3) Postflexion 27.9 + 2.79 (21.4-32.8) 20.9 + 5.77 (23.9-34.8) 26.3 + 2.82 (22.4-33.0) 30.3* Juvenile 21.6 + 2.65 (18.9-24.4) 28.0 + 1.72 (20.8-21.9) 30.3 + 3.51 (17.4-22.8) - Pectoral fin length/SL: Preflexion 9.1 + 0.98 (8.2-10.9) 7.4 + 2.34 (4.4-11.1) 11.0 + 1.08 (9.6-12.2) 10.4 + 2.91 (7.1-13.3) Flexion 9.4 + 0.71 (8.4-10.4) 12.1 + 3.54 (9.9-15.4) 12.3 + 1.98 (9.6-15.9) 11.2 + 3.50 (7.4-15.1) Postflexion 22.6 + 6.21 (9.7-29.7) 24.6 + 7.59 (10.3-34.6) 19.8 + 4.66 (13.3-27.6) 21.0* Juvenile 26.2 + 1.53 (27.1-28.4) 34.1 + 3.61 (29.9-36.6) 30.3 + 3.51 (26.8-34.2) - — = Not present at this stage. * = Only one specimen available in this stage. 180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 lanophores develop on the anteriormost portion of the spinous dorsal fin, then extend ventrally as a band of pigment stretching from the fourth or fifth dorsal spine to the pigmentation over the dorsal surface of the gut just posterior to the pectoral fin base. Juvenile pigmentation increases markedly in newly settled individuals 13 mm SL. Numerous melanophores are added over the dorsolateral surface of the head and become concentrated in the parietal-interorbital region. Additional me- lanophores extend down onto the snout and lips. Laterally, melanophores are added in the cheek region between the eye and the preopercle and the dorsal portion of the opercle. Several mela- nophores are clustered at the posterior edge of the lower jaw. The ventral surface of the head remains unpigmented. Pigmentation gradually extends from the head posteriorly across the dor- solateral surface of the body until it fuses with bands of pigment reaching from the middle of the spinous dorsal fin to the gut. Pigmentation increases on the anterior end of the dorsal fin creating a dark blotch of pigment across the first four dorsal spines. Melanistic pigmentation also increases on the pectoral fin base with melano- phores extending onto the pectoral fin rays and eventually forming several bands of pigmenta- tion. Several irregular clusters of melanophores appear along the lateral midline and gradually form a band of pigment reaching from the gut to the caudal peduncle. As juvenile pigmentation develops, saddles of pigment form along the dorsum in an anterior to posterior sequence. The first saddle or band of pigment forms under the 4th-7th dorsal fin rays. Gradually, melanophores extend ventrally from the pigment saddle and merge with the lat- eral midline melanophores. Concurrently, a sec- ond saddle of pigment forms under the 9th-10th dorsal fin rays, while a third saddle of pigment begins to develop under the 13th—1 5th dorsal fin rays. Melanophores from these pigment saddles also extend ventrally and fuse with the lateral midline pigment. At the same time, melano- phores are added on the dorsal fin forming three to four bands. Melanophores extend ventrally from the lateral midline band and form a series of five to eight scallops which reach just below the lateral midline. The rest of the ventrolateral surface of the body remains characteristically un- pigmented until juveniles reach about 19-20 mm. As the dorsal pigment saddles are forming, the lateral midline melanophores extend posteriorly to the base of the caudal fin where they form a dark band. Gradually, melanophores extend onto the caudal fin rays forming three or five indistinct bands of pigment. Approximately 13-16 ventral midline melanophores remain visible in juve- niles up to ~20 mm long. MorpuHo.ocy.—Larvae of Artedius fenestralis hatch at ~3.5-3.8 mm NL. Flexion of the no- tochord occurs between 5.9 and 6.8 mm NL. The largest planktonic larva collected is 13.9 mm and is beginning to undergo transformation. The smallest benthic juvenile examined is 13.1 mm. Thirty-four selected specimens, 3.2-21.2 mm, were examined for developmental morphology. Larval A. fenestralis have stubby bodies with a humped appearance in the nape region. Dis- tinctive diverticula extend dorsolaterally from the dorsal surface of the gut just posterior to the origin of the pectoral fin base. These diverticula are present in newly hatched larvae and remain prominent in the largest planktonic larvae. The diverticula completely disappear in benthic ju- veniles shortly after settling. The gut itself is moderately long and the posterior portion of the hindgut trails well below the rest of the body. Snout to anus length increases from 43% to 45% SL during larval development, then increases to 49% SL in benthic juveniles. Artedius fenestralis larvae have a short, rounded snout with snout length increasing from 15% HL in preflexion lar- vae to 23% HL in postflexion larvae and juve- niles. FIN DEVELOPMENT. —Caudal fin rays begin to form at ~6 mm. The adult complement of prin- cipal caudal rays is present in larvae ~7 mm long. The bases of the dorsal and anal fin rays appear in 7—-7.5 mm larvae. The full complement of fin rays is formed by ~8.5-9 mm. Dorsal fin spines begin to form at ~8 mm, and the full complement of spines (VIII-IX) is present by ~9.5 mm. Although pectoral fin rays are visible by ~7 mm, the adult complement (14-16) is not formed until ~9 mm. Pelvic buds form between 6.5 and 7 mm and the adult complement of I,3 pelvic fin rays is formed in larvae ~ 10 mm long. SPINATION. —Seven to 13 tiny spines begin to form along the posterior margin of the preopercle in larvae ~4.7 mm NL. The preopercle appears to develop in two arc-shaped sections, which overlap slightly at the angle of the preopercle. Three to 7 spines are present along the dorsal- most section and 6-8 spines occur on the lower WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 181 Ficure 9. Larvae of Artedius harringtoni: A) 3.0 mm NL, B) 4.7 mm NL, C) 6.9 mm NL (from Richardson and Washington 1980). section. The two sections fuse together in post- flexion 7 mm long. Two spines located at the site of fusion begin to increase in length relative to the other preopercular spines. Concurrently, the preopercular spines increase in number during larval development and range between 18 and 21 in larvae >8 mm long. The dorsalmost and middle two spines continue to increase in size relative to the other spines, becoming nearly three times as long. The ventralmost 2 or 3 spines also increase in size, becoming 1.5 to 2 times as long as the other spines. The number of preopercular 182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 spines decreases in transforming larvae > 13 mm long. The smaller spines (4-6, 8-10, and 13-15) begin to regress first. Newly settled juveniles pos- sess one large dorsal preopercular spine. The low- er spines are visible only as serration, or bumps, on the preopercular margin. The dorsalmost spine continues to increase in size while the lower bumps eventually disappear in juveniles ~ 19 mm long. Clusters of spines also develop in the parietal and supracleithral-posttemporal regions. One or two spines form at the posterior end of the pa- rietal in larvae ~6 or 7 mm long. A third spine is added in 7-8-mm larvae. Larvae > 8 mm have four to six spines located in two rows on each side of the head. Usually, three spines occur in the anterior row while two or three spines are present in a second row posterior to and parallel to the first row. These spines begin to regress in size in transforming larvae >12-13 mm long. The anterior spines curve posteriorly, eventually fusing with spines from the posterior row form- ing a hollow arch and canal. This canal develops into part of the cranial lateral line system in ju- veniles ~ 14-15 mm long. Two small spines develop on the ventral por- tion of the posttemporal in larvae 6-7 mm long. A third spine is added on the posttemporal in larvae >8 mm long. Concurrently, another spine develops on the dorsal tip of the supracleithrum. These spines remain prominent in planktonic larvae <12 mm long; however, in transforming juveniles the spines gradually curve dorsally and ventrally and fuse together forming a bony tube or canal. This canal becomes the anteriormost juncture of the lateral line and cephalic lateral line systems in juveniles. Artedius harringtoni (Figures 9-11; Table 4) LITERATURE.— Blackburn (1973) described a 4.6 mm larva that he called Cottid 6 that is sim- ilar to A. harringtoni. Richardson and Washing- ton (1980) illustrated and described specimens 3.0, 4.7, 6.9, 7.3, 9.3, and 13.6 mm long. IDENTIFICATION. —Juveniles and adults were identified primarily on the basis of the following characters: high dorsal fin ray counts (16-18), low pectoral fin ray counts (usually 14), presence of seven branchiostegals, presence of a preorbital cirrus, scales extending onto the head under only the posterior portion of the orbit, and scales ab- sent on the snout. The developmental series of larvae was linked together by pigmentation, pre- opercular spination, absence of gut diverticula, body shape, and the possession of seven bran- chiostegals. Postflexion and transforming larvae were linked with juveniles primarily on the basis of pigmentation, meristics, and presence of a preorbital cirrus. DISTINGUISHING FEATURES.—Characters use- ful in distinguishing small larval 4. harringtoni are a combination of presence of melanistic nape pigment, lack of head pigmentation, a series of 21-33 pigment slashes along the ventral midline of the tail, and a humped appearance in the nape region. Absence of dorsal gut diverticula distin- guishes larval 4. harringtoni from similarly pig- mented larvae of A. /ateralis, A. fenestralis, and A. Type 3. Postflexion larvae 6.5 mm are distinguished by the presence of 18-22 spines along the pos- terior margin of the preopercle. The dorsalmost and middle preopercular spines are characteris- tically larger than the other spines. Larvae >7 mm have seven branchiostegal rays. Larvae of all other species of Artedius have only six bran- chiostegal rays. Juvenile A. harringtoni may be recognized by the dark pigmentation over the head and nape, possession of seven branchiostegals, retention of 18-22 ventral midline melanophores, possession of a preorbital cirrus, and dorsal and pectoral fin ray counts. PIGMENTATION. —Preflexion larvae have no melanistic pigmentation on the head; however, 3-5 small, external melanophores are concen- trated in a dense patch on the nape. The dor- solateral surface of the gut is covered with nu- merous large, intense melanophores. One to 8 tiny melanophores encircle the anus. Posterior to the anus, the only pigmentation consists of a series of 23-33 melanophores positioned along the ventral midline. This series originates under the first to third postanal myomere and extends posteriorly toward the tail tip with 1 or 2 me- lanophores positioned under each myomere. An additional | to 3 melanophores frequently occur on the caudal finfold near the tail tip. During larval development the head region re- mains unpigmented. The nape melanophores be- come embedded in the musculature over the no- tochord in larvae >7 mm. Concurrently, the number of ventral midline melanophores de- creases to between 21 and 30, and the posterior WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 183 At Wa “ Ficure 10. Larvae of Artedius harringtoni: A) 7.3 mm SL, B) 9.3 mm SL, C) 13.6 mm SL (from Richardson and Washington 1980). half of the series appears as characteristic pig- ment slashes that extend onto the ventral finfold. During transformation, planktonic larvae > 10 mm begin to develop juvenile pigmentation. Me- lanophores are added on the tip and base of the lower jaw, on the cheek between the eye and the dorsalmost preopercular spine, on the opercu- lum, and on the isthmus. Pigmentation increases markedly over the head in newly settled benthic juveniles. Melanophores develop on the snout and upper lip and on the dorsal surface of the head over the brain. Me- lanophores gradually extend posteriorly from the head and eventually join with the nape pigmen- tation. Concurrently, melanophores extend pos- teroventrally from the posttemporal region to- ward the dorsal gut pigment. Numerous large melanophores form over the base of the pectoral 184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 11. fin, and subsequently extend onto the pectoral fin rays forming four or five distinct pigment bands across the fin. A band of melanophores also extends ventrally from the pectoral fin base and covers the isthmus. In juveniles >13 mm long, the entire head is heavily pigmented. Melanophores extend pos- teriorly from the head to a vertical line under the seventh dorsal fin spine. A dense patch of me- lanophores develops at the anterior end of the spinous dorsal fin forming a dark blotch across the fin membrane between the first four dorsal spines. Scattered melanophores are added along the rest of the dorsal fin eventually forming three or four bands of pigment. Posterior to the head, pigmentation is added in three saddles along the dorsum. The first sad- dle of pigment forms under the 2nd—4th dorsal fin rays, the second saddle forms under the 7th— 10th dorsal fin rays, and the third forms under the 13th—1 5th fin rays. Concurrently, an irregular band of faint melanophores develops along the lateral midline. This lateral pigmentation grad- ually extends posteriorly from the abdominal re- gion to the caudal peduncle. As development proceeds, bands of melanophores extend ven- trally from each of the saddles on the dorsum and merge with the lateral midline pigment. As a result, the dorsolateral surface of the tail is covered by bands of pigmentation, which enclose small unpigmented saddles and circles creating a characteristic pattern. Subsequently, groups of melanophores extend ventrally from the lateral midline pigmentation creating a scalloped edge of pigment along the ventrolateral body surface. Eventually, melano- phores from the tips of each scallop extend lat- erally and join together enclosing four to six dis- tinctive unpigmented circles, characteristic of juvenile A. harringtoni. Juvenile of Artedius harringtoni, 13.9 mm SL. In late stages of juvenile pigmentation, the lat- eral band of melanophores extends posteriorly to the base of the caudal fin. Melanophores are added on the caudal fin rays forming five to seven bands of pigmentation. Between 18 and 21 ventral midline melano- phores remain visible in juveniles <15 mm. MorpPHoLoGy.—The smallest A. harringtoni larva from plankton collections is 3.0 mm NL and still retains remnants of its yolk. Larvae undergo flexion of the notochord between 5.2 and 6.4 mm NL. The largest planktonic larva examined is 13.6 mm long and beginnning to undergo transformation. The smallest benthic juvenile collected in tidepools is 12.9 mm and is just beginning to develop juvenile pigmenta- tion on the head and pectoral fin base. Thirty- five selected specimens, 3.0-13.7 mm, were ex- amined for morphometrics. Larvae of A. harringtoni are stubby with a dis- tinctive humped appearance in the nape region. Unlike larval A. fenestralis, A. lateralis, and A. Type 3, larval A. harringtoni have no dorsal gut diverticula. The gut is moderately long with snout to anus length ranging from 42% in preflexion larvae to 50% SL in postflexion larvae. Relative snout to anus length decreases slightly in benthic juveniles. The hindgut appears to trail below the rest of the body. Relative body depth at the pec- toral fin base increases from 23% in preflexion larvae to 30% in flexion and postflexion larvae. Artedius harringtoni have blunt heads and rounded snouts. Head length increases relative to body length during development, averaging 21% in preflexion larvae, and 34% SL in juve- niles. Snout length increases from 19% to 22% HL during larval development. Fin DEVELOPMENT.—A thickening in the hy- pural region of the developing caudal fin is first visible at 4.7 mm NL, just prior to the onset of WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 185 notochord flexion which occurs at ~5.2 mm NL. Caudal fin rays begin to form in larvae ~6 mm; however, the adult complement of principal cau- dal rays is not complete until larvae reach ~7 mm long. Bases of the dorsal and anal fin rays form in larvae ~6-7 mm long. Dorsal spines begin to form in larvae ~7-8 mm long. The adult com- plement of dorsal and anal fin rays is complete at 9.3 mm. Pectoral fin rays are first visible be- tween 6 and 7 mm, and the adult complement (13-15) is countable at ~7.5 mm. The pelvic fin bud begins to form at ~7.1 mm, and the adult complement of I,3 is complete by ~10 mm. SPINATION.— Eight to ten tiny spines begin to form along the posterior margin of the preopercle in larvae ~4.5 mm NL. The number of spines increases to 18-22 in flexion and postflexion lar- vae. By the end of flexion, ~6.7 mm, the middle two spines (7-9) begin to increase in size relative to the other preopercular spines. In larvae >7.5 mm, the dorsalmost two or three spines also in- crease in size relative to other spines. As devel- opment proceeds, the dorsalmost and middle spines increase in length and diameter creating a characteristic pattern with small, inconspic- uous spines situated between the dorsalmost and middle spines, and ventral to the middle spines. In larvae >8.5 mm, the ventralmost four or five spines also become somewhat larger than the spines directly above them. When larvae reach ~10-11 mm SL, the preopercular spines begin to regress with the small, inconspicuous spines disappearing first. At the onset of transformation (~12-13 mm) only four spines remain in the approximate position of the original spines (i- 2, 4-9, 12-14, and 18-22). In newly settled ju- veniles, the dorsalmost preopercular spine be- comes quite long and stout while the lower three spines gradually become smaller and visible only as slight bumps on the margin of the preopercle. Spines never develop in the parietal and post- temporal region of the head. However, in cleared and stained larvae, bony thickenings are visible in the parietal region at the same position as parietal spines found in other cottid larvae. Artedius lateralis (Figures 12, 13; Table 4) LITERATURE. — Budd (1940) described and il- lustrated a newly hatched larva of A. lateralis 4.1 mm TL. Marliave (1975) described larvae of A. lateralis and illustrated specimens 4 mm TL, 8 mm TL, 11 mm TL, and 14 mm TL long. IDENTIFICATION. — Small larval A. /ateralis were reared from eggs spawned from known adults. Juveniles and adults were identified using the following characters: pigmentation, absence of scales on the head and caudal peduncle, absence of nasal and preorbital cirri, and the presence of 3-11 scales in the longest row in the dorsal scale band. The developmental series was linked to- gether primarily on the basis of pigmentation, preopercular spination, presence of gut divertic- ula, and body shape. Postflexion and transform- ing larvae were linked to juveniles by pigmen- tation, cirri patterns, spination, and meristics. DISTINGUISHING FEATURES.—Characters use- ful in distinguishing preflexion larvae of Artedius lateralis are prominent diverticula which extend dorsolaterally from the dorsal surface of the gut just posterior to the pectoral fin bases, the lack of head and nape pigment in larvae <6 mm NL, and a series of 22-32 melanophores that lie along the ventral midline posterior to the anus. The anterior half of the series is characterized by one large melanophore per myomere while the pos- terior half of the series consists of two or three smaller pigment slashes per myomere. Postflexion larvae of A. /ateralis >6.2 mm can be distinguished from other Artedius larvae by melanistic pigmentation over the brain. Ju- venile A. /ateralis are distinguished by two dark bars of melanophores extending ventrally from the dorsal fins across the lateral surface of the body trunk, the series of 11-21 ventral midline melanophores, and meristics. PIGMENTATION. — Newly hatched larvae of A. lateralis have no melanistic pigmentation on the head or nape. Dense, round melanophores are concentrated over the dorsolateral surface of the gut and extend dorsally onto the gut diverticula. A cluster of 4 to 6 small melanophores sur- rounds the anus. Posterior to the anus, a series of 22-32 melanophores lies along the ventral midline of the body. These melanophores orig- inate under the third or fourth postanal myomere and extend posteriorly toward the tail tip where several additional melanophores extend onto the caudal finfold. Melanophores in the anterior half of this series are relatively large and spaced one per myomere. The posteriormost melanophores appear as small pigment slashes, which extend onto the ventral finfold and are closely spaced two or three to every myomere. During larval development, melanophores form on the dorsal surface of the head in larvae >6.3 mm. Two to five melanophores also form i. on = — 186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 ne tae ep eared cio a a B eed vA eS Ficure 12. Larvae of Artedius lateralis: A) 4.6 mm NL, B) 6.4 mm SL, C) 7.1 mm SL. at the base of the cleithrum and along the ventral midline of the gut in larvae >5.2 mm. These melanophores are arranged in a characteristic T shape with two melanophores positioned as hor- izontal slashes at the base of the cleithrum and one to three melanophores extending posteriorly along the ventral midline of the gut. During transformation, in planktonic larvae >8 mm, melanistic pigmentation increases markedly on the dorsal surface of the head with 33-44 dark melanophores covering the brain. Melanophores also form just posterior to the lower jaw, on the cheek between the eye and the preopercle and on the operculum. Ventral mid- line melanophores remain unchanged in number and spacing. Pigmentation increases markedly in newly set- tled juveniles > 10 mm long. Dark melanophores form on the dorsolateral surfaces of the head and extend anteriorly onto the snout and upper and lower lips. Several melanophores are added to the gular region beneath the lower jaw. Intense pigment forms on the bases of the pectoral fins and several large melanophores extend onto the pectoral fin rays. Gradually, melanophores from the base of the pectoral fin extend ventrally form- ing a band of pigment across the isthmus. With development, pigmentation increases on the head WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 187 Sa CS "Ea Ficure 13. Young of Artedius lateralis: A) 9.1 mm SL, B) 13.3 mm SL. so that in larvae =>12 mm SL, the entire head is darkly pigmented. Shortly after settling, in larvae between 10 and 11 mm, a patch of melanophores is added to the dorsal fin between the fourth and sixth spines. These melanophores extend ventrally across the dorsum toward the pigmentation on the pectoral fin base. A second band of melanophores forms on the second dorsal fin membrane between the 2nd and 4th fin rays. Gradually this band extends anteroventrally below the lateral midline. With development, the two vertical bands of pigment become very dark and intense. Melanophores from these bands extend dorsally across the dor- sal fins. Concurrently, three saddles of faint me- lanophores are added posteriorly along the dor- sum. The first saddle of pigment forms under the 8th-10th dorsal fin rays, the second saddle is added under the 1 2th—1 5th fin rays, and the third saddle forms on the dorsal surface of the caudal peduncle. Gradually, melanophores from these pigmented saddles extend ventrolaterally and join together forming a band along the lateral mid- line. Lateral pigmentation extends posteriorly and forms a band along the base of the caudal fin. Melanophores extend onto the caudal fin rays, gradually forming two or four distinct bands across the caudal fin. Ventral midline melano- phores decrease in number in juveniles from 11 to 21. These melanophores remain visible in ju- veniles <15 mm long. MorpHo.ocy.—Artedius lateralis larvae are 3.9-4.5 mm long at hatching. Flexion of the no- tochord occurs between 5.0 and 6.3 mm NL. The largest planktonic specimen observed is 9.2 mm and beginning to develop juvenile pigmentation. A. lateralis settle at a relatively small size, ~9.5 to 10.5 mm. Thirty-three specimens (4.1-12.1 mm) were examined for developmental mor- phometrics. Larvae of A. /ateralis are rather stubby with a moderately short gut. Snout to anus length av- erages 40% in preflexion larvae, then increases to 48% SL in postflexion larvae and juveniles. Pronounced diverticula extend dorsally from each side of the gut just posterior to the pectoral fin base. The diverticula are present in newly hatched larvae and remain prominent throughout larval development. Tiny remnants of the diverticula are present in newly settled juveniles between 9 and 10.5 mm long. Larvae of A. /ateralis have a long rounded snout 188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 relative to other Artedius Group A larvae. Snout length decreases from 28% to 24% HL during larval development. Fin DEVELOPMENT. — The notochord begins to flex in larvae ~5 mm long, and is fully flexed in larvae between 5.7 and 6.3 mm. Caudal rays begin to form during flexion in larvae ~5.5 mm; however, the adult complement of 6 + 6 prin- cipal caudal rays is not complete until ~6 mm. Bases of the dorsal and anal fins are first visible in larvae between 6 and 6.5 mm and the adult complement of dorsal (15-17) and anal (12-14) rays is complete by about 7.5-8 mm. Pectoral fin rays begin to form between 7 and 8 mm, but the full complement of rays (14-16) is not formed until larvae are ~8 mm long. Pel- vic fin buds are first visible in a 7.4 mm larva; however, the fin rays are not countable until 9 mm. SPINATION.—In preflexion larvae ~4.5 mm NL 8-9 tiny spines are visible on the posterior mar- gin of the preopercle. As larvae undergo flexion of the notochord, the number of preopercular spines increases to 9-14. By the end of flexion, larval A. /ateralis have 14-16 preopercular spines. In larvae >7 mm, the dorsalmost and middle (spines 6-9 from the top of the preopercle) be- come slightly longer than the other preopercular spines. These spines never become more than 1.5 times larger than the other preopercular spines, in contrast to the situation in larvae of A. harringtoni, A. fenestralis, and A. Type 3 in which the dorsalmost and middle preopercular spines may be nearly 2.5 times larger than the other spines. Preopercular spines begin to regress in transforming specimens >9 mm. The dorsal- most spine increases in size while the lower spines (4-6 and 9-12) decrease in size becoming visible only as small serrations or irregularities on the preopercular margin. Spines 7-8, 12-13, and 16- 18 fuse together to form blunt bumps along the preopercular margin. In juveniles > 13 mm, only the large, dorsalmost spine remains. Transform- ing larvae reared in the laboratory possess 4-5 small spines at the posterior margin of the pa- rietals. These spines are not present in planktonic larvae from field collections, nor are they visible in newly settled juveniles from tidepools. Artedius Type 3 (Figures 14, 15; Table 4) LITERATURE. — Larvae of Artedius Type 3 have not been previously described. IDENTIFICATION.—Only a partial size series (2.9-7.6 mm) of Artedius Type 3 larvae are avail- able, all from California collections. The pres- ence of prominent gut diverticula and the char- acteristic Artedius-type preopercular spine pattern (dorsalmost, middle, and ventralmost spines larger than the others) identifies this larval type as an Artedius. Larvae remain unknown for only two species of Artedius, A. corallinus and A. no- tospilotus. Meristics of the largest larva of Ar- tedius Type 3 coincide with those recorded for both A. corallinus and A. notospilotus. However, pectoral counts fit those of A. notospilotus most closely. The 7.6 mm larval A. Type 3 possesses 16 pectoral fin rays. Ninety % of the A. noto- spilotus examined by Howe and Richardson (1978) possessed 16 pectoral fin rays while only 10% of A. corallinus specimens possessed 16 pec- toral fin rays. Pigmentation along the ventral midline pos- terior to the anus of A. Type 3 larvae (9-13 widely spaced melanophores) coincides most closely with that of juvenile A. corallinus. Several A. coral- linus 13.5-14 mm long, possess 3-6 widely spaced ventral midline melanophores. In con- trast, a 16-mm juvenile A. notospilotus possesses 24 ventral midline melanophores spaced one every one or two myomeres. Adult A. corallinus are common in the inter- tidal areas of the southern California coast where Artedius Type 3 larvae were collected (Miller and Lea 1972). Artedius notospilotus adults are rare in the same area. Additional larger specimens are needed before larvae of Artedius Type 3 can be specifically iden- tified. DIsTINGUISHING FEATURES.—Artedius Type 3 larvae are distinguished as an Artedius by the distinctive diverticula that extend dorsolaterally from the dorsal surface of the gut just posterior to the pectoral fin base. Artedius Type 3 larvae are distinguished from small larvae of A. fenes- tralis, which possess similar diverticula, by the low number (9-13) of ventral midline melano- phores posterior to the anus. Other characters useful in distinguishing small A. Type 3 larvae are absence of head pigmentation and presence of a cluster of 2-4 melanophores in the nape region. Preopercular spines begin to form in lar- vae <4.1 mm NL. Preopercular spines do not form in other Artedius larvae with multiple pre- opercular spines until ~4.5 mm NL. Flexion and postflexion larval Artedius Type 3 possess 21- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 189 Ficure 14. Larvae of Artedius Type 3: A) 3.2 mm NL, B) 4.1 mm NL, C) 4.9 mm NL. 24 preopercular spines, more than other larval Artedius (groups A and D), all of which have <21 preopercular spines. PIGMENTATION.—Small preflexion larvae of Artedius Type 3 possess no melanistic head pig- mentation. Two to four small external melano- phores are clustered on the surface of the nape. Numerous dark, rounded melanophores are con- centrated over the dorsolateral surface of the gut and extend dorsally onto the gut diverticula. One to four small melanophores are clustered around the anus. Posterior to the abdominal cavity, the only pigmentation consists of a series of 9-13 mela- nophores located along the ventral midline. This series of melanophores originates under the third to fourth postanal myomere and extends poste- riorly toward the tail tip. Each melanophore is 190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 15. spaced approximately two to three myomeres apart. An additional one to five pigment slashes extend onto the caudal finfold near the tail tip. Pigmentation changes little during larval de- velopment. Melanophores are added in the nape region and become embedded in the musculature over the notochord in larvae <5.5 mm. Mela- nophores are also added in the isthmus region and along the ventral midline of the gut. MorpHo.LoGy.—The smallest larval Artedius Type 3 is 2.9 mm NL and possesses remnants of the yolksac. Larvae undergo flexion of the no- tochord between 5.6 and 6.9 mm NL. The largest specimen examined is 7.6 mm long and has re- cently completed notochord flexion. Size at transformation is unknown. Thirteen larvae (2.9- 7.6 mm long) were examined for developmental morphology. Larvae of Artedius Type 3 are rather stubby with a moderately long gut; the posteriormost portion of this gut trails somewhat below the rest of the body. A prominent diverticulum extends dorsally from each side of the gut just posterior to the base of the pectoral fin. Diverticula are present in the smallest larva examined (2.9 mm Larvae of Artedius Type 3: A) 6.8 mm SL, B) 7.3 mm SL. NL) and remain pronounced in the largest spec- imen. Snout to anus length averages 45% SL in both preflexion and flexion larvae. Body depth at the pectoral fin base increases during development from 26% in preflexion larvae, to 28% in flexion larvae, and 32% SL in the single postflexion larva. Relative body depth at the anus also in- creases with development, from 23 to 30% SL. The distances from the snout to the origin of the pelvic fins and from the origin of the pelvic fins to the anus averages 26 and 22% SL, respectively, in late flexion and early postflexion larvae. Artedius Type 3 larvae have a rather large head with a blunt, rounded snout. With development relative head length increases from an average of 22% in preflexion larvae to 25% in larvae undergoing flexion of the notochord, and 29% SL in the postflexion larva. Jaw length averages about 43% HL throughout early larval devel- opment. In contrast, eye diameter decreases dur- ing development from an average of 47% in pre- flexion larvae to 37% HL in flexion and early postflexion larvae. Fin DEVELOPMENT.—A 4.9-mm NL larva ex- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 191 hibits a slight thickening of the hypural region of the forming caudal fin. By 5.6 mm, the no- tochord of larval A. Type 3 is strongly flexed and caudal rays are beginning to form. Notochord flexion is nearly complete by ~7 mm and the adult complement (6 + 6) of principal caudal rays is countable. The dorsal and anal fin bases are first visible in larvae between 6.8 and 6.9 mm long. In the largest specimen examined, 7.6 mm, the adult complement of dorsal spines (IX), dorsal rays (13-15), and anal fin rays (12) is complete. Pec- toral fin rays begin to form in a larva 6.8 mm NL, and 16 pectoral fin rays are countable in a larva 7.6 mm. Pelvic fin buds are first visible at ~6.8 mm NL; however, pelvic fin rays are not yet formed in the largest specimen. SPINATION. —Preopercular spines begin to form in small preflexion larvae of A. Type 3 at ~4.1 mm NL. A series of 15-17 tiny, equal-sized spines is visible along the posterior margin of the pre- opercle in preflexion larvae between 4.1 and 5 mm NL. During development, preopercular spines increase in number ranging from 21 to 24 in flexion and early postflexion larvae. In late flexion larvae (~6.8-6.9 mm NL) the middle 2 or 3 preopercular spines (the 8th—1 1th spine from the dorsal margin of the preopercle) begin to increase in size relative to other pre- opercular spines. In the 7.6-mm larva, the dor- salmost and ventralmost | or 2 spines are also larger than other preopercular spines. This forms the characteristic preopercular spine pattern found in Artedius larvae with multiple preoper- cular spines: the dorsalmost, middle, and ven- tralmost spines are markedly larger than the oth- er preopercular spines. No other spines develop on the head in larvae <7.6 mm. Head spination in larger larvae re- mains unknown. Oligocottus maculosus (Figures 16, 17; Table 5) LITERATURE. — Stein (1972, 1973) described O. maculosus larvae and illustrated specimens 4.6, 6.0, 6.6, and 9.2 mm TL. IDENTIFICATION.— Larvae in this series were reared from eggs spawned from known adults. Adults and juveniles were identified by the fol- lowing combination of characters: high vertebral (33-34) and dorsal fin ray (15-18) counts, small size at transformation (8-9 mm), absence of cirri me on the nasal spines and along the base of the dorsal fins, and pigmentation. The develop- mental series was linked together primarily on the basis of pigmentation, preopercular and pa- rietal spination, and body shape. Postflexion and transforming larvae were linked to juveniles by the serial method utilizing pigmentation, spi- nation, and size at transformation. DISTINGUISHING FEATURES.— Newly hatched larvae of O. maculosus reared in the laboratory are distinguished by the following pigmentation characters: intense melanistic nape pigment, dark dendritic melanophores that extend onto a prominent bubble of skin in the nape region just anterior to the origin of the dorsal finfold, | or 2 melanophores situated anteriorly on the vis- ceral mass beneath the pectoral fins, and a series of 18-36 ventral midline melanophores poste- rior to the anus. In addition to distinctive pig- mentation, larvae possess two rounded humps or protrusions that extend dorsally on either side of the gut just posterior to the pectoral fin bases. These protrusions are similarly positioned and reminiscent of the gut diverticula found in larvae of Artedius; however, they never develop into distinct diverticula. These protrusions disappear at the completion of yolk absorption about five to ten days after hatching. Oligocottus maculosus larvae also possess a distinctive bubble of skin in the nape region just anterior to the origin of the dorsal finfold. This bubble persists in larvae up to 7.5 mm SL. Flexion and postflexion larvae >6.5 mm pos- sess a relatively low number of preopercular spines (9-13). Postflexion larvae and juveniles may be dis- tinguished by meristics, especially the high ver- tebral and dorsal fin ray counts, and the small size at transformation (8—9 mm SL). In addition, juveniles possess a slender postorbital cirrus and two frontoparietal cirri. PIGMENTATION. — Newly hatched larvae of O. maculosus possess no melanistic head pigmen- tation. Fourteen to 16 intense, stellate melano- phores are concentrated in the nape region. One to 3 dendritic melanophores extend anteriorly from the nape pigment patch onto a prominent elevation or bubble of skin located just anterior to the origin of the dorsal finfold. In live larvae, xanthophores cover the bubble of skin and the nape. Three to 4 dendritic, embedded melano- phores are positioned in the otic capsule. The dorsal surface of the gut is darkly pig- 192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 16. Larvae of Oligocottus maculosus: A) 4.3 mm NL, B) 7.2 mm NL, C) 6.9 mm NL. mented with 100-150 dark melanophores. Two to 4 pale, dendritic melanophores are located along the anteroventral margin of the gut, just beneath the pectoral fins. These melanophores are frequently embedded in the gut musculature and are difficult to see. One to 5 small melano- phores are clustered around the anus. Posterior to the anus, larvae of O. maculosus possess a relatively high number of ventral mid- line melanophores. The actual number of me- lanophores appears to vary with the geographic location at which the larvae were collected. Stein (1973) recorded between 11 and 20 ventral mid- line melanophores in his reared larvae, while lar- vae reared in Oregon possessed between 16 and 20 melanophores along the ventral midline. J. B. Marliave (Vancouver Public Aquarium, Van- couver, B.C., Canada, pers. comm.) found be- tween 26 and 36 ventral midline melanophores in reared larvae from the Straits of Georgia in British Columbia. Regardless of the number of melanophores, this series begins under the third or fourth myomere posterior to the anus and extends toward the tail tip. The first four mela- nophores in the series are usually spaced one every two to three myomeres, while the remain- der of the melanophores are spaced one per myo- mere. Five or nine additional pigment slashes extend onto the ventral finfold near the tail tip. During larval development in larvae <6 mm, 15-20 melanophores form over the midbrain and interorbital region of the head. Two to 5 mela- nophores form on the snout and 1-3 melano- phores form on the cheek just anterior to the WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 193 NS | J / yy SIO Vg eR Nh ei OAR ERI oe ss 5 eee ( Ficure 17. Young of Oligocottus maculosus: A) 7.8 mm SL, B) 10.4 mm SL. dorsalmost preopercular spine in larvae >7 mm. At this size, melanophores are also added in the otic capsule; however, they become obscured by the developing musculature and are difficult to see. By ~6 mm, several melanophores are added just ventral to the nape pigment patch. Five to 7 of the centrally positioned melanophores be- come embedded while the other nape melano- phores form a prominent U-shape anterolater- ally around the central melanophores. During transformation, ~7-8 mm, melanistic pigmentation increases markedly over the dorsal surface of the head. Melanophores are added on the snout, on the cheek region anterior to the preopercle and on the dorsal portion of the oper- culum. Melanophores also form on the pectoral fin base and gradually extend ventrally onto the isthmus. Pigmentation over the dorsal surface of the head is intense in benthic juveniles. A band of melanophores forms on either side of the snout, extending from the upper lip to the ventral mar- gin of each eye. Each band continues posteriorly reaching from the eye to the dorsalmost pre- opercular spine. Melanophores are also added ventrally along the entire margin of the pre- operculum and on the anterior tip of the lower lip. In juveniles >8.5 mm, tiny melanophores cover the entire dorsolateral surface of the head; however, the bands of pigment extending through each eye remain prominent. An irregular band of tiny melanophores forms along the surface of the lateral midline in juveniles >9 mm. This band gradually extends posteriorly to the caudal fin base. Two additional bands of pigment form along the dorsum. A third band forms under the 8th—10th dorsal rays and a fourth band develops under the 14th—16th dorsal fin rays. These pig- ment bands eventually extend ventrally and fuse with the lateral midline pigmentation. Tiny me- lanophores are added over the dorsolateral body surface in juveniles ~13 mm; however, the in- tense pigment bands along the dorsum remain distinct. Melanophores extend out onto the dor- sal and caudal fin rays, forming three or five bands of pigment. MorrpHoLocy.—Newly hatched Oligocottus maculosus larvae range in length from 4.2 to 4.5 mm NL. Larvae undergo flexion of the noto- chord at 7.2-7.6 mm NL. Transformation occurs atarelatively small size, ~7.5-8 mm. The small- est benthic juvenile examined was 8 mm long. SS St ge 194 Taste 5. Bopy Proportions OF LARVAE AND JUVENILES OF OLIGOCOTTUS MACULOSUS AND O. SNYDERI. Values are percent PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 standard length (SL) or head length (HL) including mean, standard deviation, and range in parentheses. Item Head length/SL: Preflexion Flexion Postflexion Juvenile Snout length/HL: Preflexion Flexion Postflexion Juvenile Eye diameter/HL: Preflexion Flexion Postflexion Juvenile Snout to anus length/SL: Preflexion Flexion Postflexion Juvenile Snout to pelvic fin origin/SL: Preflexion Flexion Postflexion Juvenile Pelvic fin origin to anus/SL: Preflexion Flexion Postflexion Juvenile Body depth at pectoral fin base/SL: Preflexion Flexion Postflexion Juvenile Body depth at anus/SL: Preflexion Flexion Postflexion Juvenile Pectoral fin length/SL: Preflexion Flexion Postflexion Juvenile — = Not present at this stage. Oligocottus maculosus 17.3 + 0.61 (17.2-18.3) 19.7 + 1.84 (18.4-21.0) 25.3 + 1.89 (22.3-26.6) 30.1 + 3.06 (27.1-32.8) 2.08 (24.3-27.9) 1.08 (26.0-28.1) 2.83 (24.4-33.0) 3.22 (24.9-31.4) nN oO w Ge lhe theelbe 55.6 + 2.08 (53.9-57.2) 44.9 + 4.24 (42.3-48.2) 41.2 + 4.03 (33.1-45.0) + 2.01 (28.9-34.1) + 1.53 (37.2-40.4) 39.8 + 2.83 (38.1-42.3) 43.9 + 3.39 (40.8-48.1) 45.0 + 2.65 (42.7-48.5) 24.9 + 2.15 (23.1-26.9) + 3.61 (25.3-32.1) 18.0* 20.2 + 1.89 (18.1-22.8) 16.9 + 1.73 (16.1-19.4) 19.4 + 1.00 (17.8-20.2) 24.0 + 0.00 (24.0-24.0) 25.9 + 2.17 (23.7-29.1) 23.9 + 3.06 (20.6-27.2) 15.5 + 0.61 (14.9-16.2) 18.0 + 0.00 (18.0-18.0) 25.9 + 2.17 (23.7-29.1) 20.3 + 2.08 (18.1-22.3) 10.1 + 0.61 (9.8-11.2) 18.0 + 0.00 (18.0-18.0) 21.7 + 2.96 (17.3-26.3) 29.1 + 2.65 (26.8-31.9) * = Only one specimen available in this stage. Oligocottus snyderi 21.4 + 1.26 (20.1-23.4) 20.8 + 1.34 (18.9-23.1) 23.4 + 1.71 (21.4-28.2) 26.9 + 1.00 (26.5—28.0) 23.7 + 3.37 (21.2-25.9) 21.4 + 2.43 (16.9-23.3) 16.1 + 2.87 (13.9-20.4) 32.0 + 0.00 (32.0-32.0) 47.8 + 4.80 (40.1-52.3) 42.1 + 2.95 (37.6-46.1) 33.3 + 3.21 (31.2-37.9) 31.0 + 2.52 (29.0-34.3) 42.1 + 1.47 (40.1-44.4) 41.8 + 2.30 (39.2-44.9) 44.5 + 1.83 (42.1-46.4) 42.2 + 1.53 (40.3-43.1) 21.0 + 1.00 (20.0-22.0) 23.3 + 1.71 (21.0-25.5) 24.6 + 1.53 (22.9-26.1) 20.0 + 1.15 (19.1-21.1) 21.2 + 1.50 (20.1-23.3) + 3.21 (16.0-22.1) 23.2 + 1.17 (21.8-25.5) 23.4 + 2.00 (20.2-26.6) 20.6 + 1.50 (19.4-21.8) 21.2 + 2.00 (19.0-23.4) 21.0 + 2.37 (18.1-24.9) 24.3 + 2.91 (17.9-28.1) 26.2 + 1.50 (21.9-27.3) 22.1 + 2.31 (19.0-23.0) 8.2 + 2.53 (6.1-12.1) 10.1 + 1.77 (9.3-13.4) 14.4 + 1.41 (13.1-16.0) 24.3 + 1.53 (22.9-26.3) Eighteen specimens of O. maculosus (4.3-10.8 or protrusions appear on the dorsal surface of mm) were examined for developmental mor- the gut just posterior to the pectoral fin base. phology. These bulges are similar to the dorsal gut diver- In newly hatched larvae two prominent bumps __ ticula of Artedius larvae; however, they never WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 195 develop into distinct diverticula. The gut pro- trusions disappear approximately 5—10 days af- ter hatching. Oligocottus maculosus larvae also possess a distinctive bubble of skin in the nape region just anterior to the origin of the dorsal finfold. This bubble persists in larvae =<7.5 mm. Small O. maculosus are slender with a rela- tively short gut. Snout to anus length averages 39% in preflexion larvae and increases to 44% in postflexion larvae and 45% SL in juveniles. Body depth at both the pectoral fin base and the anus increases during larval development from 19% and 15%, respectively, to 25% SL. Fin DevELopMENT.—Larval Oligocottus mac- ulosus begin to undergo notochord flexion at ~6— 7 mm. The adult complement of 12 principal caudal rays is complete in larvae ~6.8-7 mm long at about the completion of notochord flex- ion. Dorsal and anal fin rays begin to form in larvae ~6.6 mm long; however, the full complement of fin rays (15-18, 12-14, respectively) is not com- plete until larvae are 8 mm long. Dorsal spines (VIII-IX) also form between 7 and 8 mm. AI- though pectoral fin rays are visible in larvae by 6.6 mm, the adult complement (12-15) is not fully formed until larvae reach about 7.6 mm. Pelvic buds are first visible in 7 mm larvae but the fin rays are not formed until ~8.5 mm. SPINATION. —Six to 7 tiny spines are first vis- ible on the posterior margin of the preopercle in larvae ~5.8 mm long. Spines increase in number to 9 or 10 in larvae undergoing notochord flex- ion. In postflexion larvae 6.9-7.8 mm long, pre- opercular spines number 10 or 11. Two or 3 of these spines appear as tiny accessory spines that form just anterior to the bases of the other spines. The dorsalmost spine becomes slightly larger than the lower spines. The 3rd, 4th, and Sth preoper- cular spines also increase slightly in size relative to the lower spines. In the largest planktonic lar- vae (~8 mm long) the preopercular spines begin to decrease in size and number and are covered with skin. In newly settled benthic juveniles ~8- 10 mm long, the dorsalmost spine is quite large and stout with a strong upward curvature. The lower spines persist only as three blunt, bony protrusions on the preopercular margin. Three tiny spines also form in the parietal re- gion in larvae ~6-7 mm long. Two spines de- velop anteriorly with a third nuchal spine form- ing just posterior to them. These parietal spines persist through the larval period, but they regress in benthic juveniles. During regression, the an- terior spines decrease in size and their tips bend posteriorly and fuse with the nuchal spine, form- ing an arch. This canal and arch become incor- porated into the cephalic lateral line system. Two spines also form on the posttemporal in larvae ~6-7 mm long. By ~7-8 mm, a third spine forms on the posttemporal and a fourth spine forms on the supracleithrum. These su- pracleithral-posttemporal spines persist through larval development and eventually form the junction of the cephalic and lateral line systems. Oligocottus snyderi (Figures 18, 19; Table 5) LITERATURE. — Stein (1973) described and il- lustrated 4.5- and 5.5-mm TL larvae of O. sny- deri. Richardson and Washington (1980) called these larvae Cottidae Type | and illustrated spec- imens 4.2, 6.7, and 9 mm long. IDENTIFICATION.—Small larvae in this series were reared from eggs spawned from known adults. Adults and juveniles were identified by the following combination of characters: high vertebral (34-37) and dorsal fin ray (17-20) counts, light pigmentation, the presence of cirri on the nasal spines and along the bases of the dorsal fins, and the absence of scales (prickles). The developmental series was linked together primarily on the basis of pigmentation, body shape, and preopercular and parietal spination. Postflexion and transforming larvae were linked to juveniles by pigmentation, meristics, and pre- opercular and parietal spination. DISTINGUISHING FEATURES.— Distinguishing pigmentation of preflexion larval O. snyderi are melanistic nape pigmentation, relatively light pigmentation over the dorsolateral surfaces of the gut, and a low number of ventral midline melanophores (5-9) situated posterior to the anus. This series of ventral midline melanophores originates beneath the fifth to seventh postanal myomeres and extends posteriorly toward the tail tip. One melanophore is spaced approxi- mately every four or five myomeres. This char- acteristic pigmentation changes little during lar- val development. In newly hatched larvae, a hump or bubble of skin is present just anterior to the origin of the dorsal finfold. Although diffuse xanthophores are present over this bump in laboratory-reared lar- vae, no melanophores extend onto this bubble of skin. In contrast, O. maculosus larvae, which 196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 18. 1980). also possess this distinctive bubble of skin at the nape, have one to three large dendritic mela- nophores that extend up onto the bubble of skin from the nape pigment patch. Larvae of O. snyderi >4.2 mm NL may be further distinguished from all other known cottid larvae by the presence of a cluster of 10-20 mi- nute prickles situated in the parietal region of the head. Larvae undergoing notochord flexion, >6 mm long, possess a distinctive pattern of multiple preopercular spination, in which approximately 15 equal-sized spines are positioned along the Larvae of Oligocottus snyderi: A) 4.7 mm NL, B) 5.1 mm NL, C) 6.7 mm NL (C from Richardson and Washington posterior margin of the preopercle. Ten to 11 small, accessory spines are situated at the ante- rior bases of the other spines and point antero- laterally. Postflexion larvae and juveniles may be dis- tinguished by their relatively light pigmentation, the prominent bands of pigment through the eye, and the low number of widely spaced ventral midline melanophores. In addition to pigmen- tation, juvenile O. snyderi are characterized by high vertebral and dorsal fin ray counts (34-37 and 17-20, respectively), and by the presence of very long, slender nasal, postorbital, and fron- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 197 Ficure 19. Young of Oligocottus snyderi: A) 8.2 mm SL, B) 10.2 mm SL, C) 14.4 mm SL. toparietal cirri. In juveniles > 15 mm, a row of distinctive cirri develop along either side of the bases of the dorsal fins. PIGMENTATION. — Newly hatched, reared lar- val O. snyderi are lightly pigmented with no mel- anistic head pigmentation. Several xanthophores are situated over the midbrain in live larvae. Two to 8 external melanophores are clustered over the notochord in the nape region. In live specimens several diffuse xanthophores extend dorsally from the nape onto a distinctive bump or bubble of skin just anterior to the origin of the dorsal finfold. In contrast to larvae of O. maculosus, however, melanophores never ex- tend onto this bubble of skin in larvae of O. snyderi. The dorsolateral surface of the gut is lightly pigmented with 50-60 small melano- phores forming an elliptical patch over the body cavity. Intense xanthophores also cover the dor- solateral surfaces of the gut. The only pigmen- tation posterior to the anus consists of a series of 5-9 melanophores that originates under the fifth to seventh postanal myomere and extends posteriorly. Each melanophore is positioned four to six myomeres apart. Occasionally, one or two melanistic pigment slashes extend onto the cau- dal finfold just beneath the notochord tip. During larval development, several melano- phores form on the dorsal surface of the head. Size of larvae at formation of this melanistic pig- 198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 mentation appears somewhat variable. One or two melanophores are present over the midbrain in 21-day-old, laboratory-reared larvae (~6.5 mm long). Morris (n.d.) reported that four or five melanophores develop in the midbrain region of larvae at four weeks of age (~8.5 mm). Mela- nistic pigmentation does not appear over the brain in field-caught larvae until about 10 mm. During transformation, head pigmentation in- creases markedly in late postflexion larvae 10— 14 mm long. Thirteen to 18 large, stellate me- lanophores form over the midbrain and inter- orbital regions of the head. Concurrently, several small melanophores form anterior to the orbit and extend anteriorly across the snout onto the upper lip forming a distinct band. Several intense melanophores develop posterior to the orbit forming a dark band extending from the orbit to the dorsalmost preopercular spine. Melano- phores are also added at the posterior margin of the lower jaw just ventral to the preopercle, along the dorsal margin of the operculum, and along the pectoral fin rays. A row of intense, embedded melanophores forms just above the spinal cord and extends posteriorly from the nape region two- thirds of the way to the caudal fin. The ventral midline melanophores remain unchanged. In juveniles between 13 and 15 mm long, nu- merous tiny melanophores form over the dor- solateral surfaces of the head and extend ante- riorly onto the snout between the eyes, ventrally along the preopercle, and along the opercular margin. These diffuse melanophores extend pos- teriorly to the seventh dorsal spine. Numerous small melanophores also form posterior to the pectoral fin base in an irregular band of pigment along the lateral midline. With development, melanophores are added along the dorsum in an anterior to posterior sequence. Concurrently, melanophores extend dorsally onto the dorsal fins forming four to five distinct bands of pig- ment. Gradually, the melanophores along the dorsal midline extend ventrally and posteriorly and join the dorsal and lateral areas of pigmen- tation. This lateral pigmentation extends poste- riorly and forms a dark band at the base of the caudal fin. Melanophores extend onto the caudal fin rays forming four to five bands of pigment. Juvenile O. snyderi are characterized by uniform diffuse pigmentation over the head and dorso- lateral surfaces of the body with a distinct, dark band of pigment extending from the snout, through the orbit, to the dorsalmost preopercular spine. The characteristic low number of widely spaced ventral midline melanophores remains visible in juveniles <18 mm. MorpHoLocy.—Newly hatched O. snyderi larvae range in size from 4 to 4.5 mm NL. No- tochord flexion occurs between 6.2 and 8.4 mm NL. The largest planktonic specimen taken in the field is 10.2 mm and has not yet begun to undergo transformation. The smallest benthic juvenile examined is 12.4 mm. Twenty-four specimens, ranging in length from 4 to 15.1 mm, were examined for development morphology. Newly hatched O. snyderi larvae are rather slender with a relatively short gut, the poste- riormost portion of which trails well below the body. Snout to anus length averages 42% in pre- flexion and flexion larvae, then increases slightly to 44% SL in postflexion larvae. Relative body depth at the pectoral fin base increases from 23% to 25% SL during larval development. A small, rounded protrusion extends dorsally from the dorsal surface of the gut just posterior to the pectoral fin base in newly hatched larvae. This protrusion is reminiscent of the gut diverticula found in larvae of several species of Artedius but is much less pronounced and never develops into distinct diverticula. This protrusion decreases in size shortly after hatching and is no longer visible by yolk absorption five days after hatching. In addition, O. snyderi larvae possess a prominent bump or bubble of skin that protrudes dorsally in the nape region just anterior to the origin of the dorsal finfold. This bubble persists in larvae up to ~6.5-7 mm. Fin DEvVELOPMENT.—Larvae of O. snyderi undergo notochord flexion between 6.2 and 8.4 mm. Caudal fin rays first appear at 7.8 mm; how- ever, the full adult complement of 6 + 6 prin- cipal caudal rays is not complete until larvae reach ~9-10 mm. Rays begin to form in the dorsal and anal fins of larvae between 7.5 and 8 mm long; however, these rays are not fully formed in larvae <9 mm. Adult complements are 17— 20 and 12-15, respectively. Dorsal fin spines be- gin forming in larvae 9-10 mm long, and the adult complement of spines (VIII-IX) is count- able in a 10.2 mm specimen. Pectoral fin rays (12-15) form at 9 mm and are complete by 10 mm. Pelvic buds are first visible in larvae be- tween 8.2 and 9 mm, but the fin rays are not fully formed until larvae reach 10-12 mm. SPINATION.—Five to nine tiny bumps form along the posterior margin of the preopercle WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 199 in larvae >4.2-5 mm. By ~5.1 mm, 10-15 tiny equal-sized spines are visible. During notochord flexion the preopercular spines increase in size and number, ranging between 17 and 22. The preopercular spines of O. snyderi larvae are unique in that 10-12 spines form along the pos- terior margin of the preopercle as in other cottids with preopercular spines, yet by ~7 mm NL, between 8 and 10 small accessory spines form anteriorly at the bases of the original spines. In larvae >9 mm, the dorsalmost preopercular spine becomes stouter and longer than the other spines and is separated from the lower spines by a short gap on the preopercular margin. The 5-8 spines just ventral to the dorsalmost spine also become slightly larger relative to the lower preopercular spines. Between 12 and 14 preopercular spines are visible in newly settled benthic juveniles. The dorsalmost spine becomes much larger relative to the other spines. The smaller, accessory spines begin to atrophy and are represented only by small bumps or irregularities on the preopercle. By ~14 mm, only the dorsalmost spine persists. Distinctive spines also form in the parietal re- gion of the head of young O. snyderi. Larvae as small as 4.2 mm have 7-10 small bumps or prickles visible over the parietals. These prickles increase in number during development; 10-20 prickles are present in the parietal region in lar- vae =>6.2 mm. Eight to 12 tiny prickles remain visible along the posterolateral margin of the pa- rietal bones in 12—13-mm cleared and stained benthic juveniles. A cluster of spines also develops in the supra- cleithral-posttemporal region in larvae >8 mm. One spine forms on the supracleithrum and five spines, situated in two rows, form on the dorsal portion of the posttemporal. These persist throughout larval development but atrophy dur- ing transformation until only three bony projec- tions are present in benthic juveniles. These bony projections represent the rudiments of the incip- ient lateral line system. Clinocottus acuticeps (Figures 20-22; Table 6) LiTeRATURE.— Blackburn (1973) illustrated and described an 8.6-mm specimen, which he called Cottid 1 “Biramous anus.” Richardson (1977) and Richardson and Pearcy (1977) listed larvae of C. acuticeps as Cottidae sp. 12. Larvae of this species were described by Richardson and Wash- NN Sse ——a ington (1980). They illustrated specimens 3.7, 3.9, 6.9, 7.6, 10.4, 13.8, and 16.5 mm long. IDENTIFICATION.—Small larval C. acuticeps were reared from eggs spawned from known adults. Adults and juveniles were identified by low dorsal fin ray (13-17) and anal fin ray (9- 13) counts, the presence of nasal cirri, and a membrane connecting the innermost pelvic fin ray with the abdomen. The developmental series was linked together primarily by pigmentation, body shape, and hindgut diverticula. Postflexion and transforming larvae were linked with juve- niles by pigmentation, meristics, and the mem- brane attaching the pelvic fin rays to the abdo- men. DISTINGUISHING FEATURES. — Clinocottus acu- ticeps larvae are distinguished from all other known cottid larvae by long protrusions (diver- ticula) that extend posteriorly from the gut on either side of the anus. These diverticula are pres- ent in yolk-sac larvae and persist in the largest pelagic specimens (14.5 mm). The gut itself is distinctively long and the posterior portion trails well below the body. Snout to anus length, av- eraging 62.5% SL, is greater than in other known larvae of Artedius, Clinocottus, or Oligocottus. In addition, these larvae have a flabby appear- ance with an outer bubble of skin, which is es- pecially pronounced in the head region. Other characters useful in distinguishing C. acuticeps larvae are melanistic pigmentation on the snout and head, and relatively few ventral midline melanophores (4-10). Transforming and juvenile C. acuticeps are distinguishable from all other known cottids by the presence of a membrane attaching the inner pelvic fin ray to the belly. Other characters useful in separating juveniles are the relatively light, uniform pigmentation over the body; a band of pigment extending from the snout posteriorly through the orbit toward the preopercle; a dark blotch of pigment at the anterior end of the spinous dorsal; and a low number of ventral mid- line melanophores. PIGMENTATION. — Newly hatched larvae reared in the laboratory exhibit 4 or 5 dendritic mela- nophores on the snout and 2 faint melanophores in each otic capsule. In field-collected larvae <3.7 mm NL, the presence of snout pigment varies; however, all larvae > 3.7 mm NL possess at least 2 melanophores on the snout. Eight to 15 me- lanophores are clustered in the nape region of even the smallest larvae. Numerous melano- 200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 20. Larvae of Clinocottus acuticeps: A) 3.7 mm NL, B) 3.7 mm NL, C) 3.9 mm NL, D) 6.9 mm NL (from Richardson and Washington 1980). WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 201 Ficure 21. 1980). phores are scattered over the dorsolateral surface of the gut extending posterolaterally over the sur- face of the gut diverticula. These melanophores are much fainter and more irregular in shape than in larvae of other species of Clinocottus. A series of 4-10 inconspicuous ventral midline melanophores originates beneath the 7th-10th postanal myomeres and extends posteriorly to- ward the tail tip. Several additional melano- phores appear as streaks of pigment on the ven- tral finfold near the tail tip. Larvae of Clinocottus acuticeps: A) 7.6 mm SL, B) 10.4 mm SL, C) 13.8 mm SL (from Richardson and Washington Pigmentation increases on the head during lar- val development. Melanophores form first on the head over the midbrain in larvae 5.5 mm NL. Concurrently, several embedded melanophores appear on the nape and extend anteriorly onto the head. Four to five internal melanophores oc- cur in or near the otic capsule. In larvae >6.5 mm, scattered melanophores extend continu- ously from the snout to the nape region. Ventral midline melanophores persist in flexion and postflexion larvae, and the posteriormost mela- 202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 22. Young of Clinocottus acuticeps: A) 13.8 mm SL, B) 16.5 mm SL (B from Richardson and Washington 1980). nophore, which is located near the notochord tip in small larvae, occurs near the middle of the caudal fin base between the forming hypural plates. The posteriormost melanophores which extended onto the ventral finfold now occur on the caudal fin. In the largest planktonic larvae, which are beginning to undergo transformation, melanophores are added in a patch just posterior to the orbit. Melanophores also are added along the bases of the pectoral fins and extend onto the pectoral fin rays. In newly settled benthic juveniles (13-14 mm) head pigmentation increases markedly. Mela- nophores extend anteriorly across the dorsal sur- face of the snout and onto the upper lip. The melanophores at the ventral margin of the snout are especially intense and closely spaced, forming a prominent band that extends from the upper lip to the ventral margin of the orbit. This band continues from the posterior margin of the orbit to the dorsal margin of the preopercle. Addi- tional melanophores are added along the ventral edge of the lower lip, at the base of the preopercle, and on the dorsal portion of the operculum. As development proceeds, a second band of pig- ment forms between the ventral margin of the orbit and the melanophores at the base of the preopercle. Simultaneously, pigmentation in- creases on the pectoral fin bases while two to three bands of pigment form across each fin. Between 14 and 15 mm, a dense patch of me- lanophores forms at the anterior end of the first dorsal fin between the first and third spines. As juvenile pigmentation progresses this patch ex- pands posteriorly to include the fourth dorsal spine, and a second patch of melanophores forms between the seventh and eighth dorsal spines. Melanophores extend ventrally from these two pigment patches forming two distinct bands across the dorsum. Pigmentation proceeds pos- teriorly along the dorsum. In juveniles between 15 and 16 mm long, a third band (or saddle) of pigment forms under the second to sixth dorsal fin rays; a fourth band forms under the 8th to 11th dorsal fin rays; and a fifth band forms under the last two dorsal fin rays. As these bands of pigment develop along the dorsum, they extend ventrally and eventually unite into a uniform band of pigment above the lateral midline. Con- currently, another band of pigment extends pos- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 203 TasLe 6. Bopy PRoporTIONS OF LARVAE AND JUVENILES OF CLINOCOTTUS ACUTICEPS, C. EMBRYUM, C. GLOBICEPS, AND C. ANaLis. Walues given are percent standard length (SL) or head length (HL) including mean, standard deviation, and range in parentheses. Item Head length/SL: Clinocottus acuticeps Clinocottus embryum Clinocottus globiceps Clinocottus analis Preflexion 27.1 + 3.44 (22.3-30.1) 26.1 + 2.65 (26.0-30.1) 20.6 + 2.21 (17.0-25.0) - Flexion 29.2 + 1.92 (26.8-32.4) 23.9 + 1.15 (21.8-25.3) 23.0 + 1.79 (21.0-27.2) - Postflexion 27.9 + 2.71 (22.9-32.6) 24.0 + 0.00 (24.0-24.0) 27.3 + 4.09 (22.4-33.3) 30.1 + 1.25 (27.8-32.1) Juvenile 32.2 + 0.65 (31.5-32.7) 31.5 + 1.13 (30.2-32.2) 31.6 + 0.46 (31.2-32.1) - Snout length/HL: Preflexion 24.6 + 4.80 (21.3-25.9) 21.3 + 1.53 (19.9-23.7) 21.3 + 4.40 (14.5-26.7) - Flexion 26.4 + 2.97 (22.9-30.6) 24.1 + 3.83 (18.4-30.8) 25.7 + 3.39 (20.0-30.5) - Postflexion 23.6 + 4.38 (16.3-30.9) 22.2 + 0.58 (21.8-23.6) 25.3 + 4.84 (20.0-33.0) 28.3 + 2.70 (22.3-32.2) Juvenile 27.5 + 4.36 (26.8-29.4) 27.5 + 1.47(22.7-31.2) 27.1 + 0.92 (26.1-27.9) - Eye diameter/HL: Preflexion 39.7 + 6.63 (31.5-54.2) 39.3 + 3.06 (35.8-42.1) 50.4 + 9.16 (46.2-63.1) - Flexion 34.6 + 2.41 (31.9-36.4) 35.7 + 3.79 (34.8-39.5) 43.7 + 7.39 (37.9-51.4) - Postflexion 32.7 + 3.30 (27.1-38.2) 32.1 + 3.06 (29.0-35.3) 38.2 + 8.09 (24.4-46.0) 31.3 + 1.41 (27.8-33.0) Juvenile 27.9 + 2.06 (25.5-29.3) 29.8 + 0.74 (29.2-30.6) 27.1 + 0.91 (26.1-27.9) - Snout to anus length/SL: Preflexion 60.7 + 4.64 (54.4-67.2) 51.6 + 3.51 (48.2-55.1) 44.0 + 3.59 (39.6-52.9) - Flexion 62.8 + 2.74 (60.3-67.1) 49.9 + 3.87 (43.9-54.4) 48.2 + 3.27 (44.4-56.0) - Postflexion 62.5 + 4.61 (57.5-70.3) 50.0 + 3.06 (47.3-53.8) 50.0 + 3.48 (44.7-56.8) 48.9 + 2.40 (46.4-54.3) Juvenile 50.2 + 1.64 (48.4-51.6) 49.0 + 3.10 (47.0-52.6) - - Snout to pelvic fin origin/SL: Preflexion - - - - Flexion - 28.0* - 24.6 + 2.94 (21.4-28.0) - Postflexion 33.4 + 2.63 (29.4-39.5) 32.1 + 3.54 (29.0-34.2) 26.7 + 2.33 (23.4-30.9) 29.3 + 1.34 (27.2-31.4) Juvenile 33.3 + 1.16 (32.5-34.6) 31.1 + 1.31 (29.6-32.1) 30.7 + 1.04 (29.5-31.4) - Pelvic fin origin to anus/SL: Preflexion - - - - Flexion - 26r1= 21.5 + 1.77 (18.7-23.4) - Postflexion 29.4 + 4.56 (23.1-37.3) 18.2 + 0.71 (17.1-18.4) 22.9 + 1.42 (21.4-25.2) 19.9 + 2.26 (17.3-22.3) Juvenile 16.9 + 1.70 (15.9-18.9) 17.9 + 2.50 (15.4-20.4) 20.7 + 0.64 (20.0-21.2) - Body depth at pectoral fin base/SL: Preflexion 24.3 + 3.25 (17.8-29.1) 25.8 + 4.36 (21.2-29.4) 20.8 + 2.87 (15.8-25.5) - Flexion 27.6 + 2.30 (24.3-30.2) 26.3 + 2.29 (23.7-30.2) 23.9 + 2.02 (22.2-29.3) - Postflexion 31.3 + 2.29 (28.4-35.0) 26.1 + 1.53 (25.0-28.2) 26.8 + 2.78 (22.1-30.9) 28.1 + 1.36 (25.4-29.1) Juvenile 26.1 + 1.99 (23.9-27.8) 21.6 + 1.01 (23.5-25.5) 27.3 + 0.53 (26.7-27.7) - Body depth at anus/SL: Preflexion 21.4 + 2.94 (18.0-24.5) 22.8 + 4.36 (18.2-26.5) 17.5 + 2.46 (13.6-21.6) - Flexion 25.4 + 2.07 (23.3-28.1) 25.1 + 1.51 (22.9-27.3) 21.2 + 2.07 (17.7-24.0) - Postflexion 28.4 + 2.43 (26.4-35.2) 27.0 + 0.00 (27.0-27.0) 25.7 + 3.04 (21.1-30.4) 25.7 + 1.50 (25.0-29.1) Juvenile 24.0 + 2.28 (21.4-25.4) 23.3 + 1.93 (22.1-25.5) 25.7 + 2.08 (23.3-27.0) - Pectoral fin length/SL: Preflexion 11.4 + 1.27 (9.6-13.3) 9.9 + 3.46 (6.3-12.1) 12.1 + 1.19 (9.6-13.8) - Flexion 11.0 + 1.87 (9.9-14.5) 11.2 + 2.75 (7.1-17.5) 12.3 + 4.40 (7.5-22.0) - Postflexion 26.4 + 5.95 (18.2-35.0) 32.0 + 4.04 (30.1-37.0) 22.7 + 6.00 (11.1-30.4) 29.3 + 2.15 (24.8-32.3) Juvenile 32.1 + 1.74 (30.8-34.1) 33.4 + 1.16 (32.1-34.2) 29.0 + 0.82 (28.1-29.7) - — = Not present at this stage. * = Only one specimen available in this stage. 204 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 teriorly along the lateral midline to the caudal fin where melanophores form a dark band at the base of the caudal fin. Melanophores extend onto the caudal fin rays where they form four or five bands across the fin. Clusters of melanophores extend first ventrally and then laterally from the lateral midline pigment band and gradually unite enclosing three to five small unpigmented circles below the lateral line. Three to six tiny mela- nophores remain visible along the ventral mid- line posterior to the anus in juveniles <17 mm long. MorpuHo.oGy.—Clinocottus acuticeps larvae hatch at the smallest length (3.1-3.3 mm NL) of any member of the genus. Notochord flexion oc- curs between 5.6 and 7.3 mm NL). The largest planktonic specimen collected is 14.5 mm and is beginning to undergo transformation. The smallest benthic juvenile examined is 12.6 mm. Thirty-seven selected specimens of C. acuticeps, ranging in length from 3.1-16.2 mm, were ex- amined for developmental morphology. Larvae of C. acuticeps have a distinctive, flab- by appearance as if a loose bubble of outer skin surrounds the anterior part of the body. Larvae are deep-bodied with a long, distinc- tive gut, the posterior portion of which trails well below the body. Snout to anus length remains relatively constant during larval development, averaging 63% SL. Prominent diverticula extend posteroventrally from the hindgut on either side of the anus. These diverticula are well developed throughout larval development but are not vis- ible in benthic juveniles. Fin DEvVELOPMENT.—A 5.6 mm NL larva is just beginning notochord flexion and a concur- rent thickening of the hypural region of the form- ing caudal fin. The adult complement of 6 + 6 principal caudal rays is present in a 6.8 mm spec- imen prior to completion of notochord flexion at ~7.5 mm. Bases of the dorsal and anal fin rays begin forming on a 6.9 mm larva. The full adult complement of dorsal (13-17) and anal (9-13) fin rays is present by ~8 mm. The adult com- plement of dorsal fin spines (VII-IX), however, is not present until ~8.7 mm. Although pectoral fin rays are visible on a 6.9 mm larva, the full adult complement (13-15) is not complete until >7.6 mm. Pelvic fin buds appear just after completion of notochord flexion in a 7.6 mm larva; however, the fin rays are not fully formed until ~10 mm. SPINATION.—Preopercular spines first appear as small bumps at 5.2 mm NL. Nine to 11 small spines are present by the onset of notochord flex- ion at ~6 mm. During flexion, spines remain small and evenly spaced with the 2nd and 3rd spines becoming slightly longer than the others. By completion of flexion, at 7.6 mm, | 1-12 spines are present along the margin of the preopercle. The dorsalmost 3 spines are beginning to elon- gate and point dorsally. In a 10-mm cleared and stained specimen, the dorsalmost 3 spines are nearly four times as long as the ventral spines. In the largest planktonic larvae (13-14 mm long) the ventralmost spines are beginning to atrophy. The 3 dorsalmost spines are still prominent in a 15.2 mm juvenile, but the 8 ventral spines are minuscule, with their tips twisted and bent an- teriorly. By ~ 19 mm, the lower spines have atro- phied completely, and only the single large dor- salmost spine persists. No spines develop in the parietal or supra- cleithral-posttemporal regions of the head in lar- vae or juveniles of this species. Clinocottus embryum (Figures 23-25; Table 6) LITERATURE. —Richardson (1977) and Rich- ardson and Pearcy (1977) listed larvae of this species as Cottidae sp. 20. Richardson and Washington (1980) described these larvae as Cottidae Type 2 and illustrated specimens 4.0, 6.4, and 7.4 mm long. IDENTIFICATION.—Juveniles and adults were identified using a combination of the following characters: an advanced anus, light pigmenta- tion, presence of a nasal cirrus, low anal fin ray counts (9-12), and absence of a membrane at- taching the pelvic fin rays to the abdomen. The developmental series was linked together pri- marily on the basis of pigmentation, body shape, and preopercular spination. Postflexion and transforming larvae were linked to juveniles by pigmentation, cirri patterns, meristics, and pre- opercular spination. DISTINGUISHING FEATURES.—Characters use- ful in distinguishing preflexion larvae of C. em- bryum are lack of head pigment, relatively light gut pigmentation, large number of ventral mid- line melanophores (15-21), and relatively long, trailing gut. Head and/or snout pigment is pres- ent in larvae of all other species of Clinocottus. C. embryum larvae are further distinguished from WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 205 yolk-sac C. acuticeps larvae (in which snout pig- ment is sometimes absent) by the absence of dis- tinct hindgut diverticula. In addition to the pigmentation characters mentioned above, flexion and postflexion larvae of C. embryum are distinguished by their head spination. Larvae 7.3 mm have 11-14 preoper- cular spines; the dorsalmost spine is the largest. In benthic juveniles of C. embryum, the anus is advanced midway between the origin of the pelvic fins and the anal fin, as in other members of the genus. Juvenile C. embryum are distin- guished from C. globiceps, C. analis, and C. re- calvus by relatively light, mottled body pigmen- tation and long and slender nasal, postorbital, and frontoparietal cirri. Clinocottus embryum juveniles are distinguished from C. acuticeps by the presence ofa large number of ventral midline melanophores (15-21) and the absence of a membrane connecting the inner pelvic fin ray to the abdomen. PIGMENTATION. —Melanistic pigmentation is absent on the head in preflexion C. embryum larvae. One to 5 melanophores are scattered over the nape region. The dorsolateral surface of the gut is relatively lightly pigmented and occasion- ally several faint melanophores are present on the anterolateral surface of the gut below the pec- toral fins. Posterior to the anus, a series of 15- 19 melanophores extends along the ventral mid- line. This series begins on the fourth or fifth myo- mere posterior to a vertical line through the anus; the melanophores are spaced approximately 1 per myomere. Several specimens have | or 2 melanophores on the ventral finfold near the no- tochord tip. During larval development, the formation of head pigmentation varies in larvae between 6 and 9 mm long. Three out of eight larvae ob- served possess one to five tiny melanophores over the brain. One or two melanophores are consis- tently present beneath the pectoral fin on the anterolateral surface of the gut in larvae >6.5 mm. Otherwise, pigmentation remains un- changed. In transforming larvae >9.6 mm long, nu- merous melanophores appear over the brain. Several melanophores appear on the cheek re- gion between the orbit and the preopercle. Me- lanophores are also added on the pectoral fin base. Melanistic pigmentation increases over the head in newly settled juveniles. Large melano- phores cover the surfaces of the head over the midbrain and interorbital regions. Several large, intense melanophores are embedded at the pos- terior margin of the parietal region. A distinct, dense band of melanophores extends from the orbit anteriorly onto the upper lip and posteriorly from the orbit to the dorsal tip of the preopercle. Several melanophores form a dark patch on the cheek beneath the orbit. Melanophores are also added to the dorsal surface of the operculum and to the pectoral fin base with several melano- phores extending onto the pectoral fin rays. As development proceeds, pigmentation in- creases markedly over the head. In a 16-mm juvenile, the bands of pigment extending through the eye are prominent, but numerous small me- lanophores cover the entire dorsal surface of the head above these bands of pigment. Pigmenta- tion increases on the operculum and pectoral fin base. Three to four bands of melanophores are added across the pectoral fin rays. Five bands of pigment develop on the body along the dorsal midline in an anterior to pos- terior sequence. The first band of pigment forms under the third to fifth dorsal fin spines, and a second smaller band begins to form under the seventh to ninth dorsal spines in juveniles be- tween 13 and 14 mm long. By ~15 mn, three additional bands of pigment are present on the dorsum beneath the second dorsal fin. The third band forms under the 2nd—4th dorsal fin rays, the fourth band forms under the 7th—9th fin rays, and the fifth band forms under the 12th—15th rays. At the same time, a series of embedded melanophores develops in a row just above the notochord, extending from the nape region to- ward the caudal fin. A few diffuse patches of external melanophores also form along the lat- eral midline posterior to the gut. As juvenile pigmentation develops, the dorsal bands of pigment extend ventrally until they unite above the lateral line, forming four unpigmented saddles between the bands. The melanophores lying along the lateral midline increase in number and extend posteriorly and ventrally toward the caudal fin. As this lateral pigmentation extends posteriorly, it fuses dorsally with the pigment bands. As pigmentation expands and unites over the lateral surface of the body, numerous, irreg- ular, unpigmented circles remain above and be- low the lateral line, giving juvenile C. embryum 206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 23. Larvae of Clinocottus embryum: A) 4.0 mm NL, B) 5.4 mm NL, C) 6.4 mm NL (A and B from Richardson and Washington 1980). a distinctively mottled appearance. Eighteen to 21 small melanophores remain visible along the ventral midline in juveniles up to ~ 19 mm long. MorpHoLtocy.—The smallest C. embryum examined is 4.0 mm NL and is recently hatched. Larvae undergo flexion of the notochord between 6.4 and 9.6 mm NL. The largest specimen taken in the plankton is 14.0 mm and is beginning to undergo transformation. The smallest benthic juvenile is 13.7 mm. Eighteen C. embryum, ranging from 4 to 14 mm long, were examined for developmental morphology. Larval C. embryum have a distinctively shaped gut with the posterior portion trailing well below WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 207 Ficure 24. Larvae of Clinocottus embryum: A) 7.4 mm SL, B) 9.6 mm SL, C) 13.9 mm SL (A from Richardson and Washington 1980). the body. The walls of the hindgut protrude on either side of the anus, reminiscent of the hindgut diverticula of C. acuticeps; however, these bulges never develop into pronounced diverticula. Snout to anus length is relatively long throughout larval development, averaging 50% SL. Fin DEvVELOPMENT.—The onset of notochord flexion is first apparent in a 6.4 mm larva. Caudal rays are present by ~7.4 mm but the adult com- plement of 6 + 6 principal caudal rays is not complete until about 8.4 mm. Bases of the form- ing dorsal and anal fin rays are first visible at ~7.4 mm; however, the adult complement of dorsal (14-17) and anal (9-12) fin rays is not present until ~8.3 mm. Dorsal spines (VIII-X) SS eer are beginning to form at ~8.3 mm but are not fully formed until 9.6 mm. Pectoral fin rays begin to form at ~8 mm, and the adult complement of fin rays (12-15) is present by 9.6 mm. Pelvic fin buds are first apparent at ~9.6 mm, and the adult pelvic fin complement (1,3) is present in postflexion larvae > 12.4 mm long. SPINATION.—Eight to 10 tiny, evenly spaced spines increases, ranging in number from 11 to 14. opercle at ~5.2 mm NL. In larvae undergoing notochord flexion, the number of preopercular spines increases, ranging in number from 11 to 14. During the flexion stage, the dorsalmost pre- opercular spine increases in size relative to the rest of the preopercular spines so that by the end 208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 25. Juveniles of Clinocottus embryum: A) 13.7 mm SL, B) 16.2 mm SL. of flexion, the dorsalmost spine is much longer and stouter than the other spines. In the largest planktonic larvae, 13-14 mm long, the upper spine is =2.5 times longer than the other spines and is separated from them by a slight gap. In newly settled benthic juveniles, the number and size of the lower preopercular spines are reduced. By >15 mm, only a second, tiny spine persists just ventral to the large dorsal spine. The other spines appear as five to seven small bumps or irregularities along the preopercular margin. In completely transformed juveniles = 16 mm long, only the uppermost spine is visible. Two spines develop in the parietal region of C. embryum larvae. A single small spine is first present at the posterior margin of each parietal at ~6.7 mm. By 9.6 mm, this parietal spine has increased in size and a second smaller parietal spine is present just behind it. As larvae undergo transformation, between 12 and 14 mm, these spines undergo a reduction in size, and the first parietal spine eventually fuses with the second parietal spine, forming a hollow central canal between the spines. This canal is part of the in- cipient cephalic lateral line system. In newly set- tled juveniles, only a skin-covered bony protu- berance is visible in the parietal region. Three spines also form in the supracleithral- posttemporal region of the head at ~9.6 mm. These spines persist through transformation and eventually become associated with the lateral line system in juveniles >15 mm long. Clinocottus globiceps (Figures 26-28; Table 6) LITERATURE. — Larvae of this species were list- ed by Richardson (1977) and Richardson and Pearcy (1977) as Oligocottus sp. 1. Richardson and Washington (1980) described larvae of this species as Cottidae Type 3. They illustrated spec- imens 6.3, 7.5, and 12.5 mm long. IDENTIFICATION.— Larvae were reared from eggs spawned from known adults. Field-collected larvae were identified through comparison with reared larvae. Identification of larvae and juve- niles was further confirmed by the following characters: pigmentation, body shape, an ad- vanced anus, and absence of a nasal cirrus. DISTINGUISHING FEATURES.—Preflexion and flexion larvae of C. globiceps may be distin- a i es ne WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 209 guished from other cottid larvae, except C. re- calvus and C. analis, by the presence of heavy pigmentation on the head, nape, and dorsolateral surface of the gut. Larval C. globiceps are distin- guished from C. recalvus and C. analis by the number (4-8) and spacing of ventral midline me- lanophores. Late flexion and postflexion larvae of C. globiceps differ from all other Clinocottus larvae in preopercular and parietal spination. Transforming and juvenile C. globiceps are distinguished from other cottid larvae by the combination ofa blunt, rounded snout and head, heavy pigmentation over the anterior third of the body, and two or three inconspicuous ventral midline melanophores which persist on the cau- dal peduncle. PIGMENTATION. — Newly hatched and preflex- ion larvae of C. globiceps have intense melanistic pigmentation on the head and nape. Eight to 11 large stellate melanophores are present over the midbrain and 21-30 melanophores are concen- trated in the nape region. These nape melano- phores are arranged in a distinctive pattern in which 7-10 melanophores are embedded along the dorsal midline of the nape and are surround- ed anteriorly and laterally by 14-23 dark mela- nophores lying on the external surface of the nape. Eight to 10 dendritic melanophores occur on both the anterior and posterior walls of the otic cap- sules. The dorsolateral surface of the gut is heavi- ly pigmented with 100-150 large, round mela- nophores. The only pigmentation occurring posterior to the anus is a series of 4-8 discrete, ventral midline melanophores. These are situ- ated under the 10 posteriormost myomeres near the tail tip. Frequently, 2-5 additional small me- lanophores extend beyond the tail tip onto the caudal finfold. Pigmentation changes little during larval de- velopment. The midbrain melanophores in- crease in number ranging from 12 to 16 in larvae >6 mm. By about 8 mm, melanophores are densely concentrated over the nape and extend anteriorly onto the head. Melanophores are added in the midbrain region and several melanophores extend anteriorly over the forebrain onto the snout. As head musculature develops, melano- phores in the otic region become obscured so that only 5 or 6 melanophores are visible on the posterior wall of the otic capsule. During transformation, in planktonic larvae 12-14 mm long, head pigmentation increases markedly. Several melanophores are added on the upper lip and beneath the orbit. Melano- phores are also added in a row along the pre- opercle and on the dorsal portion of the oper- culum. Pigmentation over the brain intensifies and expands posteriorly, merging with the nape pigmentation. Concurrently, nape melanophores extend ventrally from the nape forming a con- tinuous band of pigment between the nape and gut. Pigmentation also increases over the body cavity as melanophores extend ventrally over the lateral surfaces of the gut. Several melanophores also are added on the pectoral fin base. Ventral midline melanophores decrease in size and num- ber, until only two to four inconspicuous mela- nophores persist beneath the caudal peduncle. Newly settled benthic juveniles of C. globiceps are distinctively pigmented with the anterior third of the body covered with dark melanophores ex- tending posteriorly to about a vertical through the seventh dorsal spine. Only the posterior two- thirds of the pelvic fin rays remain unpigmented. Posterior to the intense head pigment, the two to four small, ventral midline melanophores con- stitute the only pigment. Between 14 and 16 mm SL, juvenile pigmentation is added posteriorly along the dorsum. By about 14 mm, a dark ver- tical bar of melanophores forms under the sec- ond to fourth dorsal fin rays and extends ven- trally two-thirds of the way below the lateral midline. Between 15 and 16 mm, three addi- tional saddles of melanophores are added pos- teriorly along the dorsum. The first saddle forms under the 8th-10th dorsal fin rays, the second forms under the 1 4th—1 5th fin rays, and the third saddle forms on the dorsal surface of the caudal peduncle. Concurrently, melanophores are added posteriorly along the lateral midline forming a dark band of pigment at the base of the caudal fin. Several melanophores appear on the pector- al, dorsal, and caudal fin rays. MorpHoLoGy.—Larval C. globiceps hatch at a relatively large size, 5.1-5.4 mm NL. Flexion of the notochord occurs between 6.2 and 8.1 mm NL. The largest planktonic larva taken in field collection is 12.9 mm and is beginning to undergo transformation. The smallest benthic juvenile is 13.5 mm long. Thirty-eight specimens (5.1-14.6 mm) were examined for developmental mor- phometrics. Because only 10 larvae were avail- able from field collections, this morphometric series includes 25 laboratory-reared larvae. 210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 26. Larvae of Clinocottus globiceps: A) 5.0 mm NL, B) 6.3 mm SL, C) 7.5 mm NL (B and C from Richardson and Washington 1980). Larval C. globiceps are relatively deep-bodied, and the posterior portion of the gut trails below the rest of the body. When viewed ventrally, the hindgut bulges slightly on either side of the anus similar to, but less pronounced than, the bulges in C. embryum. Relative body depth at the pectoral fin base increases during larval development from 20.7% in preflexion larvae to 28.5% SL in transforming larvae and juveniles. Larval C. globiceps have a notably blunt, rounded head and snout with relative head length increasing from 17.0% in preflexion larvae to about 31% SL in transforming juveniles. Fin DEVELOPMENT. — The smallest larva begin- ning to undergo flexion of the notochord is 6.2 mm long. Notochord flexion is complete in lar- vae between 7.5 and 8.0 mm long. Although cau- dal rays are present in late-flexion stage larvae (7.0-7.5 mm NL), the adult complement of 6 + 6 principal caudal rays is not countable until the completion of flexion at 7.4 mm. WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 211 ge Lop DDE ae, Ficure 27. Larvae of Clinocottus globiceps: A) 8.5 mm SL, B) 12.5 mm SL, C) 12.9 mm SL (B from Richardson and Washington 1980). Dorsal and anal fin bases are just beginning to form at completion of notochord flexion. The full complement of dorsal (13-17) and anal (11- 12) fin rays is complete at ~9.5 mm. The dorsal spines (VIII-X) are completely formed at ~10 mm. Development of the pectoral fin corre- sponds to that of the dorsal and anal fins. Pectoral fin rays are visible on a 7.5 mm larva. The adult complement of rays (13-15) is fully formed by ~9-9.5 mm. Pelvic fin buds are first visible in larvae between 6.5 and 7 mm long. The adult complement (1,3) is present between 9.5 and 10 mm. SPINATION.—Preopercular spines first appear as seven to nine small bumps along the posterior margin of the preopercle in larvae 5.5-6 mm. Larvae undergoing notochord flexion have 9-14 small, evenly spaced spines along the preoper- cular margin. During postflexion, spines increase in number from 16 to 22, and the dorsalmost spine becomes separated from the rest of the preoper- cular spines by a short gap. Simultaneously, this dorsalmost spine becomes longer and stouter than the other preopercular spines. In the largest planktonic larvae (12.5 mm SL) this dorsalmost spine is about 2.5 times as long as the other spines. The lower preopercular spines decrease in size and number during transformation. The uppermost spine continues to become longer and stouter in benthic juveniles and is over four times a 212 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 28. Juveniles of Clinocottus globiceps: A) 13.7 mm SL, B) 14.6 mm SL. as long as the lower spines in a 14.5 mm speci- men. The other preopercular spines are reduced to small bumps or serrations along the lower preopercular margin. In a 17-mm juvenile all remnants of the lower spines have disappeared and only the large dorsal spine persists. Clusters of spines develop on the head in the parietal region of C. globiceps larvae. One tiny spine is visible on each side of the head in larvae 6-7 mm and two spines are present on each side of the head in larvae 7-8 mm. By ~9-10 mm, five to six spines are present on each side of the head, arranged in a parallel pair of rows with two to three spines in the anterior row and three spines in the posterior row. These spines persist in the largest planktonic specimens examined (12.9 mm). In newly settled benthic juveniles, how- ever, these spines appear reduced and are present only as bony protuberances situated at the pos- terior margin of the parietals. Each protuberance has a hollow canal running through it which eventually forms the incipient cranial lateral line system in the parietal region of the head. Similar spine clusters also form in the supra- cleithral-posttemporal region. One or two small spines are first visible in larvae ~9 mm long. Five or six spines, arranged in two rows of three spines each, are present in both of the 12-mm specimens. These spines eventually become as- sociated with the lateral line system in benthic juveniles. Clinocottus analis (Figure 29; Table 6) LITERATURE.—Eigenmann (1892) and Budd (1940) briefly described and illustrated 4-mm specimens of C. analis. IDENTIFICATION. —Juveniles and adults were identified by the following combination of char- acters: an advanced anus, cirri, head shape, and pigmentation. Pigmentation, preopercular spi- nation, and body shape linked postflexion larvae of C. analis with juveniles and adults. DISTINGUISHING FEATURES. — Late postflexion and transforming specimens of C. analis were identified in collections from southern Califor- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 213 Ficure 29. Young of Clinocottus analis: 10.5 mm SL. nia. Apart from the two descriptions of newly hatched larvae, intermediate larval stages of C. analis are unknown. A brief diagnosis of post- flexion larval C. analis is presented in the hope that this information may facilitate the identi- fication of a complete developmental series of C. analis. Eleven postflexion larval C. analis (9.9-11.4 mm SL) were examined for developmental mor- phology, pigmentation, and spination. Clinocot- tus analis postflexion larvae may be distin- guished from all other larvae belonging to the Artedius, Clinocottus, Oligocottus groups by the intense band of melanistic pigmentation on the lateral body surface between the bases of the sec- ond dorsal and anal fins. Intense melanophores are also present on the dorsolateral surface of the head, the snout, the tips of the lips, and on the operculum. A patch of melanophores is present on the pectoral fin base and in a band on the dorsum beneath the spinous dorsal fin. Sixteen to 22 small, round melanophores are situated on the ventral midline posterior to the anus. Postflexion larval C. analis have blunt, round- ed snouts and relatively large heads. Snout length and head length are 28% HL and 30% SL, re- spectively, longer than in other Clinocottus lar- vae. In addition, C. analis larvae have moder- ately long, bulging guts. Snout to anus length averages 49% SL in postflexion larvae. Body depth at the pectoral fin base is 28% SL, while body depth at the anus is 26% SL. Six to 11 spines are present on the posterior margin of the preopercle. The dorsalmost spine is longer and stouter than the other spines. In the smallest specimens (9.9—1 1.0 mm) the spines are situated in two groups of three to five spines. The ventralmost spines begin to regress in larvae > 11 mm long; these spines decrease in size and num- ber and gradually become covered by skin. Two small spines are also present in the parietal region of the head in the 9.9 mm specimen. These spines decrease in size and remain only as bony bumps by ~11 mm. Artedius creaseri (Figures 30, 31; Table 7) LITERATURE.—Larval Artedius creaseri have not been previously described. IDENTIFICATION.—Juveniles and adults were identified by the following combination of char- acters: low dorsal fin ray (12-14) and anal fin ray (9-10) counts, low vertebral counts (30-31), scales extending onto head under the orbit and on the snout, and the presence of a preorbital cirrus. The developmental series was linked together primarily by preopercular and parietal spination, pigmentation, body shape, and meristics. Post- flexion and transforming larvae were linked to juveniles by the cirri pattern, pigmentation, body shape, and meristics. DISTINGUISHING FEATURES.—Preflexion lar- vae of A. creaseri are characterized by a pointed snout, large head, and relatively deep body. Dis- tinguishing pigmentation includes intense, large, round melanophores covering the dorsolateral surface of the gut, 1-3 large melanophores at the anteroventral margin of the gut, and a series of 7-11 large, evenly spaced melanophores along the ventral midline posterior to the anus. A large, distinctive, blotch-like melanophore is located on the ventral finfold near the tail tip, and another smaller melanophore occurs just beneath the tail tip. Larvae of A. creaseri >7 mm are further dis- tinguished by the presence of four large, evenly 214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 30. Larvae of Artedius creaseri: A) 5.0 mm NL, B) 6.6 mm NL, C) 7.9 mm SL. spaced preopercular spines and a prominent pa- rietal and nuchal spine. Late postflexion larvae may be recognized by their pointed snout and long jaw, large head, and the characteristic ven- tral midline pigmentation. In addition, meristics, especially the low dorsal fin, anal fin, and ver- tebral counts are diagnostic of this species. Small juveniles possess a long, slender nasal cirrus, a broad, ribbonlike postorbital cirrus with a fringed tip, and two pairs of frontoparietal cirri. PIGMENTATION. — Small preflexion larvae of A. creaseri are relatively lightly pigmented. They possess no melanistic pigmentation on either the head or the nape. Pigmentation over the dor- solateral surface of the gut is heavy and intense. Melanophores are large, round, and closely packed together. One or 2 melanophores are present on the ventral surface of the gut lying just posterior to the cleithrum. Posterior to the anus, the sole pigmentation consists of a series of 7-11 large, rounded melanophores evenly spaced along the ventral midline, positioned ap- proximately 1 to every three myomeres. This series originates under the third or fourth post- anal myomere and extends posteriorly toward the tail tip. The posteriormost 1 or 2 melano- phores in this series lie on the ventral finfold and are notable: large and blotchlike. Pigmentation increases markedly during larval development. Two melanophores form over the WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 215 Ficure 31. midbrain in larvae >5.7 mm. During flexion, melanophores increase in number and extend anteriorly onto the forebrain. By ~8.0 mm, the dorsal surface of the head is entirely pigmented. Pigmentation extends dorsally along the anterior wall of the gut so that 3 or 4 large melanophores lie just posterior to the cleithrum. Several me- lanophores form at the posterior margin of the gut in larvae >6 mm, frequently forming a ring around the anus. The number of postanal ventral midline melanophores ranges from 6 to 12, and the 4th or Sth melanophore in the series increases markedly in size and extends below the body wall onto the ventral finfold. The posteriormost 2 me- lanophores in the series move up onto the base of the caudal finfold. One is positioned just pos- terior to the lower hypural plate and the other lies just below the tip of the notochord at the dorsal base of the upper hypural plate. Transforming larvae (>10 mm) have mela- nophores extending ventrally along the preoper- cle and opercle and two to four melanophores Larvae of Artedius creaseri: A) 9.1 mm SL, B) 13.0 mm SL. on the lower jaw. Pigment is also added on the pectoral fin. MorpHoLoGy.—The smallest larval 4. crea- seri examined is 3.5 mm NL and recently hatched. Flexion of the notochord occurs between 5.7 and 7.9mm NL. The largest planktonic specimen is 13 mm and beginning to develop juvenile pig- mentation. The smallest benthic juvenile is 13.5 mm and still undergoing transformation. Thirty- two specimens ranging from 3.5 to 13.6 mm were measured for developmental morphology. Artedius creaseri larvae are relatively deep- bodied with distinctively large heads and pointed snouts. Body depth at the pectoral fin base av- erages 26% in preflexion larvae and increases to 29% SL in postflexion larvae. Relative head length averages about 25% in preflexion and flexion lar- vae and increases to 33% SL in postflexion lar- vae. Snout length remains 30% HL during larval development. Fin DEVELOPMENT. — Initiation of a thickening in the hypural region of the developing caudal TABLE 7. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Item Head length/SL: Artedius creaseri Preflexion 24.4 + 1.25 (22.9-26.5) Flexion 26.0 + 2.46 (22.8-28.8) Postflexion 32.6 + 3.80 (26.5—38.5) Juvenile 40.3 + 2.97 (38.2-42.4) Snout length/HL: Preflexion 29.7 + 2.74 (25.0-33.0) Flexion 31.9 + 3.58 (26.7-36.9) Postflexion 30.4 + 2.95 (25.1-35.5) Juvenile 23.2 + 1.20 (22.4-24.1) Eye diameter/HL: Preflexion 42.0 + 4.42 (34.7-47.3) Flexion 37.4 + 3.11 (32.5-43.1) Postflexion 33.2 + 4.01 (27.5-38.6) Juvenile 30.4 + 1.20 (29.6-31.3) Snout to anus length/SL: Preflexion 44.6 + 3.01 (41.5—-50.0) Flexion 43.5 + 3.83 (39.7-50.0) Postflexion 51.5 + 3.66 (44.6-55.8) Juvenile 54.4 + 1.77 (53.2-55.7) Snout to pelvic fin origin/SL: Preflexion - Flexion - Postflexion .5 + 2.17 (27.0-33.1) Juvenile 2.0 + 2.12 (30.5-33.5) Pelvic fin origin to anus/SL: Preflexion = Flexion - Postflexion 24.6 + 4.96 (17.5-33.9) Juvenile 22.4 + 0.35 (22.2-22.7) Body depth at pectoral fin base/SL: Preflexion 26.0 + 2.54 (23.4-29.7) Flexion 26.0 + 1.78 (22.7-28.1) Postflexion 28.8 + 2.67 (25.3-33.8) Juvenile 24.8 + 1.98 (23.4-26.2) Body depth at anus/SL: Preflexion 21.8 + 2.35 (19.4-26.0) Flexion 24.0 + 3.08 (19.3-28.1) Postflexion 28.2 + 3.07 (25.4-36.2) Juvenile 21.2 + 2.19 (19.6—22.7) Pectoral fin length/SL: Preflexion - Flexion 12.2 + 1.97 (9.5-15.1) Postflexion 23.7 + 6.11 (14.5-35.2) Juvenile 30.1 + 1.34 (29.1-31.0) — = Not present at this stage. * = Only one specimen available at this stage. fin is first evident at 5.7 mm, coincident with the 6 principal caudal rays is not present until larvae onset of notochord flexion. Caudal rays are pres- reach ~8.0 mm. Bases of the second dorsal (12-14) and anal ent at 6.4 mm, but the adult complement of 6 + Bopy Proportions OF LARVAE AND JUVENILES OF ARTEDIUS CREASERI AND A. MEANYI. Values are percent standard length (SL) or head length (HL) including mean, standard deviation, and range in parentheses. Artedius meanyi 18.6 + 1.77 (17.9-21.3) 22.7 + 3.53 (19.4-38.0) 32.1 + 4.68 (25.4-38.0) 27.5 + 7.50 (15.1-31.0) 29.5 + 5.01 (22.3-35.4) 29.4 + 4.34 (23.1-37.3) + i+ 44.0 + 3.39 (35.0-47.7) 36.6 + 4.93 (30.4-43.1) 29.0 + 3.31 (23.2-35.3) 33.0 + 2.86 (27.2-35.4) 39.6 + 2.97 (36.1-44.3) 48.3 + 4.43 (38.0-55.2) 23.0* 29.1 + 3.16 (21.9-33.6) 20.1* 19.4 + 1.60 (15.7-22.1) 18.0 + 1.64 (16.2-20.0) 19.9 + 1.25 (17.7-22.2) 24.6 + 3.65 (17.2-30.1) 15.2 + 3.11 (11.9-19.1) 18.9 + 1.55 (15.8-21.3) 24.6 + 2.73 (19.3-28.4) 8.4 + 1.35 (6.4-9.1) 9.2 + 1.77 (6.5-12.1) 19.0 + 6.21 (10.1-28.3) WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 217 (9-11) fins are countable in larvae ~7-8 mm long, and fin rays are formed between 9 and 10 mm. The adult complement of dorsal fin spines (IX-X) is first countable at 9.7 mm. Pectoral fin rays begin forming at ~7—8 mm, and the adult complement is present at 8.6 mm. Pelvic buds begin to form at >8 mm; however, the adult complement of rays is not present until larvae are >11 mm. SPINATION. —Artedius creaseri larvae develop prominent head spines. In contrast to Artedius, Clinocottus, and Oligocottus larvae which have multiple preopercular spines, 4. creaseri larvae develop four equal-sized preopercular spines. Two spines develop first on the posterior margin of the preopercle at >5.7 mm. At a state of growth between ~6.4 and 7 mm, two additional spines develop, one dorsal and one ventral to the original two spines. The middle two spines re- main slightly longer than the outer two through- out larval development. These spines persist through transformation and are present in ju- veniles. In larvae >10 mm, small basal spines or projections form on the base of each of the four main preopercular spines. With develop- ment, four bony ridges form on the inner shelf of the preopercle parallel to each basal spine. These ridges grow toward the basal spines and gradually fuse with them forming bony arches over the incipient lateral line canal of the pre- opercle. Prominent spines also form in the pa- rietal region of the head. A single parietal spine is first visible at 5.7 mm. By >9 mm, a second smaller parietal spine forms just posterior to the first. These spines are quite large and distinctive, and they are present in the largest planktonic larvae (> 13 mm long). When larvae reach ~8 mm, a spine forms in the supracleithral-posttemporal region. The su- pracleithral spine points dorsolaterally. A second supracleithral spine forms just ventral to the first and points dorsally. Larvae >9.5 mm form one posttemporal spine. These spines persist in the largest planktonic larvae (13.6 mm) but regress in young juveniles becoming incorporated into the developing lateral line canal system. Artedius meanyi (Figures 32, 33; Table 7) LiTERATURE.—Blackburn (1973) described a 4.3-mm larva resembling 4. meanyi, which he called Cottid 3. Richardson (1977) and Richard- son and Pearcy (1977) listed three larvae as Ice- linus sp. 1. Richardson and Washington (1980) illustrated and described specimens 3.3, 8.6, 10.9, 12.5, 13.5, 15.2, 16.5, and 16.6 mm long as Ice- linus spp. IDENTIFICATION. — Larval A. meanyi were mis- identified as Icelinus spp. by Richardson and Washington (1980) on the basis of meristics and the pelvic fin ray count of I,2, which is charac- teristic of Jcelinus. Meristics also match those of A. meanyi, which possess I,3 (rarely I,2) pelvic fin rays (Rosenblatt and Wilkie 1963; Lea 1974). Recently, Howe and Richardson (1978) reex- amined Lea’s specimens of 4. meanyi and re- ported that “... only one small specimen ap- peared to have two rays—all others had three rays.” Lea’s specimens were reexamined in this study. Cleared and stained specimens clearly have I,2 pelvic fin rays. The outermost ray is greatly thickened and branched at the tip in all speci- mens examined. All of the misidentified ‘‘/ce/i- nus” larvae possess this distinctive, thickened outer ray. In addition, during the present study, large transforming specimens of 4. meanyi were ob- tained that possess scales on the dorsal surface of the head, the opercle, and in four or five rows on either side of the dorsal fins. Specimens also possessed preorbital cirri and distinctive post- ocular cirri with three tentacles arising from a single base. The combination of these morpho- logical and meristic characters conclusively iden- tifies these transforming larvae and juveniles as A. meanyi. The developmental series was linked together primarily on the basis of pigmentation and body shape. DISTINGUISHING FEATURES. — Small preflexion larval A. meanyi are distinguished by their short, compact guts (snout to anus length averages 33% SL) and pointed snouts. Characteristic pigmen- tation includes a low number of ventral midline melanophores posterior to the anus (<13), sev- eral large melanophores situated anteriorly on the visceral mass at the base of the cleithrum, and two distinctive blotches of pigment on both the dorsal and anal finfolds. Notochord flexion begins at a relatively large size, ~6.2 mm in 4. meanyi larvae, and is com- plete by ~9.4 mm. Four large, evenly spaced spines form along the margin of the preopercle in postflexion larvae >9 mm. Two parietal spines develop at the posterior margin of each parietal in larvae >11 mm. Postflexion and juvenile 4. meanyi (13-18 mm 218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 Ficure 32. 1980). long) are distinguished by a low number of blotchy ventral midline melanophores posterior to the anus, a relatively pointed snout and large head (33% SL), a pelvic fin ray count of I,2 with the outermost ray thickened as if two rays are fused together, and other meristics. In addition, ju- venile A. meanyi possess a single, slender preor- bital cirrus, an eyeball cirrus, and a distinctive postorbital cirrus having three tentacles that arise from a single base. The largest specimens (16-— 18 mm) possess rows of prickle-like scales on the parietal, cheek, and opercular regions of the head and on the dorsal surface of the body and caudal peduncle. Larvae of Artedius meanyi: A) 3.3 mm NL, B) 8.6 mm NL, C) 10.9 mm NL (from Richardson and Washington PIGMENTATION. —Small A. meanyi larvae are relatively lightly pigmented. Melanistic pigmen- tation is absent on the head of preflexion larvae. Two to 5 round, external melanophores are clus- tered on the nape. The dorsolateral surface of the gut is lightly pigmented. Two or 3 large dendritic melanophores are embedded in the anterior musculature of the body cavity just posterior to the cleithrum. Posterior to the anus, a series of 7-13 large, blotch-like melanophores is posi- tioned along the ventral midline originating un- der the second to fourth postanal myomere and extending toward the tail tip. These melano- phores vary in size with the 3rd or 4th and the WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 219 Ficure 33. Young of Artedius meanyi: A) 13.8 mm SL, B) 15.2 mm SL, C) 16.5 mm SL (from Richardson and Washington 1980). posteriormost melanophores of the series being markedly larger and frequently extending onto the ventral finfold. Two large distinct pigment blotches are present on both the dorsal and ven- tral finfolds in small larvae. One specimen out of 45 examined possessed three pigment spots on both the dorsal and anal finfolds. Melanistic pigmentation increases during lar- val development. Several melanophores are added over the brain in larvae between 7.4 and 8 mm. Melanophores in the nape region become embedded in larvae >6 mm as body musculature develops. Pigmentation increases slightly over the lateral surfaces of the gut. With the onset of notochord flexion and development of the caudal fin, the posteriormost melanophore of the ven- tral midline series is characteristically positioned at the ventral margin of the forming caudal fin. A second, large melanophore is frequently added at the dorsal margin of the caudal fin base dorsal to the notochord tip. The blotches of pigment on the dorsal and ventral finfolds disappear in lar- vae >9 mm as fin rays begin to form. During transformation, between 13 and 19 mm, head pigmentation increases markedly. Me- lanophores extend anteriorly over the interor- 220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 9 bital region and onto the snout. Several mela- nophores are added just ventral to the orbit, between the eye and preopercle, and along the dorsal margin to the opercle. With development, melanophores are also added along the pectoral fin base, in a band across the dorsum, and on the dorsal fin around the first four dorsal spines. Melanophores are also added to the upper and lower lips, across the cheek, along the ventral margin of the opercle, and on the dorsal surface of the head. MorPHoLoGy.—The smallest larval 4. mean- yl collected from plankton samples are ~3 mm NL long and appear recently hatched. Larvae undergo notochord flexion between 6.2 and 9.4 mm NL. Specimens as large as 18-19 mm were collected in plankton and neuston tows. Plank- tonic larvae >15 mm are beginning to undergo transformation indicated by the development of juvenile pigmentation and the formation of scales on the head and dorsum. The smallest benthic juveniles examined were 15-16 mm long and were fully transformed. Thirty specimens, rang- ing in size from 3.3 to 17.9 mm, were examined for morphometrics. Small larval A. meanyi are relatively slender with a characteristic body shape. Body depth is constricted just posterior to the anus; the body bulges slightly in the midtail region and narrows again near the tail tip or caudal peduncle. This distinctive body shape remains apparent throughout larval development. The gut of 4. meanyi is short and tightly coiled. Snout to anus length averages 33% SL in preflexion larvae in- creasing markedly to 48% in postflexion larvae. Prior to flexion of the notochord, body depth averages 18% at the pectoral fin base and 15% SL at the anus and increases to 25% SL at both the pectoral fin base and the anus in postflexion larvae and juveniles. Artedius meanyi larvae have small heads with a distinctively pointed snout. Head length av- erages 19% in preflexion larvae, then increases dramatically to 33% SL in late postflexion larvae and juveniles. Snout length remains relatively constant throughout larval development, ranging from 28 to 30% HL. Fin DeEvELOPMENT.—The fins develop rela- tively late in A. meanyi. Initiation of notochord flexion begins at ~6.2 mm NL. Although caudal rays are first visible in larvae >7 mm long, prin- cipal caudal ray number (6 + 6) is not complete until after notochord flexion at ~11 mm. Dorsal and anal soft rays begin to form in larvae 9.5- 10 mm long. The full complement of fin rays is visible in larvae ~ 12 mm. Dorsal spines (IX—X) begin to form at ~11 mm and are all present by 12-13 mm. Pelvic fin buds form in larvae >9.5 mm; however, the adult complement of pelvic fin rays (I,2) is not complete until larvae reach ~12-13 mm. SPINATION.—Preopercular spines form rela- tively late in the development of A. meanyi lar- vae. Two tiny spines are first visible along the central portion of the preopercle in larvae >6.2 mm with a third spine forming dorsal to these spines between 8 and 8.5 mm. By 9.4 mm, a fourth spine is added at the ventral margin of the preopercle. These four spines remain prominent and approximately equal-sized throughout larval development. In larvae >13 mm, small basal spines or projections form on the base of each of the four main preopercular spines. With de- velopment, four bony ridges form on the inner preopercular shelf parallel to each basal spine. These ridges grow toward the basal spines and gradually fuse with them forming bony arches over the incipient lateral line canal of the pre- opercle. During transformation, between 15 and 17 mm, the dorsalmost preopercular spine be- comes longer and stouter than the other spines; however, all four preopercular spines remain clearly visible on the largest pelagic juveniles ex- amined (~19 mm long). Spines also develop in the parietal and supra- cleithral-posttemporal regions of the head. A sin- gle tiny spine first forms at the posterior margin of the parietal in larvae >7 mm long. This spine gradually becomes longer, and in larvae between 12 and 13 mm a second, smaller nuchal spine forms immediately posterior to it. A small spine forms on the dorsal margin of the posttemporal bone between 9 and 10 mm. A second, similarly sized spine is added ventrally on the posttemporal in larvae ~11 mm. Atabout the same time, a third spine forms posteroven- trally to the two posttemporal spines on the dor- sal portion of the supracleithrum. These three spines increase in size during transformation and eventually become associated with the junction of the cephalic and lateral line systems. ADDENDUM Since this study was accepted for publication, a subsequent review of cottid relationships has been published (Washington et al. 1984). Wash- WASHINGTON: RELATIONSHIPS AND ONTOGENY OF THE SCULPINS 2 ington et al. (1984) presented a hypothesis of phylogenetic relationships of cottoids and briefly summarized characters that supported eight pro- posed monophyletic groups of cottid genera. Much of this work was based on an unpublished manuscript (Washington and Richardson n.d.) in which a hypothesis of cottid relationships based on osteological characters of early life history stages was presented. Results of Washington et al. (1984) support the proposed monophyly of Artedius Group A, Clinocottus, and Oligocottus and the exclusion of Artedius creaseri and A. meanyi from this group. They listed five syn- apomorphic characters in addition to the multi- ple preopercular spines which support the mono- phyly of Artedius Group A, Clinocottus, and Oligocottus. These characters include anterior neural arches enlarged and elevated, arms of the anterior neural arches open in a broad u-shape un- til late in the juvenile period, a greatly expanded cleithrum base, posterior extensions or bony plates at the cleithrum base which enclose the pelvic bones, and loss of ventral postcleithrum and reduction or loss of dorsal postcleithrum. Washington et al. (1984) also placed Artedius creaseri and A. meanyi in a proposed monophy- letic group of genera including Jcelinus and Myoxocephalus as well as 10 other cottid genera. This placement gives additional evidence for the exclusion of Artedius creaseri and A. meanyi from the genus Artedius (sensu Bolin 1944) and sup- ports their close relationship to Jcelinus and Myoxocephalus. Further results of Washington et al. (1984) provide additional characters that strengthen the hypothesis of monophyly of Ar- tedius Group A, Clinocottus, and Oligocottus. ACKNOWLEDGMENTS I am indebted to many people for the loan of materials. Dr. Geoffrey Moser, National Marine Fisheries Service, La Jolla; Dr. Richard Rosen- blatt, Scripps Institution of Oceanography, La Jolla; Dr. Robert Lavenberg, Los Angeles Coun- ty Museum, Los Angeles; H. J. Walker, Ecolog- ical Marine Consultants, Solano Beach; Dr. Wil- liam Eschmeyer, California Academy of Sciences, San Francisco; David Rice, Lawrence Livermore Laboratory, Livermore; Dr. Arthur Kendall, Na- tional Marine Fisheries Service, Seattle; Dr. Theodore Pietsch, University of Washington, Se- attle,; Dr. Norman Wilimovsky, University of British Columbia, Vancouver; and Dr. Jeffrey Marliave, Vancouver Public Aquarium, Van- nN couver; all were helpful in making materials in their collections available to me. Special thanks are due Dr. Robert Morris for generously allowing me access to an unpublished manuscript on laboratory-reared Oligocottus snyderi larvae. I would also like to thank Kevin Howe, Joanne Laroche, Wayne Laroche, James Long, Bruce Mundy, Dorinda Ostermann, and Waldo Wakefield for volunteering their help in tidepool sampling. I gratefully acknowledge Dr. William Pearcy for his encouragement and for providing laboratory space. Special thanks are due to Kevin Howe for stimulating conversa- tions about cottid relationships, for reviewing parts of this manuscript, and for first teaching me about sculpins. I wish to thank Dr. Carl Bond, Wayne Laroche, and Dr. Sally Richardson for helpful discussions about cottids and for review- ing initial drafts of this manuscript. I gratefully acknowledge Margaret Snider and Lucia O’Toole for carefully typing drafts of this paper. 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Thesis, California State Univer- sity, Fullerton, California. 107 pp. Note: Since this paper went to press, a phylogenetic study of the Cottoidea has been published that addresses relation- ships of 42 cottid genera including Artedius, Clinocottus, and Oligocottus (Yabe 1985). A discussion of Yabe’s work will be included in a forthcoming paper (Washington and Rich- ardson n.d.). Yase, M. 1985. Comparative osteology and myology of the super-family Cottoidea (Pisces: Scorpaeniformes), and its phylogenetic classification. Mem. Fac. Fish. Hokkaido Univ., 32(1):1-130. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 4 as iv? « vcr se ro il Te Y 4 4 ii c255XxX NH PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 10, pp. 225-236, 5 figs., 2 tables. May 6, 1986 BILATERAL ASYMMETRY IN PHALLOSTETHID FISHES (ATHERINOMORPHA) WITH ee OF _ A NEW SPECIES FROM SELES By Lynne R. Parenti California-Academy of Sciences, Golden Gate Park, San Francisco, Californ 1° 4118 Apstract: Phenacostethus trewavasae, the first phallostethine from Borneo, is described from clay- and gravel-bottom freshwater streams of the Baram River, Sarawak. One of the distinguishing characteristics of P. trewavasae is a minute eye-lens. The subfamily Phallostethinae comprises Phallostethus Regan, known only from a single collection of one species; P. dunckeri Regan, from the mouth of the Muar River, in Johore on the Malay Peninsula; and Phenacostethus Myers, known previously from two species, P. smithi Myers and P. posthon Roberts, from coastal peninsular Malaysia and Thailand. Male phallostethids are bilaterally asymmetric. The subcephalic copulatory organ, the priapium, is oriented so that the aproctal side of the body is either the left or right; hence, males are termed sinistral or dextral, respectively. Both Phallostethus dunckeri and Phenacostethus smithi have, in about equal numbers, males that are either sinistral or dextral. In P. posthon, all males are dextral, whereas, in P. trewavasae, all males are sinistral. One species of neostethine, Mirophallus bikolanus Herre, is known in which all males are dextral. Bilateral asymmetry is compared among phallostethids to assess better the nature of this phenomenon, and its importance in determining the homology of priapial structures. INTRODUCTION Phallostethids are a group of some 20 known species of Indo-Australian atherinomorph fishes distinguished from all other teleosts by the pres- ence in males of the priapium, a complex, sub- cephalic copulatory organ (Regan 1913, 1916). Phallostethids have been divided into two groups (and classified traditionally in two families, Phal- lostethidae and Neostethidae, as in Roberts 1971a, b) on the basis of gross differences in priapial morphology. Rosen and Parenti (1981) and Parenti (1984) treated the entire group as one family, the Phallostethidae sensu lato, and that convention is followed here; the two groups are referred to as the subfamilies Phallostethinae and Neostethinae. The closest living relative of the freshwater, brackish, and occasionally salt- water phallostethids is hypothesized to be the monotypic western Pacific marine silverside (or hardyhead) Dentatherina Patten and Ivantsoff (Parenti 1984). The subfamily Phallostethinae includes two genera: Phallostethus Regan with one species, P. dunckeri Regan, 1913, and Phenacostethus Myers, with three species, P. smithi Myers, 1928, P. pos- thon Roberts, 197la, and P. trewavasae, de- scribed herein. Phallostethids are found throughout coastal peninsular Malaysia, Thailand, Borneo, the Phil- ippines, and Java. Phallostethus and Phenaco- stethus were known previously only from penin- [225] 226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 sular Malaysia and Thailand (see Roberts 197 1a, fig. 1). The third species of Phenacostethus, P. trewavasae, was collected from the Baram River in Sarawak, Malaysian Borneo. Fowler (1937) described Phenacostethus thai from a series of nine specimens, four males and five females (Academy of Natural Sciences, Philadelphia, ANSP 51352-51360). Roberts (1971a) followed Herre (1942) in treating Phenacostethus thai as a synonym of Phenacostethus smithi, and I ten- tatively concur. All Phallostethinae and Neostethinae males are bilaterally asymmetric with regard to orientation of the major supporting bones and other struc- tures associated with the priapium. Most females are bilaterally symmetric; the anus is just ante- rior to the urogenital opening along the ventral midline, under the throat (see Regan 1913, fig. 2). In just one species of neostethine are bilat- erally asymmetric females found (see Parenti, in press). Bilateral asymmetry is well documented in fishes (see Hubbs and Hubbs 1945, and refer- ences therein, fora comprehensive review). Most phallostethid species have both sinistral and dex- tral males in more or less equal numbers. Roberts (1971a) described Phenacostethus posthon, the first known species in which males are exclu- sively dextral. Phenacostethus trewavasae males are exclusively sinistral. Both species are known from relatively large numbers of specimens, such that unique or fixed asymmetry can only be in- terpreted as a natural phenomenon in some phal- lostethids. Phenacostethus trewavasae is also distin- guished by a minute eye-lens. The structure and function of the retina and, hence, the quality of vision, is unknown. MATERIALS AND METHODS The material on which the description of Phe- nacostethus trewavasae is based was made avail- able for study by Dr. E. J. Crossman, of the Royal Ontario Museum (ROM), where the holotype and majority of paratypes and additional specimens are deposited. Remaining paratypes have been deposited in the California Academy of Sciences (CAS), American Museum of Natural History (AMNH), British Museum (Natural History) (BMNH), and the United States National Mu- seum of Natural History (USNM), through the courtesy of Dr. Crossman. Included in the comparative material are spec- imens of Phallostethus dunckeri Regan from the single known collection by G. Duncker from the Muar River, in Johore on the Malay Peninsula (Duncker 1904). Regan’s (1913) description was based on seven specimens from this collection. Both Roberts (1971) and Parenti (1984) be- lieved that the only known specimens were the BMNH syntypes. However, additional speci- mens from the single collection by Duncker have been discovered in the Zoologisches Museum, Hamburg (ZMH) and have been made available for study through the courtesy of Prof. H. Wil- kens. Some of the ZMH material was given lec- totype and paralectotype status erroneously by Ladiges et al. (1958), who did not refer to the BMNH syntypes. Osteological structures were examined in, and counts made on, material counterstained with alcian blue and alizarin red S following the pro- cedure of Dingerkus and Uhler (1977), or solely alizarin stained. See text and Table 2 for catalog numbers of phallostethid material examined. Al- cohol-preserved (USNM 230367, USNM 230181), solely alizarin-stained (USNM 230371, USNM 230366), and counterstained prepara- tions (USNM 230374) of the western Pacific Dentatherina merceri were used for outgroup comparison. Additional comparative material was obtained on loan from the University of Michigan, Museum of Zoology (UMMZ) and the Museum of Comparative Zoology, Harvard Uni- versity (MCZ). A Zeiss SV8 stereomicroscope with drawing tube and photomicrography apparatus was used for dissection of specimens and recording of data. Phenacostethus trewavasae, new species (Figures 1-3, 4b) Ho.otyre.—ROM 41826, a mature, sinistral male, 14.1 mm standard length, collected 3 August 1981, by Dwight Wat- son, from Malaysia: Sarawak (Fourth Division), Baram River, Sungei Kejin Tugang, tributary of Sungei Kejin, depth to | m, clay- and gravel-bottom stream (03°41'30’N, 114°27'15”E). PARATYPES.—ROM 44289 (8 sinistral males, 11 females); ROM CS 812 (2 sinistral males, 1 female, all cleared and stained with alizarin red S); ROM 41827 (1 adult female), taken with the holotype. ROM 41829 (1 sinistral male); ROM 41830 (7 sinistral males, 6 females); CAS 55454 (3 sinistral males, 2 females); BMNH 1984.7.12:1-5 (3 sinistral males, 2 females); AMNH 55570 (3 sinistral males, 2 females); USNM 267266 (2 sinistral males, 3 females), all collected 11 February 1980, by Dwight Watson, from Malaysia: Sarawak (Fourth Division), Baram River, Sun- gei Kejin, station at confluence of Kejin Tugang and Kejin PARENTI: PHALLOSTETHID FISHES River, depth to | m, clay- and gravel-bottom stream, no vege- tation (03°41'30’N, 114°27'15”E). ADDITIONAL MATERIAL EXAMINED (no type status). -ROM 41828 (11 juveniles), collected 27-30 July 1981 from Malaysia: Sarawak (Fourth Division), Baram River, Loagan Titad. ROM 44290 (9 sinistral males, 17 females, 3 juveniles or of undetermined gender, of which 2 sinistral males and 2 females have been counterstained with alcian blue and alizarin red S); ROM 44291 (24 sinistral males, 12 females, 14 juveniles or of undetermined gender) taken with the holotype. DiaGnosis.— Phenacostethus trewavasae is distinguished from its sister species, P. posthon, by having only sinistral males, that is, with aproctal side of body on the left. The hooklike Ficure 1. Phenacostethus trewavasae, 14.1-mm holotype (ROM 41826). 227 oe pews af ‘ioe toxactinium arises on right side of head and curves very strongly under head towards left side of body. Males of P. posthon are exclusively dex- tral. Four characters distinguish the sister species from Phenacostethus smithi: distal portion of pe- nis smooth; penial bone absent; ctenactinium small or absent; and stout and distinctly curved, hooklike toxactinium (see Fig. 5). Males of P. smithi are either sinistral or dextral and occur in about equal numbers (see Introduction, Rela- tionships, Bilateral Asymmetry, and Table 2). A second diagnostic character is a minute eye- lens (Fig. 4b), as compared with the relatively large eye-lens of P. posthon and P. smithi (Fig. Ficure 2. Phenacostethus trewavasae, left lateral view of head and anterior portion of body of 14.1-mm holotype (ROM 41826). tN N oo i@ ® Ficure 3. Diagrammatic representation of dorsal mela- nophore pattern, Phenacostethus trewavasae, male paratype (ROM 44289). 4a), more typical of that found in diminutive teleost fishes (see below). DESCRIPTION. — Slender, laterally compressed diminutive phallostethid fish, distinguished by minute eye-lens and by males with sinistral pria- pium. Phenacostethus trewavasae like congeners in meristic data (Table 1). No vestigial pelvic fin rays or supports in females; males with pelvic and parts of pectoral fins modified into priapium that is invariably sinistral: prominent external- ized subcephalic bone a toxactinium arising on right side of body, articulating with right axial bone, and curving strongly under head towards left side of body (Fig. 4b). Cartilaginous pulvin- ular pad lateral to and covering articulation point of, toxactinium and axial bone; small antepleural bone just posterior to point of articulation. Mi- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 nute ctenactinium articulates with posterior base of right axial bone. Penis smooth not ruffled as in P. smithi (see Roberts 1971a). Pleural ribs of fifth? vertebra, each with a posterior flange, elongate dorsoventrally, meeting just dorsal to right axial bone (Fig. 4a). First pleural rib on fourth vertebra in females, fifth vertebra in males. Skull, gill arches, and jaws like those illustrated for Phallostethus dunckeri Regan by Parenti (1984), with following qualifications. Frontals project above dorsal head profile (Fig. 1, 2, 4b). Three infraorbital bones: preorbital, second in- fraorbital, and dermosphenotic. Outer jaws with few unicuspid teeth; paradentary with cartilagi- nous core and slight perichondral ossification, no teeth. Lower jaw protrudes beyond anterior ex- tent of upper jaw. Submaxillary element cartilag- inous. Rostral cartilage pear-shaped, wider pos- teriorly. Two small accessory cartilages between medial ramus of maxilla and rostral cartilage (as in Ceratostethus bicornis, Roberts 19715, fig. 5). Gill arch skeleton highly cartilaginous. Unicus- pid teeth on fourth ceratobranchial and infra- pharyngobranchial toothplates. Three cartilagi- nous basibranchials posterior to cartilaginous basihyal. Caudal skeleton with two epurals and autoge- nous parhypural. Caudal fin forked, dorsal and ventral rays forming incipient lobes. Pectoral fin narrow and elongate. Two dorsal fins, the first with a single spine or ray supported by single pterygiophore. Ventral dermal keel extending from base of priapium in males or urogenital opening in fe- males, to anal fin origin. Scales on body small and deciduous, absent from dorsal surface of head. Color pattern in alcohol similar to that of congeners (as in Rob- erts, 1971a, confirmed by personal observation): melanophores scattered on dorsal surface of head and anterior portion of body (Fig. 3), along mid- lateral intermuscular septum, around orbit, on operculum and priapium, and along basal por- tion of anal fin, dorsal midline, and ventral mid- line. Ground coloration very pale yellow or light brownish in alcohol; coloration in life unknown, although P. trewavasae is probably nearly trans- lucent in life, as are its congeners. Largest spec- imens reported by Roberts (1971a:13-14) of P. posthonand P. smithi with a bright orange yellow bar on caudal peduncle and a smaller orange yellow bar on the body “next to the anal fin origin.” PARENTI: PHALLOSTETHID FISHES EtymMoLocy.—trewavasae, in honor of Dr. Ethelwynn Trewavas, British Museum (Natural History), to express my deep appreciation of her continued contribution to the field of ichthyol- ogy. Eye-Lens SIzE The eye-lens is a nearly perfect sphere at the center of the eyeball (Fig. 4). In P. trewavasae (Fig. 4b), the eye-lens is minute compared with that of P. smithi (Fig. 4a). A minute eye-lens has been observed in all seven of the cleared and stained specimens of P. trewavasae. Four of these specimens were chosen at random from a lot of alcohol-preserved specimens for study: a mature male, an immature male, and two adult females (ROM 44290) that were counterstained for bone and cartilage. Three of the seven specimens— two mature males and one adult female (ROM CS 812)—stained solely with alizarin, were not prepared by me, but were probably chosen at random for preliminary identification at ROM. The presence of a minute eye-lens has been con- firmed, by dissection, in alcohol-preserved spec- imens. Size of the eye-lens varies from minute to barely detectable with a dissecting microscope, so that the character of a minute eye-lens may represent a stage in a transition series from a small eye-lens to eye-lens absent. The ratio of the distance between the center of the lens and the retina to the radius of the lens is nearly a constant in adult teleost fishes. This constant of 2.55, known as Matthiessen’s ratio, has been demonstrated in numerous teleosts, and has been confirmed in the cichlid Haplochromis elegans Trewavas (Otten 1981). During growth of H. elegans, the ratio increases rapidly from about 2.2 to 2.8, then decreases slowly to about 2.5 before leveling off at about 2.55 in the adult (Otten 1981, fig. 15). Matthiessen’s ratio in P. trewavasae could not be measured directly as part of this study. How- ever, a minute eye-lens at the center of the eyeball and a normal retina will not affect the distance from the center of the lens to the retina. But, obviously, Matthiessen’s ratio will be greatly in- creased by a small eye-lens radius, and the dis- tance from the center of the lens to the retina will be greater than the focal-length of the eye- lens. Visual acuity at any given stage in ontogeny is a function of retinal structure as well as shape of 229 the eyeball (Otten 1981; Levine and MacNichol 1982: Fernald 1985). Growth of the eye-lens is probably retarded very early in ontogeny. Struc- ture and function of the retina, as well as other accommodation made during ontogeny for a mi- nute eye-lens, is unknown. Without such infor- mation, the quality of natural vision in P. tre- wavasae will remain open to speculation, but several statements can be made. First, P. trewavasae may have poor visual acu- ity simply because of the optical properties of a minute eye-lens (Kirschfeld 1976). The short fo- cal length of the eye-lens can be correlated with low resolving power, decreased ability to distin- guish among wavelengths of light, and high chro- matic aberration. In very small lenses, absolute aperture limits resolving power (Otten 1981:681). Second, P. trewavasae lives in clay- and gravel- bottom, freshwater streams of the Baram River, Sarawak. The species is apparently omnivorous, with sample gut-contents including, for example, larval or juvenile P. trewavasae and adult dip- terans. Field notes state that there was no vege- tation at P. trewavasae collecting sites. If the mi- nute eye-lens limits the visual acuity of P. trewavasae, then we may assume that the species does not seek out prey visually. RELATIONSHIPS Myers (1928) distinguished Phenacostethus from Phallostethus by a shorter anal fin (Table 1), a protruding lower jaw, and the absence in the female of a groove on the abdomen. Regan (1913, 1916) did not state whether Phallostethus dunckeri has a spinous first dorsal fin, and Myers (1928) and Roberts (1971a, b) could only spec- ulate about its presence. The first dorsal fin is absent in the syntypes in the BMNH (Parenti 1984) and absent in the material in the ZMH. Further, Myers (1928) said that Phenacoste- thus resembled Phallostethus and differed from Neostethus (and, in fact, from all Neostethinae) in having a priapium that has a prominent hook- like anterior element (the toxactinium) and a shieldlike pulvinular pad. Nevertheless, these elements, or their homologs, are present in most phallostethids. They are well developed and hence are the prominent priapial elements in Phallo- stethus and Phenacostethus. Division of phallostethids into two groups em- phasized gross differences between the types of priapia. However, no assessment of whether one 230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 cs Figure 4. Left lateral view of head and anterior portion of body of cleared and stained preparations of male a. Phenacostethus smithi (MCZ 47299), and b. Phenacostethus trewavasae (ROM 44290), focused on eye-lens. Abbreviations: tox = toxactinium; pul = pulvinular pad; ant = antepleural; ax = axial bone. PARENTI: PHALLOSTETHID FISHES or both represented a derived priapium has been incorporated into a classification. For example, Roberts’s (19715:396) branching diagram of phallostethid genera clearly indicates a paraphy- letic Neostethinae. Furthermore, he interprets the neostethinae priapium as primitive (Roberts 1971b:395): “The priapium of Neostethinae, in which the only externalized elements are derived from pelvic spines and rays, is evidently more primitive than [the priapia of all other phallo- stethids].” Phallostethus and Phenacostethus together can be defined as monophyletic by the following shared derived characters (some characters mod- ified from Roberts 197 1a:5—6; his numbering not followed): 1. Slender, elongate atherinomorph fishes, with deciduous scales; diminutive—maximum stan- dard length recorded 17.0 mm (Roberts 1971a). 2. Body translucent or transparent, melano- phores scattered on top of head (Fig. 3), middle of dorsum, midlateral intermuscular septum, priapium, and basal and distal portion of fin rays (Fig. 1, 2). 3. Dorsum of head with translucent, membra- nous dome (not as noticeable in alcohol-pre- served specimens owing to dehydration). 4. Teeth on premaxilla, paradentary, and den- tary small, fewer in number than in other phal- lostethids and atherinomorphs; no large outer teeth on lateral ramus of premaxilla. 5. Main externalized bone of priapium a tox- actinium (Regan 1916; Myers 1928). 6. Large, oval, concave, cartilaginous pulvin- ular pad covering point of articulation of tox- actinium with axial bone (Regan 1916; Myers 1928). 7. Coiled vas deferens terminates in fleshy gen- ital pore or penis that projects from posterior section of priapium. 8. Pelvic spines or rays reduced or absent. 9. Vas deferens highly coiled, forming what has been termed an “‘epididymus”’ (Regan 1913, 1916). Most of these characters represent reductions; that is, we might think of Phallostethus and Phenacostethus as diminutive phallostethids that, perhaps because of small size, have lost or re- duced characters such as pelvic spines and fin rays, complete squamation and heavy pigmen- tation, and more complete, fuller outer dentition. However, the priapium of Phallostethus and Phenacostethus cannot be regarded as a reduced 231 9 ry v a > O = 3 2 s = rs rS) 3 ce e & a rr a 14 13 12 11 10 1-9 Ficure 5. Cladogram ofrelationships among the four species of Phallostethus and Phenacostethus. Black squares represent one or more synapomorphies; open squares represent one or more symplesiomorphies. Character 1) slender, elongate, di- minutive fishes with deciduous scales; 2) body translucent or transparent, melanophores scattered on top of head, middle of dorsum, midlateral intermuscular septum, priapium, and basal and distal portion of fin rays; 3) dorsum of head with translucent, membranous dome; 4) teeth on premaxilla, para- dentary and dentary small; 5) main externalized bone of pria- pium a toxactinium; 6) large, oval, concave, cartilaginous pul- vinular pad covering point of articulation of toxactinium with axial bone; 7) coiled vas deferens terminates in fleshy genital pore or penis; 8) pelvic spines or rays reduced or absent; 9) vas deferens highly coiled; 10) protruding lower jaw; 11) distal portion of penis smooth; 12) penial bone absent; 13) ctenac- tinium small or absent; 14) stout and distinctly curved hooklike toxactinium. See text for defining characters of each species. character complex. Several elements, including the pulvinular and the externalized toxactinium (homologous with the internalized secondary pulvinular; Roberts 1971), are more well de- veloped than in other phallostethids. The fact that pelvic fin rays at the base of the priapium are absent in Phallostethus and Phenacostethus, whereas rudimentary rays are present in neo- stethines, is interpreted as a derived condition representing a further modification of the pelvic fin supports and rays. Relationships among the four species of phal- lostethines are summarized in the cladogram in TABLE |. First Second dorsal rays dorsal rays Phallostethus dunckeri 0 8-10 Phenacostethus trewavasae 1 6 Phenacostethus posthon 1 5-6 Phenacostethus smithi 1 6-7 Figure 5. Phallostethus is readily defined by its extremely long anal fin, relatively high number of vertebrae (Table 1), serrated ctenactinium (Regan 1913), and absence of the first dorsal fin (Parenti 1984). The last two characters are also found in some neostethines. Phenacostethus is defined as monophyletic by the protruding lower jaw (Myers 1928). Myer’s additional character, absence of a groove on the abdomen of females, seems to be correlated with quality of preservation, and therefore is not used here. Phallostethus and Phenacostethus are not syn- onymized here solely for reasons of tradition. Phenacostethus trewavasae resembles P. smithi in that the first and second dorsal fins are sep- arated by a relatively large distance, as opposed to being rather close together, as in P. posthon (see Roberts 1971a, fig. 2, 3). The distance be- tween the first and second dorsal fin is a primitive character in P. smithi and P. trewavasae, and serves as a defining character of P. posthon. Roberts (1971a) described P. posthon as re- duced in a number of character states relative to P. smithi, then its only congener. Three character states that may be described as shared reductions in P. posthon and P. trewavasae are: (1) distal portion of penis smooth, as opposed to being TABLE 2. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 Meristic CHARACTERS OF PHALLOSTETHUS AND PHENACOSTETHUS Branchi- Pectoral ostegal Anal rays rays rays Vertebrae 26-28 9-10 4 40 14-15 9-10 5 34 14-15 9-10 4 34-35 14-15 9-11 4-5 33-35 ruffled as in P. smithi; (2) penial bone absent; (3) ctenactinium, if present, small and barely de- tectable. These characters were illustrated in both P. posthon (Roberts 1971a, fig. 6) and P. smithi (Roberts 197 1a, fig. 7). (Figures and captions were switched inadvertently when printed, as noted in a published erratum.) These three reductions are considered synapo- morphies of P. posthon and P. trewavasae. They are correlated with a stouter and more distinctly curved hooklike toxactinitum (compare Fig. 4a and 4b). Thus, even though the two species have characters that can only be described as reduc- tions, these characters are treated as synapo- morphies in phylogenetic reconstruction because of their correlation with uniquely derived char- acters (Fig. 5). BILATERAL ASYMMETRY A. Anatomical Homology Bilateral asymmetry is well documented in fishes (Hubbs and Hubbs 1945). The phenom- enon usually concerns reproductive structures offset to, or more complex on, one side of the body; although, the well-known asymmetry of flatfishes (pleuronectiforms) is not necessarily as- sociated with reproduction. BILATERAL ASYMMETRY IN FIvE SPECIES OF PHALLOSTETHIDAE (SENSU LATO) Dextral Sinistral Juveniles or Total males males Females undetermined Phallostethus dunckeri (a) 25 4 i 16 4 Phenacostethus trewavasae (b) 148 0 63 57 28 Phenacostethus posthon (c) 237 107 0 99 31 Phenacostethus smithi (d) 334 179 155 0 0 (e) 31 4 6 10 11 Mirophallus bikolanus (f) 300 L73 0 108 19 (a) BMNH 1913.5.24:18-20,21,22, ZMH 193-195; (b) total of known specimens, catalog numbers in text; (c) MCZ 47300, 47301 A,B, USNM 229302; (d) Hubbs and Hubbs 1945:290, table XIX; (e) BMNH 1927.12.29:1-10; (f) AMNH 50592, CAS 50722, CAS 53165, UMMZ 211665. PARENTI: PHALLOSTETHID FISHES Regan (1916:23) was the first to use the terms “sinistral’’ and ‘“‘dextral” to refer to the orien- tation of the priapium: “In its asymmetry and in being either dextral or sinistral the priapium agrees with the copulatory organ of Anableps,” a possibly bilaterally asymmetric killifish genus (see Hubbs and Hubbs 1945:289-291). Accord- ing to the convention established by Regan, in a sinistral male, the anal opening is on the right, what he termed the “proctal side.”” Hence, the left side ofa sinistral male is termed the “aproctal side.” The opposite is true for a dextral male, which may be thought of as the mirror image of a sinistral conspecific male. All female phallo- stethines examined exhibit no bilateral asym- metry; the anus is just anterior to the urogenital opening along the ventral midline under the throat. (See Parenti in press for report of bilateral asymmetric females in one neostethine species.) Regan (1916:21) also pointed out marked dif- ferences between priapia of phallostethines and neostethines: “The approximate symmetry of the priapial ribs and cleithra in Phallostethus, as compared with their marked asymmetry in Neo- stethus, is no doubt due to the symmetrical at- tachment of the priapium in the former... and its asymmetrical attachment . . . in the latter.” Proportions of sinistral and dextral males of Phenacostethus smithi were shown to be equal (Hubbs and Hubbs 1945; Table 2, herein). It was assumed that, with sufficient sample sizes, phal- lostethid species would be represented by more or less equal numbers of sinistral and dextral males, until Roberts (1971a) described Phenac- ostethus posthon, an exclusively dextral species. Roberts (1971a) termed P. posthon sinistral. However, it is more properly called dextral in keeping with homologies of priapia among all phallostethids. Confusion about whether to call a male sinistral (a left) or dextral (a right), may be traced to a casual statement by Myers (1928: 5): “The proctal side may be indifferently either the right or the left of the fish; in other words, the males are either ‘rights’ or ‘lefts.’”’ In inter- preting this statement strictly, one would assume Myers meant that a male with the proctal side on the right should be called a dextral, or a right. However, this is contrary to the terminology es- tablished by Regan and followed by most phal- lostethid systematists (e.g., Herre 1942; and Myers 1928 himself). Regan emphasized that the proctal side was away from the female during 233 copulation, and that the aproctal side, the side with the fleshy genital pore or penis, was the obviously functional side of the male with regard to internal fertilization (see also Villadolid and Manacop 1934). Herre (1942:139) followed Regan’s conven- tion but was ambiguous when describing asym- metry: “The coiled, enlarged vas deferens lies within the posterior end of the priapium, from which its penis-like tip projects. The proctal side may be either side, so that males of the same species may be either ‘rights’ or ‘lefts.’’? Hubbs and Hubbs (1945:290, table XIX) followed Re- gan strictly and documented bilateral asymmetry by tabulating the “location, left (L) or right (R), of aproctal side of males of Phallostethidae Phe- nacostethus smithi.” Division of phallostethids into two families was based primarily on the type of prominent external priapial bones. Phallostethines have a hooklike toxactinium that articulates with the axial bone (Fig. 4), a homolog of the pelvic fin girdle (Bailey 1936; Aurich 1937), and curves underneath the head toward the aproctal side (Regan 1916; Herre 1942; Fig. 2, 4 herein). Hence, a male phallostethine with a toxactinium arising on the right side of the head and curving toward the left, aproctal side, is called sinistral because the aproctal side is the left. Such sinistral phal- lostethine males also have a rudimentary cten- actinium that articulates with the left, aproctal axial bone at its posterior extent. The prominent externalized priapial bones of neostethines are the one or two ctenactinia that arise on the aproctal side of the body. Hence, a male neostethine with one or two ctenactinia arising on the left side of the body is termed sinistral not because of the position of these prominent priapial bones, but because the aproc- tal side is the left. This terminology should be adhered to strictly because of the consistency with the inferred homology of priapial structures. Fur- thermore, this convention for describing bilat- eral asymmetry of male phallostethids should be followed because the division between phallo- stethines and neostethines is not supported by unambiguous, derived characters. Roberts (1971a:13), in describing Phenaco- stethus posthon, was explicit in describing bilat- eral asymmetry: “. .. the priapium is invariably sinistral (toxactinium arising on left side) in the material examined.” It is clear, therefore, that 234 Roberts did not follow the convention estab- lished by Regan. Hence, I recommend that P. posthon be referred to as an exclusively dextral species, not sinistral as Roberts described it. This correction need not be made to Roberts’s (1971) discussion of the anatomy of Ceratostethus bi- cornis (Regan) because that species is a neo- stethine and the prominent external priapial bones are the ctenactinia, which arise on the aproctal side in every known example. B. Unique or Fixed Asymmetry Of the 148 known specimens of Phenacoste- thus trewavasae, 63 are sinistral males, 57 are females, and 28 are juveniles or of otherwise undetermined gender (see Diagnosis and Table 2). The collection of large samples of the exclu- sively sinistral Phenacostethus trewavasae allows us to conclude with certainty that unique or fixed asymmetry is a natural phenomenon in some phallostethids. In addition to the phallostethines P. trewavasae and P. posthon, the neostethine Mirophallus bikolanus Herre has males of fixed asymmetry (Tyson R. Roberts, personal com- munication). Three large lots of M. bikolanus, all collected from the Cabangan River, Albay Province in the Bicol (Bikol) region of Luzon, Philippine Islands, contain dextral males only. Of 300 specimens, 173 are dextral males, 108 are females, and 19 are juveniles or of undeter- mined gender (Table 2). The relatively high num- ber of male and low number of juvenile or un- determined M. bikolanus is probably related to the fact that immature males are readily iden- tifiable as such by the presence of a heavily pig- mented anal region. Unique or fixed asymmetry is a natural phe- nomenon in other atherinomorph fishes, as re- viewed by Hubbs and Hubbs (1945). Females of the ricefish Horaichthys setnai Kulkarni may be considered sinistral in that the urogenital open- ing is to the left of the midline in a majority of females, and the right pelvic fin girdle and rays are absent in females (Kulkarni 1940; Hubbs 1941). Males of H. setnai have an anal fin mod- ified into an elaborate gonopodium that is not bilaterally asymmetric as far as known. The vi- viparous poeciliids, Carlhubbsia kidderi (Hubbs) and Xenodexia ctenolepis Hubbs have a concav- ity on the right side of the gonopodium (Hubbs and Hubbs 1945). Males of the latter species also have a right pelvic fin modified into a so-called PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 “pectoral clasper” (Hubbs 1950), and a “... thickened fleshy ridge along the ventromesial edge of the proximal third of the outer ray of the right pelvic fin’ (Rosen and Bailey 1963:143). Phenacostethus posthon and P. trewavasae are sister species, the males of which are nearly mir- ror images of each other. Apart from type of bilateral asymmetry, they differ in several char- acteristics of priapial structure, placement of fins, and relative size of eye-lens (see Diagnosis, De- scription, and Relationships). One might assume that the common ancestor of these sister species, like most other phallostethids, contained both sinistral and dextral males (Fig. 5). One might assume further that it was the separation of the ancestral species into a sinistral and dextral pop- ulation that precipitated (or, in fact, was) the speciation event. The problem with such a spe- ciation hypothesis is that it presents a series of untestable statements, the first concerning states of the ancestral species, the second concerning isolation of sinistral and dextral subgroups. Experimental data are needed to answer the questions: What is the genetic basis of bilateral asymmetry in phallostethids? Does a male phal- lostethid determine type of bilateral asymmetry of offspring? That is, does a sinistral male have only sinistral male offspring, and likewise, does a dextral male have only dextral male offspring? Breeding experiments to answer these questions, performed when live phallostethids are available for study, will further our understanding of the evolution of bilateral asymmetry, and the special case of fixed or unique asymmetry, in phallo- stethids. CONCLUSIONS Phenacostethus trewavasae new species, is de- scribed from the Baram River, Sarawak. It is the first phallostethine species known from Borneo. This subfamily had been reported previously from Thailand and peninsular Malaysia. Two characters distinguish P. trewavasae from all other phallostethines: a minute eye-lens and males that are exclusively sinistral with regard to orientation of priapial structures. We may hy- pothesize reduced visual acuity in P. trewavasae because of the size of the eye-lens; however, a clear statement on vision awaits knowledge of structure of the retina. All phallostethid males are bilaterally asym- metric, described by position of the anus: in si- PARENTI: PHALLOSTETHID FISHES nistral males, the aproctal side is the left; in dex- tral males, the aproctal side is the right. Females exhibit no apparent asymmetry. In most species, sinistral and dextral males are represented in more or less equal numbers. Sam- ple sizes of the exclusively sinistral Phenac- ostethus trewavasae and the exclusively dextral P. posthon and Mirophallus bikolanus allow us to conclude with certainty that unique or fixed asymmetry is a natural phenomenon in some phallostethids. ACKNOWLEDGMENTS This study would not have been possible with- out the loan of recently collected phallostethid fishes from Sarawak generously provided by Dr. E. J. Crossman, ROM. For loans of additional comparative material, I thank Ms. M. Norma Feinberg and Dr. Gareth Nelson, AMNH; Dr. Barry Chernoff and Mr. William Saul, ANSP; Ms. Bernice Brewster, Dr. P. Humphry Green- wood, and Mr. Gordon Howes, BMNH; Mr. Karsten Hartel and Dr. Karel Liem, MCZ; Dr. Robert R. Miller and Mr. Douglas Nelson, UMMZ; Ms. Susan Jewett and Dr. Richard P. Vari, USNM; and Prof. H. Wilkens, ZMH. Dr. Tyson R. Roberts has continued to be a source of information on Southeast Asian fishes, and a willing discussant of all aspects of fish bi- ology and systematics. His comments and those of several reviewers greatly improved the manu- script. Members of the CAS Department of Ich- thyology staff provided assistance during the course of this study, including Dr. M. Eric An- derson, Dr. Stuart G. Poss, and Ms. Pearl M. Sonoda. Mr. Jim Patton, CAS Department of Photog- raphy, prepared the photographs in Figures | and 2. My studies of phallostethid fishes began during the tenure of a North Atlantic Treaty Organi- zation postdoctoral fellowship at the BMNH where I had the opportunity to benefit from the wisdom and experience of Dr. Ethelwynn Tre- wavas. Support of this project by the National Science Foundation through grant DEB 83-15258 is gratefully acknowledged. LITERATURE CITED Auricu, H. 1937. Die Phallostethiden (Unterordnung Phal- 235 lostethoidea Myers). Int. Rev. Gesamten. Hydrobiol. Hy- drogr. 34:263-286. Baey, R. J. 1936. The osteology and relationships of the phallostethid fishes. J. Morphol. 59(3):453-483. Dincerkus, G. AND L. D. Unter. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for demonstra- tion of cartilage. Stain. Tech. 52(4):229-232. Duncker, G. 1904. Die Fische der Malayischen Halbinsel. Mitt. Naturh. Mus. Hamburg 21:135-207. FERNALD, R. D. 1985. Growth of the teleost eye: novel so- lutions to complex constraints. Environ. Biol. Fishes 13: 113-123. Fow er, H. W. 1937. Zoological results of the third de Schauensee Siamese Expedition, Part VIII—fishes obtained in 1936. Proc. Acad. Nat. Sci. Philadelphia 89:125-264. Herre, A. W. 1942. New and little known phallostethids, with keys to the genera and Philippine species. Stanford Ichthyol. Bull. 2(5):137-156. Huss, C. L. 1941. A new family of fishes. J. Bombay Nat. Hist. Soc. 42:446-447. 1950. Studies of cyprinodont fishes. XX. A new subfamily from Guatemala, with ctenoid scales and a uni- lateral pectoral clasper. Misc. Publ. Mus. Zool. Univ. Mich- igan, no. 78:1-18. Huss, C. L. anp L. C. Husss. 1945. Bilateral asymmetry and bilateral variation in fishes. Pap. Michigan Acad. Sci. Arts Letters 30:229-310. KirscHFELD, K. 1976. The resolution of lens and compound eyes. Pp. 354-360 in Neural principles in vision, F. Zettler and R. Weiler, eds. Springer Verlag, Berlin. Ku karnl, C. V. 1940. On the systematic position, structural modifications, bionomics and development of a remarkable new family of cyprinodont fishes from the province of Bom- bay. Rec. Indian Mus. 42:379-423. LapiGes, W., G. VON WAHLERT, AND E. Mone. 1958. Die Typen und Typoide der Fischsammlung des Hamburgischen Zoologischen Staatsinstituts und Zoologischen Museums. Mitt. Hamburg Zool. Inst. 56:155-167. Levine, J. AND E. F. MAcNicHot. 1982. fishes. Sci. Am. 246(2):140-149. Myers, G. S. 1928. The systematic position of the phallo- stethid fishes, with diagnosis of a new genus from Siam. Am. Mus. Novitates, no. 295:1-12. Orten, E. 1981. Vision during growth of a generalized Hap- lochromis species: H. elegans Trewavas 1933 (Pisces, Cich- lidae). Neth. J. Zool. 31(4):650-700. Parenti, L. R. 1984. On the relationships of phallostethid fishes (Atherinomorpha), with notes on the anatomy of Phal- lostethus dunckeri Regan, 1913. Am. Mus. Novitates, no. 2779:1-12. . In press. Homology of pelvic fin structures in female Phallostethid fishes (Atherinomorpha, Phallostethidae). Co- peia. Recan, C.T. 1913. Phallostethus dunckeri, a remarkable new cyprinodont fish from Johore. Ann. Mag. Nat. Hist. 12:548- 555. Color vision in 1916. The morphology of the cyprinodont fishes of the subfamily Phallostethinae, with descriptions of a new genus and two new species. Proc. London Zool. Soc. 1916: 1-26. Roserts, T. R. 1971a. The fishes of the Malaysian family Phallostethidae (Atheriniformes). Breviora, no. 374:1-27. 1971b. Osteology of the Malaysian phallostethoid fish Ceratostethus bicornis, with a discussion of the evolution 236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 10 of remarkable structural novelties in its jaws and external genitalia. Bull. Mus. Comp. Zool. 142(4):393-418. Rosen, D. E. AND R. M. BaiLtey. 1963. The poeciliid fishes (Cyprinodontiformes), their structure, zoogeography, and systematics. Bull. Am. Mus. Nat. Hist. 126(1):1-176. Rosen, D. E. AND L. R. PARENTI. 1981. Relationships of Oryzias, and the groups of atherinomorph fishes. Am. Mus. Novitates, no. 2719:1-25. VILLADOLID, D. V. AND P. R. MANAcop. 1934 (Issued 1935). The Philippine Phallostethidae, a description ofa new species, and a report on the biology of Gulaphallus mirabilis Herre. Philippine J. Sci. 55(3):193-220. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 aoe PROCEEDINGS NH an OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 44, No. 11, pp. 237-267, 40 figs., 2 tables. May 6, 1986 LAND MOLLUSKS (GASTROPODA: PULMONATA) FROM EARLY TERTIARY BOZEMAN GROUP, MONTANA Barry Roth f i boc ae Museum of Paleontology, University of California, Berkeley, California 94720 | | y — * a cen ‘ Asstract: The Bozeman Group consists of fluvial, eolian, and lacustrine rocks deposited in intermontane basins of western Montana after the Laramide Orogeny. In the Three Forks Quadrangle, three early Tertiary formations have yielded land mollusk fossils. The Milligan Creek Formation (of probable Eocene age) con- tains snails of the genera Gastrocopta (two species), Radiocentrum, and Helminthoglypta; the Climbing Arrow Formation (middle or late Eocene) contains Gastrocopta and Polygyrella; the Dunbar Creek Formation (latest Eocene or early Oligocene) contains Gastrocopta (two species), Pupoides (two species), Radiocentrum, and Helminthoglypta. Three species of Gastrocopta, one of Pupoides (Ischnopupoides), two of Radiocentrum, and one of Helminthoglypta are described as new. Two others (Pupoides and Polygyrella) are scarcely distinguish- able from extant species. No interregional correlations are suggested because the land mollusk faunas of the western interior are too spottily known at present. In the Bozeman Group, genera and species groups that are now allopatric occur together. The land mollusks indicate a change in terrain through time: from sparsely vegetated to forested and back again. Climates were temperate and, at least toward the end of the interval, seasonally variable. Numerous land mollusk taxa in upper Cretaceous and Tertiary rocks of western North America occur outside the Holocene ranges of their families and genera. It is suggested that the evolutionary and biogeo- graphic history of North American land mollusks through the Tertiary has involved (1) the sorting out of component taxa into different geographic/adaptive zones; (2) restriction of many forms to lower latitudes, concurrent with climatic cooling; and (3) eastward and westward displacements, probably related to avail- ability of rainfall. For land mollusks, the late Eocene-early Oligocene was a time not so much of evolutionary innovation as of local extinction and biogeographic rearrangement. INTRODUCTION Land and freshwater mollusks from early Ter- tiary continental sediments of the Bozeman Group in the Three Forks Quadrangle, southwest Montana (Fig. 1), have been reported in check- lists by Taylor (in Robinson 1963; Taylor 1975). The terrestrial gastropods were not figured or discussed taxonomically; many were not iden- tified beyond family. However, the assemblage is an unusual one and bears strongly on the origins of present-day American land mollusk faunas. Preservation of the fossils ranges from fair to excellent. Seven species are represented by ma- terial good enough to permit description of them as new herein, and nearly all taxa can be char- acterized in greater detail. Table 1 presents a summary of the fauna. Robinson (1963) presented a detailed account of the geology of the Three Forks Quadrangle. He defined the Bozeman Group—which he an- ticipated would be recognizable on a regional scale—as the Tertiary fluvial, eolian, and lacus- trine rocks that accumulated in the basins of western Montana after the Laramide Orogeny. In the Three Forks Quadrangle the group consists mainly of four formations (Fig. 2). The Sphinx Conglomerate, stratigraphically the lowest, is a limestone conglomerate probably originating as an alluvial apron; it is not fossiliferous. Next lowest is the Milligan Creek Formation, consisting of light-colored, fine-grained, tuffa- [237] 238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 11 Index map of southwestern Montana. Three Forks Ficure 1. Basin diagonally hatched; other intermontane basins stippled. After Robinson (1961). ceous lake deposits that range from limestone to calcareous mudstone. Where it crops out, in the southwest part of the quadrangle, it produces whitish dissected benchlands. Its maximum TABLE |. thickness is probably not much greater than 90 m. Fossils from the Milligan Creek Formation include five taxa of land snails, all from one lo- cality (USGS 20007). Freshwater mollusks are evidently more widespread and are reported from five additional localities (Taylor 1975). The freshwater gastropod Lymnaea is also present at USGS 20007. Ostracods and charophyte re- mains occur in the formation. The age is prob- ably Eocene but none of the fossils is age-diag- nostic. The limestone and other fine-grained rocks of the Milligan Creek Formation were deposited in a perennial lake. Partly based on the snail here described as Radiocentrum laevidomus, Robin- son (1963) inferred that the lake basin lay in mountainous terrain not much different from that of the present. The Climbing Arrow Formation conformably overlies the Milligan Creek Formation in places but elsewhere may have formed contempora- neously with it. The Climbing Arrow is made up of olive, thick-bedded, sandy bentonitic clay and coarse sand, with subordinate light-colored silt- stone, sandstone, conglomerate, and limestone. OccCURRENCE OF LAND MOLLUSKS IN EARLY TERTIARY BOZEMAN GROUP IN THREE FORKS QUADRANGLE, SOUTHWEST Montana. Locality numbers are those of U.S. Geological Survey Cenozoic Series. Mil- ligan Creek Climbing Forma- Arrow tion Formation Dunbar Creek Formation Taxon 20007 20008 20009 20011 20012 20013 20014 20015 20016 20017 Class Gastropoda Subclass Pulmonata Family Pupillidae Gastrocopta (Albinula) montana n. sp. > 4 = - = ef. cf. - - — cf. G. (A.) sp. a >< = = — = = = — = = G. (A.) sagittaria n. sp. — a x _ — — — — - - G. cordillerae n. sp. = — = - X x = >, « ».¢ », « Pupoides (Ischnopupoides) tephrodes Nn. sp. _ — _ — — x — x x — P. (1) sp., cf. P. (1.) hordaceus (Gabb, 1866) = - - - xX - — = = — Family Oreohelicidae Radiocentrum taylori n. sp. - a - - = x », 4 Xx Xx xX R. laevidomus n. sp. x — — _ — — = — —- = Family Ammonitellidae Polygyrella sp., cf. P. polygyrella (Bland and Cooper, 1861) — xX xX — — - = — - - Family Helminthoglyptidae Helminthoglypta bozemanensis n. sp. —- — ? », 4 —- = - cf. cf. — ROTH: EARLY TERTIARY LAND MOLLUSKS It is extensively exposed in the Three Forks Quadrangle and tends to form subdued topog- raphy of low, rounded hills separated by broad, smooth valleys. The formation is not less than 220 m thick, and may be fully twice that. Two fossil localities (USGS 20008, 20009) have yield- ed three taxa of land snails. Freshwater gastro- pods are also present in the formation, including one, Physa?, from USGS 20009 (Taylor 1975). The Climbing Arrow Formation ranges in age from middle or late Eocene to early Oligocene. A small assemblage of vertebrates from the lower part of the formation was assigned a probable Uintan age (middle to late, but not latest, Eocene) (G. E. Lewis in Robinson 1963). Pipestone Springs (late Eocene) and Chadronian (latest Eocene to early Oligocene) vertebrates are known from higher in the formation (Hough and Lewis in Robinson 1963). A single locality well down in the lowest stratigraphic unit of the Climbing Arrow has yielded the freshwater planorbid gas- tropod Biomphalaria pseudammonius (Schlot- heim), diagnostic of middle to late Eocene age and a tropical climate (McKenna et al. 1962; Taylor 1985). This locality is several hundred meters stratigraphically below the Uintan ver- tebrate locality. The terrestrial snail localities are probably in the Uintan rather than the Chad- ronian part of the formation. A diverse fossil microflora exists but has not been studied (Rob- inson 1963). The Climbing Arrow originated largely as the product of an aggrading stream system. The coarser sediments are stream-chan- nel deposits; the finer-grained ones, evidently overflow deposits that accumulated on the flood- plains in short-lived ponds and lakes. The Dunbar Creek Formation, stratigraph- ically the highest named formation of the group, consists of white to grayish yellow thick-bedded tuffaceous siltstone, partly lacustrine and partly eolian in origin, laced with fluvial sandstone and conglomerate; minor limestone and bentonitic clay are also present. The formation is 80-240 m thick in the Three Forks Quadrangle and forms a topography much like that of the Milligan Creek—white benchlands rising steplike from the floodplain, dissected by many steep-walled can- yons. The constituent sediments were evidently deposited in a more or less enclosed local basin (part of the larger Three Forks structural basin) which, at least part of the time, contained stand- ing water. Whether the ash of a particular stra- 239 Land Mammal “Ages” Rock Units Series N. Amer. Dunbar Creek Formation Ww z Ww 1) (2) (2) 4d oO cS mo} & 3 = 750 JFMAMJJASOND Month 345 JFMAMJJASOND Ficure 1. Left: Seasonal distribution of Anepsius delicatulus at Owens Lake, Inyo County, California. (Based on data from Octo- ber, 1977 to January, 1979, courtesy of F. Andrews and A. Hardy, California Department of Food and Agriculture.) Right: Seasonal distribution of Batuliodes rotundicollis in the Eureka Valley, Mono County, California. (Based on data from October, 1977 to Janu- ary, 1979, from Andrews, et al. 1979.) Lawrence (1979) redefined Anepsiini to include Batuliini and suggested that Batuliodes should be placed in synonymy under Batulius. That classi- fication is followed here, except that I resurrect Batuliodes as a valid name for the reasons detailed below. Blackwelder (1945), and later Papp (1961), rec- ognized the subfamily Batuliinae without formal definition. From their checklists it is impossible to determine which taxa, if any, they intended to in- clude other than Batuliini. Relationships of Anepsiini to other groups of Tentyriinae are uncertain. The relatively small mentum is shared with tribes such as Stenosini, Coniontini, Lachnogyini, Cnemoplatiini, and Cryptochilini, as well as difficult to place genera such as /disia Pascoe. Divided eyes are wide- spread in Stenosini (undivided in Stenosis) and also occur in Typhlusechus, Alaudes, and Boro- morphus. A further character shared by Anep- siini and most Stenosini is the integration of the elytral bases and the scutellum into the collarlike mesothorax, which is amplected into the pro- thorax. The anterior, amplected parts of the ely- tra and the scutellum are depressed below the level of the elytral disk. The scutellum is visible only if the prothorax is moved forward, relative to the elytra. In tribes such as Coniontini a relatively small portion of the elytra is incorporated into the mesothoracic collar. In Cryptoglossini, a rela- tively large part of the elytra forms the dorsolat- eral parts of the collar, but the scutellum remains exposed on the elytral disk. I have not surveyed these complex structures extensively in tentyrioid Tenebrionidae, and their taxonomic value is un- certain. Final resolution of the relationships of the tentyrioid tribes will require detailed morphologi- cal comparisons across the entire subfamily. At present the Anepsiuni, Stenosini, and probably the Eurychorini (Koch 1955) may be considered to comprise a somewhat isolated clade within Tentyriinae. Relationships within Anepsiini were assessed by analyzing 27 characters across all the species. Character polarities were determined by out- group comparison with other tribes of Tenty- riinae. This procedure is rendered difficult by the large size and structural diversity of many of these tribes, which may themselves show more than one state for some characters. Characters and character-state polarities are discussed below and listed in Appendix 1. Antennal configuration (Characters 1-5). In Anepsius delicatulus the antennae are gradually enlarged to the 10th segment (Fig. 9), have no dis- tinct club, and are relatively long. In all other spe- 346 cies the last 3 or 4 segments are abruptly enlarged as a slight but distinct club (Fig. 7-8). Clavate an- tennae are commonplace in Tentyriinae and are therefore considered the primitive condition in Anepsiini. The two types of clubs probably devel- oped independently, since the lineages in which they occur differ in numerous other features. In burrowing species antennae are variably shortened. This derived condition, along with sev- eral other features correlated with a strongly psammophilous mode of life, has apparently arisen independently several times. Analogous shortening of antennae occurs in many other psammophilous Tenebrionidae. It is difficult to assign polarity to Character 5 (shape of apical antennal segment). Probably the subquadrate condition is associated with psam- mophily and shortening of the antennae. Tentorial configuration (Character 6). In most Anepsuni the tentorium consists of subvertical lateral laminae, connected posteriorly by a trans- verse bridge. This primitive condition prevails through a great majority of Tenebrionidae. In Ba- tuliomorpha, the posterior space between the transverse bridge and the ventral part of the oc- cipital foramen is closed by a sheet of cuticle (Fig. 2). As far as known, this state is unique to Batulio- morpha. Possibly it is correlated to the function- ing of the mouthparts, whose muscles attach in part to the tentorium. Epistomal margin (Character 7). Both truncate and emarginate epistoma are common in Tenty- riinae. However, the configuration in Batuliodes, with rather prominent lateral lobes and a nearly straight medial portion is distinctive and consid- ered derived. Submental gland (Character 8). This secondary sexual feature was discussed in some detail and il- lustrated by Doyen and Lawrence (1979). It is al- most certainly a synapomorphy in Anepsiini, but it is unclear whether its absence in Batuliodes rep- resents a primitive lack or a secondary loss. Pronotal configuration (Characters 9-12). In most Tentyriinae the pronotum has distinct angles and carinate lateral margins without fimbriation. This plesiomorphous condition pertains in Anep- sius. In Batuliodes, the posterior angles are strongly obtuse basally but are produced and acute or nearly right angled near the apex. In psammophilous species the pronotal margins be- come fimbriate, and the lateral carinae may be DOYEN: REVISION OF ANEPSIINI Ficure 2. Tentorium of Batuliomorpha comata, posterior oblique aspect; surrounding cranium removed. faint or lacking. These latter modifications appear in diverse groups of psammophilous Tentyriinae. Sculpture of hypomeron (Characters 13-14). Polarity of these characters is uncertain. Elytral fimbriation (Character 15). The ple- siomorphous, glabrous state occurs in nonburro- wing forms. Fimbriae very likely developed inde- pendently in all psammophilous lineages. Elytral sculpture (Character 16). As noted by Casey (1907:503), Anepsiinae are characterized by a peculiar pattern of elytral sculpturing. In each row the punctures are intersected anteriorly by a narrow longitudinal carinule. In Anepsius de- licatulus the anterior edges of the punctures are elevated, producing a characteristic pattern. The same pattern, albeit somewhat modified, is dis- cernable in all except the strongly fossorial spe- cies, suggesting that it is primitively present in the anepsiine lineage. Ventral setation (Character 17). Presence of se- tae is a derived condition correlated with psam- mophily. Mesocoxal closure (Characters 18-20). These characters were discussed in some detail by Doyen and Lawrence (1979). Mesocoxal cavities bounded laterally by the epimeron, with exposed trochantins, are primitive. Those closed by lobes of the meso- and metasterna are derived. How- ever, it Should be noted that, even in those Anep- siini where the epimeron reaches the coxal cavity, ius PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 347 the sternite lobes are closely apposed, constrict- ing the epimeron to a narrow strip. In some other groups of Tenebrionidae, notably Diaperini, me- socoxal closure is variable, possibly related to re- duction in body size (Doyen 1984). Casey (1907:503) previously realized the instability of this character in Anepsiini. Metasternum length (Character 21). In flight- less Coleoptera, the metasternum commonly be- comes much shortened. Although wing loss is ple- siomorphous in Anepsiini, the metasternum is relatively long in all genera but Anepsius, which is relatively primitive in most other features. Protibial configuration (Characters 22, 24). Ap- ical expansion of the tibiae is frequently associ- ated with psammophily in Tenebrionidae. Modi- fication of the tibial spurs is often an associated feature, and in extreme cases one spur may be en- tirely lost, as in Uniungulum Koch (1962:113). Likewise, fimbriation of the posterior protibial margin often accompanies these other changes. Aedeagal configuration (Character 25). The po- larity of this character is uncertain. The ventral surface (primitively dorsal in the noninverted ae- deagus) of the tegmen is often only lightly sclerotized; the amount of sclerotization some- times varies among individuals. Body proportions and setation (Characters 26, 27). Relatively slender, subglabrous bodies are associated with surface dwelling. Obese, fore- shortened bodies and long, projecting setae are associated with psammophilous, burrowing habits in many groups of tenebrionidae. These charac- teristics have probably evolved independently several times in Anepsiini. CLADISTIC RELATIONSHIPS One possible (hand-generated) cladistic ar- rangement of Anepsiini appears in Figure 3. This diagram includes the distribution of all characters for which the derived state is shared by at least two species even if the character polarity is not ab- solutely certain. Autapomorphous characters are described in the species treatments. At the outset it should be explained that psammophilous tene- brionidae that burrow in loose sand commonly share a syndrome of morphological specializa- tions. Important modifications, as discussed by Koch (1961), Doyen (1984:15), and others in- clude the following: enlargement of the foretibiae for digging; shortening of the antennae; increase in body pilosity, especially as lateral fimbriae; and development of obese, rotund body shapes. In ad- dition there is often development of protibial setal fringes, apparently to increase tibial area for dig- ging, and modification of the protarsi or protibial spurs. Parallelism in these characters is particu- larly evident in the Anepsiini, where most of the same apomorphies characterize psammophilous species of all four genera. Since the generalized body plan is similar throughout Anepsiini, the apomorphous states of these characters are dif- ficult or impossible to distinguish. However, sev- eral distinctive structural features, unrelated to psammophily, support the diagram in Figure 3. These are discussed where appropriate below. The characters subject to parallelism should ordi- narily be removed before computer analysis. I re- tain them here to emphasize the extensive level of homoplasy present in Anepsiini. The primary dichotomy separating Batuliodes from the remaining genera is based on differences in the mesocoxal region (Characters 18-20), an- tennae (Character 2), gular region (Character 8), and epistomum (Character 7). The sculpture of the hypomeron and the configuration of the poste- rior pronotal angles show derived states in the more generalized, surface-dwelling species. The absence of the derived states of these characters in some of the specialized burrowing species proba- bly represents a secondary absence. Exserted pos- terior pronotal angles also occur in Batulius seto- sus, but the majority of characters clearly show that B. setosus belongs in the Anepsius lineage. Batuliodes rotundicollis and B. confluens, the relatively generalized, surface-dwelling species, do not differ in important structural features. However, they differ in several details of cuticular sculpturing and diverge greatly in size and shape of the aedeagus (see species descriptions). The clade comprising B. wasbaueri, B. spatulatus, and B. obesus is united by a series of apomorphous character states related to a psammophilous mode of life. Batuliodes obesus is the most specialized member of this clade, having lost the pronotal ca- rina (Character 10), developed dense, long seta- tion on the venter (Character 17), and having en- tirely lost the elytral carinae (Character 16). In all other characters it is exceedingly similar to B. spa- tulatus. Batuliodes wasbaueri lacks apomorphies of the antennae (Character 5) and hypomeral ; Anepsius Batuliomorpha delicatulus 7) ) 3 2 2 2 =) © w oO co ———— o ) 7) = = 6» «© 2 = Ce o ¢€¢€ Oo 3 a ie CUS Cl a aes Se ie a oe A ee gw a = = 3 =} -“ 1c ce Do & OW « = a Ss > ¢ § oe = 2 2 8 & 6 = §—. 2a ~~ 2 5 gs 2 ao ©. 2 ro) a Ee 3 © ‘= £ @ © oe 2 -a 6 q (m2 4 4 DOYEN: REVISION OF ANEPSIINI ANTENNAL LENGTH (4) TENTORIUM (6) POST.PRONOTAL ANGLES (12) VENTRAL SETATION (17) SETAL LENGTH (27) PRONOTAL MARGIN (9) APICAL ANTENNAL SEGMENT (5) HYPOMERON SCULPTURE (14) ANTENNAL LENGTH (3) ELYTRAL MARGIN (15) PROTIBIAL SHAPE (22) PROTIBIAL SETATION (24) PROTIBIAL SPURS (23) ELYTRAL SCULPTURE (16) BODY PROPORTIONS (26) ANTENNAL SHAPE (1) METASTERNUM LENGTH (21) SUBMENTAL GLAND (8) PRONOTAL CARINA (10) POST. PRONOTAL ANGLES (11) AEDEAGUS (25) EPISTOMAL MARGIN (7) STERNITE LOBES (19) ANTENNAL SHAPE (2) MESOCOXAL CLOSURE (18) MESOTROCHANTIN (20) HYPOMERON SCULPTURE (13) 1 minute, tuberculate 2 geographically variable 3 short, bristling 4 intermediate Figure 3. Distribution of character states across genera and species of Anepsiini. One possible (hand-generated) cladistic arrange- ment is indicated by the basal connecting lines. Black dots indicate character states presumed to be apomorphic. Note the numerous parallelisms among features adapted for life in aeolian sand PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 349 sculpture (Character 14), but shares the remain- ing apomorphies. The alternate branch, comprising Anepsius, Batuliomorphus and Batulius, of the cladogram is unified by only a single character, the presence of a submental gland (Character 8) in males. This is shown as an apomorphy in Figure 3, but could also be a primitive feature of Anepsiini which has been lost in Batuliodes. Anepsius is distinguished by a single apo- morphy, the short metasternum (Character 21). Commonly, shortening of the metasternum Is cor- related with winglessness in Coleoptera. Anep- siini are primitively wingless, and in most features Anepsius (especially A. delicatulus) is the most primitive member of the tribe. Unexpectedly the metasternum is longest in the most specialized burrowing species, and it is possible that the short condition is actually primitive to Anepsiini. Within Anepsius, A. valens and especially A. montanus show fossorial specializations of the protibiae and presumably represent a monophy- letic clade. Batulius and Batuliomorpha, which constitute the remaining lineage, share numerous apo- morphic states of characters involved in psam- mophilous life. On this basis one might recognize only a single genus. However, Batuliomorpha is further characterized by a distinctive, obese body silhouette (Character 26), reduction or loss of the posterior pronotal angles (Character 12), and the uniquely apomorphic tentorium (Character 6). In contrast, in Batulius the body shape is not mark- edly different from that in Anepsius, and the ten- torium is unmodified, while the posterior pronotal angles are strongly exserted, resembling those of Batuliodes. These characters suggest that the apo- morphic protibial, setal, and antennal character states shared with Batuliomorpha arose by con- vergence. An alternative arrangement might derive the lineage containing Batulius and Batuliomorpha from one of the fossorial species of Anepsius. Both A. montanus and A. valens show a few of the apomorphic features related to psammophilous life. However, such an arrangement would re- quire that metasternal length, presumably de- rived in Anepsius, be reversed in Batulius and Ba- tuliomorpha. It seems more likely that the specializations obviously related to psammophily arose independently in the two lineages. Within Batuliomorpha, the species comata and tibiodentata share a single, apomorphous feature, the smooth, subglabrous hypomeron. However, this is acommon characteristic of psammophilous forms, and in the Anepsiini this characteristic oc- curs independently in Batuliodes. Apomorphies distinguish B. comata (loss of pronotal carina, Character 10) and B. tibiodentata (coarsely tri- dentate protibia; unique within Anepsiini, not shown in Fig. 3). Furthermore, B. comata occurs in the southern Mojave Desert, while B. tibioden- tata inhabits dunes in central and southern Baja California. Without more convincing evidence, it seems preferable to leave the relationships be- tween the three Batuliomorpha species unre- solved. Anepsiini LeConte Anepstini LeConte, 1862:215; Horn 1870:276; LeConte and Horn 1883:367; Casey 1907:503; Arnett 1960:670; Doyen and Lawrence 1979:346 Batuliini Horn, 1870:270; Casey 1907:497; Arnett 1960:670; Doyen and Lawrence 1979:346 (synonymy). Batulinae Papp, 1961:105. Batulini Papp, 1961:105. Wingless Tentyriinae 2-6 mm long with globu- lar prothorax and oval abdomen. FemMALEsS.—Lateral epistomal sutures usually distinct, medial suture obliterated. Eye com- pletely divided by epistomal canthus and super- tended by low carina; antenna with 11 segments, clavate or with apical 3 or 4 segments forming club; labrum transverse with long, slender tormae with medial processes directed obliquely poste- riad; mandible with small, smooth mola remote from bidentate incisor lobe; maxilla with galea densely setose; lacinia with bidentate uncus; la- bium subhexagonal, moderate in size, exposing maxillary articulations; tentorium with sides short, bridge thick, or closed posteriorly (Fig. 2). Prosternal process unmargined, horizontal a short distance behind coxae, then abruptly decli- vous; mesosternum scarcely excavated. Elytra constricted basally, collarlike, amplected into prothorax; scutellum angulate or rounded poste- riorly, retracted into prothorax, not visible on ely- tral disk. Mesocoxal cavities closed by sterna or nearly so; mesotrochantin exposed or concealed. Metasternum about one to two times length of mesocoxa. Metendosternite with short, thick stalk; long stout, tapering arms with tendons api- cal or subapical; median metasternal sulcus and 6 DOYEN: REVISION OF ANEPSIINI MESOSTERNUM / / — MESEPIMERON TROCHANTIN a“ ME TASTERNUM 5 Ficures 4-6. Ventral aspect of pterothoraces, showing variation in mesocoxal structure in Anepsiini. 4) Batuliomorpha comata, 5) Anepsius valens, 6) Batuliodes rotundicollis. internal ridge absent. Femurs stout; foretibia di- lated, triangular; tarsi short with few spinose se- tae beneath. Ovipositor short, thick, with para- proct and coxite subequal; coxite lobes indistinct; gonostyles minute, papilliform, inserted dorsola- terally very near apex of coxite. Maces.—Aedeagus inverted with paramere 1.1-1.7 times longer than tegmen; median lobe free, its lateral baculi fused proximally (Fig. 12- 15). Average length about 10% less than females; submentum perforated by circular opening with tuft of protruding setae (Doyen and Lawrence 1979; Fig. 16-19), except in Batuliodes. LarvAE.—Unknown. Key to Genera of Anepsiini i Middle coxal cavity with trochantin ex- posed; sternal lobes separated by nar- row space laterad of coxal cavity (Fig. — Middle coxal cavity with trochantin con- cealed; lobes of sternites meeting laterad of coxal cavity (Fig. 6) ......... af vied alorayrsrdeagseiors cee Ske Batuliodes Casey 2(1). Metasternum about twice as long as meso- coxa; pronotum and elytra fimbriate along lateral margins; apical antennal segment subquadrate or wider than long (BIS S78) ser csisce ond oetnsce scents — Metasternum about one to one and one- half times as long as mesocoxa; prono- tum and elytra subglabrous or with short, appressed setae; apical antennal seg- ment longer than broad (Fig. 9)........ hipehashisalague tom omeuees Anepsius LeConte 3(2). Posterior pronotal angles obliterated or represented by minute tubercles; anten- nal club with three segments; lateral fim- briation long, flying (Fig. 31); venter with numerous long setae .......... Batulio- morpha, new genus — Posterior pronotal angles strongly angu- late; antennal club with four segments; lateral fimbriation short, stiff (Fig. 21); venter with few short setae ............ Anepsius LeConte Anepsius LeConte, 1851:147, 1862:215; Horn 1870:277; Le- Conte and Horn 1883:367; Casey 1907:503. Type Spectes.—Anepsius delicatulus LeConte; designated by Casey 1907:501. Relatively slender to moderately obese (0.55 = EL/EW = 0.72) beetles devoid of long flying setae. Epistomal margin arcuately truncate or very feebly and gradually emarginate; lateral epi- stomal sutures obscured by sculpturing. Antennal length at least three-fourths head width; flagellum gradually enlarged to 10th segment (Fig. 9) or with terminal 3 segments enlarged to form slight but distinct club (Fig. 10); apical segment longer than broad. Submentum of males perforated by circular opening with tuft of protruding setae. Tentorium consisting of subparallel lateral lami- nae joined by posterior transverse bridge. Prono- tum moderately convex, about one and one-half times broader than long; anterior angles nearly right angled or slightly obtuse, angulate or with apexes briefly rounded; posterior angles broadly obtuse, angulate to briefly rounded, but always PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 351 distinct; lateral pronotal carina complete, never fimbriate. Elytra minutely carinate or punctate (montanus), with short, appressed setae (muinu- tus) or subglabrous. Epipleural carina very nar- rowly margined; epipleuron narrowed just behind humerus, then subparallel almost to apex. Meso- coxal cavities nearly closed by opposed meso- and metasterna; sternites subequal, not offset (Fig. 5); trochantins exposed. Metasternum length one to one and one-half times length of mesocoxa; venter subglabrous or with short, appressed se- tae. Foretibia moderately to broadly triangular (Fig. 16-18); protibial spurs subequal or mesal spur enlarged, curved posteriad. Key to the species of Anepsius ile Antenna about as long as greatest head width; anterior tibia narrowly tri- anpulan (Figs 16508) a. ese ee soe 2 — Antenna about three-fourths as long as greatest head width; anterior tibia broadly triangular (Fig. 17) ........... Jc. ea CIB Smother montanus Casey . Elytra and venter glabrous or nearly so. . .3 — _ Elytra and venter sparsely, evenly covered by short but evident setae ..... minutus, new species . Antenna with last 3 segments enlarged as distinct club; anterior tibia with lateral margin broadly scalloped; spines in cen- tral region of anterior margin separated by four to six spine widths (Fig. 18)..... Se ESR Ey ae nee valens Casey — Antenna gradually enlarged to segment 10; anterior tibia with anterior margin en- tire, spines in central region separated by about two spine widths (Fig. 16)..... PROC uate enol delicatulus LeConte Anepsius delicatulus LeConte (Figure 11) Anepsius delicatulus LeConte, 1851:148. Anepsius catenulosus Casey, 1907:505 (new synonymy) Anepsius atratus Casey, 1907:506. (new synonymy) Anepsius brunneus Casey, 1907:506. (new synonymy) Anepsius nebulosus Casey, 1907:507, (new synonymy) Anepsius bicolor Casey, 1907:507. (new synonymy) Anepsius deficiens Casey, 1907:507. (new synonymy) Relatively slender, subglabrous, reddish-black to black beetles with narrowly triangular proti- biae. FEMALE.—Cranium with elongate tubercles on epistomum, becoming tuberculopunctate poste- Olmm Ficures 7-10. Antennal configuration in Anepsiini. 7) Batu- liomorpha comata, 8) Batuliodes spatulatus, 9) Anepsius delica- tulus, 10) Batuliodes rotundicollis. riorly and punctate on back of vertex; postgena shallowly, reticulately punctate; mentum deeply, reticulately punctate. Antenna about as long as head width, gradually enlarged to 10th segment, without distinct club. Pronotal disk medially with punctures slightly larger than eye facets, separated by about one puncture diameter; laterally punctures becoming attended ectally by short, longitudinal carinules, producing tuberculopunctate appearance. Lat- eral carina transversely rugulose. Hypomeron sparsely, coarsely tuberculate, finely, longitudi- nally strigose or scabrous; prosternum and pro- sternal process coarsely punctate, asetose. Elytra seriately punctate with minute carinules intersecting punctures anteriorly; carinules faint, short medially, becoming longer and stronger in lateral two-thirds; interstrial surfaces minutely alutaceous. Metasternum and metepisternum shallowly, coarsely punctate; punctures separated 4B Ficure 11. Anepsius delicatulus, Kern County, California by about one puncture diameter medially, becom- ing closer laterally and subcontiguous on epister- num. Abdominal sternites sculpted like metaster- num; punctures denser on last two sternites. Femurs with sparse, short, appressed setae; polished or finely scabrous. Protibia narrowly tri- angular; lateral margin bearing row of coarse, blunt spines, densest near apical angle (Fig. 16); mesial margin with four to five coarse spines; pos- terior surface scabrous or rugulose, irregularly set with several coarse spines; apical spurs subequal. Meso- and metatibia with short, sharp spines on all but posterior surfaces. MaLe.—Differs as stated in tribal description. Aedeagus as in Figure 12. MEASUREMENTS.—EL 1,9-3.1 mm, EW 1.20-1.8 mm, PL 0.8-1.2 mm, PW 1.1-1.8 mm Ho.otype.—Sex undetermined; in the LeConte Collection (MCZ). Type Locaities.—Colorado River Valley (A. delicatulus), southern California (A. catenulosus); San Diego, California (A atratus); southwestern Utah (A. brunneus); southern California (A. nebulosus); Kern County, California (A. bicolor); near San Diego, California (A. deficiens). DOYEN: REVISION OF ANEPSIINI DiaGnosis.—Anepsius delicatulus differs from all other species in not having a distinct antennal club. Itis most similar to. A. minutus Doyen, but is nearly devoid of visible setae on the pronotum and elytra, whereas A. minutus is sparsely pubes- cent. In addition, the male genitalia are different in shape (Fig. 12, 13). The distribution of A. delicatulus (Fig. 20) is in arid and subarid habitats from Contra Costa County and northern Inyo County, California south through the Central Valley and Owens Val- ley to northern Baja, California and east to south- western Utah, central Arizona, and northern Sonora, Mexico. It occurs on rocky or sandy sub- strates and sometimes on aeolian dunes, as at the Ciervo Hills, Fresno County, California. The color varies from reddish black to black. Bicolored individuals, corresponding to A. bi- color Casey, with reddish prothorax and black ab- domen occur in several areas sympatrically with uniformly dark individuals. These color differ- ences are not correlated with differences in other features and may be partly related to age. Anepsius minutus, new species Slightly obese, brown, sparsely pubescent bee- tles with narrowly triangular protibiae. FeMALE.—Cranium set dorsally with nearly round tubercles slightly larger than eye facets; postgena finely scabrous posteriorly, becoming obscurely, coarsely punctate anteriorly; mentum coarsely, shallowly punctate; antenna about as long as head width; apical three segments en- larged as distinct club. Pronotal disk medially with punctures slightly larger than eye facets, separated by about one to two puncture diameters; laterally punctures at- tended ectally by short, sharp, longitudinal cari- nules, producing reticulate appearance; punc- tures laterally with short, appressed setae. Lateral carina narrowly margined, with sparse row of short, appressed setae. Hypomeron and pro- sternum finely scabrous with a few coarse, ob- scure punctures at base of prosternal process; ase- tose or with two to three short setae. Elytra with fine seriate punctures, each at- tended anteriorly by minute tubercle and short, appressed seta; interstrial surfaces minutely alu- taceous. Metasternum and metepisternum shal- lowly, coarsely and sparsely, setosely punctate, the latter obscurely so. Abdomen sculpted like metasternum, punctures denser on fifth sternite PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 353 Ficures 12-15. Male genitalia of species of Anepsius (ventral aspect, left; lateral aspect, right; median lobe, center). 12) A. delica- tulus, 13) A. minutus, 14) A. montanus, 15) A. valens. 354 DOYEN: REVISION OF ANEPSIINI Ficures 16-18. Right foretibiae of species of Anepsius, anterior aspect. 16) A. delicatulus, 17) A. montanus, 18) A. valens. and posteriorly on first four, forming subcontigu- ous rows near margins. Femora shining, with few obscure punctures and short, appressed setae. Protibia narrowly tri- angular; lateroapical margin with row of six to eight coarse, blunt spines, densest along apical angle; mesial margin with few fine setae; posterior surface finely sculpted, without spines; apical spurs subequal. Meso- and metatibia with sparse, short setae. Ma.e.—Differs as stated in tribal description. Aedeagus as in Fig. 13. MEasuREMENTS.—EL 1.6-1.7 mm, EW 1.1 mm, PL 0.6 mm, PW 1.0mm HoLotype.—Female from Laredo, Texas, Hubbard and Schwarz, Coll. (USNM) Paratype.—Male from Mexico, Nuevo Leén, 2 mi NNE China, V-24-1981, J. T. Doyen, on ground at night. DIaGNosis.—A. minutus is similar to A. delica- tulus LeConte in general appearance, but differs in its setose body (subglabrous in A. delicatulus). The holotype is mounted with a specimen of Myrmecocystus placodops Forel (Formicidae), but it seems unlikely that A. minutus is myrme- cophilous, since there are no obvious morphologi- cal modifications and no other species of Anep- siini are known to be closely associated with ants. Anepsius montanus Casey (Figure 19) Anepsius montanus Casey, 1907:504. Moderately obese, dark brown to black, sub- glabrous beetles with broadly triangular proti- biae. FeMALES.—Cranium with rounded, coarse tu- bercles on epistomum, becoming tuberculopunc- tate on vertex or punctate just before pronotum and with tubercles coalescing into carinules above eyes, producing reticulate appearance; postgena and mentum finely, closely punctate or puncta- torugose. Antenna about three-fourths as long as head width; apical three segments enlarged as dis- tinct club. Pronotal disk medially with irregular punctures slightly larger than eye facets, separated by one to three puncture diameters; laterally punctures be- coming about twice as large and, in lateral quar- ters, subcontiguous or contiguous. Lateral carina weakly crenate. Hypomeron finely scabrous with sparse, larger tubercles; prosternum and pro- sternal process shallowly, coarsely punctate with few long, projecting setae anteriorly. Elytra with punctures two to three times eye facet in diameter, separated by one to two punc- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 355 Ficure 19, Anepsius montanus, Lea County, New Mexico. ture diameters, arranged in rough striae but con- fused, especially near suture; interstrial surface smooth, polished. Metasternum shallowly, some- times obscurely punctate; punctures separated by about two puncture diameters medially, by less than one diameter on metepisternum. Abdominal Sternites sculpted like metasternum; punctures denser laterally near posterior margin of sternite four and on sternite five. Femora smooth, polished, with few small punc- tures and setae. Protibia broadly triangular (Fig. 17); lateral margin weakly crenate basally, abruptly broadened about two-thirds distance to apex, bearing about 12-14 very coarse, blunt spines, separated by two to three spine widths ba- sally, subcontiguous apically; mesial margin with sparse fringe of inclined setae about half length of tarsus; posterior surface concave, with few small irregularities and minute carina near posterior margin; mesial tibial spur much longer, stouter than lateral. Meso- and metatibia with few stout, sharp spines on lateral and posterior surfaces; slender setae on anterior and mesial surfaces. Mae.—Differs as stated in description of tribe. Aedeagus as in Fig. 14. MEasuREMENTS.—EL 1.9-2.4 mm, EW 1.4-1.7 mm, PL 0.8 1.0mm, PW 1.4-1.6mm Ho.otype.—Sex undetermined, from Greeley (Weld County), Colorado (USNM) ADDITIONAL MATERIAL EXAMINED (Fig. 20). Colorado, Otero County: La Junta, VI-24/25-1885 (1), VI-15-1896 (1). Fremont County: Florence, 12-10 (1). Larimer County: Fort Collins, 24-4 (2). Las Animas County: Trinidad, 6-25 (1). Pueblo County: Pueblo, VII-II-1934 (1). Weld County: Greeley (1), Nunn, VIII-22-1971 (6). Montana. (no additional data) (1). Nebraska. Sioux County: Glen, VIII-1903 (1). New Mexico. Lea County: just east of Caprock (1). Wyoming. Albany County: Laramie, III-18-1894 (1). Laramie County: Cheyenne, IV-23-1888 (2) Niobrara County: Lusk, VII-14-1937 (1); (on adiiona data) (1). Mexico. (no additional data; intercepted With cacti at No- gales, Arizona, II-21-1966) (1). J D1aGnosis.—A. montanus is superficially simi- lar to A. valens Casey, differing as indicated in the diagnosis for the latter. Very likely A. montanus inhabits aeolian sand. This habit is suggested by morphological adapta- tions such as the enlarged foretibiae, unequal pro- tibial spurs, and shortened antennae. Some of the collection sites are areas of extensive dune forma- tion, as at Caprock, New Mexico. Anepsius valens Casey Anepsius valens Casey, 1907:504 Moderately obese, dark brown or black, sub- glabrous beetles with narrowly triangular proti- biae. FeMALE.—Cranium tuberculate on epistomum, becoming tuberculopunctate posteriorly and punctate on back of vertex; postgena and mentum finely scabrous or rugulose; antenna about as long as head width; apical three segments enlarged as distinct club. Pronotal disk medially with punctures about one to two times eye facets in diameter, separated by about one puncture diameter; in lateral quar- ters punctures becoming attended ectally by short carinules, these carinules strongest and punctures shallowest near lateral margins; lateral carina even, punctate. Hypomeron finely scabrous, finely, longitudinally strigose; prosternum and prosternal process finely scabrous, sparsely, ob- scurely punctate, asetose or nearly so. Elytra seriately, somewhat irregularly punctate medially; punctures becoming attended anteri- orly by tubercles in lateral thirds, and then by minute carinules in lateral quarters; interstrial surfaces weakly undose. Metasternum shallowly, coarsely punctate; punctures separated by about 356 DOYEN: REVISION OF ANEPSIINI : 4. ——— eee = i *f a ee oy) oral ae = Y x Tae : bes : ac i ss if J ' : Aen : es ' ae eR Sea el : eee : $a L wena wh. : a : ° “ ' th + ' me ' . ¥ , meso. ; &: ; ee } i Fer ne fy : ’ a} ' Per ee ae : Og 8 Z é f ¢q {ag BF See “ : “Ss Hl ee = a P Tg UB me SR TE mie : i i B ° : : 9, ' @ °@ e €. ; . ty Bar eae °? é e ee ' @ : : ay a, v ' £ ’ ee a) a ; Pam? eo, 0 eee oe eS ee . ep “rena : @6@ ‘ 4 ee H 2 -N ‘ " e cy -4 MD ee. eo : ; Ep teste oo a i ie ae ey &% % . ‘SS ie} q H e® e-e : : i s H ' ' @ ao : B i 3 ' Es -. e : R i P ' : i ' axe. @ ge ; ' al / e @o i. i we RF , Siete besa ' pan keer a Re linea se @ delicatulus " *% _ O valens Sa a ® montanus H 4 & minutus ¥ XN \ A Ficure 20, Distribution of the species of Anepsius one to two puncture diameters medially, becom- ing closer, coarser, and shallower laterally; often obscure on metepisternum. Abdominal sternites sculpted like metasternum; punctures denser along posterior margin of third and fourth sterni- tes and on entire fifth sternite. Femurs shining, smooth, with few small punc- tures and appressed setae. Protibia narrowly tri- angular; lateral margin shallowly crenulate with about seven to eight coarse, blunt spines, densest along apical angle (Fig. 18); mesial margin with few short setae, sometimes with one to two PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 357 spines; posterior surface with few irregular cari- nules or tubercles; mesial tibial spur slightly larger, stouter than lateral. Meso- and metatibia with short, sharp spines set sparsely on all but pos- terior surfaces; posterior surfaces sparsely setose. Ma.e.—Differs as stated in description of tribe. Aedeagus as in Fig. 15. MEASUREMENTS.—EL 2.2-2.9 mm, EW 1.4-2.0 mm, PL 0.8- 1.1 mm, PW 1.3-1.7 mm. Ho.otype.—Sex undetermined, from Holbrook (Navajo County), Arizona, Wickham (USNM). ADDITIONAL MATERIAL EXAMINED (Fig. 20).—Arizona. (no further data) (6). Cochise County: Dragoon (4). Navajo County: Holbrook, VH-17-1940 (1), V-23-1941 (3), (no date) (12). DiaGnosis.—A valens is superficially similar to A. montanus Casey, but has much narrower ante- rior tibiae (Fig. 17, 18). In A. valens the antennae are about as long as the head is wide. In A. mon- tanus the antennae are no longer than three- fourths the head width. The habitat of A. valens is unknown, but the lack of structural modifications suggests that it is a surface-dwelling or litter-dwelling species, rather than psammophilous. Batulius LeConte Batulius LeConte, 1851:148; 1862:215; Horn 1870:270; Le- Conte and Horn 1883:364; Casey 1907:497, 498; Arnett 1960:670, Batulinus Papp, 1961:105 (misspelling). Type Species.—Batulius setosus LeConte [1851]; designated by Casey 1907:497. Relatively slender to moderately obese (0.61 = EW/EL = 0.72) beetles with pronotum and elytra fringed laterally with stiff, projecting setae. Epistomal margin truncate or very feebly and gradually emarginate; lateral epistomal sutures moderately impressed. Antennal length subequal to head width; terminal three segments enlarged as distinct club; apical antennal segment subqua- drate. Submentum of males perforated by circular opening with tuft of protuding setae. Tentorium consisting of subparallel lateral laminae joined by posterior transverse bridge. Pronotum moder- ately convex, about 1.65 times broader than long; anterior angles nearly right angled, apices briefly rounded; posterior angles broadly obtuse with apices exserted, slightly obtuse; disk abruptly de- clivous near margin; lateral carina complete, fringed with stiff, projecting setae. Elytra seri- ately tuberculopunctate, epipleural margin fringed with row of stiff, projecting setae about as long as protarsus. Mesocoxal cavities nearly closed by apposed meso- and metasterna; ster- nites not offset (as in Fig. 5); trochantins exposed. Metasternum length about twice length of meso- coxa; venter subglabrous except for few setae on prosternum. Foretibia broadly triangular (Fig. 23); mesial protibial spur larger than lateral and strongly curved posteriad. Batulius setosus LeConte Batulius setosus LeConte, 1851:148 Batulinus setosus Papp, 1961: 105 (misspelling) FeEMALE.—Cranium tuberculate on epistomum, tubercles becoming elongate posteriorly and an- teriorly or anterolaterally attending punctures, producing tuberculopunctate or reticulate ap- pearance on vertex; vertex often simply punctate posteriorly; postgena scabrous; mentum with few large, shallow punctures. Pronotal disk punctate or weakly tuberculo- punctate medially, becoming strongly tuberculo- punctate laterally, each puncture attended anteri- orly or anterolaterally by elongate tubercle or carinule; lateral carina finely serrate or crenulate, with sparse fringe of stiff setae about as long as protarsus; anterior border narrowly margined and setose but not serrate in lateral quarters. Hy- pomeron shining, very finely alutaceous with few, scattered setigerous punctures and sparse row of Ficure 21. Batulius setosus from Imperial County, California. 358 setigerous tubercles just below pronotal carina; prosternum sculpted like hypomeron, setae long- est medially; prosternal process becoming more densely punctate, especially along margins. Elytra seriately tuberculopunctate, alternate rows more strongly developed; tubercles near su- ture very small, and punctures very shallow, ill defined; laterally tubercles become larger and punctures become distinct though shallow; disk asetose medially; laterally and on declivity tuber- cles of alternate rows supertending stiff, inclined setae about one-third to one-half length of protar- sus; epipleural carina serrate anteriorly, becom- ing crenulate posteriorly; epipleuron asetose, smooth or faintly scabrous. Metasternum with sparse, coarse, shallow punctures, densest medi- ally, and each bearing a short, appressed seta and attended anteriorly by a very small tubercle; met- episternum very shallowly, obscurely punctate. Abdominal sternites one to three sculpted like metasternum, but punctures denser laterally; sternite four with punctures crowded near poste- DOYEN: REVISION OF ANEPSIINI rior margin; sternite five more densely punctate, except near anterior margin. Femora shining, with few small, shallow punctures. Protibia with irreg- ularly crenulate lateral margin, set with about 10— 12 coarse, blunt spines, sparse basally, becoming subcontiguous around angle; mesial margin bisin- uate, bearing three to five slender setae and one to several coarse spines; posterior surface rough, ir- regularly set with spines and short setae. Meso- and metatibia flattened, bearing short, sharp spines on anterior surface; slightly longer setae on posterior surface. Mace.—Differs as stated in tribal description. Aedeagus as in Figure 22. MEASUREMENTS.—EL 1.9-3.1 mm, EW 1.2-2.1 mm, PL 0.8- 1.3 mm, PW 1.1-1.9mm. Ho.otype.—Male from Gila River valley (MCZ). ADDITIONAL MATERIAL EXAMINED (Fig. 24).—Arizona. Yuma County: Fort Yuma, I-21 (3); 9 mi E San Luis, III-18-1980 (14); Tacna, IV-14 (2). California. Imperial County: Brawley, V-17-1967 (1); Glamis, IV-23-1972 (2), V-29-1971 (2); 1 mi N Glamis, IV-27/28-1972 (1); 3 mi N Glamis, IV-12-1973 (9); 7 mi SE Glamis, III-25/IV-8-1979 (18); Algodones Dunes, 2.5 mi NE O.3mm Ficures 22-23. Batulius setosus. 22) Aedeagus, ventral aspect (left), lateral aspect (right), and median lobe (center); 23) Right foretibia, anterior aspect. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 359 Coachella Bridge #1, IV-17-1979 (12). Riverside County: Blythe, V-6-1970 (1), IfI-10-1971 (1), IV-10-1971 (3); 1 mi W Blythe, V-23/24-1970 (6); 3 mi W Blythe, V-27-1971 (1); 18 mi W Blythe, I-29-1965 (1); Thousand Palms, III-12/24-1955 (3); Palen Dunes, IV-27-1978 (3). San Bernardino County: Amboy Crater, 1V-24-1960 (1). Mexico. Baja California (Norte): La- guna Salada, I-28-1960 (1); San Felipe, II-20-1954 (1). DiaGnosis.—Batulius setosus differs from all other Anepsiini in the bisinuate posterior margin of its protibia. It inhabits aeolian sand dunes or extremely sandy substrates in the southern Mojave and Col- orado Deserts. Collection records range from January to late May. Most specimens have been collected in pitfalls. Batuliomorpha new genus Relatively obese (0.70 = EW/EL = 0.80) bee- tles with long, flying setae. Epistomal margin feebly emarginate to slightly arcuate; lateral epistomal sutures weakly to mod- erately impressed. Antennal length no more than Ficure 24. Distribution of Batulius setosus. one-half head width; terminal three segments en- larged as distinct club (Fig. 7), apical segment subquadrate. Submentum of males perforated by circular opening with tuft of protruding setae. Tentorium consisting of subparallel laminae, closed posteriorly between transverse bridge and occiput (Fig. 2). Pronotum about 1.6-1.7 times broader than long, strongly convex, posterior margin depressed; anterior angles nearly right an- gled, rounded apically; posterior angles obliter- ated or marked by slight obtuse irregularity; lat- eral pronotal carina fimbriate, very narrow, carinate, or rudimentary, hypomeron sparsely se- tose and tuberculate laterally. Elytra tuberculate to tuberculopunctate, setose; epipleural carina in- dicated by regular row of closely set, setose tuber- cles. Mesocoxal cavities nearly closed by apposed meso- and metasterna; sternite lobes subequal or metasternal lobe about twice as broad as meso- sternal lobe at apex (Fig. 4); trochanters exposed. Metasternum length about twice length of meso- coxa; venter setose. Foretibia broadly triangular or macrodentate (Fig. 28-30); mesial protibial spur much larger than lateral, strongly curved posteriad. Type Species.—Batuliomorpha comata Doyen. Key to the species of Batuliomorpha ig Anterior tibia with lateral margin scal- loped (Fig. 28, 29), bearing row of coarse blunt'spines< «2... seaen ne oe — Anterior tibia with lateral margin produced as two very large, spatulate teeth (Fig. 30); margin without spines ............ CE reine eae tibiodentata, new species 1). Pronotum with lateral carina represented by row of small, discrete tubercles; lobe of mesosternum laterad of mesocoxal cavity much narrower than corre- sponding lobe of metasternum (Fig. 4) .. bo uttew) At aoe comata, new species — Pronotum with lateral margin finely cari- nate; lobe of mesosternum laterad of mesocoxal cavity subequal to corre- sponding metasternal lobe ............ Seeker imperialis, new species Batuliomorpha imperialis, new species Obese, dark brown, setose and laterally fimbri- ate beetles with proximal ends of lateral epistomal sutures subfoveate. 360 AA al DOYEN: REVISION OF ANEPSIINI 25 ees Olmm Ficures 25-27. Male genitalia of species of Batuliomorpha (ventral aspect, left; lateral aspect, right; median lobe, center). 25) B. comata, 26) B. imperialis, 27) B. tibiodentata FEMALE.—Epistomum arcuately convex or briefly truncate in middle; lateral epistomal su- tures deeply impressed, subfoveate at proximal ends; epistomum scabrous, sculpture becoming finely, rugosely punctate on vertex; epistomal canthus and postgena asetose; postgena coarsely, shallowly punctate. Pronotal disk colliculate-punctate, with short, sparse, appressed setae in lateral eighths and along posterior margin; lateral borders narrowly margined, crenulate, bearing sparse fringe of pro- jecting setae about one-half length of protibia; posterior angles very broadly obtuse, angulate; posterior border narrowly margined; prosternum and prosternal process coarsely, shallowly punc- tate, medially with few setae about one-half length of protibia. Elytral disk weakly undose and finely, seriately punctate, becoming more coarsely so laterally; al- ternate rows with tubercles supertending fine, in- clined setae about one-half length of protarsus medially to one-third to one-half length of proti- bia laterally; epipleuron narrowed abruptly about one-fourth distance from humerus, asetose. Meso- and metasternal lobes subequal laterad of mesocoxal cavity (as in Fig. 5); metasternum with large, shallow punctures, with few setae about one-half length of protarsus; metepimeron with large, shallow punctures; abdominal sternites sculpted like metasternum, but more densely se- tose; punctures denser on last two sternites. Profemur with sparse irregular posteroventral fringe of setae about one-half length of protarsus; meso- and metafemora with sparse anteroventral fringe of slightly longer setae; similar setae scat- tered on dorsal surface. Protibia with lateral mar- gin coarsely, shallowly crenate, bearing about five coarse blunt spines and about four closely set blunt spines at apical angle (Fig. 28); few coarse spines on posterior surface; row of spinose setae on mesial margin; mesial angle scarcely pro- duced; mesial spur much longer, stouter than lat- eral. Meso- and metatibia bearing longitudinal bands of sharp, coarse, projecting spines on an- terolateral surface; sparse fringe of long, inclined setae on posterior surface. MaLe.—Differs as stated in description of tribe. Aedeagus as in Fig. 26. MEASUREMENTS.—EL 1.7-2.4 mm, EW 1.35-1.8 mm, PL 0.7-0.9 mm, PW 1.2-1.6mm. Hototype.—Female (CAS) and 38 paratypes from Califor- nia, Imperial County, Algodones Dunes, 3.5 mi SE Glamis, II- 26-1978, F. Andrews, A. Hardy. ADDITIONAL PaRaTyPES.—Same data, IV-28-1978 (28); Glamis, IV-24-1972, pit trap, M. Wasbauer (1); 7.0-7.4 mi SE Glamis, II-19/TV-14-1979 (33). DiaGnosts.—Batuliomorpha imperialis is most similar to B. comata Doyen, but has the meso- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 361 0.3 mm Ficures 28-30. Right foretibiae of species of Batuliomorpha, anterior aspect. 28) B. imperialis, 29) B. comata, 30) B. tibiodentata. and metasternal lobes subequal and aligned la- terad of the mesocoxal cavities and has the ventral surface shallowly, coarsely punctate (sternal lobes offset in B. comata; ventral surface tubercu- late). The distribution of B. imperialis (Fig. 32) likely extends into the Gran Desierto, Sonora, Mexico, where some other endemics of the Algodones Dune system occur. Adult activity appears to be restricted to the winter and spring months. Batuliomorpha comata, new species (Figure 31) Obese, dark brown, setose and laterally fimbri- ate beetles with the sternal lobes laterad of the mesocoxal cavity unequal in width. FEMALE.—Epistomum shallowly emarginate; lateral epistomal sutures slightly impressed, sometimes subfoveate at proximal ends; episto- mum with short, longitudinal carinules, trans- forming into tuberculopunctate sculpture on ver- tex; epistomal canthus and postgena near eye with projecting setae about as long as protarsus; postgena faintly scabrous, asetose. Pronotal disk irregularly tuberculopunctate medially, each crescentic puncture impressed an- teriorly by small tubercle; becoming more Ficure 31. Batuliomorpha comata from Kelso Sand Dunes, San Bernardino County, California. 362 coarsely tuberculate in lateral sixths, along poste- rior margin and lateral two-thirds of anterior mar- gin; tubercles each subtending inclined, fulvous seta about as long as tibia; lateral carina indicated by regular row of setose tubercles; posterior an- gles obliterated; posterior border narrowly and obscurely margined. Prosternum smooth later- ally, finely tuberculate and on prosternal process set with setae about as long as tibia. Elytra seriately tuberculopunctate; elongate tu- bercles anteriad of shallow punctures, larger in al- ternate rows and supertending declined setae about as long as protarsus; becoming coarser and supertending longer setae laterally; epipleuron abruptly narrowed one-fourth distance from hu- merus, asetose. Metasternal lobe extending la- terad of mesosternal lobe at mesocoxal cavity (Fig. 4). Metasternum regularly set with round tu- bercles, each supertending long, declined seta set in very shallow puncture. Metepimeron glabrous. Abdominal sternites sculpted like metasternum, but more clearly tuberculopunctate, most densely so on fifth sternite. Femora finely, irregularly tu- berculate, tubercles supertending long fulvous se- tae. Protibia with serrate lateral margin (Fig. 29), each serration bearing large, blunt spine; five to six subcontiguous spines fringing outer apex; me- sial margin with row of short, stiff setae; mesial angle prominent; mesial spur much stouter and longer than lateral. Mesotibia bearing irregular double row of long, blunt spines on lateral mar- gin; long, fine, inclined setae on lateral and poste- rior surfaces. Metatibia bearing long, blunt spines on anterior and lateral surfaces, whorled with long, fine, inclined setae. Mate.—Differs as stated in description of tribe. Aedeagus as in Figure 25. MEASUREMENTS.—EL 1.7-2.7 mm, EW 1.2-1.9 mm, PL 0.6— 1.0mm, PW 1.0-1.6mm Ho.otype.—Female (CAS) and 8 paratypes from Califor- nia, San Bernardino County, Kelso Sand Dunes (8 mi SW Kelso), II-1-1977. J. Doyen, P. Rude, M. Bentzien (J. Doyen Lot # 77B1.1). ADDITIONAL ParatyPEs.—Kelso Dunes, I-13-1965, M. Irwin (5), IV-16/18-1974, F. Andrews, M. Wasbauer (19), II-8-1974, D. Giuliani (22), 11-13-1976, F. G. Andrews, A. Hardy (5); 2 mi S Kelso, XII-18-1977, J. Doyen (J. Doyen Lot # 77L4) (2); 9 air mi SW Kelso, VI-29/30-1978, J. Doyen, P. Rude (J. Doyen Lot # 78F4) (3); 10 air mi SW Kelso, IV-23-1977, J. Doyen (2); 2.5 mi E Kelso, II-13-1965, R. Dickson, M. Irwin (3) ADDITIONAL MaTERIAL EXAMINED (Fig. 32).—California. San Bernardino County: Dumont Dunes, 600’, V-1-1974, Creosote Assoc., T. Eichlin and A. Hardy (1), II-21-1974, D. Giuliani (1). Arizona. Mohave County: 3 mi SE Parker, VI-28-78, J. Doyen (J. Doyen Lot # 78F3) (1). DOYEN: REVISION OF ANEPSIINI DiaGnosis.—The unequal meso- and metaster- nal lobes and the rudimentary lateral carina of the pronotum distinguish Batuliomorpha comata from other Batuliomorpha, where the sternal lobes are equal and the carina distinct. Nearly all specimens of B. comata have been collected between December and April, suggest- ing winter activity of the adults. Most individuals have come from pitfall traps. One large collection was made from about the roots of sparse peren- nial grass (J. Doyen Lot # 77B1.1). At the Kelso Sand Dunes, where B. comata is common, collec- tion sites range from the flat apron of sand sur- rounding the main dune mass to the highest crests. Batuliomorpha Comata is known from three sets of dunes in the northeastern Mojave Desert (Fig. 32). Batuliomorpha tibiodentata, new species Obese, dark brown, setose and laterally fimbri- ate beetles with the anterior tibiae produced into two large teeth. FEMALE.—Epistomum arcuately convex or B® tibiodentatus O Imperiatis @ comatus Figure 32. Distribution of the species ot Batuliomorpha. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 363 nearly truncate medially; lateral epistomal su- tures lightly impressed; epistomum and vertex with short, longitudinal carinules between eyes, becoming more oblique anteriorly and poste- riorly. Epistomal canthus asetose; postgena with few projecting setae about as long as protarsus. Postgena finely tuberculopunctate, bearing a few long setae medially. Pronotal disk coarsely, closely punctate or weakly tuberculopunctate in medial one-fourth to one-third, becoming tuberculopunctate, then tu- berculate laterally and along posterior margin; tu- bercles in lateral one-sixth to one-third and in pos- terior one-sixth to one-fifth supertending erect, fulvous setae about two-thirds as long as protibia; lateral carina narrowly margined, finely crenate, bearing fringe of closely set, long, projecting se- tae; posterior angles obliterated; posterior border very narrowly margined. Prosternum and pro- sternal process shallowly tuberculopunctate, set with setae about two-thirds as long as protibia. Elytra very finely, seriately tuberculate; tuber- cles supertending inclined, fulvous setae about one-half to two-thirds length of protibia, slightly longer laterally; epipleural carina narrow, cre- nate, bearing row of closely set long setae; epi- pleuron gradually narrowed from base to apex, setose. Meso- and metasternal lobes aligned la- terad of mesocoxal cavity (as in Fig. 5); metaster- num closely set with coarse, shallow punctures bearing setae about one-third to one-half length of protibia. Metepimeron obscurely punctate, asetose; abdominal sternites sculpted like meta- sternum, but setae slightly longer. Profemur sparsely set with long fulvous setae on anterodorsal surface; set with shorter setae on basal anteroventral surface; meso- and metafe- mora sparsely set with long setae on anterior sur- face. Protibia with small basal, large medial, and very large apical scallop (Fig. 30); few coarse, blunt spines on basal posterior surface, two to three coarse, truncate spines on distal margin; two to three long setae on mesial margin; mesial angle slightly produced; mesial spur slightly larger than lateral. Meso- and metatibia bearing irregu- lar row of long coarse spines on lateral margins; posterior surfaces set with long, inclined setae. Mate.—Differs as stated in description of tribe. Aedeagus as in Fig. 27. MEASUREMENTS.—EL 1.6-2.1 mm, EW 1.2-1.5 mm, PL 0.7- 0.8 mm, PW 1.1-1.4 mm. Ho.otypr.—Female (CAS) and five paratypes from Mexico, Baja California del Sur, 7 mi SE Guerrero Negro, IV-8-1976, J. Doyen, P. Rude, R. Morrison; on dunes at night, J. Doyen Lot # 76D5. Two paratypes from Baja California del Sur, San Carlos, IX-25-1981, D. Faulkner, F. Andrews, sifted from sand dunes. DiaGnosis.—Batuliomorpha tibiodentata is dis- tinguished from all other Anepsiini by its very coarsely tridentate protibiae. The specimens from near Guerrero Negro were found crawling slowly on the surface of sand hum- mocks at night. Batuliomorpha tibiodentata is known from sand dunes at two localities in south central and central Baja California (Fig. 32). Batuliodes Casey Batuloides Casey, 1907:499; Arnett 1960:670. Batulius Doyen and Lawrence, 1979:347 (in part). Type Species.—Batulius rotundicollis LeConte, 1851; desig- nated by Casey 1907:498. Relatively slender to moderately obese (0.63 = EW/EL =<0.75) beetles with mesocoxal cavities closed by apposed meso- and metasterna. Epistomal margin feebly to distinctly emargin- ate (Fig. 33). Lateral epistomal sutures weakly to not impressed, often faint, obscured by sculptur- ing. Antenna 0.7-1.2 times longer than head width; terminal four segments enlarged as distinct club; apical segment longer than broad or subqua- drate. Tentorium consisting of subparallel lami- nae joined by posterior transverse bridge. Prono- tum moderately convex, about 1.4-1.6 times broader than long; anterior angles nearly right an- gled or obtuse, angulate to rounded; posterior an- gles very broadly obtuse basally, usually exserted and nearly right angled or slightly acute just be- fore apex; lateral pronotal carina complete or ab- sent, subglabrous or fringed with setae. Elytra mi- nutely carinate or tuberculate with seta arising behind each tubercle. Mesocoxal cavities closed by apposed meso- and metasterna; metasternal lobe laterad of coxal cavity broader than meso- sternal lobe (Fig. 6); trochantins concealed. Meta- sternum length about twice length of mesocoxa; venter subglabrous or sparsely setose. Foretibia moderately to very broadly triangular (Fig. 39, 40); protibial spurs subequal or mesial spur slightly larger, curved posteriad. Key to the Species of Batuliodes ile Anterior tibia narrowly triangular (Fig. 39); epipleural carina appearing es TOUS ONE ATLY!/ SO ta nrapenersceyaretetsye anes 2 — Anterior tibia broadly triangular (Fig. Hee 364 Ficure 33. Batuliodes rotundicollis from Inyo County, Cali- fornia. epipleural carina sparsely fimbriate with moderate to long setae, at least an- tETLONY .scc nor ososioms shematio ewes eens 3 2(1). Pronotal disk punctate in lateral quarters; intercarinal areas of elytra near epi- pleuron impunctate or nearly so; aedea- gus with paramere attenuate in apical one- third; much longer than tegmen (Fig. 34) asa eueaueryeus ey tants rotundicollis LeConte — Pronotal disk reticulate or reticulopunctate in lateral quarters; intercarinal areas of elytra distinctly punctate, even near epi- pleuron; aedeagus with paramere grad- ually attenuate to very acute apex; sub- equal in length to tegmen (Fig. 35) ..... «Mites everaystnw wees confluens (Blaisdell) 3(1). Antenna about three-fourths as long as head width; pronotum with lateral fringe DOYEN: REVISION OF ANEPSIINI of setae; pronotum with posterior angles strongly obtuse or rounded, ill de- fined ah. recast re cea eee 4 — Antenna about as long as head width; pro- notum without lateral fringe of setae; pronotum with posterior angles exserted at apex, nearly right angled............ aisles tendocuoeyestmee tne wasbaueri new species 4(3). Pronotum with lateral carina absent; hypo- meron with lateral row of setae about as long as protarsus............. obesus new species — Pronotum with lateral margin finely, nar- rowly carinate; hypomeron with lateral row of setae about half as long as pro- PATSUScir one seeks spatulatus new species Batuliodes rotundicollis (LeConte) (Figure 33) Batulius rotundicollis LeConte, 1851:148. Batuliodes rotundicollis Casey, 1907:499. Slender, pale to dark brown, subglabrous bee- tles with finely carinate elytra. FeMALE.—Epistomum with lateral lobes promi- nent, medial portion truncate (Fig. 33); set with nearly round to elongate tubercles anterome- dially, these becoming short carinules laterally and posteriorly; vertex tuberculopunctate; postgena and mentum with very shallow, obscure, nearly contiguous, coarse punctures. Antenna about as long as head width; apical segment longer than broad. Pronotal disk medially with punctures slightly larger than eye facets, separated by one to two puncture diameters; in lateral quarters ectal rims of punctures becoming raised as slight tubercles; lateral carina crenulate, asetose; posterior angles acute or nearly right angled, exserted at apex; an- terior angles angulate, nearly right angled. Hypo- meron with coarse, exceedingly shallow, often obscure, subcontiguous punctures, sometimes be- coming reticulate; prosternum shallowly, coarsely punctate; prosternal process with punc- tures mostly along margins. Elytra seriately, shallowly punctate, punctures interrupting fine, longitudinal carinae; near su- ture rows of coarser punctures with lower carinae alternating with rows of smaller punctures with more prominent carinae; laterally rows of coarser punctures becoming shallower, obscure or absent PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 36 near epipleuron; carinae becoming less pro- nounced laterally, obscure or absent near epi- pleuron. Epipleural carina weakly crenulate ante- riorly, becoming nearly straight posteriorly; supertended by irregular row of shallow punc- tures, these disappearing in posterior third. Meta- sternum with coarse punctures separated by about one puncture diameter; metepisternum glabrous; abdominal sternites more finely punctate than metasternum; sternites four and five with punc- tures mainly confined to posterior margins. Femora finely, sparsely tuberculate. Protibia (Fig. 39) rather narrowly triangular with about 8— 10 coarse, blunt spines along lateral margin; me- sial margin with few coarse spines and setae; pos- terior surface with irregular ridge and row of spines near mesial margin; tibial spurs subequal. Meso- and metatibia each with sparse row of sharp spines along lateral margin, few setae on an- terior and posterior surfaces. nm Mate.—Differs as stated in description of tribe. Aedeagus as in Figure 34. MEASUREMENTS.—EL 1.3-2.0 mm, EW 0.1-1.4 mm, PL 0.5- 0.8 mm, PW 0.8-1.2 mm Ho.otype.—Sex not determined; from Gila River valley (MCZ). ADDITIONAL MATERIAL EXAMINED (Fig. 41).—Arizona. Mari- copa County: Buckeye, 2-4-1942 (1); Phoenix, IT-9/III-16-1941 (5), H-2-1945 (1); 18 mi W Tonopah, IX-5-1978 (3). Mohave County: 3 mi N, 7 miE Littlefield, Virgin River, III-28/X-1-1982 (3). Pima County: Lukeville, X-26-1969 (2); Tucson, April (1) Yuma County: Ehrenberg, II-8-1939 (1), II-15-1940 (1); Fort Yuma, I-28 (2); Quartzite, II-27-1940 (4). California. Imperial County: Cargo Muchacho Mtns, Mesquite-Creosote, 480’, IV- 19/V-27-1979 (3); 3.9 mi N Walter’s Camp (1). Inyo County: Eu- teka Valley, 1978, March, (4), April, (13), May, (58), June, (36), July, (40), August (31), September (13), November/ December (1); Eureka Valley, 8 mi N, 4 mi W dunes, 3300’, IX-1-78 to V-5-1980 (77); Saline Valley, VI-21-1978/V-18-1979, 1100’ (6), 1200’ (37), 1360’ (28); Grapevine Canyon, IV-20- 1978/V-18-1979, 2500’ (4), 2700’ (5), 3800’ (1); NW end Saline Valley, Sand Dunes, 1150’, VI-6-1976 (1); Saline Valley Dunes, VI-6-1976 (14), IV-20-1975 (4); Inyo Mountains, Lead Canyon, if 34 35 : | gee 38 37 Ficures 34-38. Male genitalia of species of Batuliodes (ventral aspect, left; lateral aspect, right; median lobe, center). 34) B rotundicollis, 35) B. confluens, 36) B. wasbaueri, 37) B. spatulatus, 38) B. obesus. 3300’ V-5/VIL-13-1982 (12). Riverside County: Painted Can- yon, IV-15-1974 (3), various dates, V-18-1978/I-7-1979 (4); Riv- erside Mountains, Crest, Riverside Pass Rd., I[V-27/VII-18- 1978 (71). San Bernardino County: Daggett, X-17-1951 (1), San Diego County: Borrego, II-3-1939 (5); Carter Lake, III-23-1959 (1). Nevada. Nye County: Rock Valley, [V-11-1975 (1). DraGnosis.—Batuliodes rotundicollis is most similar to B. confluens (Blaisdell). In B. rotundi- collis the lateral areas of the pronotal disk are punctate and the intercarinal areas of the elytra near the epipleuron are impunctate or nearly so. In B. confluens, the lateral areas of the pronotum are irregularly set with short carinules, causing a reticulate appearance and the intercarinal spaces of the elytra are distinctly punctate, even later- ally. In addition, the aedeagi of these two species are different in shape and very different in size (Fig. 34, 35). Batuliodes rotundicollis occupies arid habitats from east central California and southern Nevada south to extreme southern California and east to central Arizona (Fig. 41). No collection records exist, but the beetles undoubtedly inhabit ex- treme northern Baja California and Sonora. Many collections are from aeolian dunes, but DOYEN: REVISION OF ANEPSIINI others are from sandy washes, areas of desert pavement, or stoney regions. Nearly all speci- mens have been collected in pitfalls. In contrast to the predominantly winter activity of most of the species inhabiting dunes, rotundicollis is most abundant during the warm season. Batuliodes confluens (Blaisdell) (new combina- tion) Anepsius confluens Blaisdell, 1923:243; 1943:218. Anepsius angulatus Blaisdell, 1923:244; 1943:218. (new synon- ymy) Slender, brown to black beetles with finely cari- nate elytra. FeMALE.—Epistomum truncate anteriorly or with lateral lobes slightly more prominent than middle; sparsely set with round or elongate tuber- cles anteriorly, becoming tuberculopunctate pos- teriorly on vertex and tubercles becoming short, sometimes anastomosing carinules, producing re- ticulate appearance, especially laterally above eyes; postgena and mentum with shallow, nearly contiguous, coarse punctures. Antenna about as long as head width; apical segment longer than broad. 0.3 mm Ficures 39-40. Right foretibiae of species of Batuliodes, anterior aspect. 39) B. rotundicollis, 40) B. spatulatus. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 367 Pronotal disk medially with punctures about two eye facets in diameter, separated by about one puncture diameter or less; laterally, ectal rims becoming raised as low carinules, these becoming strong and anastomosing in lateral thirds, produc- ing reticulate surface; lateral carina crenulate, asetose. Posterior angles acute or nearly right an- gled, exserted at apex; anterior angles angulate, nearly right angled. Hypomeron scabrous, longi- tudinally strigose, with scattered, coarse, shallow punctures, often obscure; prosternum and pro- sternal process shallowly, coarsely, subconti- guously punctate. Elytra seriately punctate; in alternate rows punctures interrupting fine longitudinal carinae; carinae weakest near suture, becoming very dis- tinct laterally, where anterior rims of punctures are raised; rows between carinae with each punc- ture attended anteriorly by slight tubercle, ante- rior rim slightly raised laterally; intercarinal areas becoming more weakly sculpted posteriorly and usually smooth on declivity. Epipleural carina weakly serrate or crenulate near humerus, be- coming nearly straight posteriorly, bearing sparse line of short, appressed setae; supertended by ir- regular row of obscure, shallow punctures. Meta- sternum with coarse, setigerous punctures sepa- rated by about one puncture diameter; setae short, appressed or declined; metepimeron ob- scurely, shallowly, and coarsely punctate; abdom- inal sternites sculpted as metasternum, some- times more finely so; sternites four and five usually with punctures mostly near posterior mar- gin. Femora with few fine tubercles or obscure punctures. Protibia nearly as in Batuliodes rotun- dicollis (Fig. 39), posterior surface with scattered tubercles, ridge near mesial margin usually indis- tinct; meso- and metatibia with few short spines and scattered tubercles on ectal surface and sparse, short, appressed setae. Mace.—Differs as stated in description of tribe. Aedeagus as in Figure 35. MEASUREMENTS.—EL 1.6-2.4 mm, EW 1.0-1.4 mm, PL 0.6-— 0.9mm, PW 0.9-1.2 mm. Hototype.—Male in the California Academy of Sciences, San Francisco. Type Locatities.—Mexico, Baja California Sur, Isla Par- tida, (B. confluens); Mexico, Baja California Sur, Loreto, Ar- royo Gua (B. angulatus). ADDITIONAL MATERIAL EXAMINED (Fig. 41).—Mexico. Baja California (Norte): Tijuana, [V-14-1942 (2); Arroyo Catavina, 35 mi S El Progresso, IV-2-1976 (1); 2.7 mi SE Catavina, VII-4- 1979 (8); 6.2 mi W Bahia de Los Angeles (31); 10 mi S Punta Prieta, VI-21-38 (2); 6.2 mi NE Rosarito, VII-10-1979 (20); 57 mi E El Rosario, 2 mi E San Fernando Velicata, VII-2-1979 (2); 2 mi NW Rancho Santa Ynez, III-27-1973 (3). Baja California Sur: 2 mi E San Ignacio, VII-6-1979 (6); Rancho Mesquital, 21.4 mi E San Ignacio, VII-9-1979 (11); Rancho Tablon, 13 km S Guillermo Prieto, [V-16/18-1983 (11); 12 mi S Guillermo Prieto, IV-7-1982 (1); 34.4 mi SE Guerrero Negro, IX-22-1981 (1); Isla Mejia, IV-20-1921 (1); Isla Carmen, V-23/VI-6-1978 (7); Isla Estanque, VII-1-1921 (1); Isla Monserrate, VI-11/23-1978 (3); Isla Raza, [V-21-1921 (1). DiaGnosis.—Batuliodes confluens is very simi- lar to B. rotundicollis LeConte, differing as stated in the discussion of the latter. The habits of B. confluens seem to be similar to those of B. rotun- dicollis, with occupation of many different sub- strates and adult activity through the hot part of the year. Batuliodes wasbaueri new species Moderately obese, brown, subglabrous beetles with very broadly triangular protibiae. FeMALE.—Epistomum with lateral lobes promi- nent, usually extending well beyond truncate mid- dle; sparsely, evenly set with round tubercles @ rotundicollis O confluens Ficure 41. Distribution of Batuliodes rotundicollis and B. confluens. 368 slightly larger than eye facets; tubercles becoming elongate posteriorly and carinulate posterola- terally near eyes; postgena coarsely, subconti- guously punctate; mentum scabrous; antenna about as long as head width; apical segment longer than broad. Pronotum medially with punctures about 1.5 times diameter of eye facets, separated by one to several puncture diameters; becoming denser lat- erally; punctures attended ectally by longitudinal carinules, strongest in lateral quarters; lateral ca- rina distinct, fine, immediately subtended by row of asperities, producing thickened appearance; posterior angles nearly right angled, exserted at apex; anterior angles angulate, slightly obtuse; hypomeron, prosternum, and prosternal process coarsely punctate; punctures separated by less than one puncture diameter. Elytra confusedly set near suture with shallow, ill-defined punctures about as large as eye facets, punctures attended anteriorly by small, slightly elongate tubercles; laterally punctures becoming smaller, very poorly defined, and tubercles be- coming more elongate, then carinulate in lateral thirds; carinules and punctures becoming obso- lete in alternate one to two rows adjacent to epi- pleuron; epipleural carina evenly margined, forming prominent, explanate humerus; sparsely fringed by short, declined setae; metasternum with coarse punctures separated by about one puncture diameter or less; metepisternum with few coarse punctures; abdominal sternites more finely punctured than metasternum; punctures mostly on posterior margins of sternites four and five. Femora finely, sparsely punctate: Protibia very broadly triangular (as in Fig. 40) with about 9-11 very coarse, blunt spines along lateral margin, subcontiguous around angle; mesial margin with four to five slender setae about one-fourth to one- third length of protarsus; posterior surface sparsely tuberculate; tibial spurs subequal. Me- sotibia with row of 2-4 stout spines along ectal margin, scattered spinose setae;, metatibia with few smaller spines on ectal margin, scattered spinose setae. Ma.e.—Differs as stated in description of tribe. Aedeagus as in Figure 36. MEASUREMENTS.—EL 1.7-2.4 mm, EW 1.2-1.7 mm, PL 0.7— 0.9 mm, PW 1.1-1.4mm. Ho.otype.—Female (CAS) and one paratype from Califor- nia, Imperial County, 5 mi N Glamis, IX-10-1974, M. Was- bauer, R. McMaster, pit trap. Additional paratypes as follows. DOYEN: REVISION OF ANEPSIINI California. Imperial County: Glamis, 1V-24-1972, M. Was- bauer, pit trap (3), V-29-1971, M. Wasbauer, pit trap (11); Al- godones Dunes, 12.4 mi ESE Holtville, 1V-13-1979 (1); Algo- dones Dunes, 2.5 mi NE Coachella Bridge #1, IV-17-1979 (1); Seeley, V-8-1970 (1) Paddock, Flock and Johnson. Mexico. Baja California (Norte), 20 mi S Palacio [= Palaco?], 1V-4-1939, E. Ross (2). DiaGnosis.—Batuliodes wasbaueri is similar to B. confluens (Blaisdell), but has the protibiae very broadly triangular (much narrower in B. con- fluens) (Fig. 39, 40). Batuliodes wasbaueri is simi- lar to B. spatulatus Doyen, differing as indicated in the discussion of the latter. Batuliodes was- baueri appears to be endemic to the Algodones Dunes, although it lacks most of the morphologi- cal specializations that distinguish the truly psam- mophilous species such as Batuliodes obesus and the species of Batuliomorpha. Batuliodes spatulatus new species Brown, strongly convex, obese beetles, with in- conspicuously setose elytra and very broadly tri- angular protibiae. @ spatulatue O ocbesus O waebauer! Ficure 42. Distribution of Batuliodes spatulatus, B. wasbaueri, and B. obesus. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 369 FEMALE.—Epistomum with lateral lobes slightly more prominent than middle, which is truncate or concavely arcuate; set with round tu- bercles slightly larger than eye facets, these be- coming elongate posteriorly on vertex and often carinulate; postgena and mentum shallowly, coarsely punctate. Antenna about three-fourths as long as head width; apical segment about as long as broad. Pronotum medially with shallow punctures slightly larger than eye facets, separated by one to several puncture diameters; attended ectally by low tubercles, these becoming stronger and longi- tudinally elongate laterally and forming short car- inules near lateral margins; lateral carina very fine, narrow and even; posterior angles strongly obtuse but definitely angulate; anterior angles rounded. Hypomeron glabrous except for few punctures on coxal cowling and band of setigerous tubercles just below pronotal carina; setae stiff, slightly curved, about one-third to one-half as long as protarsus; prosternum with few setigerous punctures medially with anterior rims tubercu- lately raised; setae about as long as protarsus; prosternal process with few marginal punctures. Elytra seriately tuberculate, tubercles anteri- orly attending shallow, ill-defined punctures; tu- bercles smaller, ill defined, less regular near su- ture, often appearing confused, becoming slightly stronger, more elongate laterally, sometimes forming interupted carinules; punctures disap- pearing near epipleuron and on declivity; alter- nate rows of tubercles larger, especially laterally; sometimes supertending short, declined setae, es- pecially laterally; rows of smaller tubercles be- coming obsolete laterally and posteriorly, disap- pearing near epipleuron and on declivity; epipleural carina narrow, weakly crenate, bearing row of slightly curved setae about one-third to one-half length of protarsus. Metasternum with coarse, shallow, setigerous punctures separated by about one puncture diameter; setae exceed- ingly fine, very short; metepisternum with few ob- scure punctures; abdominal sternites one to two sculpted like metasternum, but more finely, sparsely so; punctures sparser medially and on last three sternites mostly on posterior margins. Femora polished, with scattered minute setae. Protibia very broadly triangular (Fig. 40), with row of very short, blunt spines along lateral mar- gin, becoming subcontiguous around angle; me- sial margin with row of about five curved setae about two-thirds length of protarsus; posterior surface with few scattered tubercles, short spinose setae; mesial tibial spur longer, stouter than lat- eral; strongly curved. Meso- and metatibia with row of three to four coarse, spines on ectal mar- gins; posterior surfaces sparsely, irregularly set with inclined setae about one-half to two-thirds length protarsus. Ma e.—Differs as stated in tribal description. Aedeagus as in Figure 37. MEASUREMENTS.—EL 1.6-2.2 mm, EW 1.1-1.6 mm, PL 0.6- 0.9mm, PW 0.9-1.3 mm. Hototyre.—Female (CAS) and six paratypes from Califor- nia, San Bernardino County, 9 air mi S Baker, VII-1-1978, J Doyen, P. Rude (J. Doyen Lot # 78F5). ADDITIONAL ParaTyPEsS.—Same locality IV-27-1977, J. Doyen (J. Doyen Lot # 77D2) (2). Riverside County: Sand dunes 1 mi W Blythe, V-23/24-1970, pit trap, J. Johnson, M Wasbauer (19). Utah. Washington County, 2 mi E Washington, V-20/VI-8-1980, R. Hardy (3). ADDITIONAL MATERIAL EXAMINED (Fig. 42).—Arizona. Mo- have County: 3miN, 7 miE Littlefield, Virgin River, II-28/X- 1-1982 (2). Mexico. Sonora: Cholla Bay, 6 mi N Puerto Penasco, III-18/19-1980 (1); El Desemboque, III-22-1980 (6); 1 mi W Ba- hia de San Carlos, II-23-1980 (7). DiaGnosis.—Batuliodes spatulatus is very simi- lar to B. obesus Doyen, differing as described in the discussion of the latter. It is also similar to B. wasbaueri Doyen, but B. wasbaueri entirely lacks the fringe of pronotal setae present in B. spatula- tus and has the antennae as long as the head width (about three-fourths head width in B. spatulatus). Batuliodes spatulatus is geographically varia- ble. Specimens from California and Utah have the elytral disk very sparsely setose or subglabrous; if setae are present, they are only about as long as the basal protarsal segment. The pronotal and epipleural setae are about one-third the length of the protarsus. Specimens from Sonora have the elytra setose and the pronotal and epipleural se- tae are about one-half the length of the protarsus. Batuliodes obesus new species Pale tan, strongly convex, obese beetles with setose elytra and with pronotal carina obliterated. FemaLe.—Epistomum with lateral lobes promi- nent, extending well beyond truncate middle; sparsely, evenly set with round tubercles slightly larger than eye facets, occasionally becoming ca- rinulate posteriorly on vertex; postgena and men- tum nearly smooth with few, obscure punctures. Antenna about three-fourths as long as head width; apical segment about as long as broad. Pronotum medially with shallow punctures about twice eye facet diameter, separated by about one puncture diameter; becoming denser, 370 smaller, and less well defined laterally and at- tended ectally by slightly elongate tubercles in lat- eral thirds of disk; few short, declined setae near margins of disk; lateral carina absent; posterior angles rounded, strongly obtuse, scarcely indi- cated; anterior angles rounded. Hypomeron pol- ished, glabrous except for few punctures on coxal cowling and somewhat irregular band of postero- dorsally curved setae about as long as protarsus near dorsal margin and on lateral sixths of ante- rior margin; prosternum and prosternal process glabrous except for few long, curved setae. Elytra seriately tuberculate; rounded tubercles anteriorly attending shallow, ill-defined punc- tures near suture, these becoming smaller later- ally and posteriorly and virtually disappearing near epipleuron and on declivity; alternate rows of tubercles larger, setigerous; setae about one- half length of protarsus, inclined, slightly curved; rows of smaller tubercles becoming obsolete lat- erally and posteriorly and disappearing near epi- pleuron and on declivity; epipleural carina scarcely elevated, indicated by row of small, round setigerous tubercles anteriorly, becoming carinate in anterior and posterior one-fifth but not prominent; tubercles separated by about one tu- bercle diameter; setae about as long as protarsus, slightly curved posterodorsad. Metasternum with coarse, very shallow, setigerous punctures, sepa- rated by one to several puncture diameters; setae about as long as basal metatarsal segment; met- episternum with few obscure punctures; abdomi- nal sternites sculpted like metasternum, except for setae being about one and one-half to two times longer; punctures sparser medially and on sternites three to five mostly near posterior bor- ders. Legs essentially as in spatulatus (Fig. 40). Mate.—Differs as stated in tribal description. Aedeagus as in Figure 38. MEasuREMENTS.—EL 1.4-1.8 mm, EW 1.0-1.4 mm, PL 0.5- 0.8mm, PW 0.9-1.2 mm HoLotypre.—Female (CAS) from California, Imperial County, 1 mi S Glamis, I[I-31-1978, J. Powell, in pitfall. Para- types: Imperial County: 2 mi N Glamis, I-27-1977, J. Doyen, on sand at night (1); Glamis, V-29-1971, pit trap, M. Wasbauer (1); 5.5 mi SE Glamis, VII-19-1978, A. Hardy, F. Andrews, pit trap (1); 7 mi SE Glamis, III-25/IV-8-1979, pit trap (1). Riverside County: 5 mi NW Indio, III-4-1972, F. Andrews, E. Kane, A. Hardy (1). Inyo County: Death Valley, Stovepipe Wells Sand Dunes, IX-14-1972, D. Giuliani DiaGnosis.—Batuliodes obesus is very similar to B. spatulatus Doyen. In B. obesus the lateral pronotal carina is absent and the hypomeral setae DOYEN: REVISION OF ANEPSIINI are about as long as the protarsus and curve strongly posterodorsad. In spatulatus the prono- tal carina is fine but complete, and the hypomeral setae are about half as long as the protarsus. Batuliodes obesus is restricted to aeolian sand dunes, with morphological adaptations for a psammophilous mode of life similar to that of Ba- tuliomorpha. All specimens have been collected during the winter months, save that from Death Valley, which is badly abraded, lacking setae, and may have been found dead. ACKNOWLEDGMENTS The following individuals and institutions pro- vided specimens for study: R. Aalbu (private col- lection); L. Herman, American Museum of Natu- ral History, New York; F. Hasbrouck, Arizona State University, Tempe; M. Campbell, Biosys- tematics Research Institute, Ottawa, Ontario, Canada; D. Kavanaugh, California Academy of Sciences (CAS), San Francisco; F. Andrews, Cal- ifornia Department of Food and Agriculture, Sac- ramento; E. Smith, Field Museum of Natural His- tory, Chicago, Illinois; J. Johnson, University of Idaho, Moscow; A. Newton, Museum of Com- parative Zoology (MCZ), Harvard University, Cambridge, Massachusetts; R. Snelling, Natural History Museum of Los Angeles County, Los Angeles, California; R. C. Bechtel, Nevada De- partment of Agriculture, Reno; C. Triplehorn, Ohio State University, Columbus; S. Frommer, University of California, Riverside; J. Chemsak, University of California, Berkeley; P. Ashlock, University of Kansas, Lawrence; R. Rust, Uni- versity of Nevada, Reno; T. Spilman, United States National Museum (USNM), Washington, D.C. Details of results of a pitfall survey at the sand dunes at Owens Dry Lake, California were made available by F. Andrews and A. Hardy, California Department of Food and Agriculture. The illustrations of beetles and legs were pre- pared by C. M. Tibbets. R. Snelling, Museum of Natural History of Los Angeles County, iden- tified the ant associate of Anepsius minutus. Of special note is the material housed in the col- lection of the California Department of Food and Agriculture. This is the largest collection of sand dune Coleoptera from western North America, and contains more than one-half the known speci- mens of Anepsiini. About a quarter of the known specimens are housed in the Essig Museum of En- tomology, University of California, Berkeley. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44, No. 15 371 LITERATURE CITED Anprews, F. G., A. R. Harpy, AND D. GIULIANI. 1979. The co- leopterous fauna of selected California sand dunes. Calif. Dep. Food and Agric., Sacramento, California. 142 pp. ArnETT, R. W., Jr. 1960. The beetles of the United States (A manual for identification). Catholic University Press, Wash- ington, D.C. 1112 pp. BiackweLper, R. E. 1945. Checklist of the Coleopterous in- sects of Mexico, Central America, the West Indies, and South America, Part 3. Bull. U.S. Nat. Mus. 185:343-550. BLAISDELL, F. E., Sr. 1923. Expedition of the California Acad- emy of Sciences to the Gulf of California in 1921. The Tene- brionidae. Proc. Calif. Acad. Sci. 4th Ser. , 12:201-288. . 1943. Contributions toward a knowledge of the insect fauna of Lower California, No. 7, Coleoptera: Tenebrioni- dae. 4th Ser., Proc. Calif. Acad. Sci. 24:171-287. Casey, T. L. 1907. A revision of the American components of the tenebrionid subfamily Tentyriinae. Proc. Wash. Acad. Sci. 9:275-522. Doyen, J. T. 1966. The skeletal anatomy of Tenebrio molitor (Coleoptera: Tenebrionidae). Misc. Publ. Entomol. Soc. Am. 5:103-150. . 1984. Systematics of Eusattus and Conisattus (Coleop- tera: Tenebrionidae; Coniontini; Eusatti). Occas. Pap. Calif. Acad. Sci. 141:1-104. Doyen, J. T. anDJ. F. Lawrence. 1979. Relationships and clas- sification of some Tenebrionidae and Zopheridae (Coleop- tera). Syst. Entomol. 4:333-377. GesieN, H. 1937. Katalog der Tenebrioniden—I. Teil. Pubbl. Mus. Pietro Rossi, Duino, 2:1-381. Horn, G. H. 1870. Revision of the Tenebrionidae of America north of Mexico. Trans. Am. Philos. Soc. 14:253-404. Kocu, C. 1955. Monograph of the Tenebrionidae of Southern Africa. Vol. I (Tentyriinae, Molurini.—Trachynotina: So- maticus Hope) Transvaal Museum, Pretoria. . 1961. Some aspects of abundant life in the vegetation- less sand of the Namib Desert Dunes. J. S.W. Afr. Sci. Soc. 15:8-92. . 1962. The Tenebrionidae of southern Africa XXXII. New psammophilous species from the Namib Desert. Ann. Transvaal Mus. 24:107-159. LacorpalirE, T. 1859. Histoire Naturelle des Insectes. Genera des Coléoptéres . . . Tome 5. Roret, Paris. LeConte, J. L. 1851. Descriptions of new species of Coleop- tera, from California. Ann. Lyc. Nat. Hist. New York 5:125- 184. . 1862. Classification of the Coleoptera of North America prepared for the Smithsonian Institution. Smithson. Misc. Coll. 136:209-286. LeConte, J. L. anpG. H. Horn, 1883. Classification of the Co- leoptera of North America. Smithson. Misc. Coll. 507: XXX-Vili + 567 pp. Lena, C. W. 1920. Catalogue of the Coleoptera of America, North of Mexico. John D. Sherman, Mount Vernon, New York. Matson, J. O. 1976. The distribution of rodents in Owens Lake region, Inyo County, California. Contnb. Sci. Nat. Hist. Mus. Los Angeles County, 276:1-27. Papp, C. S. 1961. Checklist of the Tenebrionidae of America, north of the Panama Canal (Notes on North American Cole- optera, No. 14). Opusc. Entomol. 26:97-140. APPENDIX | Characters and Character States Plesiomorphic character states are listed first, apomorphic states last. Some characters, such as antennal shape (1, 2), posterior pro- notal angles (11, 12) and hypomeron sculpture (13, 14) are listed twice because more than 2 character states are recognized. A double listing allows such characters to be coded in an additive binary fashion. a . Antennal shape: a) clavate (or four-segmented club); b) three-segmented club. 2. Antennal shape: a) clavate (or three-segmented club); b) four-segmented club. 3. Antennal length: a) at least as wide as head; b) three-fourths as wide as head or less. 4. Antennal length: a) at least three-fourths as wide as head; one-half as wide as head or less. 5. Appical antennal segment: a) apical segment much longer than wide; b) apical segment subquadrate or nearly round. 6. Tentorium: a) open posteriorly; b) closed posteriorly be- tween bridge and occiput (Fig. 2). . Epistomal margin: a) shallowly emarginate or truncate; b) lateral lobes prominent; middle recessed, + straight (Fig. 33). . Submental gland: a) present; b) absent. 9. Pronotal margin: a) glabrous or with short, appressed setae; b) fimbriate. Pronotal lateral carina: a) distinct, carinate; b) absent. ~ wo 10. i=) 11. Posterior pronotal angles: a) obtuse (or rounded, absent); b) exserted, about 90° or acute. 12. Posterior pronotal angles: a) angulate, distinct; b) absent. 13. Hypomeron sculpture: a) scabrous, rugulose, or smooth; b) coarsely punctate. 14. Hypomeron sculpture: a) scabrous, rugulose (or punctate); b) smooth polished. 15. Elytral margin: a) glabrous or with short, appressed setae; b) fimbriate. 16. Elytral sculpture: a) carinae interrupted by punctures; b) tu- berculate, punctate. 17, Ventral setation: a) subglabrous or short, sparse setae; b) long semi-erect setae. 18. Mesocoxal closure (Fig. 4-6): a) narrow space between lobes of meso- and metasternites; b) sternite lobes touching. 19. Sternite lobes (Fig. 4-6): a) mesosternal and metasternal lobe subequal in width laterad of coxal cavity; b) metaster- nal lobe much broader than mesosternal. 20. Mesotrochantin (Fig. 4-6): a) exposed; b) concealed. 372 . Metasternum length: a) about twice mesocoxal length; b) about mesocoxal length (occasionally 1.5 times mesocoxal length). . Protibial shape: a) narrowly triangular (Fig. 18); b) broadly triangular (Fig. 17). 23. Protibial spurs: a) subequal, straight; b) mesial spur much larger than lateral, curved posteriad (Fig. 17). 24. Protibial setation: a) posterior margin glabrous or with short, appressed setae; b) posterior margin with three to seven long, erect or semi-erect setae (Fig. 17). DOYEN: REVISION OF ANEPSIINI . Aedeagus: a) Tegmen entire (Fig. 14); b) tegmen with mem- branous or submembranous central region on ventral sur- face (Fig. 13). . Body proportions: a) slender (EW/EL < 0.65); b) obese (EW/EL = 0.72). . Setal length: a) pronotal and elytral fimbriae short (or ab- sent); b) setae long, projecting. INDEX TO VOLUME 44 Additions to Index, Proceedings vol. 44 (nos. 13-15, pp. 283-372) New names Anepsius minutus Batuliomorpha comata Batuliodes obesus Batuliomorpha imperialis Batuliodes spatulatus Batuliomorpha tibiodentata Batuliodes wasbaueri Rhinobatos punctifer New names in boldface type Abies 303, 305 Aneflus 325-326 Abutilon 289, 298, 319, 328 prolixus insoletus 301 Acacia 291, 295-297, 299-308, 311-312, 314-316, 318 protensus protensus 301 berlandieri 301 sonoranus 301 farnesiana 289, 323, 334 Anelaphus 324-325 Acanthocinini 289 debilis 303, 323 Acanthoderes 316 inermis 303, 324 Acer 294, 302-303, 312, 316-317, 320 moestus moestus 303 Achryson 327 niveivestitus 303 surinamum 295, 323 spurcus 292, 303 Adetus 327 truncatus 303 brousi 311 Anepsiinae 346 Aegomorphus 316 Anepsiini 343-350 quadrigibbus 316 Anepsius 343-344, 346-351, 353-356 quadrigibbus form lucidus 316 angulatus 366 Aesculus 295 atratus 351-352 Aetobatus bicolor 351-352 narinari 341 brunneus 351-352 Aetomylaeus catenulosus 351-352 milvus 341 confluens 343, 366 Aetoplatea deficiens 351-352 tentaculata 341 delicatulus 343-346, 348-354, 356 Agallisini 308 minutus 343-344, 348, 351-354, 356, 370 Agallisus montanus 344, 348-349, 351, 353-357 lepturoides 308 nebulosus 351-352 Agave 309, 319 valens 344, 348-351, 353-357 Ageratum 315 Anoplium Alaudes 345 truncatum 303 Albizzia Anoxypristis Julibrissin 314 cuspidata 341 Alnus 305, 310 Antecrurisa Amaranthus 315 apicalis 316 Ambrosia 311-312, 315, 321, 325 Apocynum 312 Amelanchier 310 Aporataxia 315 Amphionycha 322 Archodontes 327 Ancylocera 325 melanopus serrulatus 293, 323 bicolor 308, 323, 325 Aromia macrotela 325 moschata 284 Ancylocerini 325 Arundo Aneflomorpha 324-326 donax 291 opacicornis 301 Asclepias 321-322 seminuda 301 spp. 321 tenuis 301 Asidinae 344 [381] 382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Asimina 312 Aspidosperma 295 Aster 301-321, 325 spinosus 316, 321 Asteraceae 289 Astyleiopus 326 variegatus 318 Astylidius 327 leiopinus 317 parvus 317 Ataxia 327 crypta 312, 323 hubbardi 311-312 spinicauda 323 tibialis 309, 312 Atriplex 344 confertifolia 344 Axestinus obscurus 301 Baccharis 291, 297, 299, 301-304, 306-308, 315-316, 318 Baileya 320 Batuliinae 345 Batuliini 344-345, 349 Batulinae 349 Batulini 349 Batulinus 357 setosus 357 Batuliodes 343, 345-350, 363-369 confluens 344, 347-348, 364-368 obesus 343-344, 347-348, 364-365, 368-370 rotundicollis 344-345, 347-348, 350-351, 363- 367 spatulatus 343-344, 347-348, 351, 364-370 wasbaueri 343, 347-348, 364-365, 367-369 Batuliomorpha 343-344, 346, 348-350, 359-362, 368, 370 comata 343-344, 346, 348-350, 359-362 imperialis 343, 348, 359-362 tibiodentata 343, 349, 359-363 tibiodentatus 348 Batulius 343-345, 348-350, 357, 363 rotundicollis 364 setosus 344, 347-348, 357-359 Batyle suturalis cylindrella 298 Bernardia 322 myricaefolia 322 Betula 295-296, 310, 316 Bidens 315 Boromorphus 345 Brya 295 Bumelia 305-306, 309 Bursera 294 Cacostola 327 salicicola 315, 323 Callancyla 325 Callidium 326 texanum 322-323 Callipogonius cornutus 313, 323 Callona 293 rimosa 297 Cambaia 326 Carpinus 310, 312 Carya 293-294, 296, 298, 300, 302, 303, 305-306, 308, 312, 316, 320 Cascostola lineata 315 Castanea 294, 298, 302-303, 305, 310, 312, 316, 318 Cathetopteron 326, 328 amoena 322-323 Ceanothus 308 Celastrus 312, 318 Celtis 290-292, 294-300, 302-305, 307-308, 310-320, 822) laevigata 323 laevigata var. texana 334 pallida 323, 334 Ceralocyna 325 Cerambycidae 283-285, 287-290, 293, 323-326, 328 Cerambycinae 294, 328 Ceratonia 295 Cercidium 294-295, 302 Cercis 305, 308, 312, 316 Cercoptera 325 Chamaerops 302 Championa 325 Chlorophora 295, 310 Chrysothamnus 325 Cirsium 311 Cissus 290, 297, 301, 305, 308-309 Citrus 283, 290, 293-298, 302-303, 305-308, 311, 314— 3'1'5;/323 paradisi 323 sinensis 323 spp. 328 Caldrastis 303 Clematis 290, 297, 301, 306, 308-309, 311, 322 Cnemoplatiini 345 Coleoptera 283, 343, 347 Compositae 289, 293, 299, 316, 319, 320 Compsa alacris 304 quadriplagiata 304 textilis var. alacris 304 Condalia 291, 297, 299, 301, 305-307, 311, 315 Hookeri 334 Coniontini 345 Conium 306 Corallancyla 325 Cordia 300, 312 boisseri 334 Coreopsis 315 INDEX Cornus 308, 312, 320 Corylus 295, 305 Cotinis mutabilis 297 Crossidius 289, 325 humeralis 298 humeralis quadrivittatus 298-299 pulchellus 299 suturalis 298 suturalis melanipennis 298-299 Cryptochilini 345 Cryptoglossini 345 Cucumis 311 Cucurbita 311 Cylindrataxia 315 Cyphonotida 327 laevicollis 327 laevicollis laevicollis 309 Cyrtinus 283, 327 pygmaeus 320, 326 Dasyatididae 341 Datura 311 Dectes 325 texanus 319 texanus aridus 319 Deltaspis 324 Dendrobias 300, 306, 327 mandibularis ssp. 323 mandibularis virens 323 Derobrachus 283 geminatus 294 Desmiphora 327 aegrota 311-312 hirticollis 312-313, 324 Desmodium 315 Diaperini 347 Dihammophora 327 dispar 307 Diospyros 292, 305, 308 texana 334 Dorcaschema 326 alternatum 311 alternatus octovittata 311 wildii 311 Dorcasta 327 cinerea 311 Eburia 327 haldemani 296 mutica 296, 323 mutica var. manca 296 ovicollis 296, 323 stigmatica 296, 323 tumida 296 Ecyrus arcuatus 313-314 383 arcuatus texanus 314 dasycerus 314 fasciatus 313 penicillatus 313 texanus 323 Ehretia 292 anacua 334 Elaphidini 325 Elaphidion 325, 327 irroratum 303 linsleyi 302, 323 linsleyi x mimeticum 302 mimeticum 302, 323 mucronatum 302 Elaphidionini 325 Elaphidionoides 325 aspersus 303 incertus 302-303 incertus-aspersus complex 303 linsleyi 303 spp. 326 villosus 303, 323 Elaphidionopsis 325 fasciatipennis 304 Elytroleptus 284, 299, 327 divisus 299 Enaphalodes 325 atomarius 302, 326 cortiphagus 302 hispicornis 302 rufulus 302, 326 taeniatus 302, 323 Enustromula validum 302 Epectasis hiekei 315 Erechtites 315 Ergates 326 spiculatus neomexicanus 322 Erigeron 311, 315 Eriogonum 306 Euderces reichei exilis 292, 307, 323 Eupogonius 327 fulvovestitus 313 pauper 312, 326 vestitus 313 Eurychorini 345 Eustromula 326 Evander 299 Eysenhardtia 297, 308 texana 334 Fagus 293-294, 310, 316 Ficus 295, 297, 305, 310, 316-317, 319 Franseria 344 dumosa 344 Fraxinus 291, 296, 300, 305-306, 312-313 384 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 berlandieriana 334 pennsylvanica var. subintegerrima 334 Gaillardia 321 Genista 306 Geropa 327 concolor 295, 323 Gleditsia 303, 306, 312, 315 Gliditala 315 Glycine 315 Gnaphalodes 327 trachyderoides 297, 323 Goes fisheri 310 pulverulentus 310 tesselatus 310 Gossypium 311-312 Gracilia minuta 295, 323, 326, 328 Graphisurus 327 triangulifer 316-317 Gutierrezia 325 spp. 299 Gymnopsyra 325 Gymnosperma 299, 325 glutinosa 299 Gymnura poecilura 341 Gymnuridae 341 Hamamelis 312 Haplopappus 289, 291, 297-298, 308, 325, 328 drummondi 298 Helenium 320 Helianthus 289, 296, 298-299, 301, 308, 311, 315, 319, 321, 325 Hemierana 325 marginata 322 suturalis 322 Hemilophini 326 Heterachthes 327 ebenus 304, 324 nobilis 304, 323 Heterotheca 320-321, 325 sp. 320 Himantura gerrardi 341 imbricata 341 uarnak 341 Hippopsis 327 lemniscata 315 Hostelezkya 316 Hylotrupes bajulus 305, 328 Hypexilis 327 pallida 296, 323 Hypolophus sephen 341 Ibidion pubescens 304 Ichneumonidae 324 Ichyomethia 303 Idisia 345 Inga 295 Inocarpus 310 Ischnocnemis 297 Isocoma 298 Jatropha 297 Juglans 296, 298, 302-303, 306, 310, 312, 318 Juniperus 322 Karwinskia 291, 297, 299, 301 humboldtiana 334 Knulliana 327 cincta cincta 298, 323 Koeberlinia 306 Lachnogyini 345 Lagocheirus procerus 317 texensis 317 undatus 317 Lallancyca 325 Lamiinae 283, 309, 328 Larrea 344 Leguminoseae 323 Leiobatus 336, 339 (Rhinobatos) 336 Leiopus alpha 318 Leptostylopsis 325, 327 luteus 318 Leptostylus 325, 327 biustus 318 gibulossus vogti 317 transversus asperatus 317 transyersus dietrichi 317 transversus spp. 317 Leptura 326, 328 (Stenura) emarginata 326 (Stenura) gigas 309, 326 (Stenura) splendens 326 emarginata 326 gigas 323, 326 splendens 326 Lepturges 327 angulatus canus 319, 323 infilatus 319, 323 minutus 320 sp. near confluens 319 subglaber 320 symmetricus 319 vogti 319 yucca 320 Lepturina 308 INDEX Lepturinae 328 Leucaena 284, 290-291, 295-301, 303-306, 308, 314— 316, 319-320 flavocinctus 300 flavocinctus puncticollis 300 pulverulenta 292, 323, 328, 334 Leucophyllum 292, 334 Ligustrum 296 Liopus houstoni 318 Liquidambar 294 Liriodendron 293, 296, 320 Lissonotus 327 Lochmaeocles cornuticeps cornuticeps 292, 314, 323 Lophalia 289, 327 cyanicollis 297 Lupinus 308 Maclura 305-306, 311 Magnolia 300 Malacopterus tenellus 295 Malus 303 Malvaviscus 312 arboreus var. drummondi 312 Mannophorus 327 laetus 296, 299 Manta birostris 341 ehrenbergi 341 Mecas 321, 325, 328 (Dylobolus) rotundicollis 320 (Mecas) cana saturnina 321 (Mecas) cineracea 320 (Mecas) confusa 320 (Mecas) linsleyi 321 (Mecas) marginella 320 (Mecas) pergrata 321 cana saturnina 321 cineracea 321 confusa 320-321 inornata 321 linsleyi 316, 321 pergrata 320-321 spp. 293 Megacyllene (Cyllene) erythropa 306 caryae 306, 323, 326 Megaderus 322, 327 bifasciatus 322 Meganeflus 325 Melochia 297 Melothria 290, 315 Methia 326 constricticollis 295, 323 Micraneflus 325 Micropsyrassa 325 Mimosa 295, 304, 307-308, 314 Mobula diabolus 341 kuhlii 341 Mobulidae 341 Monarda punctata 309 Moneilema 309, 326 armatum 309 armatum form rugosipenne 309 armatum punctatum 309 blapsides ulkei 309-310 mundelli 310 Morus 294, 305-306, 310-312, 318 rubra 334 Myliobatidae 341 Myliobatiformes 341 Myrmecocystus placodops 354 Nathriobrium 326, 328 methioides 305 Neaneflus 325 Nectandra 295 Necydaliella 326 Neoclytus 327 abbreviatus 307 acuminatus acuminatus 307 acuminatus hesperus 307, 323 augusti 307, 323 mucronatus vogti 306-307, 323 Neocompsa 304, 327 alacris 304 exclamationis 304, 323 Intricata 304 mexicana 304, 323 puncticollis orientalis 304 quadriplagiata 304 Neoptychodes 327 trilineatus 310 Nicotiana 311 trigonophylla 311 Obrium maculatum 305, 323 mozinnae 305, 323 rufulum 305, 326 Ochraethes 327 citrinus 306 Oncideres 311, 315, 318, 327 cingulata 315 cingulata texana 314-316, 323 cingulator 315 pustulata 310 pustulatus 290, 292, 307, 314, 318, 323 putator 314 texana 315, 318 Opuntia 291, 298, 304, 309-310 386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Ornithia 327 mexicana form zapotensis 305 mexicana mexicana 305 Osmopleura 308 Ostrya 310 Paraleptidea 326 Parandra 327-328 (Archandra) polita 293 Parandrinae 293, 328 Parevander 289, 299 Aurivillius 299 hovorei 299 xanthomelas 299 Parkinsonia 297, 302, 306, 314-315 aculeata 334 Parmenonta wickhami 311 Parmenosoma griseum 309 Parthenium 319 Pentanodes 327 albofasciata 327 dietzii 307, 327 Pepsis 309 Picea 305 Piezocera 327 monochroa 305 serraticollis 304-305, 323 Pinus 293, 304-305, 322 Pithecellobium 290-291, 295-300, 303-308, 313-317, 319 flexicaule 334 Placosternus 306 difficilis 292, 306, 323-324 erythropus 306 Platanus 294, 310 Plectrodera 327 scalator 310 Plinthocoelium schwarzi 305 suaveolens plicatum 305 Plionoma 326 suturalis 298 Populus 294-295, 305, 310, 326 spp. 293 Prioninae 293, 328 Prionus 327 (Antennalia) fissicornis 294 (Neopolyarthron) imbricornis 294 Pristidae 341 Pristiformes 341 Pristis pectinata 341 zijsron 341 Prosopis 291-298, 301-307, 314-316 glandulosa 289, 323, 334 Prunus 300, 303, 310, 312 Psapharochrus 316 Pseudostrangalia 326 cruentata 309 Psyrassa 324-325 brevicornis 300 castanea 300 pertenuis 300 sallaei 300 texana 300 Purpuricenini 289 Pygmaeopsis 326-328 viticola 313 Pyrus 294, 298, 312 Quercus 293, 295-296, 298, 302-303, 306-308, 310, 312, 316, 320, 326 Rajiformes 341 Rhamnus 295 Rhina ancylostoma 340 Rhinobatidae 335, 340 Rhinobatiformes 335, 340 Rhinobatos 335-336, 339-340 (Leiobatus) 336 (Rhinobatos) 335-336 albomaculatus 339-340 annandalei 336 formosensis 336, 340 granulatus 335 halavi 335-337, 340 holcorhynchus 336, 340 hynnicephalus 336, 340 irvinei 336, 339-340 lionotus 336 punctifer 335-336, 339-341 rhinobatos 336, 339 schlegelii 335-336, 340 thouin 335-337, 340 Rhopalophora 308, 327 angustata 308, 323 bicolorella 308 laevicollis 294, 308, 323 longipes 308, 326 longipes longipes 308 longipes meeskei 308 rugicollis 308 Rhus 303, 312 Rhynchobatidae 340 Rhynchobatus 340 djiddensis 340 Robinia 294-295, 318, 320 Rosa 295 Rubus 295 Rudbeckia 315 Rutaceae 323 INDEX Sabal texana 290, 334 Salicaceae 323 Salix 291, 294-296, 298, 302, 309-310, 312-315, 326 nigra 323, 334 Sambucus 308 Sapindus 298, 300, 305, 307, 317, 320 Saponaria var. drummondi 334 Sapium 312 Schnopsis 295 Senecio 297 Serjania 290, 297, 301, 308-309 Sesamum 315 Silphium 311 Smilax 311-312 Smodicum 295, 327 cucujiforme 294-295 texanum 294-295 Solanum 311 Solidago 301, 306, 319 Spalacopsis 327 texana 316 Sphaenothecus 298 Sphaeralcea 297 sp. 298 Sphaerion 327 exutum 301 Sphaerionini 325 Spondias 310 Stenaspis 297, 306 solitaria 297 verticalis arizonicus 297 verticalis insignis 297 Stenodontes 327 (Orthomallodon) dasytomus dasytomus 293-294 dasytomus dasytomus 323 Stenosini 345 Stenosis 345 Stenosphenus 325 dolosus 301, 323 lugens 300, 323 notatus 300 novatus 301 Sternidius 318 alpha 318 alpha misellus 318 crassulus 318 mimeticus 318, 323 naeviicornis 318 texanus 318, 323 wiltii 318, 323 Strangalepta abbreviata 323 vittata 323 Strangalia virilis 308 Styloxus 295, 326 fulleri 295 fulleri californicus 295 fulleri fulleri 295 texanus 295 Taeniura grabata 341 lymma 341 melanospilos 341 Tamarindus 295 Tamarix 306 Tanyochraethes tildeni 306 Taranomis 326 bivittata bivittata 296, 323 Taricanus truquil 323 Tenebrionidae 345-347 Tentyriinae 344-346, 349 Tetranodus 327 niveicollis 307 Tetraopes 321, 325, 327 discoideus 321 femoratus 321-322 texanus 321 thermophilus 321-322 Thryallis undatus 316-317, 323 Thurberia 312 Tilia 303, 312, 316 Torpedinidae 341 Torpediniformes 341 Torpedo panthera 341 sinuspersici 341 Toxylon 311 Trachyderes 300 (Dendrobias) mandibularis 299 mandibularis mandibularis 300 mandibularis reductus 299-300 mandibularis virens 299-300 Tragidion coquus 298, 326 Triboliides 344 Tylonotus 326 bimaculatus 296 Tylosis 289, 298, 327 Jiminezi 298 oculatus 298 Typhlusechus 345 Ulmaceae 323 Ulmus 290-292, 294-296, 299, 306-307, 310, 312- 313, 316, 318 crassifolia 323, 334 Ulomides 344 Ungnadia 302 388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 44 Uniungulum 347 Wisteria 303 Urgleptes 327 celtis 320, 323 Xanthium 311-312, 315, 321 knulli 320 Xanthocephalum 299 Urogymnus africanus 341 Yucca 286, 291-292, 309, 312, 317, 319-320 asperrimus 341 treculeana 319, 334 Valenus 326 Zagymnus 308 inornata 319 Zanthoxylum 292, 295, 300, 304-305, 307-308, 312, Verbesina 289, 296-297, 299, 308, 311-312, 322 S17. microptera 320 fagara 334 Vernonia 311, 315, 322 Zaplous 327 Viguiera 289, 296, 299, 328 Zizyphus 291 Vitis 294, 303, 306 obtusifolia 334 5 i ae i HECKMAN [= BINDERY INC. [= AUG 99 “to Pka®’ N. MANCHESTER, Bound -To-Pleas® Td aNA 46962 IM 3 9088 01302 7131 mV 2 w c < c 2 a z re} r =] E = @ 2 _ < z °o n x —E = ” are cole Serene oreo