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PROCEEDINGS 

of the 

Biological Society of 
Washington 



VOLUME 112 
1999 



Vol. 112(1) published 23 March 1999 Vol. 112(3) pubhshed 17 September 1999 

Vol. 112(2) published 15 June 1999 Vol. 112(4) pubhshed 29 December 1999 



WASHINGTON 
PRINTED FOR THE SOCIETY 



EDITOR 
C. Brian Robbins 



ASSOCIATE EDITORS 

Classical Languages Invertebrates 

Frederick M. Bayer Stephen L. Gardiner 

Frank D. Ferrari 
Rafael Lemaitre 

Plants Vertebrates 

David B. Lellinger Gary R. Graves 

Insects 

Wayne N. Mathis 



All correspondence should be addressed to the 

Biological Society of Washington, Smithsonian Institution 

Washington, D.C. 20560 



Allen Press Inc. 
Lawrence, Kansas 66044 



OFFICERS AND COUNCIL 

of the 

BIOLOGICAL SOCIETY OF WASHINGTON 

FOR 1998-1999 



OFFICERS 

President 
RICHARD P. VARI 

P resident-Elect 
BRIAN F. KENSLEY 

Secretary 
CAROLE C. BALDWIN 

Treasurer 
T. CHAD WALTER 



COUNCIL 

Elected Members 
MICHAEL D. CARLETON RAFAEL LEMAITEE 

W. DUANE HOPE ROY W. MCDIARMID 

SUSAN L. JEWETT JAMES N. NORRIS 



TABLE OF CONTENTS 

Volume 112 

Asakura, Akira. Icelopagurus tuberculosis, a distinct new hermit crab species (Crustacea: Decap- 

oda: Paguridae) from Japan 38 1-395 

Blazewicz, Magadalena, and Richard W. Heard. First record of the family Gynodiastyldae Stebbing, 
1912 (Crustacea: Malacostraca: Cumacea) from Antarctic waters with the description of Gyno- 
diastylis jazdzewskii, a new species 362-367 

Bolstad, Kathrin S., and Brian Kensley. Two new species of Hansenium (Crustacea: Isopoda: 

Asellota) from Madang, Papua New Guinea 1 64- 1 74 

Boyko, Christopher B. The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian Islands, 

with description of a new species 145-163 

Brown, W. C, A. C. Alcala, P. S. Ong, and A. C. Diesmos. A new species of Platymantis (Am- 
phibia: Ranidae) from the Sierra Madre Mountains, Luzon Island, Philippines 51 0-514 

Cai, Yixiong, Nguyen Xuan Quynh, and Peter K. L. Ng. Caridina clinata, a new species of 

freshwater shrimp (Crustacea: Decapoda: Atyidae) from northern Vietnam 53 1-535 

Campos, Ernesto. Inclusion of the austral species Pinnotheres politus (Smith, 1869) and P. garthi 
Fenucci, 1975 within the genus Calyptraeotheres Campos, 1990 (Crustacea: Brachyura: 
Pinnotheridae) 536-540 

Campos, Martha R. A new species of freshwater crab of the genus Strengeriana Pretzmann, 1971, 

from Colombia (Crustacea: Decapoda: Pseudothelphusidae) 405-409 

Campos, Martha R., and Rafael Lemaitre. Two new freshwater crabs of the genus Ptychophallus 
Smalley, 1964 (Crustacea: Decapoda: Brachyura: Pseudothelphusidae) from Panama, with notes 
on the distribution of the genus 553-56 1 

Carleton, Michael D., Robert D. Fisher, and Alfred L. Gardner. Identification and distribution of 

cotton rats, genus Sigmodon (Muridae: Sigmodontinae), of Nayarit, Mexico 81 3-856 

Chen, H.-L., and Peter K. L. Ng. Description of a new spider crab, Maja gracilipes, from the 
South China Sea, with notes on the taxonomic validity of M. brevispinosis Dai, 1981 (Crustacea: 
Decapoda: Brachyura: Majidae) 754-758 

Erdmann, Mark V., and Raymond B. Manning. A new species of Siamosquilla from Indonesia 

(Crustacea: Stomatopoda: Protosquillidae) 94-96 

Erseus, Christer. Parvidrilus strayeri, a new genus and species, an enigmatic interstitial clitellate 

from underground waters in Alabama 327-337 

Espinasa, Luis. A new genus of the subfamily Cubacubaninae (Insecta: Zygentoma: Nicoletiidae) 

from a Mexican cave 52-58 

Espinasa, Luis. Two new species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from 

Mexican caves, with redescription of the genus 59-69 

Fautin, Daphne G., and Brian R. Barber. Maractis remicarivora, a new genus and species of sea 
anemone (Cnidaria: Anthozoa: Actiniaria: Actinostolidae) from an Atlantic hydrothermal 
vent 624-63 1 

Eraser, Thomas H. A new species of cardinalfish (Perciformes: Apogonidae) from the Bay of 

Bengal, Indian Ocean 40-44 

Gillet, Patrick. A new species of Orbiniella (Orbiniidae: Polychaeta) from Marion Island, Indian 

Ocean 593-597 

Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 7. Probable parentage of 

Calliphlox iridescens Gould, 1860 443-450 

Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 8. A provisional hypoth- 
esis for the hybrid origin of Zodalia glyceria (Gould, 1858) 49 1 -502 

Graves, Gary R. Taxonomic notes on hummingbirds (Aves: Trochilidae). 2. Popelairia letitiae 

(Bourcier & Mulsant, 1 852) is a valid species 804-8 1 2 

Gutierrez- Aguirre, M. A., and E. Suarez-Morales. The freshwater centropagid Osphranticum la- 
bronectum Forbes, 1882 (Crustacea: Copepoda: Calanoida) in Mexico with description of a new 
subspecies 687-694 

Harasewych, M. G., and Yuri I. Kantor. A revision of the Antarctic genus Chlanidota (Gastropoda: 

Neogastropoda: Buccinulidae) 253-303 

Hauswaldt, J. Susanne. and Katherine E. Pearson. Urticina mcpeaki, a new species of sea anemone 

(Anthozoa: Actiniaria: Actiniidae) from the North American Pacific coast 652-660 

Heyer, W. Ronald. A new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Lep- 

todactylidae) with comments on the zoogeography of the Brazilian campos rupestres 19-39 



vu 



Vlll 



Heyer, W. Ronald, and Anna M. Munoz. Validation of Eleutherodactylus crepitans Bokermann, 

1965, notes on the type locality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and de- 
scription of a new species of Eleutherodactylus from Mato Grosso, Brazil (Amphibia: Anura: 
Leptodactylidae) 1-1 8 

Humes, Arthur G. Collocherides brychius, a new species (Copepoda: Siphonostomatoida: Aster- 

ocheridae) from a deep-water hydrothermal site in the northeastern Pacific 181-188 

Jara, Carlos G., and Victor L. Palacios. Two new species of Aegla Leach (Crustacea: Decapoda: 

Anomura: Aeglidae) from southern Chile 106-1 19 

Jensen, Gregory C, and Rachel C. Johnson. Reinstatement and further description of Eual us subtilis 
Carvacho & Olson, and comparison with E. Imeatus Wicksten & Butler (Crustacea: Decapoda: 
Hippolytidae) 133-140 

Karanovic, Tomislav. A new stygobitic Calanoida (Crustacea: Copepoda) of the genus Stygodiap- 

tomus Petkovski, 198 1 from the Balkan Peninsula 682-686 

Kensley, Brian, and Marilyn Schotte. New records of isopods from the Indian River Lagoon, 

Florida (Crustacea: Peracarida) 695-7 1 3 

Koenemann, Stefan, and John R. Holsinger. Megagidiella azul, a new genus and species of cav- 
emicolous amphipod crustacean (Bogidiellidae) from Brazil, with remarks on its biogeographic 
and phylogenetic relationships 572-580 

Lawson, Dwight P. A new species of arboreal viper (Serpentes: Viperidae: Atheris) from Cameroon, 

Africa 793-803 

Lazo-Wasem, Eric A. A new species of Chevalia (Crustacea: Amphipoda: Corophiidae) from the 
Indian Ocean with remarks on Chevalia carpenteri and the C. aviculae superspecies 
complex 562—571 

Leon-Gonzalz, Jesus Angel de, Vivianne Solis-Weiss, and Victor Ochoa Rivera. Nereidids (Poly- 

chaeta) from the Caribbean Sea and adjacent Coral Islands of the southern Gulf of Mexico ... 667—681 

Lewis, Julian J. Caecidotea simulator, a new subterranean isopod from the Ozark Springfield Plain 

(Crustacea: Isopoda: Asellidae) 175-180 

Locke, J. M., and K. A. Coates. Redescriptions of Grania americana, G. bermudensis and descrip- 
tions of two new species of Grania (Annelida: Clitellata: Enchytraeidae) from Bermuda 598-623 

Long, Douglas J., and John E. McCosker. A new species of deep-water skate, Rajella eisenhardti, 

(Chondrichthyes: Rajidae) from the Galapagos Islands 45-5 1 

Malabarba, Luiz R., and Stanley H. Weitzman. A new genus and species of South American fishes 
(Teleostei: Characidae: Cheirodontinae) with a derived caudal fin, including comments about 
inseminating cheirodontines 410-432 

Mayer, Gregory C, John A. W. ICirsch, James M. Hutcheon, Frangois- Joseph Lapointe, and 
Jacinthe Gingras. On the valid name of the lesser New Zealand short-tailed bat (Mammalia: 
Chiroptera) 470-490 

McCranie, James R., and Larry David Wilson. Two new species of the Eleutherodactylus rugulosus 

group (Amphibia: Anura: Leptodactylidae) from Honduras 5 15— 522 

Migotto, Alvaro E., and Antonio C. Marques. Hydroid and medusa stages of the new species 

Ectopleura obypa (Cnidaria: Hydrozoa: Tubulariidae) from Brazil 303— 3 12 

Miquelarei la, Amalia M., and Adriana E. Aquino. Taxonomic status and geographic distribution 

of Bryconamericus eigenmanni Evermann & Kendall, 1906 (Characiformes: Characidae) 523—530 

Mori, Atsushi. Caprella kuroshio, a new species (Crustacea: Amphipoda: Caprellidae), with a rede- 

scription of Caprella cicur Mayer, 1903, and an evaluation of the genus Metacaprella Mayer ... 722-738 

Murano, Masaaki. A new genus, Neodoxomysis (Crustacea: Mysidacea: Mysidae: Leptomysini), 

with description of two new species 352-36 1 

Ng, Peter K. L., and H.-L. Chen. On the identities of two Pacific species of deep-water porter 
crabs, Hypsophrys longirostris Chen, 1986, and Homologenus donghaiensis Chen, 1986 (Crus- 
tacea: Decapoda: Brachyura: Homolidae) 759-767 

Ng, Peter K. L., and S. H. Tan. The Hawaiian parthenopid crabs of the genera Garthambrus Ng, 

1966, and Dairoides Stebbing, 1920 (Crustacea: Decapoda: Brachyura) 120-132 

Nishe, EijLroh, Tomoyuki Miura, and Michel Bhaud. A new species of Spiochaetopterus (Chae- 

topteridae: Polychaeta) from a cold-seep site off Hatsushima in Sagami Bay, central Japan 210-215 

Okuno, Junji. Paleomonella hachijo, a new species of shrimp (Crustacea: Decapoda: Palaemonidae) 

from a >iubmarine cave in southern Japan 739-745 

Olson, Storrs L. A new species of pelican (Aves: Pelecanidae) from the Lower Pliocene of North 

Carolina and Rorida 503-509 

Peters, Daniel J., and Jean E. Pugh. On the entocytherid ostracods of the Brazos River basin and 

adjacent coastal region of Texas 338—35 1 



Rambla, Juan Pablo Blanco, and Rafael Lemaitre. Neocallicirus raymanningi, a new species of ghost 

shrimp from the northeastern coast of Venezuela (Crustacea: Decapoda: Callianassidae) 768-777 

Reid, Janet W., and Luis Moreno D. The western and southern distribution of Mesocyclops edax 

(S. A. Forbes) (Crustacea: Copepoda: Cyclopoida) 581-592 

Richart, Eric A., Jennifer A. Mercier, and Lawrence R. Heaney. Cytogeography of Philippine bats 

(Mammalia: Chiroptera) 453^69 

Rios, Ruben, and J. Emmett Dufy. Description of Synalpheus williamsi, a new species of sponge- 
dwelling shrimp (Crustacea: Decapoda: Alpheidae), with remarks on its first larval stage 541-552 

Robinson, Harold. Two new subtribes, Stokesiinae and Pacourininae, of the Vemonieae 

( Asteraceae) 21 6-2 1 9 

Robinson, Harold. Revisions in paleotropical Vemonieae (Asteraceae) 220-247 

Rojas, Yolanda, Fernando Alvarez, and Jose Luis Villalobos. A new species of crayfish of the 

genus Procambarus (Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico 396-404 

San Martin, Guillermo, and David Bone. Two new species of Dentatisyllis and Branchiosyllis 

(Polychaeta: Syllidae: Syllinae) from Venezuela 31 9-326 

Savage, Jay M., James R. McCranie, and Larry David Wilson. A new species of rainfrog of the 

Eleutherodactylus cruentus group from eastern Honduras (Amphibia: Anura: Leptodactylidae) . . 787-792 

Smith, David G., and Carole C. Baldwin. Psilotris amblyrhynchus, a new seven-spined goby (Te- 

leostei: Gobiidae) from Brazil, with notes on settlement-stage larvae 433^442 

Solis-Marin, Francisco A., and Alfredo Laguarda-Figueras. Cucumaria flamma, a new species of 

sea cucumber from the central eastern Pacific (Echinodermata: Holothuroidea) 778-786 

Stark, Bill P. Anacroneria from northeastern South America (Insecta: Plecoptera: Perlidae) 70-93 

Suarez-Morales, E., and R. Palomares-Garcia. Cymbasoma calif omiense, a new monstrilloid (Crus- 
tacea: Copepoda: Monstrilloida) from Baja California, Mexico 189-198 

Tavares, Marcos. Deilocerus captabilis, a new species of cyclodorippid crab from southeastern 

Brazil (Crustacea: Decapoda: Brachyura: Cyclodorippidae) 141-144 

Thoma, Roger F., and Raymond F. Jezerinac. The taxonomic status and zoogeography of Cambarus 

bartonii carinirostris Hay, 1914 (Crustacea: Decapoda: Cambaridae) 97-105 

Villalobos, Jose Luis, and Fernando Alvarez. A new species of Macrobrachium (Crustacea: De- 
capoda: Palaemonidae), with abbreviated development, from Veracruz, Mexico 746-753 

Wetzer, Regina, and Niel L. Bruce. A new genus and species of sphaeromatid isopod (Crustacea) 

from Atlantic Costa Rica 368-380 

White, Tracy R., April K. Wakefield Pagels, and Daphne G. Fautin. Abyssal sea anemones (Cni- 
daria: Actiniaria) of the northeast Pacific symbiotic with molluscs: Anthosactis nomados, a new 
species, and Monactis vestita (Gravier, 1918) 637-65 1 

Williams, Jason D., and Christopher B. Boyko. A new species of Pseudostegias Shino, 1933 

(Crustacea: Isopoda: Bopyridae: Atheliginae) parasitic on hermit crabs from Bali 714-72 1 

Winston, Judith E., and Stace E. Beaulieu. Striatodoma dorothea (Cheilostomatida: Tessaradomi- 

dae), a new genus and species of bryozoan from deep water off California 31 3-3 1 8 

Young, Richard E., and Michael Vecchione. Morphological observations on a hatchling and a 

paralarva of the vampire squid, Vampyroteuthis infemalis Chun (Mollusca: Cephalopoda) 661-666 

Zottoli, Robert. Early development of the deep-sea ampharetid (Polychaeta: Ampharetidae) De- 

cemunciger apalea Zottoli 199-209 



XI 



INDEX TO NEW TAXA 



Volume 112 



(New taxa are indicated in italics: new combinations designated n.c.) 

CNIDARIA 



Anthosactis nomados 

Ectopleura obypa 

Maractis 



rimicarivora 
Urticina mcpeaki - 



640 

304 

625 
625 

653 



BRYOZOA 



Striatodonia _ _ _ _. 




313 


dorothea ___ _ _ _- 




315 


Chlanidota (Pfefferia) invenusta 


MOLLUSCA 


289 


Rranchiosyllis Inrpnof 


ANNELIDA 
Polychaeta 


.... 320 


Dentatisyllis morrocoyensis _____ 




320 


Orbiniella marionensis _ _____ 




.... 593 


Spiochaetopterus sagamiensis ... 
Grania hylae ... __ __ . _ _. . . 




.. 211 


Oligochaeta 


605 


laxartus . . . _ 




602 


Parvidrilus 




328 


strayeri 




3^8 


Aegla cholchol 


ARTHROPODA 
Crustacea 


107 


hueicollensis _.._ .. 




112 


Albunea danai 




155 


Beathcesphaera . 




369 


ruthae 




._ 371 


Caecidotea simulator 




175 


Calyptraeotheres garthi n.c. 




537 


politus n.c. . . ... 




.. 537 


Caprella kuroshio 




722 


Caridina clinata 




531 



xu 



Chevalia hirsuta 562 

Collocherides brychius 1 82 

Cymbasoma califomiense 190 

Deilocerus captabilis 141 

Gynodiastylis jazdzewskii 362 

Hansenium thomasi 169 

tropex 165 

lais floridana 697 

Icelopagurus tuberculous 382 

Macrobrachium tuxtlaense 746 

Maja gracilipes 754 

Megagidiella 572 

azul 573 

Neocallichirus raymanningi 769 

Neodoxomysis 352 

elongata 353 

sahulensis 356 

Osphranticum labronectum mexicanum 688 

Palaemonella hachijo 739 

Procambarus (Austrocambarus) Citlaltepetl 398 

Pseudostegias macdermotti 715 

Ptychophallus kuna 555 

uncinatus 554 

Siamosquilla sexava 94 

Sphaeromyopsis sanctaluciae 707 

Strengeriana casallasi 405 

Stygodiaptomus ferus 682 

Synalpheus williamsi 542 

Insecta 

Ancroneuria achagua 70 

arawak 72 

chaima 73 

claudiae 74 

karina 77 

makushi 77 

paria 80 

perija 80 

timote 83 

wapishana 8 8 



Anelpistina cuaxilotla 64 

inappendicata 60 

Squamigera 53 

latebricola 56 

ECHINODERMATA 

Cucumaria flamma 779 



Xlll 



CHORDATA 
Pisces 

Acinocheirodon 413 

me lano gramma 413 

Apogon oxina 41 

Psilotris amblyrhynchus 434 

Rajella eisenhardti 45 

Amphibia 

Eleutherodactylus dundeei 12 

olanchano 518 

operosus 788 

pechorum 515 

Rupirana 26 

cardosoi 27 

Platy mantis sierramadrensis 510 

Reptilia 

Atheris broadleyi 794 

Aves 

Pelicanus schreiberi 503 

PLANT AE 

Acilepis aspera n. c . 226 

clivorum n.c. 226 

dalzelliana n.c. 226 

dendigulensis n.c. 226 

namtcianensis n.c. 226 

saligna n.c. 226 

scariosa n.c. 227 

silhetensis n.c. 227 

spirei n.c. 227 

Bechium nudicaule n.c. 227 

rhodolepis n.c. 228 

Brenandendron 244 

donianum n.c. 244 

frondosum n.c. 244 

titanophyllum n.c. 244 

Cabobanthus 228 

bullulatus n.c. 228 

polysphaerus n.c. 228 

Centrapalinae, new subtribe 223 

Centrapalus acrocephalus n.c. 236 

africanus n.c. 236 

chthonocephalus n.c. 236 

denudatus n.c. 236 

kirkii n.c. 236 

pauciflorus n.c. 236 

praemorsus n.c. 236 

purpureus n.c. 236 



XIV 



subaphyllus n.c. 236 

Cyanthillium albicans n.c. 229 

cony zoides n.c. 229 

hookerianum n.c. 229 

vemonioides n.c. 229 

Distephanus henryi n.c. 238 

Erlangeinae, new subtribe 222 

Gymnantheminae, new subtribe 224 

Gy nmanthemum anceps n.c. 239 

andersonii n.c. 239 

andrangovalense n.c. 240 

antanalus n.c. 240 

appendiculatum n.c. 240 

arboreum n.c. 240 

baronii n.c. 240 

bellinghamii n.c. 240. 

bockianum n.c. 240 

chapelieri n.c. 240 

corymbosum n.c. 241 

coursii n.c. 241 

crataegifolium n.c. 241 

cumingianum n.c. 241 

cylindriceps n.c. 241 

dissolutum n.c . 24 1 

esculentum n.c. 241 

exsertiflorum n.c. 24 1 

exsertum n.c. 241 

glaberrimum n.c. 241 

hildebrandtii n.c. 24 1 

humblotii n.c. 242 

lou velii n.c. 242 

mespilifolium n.c. 242 

myrianthum n.c. 242 

pectiniforme n.c. 242 

pectorale n.c. 242 

pleistanthum n.c. 242 

nieppellii n.c. 242 

secundifolium n.c. 243 

solanifolium n.c. 243 

subcrassulescens n.c. 243 

theophrastifolium n.c. 243 

thomsonianum n.c. 243 

urticifoilum n.c. 243 

vidalii n.c. 243 

volkameriifolium n.c. 243 

weghtianum n.c. 243 

zanzibarense n.c. 243 

zeylanicum n.c. 243 

Hilliardiella 229 

aristata n.c. 230 

calyculata n.c. 230 

hirsuta n.c. 230 

leopoldii n.c. 230 

nudicaulis n.c. 230 

oligocephala n.c. 230 

pinifolia n.c. 230 



XV 



smithiana n.c. 230 

Koyamasia 234 

calcarea n.c. 235 

Lampropappus new status 245 

Lampropappus eremanthifolia n.c. 245 

hoffmannii n.c. 245 

turbinellus n.c. 245 

Linzia gerberiformis n.c. 237 

infundibulariformis n.c. 237 

ituriensis n.c. 237 

melleri n.c. 238 

usafuensis n.c. 238 

Manyonia 224 

peculiaris n.c. 225 

Myanmaria 244 

calycina n.c. 245 

Oocephala agrianthoides n.c. 232 

stenocephala n.c. 232 

Orbivestus 230 

cinerascens n.c. 231 

homilanthus n.c. 231 

karaguensis n.c. 231 

undulatus n.c . 23 1 

Pacourininae, new subtribe 218 

Phyllocephalum microcephalum n.c. 235 

Polydora angustifolia n.c . 232 

bainesii n.c. 232 

chloropappa n.c. 233 

jelfiae n.c. 233 

poskeana n.c. 233 

serratuloides n.c. 233 

steetziana n.c . 23 3 

sylvicola n.c. 233 

Stokesiinae, new subtribe 216 

Vemoniastrum 233 

aemulans n.c. 233 

ambiguum n.c. 233 

latifolium n.c. 234 

musofense n.c. 234 

nestor n.c. 234 

ugandense n.c. 234 

uncinatum n.c. 234 

viatorum n.c. 234 






PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY 

OF 

WASHINGTON 




VOLUME 112 NUMBER 1 
23 MARCH 1999 



ISSN 0006-324X 






THE BIOLOGICAL SOCIETY OF WASHINGTON 

1998-1999 
Officers 

President: Richard P. Vari Secretary: Carole C. Baldwin 

President-elect: Brian F. Kensley Treasurer: T. Chad Walter 

Elected Council 

Michael D. Carleton Rafael Lemaitre 

W. Duane Hope Roy W. McDiarmid 

Susan L. Jewett James N. Norris 

Custodian of Publications: Storrs L. Olson 



PROCEEDINGS 

Editor: C. Brian Robbins 



Associate Editors 

Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner 
Plants: David B. Lellinger Frank D. Ferrari 

Insects: Wayne N. Mathis Rafael Lemaitre 

Vertebrates: Gary R. Graves 

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):1-18. 1999. 

Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on 

the types and type locality of Telatrema heterodactylum Miranda- 

Ribeiro, 1937, and description of a new species of Eleutherodactylus 

from Mato Grosso, Brazil (Amphibia: Anura: Leptodactylidae) 

W. Ronald Heyer and Anna M. Munoz 

(WRH) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian 

Institution, Washington, D.C. 20560-0109, U.S.A.; 

(AMM) Department of Biology, New Mexico State University, Las Cruces, New Mexico 88003, 

U.S.A.; Current address: 1306 Alvarado, Carlsbad, New Mexico 88220, U.S.A. 

Abstract. — Examination of the types of Eleutherodactylus crepitans Boker- 
mann, 1965, indicates that they represent a valid species, which shoilld be 
removed from the synonymy of Eleutherodactylus fenestratus (Steindachner 
1864). Eleutherodactylus crepitans is only known from the type material. Cer- 
tain features of the types of Telatrema heterodactylum Miranda-Ribeiro, 1937, 
are described and the type locality clarified. Analysis of the advertisement call 
and morphology of a series of specimens from Chapada dos Guimaraes indi- 
cates that the specimens represent a species distinct from E. fenestratus (Stein- 
dachner 1864), for which no name is available. We describe this new species 
as Eleutherodactylus dundeei. The frog fauna of Chapada dos Guimaraes con- 
tains four distinct historical units: cerrado, chaco, Amazonian hylaea, and en- 
demic. 



The advertisement call of what has been 
identified as Eleutherodactylus fenestratus 
from Chapada dos Guimaraes, Mato Gros- 
so, Brazil, was recently recorded by WRH 
as part of PRODEAGRO. PRODEAGRO is 
an ecological-agricultural survey of the 
State of Mato Grosso, undertaken by the 
State Secretariat of Planning, funded by the 
World Bank, and contracted to the Sao Pau- 
lo firm of consultants "Consorcio Nacional 
de Engenheiros Consultores" (CENEC). As 
part of the National Museum of Natural 
History's Research Training Program, 
AMM analyzed the advertisement call of 
the Chapada dos Guimaraes form and com- 
pared it to the advertisement call of Eleu- 
therodactylus fenestratus from the Amazon 
basin. The results indicate that the Chapada 
dos Guimaraes form is a different species 
from the Amazonian E. fenestratus. To de- 
termine whether there is an available name 
for the Chapada dos Guimaraes form and 



whether it has a distribution beyond the re- 
gion of the town of Chapada dos Guima- 
raes, we examined other specimens of 
Eleutherodactylus from Mato Grosso, in- 
cluding the type material of Telatrema het- 
erodactylum Miranda-Ribeiro, 1937 and 
Eleutherodactylus crepitans Bokermann, 
1965. The purpose of this paper is to report 
our findings. 

Methods and Materials 

The first recording of the Chapada dos 
Guimaraes form is from USNM Tape 320, 
Cut 2, Brazil, Mato Grosso, Chapada dos 
Guimaraes, Estancia Monarca, recorded 30 
September 1996, between 1835-1930 h, air 
temperature 21.6°C, unvouchered, by 
WRH, one call recorded and analyzed. The 
second recording of the Chapada dos Gui- 
maraes form is from USNM Tape 320, Cut 
3, same data as Cut 2, except recorded at 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



1930 h, voucher USNM 507899, one call 
recorded and analyzed. The recordings for 
Eleutherodactylus fenestratus are from 
USNM Tape 206, Cut 2, Peru, Madre de 
Dios, Tambopata, recorded 6 January 1989, 
1850 h, air temperature 24.0°C, voucher 
USNM 342993, by Reginald B. Cocroft III, 
four calls analyzed, and USNM Tape 266, 
Cut 19, Peru, Madre de Dios, Tambopata, 
recorded 15 November 1990, 1815 h, air 
temperature 27.0°C, unvouchered, by Re- 
ginald B. Cocroft III, six calls analyzed. 

Call recordings were digitized using 
Sound Image software at a sampling fre- 
quency of 22.05 KHz and 16-bit resolution. 
The digitized calls were converted into au- 
dio data files using a wave converter pack- 
age and analyzed as audiospectrograms and 
wave forms using Computerized Speech 
Research Environment software (AVAAZ 
Innovations Inc., Version 4.2). 

Eleven parameters were analyzed for 
each call following the definitions of Heyer 
et al. (1990): call duration, call rate, notes 
per call, note duration, note repetition, note 
pulsation, fundamental frequency, dominant 
frequency, change in dominant frequency, 
peak frequency, and characterization of har- 
monics. 

The audiospectrograms used in Fig. 1 
were produced using Canary software (Cor- 
nell Laboratory of Ornithology, Version 
1.2). 

Color pattern and external morphological 
data were recorded using a system of 
sketches to represent all distinctive patterns 
or states. These data were then summarized 
and categorized into written character 
states. Measurements were taken with dial 
calipers and recorded to the nearest 0. 1 mm 
following Heyer (1984). Adult males were 
determined by presence of vocal slits. Adult 
females were determined by presence of 
mature ova visible through the body wall. 
Unclear sex determinations based on exter- 
nal examination were verified by dissection. 
Multivariate discriminant function analyses 
were performed using SYSTAT 7.0 for 
Windows. 



Specimens examined are listed in Appen- 
dix 1. Museum abbreviations follow Levi- 
ton et al. (1985). 

Results 

Advertisement calls. — The ideal calls to 
compare the Chapada dos Guimaraes form 
with are those from topotypic Eleuthero- 
dactylus fenestratus males. Eleutherodac- 
tylus fenestratus (Steindachner 1864) was 
described from specimens from Rio Ma- 
more, Rondonia and Borba, Amazonas, 
Brazil. The geographically most proximate 
adequate recordings we know for Amazo- 
nian E. fenestratus are from Tambopata, 
Peru. We use these recordings to represent 
E. fenestratus for our comparisons. 

As there is some variation among the 
four individual frogs recorded from Peru 
and Brazil, the data are summarized by in- 
dividual caller (Table 1). 

The frog from Tambopata, USNM Tape 
206, Cut 2, has a mean call duration of 0.31 
sec; 2.75 mean notes per call; and a mean 
note duration of 0.07 sec. The dominant fre- 
quency is the second harmonic; additional 
harmonics are present, but weak. (Fig. la). 

The second frog from Tambopata, 
USNM Tape 266, Cut 19, has a mean call 
duration of 0.18 sec; and a mean note du- 
ration of 0.07 sec. The dominant frequency 
is the second harmonic; additional harmon- 
ics are present, but weak. 

The single call of the frog from Chapada 
dos Guimaraes, USNM Tape 320, Cut 2, 
has a relatively short (0.02 sec) and weak 
first note, the other 4 notes are stronger and 
have a duration range of 0.03-0.04 sec. The 
notes are pulsatile. The dominant frequency 
is the second harmonic; additional harmon- 
ics are present, but weak. 

The single call of the second frog from 
Chapada dos Guimaraes, USNM Tape 320, 
Cut 3, has a relatively short (0.02 sec) and 
weak first note, all other notes are stronger 
with a duration of 0.04-0.05 sec. The notes 
are pulsatile. The dominant frequency is the 



VOLUME 112, NUMBER 1 



10 n 



N 
U 

O 

a 
a 

»-i 
fa 




2 - 



10 n 



8 - 



6- 



4- 



2- 



B 



^ « ^ t 




o 



0.2 



0.4 0.6 

Time in Seconds 



0.8 



— I 
1.0 



Fig. 1. A. Advertisement call of Eleutherodactylus fenestratus, USNM Tape 206, Cut 2. B. Advertisement 
call of Chapada dos Guimaraes form, USNM Tape 320, Cut 3. 



second harmonic; additional harmonics are 
present, but weak. (Fig. lb). 

There is some variation between individ- 
ual male vocalizations from the same lo- 
cality. The variation observed between the 
recordings from Tambopata might be ac- 
counted for by differences of temperature 
of calling males, at least in part. As a gen- 
eral rule for frogs, increases in temperature 
result in decreased call and note duration 



and increased note repetition rate (Schnei- 
der et al. 1984; Duellman & Trueb 1986). 
The three degree difference of ambient tem- 
perature between the two recordings of the 
Peruvian specimens is in the predicted di- 
rection for call and note duration and note 
repetition rate. Note, however, that the re- 
cordings from Chapada dos Guimeraes 
were recorded at lower temperatures than 
those of Peru, yet these Brazilian calls had 



4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Table L — Advertisement call data for four Eleutherodactylus specimens from Tambopata, Peru and Chapada 
dos Guimaraes, Brazil. 



Geographic area 
USNM Tape and Cut 


Peru 
206, 2 


Peru 
266, 19 


Brazil 
320, 2 


Brazil 
320, 3 


Call duration (sec) 


0.21-0.36 


0.16-0.20 


0.29 


0.53 


Number of notes/call 


2-3 


2 


5 


8 


Note duration (sec) 


0.06-0.09 


0.05-0.08 


0.04 


0.05 


(Mean) note repetition rate per sec 


7.6 


9.1 


14.7 


13.7 


Fundamental frequency range 


1636-2411 


1464-2497 


1894-2239 


1808-2497 


Peak fundamental frequency 


1720 


2067 


2067 


2153 


Dominant frequency range 


2842^392 


2842^220 


3014-^220 


3273-4737 


Peak dominant frequency 


3531 


3617 


3692 


3875 



shorter note durations and higher note rep- 
etition rates than the Peruvian calls, indi- 
cating that temperature differences do not 
explain the differences between the Brazil- 
ian and Peruvian calls. 

The calls from Brazil and Peru do not 
differ in terms of carrier frequencies; they 
are broadcasting in the same frequency 
band. The calls differ markedly in terms of 
temporal packaging (Table 1), such as num- 
ber of notes per call, individual note dura- 
tion, and note repetition rate. Penna (1997) 
among others, has experimentally demon- 
strated that the kinds of temporal differenc- 
es found between the Tambopata and Cha- 
pada dos Guimaraes frogs are recognized as 
species differences by frogs. 

Unfortunately, to date, there are no re- 
cordings of what we consider to be Ama- 
zonian Eleutherodactylus fenestratus from 
the hylaea of northern Mato Grosso. In fact, 
the only other published recordings for E. 
fenestratus are from Amazonian Bolivia 
(Marquez et al. 1995) and are very similar 
to the recordings from Tambopata. Ronald 
I. Crombie made three recordings in Ron- 
donia, Brazil in which a single call of an E. 
fenestratus is in the background. In two of 
them (USNM Tape 55, Cut 5 and USNM 
Tape 56, Cut 5, both from Alto Paraiso), 
the call sounds as though it consists of two 
notes, but can not be confirmed on an au- 
diospectrogram due to the relative weak- 
ness of the call and strength of the back- 
ground noise. The call on USNM Tape 56, 
Cut 8 from Santa Cruz da Serra is a bit 



stronger and the audiospectrogram indicates 
it is comprised of two notes at a dominant 
broadcast frequency of 3300 Hz. The calls 
from Rondonia, as expected, match the 
calls from Tambopata rather than the calls 
from Chapada dos Guimaraes. 

The advertisement call evidence is con- 
sistent with recognizing the Chapada dos 
Guimaraes form as a species distinct from 
Eleutherodacytlus fenestratus. 

Morphology. — There are two available 
names for Eleutherodactylus from Mato 
Grosso that potentially could apply to the 
Chapada dos Guimaraes form. There are 
also a number of Eleutherodactylus fenes- 
tratus-\\k& specimens from Mato Grosso. 
The purpose of this section is to compare 
the Chapada dos Guimaraes form with 
these specimens to determine whether the 
Chapada dos Guimaraes form occurs more 
broadly within the State of Mato Grosso 
and to determine whether there is an avail- 
able name for it. 

The materials for comparison comprise 
the following (Fig. 2 for Mato Grosso lo- 
calities; Appendix 1 for specimen lists): the 
2 types of Telatrema heterodactylum Mi- 
randa-Ribeiro, 1937; the 3 type specimens 
of Eleutherodactylus crepitans Bokermann, 
1965; 11 specimens from Chapada dos Gui- 
maraes; 2 specimens from Fazenda Santa 
Edwiges; 1 specimen from Jacubim; 2 spec- 
imens from Barra do Tapirapes; and speci- 
mens considered to be E. fenestratus', 3 
specimens from Sao Jose do Rio Claro; 8 
specimens from Apiacas, 48 specimens 



VOLUME 112, NUMBER 1 




Fig. 2. Outline map of the State of Mato Grosso. See text for key to locality numbers. Open square is the 
city of Cuiaba. 



from Juruena (measurement data taken for 
10 females and 1 male), 1 specimen from 
Alto Juruena, 46 specimens from Aripuana 
(measurement data taken for 10 males and 
females), and 28 specimens from Tambo- 
pata and Pakitza, Madre de Dios, Peru (the 
specimens from the nearby locaHty of Pak- 
itza were added to those from Tambopata 
to bring the sample sizes up to at least 10 
males and 10 females). 

Patterns: The dorsal snout patterns dem- 
onstrate as much intra- as inter-population 
variation and are not discussed further. The 
other patterns analyzed demonstrate some 
levels of interpopulation variation (Tables 
2—8). In some cases, the sample sizes are 



adequate to conclude that the variation is 
meaningful (e.g., the differences in loreal 
region pattern between the Mato Grosso 
populations of Eleutherodactylus fenestra- 
tus (Table 3, samples 7-10) and the Peru- 
vian sample of E. fenestratus. In other cas- 
es, the differences are suggestive, but any 
conclusions are compromised by sample 
sizes. For example, there are two aspects of 
light mid-dorsal stripes that are impacted by 
small sample size. The first is that there is 
variation in the presence condition of the 
stripe. It may either be narrow or broad, and 
it may extend from the tip of the snout to 
the vent or from the interocular area to the 
vent. Because there were relatively few in- 



6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Table 2. — Interorbital markings. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost 
absent; B = Outlined only, not with expansion; C = Solid, not with expansion; D = Solid with single prominent 
posterior expansion; E = Solid with bifid prominent posterior expansion. Data are number of individuals dem- 
onstrating the various states (question mark indicates specimen faded, state uncertain). 



C. dos 
Taxa E. heterodactylns E. crepitans Gui maraes 

Locality 1 2 3 



E. fenestratus 



7 


8 


9 


9a 


10 


Peru 












5 




2 


12 




9 


2 


3 


6 


9 


1 


11 


9 
6 
6 



A 
B 
C 
D 
E 



11 



From illustration 2a (Miranda-Ribeiro 1937). 



dividuals with light mid-dorsal stripes, the 
information had to be collapsed for com- 
parison. Second, the sample size for the 
Chapada dos Guimaraes form is too small 
to conclude that it lacks light mid-dorsal 
stripes, even though all 11 specimens ex- 
amined lack them. There are three adequate 
samples of E. fenestratus to compare with 
the Chapada dos Guimaraes form: Aripua- 
na, Juruena, and Peru. For Aripuana, the 
probability of an individual having a mid- 
dorsal stripe is 0.35, thus the probability of 
not having a stripe is 0.65. If we use that 
probability to ask whether the Chapada dos 
Guimaraes population could have the same 
occurrence of stripes, the answer is no: 
0.65" = P < 0.001 (exact probability test). 
However, when the population data from 
Juruena and Peru are used, the answer is 
yes: for Juruena, the probability of not hav- 
ing a stripe is 0.91, for Peru 0.92, with P 
= 0.35 and 0.40 respectively. Thus, based 
on the low frequency of occurrence of mid- 
dorsal stripes in the Juruena and Peru sam- 



ples, if the Chapada dos Guimaraes form 
has mid-dorsal stripes at those same fre- 
quencies of occurrence, then the sample 
size is not adequate to have found individ- 
uals with that condition. 

Morphological features: All but three in- 
dividuals scored for tarsal fold variation 
had discernible tarsal folds (Table 9). It is 
likely that the three individuals thus scored 
actually have them but are preserved in 
such a manner that they are not discernible. 
The two individuals of Eleutherodactylus 
crepitans with discernible tarsal folds have 
a very different morphology from all other 
specimens examined (Table 9). The varia- 
tion in toe fringe condition is continuous 
(Table 10). The fringed condition is weakly 
fringed, with noticeable basal toe webbing 
(the fringe is not as well-developed as in 
some species of Leptodactylus, for exam- 
ple). The variation in tubercles on the sole 
of the foot (Table 11) suggests that E. cre- 
pitans is distinctive (in contrast to com- 
pletely distinct) from the other samples, and 



Table 3. — Loreal region patterns. Locality numbers refer to those in Fig. 2 (Locality 9a not mappable). A = 
No pattern; B = Incomplete stripe in front of eye; C = Dark stripe from eye to nostril, entire loreal region may 
be dark. Data are number of individuals exhibiting various states. Pattern not discernible for specimen from 
locality 6. 



Taxa 


E. 


heterodacnlus 
1 


E. 


crepitans 

2 


c 

Gui 


. dos 
maraes 
3 


1 

4 


9 

5 


9 

6 






£■. 


fenestratus 






Locality 


7 


8 


9 


9a 


10 


Peru 


A 






























8 


B 




1 




2 




4 




1 








9 


1 


1 


13 


C 








1 




7 


2 






3 


8 


12 




20 


7 



VOLUME 112, NUMBER 1 7 

Table 4. — Supratympanic fold patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = 
Fold not accentuated with dark brown band; B = Fold accentuated with interrupted dark brown band; C = Fold 
accentuated with solid dark brown band. Data are number of individuals demonstrating the various states (ques- 
tion mark indicates state uncertain due to fading or perspective of illustration). 



Tax a 
Locality 


E. 


heterodactxlus 
1 


E. crepitans 

2 


C. dos 

Guimaraes 

3 


7 
4 


7 

5 


7 
6 






E. fenestratus 






7 


8 


9 9a 


10 


Peru 


A 




9a 




















2 


B 






3 


2 




1 












12 


C 








9 


2 




7 


3 


8 


21 1 


21 


14 



From illustration 2a (Miranda-Ribeiro 1937). 



the variation in the Chapada dos Guimaraes 
specimen tubercles is distinctive from the 
variation observed in the E. fenestratus 
samples. 

Measurements: Male and female sizes 
vary among the samples (Table 12). Two 
points are worth noting, although additional 
samples are necessary to verify them statis- 
tically. There is no sexual size dimorphism 
in the Eleutherodactylus crepitans sample, 
whereas sexual size dimorphism is pro- 
nounced in all other samples that contain 
both sexes. The Chapada dos Guimaraes 
form is smaller than the Eleutherodactylus 
fenestratus samples analyzed. 

The results of multivariate discriminant 
function analyses for males and females 
(analyzed separately) agree in that the 
Eleutherodactylus crepitans specimens are 
clearly distinct from all other individuals 
analyzed (Fig. 3). The rest of the popula- 
tions analyzed indicate that there is differ- 
entiation among them, but the degree of 
variation is consistent with both intra- and 
inter-specific variation based on similar 
studies in other groups of frogs. 



Taxonomy 

Telatrema heterodactylum Miranda-Ri- 
beiro, 1937. — Alipio Miranda-Ribeiro de- 
scribed this species from "Matto-Grosso — 
Caceres, 2 exemplares colhidos na gruta 
dita 'Fazendinha' Comm. Rondon (1937: 
67)." His son Paulo Miranda-Ribeiro des- 
ignated specimen 106 A in the collection of 
the Museu Nacional, Rio de Janeiro, as the 
lectotype (1955:411). The second speci- 
men, the paralectotype, which also had the 
number 106 originally, was subsequently 
recatalogued as 5089. 

Dr. R E. Vanzolini (pers. comm.) was 
able to locate A. Miranda-Ribeiro 's locality. 
Miranda-Ribeiro was a member of the Ron- 
don Commission expedition and he collect- 
ed the specimens in question himself. Cac- 
eres, as used by Miranda-Ribeiro, refers to 
the municipality, which at the time of the 
Rondon Commission expedition was very 
extensive. Vanzolini found in the Rondon 
Commission itinerary that Miranda-Ribeiro 
travelled to "Fazendinha" and returned to 
Caceres in the same day and that "Fazen- 
dinha" was near a place called Pirizal. Van- 



Table 5. — Mid-dorsal stripe patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent; 
B = Present. Data are number of individuals demonstrating the states. 

C. dos 
Taxa E. heterodactylus E. crepitans Guimaraes ? ? 

Locality 1 2 3 4 5 



A 
B 



1 



11 







E. fenestratus 






7 


8 


9 


9a 


10 


Peru 


3 


8 


44 


1 


34 


26 






4 




12 


2 



8 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 6. — Dorsal patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Indistinct or 
uniform; B = Dark symmetrical spots; C = Dark chevron in medial scapular region followed by dark blotches 
posteriorly; D = Dark W-shaped mark in medial scapular region followed by dark blotches posteriorly; E = 
Extended dark W-shaped pattern (additional strophes to next in series from inverted V to W) in medial scapular 
region followed by dark blotches posteriorly. Data are number of individuals demonstrating the various states 
(question mark indicates uncertain condition due to fading). 



C. dos 

Taxa E. heienxlactxhis E. crepitans Guimaraes 

Locality I ' 2 3 



E. fenestratus 



7 


8 


9 


9a 


10 


Peru 


1 


3 


7 


1 


9 


13 




1 


3 




4 






4 


2 




2 




2 




9 




6 


15 



A 
B 
C 
D 
E 



From illustration 2a, b (Miranda-Ribeiro 1937). 



zolini located a "Fazendinha" that not only 
meets these conditions, but is in an area 
known to have sandstone caves (grutas = 
caves) at 16°00'S, 57°36'W (Fig. 2, Local- 
ity 1). 

Lynch & Duellman (1997:225) recog- 
nized Eleutherodactylus heterodactylus 
(Miranda-Ribeiro, 1937) as a distinct spe- 
cies and included it in the Eleutherodacty- 
lus binotatus species-group. As Lynch and 
Duellman did not examine the types, we of- 
fer the following observations. 

Both types are faded such that most fea- 
tures of any color patterns are no longer 
discernible. The lectotype is in poor con- 
dition, the paralectotype is in worse condi- 
tion. The paralectotype is very brittle and 
fragile and disintegrates more each time it 
is handled. The lectotype is the (noticeably) 
larger of the two, and the only specimen 
measured for purposes of this paper. WRH 
recorded the measurements (in mm) of the 
lectotype as: SVL 24.6 (contrasted to Mi- 



randa-Ribeiro's measurement of 27); head 
length 8.6; head width 8.5; eye-nostril dis- 
tance 3.5; eye-eye distance 5.1; greatest 
tympanum diameter 2.4; thigh 13.9; shank 
14.1; foot 12.0; width of 3rd finger disk 1.4; 
width of 4th toe disk 0.7. The lectotype has 
slits in the floor of the mouth, but they ap- 
pear to be cuts, not vocal slits. The sex of 
the lectotype can not be determined with 
certainty without dissection, which given 
the poor condition of the specimen was 
considered inappropriate. 

The following support recognition of Te- 
latrema heterodactylum as a distinct and 
valid species of Eleutherodactylus, confirm- 
ing Lynch & Duellmans' (1997) assess- 
ment. 

The disks on fingers 3 and 4 are very 
large, much larger than the disk on the 
fourth toe, and the shape is ovate (Fig. 4a). 
The disks on fingers 3 and 4 of the other 
Eleutherodactylus examined are expanded, 
but either the same size or just larger than 



Table 7. — Posterior face of thigh patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = 
Uniform; B = Finely mottled around vent, rest uniform; C = Finely mottled; D = Mottled. Data are number 
of individuals demonstrating the various states. 



C. dos Guima- 
Taxa E. heterodactylus E. crepitans raes 

Locality 1 ' 2 3 







E. 


fenestratus 






7 


8 


9 




9a 


10 


Peru 
















3 


7 


18 






20 


5 




1 


3 




1 


1 


8 
15 



A 
B 
C 
D 



VOLUME 112, NUMBER 1 



9 



Table 8. — Belly patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost immaculate; 
B = Noticeably flecked or mottled. Data are number of individuals demonstrating the states. Specimen from 
locality 6 too faded to score. 



Taxa 
Locality 


E. 


heterodactxlus E. crepitans 
1 2 


C. dos 

Guimaraes 

3 


■I 

4 


5 


•7 

6 






E. 


fenestratus 






7 


8 


9 




9a 


10 


Peru 


A 
B 




1 3 


10 

1 


2 


1 




3 


6 

2 


10 

11 




1 


11 

10 


28 



the disks on toe 4 and they are broadly tri- 
angular in shape (Fig. 4b, c). 

The type illustrations (most likely that of 
the lectotype, Miranda-Ribeiro 1937, Fig. 
2a-b) show a pair of ill-defined light lon- 
gitudinal stripes from behind the eye to the 
sacrum. This feature is not seen in any of 
the other Eleutherodactylus examined for 
this study. 

The type locality is characterized by a 
very open vegetation formation lacking gal- 
lery forests. Eleutherodactylus fenestratus 
is a forest denizen and the Chapada dos 
Guimaraes form inhabits well-developed 
gallery forests. Eleutherodactylus crepitans 
also occurs in open habitats, but is morpho- 
logically very distinct from E. heterodac- 
tylus. 

One other morphological feature will re- 
quire fresh material to evaluate. There is no 
indication of any tarsal fold or other tarsal 
decoration on either the lectotype or para- 
lectotype, suggesting that, if in fact the tar- 
sus lacks a fold, that would be another fea- 
ture distinguishing the species from all oth- 
er Mato Grosso Eleutherodactylus. 



Eleutherodactylus heterodactylus (Miran- 
da-Ribeiro 1937) is thus far known only 
from the original two specimens collected 
by Miranda-Ribeiro from "Fazendinha." 
All other members of the Eleutherodactylus 
hinotatus group, to which Lynch & Duell- 
man (1997) assigned E. heterodactylus, oc- 
cur in eastern Brazil, primarily in the At- 
lantic Forest Morphoclimatic Domain. In- 
clusion of E. heterodactylus in this group 
does not make zoogeographic sense. 

Status of Eleutherodactylus crepitans 
Bokermann, 1965. — Werner C. A. Boker- 
mann described the species based on three 
specimens from Sao Vicente, Mato Grosso. 
The holotype and allotype are now in the 
collections of the Museu de Zoologia, 
Universidade de Sao Paulo and the other 
paratype is in the Museu Nacional, Rio de 
Janeiro. 

John Lynch (1980:8) synonymized 
Eleutherodactylus crepitans Bokermann, 
1965 with E. fenestratus (Steindachner 
1864). Lynch did not examine the speci- 
mens of E. crepitans (1980:6). Lynch ar- 
gued that because Bokermann was unaware 



Table 9. — Tarsal fold states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent; B = 
Short, oblique, lying more than length of inner metatarsal tubercle from inner metatarsal tubercle; C = Mod- 
erately short, less than Vi length of tarsus, lying less than length of inner metatarsal tubercle from inner metatarsal 
tubercle; D = Long, greater than Vi length of tarsus, lying less than length of inner metatarsal tubercle from 
inner metatarsal tubercle. Data are number of individuals demonstrating the various states. Data not taken for 
one desiccated individual from locality 10. 



Taxa 
Legality 


£. 


heterodactxlus 
1 


E. 


crepitans 

2 


C. dos 

Guimaraes 

3 


A 




1 




1 




B 








2 




C 










10 


D 










1 







E. fenestratus 






7 


8 


9 9a 


10 


Peru 








1 




3 


8 


21 1 


19 


28 



10 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 10. — Toe fringe states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Toes free; 
B = Toes with lateral ridges; C = Toes with modest lateral fringes. Data are number of individuals demonstrating 
the states. 





E. 


hetenxiactxlus 
1 


E. crepitans 

2 


c 

Gui 


. dos 
maraes 
3 




4 


7 
5 


7 
6 






E . 


fenestratus 






Locality 


7 


8 


9 


9a 


10 


Peru 


A 




1 


3 
















I 




5 


5 


B 










3 










3 


8 




6 


17 


C 










8 


2 


1 


2 


3 


5 


12 


1 


10 


6 



that E. fenestratus was a valid species, Bok- 
ermann compared his new species with in- 
appropriate species from Ecuador, from 
which they were very distinct. However, 
had Lynch examined the type material of E. 
crepitans, he would have found that they 
differ from E. fenestratus as well. 

The following argue for recognizing 
Eleutherodactylus crepitans as a valid spe- 
cies. 

Eleutherodactylus crepitans has short, 
oblique tarsal folds, separated from the in- 
ner metatarsal tubercle by a distance greater 
than the length of the fold. Eleutherodac- 
tylus heterodactylus apparently lacks tarsal 
folds. Eleutherodactylus fenestratus and the 
Chapada dos Guimaraes form have longer 
(but still relatively short) tarsal folds, par- 
allel to the long axis of the tarsus, separated 
by a short distance (less than length of the 
fold) from the inner metatarsal tubercle. 

Eleutherodactylus crepitans does not 
have marked sexual dimorphism in size. 
Both Eleutherodactylus fenestratus and the 
Chapada dos Guimaraes form show marked 
sexual size dimorphism. 

The overall body shape of Eleutheroda- 
cytlus crepitans differs markedly from that 



of E. fenestratus and the Chapada dos Gui- 
maraes form (Fig. 3). 

Bokermann (1965:264) made a point of 
stating that the specimens of Eleutherodac- 
tylus crepitans were obtained in grass-shrub 
vegetation in the middle of large blocks of 
granite from such a dry habitat that he 
would never have expected to find frogs 
there. The immediate vicinity of Sao Vi- 
cente does have a granitic intrusion of rath- 
er small extent and is the only such granitic 
habitat in Mato Grosso (Brasil 1982). It 
would not be surprising if E. crepitans is 
limited to this distinctive habitat. 

Bokermann (1965:264) indicated that 
males called at night from sandy hiding 
places among rocks. He described the call 
as consisting of two notes in sequence, like 
a short and coarse click, repeated many 
times but irregularly (translation by P. E. 
VanzoUni). The two note call of Eleuther- 
odactylus crepitans is more similar to 
Eleutherodactylus fenestratus than the Cha- 
pada dos Guimaraes form, but the frequent 
calling of E. crepitans is different from the 
very infrequent calls of both E. fenestratus 
and the Chapada dos Guimaraes form. 

Questionable status specimens. — The 



Table 1 1 . — Maximum number of tubercles on sole of foot. Locality numbers refer to those in Fig. 2 (9a not 
mappable). Data are number of individuals demonstrating the states. 



Taxa E. heterodactylus E. crepitans 

Locality 1 2 



C. dos 

Guimaraes 

3 







E. 


fenestratus 






7 


8 


9 


9a 


10 


Peru 










1 






5 


9 




9 


23 


1 


2 


9 




9 


5 


2 


1 


3 


1 


2 





VOLUME 112, NUMBER 1 



11 



Table 12. — Adult size. Locality numbers refer to those in Fig. 2 (9a not mappable). Data are SVL ranges in mm. 



Taxa 

Locality 


E. hetero- 

dacnlus 

\ 


E. crepitans 

2 


C. dos 

Guimaraes 

3 


9 

4 


•> 

5 


1 

6 






E. fenestratus 






7 


8 


9 


9a 


10 


Peru 


Females 




30 


34-36 






34 


38^1 




37^0 


42 


37-44 


40-52 


Males 




28-29 


22-27 




30 








30 




26-32 


27-34 


Largest juveniles 


25 






30 








29 











name Hylodes gollmeri bisignata Werner, 
1 899, which based on geography should be 
considered as an available name for the 
frogs considered in this paper, and speci- 
mens of Eleutherodactylus from three lo- 
calities in Mato Grosso can not be unequiv- 
ocally assigned to either Eleutherodactylus 
crepitans, fenestratus, heterodactylus, or 
the Chapada dos Guimaraes form. Adver- 
tisement calls are not available at present 
for any of the specimens involved. At this 
time, they are best left as indeterminate. 

Werner (1899) gave a brief description of 
Hylodes gollmeri bisignata without locality 
information in the type description. Haupl 
& Tiedemann (1978) indicated that the ho- 
lotype, NMW 16502, in the Naturhisto- 
rischen Museums Wien collection was from 
Chaco, Bohvia. Lynch & Duellman (1997) 
considered bisignatus to be a synonym of 
Eleutherodactylus fenestratus. Based on the 
fact that the Chaco and Amazonian hylaea 
frog faunas are virtually distinct from each 
other at the species level, the validity of E. 
bisignatus should be reconsidered. The 
original description is too brief to be of help 
in evaluating this suggestion. Dr. Lynch ex- 
amined the type of H. g. bisignata, but his 
original notes are not in the appropriate 
folder in his archives (pers. comm.). The 
Naturhistorisches Museum Wien now has a 
policy of not loaning types, precluding our 
examination of the type for this paper. We 
suspect that bisignatus is a species of 
Eleutherodactylus distinct from E. fenestra- 
tus. If it is the same as any species that 
occurs in Mato Grosso, it might be conspe- 
cific with E. heterodactylus, based on most 
similar (but still quite different) habitat oc- 
currences. 



The two specimens from the Panatanal 
locality of Fazenda do Santa Edwiges (Fig. 
2, Locality 4) could either be the same as 
the Chapada dos Guimaraes form or a dis- 
tinct species. The smaller specimen 
(MZUSP 71103) has a tarsal fold like that 
of the Chapada dos Guimaraes form, but 
the larger specimen (MZUSP 71104, which 
looks like it died before it was preserved) 
appears to have a much longer tarsal fold. 

The single specimen from Jacubim 
(MZUSP 4277) has completely mottled 
thighs with a large pattern, such that the 
white blotches almost cover the same total 
area as the darker ground hue. The thigh 
pattern differs markedly from other speci- 
mens discussed in this paper, but is closer 
to that of specimens identified as Eleuth- 
erodactylus cf peruvianus (e.g., MZUSP 
80854, 80856-80857, 80859, 80863) from 
the Amazon forest locality of Apiacas. Ja- 
cubim, however, is in an area of Cerrado- 
seasonal forest contact (Fig. 2, Locality 5). 

The two specimens from Barra do Tapi- 
rapes are very faded and their proper tax- 
onomic allocation may never be ascertained 
until fresh specimens from the locality are 
available. The fading extends to the poste- 
rior surfaces of the thighs, but it does ap- 
pear that the thighs are mottled in a similar 
fashion to the pattern observed in MZUSP 
4277 from Jacubim, with which they may 
be conspecific. 

Status of the Chapada dos Guiraraes 
form. — As documented above, the Chapada 
dos Guimaraes Eleutherodactylus repre- 
sents a distinct species from all other 
known Eleutherodactylus, which we de- 
scribe as: 



12 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



MALES 



4- 



CJ 2- 

O 



-4 




— r- 

-4 



— r 
-2 







2 



-r 
4 



FEMALES 



6- 
4- 
2- 
0- 
-2- 



-10 



FACTOR 




— r- 

-5 



g^ 



B 




10 



Fig. 3. Discriminant function analysis results for males and females. Minimum polygons contain all individuals 
for sample sizes >3. Dots = Eleutherodactylus crepitans; A = sample from Aripuana, Mato Grosso; B = individual 
from Barra do Tapirapes, Mato Grosso; C = sample from Chapada dos Guimaraes, Mato Grosso; J = individual 
(male) and sample (females) from Juruena, Mato Grosso; P = sample from Peru; 5 = individual from Jacubim, 
Mato Grosso. 



Eleutherodactylus dundeei, 

new species 

Figs, lb, 5 

Holotype.—MZUSV 79834, adult male 
from Brazil: Mato Grosso; Chapada dos 
Guimaraes, Veu da Noiva, Rio Coxipozin- 
ho, approximately 15°25'S, 55°47'W. Col- 
lected by Miriam H. Heyer, W. Ronald Hey- 
er, and Liliam Patricia Pinto on 25 Sep 
1996. 

Paratopotypes.—MLVS? 79835-79837 
(1 female, 2 males), same data as holotype. 

Paratypes. — All from Brazil: Mato Gros- 



B 






Fig. 4. Diagrammatic outlines of shape of third 
finger disk based on free-hand sketches. A. Holotype 
of Telatrema heterodactylum, actual width of disk 1 .4 
mm. B. Holotype of Eleutherodactylus crepitans, ac- 
tual width 0.9 mm. C. Chapada dos Guimaraes form, 
actual width 1.3 mm (based on USNM 507897). 



so; Chapada dos Guimaraes; MNRJ 19785 
(female), collected by Erminia (UFF) on 24 
Oct 1987; MZUSP 76237 (juvenile), an 
originally unnumbered specimen from the 
Werner C. A. Bokermann collection col- 
lected by A. Sebben and A. Schwartz on 15 
Jun 1988; MZUSP 85614 (male), originally 
WCAB 15546 collected by M. Alvarenga, 
F. M. Oliveira, and Werner C. A. Boker- 
mann from 15-25 Nov 1963; USNM 
507897-507898 (female, male), Casa de 
Pedra, collected by Miriam H. Heyer, W 
Ronald Heyer, Liliam Patricia Pinto on 28 
Sep 1996; USNM 507899 (male, call 
voucher for Fig. lb), Estancia Monarca, 
collected by Miriam H. Heyer, W. Ronald 
Heyer, Liliam Patricia Pinto on 30 Sep 
1996; USNM 507900 (female), Escola 
Evangelica de Buriti, collected by Miriam 
H. Heyer, W Ronald Heyer, Liliam Patricia 
Pinto on 1 Oct 1996. 

Diagnostic definition (scheme established 
by Lynch, e.g., 1979, of numbered charac- 
ter states followed here for ease of com- 
parison). — Skin of upper surfaces finely 



VOLUME 112, NUMBER 1 



13 




Fig. 5. Dorsal view of paratype of Eleutherodactylus dundeei, new species, USNM 507899. This specimen 
has the most contrasting pattern among the type series. Other specimens have similar, but fainter markings. 



pebbled with few to many scattered white 
fleshy tubercles, tubercles denser posteri- 
orly, skin of throat, chest, and anteriormost 
belly smooth, rest of belly weakly areolate; 
tympanum distinct, its horizontal diameter 
about V2-% eye opening diameter; snout su- 
belliptical in dorsal view, rounded in pro- 
file; upper eyelid width equal to or (usually) 
slightly smaller than interorbital distance. 



with same fine pebbling of back with scat- 
tered, low, flattened, light tubercles; no cra- 
nial crests; vomerine teeth in two small 
patches well posterior to choanae, separated 
from each other by about the width of a 
single tooth patch; males with vocal slits 
and single subgular vocal sac, either notice- 
ably expanded and wrinkled in preservation 
or not; male nuptial pads of two weak light 



14 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



glandular patches; first finger about same 
size or just longer than second; fingers bear- 
ing disks (largest on III and IV); largest 
disks broader than long; fingers free or with 
weak lateral keels; 2-7 light spots indicat- 
ing weakly developed ulnar tubercles (tu- 
bercles themselves visible in only some in- 
dividuals); short tarsal fold, about Vs length 
of tarsus, otherwise tarsus smooth; heel 
smooth to granular, lacking tubercle(s); 2 
metatarsal tubercles, inner oval, 2-3 times 
size of outer; 0-2 fleshy plantar tubercles; 
toes free or (usually) bearing weakly to well 
developed lateral keels; individuals with 
well-developed keels with basal toe web- 
bing; predominantly brown frog, no flash 
colors; posterior surfaces of thighs flesh 
color-purplish in life, tan/brown in preser- 
vative; adults small, males 22—27 mm, fe- 
males 34-36 mm SVL. 

Eleutherodactylus dundeei is most simi- 
lar to and most likely to be confused with 
Eleutherodactylus fenestratus. Eleuthero- 
dactylus dundeei is smaller (males 22—27 
mm, females 34-36 mm SVL) than E. fe- 
nestratus (males 26—34 mm, females 37-52 
mm SVL). The belly of E. fenestratus is 
either entirely smooth; or weakly areolate 
only on the lateralmost portions, but not 
areolate posteriorly; the belly of E. dundeei 
is weakly areolate at least posteriorly. Male 
E. dundeei have two weakly developed 
light glandular nuptial asperities on each 
thumb; male E. fenestratus have a single, 
better developed light glandular nuptial as- 
perity on each thumb. 

Description of holotype. — Snout shape 
subelliptical in dorsal outline, rounded in 
profile; canthus rostralis sharply rounded; 
lip weakly flared; upper eyelid width about 
equal interocular distance; no external in- 
dications of cranial crests; tympanum dis- 
tinct, horizontal diameter including annulus 
about Vi eye opening diameter, horizontal 
diameter < vertical diameter; vomerine 
teeth in two small patches, well posterior to 
rounded choanae, separated from each other 
by about width of a single tooth patch; vo- 
cal slits present, elongate; vocal sac single. 



subgular, slightly expanded externally in 
preservation; finger lengths I just 
>II<IV<III (left finger IV mostly miss- 
ing); finger tips on fingers I and II slightly 
expanded, small, round, lacking circumfer- 
ential grooves, fingers III and IV with ex- 
panded disks, triangular, broader than long, 
with circumferential groove and upper sur- 
face weakly to noticeably notched; fingers 
lacking noticeable lateral keels; no finger 
webbing; palmar tubercle large, ovate- 
heart-shaped, larger than proximate ovoid 
thenar tubercle, one well-developed super- 
numerary palmar tubercle associated with 
each digit; subarticular tubercles distinct, 
rounded, single, each thumb with 2 weakly 
developed but distinct patches of light glan- 
dular nuptial excrences; outer ulnar region 
with a series of 6-7 light spots, apparently 
with low tubercles in life, but only one 
weakly visible on each arm; supratympanic 
fold distinct; no other distinct glands or 
folds on body; upper eyelid weakly pebbled 
with several light tubercles, rest of back 
texture weakly pebbled with scattered light 
fleshy tubercles; throat smooth, discoidal 
belly fold distinct, chest and adjoining belly 
to about mid-belly smooth, sides of and 
posterior belly weakly areolate, ventro-pos- 
terior surfaces of thighs areolate, rest of 
ventral limb surfaces smooth; toe lengths 
I<II«III<V<IV (condition B as used in 
Lynch & Duellman 1997, appendix III); 
disks on toes I and V small, round, disks 
on toes II, III, IV increasing in size, largest 
disks triangularly ovate, broader than long, 
upper surfaces emarginate or weakly 
notched; sides of toes keeled, strongest on 
inner sides of toes II, III, FV, forming basal 
web, not encompassing the basal subarti- 
cular tubercles, best developed between 
toes II, III, IV; outer round metatarsal tu- 
bercle about Va size of ovate inner metatar- 
sal tubercle; tarsal fold distinct, short, about 
Va length of tarsus, separated from inner 
metatarsal tubercle by a distance of about 
Vi length of tarsal fold, rest of outer tarsus 
smooth; heel smooth, lacking tubercle(s) or 
calcar; plantar surface with one prominent 



VOLUME 112, NUMBER 1 



15 



fleshy tubercle; subarticular tubercles mod- 
erately developed, weakly pungent. 

Coloration in alcohol: A tan frog with 
brown markings. Tip of snout lightest, ex- 
panding into somewhat darker but light tri- 
angle bounded by canthus rostralis and dark 
interorbital mark, with a few brown dots 
and lines. Dark canthal stripe in front of eye 
only extending about Va distance to nostril. 
Loreal region almost uniform brown. Upper 
lip indistinctly barred. Supratympanic fold 
dark brown. Faint interorbital bar, defined 
by straight solid brown line anteriorly, very 
weakly defined posteriorly as a shallow U 
with a few dark dots in the interorbital 
mark. A faint dark W- shaped mark in the 
pre-scapular area followed by a median, 
faint, inverted Y-shaped mark between 
scapular and sacral region, followed by two 
median irregular sacral chevrons. Notice- 
ably dark pair of lateral spots behind eyes 
in scapular region. Upper limbs tan with ir- 
regular darker cross bands. Series of 6-7 
light spots on outer ulnar region. Throat 
speckled with melanophores. Chest and bel- 
ly light, almost immaculate, but with a very 
few scattered melanophores. Posterior sur- 
faces of thighs with very fine mottle pattern 
on area just around vent, otherwise uniform 
tan. Outer tarsus and sole of foot dark 
brown. 

Measurements of holotype. — SVL 23.6 
mm, head length 9.6 mm, head width 7.9 
mm, eye opening diameter 3.0 mm, eye- 
nostril distance 3.0 mm, eye-eye distance 
4.9 mm, horizontal tympanum diameter 1.8 
mm, vertical tympanum diameter 1.9 mm, 
thigh length 11.6 mm, shank length 13.3 
mm, foot length 11.7 mm, width of 3rd fin- 
ger disk 1.0 mm, width of 4th toe disk 0.7 
mm. 

Variation. — Most details of variation 
have been described in the preceding sec- 
tions. Additional measurement variations 
are (broken down by sex only when range 
of one not subsumed in other): head length 
39-42% SVL; head width 33-38% SVL; 
eye-nostril distance 12-14% SVL; eye-eye 
distance 20-22% SVL for males, 18-19% 



for females; tympanum diameter 7-9% 
SVL; thigh length 48-53% SVL; shank 
length 52-57% SVL; foot length 47-54% 
SVL for males, 46—53% for females; width 
of 3rd finger disk 3-4% SVL; width of 4th 
toe disk 3-4% SVL. 

Color in life. — Iris bright bronze above, 
dull bronze below; posterior face of thigh 
flesh color-purplish; belly white with yel- 
low wash; no flash colors (based on USNM 
507897-507898). 

Advertisement call. — Described in pre- 
vious section (Fig. lb). 

Etymology. — Named for Dr. Harold A. 
Dundee for his personal, financial, and sci- 
entific contributions to the field of herpe- 
tology. 

Ecological notes. — Data are available for 
eight individuals. Five of them were col- 
lected in the late morning on dead leaves 
or under 5 cm diameter rocks on the 
ground, four of these 3-6 m from a river 
bank in rainforest-like gallery forest 
(MZUSP 79834-79837), one of these on 
dead leaves on the ground on the bank of 
a tree-shaded pond (USNM 507900). Two 
individuals were taken shortly after dark in 
cerrado gallery forest 2 m from a 2 m wide 
stream either on the surface of the soil 
(USNM 507897) or on a 0.3 m diameter log 
(USNM 507898). The call voucher (USNM 
507899) was taken at night from secondary 
cerrado vegetation on a leaf of a herbaceous 
plant, less than 1 m above ground. 

Distribution. — All known specimens 
have been collected from within 25 km of 
the town of Chapada dos Guimaraes in the 
limited region identified as having meso- 
philic seasonal forest vegetation cover in 
the Cartograma de Vegeta9ao do Estado de 
Mato Grosso, produced by the Consorcio 
Nacional de Engenheiros Consultores S.A., 
published by the Govemo do Estado de 
Mato Grosso (Fig. 6). There are other is- 
lands of similar vegetation cover in the 
same region (Fig. 6) which might be ex- 
pected to contain Eleutherodactylus dun- 
deei. We predict that either E. dundeei is 
restricted to the known Chapada dos Gui- 



16 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Major vegetation types in portion of the State of Mato Grosso surrounding Chapada dos Guimaraes. 
Black areas are mesophilic seasonal forests. Stippled areas are dense cerrado (cerradao) and forest/cerrado contact 
regions. Rest of region is covered by various forms of cerrado vegetation (except cerradao). Open square is the 
city of Cuiaba. 

Eleutherodactylus dundeei is thus far only known from the mesophilic seasonal forest block identified as 3 
on the map, but other nearby blocks should be explored for the species. Map redrawn from "Cartograma de 
Vegeta^ao do Estado do Mato Grosso," produced by Consorcio Nacional de Engenheiros Consultores S.A. 



maraes vegetation type block or to it and 
one or more of the additional habitat islands 
identified in Fig. 6. 

Discussion 

Eleutherodactylus fenestratus, with the 
removal of Eleutherodactylus crepitans and 
dundeei, has an Amazonian rainforest dis- 
tribution (assuming Eleutherodactylus bi- 
signatus is not a synonym of E. fenestra- 
tus). There has not been an adequate study 
of variation for E. fenestratus. The limited 
data we analyzed indicate that there is at 
least interpopulational variation in E. fenes- 
tratus. Analyzing variation throughout the 
entire range of the species would be an ob- 



vious next step to understand the signifi- 
cance of the variation we found. 

Chapada dos Guimaraes has a very in- 
teresting frog fauna from a zoogeographical 
perspective. The landscape surrounding the 
Chapada is cerrado. There are expected cer- 
rado frog species at Chapada dos Guimar- 
aes, such as Bufo paracnemis, Scinax fus- 
covarius, Leptodactylus labyrinthicus, and 
Physalaemus nattereri. There are also Cha- 
co and Amazonian elements represented. 
For example, the Chaco species Leptodac- 
tylus chaquensis was collected syntopically 
with the Amazonian species Leptodactylus 
mystaceus in 1996. Finally, there are at least 
two species thus far known only from Cha- 
pada dos Guimaraes, Phyllomedusa cen- 



VOLUME 112, NUMBER 1 



17 



trails Bokermann, 1965 and Eleutherodac- 
tylus dundeel. Obviously, the frog fauna of 
Chapada dos Guimaraes represents a rich 
and complicated history, worthy of much 
more detailed study. There is actually a fair 
amount of information available for frogs 
from Chapada dos Guimaraes to base such 
a study upon. Herbert Huntingdon Smith 
collected materials reported on by E. D. 
Cope (1887) and housed at the Academy of 
Natural Sciences in Philadelphia. At least 
one of the species reported, Leptodactylus 
petersll, has not been collected since from 
Chapada dos Guimaraes (Heyer 1994:79, 
Cope's Leptodactylus brevlpes = L. peter- 
sii). Boulenger (1903) listed nine species 
collected by the Percy Sladen Expedition to 
central Brazil. The species Boulenger listed 
as Hylodes gollmerl is probably the species 
we describe as Eleutherodactylus dundeel. 
All other species of Boulenger's list are 
plausible (with taxonomic adjustments) ex- 
cept for Ceratophrys crlstlceps, which 
should be re-examined. Werner C. A. Bok- 
ermann and colleagues made collections 
which now (for the most part) are at 
MZUSP MZUSP also has materials col- 
lected by Drs. Rita Kloss and Francisca C. 
do Val. Adao Cardoso also collected at 
Chapada dos Guimaraes, and those speci- 
mens are housed at the Universidade Estad- 
ual de Sao Paulo in Campinas. As Chapada 
dos Guimaraes is now a tourist attraction, 
it is likely that there are other incidental 
collections waiting study in various collec- 
tions as well. 

Acknowledgments 

PRODEAGRO and CENEC made the 
fieldwork possible during which the record- 
ings of Eleutherodactylus dundeel were ob- 
tained. Without these recordings, this paper 
would not have been initiated. Drs. John D. 
Lynch and P. E. Vanzolini critically re- 
viewed the manuscript. AMM thanks the 
National Museum of Natural History for 
the opportunity to participate in the 1997 
Research Training Program. WRH thanks 



Dr. P. E. Vanzolini and the Museu de Zool- 
ogia da Universidade de Sao Paulo for char- 
acteristic help and access to the collections. 
Dr. Ulisses Caramaschi facilitated exami- 
nation of specimens in the collections of the 
Museu Nacional, Rio de Janeiro. WRH's re- 
search for this paper was supported by the 
Neotropical Lowlands Research Program, 
National Museum of Natural History, Dr. 
Richard P. Vari, Principal Investigator. 

Literature Cited 

Bokermann, W. C. A. 1965. Tres novos batraquios da 
regiao central de Mato Grosso, Brasil (Amphib- 
ia, Salientia). — Revista Brasileira de Biologia 
25:257-264. 

Boulenger, G. A. 1903. List of the batrachians and rep- 
tiles collected by M. A. Robert at Chapada, 
Matto Grosso, and presented by Mrs. Percy 
Sladen to the British Museum. (Percy Sladen 
Expedition to Central Brazil). — Proceedings of 
the Zoological Society of London 1903 II(I): 
69-70. 

Brasil. Ministerio das Minas e Energia. 1982. Projeto 
RADAMBRASIL. Folha SD.21 Cuiaba; geo- 
logia, geomorfologia, pedologia, vegeta^ao e 
uso potencial da terra. Levantamento de Recur- 
sos Naturals, Vol. 26. Rio de Janeiro, 544 pp., 
5 maps. 

Cope, E. D. 1887. Synopsis of the Batrachia and Rep- 
tilia obtained by H. H. Smith, in the Province 
of Mato Grosso, Brazil. — Proceedings of the 
American Philosophical Society, Philadelphia 
24:44-60. 

Duellman, W. E., & L. Trueb. 1986. Biology of am- 
phibians. McGraw-Hill Book Company, New 
York, 670 pp. 

Haupl, M., & E Tiedemann. 1978. Typenkatalog der 
Herpetologischen Sammlung. — Kataloge der 
wissenschaftlichen Sammlungen des Naturhis- 
torischen Museums in Wien. Vertebrata. 2:1-35. 

Heyer, W. R. 1984. Variation, systematics, and zoo- 
geography of Eleutherodactylus guentheri and 
closely related species (Amphibia: Anura: Lep- 
todactylidae). — Smithsonian Contributions to 
Zoology 402:1-42. 

. 1994. Variation within the Leptodactylus pod- 

icipinus-wagneri complex of frogs (Amphibia: 
Leptodactylidae). — Smithsonian Contributions 
to Zoology 546:1-124. 

, A. S. Rand, C. A. G. Cruz, O. L. Peixoto, & 



C. E. Nelson. 1990. Frogs of Boraceia. — Ar- 
quivos de Zoologia 31:231-410. 
Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. 
Dawson. 1985. Standards in herpetology and 



18 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ichthyology: Part I. Standard symbolic codes 
for institutional resource collections in herpe- 
tology and ichthyology. — Copeia 1985:802- 
832. 

Lynch, J. D. 1979. Leptodactylid frogs of the genus 
Eleiitherodactyhis from the Andes of southern 
Ecuador — University of Kansas Museum of 
Natural History Miscellaneous Publication 65: 
1-62. 

. 1980. A taxonomic and distributional synop- 
sis of the Amazonian frogs of the genus Eleuth- 
erodactylus. — American Museum Novitates 
2696:1-24. 

, & W. E. Duellman. 1997. Frogs of the genus 



Eleutherodactylus in western Ecuador: System- 
atics, ecology, and biogeography. — The Univer- 
sity of Kansas Natural History Museum Special 
Publication 23:1-236. 

Marquez, R., I. De la Riva, & J. Bosch. 1995. Adver- 
tisement calls of Bolivian Leptodactylidae (Am- 
phibia, Anura). — Journal of Zoology, London 
237:313-336. 

Miranda-Ribeiro, A. 1937. Alguns batrachios novos 
das collecgoes do Museu Nacional. — O Campo 
May 1937:66-69. 

Miranda-Ribeiro, P. 1955. Tipos das especies e subes- 
pecies do Prof. Alipio de Miranda Ribeiro de- 
positados no Museu Nacional (Com uma rela- 
gao dos generos, especies e subespecies descri- 
tos). — Arquivos do Museu Nacional, Rio de Ja- 
neiro 42:389-417. 

Penna, M. 1997. Selectivity of evoked vocal responses 
in the time domain by frogs of the genus Ba- 
trachyla. — Journal of Herpetology 31:202-211 . 

Schneider, H., T S. Sofianidou, & P. Kyriakopoulou- 
Sklavounou. 1984. Bioacoustic and morpho- 
metric studies in water frogs (genus Rana) of 



Lake loannina in Greece, and description of a 
new species (Anura, Amphibia). — Zeitschrift 
fiir zoologischen Systematik und Evolutionsfor- 
schung 22:349-366. 
Werner, F 1899. Beschreibung neuer Reptilien und Ba- 
trachier. — Zoologischer Anzeiger 22:479-484. 

Appendix 1 
Specimens examined 

Eleutherodactylus crepitans. — Brazil: Mato Grosso; 
Sao Vicente, 15°38'S, 55°23'W, MNRJ 3985 (para- 
type), MZUSP 73671 (allotype), 85628 (holotype). 

Eleutherodactylus dundeei. — See species account. 

Eleutherodactylus fenestratus. — Brazil: Mato Gros- 
so; [Alto Juruena — see preceding Juruena], Apiacas, 
09°34'S, 57°23'W, MZUSP 80851-80853, 80855, 
80858, 80860-80862; Aripuana, 10°24'S, 59°19'W, 
MZUSP 80643-80648, 80652-80654, 86410; Alto Ju- 
ruena (locality is the upper Rio Juruena and cannot be 
assigned specific coordinates), MZUSP 85540; Jurue- 
na, 10°18'S, 58°32'W, MZUSP 86153-86200; Sao Jose 
do Rio Claro, 13°26'S, 56°44'W, MZUSP 86358- 
86360. 

Peru: Madre de Dios; Pakitza, 11°58'S, 71°14'W, 
USNM 333014, 342618-342622, 342851-342853, 
345172; Tambopata, 12°50'S, 69°17'W, USNM 
222268, 247298-247304, 247630-247631, 268941- 
268944, 314901, 332438-332439, 342993. 

Eleutherodactylus heterodactylus. — Brazil: Mato 
Grosso; Fazendinha, near Pirizal, 16°00'S, 57°36'W, 
MNRJ 106 (lectotype), 5089 (paralectotype). 

Eleutherodactylus sp. — Brazil: Mato Grosso, Barra 
do Tapirapes, 10°39'S, 50°36'W, MZUSP 85645- 
85646; Jacubim, 13°20'S, 52°09'W, MZUSP 4277; Fa- 
zenda Santa Edwiges, 16°38'S, 55°11'W, MZUSP 
71103-71104. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):19-39. 1999. 

A new genus and species of frog from Bahia, Brazil (Amphibia: 
Anura: Leptodactylidae) with comments on the zoogeography of the 

Brazilian campos rupestres 

W. Ronald Heyer 

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0109, U.S.A. 

Abstract. — A new genus and species of frog, Rupirana cardosoi, is described 
from the northern Espinhago Range in the State of Bahia, Brazil. The new 
genus shares most character states with the genus Thoropa, but cladistic anal- 
ysis of morphological data indicates that most of these shared features are 
primitive states. The cladistic analysis indicates that Rupirana and Thoropa do 
not have a close sister-group relationship with each other. Rupirana cardosoi 
is another addition to the many species endemic to the campos rupestres of the 
Espinha90 Range. The amphibians of the campos rupestres show a much stron- 
ger biogeographical affinity with the Atlantic Forest biota than other groups 
studied; these other groups show a much stronger affinity with the biota of the 
diagonal of open formations running from northeast Brazil (caatingas-cerrados) 
to Argentina and Paraguay (Gran Chaco). 



Several years ago, Dr. Miguel T. Ro- 
drigues brought my attention to a series of 
unusual frogs he had collected at two lo- 
calities in the State of Bahia, Brazil, asking 
me if I knew what they were. I did not. Drs. 
Rodrigues and R E. Vanzolini kindly put 
the specimens at my disposal for further 
study, but for various reasons, detailed 
study was delayed. Shortly after I had ex- 
amined aspects of the myology and skele- 
ton of the taxon, I had occasion to evaluate 
some of the problematical specimens of the 
Werner C. A. Bokermann collection as they 
were being catalogued into the Museu de 
Zoologia da Universidade de Sao Paulo col- 
lection. I found a series of the same taxon 
from a third locality in Bahia. All known 
localities are in the distinctive campos ru- 
pestres formation of Brazil. The purpose of 
this paper is to demonstrate that the new 
frog differs from all other known leptodac- 
tylid genera, provide a description for it, 
and briefly comment on its relationships 
and zoogeography. 



Methods and Materials 

One male (MZUSP 65204) was superfi- 
cially dissected to obtain myological infor- 
mation on jaw, hyoid, and thigh muscula- 
ture for the characters used in a previous 
study of leptodactylid relationships (Heyer 
1975). A second female specimen (MZUSP 
68959) was cleared-and-stained using the 
double-staining technique for cartilage and 
bone (Dingerkus & Uhler 1977) to evaluate 
osteological features. 

For the species description, measurement 
data were taken with a dial calipers to the 
nearest 0.1 mm following the definitions in 
Heyer et al. (1990). 

Relationships were analyzed with PAUP 
3.1 (Swofford 1993). 

Comparison with Leptodactylid Genera 

Lynch (1971) still provides the most 
complete data set for leptodactylid genera, 
providing a baseline for comparison with 
subsequent studies. He recognized four 
New World subfamilies of the family Lep- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. I. Vertebral column of Rupirana cardosoi, 
MZUSP 68959. Cartilage stippled. Scale line = 2 mm. 



todactylidae: Ceratophryinae; Elosiinae; 
Leptodactylinae, and Telmatobiinae. The 
Australian and African leptodactyloids 
Lynch treated as subfamilies of the family 
Leptodactylidae are now recognized as the 
separate families Myobatrachidae and He- 
leophrynidae, respectively (Frost 1985; 
Duellman & Trueb 1986). 

The Bahia frog is not a member of the 
subfamilies Ceratophryinae, Elosiinae, or 
Leptodactylinae as defined by Lynch 
(1971). Rather than document this state- 
ment with a comparison of all states Lynch 
used, only a single character is used as an 
example. Ceratophryinae members have 
widely expanded transverse processes of 
the anterior presacral vertebrae; the Bahia 
frog does not (Fig. 1). Elosiinae species 
have a pair of dermal, scute-like glandular 
pads on the dorsal surface of each digital 
disk; the Bahia form lacks digital disks. 
Leptodactylinae genera have a bony sternal 
style; the sternum is a cartilaginous plate in 
the Bahia frog (Fig. 2). 

Lynch (1971:112-113) listed nine char- 



acters defining the subfamily Telmatobiinae 
(character numbers are those used by 
Lynch): (1) sternum cartilaginous; (2) ver- 
tebral shield lacking; (3) transverse pro- 
cesses of the anterior presacral vertebrae 
not widely expanded; (9) when present, 
maxillary teeth blunt and pedicellate; (35) 
mandible lacking odontoids; (39, the 38 is 
a typographical error on p. 113) m. petro- 
hyoideus anterior and m. stemohyoideus in- 
sert on the lateral edge of hyoid plate; (48) 
eggs laid in water, in terrestrial situations, 
or in bromeliads. The egg deposition site is 
unknown for the Bahia frog; it agrees in the 
rest of the character states listed for the sub- 
family. 

Lynch (1971) defined five tribes of Tel- 
matobiinae and one genus he was unable to 
assign to any of these tribes: Alsodini; 
Eleutherodactylini; Grypiscini; Odonto- 
phrynini; Telmatobiini; and the genus Scy- 
throphrys. 

The Bahia frog cannot be assigned to the 
Eleutherodactylini, Grypiscini, Odonto- 
phrynini, Telmatobiini, or Scythrophrys. 
Again, for sake of brevity, only examples 
are used to document this statement. Mem- 
bers of the Eleutherodactylini have rounded 
sacral diapophyses and males lack comified 
nuptial asperities; the Bahia frog has flat- 
tened sacral diapophyses and the males 
have comified nuptial asperities. In species 
of the Grypiscini, the frontoparietal fonta- 
nelle is not exposed; in the Bahia frog it is 
(Fig. 3). Members of the Odontophrynini 
have short transverse processes of the pos- 
terior presacral vertebrae; the Bahia frog 
has long processes (Fig. 1). Members of the 
Telmatobiini have cervical cotyles that are 
narrowly separated with two distinct artic- 
ular surfaces (Lynch's type II, 1971:54); the 
Bahia frog has widely spaced cervical co- 
tyles with two distinct articular surfaces 
(Fig. 4, Lynch's type I, 1971:53). Scythro- 
phrys has rounded sacral diapophyses and 
a concealed tympanum; the Bahia frog has 
flattened sacral diapophyses and an ex- 
posed, well-developed tympanum. 

Lynch (1971:123-124) defined the Tribe 



VOLUME 112, NUMBER 1 



21 




Fig. 2. Ventral portion of pectoral girdle of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Scale 
line = 1 mm. 



Alsodini based on 16 characters (numbers 
are those of Lynch): (3) transverse process- 
es of posterior presacral vertebrae long; (4) 
cervical cotylar arrangements type I or II; 

(5) cervical and second vertebrae not fused; 

(6) cranial bones not involved in dermos- 
tosis; (7) omostemum present, moderately 
large; (8) sacral diapophyses somewhat en- 
larged; (9) maxillary teeth blunt, pedicel- 
late; (12) facial lobe of maxilla deep, not 
exostosed; (17) nasals not in contact with 



frontoparietals; (18) frontoparietal fontanelle 
exposed, moderate-sized; (19) frontopari- 
etal not fused with prootic; (21) temporal 
arcade lacking; (37) alary processes of hy- 
oid plate on narrow stalks; (42) male with 
cornified nuptial asperities on thumb; (45) 
outer metatarsal tubercle present, inner 
metatarsal tubercle normal; (46) larvae with 
median vent. Larvae are unknown for the 
Bahia frog. In all other characters, except 
(8), the Bahia frog matches exactly the def- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 








Fig. 3. Dorsal view of skull of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Frontoparietal fon- 
tanelle hatched. Scale line = 1 mm. 



inition of the subfamily. In the Bahia frog, 
the sacral diapophyses are flattened (not 
rounded), but they are not really enlarged 
(Fig. 1). However, the differences involved 
in the shape of the sacral diapophyses are 
minor and alone do not argue for a distinct 
tribe status for the Bahia frog. 

Lynch (1971) included four genera in his 
concept of Alsodini: Batrachyla, Eupso- 
phus, Hylorina, and Thoropa (in 1978 he 
restricted the Tribe Batrachylini to the gen- 
era Batrachyla and Thoropa). Of these, the 
Bahia frog is most similar to Thoropa. Eup- 
sophus and Hylorina have the type II cer- 
vical cotylar arrangement; the Bahia frog 



has type I. The maxillary arch of Batrachy- 
la is incomplete and the quadratojugal is 
absent; the maxillary arch of the Bahia frog 
is complete (Figs. 3-4). 

Lynch (1971:129-130) defined the genus 
Thoropa by 31 characters (following his 
numbers): (4) cervical cotylar arrangement 
type I; (10) alary processes of premaxillae 
directed dorsally and slightly anteriorly, rel- 
atively narrow at base; (11) palatal shelf of 
premaxilla very narrow with elongate pal- 
atal process present [I could not find where 
Lynch defined what a palatal process of the 
maxilla was and I do not find such a struc- 
ture for either Thoropa or the Bahia frog]; 



VOLUME 112, NUMBER 1 



23 




T'tr: :■■:"' J 




W/ 




•'-i.'r.'. 



Fig. 4. Ventral view of skull of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Scale line = 1 mm. 



(14) maxillary arch complete, quadratojugal 
present; (15) nasals relatively large with 
moderately long maxillary processes, sep- 
arated medially; (16) nasals not in contact 
with maxillae or pterygoids; (22) epiotic 
eminences relatively long and narrow or 
short and stocky, carotid artery passes dor- 
sal to skull bones; (24) zygomatic ramus of 
squamosal relatively short; (25) otic ramus 
of squamosal moderately long, no otic 
plate; (26) squamosal-maxillary angle 50- 
70°; (27) columella present; (28) prevomers 
relatively small, entire, separated medially, 
toothed; (29) palatine long and narrow, ex- 
panded laterally, separated medially; (30) 
sphenethmoid entire, extending anteriorly 



to posterior edge of nasals or not reaching 
nasals; (31) anterior ramus of parasphenoid 
broad, keeled medially, extending anteriorly 
to prevomers; (32) parasphenoid alae ori- 
ented at right angles to anterior ramus of 
parasphenoid, relatively short, not over- 
lapped laterally by median ramus of pter- 
ygoids; (33) pterygoids large, anterior rami 
in long contact with maxillae, not reaching 
palatines; (34) occipital condyles large or 
small, not stalked, moderately to widely 
separated medially; (36) terminal phalanges 
T-shaped; (40) m. depressor mandibulae in 
two slips; (41) pupil horizontal; (42) males 
with median subgular vocal sac; (43) body 
lacking glands; (44) tongue large, oval, pos- 



24 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



terior edge free; (45) toes lacking webbing, 
bearing lateral fringes, digital tips bulbous, 
somewhat dilated, first finger shorter than 
second; (46) larvae with 2/3 tooth rows, la- 
bial papillae broadly interrupted anteriorly; 
(48) eggs large, few in number, laid in lotic 
situations; (49) males 19-78, females 24- 
70 mm SVL; (50) tympanum visible exter- 
nally; (51) tadpoles with greatly flattened 
and attenuate bodies and tails. 

There is no egg placement or larval in- 
formation for the Bahia frog. 

Several features require further comment. 
The Bahia frog does not have a medial keel 
of the parasphenoid, nor does the paras- 
phenoid extend anteriorly to the vomers 
(Fig. 4), as stated for character 31 in Tho- 
ropa. However, Lynch's Fig. 86 (1971:130) 
does not show the conditions he described. 
In re-examining the material used by 
Lynch, I find that in Thoropa miliaris (KU 
92855), the parasphenoid has a weak me- 
dial keel and does extend to the vomers, but 
in both T. lutzi (KU 92908), and T. petro- 
politana (KU 92862), the parasphenoid 
lacks a median keel and extends far short 
of the vomers (as in the Bahia frog). For 
character 32, in the Bahia frog the paras- 
phenoid is overlapped by the median rami 
of the pterygoids, contrasting with Lynch's 
statement for Thoropa. However, Lynch's 
Fig. 86 (1971:130) shows overlap, which I 
also find in T. lutzi (KU 92908), T. miliaris 
(KU 92855), and T. petropolitana (KU 
92862). The terminal phalanges of the Ba- 
hia frog, character 36, are not distinctly T- 
shaped as in Thoropa, but either expanded 
or weakly T-shaped (Fig. 5). Although the 
shape of the terminal phalanges in the Ba- 
hia frog differs from the distinct T-shaped 
condition in Thoropa, this condition could 
be interpreted as part of a morphological 
continuum in degree of expansion of the 
terminal phalanges. The depressor mandib- 
ulae muscle, character 40, of the Bahia frog 
is the DFSQat condition (three slips, the 
third a small slip originating from the tym- 
panic annulus), contrasting with the DFSQ 
state listed for Thoropa by Lynch. I find 



Ks a s 



Fig. 5. Terminal finger phalanges. Left to right, 
Thoropa lutzi, from cleared-and-stained specimen KU 
92908, third finger; Thoropa petropolitana, from 
cleared-and-stained specimen KU 92862, third finger; 
Rupirana cardosoi, from cleared-and-stained specimen 
MZUSP 68959, third finger; Rupirana cardosoi, from 
partially dissected wet specimen MZUSP 76035, 
fourth finger. Scale line = 1 mm. 

both the two and three slip conditions in 
Thoropa, however {T. miliaris, DFSQ 
[USNM 97765]; T petropolitana, DFSQat 
[USNM 164135]). 

There have been two additional species 
of Thoropa described since Lynch's (1971) 
publication {T. megatympanum, T saxatilis) 
which require modification of character 45 
in that genus. Thoropa lutzi and T petro- 
politana have weak lateral toe ridges that 
are less well-developed than in the Bahia 
frog. Thoropa miliaris has weak ridges, 
similar to the condition of the Bahia spe- 
cies. The toe ridges of T. megatympanum 
are even weaker than those of T. lutzi and 
T. petropolitana. Thoropa saxatilis lacks 
any toe ridge or fringe. The digital tips of 
the Bahia frog are narrow, not dilated. In 
both T. lutzi and T. saxatilis, the digit tips 
are dilated and the finger tips are more di- 
lated than the toe tips. In T. miliaris, the 
digital tips are bulbous and somewhat di- 
lated (Lynch's description), and there is no 
size difference between the finger and toe 
tips. In both T. megatympanum and T. pe- 
tropolitana, the tips of the digits are slightly 
dilated but not particularly bulbous and the 
finger and toe tips are equal sized. The first 
finger is the same length as the second in 
the Bahia species, the same condition that 
I find in T. miliaris (contra Lynch). The first 
finger is shorter than the second in T. lutzi, 
T. petropolitana, and T. saxatilis. In T. me- 
gatympanum, the first finger is either just 



VOLUME 112, NUMBER 1 



25 



shorter than the second or the first and sec- 
ond are of equal length. 

Summarizing thus far, the Bahia frog is 
most similar to species in the genus Tho- 
ropa, but differs consistently by two char- 
acters used by Lynch (1971): shape of the 
terminal phalanges (but see discussion of 
character 36 above) and dilation of digit 
tips. 

In a preliminary analysis of relationships 
of leptodactylid genera, I (Heyer 1975) 
came to quite a different conclusion from 
Lynch (1971) regarding the relationships of 
Thoropa. I did not recognize formal taxo- 
nomic groupings such as subfamilies and 
tribes, but used informal grouping names, 
which are equivalent in scope to Lynch's 
subfamilies. Specifically, my concept of a 
grypiscine clade combined the content of 
Lynch's Grypiscinae plus Elosiinae with the 
genera Paratelmatobius and Thoropa. The 
reasons for this very different viewpoint are 
due to two factors: although Lynch and I 
analyzed most of the same characters, there 
were some differences; and I attempted a 
phylogenetic analysis, whereas Lynch's 
analysis was phenetic. 

I analyzed 37 characters. For the states 
that I Usted for Thoropa (1975, table B, p. 
51), the following differ from the condi- 
tions found in the Bahia species. Character 
5, state B, disks on toes (see discussion of 
Lynch's character 45 above). Character 6, 
state A, no tarsal fold, flap, or tubercle. The 
Bahia frog has a tarsal fold, however, re- 
examination of T. miliaris indicates that 
specimens have weak tarsal folds as do 
specimens of T. saxatilis. Character 9, state 
D, toes with lateral fringe (see discussion 
of Lynch's character 45 above). Character 
12, state B, depressor mandibulae origin 
from dorsal fascia, squamosal, and otic re- 
gion only (see discussion of Lynch's char- 
acter 40 above). Character 16, state B, omo- 
hyoideus insertion on hyoid body and fascia 
between posteromedial and posterolateral 
processes. In the Bahia species, the omo- 
hyoideus inserts only on the hyoid plate. 
Character 17, state B, iliacus extemus ex- 




Fig. 6. Hyoid apparatus of Rupirana cardosoi, 
MZUSP 68959. Cartilage stippled. Scale line = 1 mm. 



tends Vi—Va- anterior on ilium. In the Bahia 
species, the extension is >Ya, about 90%. 
Character 20, state B, adductor longus 
poorly developed, inserting on adductor 
magnus, covered by sartorius. In the Bahia 
frog, the adductor longus is well-developed, 
inserting on the knee. Character 30, state C, 
anterior process of hyale absent. In re-ex- 
amining material, I found that the process 
is clearly absent in T. miliaris (USNM 
97765). In T. petropolitana (USNM 
164135), there is a medial, but not an an- 
terior swelling of the hyale at its most an- 
terior extent, contrasting markedly with the 
well-developed anterior process found in 
the Bahia species (Fig. 6). Character 34, 
state A, sacral diapophyses expanded. In 
the Bahia frog, the sacral diapophyses are 
flattened, but not expanded (Fig. 1). In re- 
examining the two Thoropa species that are 
similar in size to the Bahia species {T. lutzi, 
KU 92908, T petropolitana, KU (92862), 
the sacral diapophyses are flat, as in the Ba- 
hia species, and differ slightly from the Ba- 
hia species in being expanded. Character 
35, state B, terminal phalanges T-shaped is 
the same as Lynch's character 36 (see dis- 
cussion above). 

The Bahia species clearly shares most 
features of all known leptodactylid genera 



26 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



with the genus Thoropa. The question is 
whether the definition of Thoropa should be 
expanded to include the Bahia species, or 
whether the Bahia frog should be treated as 
a genus distinct from Thoropa. 

The Bahia species differs from all known 
Thoropa in seven characters (of the char- 
acters examined for this study). For two of 
these, the definition of the genus Thoropa 
would only have to be slightly modified to 
incorporate the Bahia frog: degree of ex- 
pansion of the flattened sacral diapophyses 
and shape of the terminal phalanges. The 
other five characters would require a more 
drastic redefinition of Thoropa: toe disks; 
insertion of the omohyoideus muscle; 
length of the ihacus extemus muscle; de- 
velopment of the adductor longus muscle; 
anterior process of the hyale. These latter 
five characters demonstrate different con- 
ditions involving toe, hyoid, and thigh mor- 
phologies. These kinds of differences are 
consistent with the Bahia species being on 
a separate evolutionary track from Thoro- 
pa. Thus, if the Bahia form were described 
as a species of Thoropa, evolutionary re- 
lationships would be obscured (also see 
section on relationships, below). Therefore, 
in order to emphasize the evolutionary dis- 
tinctiveness of the Bahia species, it is de- 
scribed as: 

Rupirana, new genus 
Figs. 1-6 

Type-species. — Rupirana cardosoi, new 
species. 

Diagnosis. — The only leptodactyHd gen- 
era that share the combination of sternum 
cartilaginous, widely spaced cervical coty- 
les with two distinct articular surfaces, long 
transverse processes of posterior presacral 
vertebrae, flattened sacral diapophyses, and 
an exposed frontoparietal fontanelle are Ba- 
trachyla, Thoropa, and Rupirana. Batra- 
chyla has an incomplete maxillary arch; in 
Rupirana the maxillary arch is complete. 
Thoropa has well-developed T-shaped ter- 
minal phalanges, dilated digit tips, and 



lacks an anterior process of the hyale; Ru- 
pirana does not have well-developed T- 
shaped terminal phalanges, has narrow dig- 
ital tips, and has an anterior process of the 
hyale. 

Definition. — Pupil horizontal; tympanum 
distinct; vocal sac single, subgular, slightly 
expanded externally; male thumb with one 
extensive and one small patch of kerati- 
nized sandpaper-like asperities; body with- 
out well-defined glands; digital tips narrow, 
not dilated; tarsus with a tarsal fold; outer 
metatarsal tubercle small; inner metatarsal 
tubercle large, ovoid, not comified; toes 
with lateral fringes, joined at base of toes. 

Adductor mandibularis muscle condition 
adductor mandibulae posterior subextemus 
only; depressor mandibulae condition 
DFSQat; geniohyoideus muscle contiguous 
medially; anterior petrohyoideus insertion 
on lateral edge of alary process and hyoid 
plate; stemohyoideus muscle insertion en- 
tirely near edge of hyoid body; omohyoi- 
deus muscle insertion entirely on hyoid 
plate; iliacus extemus muscle extending al- 
most to anterior tip of ihum; tensor fasciae 
latae muscle insertion posterior to iliacus 
extemus muscle on iliac bone; interior and 
exterior portions of semitendinosus muscle 
uniting in a common distal tendon distally, 
exterior portion larger than interior; adduc- 
tor longus muscle about same size and 
shape as sartorius muscle, inserting on knee 
and adductor magnus. 

Quadratojugal present, contacting maxil- 
la; frontoparietals not meeting medially, ex- 
posing moderate-sized fontanelle; vomerine 
teeth present; occipital condyles widely 
separated; anterior process of hyale present; 
alary processes of hyale on narrow stalks; 
posterior sternum a cartilaginous plate, ter- 
minally expanded and bifid; last presacral 
vertebra just narrower than sacrum; sacral 
diapophyses flattened, not noticeably ex- 
panded; dorsal crest of ilium present; ter- 
minal phalanges barely expanded, not or 
weakly T-shaped. 

Etymology. — From the Latin rupes, rock, 
and rana, frog. The gender is feminine. The 



VOLUME 112, NUMBER 1 



27 



• A.*-:: 



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m 



I ^:J;V~ • • fi-r**^ ' 



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Fig. 7. Dorsal view of male paratype of Rupirana cardosoi, MZUSP 76032. 



name is to highlight the association of this 
genus with the campos rupestres of Brazil. 
Content. — Monoty pic . 

Rupirana cardosoi, new species 
Figs. 7-8 

Holotype.—MZ\]SV 65203, adult male, 
27 Sep 1987, Mucuje, Bahia, Brazil, 
13°00'S, 41°23'W, Miguel T. Rodrigrues. 

Paratopotypes. — MZUSP 65202 (adult 
male), 65204 (aduh male), 65205 (juve- 
nile), same data as holotype; MZUSP 

68959 (cleared and stained adult female), 

68960 (juvenile), 2 Oct 1990, Miguel T. 
Rodrigues. 

Paratypes.—MZUS^ 68961-68962 
(adult females), Morro do Chapeu, Bahia, 



Brazil, 3 Oct 1990, 11°33'S, 41°09'W, Mi- 
guel T Rodrigues; MZUSP 76017-76018 
(aduh males), 76023-76030 (aduh males), 
76031 (aduh female), 76032-76034 (aduh 
males), 76035 (adult female, skull re- 
moved), 76036 (adult male), 76037 (juve- 
nile male), 76038-76040 (aduh males), 
USNM 519755-519757 (adult males), 
519758 (adult female), Andarai, Bahia, 
Brazil, 12°48'S, 4r20'W, 19-22 Nov 1968, 
Werner C. A. Bokermann, Francisco M. 
Oliveira, and B. D. Silva. 

Diagnosis. — As for genus. 

Description of holotype. — Snout round in 
profile and from above; nostrils anterolat- 
eral, near tip of snout; can thus rostralis in- 
distinct; loreal obtuse; tympanum distinct, 



28 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



C. ^.«o^ 




• '. •' •■"'"'■-.'. '"• :*V.^V•vV^;;>'•*Wy"'5V/. V;'V/.v. o 









Fig. 8. Profile of head of holotype of Rupirana cardosoi, MZUSP 65203. 



rounded, about Vi diameter of eye; supra- 
tympanic fold distinct from behind eye to 
shoulder, bordering tympanum dorsally; 
tongue elongate, triangular, with slight 
emargination on anterior edge; vomerine 
teeth in two small transverse patches in line 
with posterior borders of small, round cho- 
anae, vomerine tooth patches separated 
from each other by about width of single 
tooth patch; vocal slits elongate; vocal sac 
single, subgular, indicated externally by lat- 
eral skin folds/wrinkles; finger lengths 
I~II~IV<III; fingers free of web, very 
slight lateral ridges; tips of fingers rounded, 
not expanded; palmar tubercle ovoid, just 
smaller than ovoid thenar tubercle; subar- 
ticular tubercles moderately developed, 
slightly pungent; no supernumerary tuber- 
cles; accessory palmar tubercles present, 
two in line with each digit; thumb with ex- 
tensive keratinized sandpaper-appearing as- 
perity from penultimate phalanx to base of 
thumb, a second ovoid patch on inner half 
of thenar tubercle; forearm slightly hyper- 
trophied; dorsum, including upper eyelid, 
smooth with many scattered large white tu- 
bercles; body lacking any obvious glands; 
venter smooth except for areolate poster- 
o ventral thigh surfaces; belly disk weakly 



indicated; toe lengths I<II<V<ni<IV; tips 
of toes rounded, not expanded; toes with 
weak lateral ridges expanded into weak 
fringes at base of inner sides of toes II and 
III resulting in trace of strap-shaped basal 
web between toes I, II, and III; inner meta- 
tarsal tubercle small, round, about V3 size of 
small, ovoid outer metatarsal tubercle; dis- 
tinct but relatively weak tarsal fold extend- 
ing about V2 length of tarsus; sole of foot 
smooth; subarticular tubercles moderate, 
pungent. 

SVL 27.3 mm; head length 9.8 mm, head 
width 10.3 mm; horizontal diameter of tym- 
panum 2.1 mm, including annulus; distance 
from eye to posterior edge of naris 2.8 mm; 
intemarial distance 2.5 mm; thigh length 
12.5 mm; shank length 13.4 nmi; foot 
length 14.3 mm. 

In preservative, dorsal body brown with 
darker brown markings, upper limbs tan 
with brown markings; upper snout with a 
few irregular darker blotches and dots; ir- 
regular interorbital bar connecting with li- 
chenous network behind eyes, rest of dor- 
sum with mostly elongate blotches and 
broad stripes; upper limbs weakly cross- 
banded; distinct broad dark brown canthal 
stripe extending more faintly from nostril to 



VOLUME 112, NUMBER 1 



29 



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tip of snout (Fig. 8); upper lip mostly uni- 
form light tan; supratympanic fold dark 
brown, expanded into dark triangular blotch 
posteriorly followed by series of small dark 
spots on anterior half of otherwise light tan 
flanks; throat, chest, and belly light tan with 
lighter spots lacking melanophores; sole of 
foot brown; posterior surfaces of thighs uni- 
form tan. 

Variation. — Available data do not indi- 
cate any marked variation in measurements 
among sites, with a suggestion of slight 
sexual dimorphism in size (Table 1). Mea- 
surement variation of the entire sample of 
adults (male n = 23, female n = 5) is 
(means in parentheses): SVL 27.3-31.2 
(29.6) mm for males, 30.2-34.4 (32.0) mm 
for females; % head length/SVL 34-40 
(37.3) for males, 32-39 (35.0) for females; 
% head width/SVL 36-40 (37.9) for males, 
34-39 (37.0) for females; % tympanum di- 
ameter/SVL 7-9 (7.7) for males, 6-8 (7.2) 
for females; % eye-nostril distance/SVL 9- 
11 (10.0) for males, 9-12 (9.8) for females; 
% intemarial distance/SVL 8-10 (9.1) for 
males, 8-10 (8.8) for females; % thigh 
length/SVL 44-48 (45.7) for males, 42-44 
(43.2) for females; % shank length/SVL 
44-51 (48.2) for males, 44-47 (45.8) for 
females; % foot length/SVL 47-54 (51.0) 
for males, 45-49 (47.0) for females. The 
sample size for females is small, but it does 
appear as though the female legs are pro- 
portionately shorter than in males. 

The most striking variation in addition to 
the male secondary sexual characteristics 
involving vocal slits, vocal sacs, thumb as- 
perities, and forearm hypertrophy is the 
sexual variation in dorsal texture. In fe- 
males, the dorsum is either smooth or with 
a weak shagreen, whereas the males have 
many, large, white tubercles scattered pro- 
fusely over the back (Fig. 9). In the speci- 
mens from Monro do Chapeu and Mucuje, 
the male dorsal tubercles are largely limited 
to the upper eyelids and the back. The 
males from Andarai have a much more ex- 
tensive distribution of tubercles, including 
the snout, outer arms, dorsal thigh surfaces, 




dorsal shank surfaces, outer tarsus, and var- 
iably, the sole of the foot. 

Other aspects of variation noted among 
the specimens (males and females, except 
as noted) include: tongue shape also ovoid 
or round; vomerine tooth placement be- 
tween to just posterior to choanae, separat- 
ed from each other by <% distance of 
length of a single tooth patch; finger lengths 
also n just <IV just <I<in; some males 
with no external indication of a vocal sac; 
finger subarticular tubercles sometimes 
pungent; outer tarsus sometimes shagree- 
ned; tarsal fold occasionally extending % 
distance of tarsus; the inner metatarsal tu- 
bercle sometimes elevated; the sole of the 
foot sometimes with series of small, fleshy 
tubercles in line with digits; the dark supra- 
tympanic stripe sometimes extending con- 
tinuously to mid-flank; front of tympanum 
sometimes bordered by dark brown spur ex- 
tending from dark supratympanic stripe; 
dorsum relatively uniform to longitudinally 



striped; chin through the chest sometimes 
scattered uniformly with melanophores 
with anterior belly with mottled pattern and 
posterior belly lacking melanophores to the 
entire region from chin through belly Hght- 
ly brown mottled. 

MZUSP 76037, a 29.4 mm SVL speci- 
men, is a juvenile male, although not the 
smallest male in the sample. The specimen 
does not have vocal sHts, but it has a small 
thumb patch asperity (on the thumb only, 
not on the thenar mbercle) and no dorsal 
mbercles. MZUSP 76039, a 30.2 mm SVL 
male, has vocal sHts that are in early de- 
velopment, but has fuUy developed nuptial 
pads and dorsal tubercles. 

The cleared-and-stained female (MZUSP 
68959) contained mamre-appearing ova, of 
varying sizes up to about 1.5 mm diameter 
with melanophores on the animal pole. 

Etymology. — Named in honor of Dr. 
Adao Jose Cardoso, a colleague whose trag- 
ic death is deeply felt both at a personal 



VOLUME 112, NUMBER 1 



31 



level and as a premature halt to his signif- 
icant contributions to our understanding of 
the Neotropical amphibian fauna. 

Distribution and habitat. — ^At present, 
Rupirana cardosoi is known from three lo- 
calities in the northern portion of the Es- 
pinha90 Range in the State of Bahia, Brazil 
(Fig. 10). 

Dr. Rodrigues kindly provided the fol- 
lowing habitat information. MZUSP 
65202-65205 were obtained during the day 
on the bank of a small stream with white 
sand and rocks. The stream crosses the city 
of Mucuje. The water of the stream was 
very clear but with a red-brown tint (as usu- 
al in the Espinha90 Range). The vegetation 
was typical of quartzitic campos rupestres: 
dominated by Velloziaceae, Euriocaulla- 
ceae, Xiridaceae and other endemics. There 
were many large and highly eroded quartz- 
itic outcrops. MZUSP 68959-68960 were 
collected during the day at a similar stream 
habitat (2-3 m wide) less than 2 km from 
Mucuje. The general habitat was the same 
as for the Mucuje stream. Both of these 
sites were also visited at night, but no Ru- 
pirana cardosoi were found. MZUSP 
68961-68962 were collected in a relictual 
patch of white sands on a large red rocky 
mountain not far from Morro de Chapeu. 
The frogs were on the margins of two 
small, drying ponds. The vegetation, dom- 
inated by Mellastomataceae on the sand, 
was typical for campos rupestres. Several 
specimens of the sand-adapted Tropidurus 
cocorobensis were also collected at the 
same time, around 1500 h. 

Relationships 

Rupirana shares the most character states 
with the genus Thoropa, as documented in 
the section, "Comparison with leptodactyl- 
id genera." The evaluation of character 
states in that section did not differentiate 
between shared primitive and shared de- 
rived states, however. If most of the states 
shared by Rupirana and Thoropa are shared 



primitive states, then there would be little 
support for a close sister- group relationship. 

In order to undertake a first approxima- 
tion of the relationships of Rupirana within 
leptodactylid frogs, the data assembled for 
an earlier study (Heyer 1975) are used as a 
basis for a cladistic analysis (Appendix 1). 
As indicated above, the relationships of 
Thoropa within the Leptodactylidae have 
been disputed by Lynch (1971, 1978) and 
me (Heyer 1975). In order to evaluate the 
relationships of Rupirana relative to Tho- 
ropa, the genus Batrachyla is included to 
evaluate Lynch's position that Batrachyla 
and Thoropa are sister-taxa (Lynch 1978), 
and the genera Cycloramphus and Megae- 
losia are included to evaluate my position 
that Thoropa is a member of a grypiscine 
clade (Heyer 1975). Three additional rep- 
resentatives of telmatobiine genera are in- 
cluded to provide structure to test the alter- 
native hypotheses: Eleutherodactylus, Eup- 
sophus, and Hylorina. Three genera are 
used as outgroups: Ceratophrys as a rep- 
resentative of the South American cerato- 
phryine clade and a representative each of 
two subfamilies of the Australian Family 
Myobatrachidae, Adelotus and Crinia. In all 
cases except for Eleutherodactylus, the 
character states analyzed are for genera. As 
Eleutherodactylus is currently in the pro- 
cess of being broken into smaller mono- 
phyletic units, with the final configuration 
of this effort far from clear, data for a single 
species, E. coqui, are used for purposes of 
this analysis. 

The data were analyzed with as much or- 
dering of character states as unambiguous 
morphoseries would allow (Appendix 1). 

Phyletic signal in the data set. — The 
strength of the phyletic signal in the data 
set is evaluated with two tests: the indirect 
^1 statistic (Hillis 1991) and the more direct 
Permutation Tail-Probabilities (PTP) test 
(Faith and Cranston 1991). 

Hillis (1991) explained the rationale for 
the g, statistic and provided probability lev- 
els for cases using 6, 7, or 8 taxa. As there 
is no graphable correlation between the 



32 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



45 



42 






.39 



-10 



I 



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BAHIA 



Jk 



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MINAS 
?? i->GERAIS 



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r>^y 




Fig. 10. The Espinha^o Range in the States of Bahia and Minas Gerais. StippHng indicates areas above 1000 
m. Dot is the type locality of Rupirana cardosoi; triangles indicate the other two known localities for R. cardosoi. 
Map redrawn from Map 56, page 398 in Giuletti et al. 1997. 



VOLUME 112. NUMBER 1 



33 



number of taxa and g, critical values, ex- 
haustive PAUP analyses were run using 8 
taxa with Adelotus or Crinia or Cerato- 
phrys used as sole outgroup taxa and delet- 
ing Hylorina. The ^, statistics for the 8 taxa 
are 0.31, 0.29, and 0.01 respectively when 
Rupirana is coded as not having T-shaped 
phalanges and 0.17, 0.39, and 0.08 respec- 
tively when Rupirana is coded as having T- 
shaped phalanges. The critical g^ values for 
8 taxa for P of 0.05 is -0.34 and P of 0.01 
is —0.47. None of the g, statistic values ob- 
tained in the six analyses approaches those 
critical values. The distribution of trees is 
not skewed. 

The PTP test was run with Rupirana cod- 
ed as not having T-shaped phalanges. The 
P value is 0.06, which is not significant at 
the traditional 0.05 level, but indicative that 
there is likely some phylogenetic content in 
the data. 

Clearly, the phyletic signal in this data 
set is not strong and the results must be 
interpreted extremely conservatively. 

Cladistic relationships. — The results of 
PAUP analyses run on the entire data set 
with a branch-and-bound search or with the 
outgroup taxa run individually with ex- 
haustive searches (using the data matrix in 
Appendix 1) all have the same outcome: 
multiple numbers of shortest trees, for 
which the strict consensus tree is an entirely 
unresolved polytomy. The branch-and- 
bound search of the entire data set yields 
19 shortest trees of tree length 112, with a 
consistency index (CI) excluding uninfor- 
mative characters of 0.52. The exhaustive 
search of the data set using Adelotus as the 
sole outgroup taxon gives 240 shortest trees 
of tree length 86, with a CI value excluding 
uninformative characters of 0.56. Similar 
results for Crinia as the outgroup are 45 
trees of length of 91, CI = 0.54 and for 
Ceratophrys as the outgroup are 45 trees of 
length 89, CI = 0.56. 

In case the coding of the shape of the 
terminal phalanges in Rupirana might be 
pivotal, exhaustive searches were also run 
using Adelotus, Crinia, and Ceratophrys in- 



dividually as outgroup taxa with Rupirana 
coded as having T-shaped terminal phalan- 
ges. Again the analyses resulted in multiple 
shortest trees with the strict consensus tree 
of each being an entirely unresolved poly- 
tomy. The results for using Adelotus as the 
outgroup are 240 trees of shortest length 85, 
CI (excluding uninformative characters) = 
0.57; for Crinia, 150 trees of shortest length 
91, CI = 0.55; for Ceratophrys, 45 trees of 
shortest length 88, CI = 0.56. 

These results certainly do not resolve any 
questions about the relationships of Tho- 
ropa within the Family Leptodactylidae. 
However, there is one result that is so con- 
sistent that a statement about relationships 
can be made: Rupirana and Thoropa do not 
share a close relationship. In fact, Rupirana 
and Thoropa never form a sister-group re- 
lationship in any trees, and Thoropa either 
demonstrates a basal relationship with all or 
most other taxa or (usually) forms a sister- 
group relationship with Batrachyla (e.g., 17 
out of 19 trees in the trees based on the 
entire data set). This Batrachyla-Thoropa 
sister-group relationship, repeated in most, 
but not all trees, lends support for Lynch 's 
proposal of relationships. However, the po- 
tential problem of long-branch attraction 
(Swofford et al. 1996:427) would have to 
be ruled out to support the Batrachyla-Tho- 
ropa relationship. 

The results of the above analyses indicate 
that the similarities analyzed between Ru- 
pirana and Thoropa are due to shared prim- 
itive state conditions, not shared derived 
states, and that there is no close sister-group 
relationship between the two genera. 

Zoogeography 

The campos rupestres contain a notably 
endemic biota at the species level (for a bo- 
tanical introduction, see Giuletti & Pirani 
1988), including amphibians and at least 
some reptiles (e.g., Vanzolini 1982, Ro- 
drigues 1988). As I have extremely limited 
experience with the campos rupestres, the 
following comments are very superficial. 



34 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Hopefully these comments will encourage 
an honest zoogeographical study of the am- 
phibians of the campos rupestres, because I 
think the amphibians show a pattern that 
differs from those based on study of other 
groups. 

The campos rupestres occur above 800- 
900 m and, as the common name indicates, 
are rocky places with poor soils (see Giu- 
letti & Pirani 1988 and Giuletti et al. 1997 
for better characterizations and maps). 
Plants have been the most throroughly stud- 
ied campos rupestres group (Giuletti & Pir- 
ani 1988), otherwise, there has been little 
research on the distributions of other groups 
of organisms found in the campos rupestres 
with the exception of certain lizards (for 
Tropidurus see Rodrigues 1987, 1988, for 
other lizards see Vanzolini 1982). Most of 
the plant and lizard species found in the 
campos rupestres are either endemic to 
them or show their major central distribu- 
tions on them. Species such as Polychrus 
acutirostris and Tropidurus hispidus, which 
occur in both the campos rupestres and 
more broadly beyond them, seem to be ex- 
ceptional cases. Even though the endemic- 
ity at the species level is rather astounding, 
there are few endemic genera of plants 
(Giuletti & Pirani 1988). The affinities for 
the campos rupestres endemic species are 
primarily with cerrado and caatinga species 
(for plants and Tropidurus), and cerrados 
and caatingas likely served as the primary 
source of species which differentiated in the 
campos rupestres environment. Whereas the 
campos rupestres could be considered a part 
of the diagonal of open formations (caatin- 
gas-cerrados-Gran Chaco), first noticed by 
K. R Schmidt (e.g., Schmidt & Inger 1951), 
the campos rupestres do not fit comfortably 
within the diagonal from a zoogeographic 
perspective. Another common affinity for 
both plants and Tropidurus is with the res- 
tingas (sand beaches) scattered along the 
east coast of Brazil. In this case, the restin- 
ga associated elements are thought to be de- 
rived from campos rupestres ancestors 
(Giuletti & Pirani 1988, Rodrigues 1988). 



The plants also show affinities that the cam- 
pos rupestres lizards do not, i.e., with the 
mountain biotas of northern South America 
(stronger) and the Andes (weaker) (Giuletti 
& Pirani 1988, Harley 1988). There is a mi- 
nor affinity with Atlantic Forest plants and 
lizards; the plant genus Pleurostima (Velo- 
ziaceae) occurs on the tops of some granitic 
outcrops in the Atlantic Forests, and the At- 
lantic Forest lizard genera Enyalius, Pla- 
cosoma, and perhaps Heterodactylus have 
representatives in the campos rupestres. 

Most of our knowledge of campos ru- 
pestres amphibians comes from the work of 
Werner C. A. Bokermann and Ivan Sazima, 
who pubhshed several papers describing 
new species of amphibians from the Serra 
do Cipo, southern Espinhago Range, in the 
State of Minas Gerais. Many of the new 
species described from Serra do Cipo be- 
long to well delineated species groups or 
genera for which the general distributions 
are known. For most of these, the Serra do 
Cipo endemics are related to Atlantic Forest 
groups from which they must have been de- 
rived. For example all species of the genera 
Hylodes, Phasmahyla, and Thoropa and the 
Scinax catherinae species group occur only 
in the Atlantic Forests except for disjunct 
endemic species at Serra do Cipo {H. ota- 
vioi, P. jandaia, T. megatympanum, and S. 
machadoi). Determining the faunistic origin 
of several other species from the Serra do 
Cipo is not as clear, but it would seem that 
the majority of them were derived from At- 
lantic Forest lineages and a smaller number 
from cerrado groups. Some of the non-en- 
demic species that occur at Serra do Cipo 
are likely cerrado species, such as Lepto- 
dactylus fumarius, L. jolyi, and Physalae- 
mus cuvieri. Some of the Serra do Cipo en- 
demics may also have been derived from 
cerrado ancestors, for example Leptodacty- 
lus camaquara, L. cunicularius, Physalae- 
mus evangelistai and Proceratophrys cu- 
ruru. The frog fauna of the campos rupes- 
tres thus has a strong faunal element de- 
rived from Atlantic Forest ancestors and 



VOLUME 112, NUMBER 1 



35 



this pattern is different from that found in 
plants or Uzards. 

Remnants of the Atlantic Forest vegeta- 
tion occur today in the Espinha90 Range 
(e.g., Pico das Almas, Harley 1995:20-24). 
The Atlantic Forests certainly were more 
extensive in the Espinhago Range and con- 
tinuous with the coastal forests during more 
mesic times. The campos rupestres, a very 
open formation habitat, would be expected 
to show biotic affinities with adjacent open 
formation habitats, as is true for plants and 
lizards. The campos rupestres would not be 
expected to show a primary affinity with 
the Atlantic Forest biota, as the basic ad- 
aptations for those two environments are 
profoundly different. 

One aspect of anuran ecogeography 
could explain why many campos rupestres 
frogs were derived from the Atlantic Forest 
frog fauna rather than entirely from the cer- 
rado/caatinga frog fauna. There are small 
streams (mostly seasonal at this stage of the 
interglacial) in the campos rupestres land- 
scape. There are, as far as I know, no strict- 
ly stream-adapted frogs in the cerrados or 
caatingas, whereas many of the Atlantic 
Forest frogs are stream- adapted. Thus, the 
most probable source of stream frogs in the 
campos rupestres would come from the At- 
lantic Forests rather than the cerrados or 
caatingas. If, during periods of drier cli- 
mates, the Atlantic Forests underwent grad- 
ual change, first contracting to gallery for- 
ests and then degrading to even more open 
vegetation, the stream frogs may have been 
able to adapt to the more open habitat con- 
ditions and occupy the campos rupestres. 

In summary, from the available data, it 
would seem that the unique campos rupes- 
tres frog fauna has been derived from only 
two sources: Atlantic Forest (strongest); 
and cerrado/caatinga (the conclusion of 
Heyer 1988, that present data are inade- 
quate to determine whether the caatingas 
harbor a frog fauna distinct from that of the 
cerrados is unfortunately still true). 

The discovery of Rupirana cardosoi has 
one expected and one unusual zoogeo- 



graphical consequence. The Espinha^o 
Range is comprised of a series of individual 
isolated mountains (Fig. 10), with many 
species narrowly endemic to single moun- 
tain systems within the entire Range (Giu- 
letti & Pirani 1988). Finding a new species 
in the Chapada Diamantina, which is still 
poorly sampled for its frog fauna, is to be 
expected. What is unusual is that the new 
species also represents a new genus. 

The fact that all but one of the endemic 
campos rupestres frogs are endemic only at 
the species level suggests that the differ- 
entiation of the campos rupestres biota oc- 
curred in a single episode and relatively re- 
cently in terms of geological time. Discov- 
ery of an endemic campos rupestres genus 
of frogs requires that the single episode of 
faunal differentiation requires emendment 
or explanation. There seem to be three most 
likely alternatives. 

Rupirana is an endemic campos rupestres 
genus of frog. If this is true, that suggests 
that it had an earlier episode of differenti- 
ation than other campos rupestres endemic 
frogs. Although this alternative requires an 
additional episode of differentiation for 
frogs, this is not unusual for the rest of the 
campos rupestres biota, as there are endem- 
ic genera of campos rupestres plants (e.g., 
Barbacenia (Velloziaceae), Morithamnus 
(Compositae), Pseudotrimezia (Iridaceae), 
and Raylea (Sterculiaceae), Giuletti and 
Pirani 1988) and fishes {Copionodon and 
Glaphryopoma (Trichomycteridae), Pinna 
1992). 

Rupirana represents a sampling problem. 
It could be that Rupirana had or has a dis- 
tribution with other species occurring in the 
Atlantic Forests (I think this more likely 
than in the cerrados/caatingas). If this is 
true, then either the Atlantic Forest species 
of the genus have become extinct and/or 
they are still extant but have not been 
found. The frog fauna of the northern At- 
lantic Forests is not well known and I 
would not be surprised if additional species 
of Rupirana were found there. 

Recognition of Rupirana as a distinct ge- 



36 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



nus is a taxonomic error. If this is the case, 
the new species does not require an histor- 
ical zoogeographical explanation different 
from that for other campos rupestres en- 
demic frogs. 

Acknowledgments 

Drs. Miguel T. Rodrigues (MZUSP), P. E. 
Vanzolini (MZUSP), and George R. Zug 
(USNM) critically reviewed the manuscript. 
Drs. Rodrigues and Vanzolini steered me to 
the pertinent literature and made substan- 
tive contributions to my zoogeographic un- 
derstanding of the biota of the Espinhago 
Range. Steve Gotte (Biological Resources 
Division, U.S. Geological Service stationed 
at the Smithsonian Institution) made the 
cleared-and-stained preparation of Rupi- 
rana cardosoi. 

Research for this paper was facilitated by 
the Museu de Zoologia da Universidade de 
Sao Paulo, especially from Dr. P. E. Van- 
zolini. Additional support was provided by 
the Neotropical Lowland Research Pro- 
gram, Smithsonian Institution, Dr. Richard 
P. Vari, Principal Investigator. 

Figures 7, 8, and 9 were rendered by Re- 
bekah St. John. 

Literature Cited 

Dingerkus, G., & L. D. Uhler. 1977. Enzyme clearing 
of alcian blue stained whole small vertebrates 
for demonstration of cartilage. — Stain Technol- 
ogy 52:229-232. 

Duellman, W. E., & L. Trueb. 1986. Biology of am- 
phibians. McGraw-Hill Book Company, New 
York, 670 pp. 

Faith, D. P., & P. S. Cranston. 1991. Could a cladogram 
this short have arisen by chance alone?: on per- 
mutation tests for cladistic structure. — Cladis- 
tics 7:1-28. 

Frost, D. R. (ed.). 1985. Amphibian species of the 
world: A taxonomic and geographical reference. 
Allen Press, Inc. and The Association of Sys- 
tematics Collections, Lawrence, Kansas, 732 
pp. 

Giuletti, A. M., & J. R. Pirani. 1988. Patterns of geo- 
graphic distribution of some plant species from 
the Espinhago Range, Minas Gerais and Bahia, 
Brazil. Pp. 39-69 in R E. Vanzolini & W. R. 
Heyer, eds.. Proceedings of a workshop on Neo- 



tropical distribution patterns held 12-16 Janu- 
ary 1987. Academia Brasileira de Ciencias, Rio 
de Janeiro, 488 pp. 

, , & R. M. Harley. 1997. Espinha90 

range region: Eastern Brazil. Pp. 397-404 in S. 
D. Davis, V. H. Heywood, O. Herrera-Mac- 
Bryde, J. Villa-Lobos, & A. C. Hamilton, eds.. 
Centres of plant diversity. A guide and strategy 
for their conservation. Volume 3. The Ameri- 
cas. The World Wide Fund for Nature and 
lUCN-The World Conservation Union, Cam- 
bridge, UK, 562 + xiv pp. 

Harley, R. M. 1988. Evolution and distribution of Er- 
iope (Labiatae), and its relatives, in Brazil. Pp. 
71-120 in P E. VanzoHni & W. R. Heyer, eds.. 
Proceedings of a workshop on Neotropical dis- 
tribution patterns held 12-16 January 1987. Ac- 
ademia Brasileira de Ciencias, Rio de Janeiro, 
488 pp. 

. 1995. Introduction. Pp. 1-40 in B. L. Stan- 

nard, ed.. Flora of the Pico das Almas, Chapada 
Diamantina — Bahia, Brazil. Royal Botanic Gar- 
dens, Kew, 853 pp. 

Heyer, W. R. 1975. A preliminary analysis of the in- 
tergeneric relationships of the frog family Lep- 
todactylidae. — Smithsonian Contributions to 
Zoology 199:1-55. 

. 1988. On frog distribution patterns east of the 

Andes. Pp. 245-273 in P E. Vanzolini & W. R. 
Heyer, eds.. Proceedings of a workshop on Neo- 
tropical distribution patterns held 12-16 Janu- 
ary 1987. Academia Brasileira de Ciencias, Rio 
de Janeiro, 488 pp. 

, A. S. Rand, C. A. G. Cruz, O. L. Peixoto, & 



C. E. Nelson. 1990. Frogs of Boraceia. — Ar- 
quivos de Zoologia 31:231-410. 

Hillis, D. M. 1991. Discriminating between phyloge- 
netic signal and random noise in DNA sequenc- 
es. Pp. 278-294 in M. M. Miyamoto & J. Cra- 
craft, eds., Phylogenetic analysis of DNA se- 
quences. Oxford University Press, Inc., New 
York, 358 pp. 

Lynch, J. D. 1971. Evolutionary relationships, osteol- 
ogy, and zoogeography of leptodactyloid 
frogs. — University of Kansas Museum of Nat- 
ural History Miscellaneous Publication 53:1- 
238. 

. 1978. A re-assessment of the telmatobiine lep- 

todactylid frogs of Patagonia. — University of 
Kansas Museum of Natural History Occasional 
Papers 72:1-57. 

Pinna, M. C. C. de. 1992. A new subfamily of Tricho- 
mycteridae (Teleostei, Siluriformes), lower lor- 
icaroid relationships and a discussion on the im- 
pact of additional taxa for phylogenetic analy- 
sis. — ^Zoological Journal of the Linnean Society 
106:175-229. 

Rodrigues, M. T 1987. Sistematica, ecologia e zo- 



VOLUME 112, NUMBER 1 



37 



ogeografia dos Tropidurus do grupo Torquatus 
ao sul do Rio Amazonas (Sauria, Iguanidae). — 
Arquivos de Zoologia 31:105-230. 
-. 1988. Distribution of lizards of the genus Tro- 



pidurus in Brazil (Sauria, Iguanidae). Pp. 305- 
315 in P. E. Vanzolini & W. R. Heyer, eds.. Pro- 
ceedings of a workshop on Neotropical distri- 
bution patterns held 12-16 January 1987. Ac- 
ademia Brasileira de Ciencias, Rio de Janeiro, 
488 pp. 

Schmidt, K. P, & R. E Inger. 1951. Amphibians and 
reptiles of the Hopkins-Branner Expedition to 
Brazil. — Fieldiana, Zoology 31:439-465. 

Swofford, D. L. 1993. PAUP: Phylogenetic Analysis 



Using Parsimony, Version 3.1. Formerly distrib- 
uted by Champaign, Illinois, Illinois Natural 
History Survey, 257 pp. 
-, G. J. Olsen, R J. Waddell, & D. M. Hillis. 



1996. Phylogenetic inference. Pp. 407-514 in 
D. M. Hillis, C. Moritz & B. K. Mable, eds.. 
Molecular systematics. Second edition. Sinauer 
Associates, Inc., Sunderland, Massachusetts, 
655 pp. 
Vanzolini, P. E. 1982. A new Gymnodactylus from Mi- 
nas Gerais, Brasil, with remarks on the genus, 
on the area and on montane endemisms in Bras- 
il (Sauria, Gekkonidae). — Papeis Avulsos de 
Zoologia 34:403-413. 



38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Appendix 1. — Character state descriptions and data matrix for phylogenetic analysis of selected genera of 
leptodactylid frogs. 

Data from Heyer (1975) supplemented by taxa examined for this study as indicated in body of text. 

Multistate characters ordered by criterion of morphoseries when reasonable. Character states not polarized. 

Character 1. Pupil shape. State — round; State 1 — horizontal. 

Character 2. Tympanum visibility. State — external, visible; State 1 — partially concealed; State 2 — concealed. 
Character state order 0-1-2. 

Character 3. Male thumb. State — no asperities; State 1 — pad; State 2 — spines. Character states not ordered. 

Character 4. Body glands. State — none; State 1 — paratoids; State 2 — inguinals. Character states not ordered. 

Character 5. Toe disks. State — none; State 1 — disks but no grooves; State 2 — disks with circumferential 
grooves; State 3 — disks with dorsal scutes. Character states not ordered. 

Character 6. Tarsal decoration. State — none; State 1 — fold; State 2 — flap. Character state order 0-1-2. 

Character 7. Toe webbing. State — web; State 1 — fringe; State 2 — ridge; State 3 — none. Character state order 
0-1-2-3. 

Character 8. Life history. State — tadpole with 2/3 denticle formula; State 1 — tadpole with >2/3 denticle 
formula; State 2 — tadpole with <2/3 denticle formula; State 3 — direct development. Character states ordered in 
a triangular relationship with the following connections: 2-0-1; 2-3; 0-3; 1-3. 

Character 9. Depressor mandibulae muscle. State — three slips (DFSQAT to dfsqat); State 1 — two slips 
(DFSQ to dfsq); State 2 — one slip (SQ). Character state order 0-1-2. 

Character 10. Stemohyoideus muscle insertion. State — on lateral edge of hyoid plate; State 1 — on lateral 
edge and midline of hyoid body; State 2 — on midline of hyoid body. Character state order 0-1-2. 

Character 11. Omohyoideus muscle insertion. State — muscle absent; State 1 — insertion on hyoid body and 
fascia between posteromedial and posterolateral processes; State 2 — insertion on hyoid body only; State 3 — 
insertion on hyoid body adjacent to posteromedial process. Character states unordered. 

Character 12. Iliacus extemus muscle. State — extends <% ilium length; State 1 — extends Vi-Ya ilium length; 
State 2 — extends %-full length of ilium. Character state order 0-1-2. 

Character 13. Tensor fasciae latae muscle insertion. State — posterior to anterior extent of iliacus extemus 
muscle; State 1 — at same level as anterior extent of iliacus extemus muscle. 

Character 14. Semitendinosus muscle. State — Extemal head smaller than intemal head, extemal head attached 
by tendon to intemal head; State 1 — as state 0, except intemal and extemal heads displaced from each other; 
State 2 — extemal head absent. Character states unordered. 

Character 15. Adductor longus muscle. State — well developed, insertion on or near knee; State 1 — poorly 
developed, insertion on adductor magnus muscle; State 2 — absent. Character state order 0-1-2. 

Character 16. Quadratojugal. State — present, contacting maxilla; State 1 — absent. 

Character 17. Nasal contact with maxilla. State — present; State 1 — absent. 

Character 18. Nasal contact with frontoparietal. State — absent; State 1 — present, not fused; State 2 — present, 
fused. Character states unordered. 

Character 19. Frontoparietal fontanelle exposed. State — not exposed; State 1 — exposed. 

Character 20. Squamosal. State — zygomatic ramus about same length as otic ramus; State 1 — otic ramus 
developed into plate; State 2 — as State 1 and zygomatic ramus articulates with maxilla; State 3 — otic ramus 
much smaller than zygomatic ramus, latter articulates with maxilla; State 4 — otic ramus much larger than 
zygomatic ramus. Character states unordered. 

Character 21. Vomerine teeth. State — present; State 1 — absent. 

Character 22. Median contact of vomers. State — absent; State 1 — present. 

Character 23. Prootic fusion with frontoparietal. State — absent; State 1 — present. 

Character 24. Occipital condyles. State — confluent; State 1 — approximate; State 2 — widely separated. Char- 
acter state order 0-1-2. 

Character 25. Anterior process of hyale. State — present; State 1 — absent. 

Character 26. Posterior stemum. State — cartilaginous plate, broadens posteriorly; State 1 — cartilaginous plate, 
parallel or narrow; State 2 — as State 1 with mesostemal mineralization; State 3 — stemal style. Character state 
order 0-1-2-3. 

Character 27. Length of last presacral transverse processes relative to sacrum. State — about equal; State 1 — 
last presacral processes <C sacrum. 

Character 28. Sacral diapophyses. State — expanded; State 1 — rounded. 

Character 29. Terminal phalanges. State — simple, knobbed, or claw-shaped; State 1 — T-shaped. 

Character 30. Dial dorsal crest. State — absent; State 1 — present. 

Character 31. Diploid number of karyotype. State — greater than or equal to 26; State 1 — 24. 



VOLUME 112, NUMBER 1 



39 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):40-44. 1999. 

A new species of cardinalfish (Perciformes: Apogonidae) from the 

Bay of Bengal, Indian Ocean 

Thomas H. Fraser 
W. Dexter Bender and Associates, Inc., 2052 Virginia Avenue, Fort Myers, Florida 33901, U.S.A. 

Abstract. — A new species, Apogon oxina, is described from the Bay of Ben- 
gal, Indian Ocean. The preserved color pattern suggests a close relationship 
with Apogon ventrifasciatus from the West Pacific. The new species lacks the 
white spot at the posterior base of the second dorsal fin present in several other 
species and has darker and sharper stripes on the head and upper body and 
well-developed bars on the sides reaching to or above the lateral line. 



New species of the large cardinalfish ge- 
nus Apogon continue to be recognized and 
described at a surprisingly high rate. Most 
of these species are from the Indo-Pacific 
region, with the majority in the large sub- 
genus Ostorhinchus. There are about 111 
valid species in this subgenus, of which 
about 21 undescribed species are known to 
me. Relationships within this large group 
are not well understood. Gon (1996) pro- 
posed phyletic relationships within his con- 
cept of the Apogon subgenus Jaydia. His 
detailed analysis illustrates the complexities 
of developing a robust hypothesis for rela- 
tionships within species groupings of Apo- 
gon prior to testing with an inclusive anal- 
ysis all of proposed subgenera, other spe- 
cies groupings of Apogon and other genera. 
Fraser (1972) suggested that a more com- 
plete understanding of supra- specific rela- 
tionships, within and without Apogon, was 
dependant on well developed systematics of 
the species. Such a critical mass of infor- 
mation and material is emerging. 

Color patterns and variations of those 
patterns appear to be the best distinguishing 
features of closely related species of Apo- 
gon. New species usually are first recog- 
nized by unique color patterns. Consistent 
color pattern differences between closely 
related species may be accompanied by a 
few morphometric differences, such as 



body depth, caudal peduncle length, spine 
lengths, soft fin-ray lengths, jaw length and 
eye diameter, or modally different counts 
for gill rakers and pectoral fin-rays. Other 
differences have been noted by many au- 
thors, such as meristic changes in the spine 
count of the first dorsal fin and soft rays of 
the second dorsal and anal fins; loss of the 
supramaxilla, supraneurals at the anterior 
end of the dorsal fin, one pair of epipleural 
ribs and uroneurals in the caudal skeleton; 
fusion of some hypurals in the caudal skel- 
eton; character of spination of the postem- 
poral, preopercle, infraorbitals; coloration 
of the stomach, intestine and peritoneum; 
presence or absence of known or potential 
bioluminescent ability; shape of the caudal 
fin; elongation of individual vertical fin 
rays; position of the anal opening between 
the pelvic and anal fins; general scale size, 
elaboration of pore architecture on scales as 
a simple pore, complex pores, or pit in 
scale; and the degree of ossification of the 
preopercle. Apogon and the family, in gen- 
eral, are replete with internal and external 
character variation not usually seen within 
a single percoid family. Nevertheless, some 
allopatric species pairs, for example, Apo- 
gon abro gramma- Apogon exo stigma and 
Apogon taeniopterus-Apogon menesemus 
proposed by Fraser & Lachner (1985) differ 
only in color pattern. No morphometric, in- 



VOLUME 112, NUMBER 1 



41 



temal, or meristic differences within those 
species pairs are known. 

The new species belongs in the subgenus 
Ostorhinchus Lacepede, 1802 as defined by 
Fraser ( 1972) using the name Nectamia Jor- 
dan, 1917. Gon (1987) designated a neo- 
type for Apogon fleurieu (Lacepede 1802) 
and effectively relegated Nectamia as a 
synonym. 

Methods 

Methods of taking and recording meristic 
data and measurements are given in Fraser 
& Lachner (1985). All measurements are in 
millimeters to the nearest 0.1. The follow- 
ing acronyms are used to designate insti- 
tutions and collections cited and follow 
general usage given in Eschmeyer (1998); 
BPBM Bernice P. Bishop Museum, Hono- 
lulu; CAS California Academy of Sciences, 
San Francisco; MNHN Museum National 
d'Histoire Naturelle, Paris; USNM collec- 
tions of the former United States National 
Museum, deposited in the National Muse- 
um of Natural History, Smithsonian Insti- 
tution, Washington, D.C. Field station num- 
bers are listed for additional collection in- 
formation, for example PCH-69-279. 

Apogon oxina, new species 
Fig. 1 

Material examined. — Holotype: Apogon 
oxina CAS 33959, 55.9 mm SL; India, Ma- 
dras; in 15-22 m; April-June 1975; K. V. 
Rama Rao. Paratypes: India: CAS 98101; 
7(43-71); data same as holotype. Sri Lanka: 
USNM 213364 55(44-58); Trincomalee; 10 
m; PCH 69-277; 29 Sep 1969; Phillip C. 
Heemstra. USNM 213345 (43-44); PCH 
69-279; 30 Sep 1969; Phillip C. Heemstra. 
Other material: Sri Lanka: USNM 213346 
4(19-28); Trincomalee; CCK 69-135; 10- 
20 m; Christopher C. Koenig; 4 Apr 1970. 

Comparative material. — Paratypes Apo- 
gon ventrifasciatus BPBM 34136 7(35-49); 
Indonesia, Flores I., Maumere Bay; 8 m; J. 
E. Randall; 19 Sep 1988. BPBM 34085 
5(27-41); Indonesia, Flores I., Maumere 



Bay; 3-4 m.; J. E. Randall; 17 Sep 1988. 
Other material. USNM 328265 (45); Papua 
New Guinea, D' Entree asteau Is., Normand- 
by I.; 16 Dec 1993. BPBM 30132 (40-53); 
Indonesia, Lombok I., Sorongjunkung; J. E. 
Randall; 21 Feb 1974. Holotype Apogon 
moluccensis MNHN 8707; 60.3 mm SL, 76 
mm TL; Amboina; Quoy and Gaimard. 

Diagnosis. — An Apogon of the subgenus 
Ostorhinchus with 14 pectoral fin-rays, usu- 
ally 23—24 total gill rakers on the first arch, 
no white spot behind the second dorsal fin, 
dark vertical bars present on body reaching 
to or past the lateral line, dark stripes pre- 
sent on head and upper body, and caudal 
peduncle length 20-25% SL. 

Description. — For general body shape 
see Fig. 1. Range of proportions (as per- 
centages of standard length) with the ho- 
lotype in parentheses: greatest body depth 
36-39 (39); head length 38-41 (41); eye 
diameter 12-13 (12.2); snout length 8-9 
(9.5); bony interorbital width 7-8 (7.5); up- 
per jaw length 18-20 (19); caudal peduncle 
depth 15-17 (16); caudal peduncle length 
20-25 (23); first dorsal-fin spine length 
2.8-3.8 (2.8); second dorsal-fin spine length 
7.9-9.7 (8.0); third dorsal-fin spine length 
17-20 (20); fourth dorsal-fin spine length 
16-21 (21); second dorsal spine 12-16 
(15); first anal-fin spine length 1.9-3.3 
(2.3); second anal-fin spine length 11-12 
(11); pectoral fin length 23-27 (23); pelvic 
fin length 23-26 (26). 

Dorsal fin VII-1,9; anal fin 11,8, last anal 
ray much longer than preceding fin-ray; 
pectoral fin-rays 14-14; pelvic fin 1,5; prin- 
cipal caudal fin-rays 9 + 8; pored lateral 
line scales 24; transverse scale rows above 
lateral line 2; transverse scale rows below 
lateral line 6; median predorsal scales 3-4 
(4); circumpeduncular scale rows 12 
(5 + 2+5); total gill rakers 23-24 (24), usu- 
ally 20-22 (21) well developed 1-2+4-5 
upper, 15-17+0-2 lower (2+5-16+1). 

Villiform teeth in a band on premaxilla; 
two to three rows on dentary; one row on 
palatine and vomer; none on ectopterygoid, 
endopterygoid or basihyal. 



42 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L The holotype of Apogon oxina, 55.9 mm SL, from Madras, India, CAS 33959. 



Vertebrae 10 + 14. Five free hypurals, 
one pair of slender uroneurals, three epur- 
als, a free parhypural. Three supraneurals, 
two supranumerary spines on first dorsal 
pterygiophore. Basisphenoid present. Su- 
pramaxilla absent. Posttemporal serrate on 
posterior margin. Preopercle serrate on ver- 
tical and horizontal margins. Infraorbital 
shelf present on third bone. Scales ctenoid. 
Simple pored lateral-Une scales from post- 
temporal to caudal fin, pored scales usually 
24. 

Life colors. — Unknown. 

Preserved color pattern. — In 70% ethyl 
alcohol: Adult pattern: five dark stripes on 
each side of head with one (unpaired) mid- 
nape from snout to origin of first dorsal fin, 
two stripes over eye extending onto body 
above lateral line, the dorsal stripe along 
base of first dorsal fin reaching to or just 
beyond origin of second dorsal fin, the ven- 
tral one reaching well past origin of second 
dorsal fin; two stripes behind eye, neither 
extending onto body, the upper one reach- 
ing below the posttemporal and the lower 
one from snout, continuing behind eye 
along midline of head to edge of opercle 
above pectoral fin; one stripe extending 
from lower lip below eye onto opercle end- 
ing near pectoral fin base; four or five nar- 
row dark bars from abdomen to first stripe 



above lateral line on body, first bar just be- 
hind and below pectoral fin reaching to lat- 
eral line, second bar beginning on abdomen 
near mid-pelvic fin length reaching past lat- 
eral line merging with first stripe above lat- 
eral line, third bar beginning on abdomen 
near posterior tip of pelvic fin reaching 
above lateral line merging with first stripe 
above lateral line, fourth bar beginning 
above anterior end of anal fin reaching 
above lateral line, and a faint indication of 
a fifth bar beginning about mid-anal fin 
base reaching above lateral line; first dorsal 
fin dusky; melanophore patterns indicate 
stripe may be present in second dorsal fin 
and in anal fin; faint dark mark at posterior 
base of second dorsal; edges of caudal fin 
pale, melanophore patterns indicate remains 
of mid-stripe on caudal fin; outer edge of 
pelvic fin dusky; stomach and intestine 
black, peritoneum pale. Juvenile pattern: 
mid-body lateral stripe extending from 
snout, continuing behind eye along midline 
of head and body onto caudal fin; 3-5 ver- 
tical bars reaching nearly to stripe just 
above lateral line; no melanophore patterns 
in the second dorsal or anal fin; other pre- 
served patterns similar to adults. 

Distribution. — Known from Madras, In- 
dia and Sri Lanka. 

Etymology. — The Greek word oxina 



VOLUME 112, NUMBER 1 



43 



meaning rake or harrow, a feminine noun 
in apposition, referring to the vertical bars 
on the side reaching the stripe. 

Remarks. — This new species of Apogon 
has a combination of characters suggesting 
a close relationships with Apogon ventrifas- 
ciatus Allen, Kuiter & Randall, 1994. Apo- 
gon oxina differs in lacking the white spot 
behind the second dorsal fin, having vertical 
bars reaching to or past the lateral line, a 
shorter caudal peduncle length (20-25% SL 
versus 25-31% SL), stronger and darker 
stripes and bars at all sizes, and an appar- 
ently larger size (71 mm SL versus 53 mm 
SL). Total gill-raker counts for the first arch 
overlap at 23—24 for Apogon oxina and 22- 
25 for Apogon ventrifasciatus. Both species 
have 20-22 well developed gill-rakers and 
have 15-17 well developed lower arch gill 
rakers. 

Allen et al. (1994) compared their new 
species, Apogon ventrifasciatus, with Apo- 
gon moluccensis Valenciennes, 1832 men- 
tioning some similarities in color pattern 
and a difference in the well developed low- 
er arch gill raker count of 15-17 for Apo- 
gon ventrifasciatus and 20—21 for Apogon 
moluccensis. However, the type of Apogon 
moluccensis has 16 well developed lower 
arch gill rakers with no remaining color pat- 
tern other than a white spot at the posterior 
base of the second dorsal fin based on my 
examination. Valenciennes (1832) describes 
the type of Apogon moluccensis as having 
stripes on the nape, blackish lips, first dor- 
sal fin with blackish tip and a reddish body 
lacking any marks or spots on or near the 
caudal fin. Two species with basal second 
dorsal-fin white spots and differing gill-rak- 
er counts exist and more species may be 
present based on color patterns. There are 
two Bleeker names which need to be con- 
sidered in this group besides Apogon ven- 
trifasciatus: Apogon chrysosoma Bleeker, 
1852 and Apogon monochrous Bleeker, 
1856. Gon (1987) identified Apogon mol- 
uccensis from the Maldives at Rasdu Atoll. 
He described the 38 mmSL specimen as 
having 6-7 vertical rows of spots on the 



body from the lateral line to the abdomen, 
stripes in the soft dorsal and anal fin, a 
white spot at the posterior base of the sec- 
ond dorsal fin, 21 well developed gillrakers 
on the first arch and 24(6+18) total gill rak- 
ers. The redescription of types and valid 
species of this group are under study by J. 
E. Randall, G. R. Allen and me. None of 
these white-spot nominal or possibly un- 
described species have the combination of 
characters of Apogon oxina. 

Acknowledgments 

For the loan of material and the use of 
museum facilities I extend thanks to Arnold 
Y. Suzumoto (BPBM), David Catania, Wil- 
liam N. Eschmeyer and Tomio Iwamoto 
(CAS), M. L. Bauchot (MNHN), Susan L. 
Jewett, David G. Smith, Jeffery T Williams 
(USNM) aided in curatorial processes. Jef- 
fery T. Williams helped with photography 
and David G. Smith took radiographs. 
Leonard P. Schultz funds were provided by 
Victor G. Springer (USNM) for several 
study trips to the National Museum. John 
E. Randall (BPBM), Gerald R. Allen 
(WAM) and anonymous reviewers provided 
useful comments on drafts of this manu- 
script. 

Literature Cited 

Allen, G. R., R. H. Kuiter, & J. E. Randall. 1994. De- 
scriptions of five new species of cardinalfishes 
(Apogonidae: Apogon) from Maumere Bay, 
Flores, Indonesia and surrounding regions. — 
Revue fran9aise d'Aquariologie 21(1994)1-2: 
27-38. 

Bleeker, R 1852. Bijdrage tot de kennis der ichthijol- 
ogische fauna van de Moluksche Eilandan. 
Visschen van Amboia en Ceram. — Natuurkun- 
dig Tijdschrift voor Nederlandsch-Indie 3:229- 
309. 

. 1856. Beschrijvingen van nieuwe of weinig 

bekende vischsoorten van Manado en Makassar 
grotendeels verzameld op een rijs naar den Mo- 
luksen Archipel in het gevolg van den Goov- 
erneur-Generaal Duymaer van Twist. — Acta So- 
cietatis Scientiarum Indo-Neerlandicae 1:1-80. 

Eschmeyer, W. N., 1998. Introduction. Pp. 16-22 in 
Catalog of Fishes. Volume I. W. N. Eschmeyer, 
ed., California Academy of Sciences. 



44 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Eraser, T. H. 1972. Comparative osteology of the shal- 
low water cardinal fishes (Perciformes: Apo- 
gonidae) with references to the systematics and 
evolution of the family. — Ichthyological Bulle- 
tin of the J. L. B. Smith Institute of Ichthyology 
34:1-105. 

, & E. A. Lachner. 1985. A revision of the car- 

dinalfish subgenera Pristiapogon and Zoramia 
of the Indo-Pacific region (Teleostei: Apogoni- 
dae). — Smithsonian Contributions to Zoology 
412:1-47. 

Gon, O. 1987. Redescription of Apogon (Ostorhin- 
chiis) fleurieu (Lacepede, 1802) with notes on 
its synonymy. — Japanese Journal of Ichthyolo- 
gy 34(2): 138-145. 

. 1987. The cardinal fishes (Perciformes: Apo- 



gonidae) collected in the Maldive Islands during 
the Xarifa expedition (1957/58). — Special Pub- 
lication, J. L. B. Smith Institute of Ichthyology 
42:1-18. 
-. 1996. Revision of the cardinalfish subgenus 



Jaydia (Perciformes, Apogonidae, Apogon). — 
Transactions of the Royal Society of South Af- 
rica 51:147-194. 

Jordan, D. S. 1917. Notes on Glossamia and related 
genera of cardinal fishes. — Copeia 44:46-47. 

Lacepede, B. G. E. 1802. — histoire Naturelle des pois- 
sons. Paris 4:1-728. 

Valenciennes, A. 1832. Descriptions de plusieurs es- 
peces nouvelles de poissons du genre Apo- 
gon. — Nouvelles Annales Museum Histoire Na- 
turelle, Paris 1:51-60. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):45-5L 1999. 

A new species of deep-water skate, Rajella eisenhardti, 
(Chondrichthyes: Rajidae) from the Galapagos Islands 

Douglas J. Long and John E. McCosker 

(DJL) Department of Ichthyology, (JEM) Department of Aquatic Biology, California Academy of 
Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. 

Abstract. — A new species of skate, Rajella eisenhardti, is described from 
two specimens collected from deep-slope (757-907 m) areas off the Galapagos 
Islands. It is similar to another eastern Pacific species, Rajella nigerrima de 
Buen 1960, but differs in part by having more precaudal vertebrae and more 
pectoral radials, a wider and longer disc, a longer snout, larger gill openings 
with wider interspaces, longer anterior and shorter posterior pelvic lobes, and 
in coloration and spination. 



The skate fauna of the eastern central Pa- 
cific, extending from southern Mexico to 
northern Peru, contains at least 12 species 
(McEachran & Miyake 1984, 1990; Mc- 
Eachran 1995). Galapagos records are lim- 
ited to those of Bathyraja spinossima, Gur- 
gesiella furvescens and Raja velezi (Mc- 
Eachran & Compagno 1979; McEachran 
1995; Grove & Lavenberg 1997). Mc- 
Eachran and Miyake (1984) also reported a 
specimen of Bathyraja cf. richardsoni from 
near the Galapagos Islands (00°11.3'S, 
97°27.7'W, 1710 m), but not within the ar- 
chipelago. The recent capture of two spec- 
imens of an undescribed Rajella using the 
manned submersible Johnson Sea-Link dur- 
ing deep-water dives around the Galapagos 
Islands (McCosker et al. 1997) has prompt- 
ed this paper. 

Methods 

Abbreviations: CAS = California Acad- 
emy of Sciences, ichthyological collections; 
TCWC = Texas Cooperative Wildlife Col- 
lection; TL = total length. 

Measurements and meristic counts fol- 
low current methodology in McEachran & 
Miyake (1984, 1988) and Stehmann (1995). 
Measurements were made with dial calipers 
and recorded to the nearest 0.1 mm. Ver- 



tebral and radial counts were made from ra- 
diographs. 

Rajella eisenhardti, new species 
Figs. 1-2; Table 1 

Diagnosis. — The new species can be dis- 
tinguished from other Rajella species by a 
combination of its moderately long tail (tail 
length greater than distance from snout to 
cloaca) that is largely free of denticles on 
the ventral side; three rows of caudal thorns 
that begin at level of anterior pelvic lobe; 
67—68 predorsal caudal vertebrae; 68-69 
pectoral radials; disc width 55.0-58.5% of 
TL and disc length 50.0-51.2% of TL; 
snout- vent length 44.1-46.4% of TL; pre- 
orbital snout length 13.4-14.1% of TL; and 
ventrum darker than dorsum, with distinct 
light areas around mouth and nasal areas, 
gill openings, abdomen, cloaca, and tips of 
rostrum, tail, and pectoral fins. 

Holotype: CAS 86817, an immature male 
(384.5 mm TL) caught with a siphon hose 
on the bottom at 757 m off Cabo Rosa 
(01°04.74'S,91°11.08'W),Isla Isabella, Ga- 
lapagos Islands, by J. E. McCosker abroad 
the submersible Johnson Sea-Link on 1 1 
Nov 1995. 

Paratype: CAS 86561, an immature male 
(254.1 mm TL) caught with a siphon hose 



46 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. L Preserved types of Rajella eisenhardti. A-B, dorsal and ventral views of holotype, CAS 86817 (384.5 
mm TL); C-D dorsal and ventral views of paratype, CAS 86561 (254.1 mm TL). 



on the bottom at 907 m off James Bay 
(0°26.46'S, 90°19.04'W), Isla San Salvador 
(Santiago Island), Galapagos Islands, by J. 
E. McCosker aboard the submersible John- 
son Sea-Link on 26 Nov 1995. 

Comparative material: Rajella nigerrima. 



TCWC 3881-01, 273 mm TL, juvenile fe- 
male, trawled off Chile (35°53.5'S, 
72°44'W) at 780-925 m, Anton Brunn 
cruise 18A, sta. 40, 9-10 Aug 1966; TCWC 
3885-01, 239 mm TL, juvenile female, 
trawled off Peru (03°15'S, 80°55'W) at 



VOLUME 112, NUMBER 1 



47 




Fig. 2. Rajella eisenhardti. Underwater photo- 
graph of living paratype (CAS 86561) taken from the 
submersible Johnson Sea-Link on the bottom at 907 m 
off James Bay, Isla San Salvador (Santiago Island) Ga- 
lapagos Islands. 



945-960 m, Anton Brunn cruise 18 A, sta. 
120, 10 Sep 1966. 

Description. — See Table 1 for meristics 
and measurements. Meristic values de- 
scribed below are listed for the holotype, 
and values for the paratype are shown in 
parentheses. 

A medium-sized species of Rajella (sen- 
su Stehmann 1970, 1978; McEachran & 
Dunn 1998), its disc heart-shaped and mod- 
erately broad, with broadly-rounded outer 
comers. Disc length 0.88 (0.91) in disc 
breadth. Anterior margin of disc weakly 
concave near tip of rostrum, straight to 
slightly convex to widest part of pectorals. 
Axis of greatest width 0.83 (0.84) times dis- 
tance from tip of snout to axil of pectoral 
fins. Snout to mid-cloaca 2.15 (2.26) times 
in total length. 

Preorbital length 4.0 (3.5) times orbit 
length; preoral length 1.87 (1.98) times in- 
temarial distance. Interorbital distance 1.16 
(0.96) times orbit length; orbit length 1.48 
(1.85) times spiracle length. Anterior nasal 
lobe laterally expanded to form a nasal cur- 
tain, weakly convex anteriorly and finely- 
fringed laterally and posteriorly. Posterior 
nasal flap well-developed, extending to near 
comer of mouth; a single projection on lat- 
eral side of flap with fine fringes on the 
lateral posterior edges, with a smooth mar- 
gin medially; length of flap 0.32 (0.29), and 



width 0.59 (0.57), into preoral length; space 
between fimbriae 0.30 (0.29) into preoral 
length. 

Distance between first gill slits 1.95 
(2.15) times distance between nares; dis- 
tance between fifth gill slits 1.22 (1.33) 
times distance between nares; length of first 
gill slit 1.20 (1.11) times length of fifth gifl 
slit. Third gill slit largest, 0.26 (0.27) times 
distance between nares, fifth gill slit 0.16 
(0.19) times distance between nares, only 
slightly smaller than first, 0.19 (0.21) times 
distance between nares. 

Pelvic fins deeply incised; anterior lobe 
long and moderately narrow, bluntly ta- 
pered and rounded distally; length of ante- 
rior lobe 1.59 (1.46) times length of pos- 
terior lobe. Tail long, gradually tapering to 
tip; distance from mid-cloaca to caudal tip 
1.15 (1.26) times distance from snout to 
cloaca. Base of tail moderately narrow, 
9.3% (9.2%) of cloaca to caudal tip length, 
and moderately convex dorsally, weakly 
concave ventrally becomming flattened dis- 
tally. Lateral folds originating well behind 
the base of tail and continue along length 
to just anterior of caudal tip where it ends 
as small lobed flap. Dorsal fins low and 
rounded; first fin only slightly larger than 
second and similarly shaped, no space be- 
tween dorsal fins or between second dorsal 
and epichordal caudal lobe. 

Dorsum entirely covered with fine den- 
ticles, including tail and dorsal fins, except 
on the lateral and posterior margins of the 
pectoral fins; pelvic fins free of denticles 
and anterior pelvic lobes smooth, but base 
of pelvic with fine denticles in the paratype. 
Larger denticles and thorns with oval or ob- 
long bases, often with weak longitudinal 
ridges and anteriorly curving tips. Larger 
denticles on rostral shaft; one to two thorns 
anterior to each orbit, three posterior to 
each orbit, and one thorn on each side of 
the inner orbit. Five thorns on nuchal mid- 
line, three on each scapular lateral to the 
midline, 6-7 thorns on post-scapular mid- 
line anterior to the tail. Three rows of 
thorns on the dorsal surface of the tail be- 



48 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Measurements (in mm) and percentages of total length (in parentheses) of Rajella eisenhardti 
compared to percentages of total length of R. nigerrima (5 males, 6 females; data from McEachran & Miyake, 
1984). 







Raja 


eisenhardti 


R. nigerrima (n = 


U) 




Holotype 
CAS 86817 


Paratype 
CAS 86561 


Range 


Mean 


Total length 


384.5 


254.1 


139-374 





Snout to mid-cloaca 


178.5 ( 


46.4) 


112.3 (44.1) 


40-44 


43 


Disc width 


225.0 ( 


58.5) 


140 (55.0) 


51-53 


52 


Disc length 


197.0 ( 


51.2) 


127.0 (50.0) 


46-49 


48 


Head length 


104.8 ( 


27.3) 


63.0 (24.8) 


NA 


NA 


Snout length (preorbital) 


54.3 ( 


14.1) 


34.3 (13.4) 


10.2-12.8 


11.0 


Snout length (preoral) 


57.5 ( 


14.9) 


35.9 (14.1) 


12.6-14.5 


13.9 


Snout to maximum width 


132.0 ( 


.34.3) 


78.0 (30.6) 


31-35 


32 


Snout to pectoral axis 


164.5 ( 


:42.8) 


104.8 (41.2) 


NA 


NA 


Prenasal length 


47.7 ( 


T2.4) 


31.4 (12.3) 


9.6-11.5 


11.0 


Orbit diameter 


13.5 ( 


3.5) 


9.8 (3.8) 


3.7^.8 


4.3 


Interorbital distance 


15.7 ( 


:4.0) 


9.5 (3.7) 


3.3-3.6 


3.5 


Spiracle length 


9.1 ( 


:2.3) 


5.8 (2.2) 


1.5-2.3 


1.8 


Orbit and spiracle length 


19.4 ( 


:5.o) 


12.3 (4.8) 


3.4-5.2 


4.6 


Distance between spiracles 


28.0 ( 


7.2) 


17.7 (6.9) 


6.1-6.8 


6.3 


Mouth width 


28.7 ( 


7.4) 


18.7 (7.3) 


6.2-7.2 


6.9 


Distance between nostrils 


30.8 ( 


8.0) 


18.1 (7.1) 


6.5-7.1 


7.0 


Nasal curtail length 


18.6 ( 


4.8) 


10.3 (4.1) 


NA 


NA 


Nasal curtail width 


34.0 ( 


'8.8) 


20.5 (8.1) 


NA 


NA 


Width of 1st gill opening 


5.9 ( 


:i.5) 


3.8 (1.4) 


1.0-1.5 


1.2 


Width of 3rd gill opening 


8.1 ( 


:2.i) 


4.9 (1.9) 


1.0-1.5 


1.2 


Width of 5th gill opening 


4.9 ( 


.1.2) 


3.4 (1.3) 


0.6-1.1 


0.8 


Distance betw. 1st gill openings 


60.3 ( 


T5.6) 


38.9 (15.3) 


13.0-14.5 


13.8 


Distance betw. 5th gill openings 


37.6 ( 


:9.7) 


24.1 (9.4) 


8.1-8.8 


8.5 


Length of anterior pelvic lobe 


63.9 ( 


:i6.6) 


41.3 (16.2) 


10.4-14.7 


12.9 


Length of posterior pelvic lobe 


40.2 ( 


T0.4) 


28.3 (11.1) 


11.8-16.0 


14.9 


Cloaca to 1st dorsal fin 


142.8 ( 


'37.1) 


100.2 (39.4) 


41-44 


42.0 


Cloaca to caudal origin 


188.5 ( 


:49.0) 


134.3 (52.8) 


47-57 


54.0 


Cloaca to caudal tip 


206.1 ( 


;52.5) 


141.8 (55.8) 


50-60 


57.0 


Caudal lobe length 


11.6 ( 


:3.o) 


8.9 (3.5) 


NA 


NA 


1st dorsal height 


21.7 ( 


;5.6) 


15.0 (5.9) 


NA 


NA 


1st dorsal base 


22.5 ( 


:5.8) 


16.1 (6.3) 


NA 


NA 


2nd dorsal height 


21.1 ( 


:5.5) 


12.6 (5.0) 


NA 


NA 


2nd dorsal base 


22.8 ( 


:5.9) 


16.5 (6.5) 


NA 


NA 


Tail width at base 


19.2 ( 


:5.o) 


13.1 (5.2) 


NA 


NA 


Tail width at 1st dorsal origin 


6.9 ( 


:i.8) 


5.0 (2.0) 


NA 


NA 


Tail thickness at base 


13.6 ( 


:3.5) 


7.8 (3.1) 


NA 


NA 


Tail width at 1st dorsal origin 


4.5 ( 


:i.2) 


3.2 (1.3) 


NA 


NA 


No. of upper/lower tooth rows 


48/ 


32 


46/42 


36-46/36-46 


42.4 


No. of trunk vertebrae 


29 




28 


27-30 


28.6 


No. of predorsal caudal vert. 


67 




68 


59-65 


62.6 


No. of pectoral radials 


69 




68 


61-65 


63.0 


No. of pelvic radials 


19 




17 


NA 


NA 



ginning at the mid-level of the pelvic fins 
and continuing to first dorsal fin; median 
caudal thorns 24 (23); lateral caudal thorns 
numbering 38-39 (19-20), thorns diminish- 
ing along the tail. Ventral surface of holo- 



type entirely smooth and bare of denticles; 
ventral surface of paratype smooth except 
for some small denticles on ventrolateral 
margins of tail. 

Upper and lower jaws slightly arched; 



VOLUME 112, NUMBER 1 



49 



teeth in quincunx arrangement with small 
short median crowns becoming more blunt 
and flattened distally; upper tooth rows 48 
(46), lower rows 32 (42). Trunk vertebrae 
29 (28), predorsal caudal vertebrae 67 (68). 
Pectoral radials 69 (68), pelvic radials 19 
(17). 

Color after preservation: Holotype pur- 
plish gray and paratype light brownish gray 
dorsally; both with whitish blotch on the 
dorsal edge of the pectoral fin tip; ventral 
surface darker except for distinct whitish 
areas on the anterior tip of the rostrum, an- 
terior and posterior to the mouth, around 
the nostrils, around each gill opening, 
around the cloaca, on the abdomen, on the 
tips of the anterior pelvic fin lobes, and at 
the tip of the tail; light area between ab- 
domen and gill openings connected by a V- 
shaped mark. Thorns and large denticles 
whitish, smaller denticles same color as sur- 
rounding skin. Color in life (based on an 
underwater photo of the paratype): dorsal 
interior of disc, pelvic base, and tail pale 
brown to grayish; rostrum, dorsal fins, and 
margins of pectoral and pelvic fins blue- 
gray. 

Etymology. — Named in honor of E. Roy 
Eisenhardt, director emeritus of the Cali- 
fornia Academy of Sciences, who has gen- 
erously assisted us and our colleagues. 

Discussion 

The subgenus Rajella was recently ele- 
vated to full generic status (McEachran & 
Dunn 1998), and includes 14 species found 
in the Atlantic, Pacific, and Indian oceans. 
These are: R. annandalei Weber, 1913; R. 
bamardi (Norman 1935); R. bathyphilia 
(Holt & Byrne 1908); R. bigelowi (Steh- 
mann 1978); R. caudaspinosa (Von Bonde 
& Swart 1923); R. dissimilis (Hulley 1970); 
R. fuliginea (Bigelow & Schroeder 1954); 
R. fyllae (Liitken 1887); R. kukujevi (Dol- 
ganov 1985); R. leopardus (Von Bonde & 
Swart 1923); R. nigerrima (de Buen 1960); 
R. purpuriventralis (Bigelow & Schroeder 
1962); R. ravidula (Hulley 1970); and R. 



sadowskii (Krefft & Stehmann 1974). Our 
specimens differ from all of these species 
by a combination of morphometries, mer- 
istics, spination, and coloration. It most 
closely resembles the only other eastern Pa- 
cific Rajella, R. nigerrima, a species that 
ranges from Peru to Chile (McEachran & 
Miyake 1984). 

Rajella nigerrima has a checkered taxo- 
nomic history. It was first placed in the ge- 
nus Breviraja (de Buen 1960; Stehmann & 
Seret 1983), later placed in the subgenus 
Malacoraja (McEachran & Compagno 
1982) which was subsequently elevated to 
generic rank (McEachran 1984; McEachran 
& Miyake 1984; Ishihara & Ishiyama 
1986), later reclassified in Raja, subgenus 
Rajella (McEachran & Miyake 1990; Pe- 
queno & Lamilla 1993), and has recently 
been elevated to generic status (McEachran 
& Dunn 1998). The types of R. nigerrima 
are believed lost and a neotype was desig- 
nated by McEachran & Miyake (1984). 

Our comparisons with the original de- 
scription of R. nigerrima, specimens de- 
scribed by McEachran & Miyake (1984), 
and preserved specimens we examined, 
show significant meristic and morphologi- 
cal differences (Table 1). Raja nigerrima 
has fewer predorsal caudal vertebrae and 
fewer pectoral radials. There are several im- 
portant proportional differences between R. 
eisenhardti and R. nigerrima including the 
latter's smaller preorbital length in relation 
to orbit length, and its first gill slit is longer 
than its fifth gill slit. And (in relation to 
total length) R. nigerrima has a shorter 
snout, a shorter snout to mid cloaca length, 
a reduced disc length and width, a narrower 
mouth, narrower interorbital and interspi- 
racular distances, shorter anterior pelvic 
lobes, larger posterior pelvic lobes, and nar- 
rower third and fifth gill openings. 

Rajella nigerrima has three rows of dor- 
sal thorns posterior to the suprascapular, but 
R. eisenhardti has three rows beginning at 
the middle edge of the pelvic fin; R. niger- 
rima has three dorsal and lateral rows of 
thorns from the base of the tail to the first 



50 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



dorsal fin, but in R. eisenhardti the caudal 
thorns begin at the level of the anterior pel- 
vic lobes. The underside of the tail in R. 
nigerrima has many small, sharp denticles 
except for a very narrow strip along the 
midline, which is bare; in R. eisenhardti, 
the underside is completely bare (holotype) 
or has spinelets only on the lateral margins 
of the tail's underside (paratype). 

The color of R. nigerrima is brown, with 
some indistinct lighter areas on the under- 
side around the mouth, vent, abdomen, and 
around the gills; in R. eisenhardti, the body 
color is gray to brownish gray with very 
distinct whitish areas around the nares, 
mouth, gills, vent, and abdomen, and on the 
tips of the rostrum, tail, and pectoral fins 
and pelvic lobes. 

Acknowledgements 

For assistance and permission to study in 
Ecuador, we sincerely thank: Ing. O. Aguir- 
re, Subdirector de Pesca de Galapagos; C. 
V. Zambrano, Biol. H. MuUer, Dr. F. Or- 
maza-Gonzalez, Instutito Nacional de Pes- 
ca; Lcdo. A. Izurieta Valery and Biol. E. 
Amador, Parque Nacional Galapagos; Ten- 
iente A. Villacis, Capitan de Puerto de Isi- 
dro Ayora; and Dr. C. Blanton, Director de 
Estacion Cientifica Charles Darwin. Many 
other individuals have assisted us in this 
project, and in particular we wish to thank: 
R. G. Gilmore and the staff of the Harbor 
Branch Oceanographic Institution for the 
operation of the Vessel Seward Johnson and 
its submersible, the Johnson Sea— Link; J. D. 
McEachran of Texas A&M University for 
loan of comparative material and critical re- 
view of an earlier draft of this manuscript; 
E. R. Eisenhardt, D. Lin, and B. E. Carlson 
for photographic assistance; W. Eschmeyer, 
D. Catania, and J. D. Fong of the California 
Academy of Sciences; and the David and 
Lucile Packard Foundation and the Discov- 
ery Channel for grants and other assistance. 

Literature Cited 

Bigelow, H. B., & S. C. Schroeder. 1954. A new fam- 
ily, a new genus, and two new species of batoid 



fishes from the Gulf of Mexico. — Breviora 24: 
1-16. 
-, & . 1962. New and little known batoid 



fishes from the western Atlantic. — Bulletin of 
the Museum of Comparative Zoology 128:162- 
244. 

de Buen, F. 1960. Tiburones, ray as y quimeras en la 
Estacion de Biologia Marina de Montemar, 
Chile. — Revista de Biologia Marina 10(l-3):3- 
50. 

Dolganov, V. N. 1985. Raja (Rajella) kukujevi sp. n. 
(Elasmobranchii, Rajidae) from the North- At- 
lantic Ridge. — Zoologicheskii Zhurnal 64(2): 
304-307. 

Grove, J. S., & R. J. Lavenberg. 1997. The Fishes of 
the Galapagos Islands. Stanford University 
Press, Stanford. 863 pp. 

Holt, E. W. L., & L. W. Byrne. 1908. Second report 
on the fishes of the Irish Atlantic slope. — Fish- 
eries Ireland Scientific Investigations 5:141- 
201. 

Hulley, R A. 1970. An investigation of the Rajidae of 
the west and south coasts of southern Africa. — 
Annals of the South African Museum 55(4): 
151-220, 13 pis. 

Ishihara, H., & R. Ishiyama. 1986. Systematics and 
distribution of the skates of the north Pacific 
(Chondrichthyes, Rajoidei). Pp. 269-280 in T. 
Uyeno, R. Arai, T Taniuchi, and K. Matsuura, 
eds., Indo-Pacific Fish Biology: Preceedings of 
the Second International Conference on Indo- 
Pacific Fishes. Ichthyological Society of Japan, 
Tokyo. 

Krefft, G., & M. Stehmann. 1974. Ergebnisse der 
Forschungsreisen des FFS "Walther Herwig" 
nach Sudamerika: XXXIII. Raja {Rajella) sa- 
dowskii spec. nov. (Chondrichthyes, Batoidei, 
Rajidae), ein weiterer neuer Roche vom siid- 
westatlantischen Kontinentalabhang. — Ar- 
chiv Fiir FischereiWeissenschaftliche 25:33- 
50. 

Liitken, C. F. 1887. Korte bidrag til nordisk ichthy- 
ographi. VI. En for Gr0nlandshavet ny Rokke- 
art. — Videnskabelige Meddelelser Naturhistorik 
Forening Kobenhaven 1887:1-4. 

McCosker, J. E., G. Merlen, D. J. Long, R. G. Gilmore, 
& C. Villon. 1997. Deepslope fishes collected 
during the 1995 eruption of Isla Femandina, 
Galapagos. — Noticias de Galapagos 58:22-26. 

McEachran, J. D. 1984. Anatomical investigations of 
the New Zealand skates Bathyraja asperula and 
B. spinifera, with an evaluation of their classi- 
fication within the rajoidei (Chondrichthyes). — 
Copeia 1984(l):45-58. 

. 1995. Rajidae. Pp. 773-777 in W. Fischer, F 

Krupp, W. Schneider, C. Sommer, K. E. Car- 
penter and V. H. Niem, eds., Guia FAO Para la 
Identificacion de Especies para los Fines de la 



VOLUME 112, NUMBER 1 



51 



Pesca. Pacifico centro-oriental. Vol. II. Verte- 
brados— Parte 1 : 647- 1 200. 

- & L. J. V. Compagno. 1979. A further de- 
scription of Gurgesiella furvescens with com- 
ments on the interrelationships of Gurgesielli- 
dae and Pseudorajidae (Pisces: Rajoidei). — Bul- 
letin of Marine Science 29(4):530-553. 

-, & . 1982. Interrelationships of and 

within Breviraja based on anatomical structures 
(Pisces: Rajoidei). — Bulletin of Marine Science 
32(2):399-425. 

-, & K. A. Dunn. 1998. Phylogenetic analysis 
of skates, a morphologically conservative clade 
of elasmobranchs (Chondrichthyes: Rajidae). — 
Copeia 1998 (2):27 1-290. 

-, & T. Miyake. 1984. Comments on the skates 
of the tropical eastern Pacific: one new species 
and three new records (Elasmobranchii: Raji- 
formes). — Proceedings of the Biological Soci- 
ety of Washington 97(4):773-787. 

-, & . 1988. A new species of skate from 

the Gulf of California (Chondrichthyes, Rajoid- 
ei).— Copeia 1988(4):877-886. 

-, & . 1990. Zoogeography and bathym- 



etry of skates (Chondrichthyes, Rajoidei). Pp. 
305-326 in H. L. Pratt, S. H. Gruber and T. 
Taniuchi, eds., Elasmobranchs As Living Re- 
sources. National Oceanographic and Atmo- 
spheric Administration Technical Report 90. 

Norman, J. R. 1935. Coast fishes. Part I. The south 
Atlantic. — "Discovery" Reports 12:1-58. 

Pequeno, G. & J. Lamilla. 1993. Batoideos comunes a 
las costas de Chile y Argentina-Uruguay (Pi- 



sces: Chondrichthyes). — Revista Biologia Ma- 
rina, Valparaiso 28(2):203-217. 

Stehmann, M. 1970. Vergleichend morphologische und 
anatomische Untersuchungen zur Neuordnung 
der Systematik der nordostatlantischen Rajidae 
(Chondrichthyes, Batoidei). — Archiv fiir Fis- 
cherei- und Meeresforschung Wissenschaftliche 
21:73-164. 

. 1978. Raja ''bathyphilia'\ eine doppelart des 

subgenus Rajella: wiederbeschreibung von R. 
bathyphilia Holt & Byrne, 1908 und Raja bi- 
gelowi spec. nov. (Pisces, Rajiformes, Raji- 
dae). — Archiv fiir Fischerei- und Meeresfor- 
schung Wissenschaftliche 29(l/2):23-58. 

. 1995. First and new records of skates (Chon- 
drichthyes, Rajiformes, Rajidae) from the west 
African continental slope (Morocco to South 
Africa), with descriptions of two new species. — 
Archive of Fishery and Marine Research 43(1): 
1-119. 

, & B. Seret. 1983. A new species of deep- 



water skate, Breviraja africana sp. n. (Pisces, 
Batoidea, Rajidae), from the eastern Central At- 
lantic slope, and remarks on the taxonomic sta- 
tus of Breviraja Bigelow & Schroeder, 1948. — 
Bulletin de Museum National d'Histoire Natu- 
relle, Paris, 4th series, section A, 3:903-925. 

Von Bonde, C, & D. B. Swart. 1923. The Plagiostoma 
(skates and rays) collected by the S. S. "Pick- 
le". — Union of South Africa Fisheries and Ma- 
rine Biological Survey 3(5): 1-22. 

Weber, M. 1913. Die fische der Siboga-Expedition. — 
Siboga Expedition 57:1-710. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):52-58. 1999. 

A new genus of the subfamily Cubacubaninae 
(Insecta: Zygentoma: Nicoletiidae) from a Mexican cave 

Luis Espinasa 

CEAMISH, Universidad Autonoma del Estado de Morelos Av. Universidad #1001, Col. 
Chamilpa, Cuemavaca, Mor. C.P. 62210, Mexico 

Abstract. — ^A new genus and species (Squamigera latebricola) of the sub- 
family Cubacubaninae from a Mexican cave is described. The new genus is 
distinguished from almost all the genera of this subfamily by having scales. It 
differs from Texoreddellia, the only other genus of this subfamily with scales, 
by the morphology of scales and other characters, which are discussed. 



The family Nicoletiidae is a frequently 
encountered group in caves of Mexico and 
Central America (Reddell 1981), but with 
few exceptions, most species await descrip- 
tion. During exploration of several caves in 
the states of Guerrero and Morelos, Mexi- 
co, several specimens of this family were 
collected. A specimen collected from "Po- 
zas Azules" cave is a previously undescri- 
bed genus with scales of the subfamily Cu- 
bacubaninae. Available material is of par- 
ticular interest because most members of 
the subfamily lack scales. Only two genera 
have them: Texoreddellia (Wygodzinsky 
1973) and the new genus described here. 
However, the morphology of the scales 
among other characteristics is drastically 
different for these two genera; therefore, 
they are probably not closely related. 

Study Area 

The material was collected in "Pozas 
Azules" cave (Taxco de Alarcon Munici- 
pality, Guerrero State, Mexico, 18°36'40"N, 
99°33'25"W). The cave is 1399 m long and 
+53 m deep. This cave is the resurgence of 
a cave system formed by Isote cave and 
Cueva de las Pozas Azules. Total length of 
system is almost three kilometers and depth 
is 230 m (for a detailed description of both 
caves' topography, see Espinasa-Pereiia 
1989). 



This cave system has a very peculiar fau- 
na. Apart from the new genus described 
here, a freshwater polychaete has been de- 
scribed as a troglobite (Solis-Weiss & Es- 
pinasa 1991). Some undescribed copepods 
living in association with the polychaete 
will probably be characterized as troglobi- 
tes. Other organisms collected in the system 
are troglophile ricinulids (Cryptocellus bo- 
ne ti), collembola of the families Entomo- 
bridae and Sminthuridae, beetles, two spe- 
cies of spiders, millipedes, mites, protozoa 
and bacteria (Salmonella, Klebsiella, and 
Bacillus). To date, physical connection be- 
tween Isote cave and Cueva de las Pozas 
Azules has not been explored because both 
ends of the caves end in sumps. Connection 
is assumed in part on geohydrological evi- 
dence (Espinasa-Pereiia 1989), by the prox- 
imity of the two final sumps found in each 
cave (less than 20 m apart), and most im- 
portantly, by the shared fauna mentioned 
above. 

Cave formation. — The limestone rock 
was formed during the lower Cretaceous 
period (Albian, Cenomanian, and Turoni- 
an). Elevation occurred at the end of Cre- 
taceous (Maastrichtian), when erosion be- 
gan. Presence of an igneous rock (Riolite 
tilzapotla) overlaying the limestone from 
which the water that formed the cave 
comes, seems to indicate that the cave was 
not formed until late Oligocene (L6pez-Ra- 



VOLUME 112, NUMBER 1 



53 



mos 1974). It is also known that the cave 
was already formed in the Quaternary be- 
cause there is evidence in cave sediments 
of climatic changes related to this period 
(Espinasa-Pereila, pers. comm.). Thus, if 
the cave was formed at some point between 
Oligocene and Quaternary, there is reason 
to assume that the troglobites in this cave 
system separated from the outside commu- 
nities at more or less the same time. 

Materials and Methods 

The specimen was found crawling on the 
cave wall. It was placed into a vial with 
70% alcohol. Dissections were made with 
the aid of a stereo microscope and the dif- 
ferent parts of the body were mounted in 
fixed preparations with Hoyer's solution. 

All illustrations were made with the aid 
of a camera lucida attached to a compound 
microscope. 

The type is deposited in the following 
collection: LESM-DB-MEX (Laboratory of 
Ecology and Systematic of Microarthro- 
pods. Department of Biology, Faculty of 
Sciences, UNAM. Mexico D. E) Catalog 
number: ZYG-2. 

Squamigera, new genus 

Diagnosis. — A member of the subfamily 
Cubacubaninae with scales. Cerci of male 
with modified spines. Parameres with a 
cleft on apex. 

Description. — Body robust, long, ap- 
proximately parallel- sided, thorax slightly 
but distinctly wider than abdomen. Scales 
present, numerous, and multiradiate. Form 
mucronate to emarginate, with borders 
slightly serrated. Head, thorax, and abdo- 
men with scales and setae. Legs with scales 
on proximal articles. Mouthparts and ab- 
dominal stylets only with setae. 

Pedicellus of male with unicellular 
glands and a small spur on its base. Mouth- 
parts not specialized. Mandibles strongly 
sclerotized apically and with usual teeth. 
Galea apically with several sensory pegs. 
Lacinia heavily sclerotized distally; first 



process of lacinia pectinate. Labium with- 
out prominent lateral lobes. 

Tarsi with 4 articles. Praetarsi with 3 sim- 
ple claws, median claws glabrous, slender, 
and smaller than lateral claws. 

Abdominal sterna II (apparently)-III-VII 
subdivided into coxites and stemite. Sterna 
VIII and IX of male entire. Sterna II-VII 
with 2 + 2 macrochaetae. Coxites on seg- 
ments II-IX with stylets. Eversible vesicles 
on segments II- VI, pseudovesicles on VII. 
Urosternum VIII with a wide and not too 
deep posterior emargination. Posterior pro- 
jections acute to slightly rounded, pointing 
slightly outward. Tergum X protruding, al- 
most straight on posterior border, posterior 
angles with 2 or 3 subequal macrochaetae. 
Urostema III and IV simples. Cerci of male 
with modified spines. 

Parameres with a cleft and specialized se- 
tae on apex. Opening of penis longitudinal. 

Type species. — Squamigera latebricola. 

Etymology. — Latin Squamigera = scale 
bearing, scaly. Here treated as a noun in the 
nominative singular. Gender, feminine. 
Makes reference to its body covered with 
scales. 

Distribution. — The only individual found 
is from "Cueva de las Pozas Azules" cave, 
Taxco de Alarcon Municipality, Guerrero 
State, Mexico, 18°36'40"N, 99°33'25"W. 

Remarks. — Squamigera belongs to a 
group of nicoletiid genera, the Cubacuban- 
inae (Mendes 1988), characterized by sub- 
divided abdominal sterna II-VII and fused 
coxites of VIII and IX abdominal segments 
of the male. Squamigera is distinguished 
from almost all genera of this subfamily by 
having scales. It differs from Texoreddellia, 
the only other genus with scales, by the 
morphology of scales, having scales on 
head, morphology of parameres, and mod- 
ified spines in male cerci. 

The genus shares with Anelpistina (Sil- 
vestri 1905 as subgenus), Prosthecina (Sil- 
vestri 1933), and Cubacubana (Wygodzin- 
sky & Hollinger 1977), the presence of 
spines in cerci and a posterior emargination 
on urosternum VIII, which are absent in 



54 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





:Mn i^.-.. k 










/ 







X\\\\ 
G 







■tv\li-M4k 




I fL 1 




Fig. L Squamigera latebricola new genus. (Male): A, Head; B, Basal article and pedicellus of antenna; C, 
Small spur of pedicellus; D, Labial palp and labium; E, Mandible; F, Thoracic tergum (scales and setae partially 
shown); G, Posterior border of metanotum (scales partially represent). 



VOLUME 112, NUMBER 1 



55 



>^j/f^iff^ 




Fig. 2. Squamigera latebricola new genus. (Male): A, Hind leg, microchaetae partially shown, scales not 
shown; B, Urostemum VII and VIII (scales and setae partially shown); C, Genital area; D, Scales of head. 



56 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






'^^'V~/m'^^ ^ 



Fig. 3. Squamigera latebricola new genus. (Male): A, Scales and spines of the posterior border of the 
metanotum; B, Scales of urotergum I; C, Urotergum X (scales and setae partially shown): D, Cercus basal 
portion. 




Texoreddellia. It is possible that Squami- 
gera has closer affinities with this group of 
species than with Texoreddellia. 

Squamigera latebricola, new species 
Figs. lA-G, 2A-D, 3A-D 

Material examined. — Mexico. Guerrero: 
Taxco de Alarcon, Cueva de las Pozas Azu- 



les (+53 meters deep, 1399 meters long). 3 
Dec 1988, R. Espinasa-Closas col. Male ho- 
lotype. 

Description. — Maximum body length 
22.0 mm. Maximum length of antennae un- 
known (broken), of caudal appendages 28.0 
mm. Body and legs robust. Head, thorax, 
abdomen, and proximal articles of legs cov- 



VOLUME 112, NUMBER 1 



57 



ered by scales (Figs. 2D, 3 A, B). General 
color light yellow to white. 

Head with macrochaetae and microchae- 
tae as shown in Fig. lA. Basal article of 
antennae in males has a small projection 
with 2 macrochaetae. Pedicellus of anten- 
nae of male as shown in Fig. IC, with 5 
clusters of unicellular glands and a small 
sclerotized spur. Mouthparts relatively long. 
Apex of maxillary palp similar to Texored- 
dellia, but with a third very small extra con- 
ule. Labial palp long, apical article barely 
longer than wide, labium and first article of 
labial palp with macrochaetae (Fig. ID). 
Mandible without very small pegs over the 
bigger tooth. Setae as shown in Fig. IE. 
Legs as shown in Fig. 2 A, relatively long; 
hind tibia approximately 5 times longer 
than wide; claws of normal size. 

Cerci of male with a longer than wider 
basal article and a very long second article, 
followed by short articles of subequal size. 
In the very long article the spines start al- 
most at its base until they become a more 
prominent group; the relatively long spines 
are inserted on distinct tubercles (Fig. 3D). 
Compared with other genus of the subfam- 
ily, spines are relatively small. 

Thoracic terga with long macrochaetae 
(Fig. IF), very abundant on all borders. 
Also, on the posterior border they show a 
series of small sclerotized spines (Figs. IG, 
3A). Tergum X protruding, almost straight 
on the posterior border (Fig. 3C). 

Urostema III and IV of male without 
modifications. Urostemum VIII with a wide 
and not too deep posterior emargination. 
Posterior projections acute to slightly 
rounded, pointing slightly outward (Fig. 
2B). 

Urostemum IX of male slightly curved 
behind and centrally on the posterior border 
with macrochaetae slightly more sclerotized 
(Fig. 2C). Point of insertion of parameres 
in urostemum IX is deep, with internal face 
of coxal processes with macrochaetae more 
sclerotized (Fig. 2C) similar to some mem- 
bers of the genus Anelpistina. 

Stylets IX bigger than the others, with 6 



macrochaetae and an extra subapical pair. 
The other stylets only have 3 macrochaetae 
and an extra subapical pair. In males, styles 
IX without other modifications. 

Penis and parameres as in Fig. 2C. Par- 
ameres curved outward, attaining less than 
Vi the length of stylets IX. Surface of par- 
ameres with a cleft and short setae. 

Etymology. — Latin latebricola = One 
who dwells in lurking places. Gender mas- 
culine. Makes reference to the cavemicole 
habitat where this species dwells. 

Remarks. — Despite several trips to Pozas 
Azules cave, only a single male was col- 
lected. Because of its size, this specimen is 
probably an adult with all the secondary 
sexual characters developed. Individuals at 
different stages of the postembryonic de- 
velopment might differ in characters, such 
as the number and size of spines in cerci. 

Many caves in this area of Guerrero, 
Mexico, have been explored and have 
yielded many individuals of the family Ni- 
coletiidae (Espinasa 1991), but all from dif- 
ferent genera. The one locality where S. la- 
tebricola has been found is in this single 
cave. Therefore it is probably a troglobite 
with a very restricted geography. 

Acknowledgments 

I thank Dr. Jose G. Palacios Vargas, di- 
rector of "Laboratorio de Ecologia y Sis- 
tematica de Microarstropodos," where most 
of the descriptive work was done and to the 
Director of CEAMISH-UAEM, Dr. Oscar 
Dorado, for the support to publish this man- 
uscript. Thanks are also due to Ramon Es- 
pinasa-Perena, geologist at the "Universi- 
dad Nacional Autonoma de Mexico" for 
providing the information regarding the 
date of formation of the cave, and to Dr. 
Luis F Mendes and Monika Baker for re- 
viewing the manuscript. 

Literature Cited 

Espinasa, L. 1991. Descripcion de una nueva especie 
del genero Ciibacubana (Zygentoma: Nicoleti- 
idae) y registro del genero para America Con- 



58 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tinental. — Folia EntomoI6gica Mexicana 82:5- 
16. 

Espinasa-Perena, R. 1989. El resumidero del Isote y la 
cueva de las Pozas azules. — TepeyoUotli: Gac- 
eta de la Sociedad Mexicana de Exploraciones 
Subterraneas 4:24-27. 

Lopez-Ramos, E. 1974. Geologia de Mexico. Third 
Edition. Volume 3. Secretaria de Educacion 
Publica, Edicion Escolar, Mexico, 453 pp. 

Mendes, L. E 1988. Sur deux nouvelles Nicoletiidae 
(Zygentoma) cavemicoles de Grece et de Tur- 
quie et remarques sur la systematique de la fam- 
ille. — Revue Suisse de Zoologie 95 (3):751- 
772. 

Reddell, J. R. 1981. A review of the cavernicole fauna 
of Mexico, Guatemala, and Belize. — Texas Me- 
morial Museum Bulletin 27. 327 pp. 

Silvestri, F. 1905. Materiali per lo studio dei Tisanuri. 
VI. Tre nuove specie di Nicoletia appartenenti 



ad un nuovo sottogenero. — Redia (Firenze) 2: 
111-120. 

. 1933. Nuovo contributto alia conoscenza dei 

Tisanuri del Messico. — Bolletino del Laborato- 
rio di Zoologia general e agraria di Portici 27: 
127-144. 

Soils- Weiss, v., & L. Espinasa. 1991. Lycastilla cav- 
ernicola, a new freshwater nereidid from an in- 
land mexican cave (Polychaeta: Nereididae: Na- 
manereidinae). — Proceedings of the Biological 
Society of Washington 104:631-639. 

Wygodzinsky, P. 1973. Description of a new genus of 
cave thysanuran from Texas (Nicoletiidae, Thy- 
sanura, Insecta). — American Museum Novitates 
2518:1-8. 

, & A. M. HoUinger. 1977. A study of Nico- 
letiidae from Cuba (Thysanura). — Resultats des 
Expeditions Biospeleologiques Cubano-Rou- 
maines h. Cuba 2:313-324. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):59-69. 1999. 

Two new species of the genus Anelpistina (Insecta: Zygentoma: 
Nicoletiidae) from Mexican caves, with redescription of the genus 

Luis Espinasa 

CEAMISH, Universidad Autonoma del Estado de Morelos, Av. Universidad #1001 Col. 
Chamilpa, Cuemavaca, Mor. C.P. 62210. Mexico 

Abstract. — Two new species of the genus Anelpistina are described, A. in- 
appendicata and A. cuaxilotla. Of these two, A. inappendicata lacks the artic- 
ulated submedian appendages found in urostemite IV of males, which are di- 
agnostic characters of genus Anelpistina. Analysis of other characters shows 
that this species has close affinities with members of genus Anelpistina, and 
therefore a new re-description of the genus is provided to include this species. 



Insects of the family Nicoletiidae are 
common inhabitants of caves in Mexico. 
Numerous species of troglobites and trog- 
lophiles have been collected from this area, 
but with few exceptions, they await descrip- 
tion (Reddell 1981). During the exploration 
of several caves in the states of Guerrero 
and Morelos, Mexico, several specimens of 
this family were collected and are described 
here, thus contributing to the knowledge of 
the cave fauna of the region. 

Definition of genus Anelpistina by Sil- 
vestri (1905) is based on the articulated 
submedian appendages found in urostemite 
IV of males. Organisms who are closely re- 
lated to species of genus Anelpistina, but 
lack such appendages, would be incorrectly 
placed in genus Cubacubana (Wygodzin- 
sky & Hollinger 1977). For this reason, ge- 
nus Anelpistina is redescribed using more 
characters to include species, such as the 
one described in this paper, that lack the 
appendages. 



taken alive to the laboratory. Dissections 
were made with aid of a stereo microscope 
and the different parts of the body were 
mounted in fixed preparations with Hoyer's 
solution. 

Comparison with A. boneti was done 
with specimens collected by author from 
their type localities of Juxtlahuaca cave, 
Colotlipa, Guerrero, Mexico, and from a 
new locality, the cave of "Iglesia-Mina su- 
perior", San Juan, Tepoztlan, Morelos, 
Mexico. Comparison with other species 
was based on published material. All illus- 
trations were made with aid of a camera 
lucida attached to a compound microscope. 

Types were deposited in the following 
collection: LESM-DB-MEX (Laboratory of 
Ecology and Systematic of Microarthro- 
pods. Department of Biology, Faculty of 
Sciences, UN AM. Mexico D. F). Catalog 
numbers: ZYG-3 for Anelpistina inappen- 
dicata and ZYG-4 for Anelpistina cuaxilot- 
la. 



Materials and Methods 

Detailed descriptions of the caves can be 
found in Hoffman et al. (1986), Diamant & 
Espinasa-Perena (1991), and in Espinasa- 
Pereiia (1989). Organisms collected were 
crawling on floor or under rocks. They were 
placed into a vial with 70% alcohol or were 



Anelpistina Silvestri, 1905 

Diagnosis. — Member of subfamily Cu- 
bacubaninae without scales. Submentum 
without conspicuous lateral lobes bearing 
glandular pores. Urostemite IV of adult 
male with 1 + 1 articulated submedian ap- 
pendages and/or with point of insertion of 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



parameres in urostemum IX deep, normally 
with internal face of coxal processes with 
macrochaetae more or less sclerotized. 

Description emending. — Body slender, 
approximately parallel- sided, thorax slight- 
ly but distinctly wider than abdomen. An- 
tennae and body are of similar length. Head 
with approximately 8 + 8 macrochaetae in 
border of insertion of antennae. Pedicellus 
of male normally with 3 clusters of unicel- 
lular glands, one of them very long. 

Mouthparts not specialized. Mandibles 
strongly sclerotized apically and with usual 
teeth, typically with more than 4 macro- 
chaetae. Galea apically with several sensory 
pegs. Lacinia heavily sclerotized distally; 
first process of lacinia pectinate. Labium 
without prominent lateral lobes. Next to last 
article of labial palp more or less with a 
rounded bulkiness. 

Tarsus longer or equal in length to tibia. 
Tarsi with 4 articles. Praetarsi with 3 simple 
claws, median claws glabrous, slender, and 
smaller than lateral claws. 

Abdominal sterna II- VII subdivided into 
coxites and stemite. Sterna VIII and IX of 
male entire. Coxites on segments II-IX 
with stylets. Eversible vesicles on segments 
11- VI, pseudovesicles on Vn. Urostema in 
simple. Urostema IV in males with or with- 
out 1 + 1 articulated submedian append- 
ages. Point of insertion of parameres in 
urostemum IX generally deep except in A. 
weyrauchi (Wygodzinsky 1959) and to cer- 
tain degree in A. ruckeri (Silvestri 1905), 
normally with internal face of coxal pro- 
cesses with macrochaetae more or less 
sclerotized. Cerci of male with modified 
spines. 

Normal parameres, not subdivided or 
somewhat constricted apically. With spe- 
cialized setae on apex. Opening of penis 
longitudinal. 

Type species. — Anelpistina wheeleri Sil- 
vestri 1905. 

Remarks. — According to Silvestri's 
(1905) definition of genus Anelpistina, the 
main difference between this genus and the 
other members of the subfamily Cubacu- 



baninae is the articulated submedian ap- 
pendages in the urostemite IV of males. 
Specimens that are closely related to spe- 
cies of the genus Anelpistina, but lack such 
appendages could incorrectly be placed in 
the genus Cubacubana. Therefore a broader 
re-description is given to include these 
specimens. 

With the new re-description, genus Anel- 
pistina differs from Texoreddellia (Wygod- 
zinsky 1973) by the absence of scales and 
of a conspicuous process of the pedicellus 
in males, as well as by the presence of mod- 
ified spines on the cerci of males. From A/- 
lonicoletia (Mendes 1992) by the presence 
of stylets in urostemite n. From Neonico- 
letia (Paclt 1979) by the aspect of the en- 
dopodium. Prosthecina (Silvestri 1933) and 
Cubacubana agree with Anelpistina in the 
possession of modified spines on the cerci 
of males, but Prosthecina has a submentum 
with conspicuous lateral lobes bearing nu- 
merous glandular pores. Cubacubana lacks 
1 + 1 articulated submedian appendages in 
urostemite IV and the point of insertion of 
parameres in urostemum IX is shallow 
without more or less sclerotized macro- 
chaetae in the internal face of coxal pro- 
cesses. Anelpistina has either or both the 
appendages and the deep insertion. 

Anelpistina inappendicata, new species 
Figs. lA-F, 2A-G 

Type material. — Mexico, Guerrero State, 
Tetipac municipality, Agua Brava cave (17 
meters deep, 819 meters long), ex soil, 7 
Aug 1988, 25 Mar 1989 and 15 Mar 1991, 
L. Espinasa col. Male holotype, 4 male par- 
atypes and ten female paratypes. 

Description. — Maximum body length 
13.5 mm. Maximum length of antennae 9.0 
mm, of caudal appendages 9.0 mm. When 
complete, antennae and caudal appendages 
measure more or less the length of body. 
General color hght yellow to white. 

Head with macrochaeta and microchaeta 
as shown in Fig. IF, with approximately 8 
+ 8 macrochaetae on border of insertion of 



VOLUME 112, NUMBER 1 



61 







&a^ 



/f 






^:r/ 



X/ 



-'-'h' 



'A 
% 








'A 



Fig. 1. Anelpistina inappendicata n. sp.: A, Mandible; B, Apical portion of maxilla; C, Labium with palp; 
D, Hind leg, microchaetae partially shown; E, Male. Basal portion of antenna; F, Head. 



antennae. Basal article of antennae in males 
without projections. Pedicellus of antennae 
of male as shown in Fig. IE, with numerous 
clusters of unicellular glands. Female basal 
articles of antennae simple. 



Mouthparts relatively long, apex of max- 
illa as shown in Fig. IB. Labial palp long, 
apical article longer than wide. Labium and 
first article of labial palp with macrochae- 
tae. Mandibles with approximately 8 ma- 



62 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 









Vvi\^ii 











u 



I 




. / If 

A ''''' 



\ 



'/ma- 



Fig. 2. Anelpistina inappendicata n. sp.: A-B, Male. Genital area; C, Ovipositor and subgenital plate; D, 
Subgenital plate; E, Basal portion of cercus; F, Modified spines of cercus, higher magnification; G, Thoracic 
nota. dorsal. 



VOLUME 112, NUMBER 1 



63 



crochaetae, some very small pegs on larger 
tooth (Fig. lA). Legs as shown in Fig. ID, 
moderately elongate; hind tibia approxi- 
mately 7X longer than wide. Claws of nor- 
mal size. 

Cerci of male with a longer than wide 
basal article, a very long second one bear- 
ing numerous spines, followed by numer- 
ous short articles of simple chaetotaxy. 
Spines along most of second article and of 
similar size. Spines with medial enlarge- 
ment in form of ring (Fig. 2E, F). Female 
cerci simple. 

Thorax with short macrochaetae, very 
abundant on lateral borders, 1 + 1 subme- 
dian distinct macrochaetae, apart from sev- 
eral setae of varied sizes, in posterior border 
of pro-, meso- and metanotum. 2 + 2 also 
on anterior border of pronotum (Fig. 2G). 

Urotergite X almost straight in both sex- 
es, posterior angles with few macrochaetae 
of varied sizes, length of inner macrochae- 
tae almost equal to distance between them. 

Abdominal terga and sterna as in other 
members of genus. Abdominal sterna II- 
VII subdivided into coxites and stemite. 
Sterna VIII and IX of male entire. Uroster- 
num ni and IV of male without modified 
coxites. Urostemum Vni of male shallowly 
emarginate on posterior margin. Uroster- 
num IX of male straight behind without a 
row of sensory cones, but point of insertion 
of parameres in urostemum IX is deep, with 
internal face of coxal processes with ma- 
crochaetae more sclerotized (Fig. 2A, B). 
Stylets II- VIII as usual for subfamily. Sty- 
lets IX larger than others, with 2 or 3 ma- 
crochaetae and an extra subapical pair. Ter- 
minal spine with small teeth. In males and 
females styles without modifications. 

Penis and parameres as shown in Fig. 
2A, B. Parameres attaining Vi of stylets IX, 
divergent and slightly concave. Surface of 
parameres with short setae. Subgenital plate 
of female triangular, twice as wide as it is 
long (Fig. 2C, D). Ovipositor barely sur- 
passing apex of stylets IX. Gonapophyses 
with approximately 12 articles. 

Postembryonic development not very 



complex, with younger instars almost iden- 
tical to older ones except for size. In small- 
er sized males (9 mm.), spines on cerci lack 
medial enlargement in form of a ring (it is 
unknown if younger instars than those col- 
lected lack spines). In females, length of 
ovipositor increases proportionally with 
body size until at body length of 9.5 mm, 
ovipositor barely surpasses apex of stylets 
IX. Longer body sizes of up to 13.5 mm. 
did not bring an increase of size of ovipos- 
itor when compared to stylets IX. 

Etymology. — In/appendicata = without/ 
appendages. Makes reference to the lack of 
articulated submedian appendages in this 
species, which are normally found in the 
urostemite IV of males in the genus Anel- 
pistina. 

Remarks. — The characters that allow us 
to identify A. inappendicata as more closely 
related to species of the genus Anelpistina 
than to Cubacubana, even though it lacks 
such appendages are: Antennae and body 
are of similar length, as in Anelpistina, 
while in Cubacubana the antennae are 
twice as long; Pedicellus of antennae with 
three clusters of unicellular glands, one of 
them very long. This is in general the case 
in Anelpistina. The norm in Cubacubana is 
four clusters bordered by a "U" shaped 
row of microchaetae; Head with approxi- 
mately 8 + 8 macrochaetae in the border of 
insertion of the antennae, as in some Anel- 
pistina, while Cubacubana normally has 5 
+ 5; Lateral borders of thoracic nota with 
approximately eight macrochaetae. This is 
closer to most Anelpistina, with approxi- 
mately five macrochaetae, than to Cubacu- 
bana, with approximately three; Mandibles 
with more than four macrochaetae, as in 
some Anelpistina, while in Cubacubana 
there are typically four; The next to last ar- 
ticle of the labial palp without a rounded 
bulkiness. In this character A. inappendi- 
cata is, on the contrary, more similar to 
species of the genus Cubacubana whose ar- 
ticle is straighter, while in Anelpistina they 
are bulky. This character could be ex- 
plained by a loss of bulkiness in A. inap- 



64 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



pendicata; Tarsus longer than the tibia. Al- 
though this is generally the case for Cuba- 
cubana, in Anelpistina it is longer to equal; 
Point of insertion of parameres in 
urostemum IX deep, with internal face of 
coxal processes with macrochaetae sclero- 
tized. This deep insertion and the sclero- 
tized macrochaetae are absent in Cubacu- 
bana. On the contrary, the deep insertion is 
present in all the species of Anelpistina, ex- 
cept A. weyrauchi, and the sclerotized ma- 
crochaetae are present in most of the spe- 
cies. 

Anelpistina inappendicata and the other 
North American species of Anelpistina have 
in common a deep insertion of the paramers 
and other intermediate characteristics, 
which are absent in the South American 
species (A. weyrauchi). It is likely therefore 
that the divergence of A. inappendicata 
from the species of this genus happened af- 
ter the separation of A. Weyrauchi from the 
group, but before the radiation of the other 
described North American species of Anel- 
pistina. 

Anelpistina decui (Wygodzinsky & Hol- 
linger 1977), new combination Cubacu- 
bana decui Wygodzinsky & Hollinger, 
1977:320-322, figs. 4-6; Mendes, 1986: 
341; Espinasa, 1991:6, 14. 

Remarks. — Although A. decui lacks the 
articulated submedian appendages on uros- 
temite IV of the male, this species has clos- 
er affinities with members of Anelpistina 
than it does to Cubacubana because its 
point of insertion of parameres in uroster- 
num IX is deep and the internal face of cox- 
al processes has sclerotized macrochaetae. 

An analysis of other characters present in 
A. decui that allow us to identify it as more 
closely related to species of genus Anelpis- 
tina than to Cubacubana, even though it 
lacks such appendages, is given based on 
the illustrations and description of Wygod- 
zinsky (1977): Antennae twice as long the 
length of the body, as in Cubacubana, 
while in Anelpistina antennae are of similar 
length. Even though this character would 
appear to make it closer to Cubacubana, its 



longer antennae may be due to a secondary 
adaptation to its cavemicole environment; 
Pedicellus of antennae with two long clus- 
ters of unicellular glands and no "U" 
shaped row of microchaetae. In Anelpistina, 
normally there are three clusters, one of 
them very long. The norm in Cubacubana 
is four clusters bordered by a "U" shaped 
row of microchaetae; Lateral borders of 
thoracic nota with approximately five ma- 
crochaetae, as in most Anelpistina, while 
Cubacubana approximately has three; The 
next to last article of the labial palp with a 
somewhat rounded bulkiness, as in Anelpis- 
tina, while in Cubacubana it is straighter; 
Cerci of male with four spines; a small one, 
a long and strong one, another small and a 
very long curved one. Although in both 
Anelpistina and Cubacubana there is diver- 
sity in morphology of spines, this particular 
arrangement of four spines is exactly the 
same for most species in genus Anelpistina. 
Other characteristics that could further 
establish to whom the species is more 
closely related, such as the number of ma- 
crochaetae on mandibles and on the border 
of the insertion of antennae, regrettably 
could not be obtained from the illustrations 
of Wygodzinsky (1977). 

Anelpistina cuaxilotla, new species 
Figs. 3A-G, 4A-D 

Type material. — Type locality: Mexico. 
Guerrero: Apetlanca, Cuaxilotla town, in 
the penumbra zone (Entrance to 120 m) of 
"Cuaxilotla" cave, under rocks. 6 Jun 1987 
and 23 Aug 1987. L. Espinasa col. Male 
holotype, ten male paratypes and 12 female 
paratypes. Other localities: Mexico, More- 
los state, Tepoztlan municipality, San Juan 
town, "San Juan" or "Sistema Ferrocarril- 
Mina inferior" volcanic cave system, under 
rocks. 21 Jan 1979. R. Garcia col. One male 
paratype. 

Description. — Maximum body length 
12.0 mm. Maximum length of antennae 6.0 
mm, of caudal appendages 5.0 mm. When 
complete, antennae and caudal appendages 



VOLUME 112, NUMBER 1 



65 





MM 



am' J 










f 












\\\ 
AS 












\ 



f 
f, 




Fig. 3. A-C. Anelpistina boneti (Wygodzinsky). D-G. Anelpistina cuaxilotla n. sp.: A. Male, basal portion 
of antenna; B, Female, basal portion of antenna; C, Mandible. D, Male, basal portion of antenna; E. Female, 
basal portion of antenna; F, Mandible; G, Male, abdominal sternum VIII; H. Male from "San Juan" cave, genital 
area; I, Male from "Cuaxilotla" cave, genital area; J. Ovipositor and subgenital plate. 



66 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 







Fig. 4. Anelpistina cuaxilotla n. sp.: A, Male from "Cuaxilotla" cave, appendages of urostemite IV; B, Male 
from "San Juan" cave, appendages of urostemite IV; C, Male from "Cuaxilotla" cave, modified spines of 
cercus; D, Male from "San Juan" cave, modified spines of cercus. 



measure less than length of the body. Setae 
on body strong, long and very abundant. 
General color light yellow to white. 

Head with macrochaetae and microchae- 



tae similar to Fig. IF. Basal article of an- 
tennae in males slightly longer than pedi- 
cellus, but shorter than in A. boneti. In the 
female it is also shorter than in A. boneti 



VOLUME 112, NUMBER 1 



67 



(Fig. 3 A, B, D-E). Pedicellus of male as 
shown in Fig. 3D, with numerous clusters 
of unicellular glands. Female basal articles 
of antennae simple. 

Mouthpart appendages relatively long, 
very similar to A. boneti. Apex of maxillary 
palp with two conules, one longer than wid- 
er and the other wider than longer. Labial 
palp long, apical article barely longer than 
wide and only slightly longer than next to 
last article. This penultimate article with 
bulkiness with two macrochaetae. Labium 
and first article of labial palp with macro- 
chaetae. Mandibles without very small pegs 
on bigger tooth and chaetotaxy as in Fig. 
3F Legs as in A. boneti, with sclerotized 
macrochaetae on tibia. Claws of normal 
size. 

Cerci of male with a longer than wide 
basal article, second article wider than lon- 
ger and a very long third one bearing 4 
spines in distal portion, followed by nu- 
merous short articles of simple chaetotaxy. 
Spines consist of a very small one, a strong, 
subacute one, another very small, and a 
long, acute and slightly curved one (Fig. 
4C, D). Cerci of female simple. 

Abdominal sterna and terga as in other 
members of the genus. Setae long and nu- 
merous. Thorax with shorter setae than on 
urostema. Lateral borders as in A. boneti, 
with 4-5 macrochaetae. Urotergite X long 
and almost straight in both sexes, posterior 
angles with 2 + 2 macrochaetae and a few 
relatively strong setae. 

Appendages of urostemum IV of male 
very long and robust, acute on apex. Their 
longest diameter equals the length of the 
stylets of this segment (Fig. 4A, B). 

Urostemum VIII of male long and shal- 
low shallowly emarginate on the posterior 
margin (Fig. 3G), slightly deeper than in A. 
boneti. 

Urostemum IX of male with setae abun- 
dant. Behind insertion of parameres, in the 
center, with a small group of short, sclero- 
tized and spiniform setae. Internal face of 
coxal processes with irregular row of 5 spi- 
niform macrochaetae highly sclerotized 



(Fig. 31). In the only male individual col- 
lected from cave of San Juan (Sistema Fer- 
rocarril-Mina inferior), this row is of ten 
macrochaetae (Fig. 3H). It is not known if 
this difference is due to population differ- 
ences or because the individual of San Juan, 
being the biggest (12.0 mm) corresponds to 
an older instar. 

Stylets similar to A. boneti. Stylets IX 
bigger than the others. Penis and parameres 
as shown in Fig. 3H, I. Parameres attain the 
length of stylets IX. Surface of parameres 
with short setae. 

Subgenital plate of female rounded, with 
the apex slightly flat (Fig. 3J). Ovipositor 
surpassing apex of stylets IX by twice the 
length of stylets. Gonapophyses with ap- 
proximately 17 articles. 

Postembryonic development not very 
complex with younger instars almost iden- 
tical to older ones except for size. In male, 
appendages of urostemite IV appear very 
small at a length of approximately 6.0 mm 
and acquire adult morphology at a length of 
7.0 mm. In the longest individual (12.0 
mm) only a slight increase in the propor- 
tions of sexual secondary characters is ob- 
served. 

Female development not well known be- 
cause all females collected measure more 
than 7.5 mm, when length of ovipositor al- 
ready surpasses apex of stylets IX by one 
and a half to two times its size. 

Etymology. — Cuaxilotla. Makes refer- 
ence to the type locality, "Cuaxilotla" cave. 

Remarks. — The closest species to Anel- 
pistina cuaxilotla are A. bolivari, A. ano- 
phtalma, and A. boneti. In males of A. cuax- 
ilotla, urostemum VIII is shallowly emar- 
ginate on posterior margin, while in A. bo- 
livari, emargination is narrow and deep. 
Other characteristics that differentiate these 
two species are the spines in the cerci and 
the abundance of setae in terga and sterna. 
From A. anophtalma it differs because in 
this species, urotergite X is deeply emar- 
ginate on the posterior margin, appendages 
of urostemite IV are thin and ovipositor is 
short, while in A. cuaxilotla, urotergite X is 



68 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



almost straight, appendages are broad and 
ovipositor is long. For A. boneti, the closest 
species to A. cuaxilotla, the main difference 
is that in A. boneti the appendages of uros- 
temite IV are thin, while in A. cuaxilotla 
they are broad. Other characteristics that 
differentiate them are that in the former, the 
cerci have three spines (although some in- 
dividuals from "Iglesia-Mina superior" 
cave in Morelos have 4) while in A. cuax- 
ilotla they have four; the basal article of the 
antennae in males and females is propor- 
tionally longer than the pedicellus in A. bo- 
neti than it is in A. cuaxilotla (Fig. 3 A, B, 
D, E); and the chaetotaxy of the mandible 
(Figs. 3C and F). To establish these differ- 
ences with precision, a large number of in- 
dividuals of A. boneti collected from the 
type locality by the author were compared 
to the individuals of A. cuaxilotla. 

The somewhat overlapping distribution 
of A. cuaxilotla and A. boneti is intriguing. 
The latter was reported to a wide distribu- 
tion comprising the states of Guerrero, Mo- 
relos, D.F and even Habana island in the 
Gulf of California (Wygodzinsky 1946), 
while A. cuaxilotla has been found in Guer- 
rero and Morelos. Wygodzinsky (1946) ini- 
tially distrusted such a big geographic dis- 
tribution of A. boneti, although repeated 
studies failed to show differences between 
samples of different origins ("Desconfia- 
mos inicialmente da grande distribu^ao da 
especie, repetidos exames detalhados fal- 
haram de demostrar diferen^as entre os ex- 
emplares de procedencia diferente"). 

Regardless of whether organisms from 
the Gulf of California are indeed A. boneti, 
the author of the present paper has found a 
locality where both species can be found in 
close proximity. Individuals that clearly be- 
long to A. boneti have been collected in the 
cave system of the "Iglesia-Mina superior" 
(Morelos, Tepoztlan, San Juan). This cave 
system is only tens of meters away from the 
cave system of Ferrocarril-Mina inferior, 
where an individual of A. cuaxilotla was 
found. Individuals of both species from 
these two caves can clearly be differentiated 



morphologically, especially because sam- 
ples of both contain mature males. 

It is currently not known if the presence 
of four spines on the cerci in males of A. 
boneti from Iglesia-Mina superior cave, in- 
stead of three spines as it is the norm for 
this species, reflects variation within the 
species, or if it instead reflects local hybrid- 
ization between both species in these neigh- 
boring caves. 

Acknowledgments 

I wish to thank Dr. Jose G. Palacios Var- 
gas, director of "Laboratorio de Ecologia y 
Sistematica de Microarstropodos", where 
most of the descriptive work was done and 
to the Director of CEAMISH-UAEM, Dr. 
Oscar Dorado, for the support to publish 
this manuscript. Thanks are also due to Dr. 
Luis F. Mendes and Monika Baker for re- 
viewing the manuscript. 

Literature Cited 

Diamant, R., & R. Espinasa-Perena. 199L La cueva 
de Agua Brava. — Tepeyollotli: Gaceta de la So- 
ciedad Mexicana de Exploraciones Subterraneas 
5:46-48. 

Espinasa, L. 1991. Descripcion de una nueva especie 
del genero Cubacubana (Zygentoma: Nicoleti- 
idae) y registro del genero para America Con- 
tinental. — Folia Entomologica Mexicana 82:5- 
16. 

Espinasa-Perena, R. 1989. Proyecto Chilacachapa, re- 
porte del progreso. — Tepeyollotli: Gaceta de la 
Sociedad Mexicana de Exploraciones Subterra- 
neas 4:32-40. 

Hoffman, A., J. G. Palacios-Vargas, & J. B. Morales- 
Malacara. 1986. Manual de bioespeleologia. 
Universidad Nacional Autonoma de Mexico, 
Mexico, 274 pp. 

Mendes, L. E 1986. Nouvelles donnees sur le Zygen- 
toma (Insecta) de I'Amerique Centrale et du 
Mexique. — Bulletin du Museum national 
d'Histoire naturelle., Paris (4)8(A)(2):333-342. 

. 1992. Novos dados sobre os tisanuros (Micro- 

coryphia e Zygentoma) da America do Norte. — 
Garcia de Orta 16(1-2):171-193. 

Paclt, J. 1979. Neue Beitrage zur Kenntnis der Apter- 
ygoten-Sammlung des Zoologischen Instituts 
und Zoologischen Museums der Universitat 
Hamburg. VL Weitere Doppel- und Borstensch- 
wanze (Diplura: Campodeidae; Thysanura: Lep- 
ismatidae und Nicoletiidae). — Entomologische 



VOLUME 112, NUMBER 1 



69 



Mitteilungen aus dem zoologischen Museum 
Hamburg 6(105):221-228. 

Reddell, J. R. 1981. A review of the cavemicole fauna 
of Mexico, Guatemala and Belize. — Texas Me- 
morial Museum Bulletin 27:204-205. 

Silvestri, E 1905. Materiali per lo studio dei Tisanuri. 
VI. Tre nuove specie di Nicoletia appartenenti 
ad un nuovo sottogenero. — Redia (Firenze) 2: 
111-120. 

. 1933. Nuovo contribute alia conoscenza dei 

tisanuri de Mesico. — Bolletino del Laboratorio 
di Zoologia general e agraria di Portici 27:127- 
144. 



Wygodzinsky, R 1946. Sobre Nicoletia {Anelpistina) 
Silvestri 1905 e Prosthecina Silvestri, 1933. — 
Ciencia 7:15-25. 

. 1959. Contribution to the knowledge of the 

"Thusanura" and "Machilidae" (Insecta). — 
Revista Brasileira de Biologia 19(4):441-457. 

. 1973. Description of a new genus of cave 

Thysanura from Texas (Nuciletiidae, Thysanu- 
ra, Insecta). — American Museum Novitates 
2518:1-8. 

Wygodzinsky, R, & A. M. Hollinger. 1977. A study of 
Nicoletiidae from Cuba (Thysanura). — Resul- 
tats des Expeditions Biospeleologiques Cubano- 
Roumaines a Cuba 2:313-324. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):70-93. 1999. 

Anacroneuria from northeastern South America 
(Insecta: Plecoptera: Perlidae) 

Bill P. Stark 
Department of Biology, Mississippi College, Clinton, Mississippi 39058, U.S.A. 

Abstract. — Descriptions are given for ten new species oi Anacroneuria from 
Venezuela, Guyana, and Suriname, and records are given for ten species pre- 
viously described from this region. The following 10 new species are described 
(type locality in parenthesis): Anacroneuria achagua (Venezuela. Portuguesa: 
Guanare), A. arawak (Suriname. Brokopondo: Brownsberg Naturpark, Maza- 
roni Plateau), A. chaima (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las Pie- 
dras de Cocollar), A. claudiae (Venezuela. Zulia: Rio Yasa (3 km E Kasmera), 
Estacion Biologica), A. karina (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las 
Piedras de Cocollar), A. makushi (Guyana. Kanuku Mountains, Moco Moco 
River), A. paria (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las Piedras de 
Cocollar), A. perija (Venezuela. Zulia: Parque Nacional Perija, Rio Negro, To- 
romo), A. timote (Venezuela. Tachira: Quebrada Los Mirtos, 8 km S El Cobre), 
and A. wapishana (Guyana. Potaro River, Kaieteur Falls). Anacroneuria phan- 
toma (Banks) and A. pictipes Klapalek are redescribed and a modified key to 
regional males is presented. 



Stark (1995) provided descriptions of Ve- 
nezuelan Anacroneuria based on collec- 
tions made primarily through 1985. Re- 
cently a series of Venezuelan and Guyanan 
specimens collected in 1994-97 were made 
available by O. S. Flint and R. Holzenthal, 
and a few specimens were provided by the 
California Academy of Science. This ma- 
terial includes nine new species, new re- 
cords for eight previously described spe- 
cies, and a few unassociated females de- 
scribed under informal designations. Also 
included are redescriptions of A. pictipes 
Klapalek and A. phantoma (Banks) from 
holotypes and an additional new species 
from Suriname. Holotypes are deposited in 
the National Museum of Natural History, 
Washington, D.C. (USNM) or the Califor- 
nia Academy of Science, San Francisco, 
CA (CAS). Paratypes and other specimens 
are deposited in the Universidad Central de 
Venezuela, Maracuy (UCV), the University 
of Minnesota, Saint Paul (UMSP) or the 
collection of the author (BPS). Additional 



material was examined from the Museum 
of Comparative Zoology, Harvard Univer- 
sity (MCZ) and the National Museum of 
Natural History, Prague (NMH). 

Anacroneuria achagua, new species 

Adult habitus. — Head with an obscure 
brown pattern over anterior half of frons; 
lappets brown. Median pronotal stripe pale, 
broad irregular lateral bands brown (Fig. 1). 
Wing membrane transparent, veins pale. 

Male. — Fore wing length 8 mm Hammer 
thimble shaped, height greater than basal 
diameter (Fig. 2). Aedeagal apex short, with 
wide, emarginate tip offset from shoulders; 
ventral aspect with an obscure pair of mem- 
branous lobes; dorsal keel triangular; shoul- 
ders somewhat quadrate, hooks stout (Figs. 
3-5). 

Female. — ^Unknown. 

Nymph. — Unknown. 

Etymology. — The name honors the 



VOLUME 112, NUMBER 1 



71 







1 



|.t rV 1 . 1; v.A \y>-;.v i oi ■■'/■A./ ^■;-, 



■V:i> 



2 







Figs. 1-5. A. achagua structures. 1. Head and pronotum. 2. Male sternum 9. 3. Aedeagus, ventral. 4. Ae- 
deagus, lateral. 5. Aedeagus, dorsal. Scales: 0.6 mm (1), 0.3 mm (2), 0.15 mm (3-5). 



72 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 6— H. A. arawak structures. 6. Head and pronotum. 7. Female sterna 8 and 9. 8. Male sternum 9. 9. 
Aedeagus, ventral. 10. Aedeagus, lateral. 11. Aedeagus, dorsal. Scales: 0.6 mm (6), 0.3 mm (7, 8), 0.15 mm 
(9-11). 



Achagua people of Venezuela and is used 
as a noun in apposition. 

Types. — Holotype S and S paratype 
from Venezuela, Portuguesa, Guanare, 10- 
13 Sep 1957, B. Malkin (CAS). 

Diagnosis. — This species has a V-shaped 
aedeagal keel and apex somewhat like that 
of A. pictipes but these species are easily 
distinguished by the distinctive wing and 
pronotal pigmentation of that species (Figs. 
48, 53). 



Anacroneuria arawak, new species 

Adult habitus. — Head with diffuse yel- 
low brown area extending forward from 
ocelli; pronotum with diffuse brown lateral 
bands and a narrow pale median band (Fig. 
6). Wing membrane transparent, veins pale. 

Male. — Fore wing length 9 mm. Hammer 
thimble shaped, height subequal to basal di- 
ameter (Fig. 8). Aedeagal apex slender, 
scoop shaped with a pair of subapical ven- 



VOLUME 112, NUMBER 1 



73 



tral membranous lobes. Apex curved slight- 
ly dorsad; hooks slender, dorsal keel long 
and narrow (Figs. 9-11). 

Female. — Forewing length 10.5 mm. Su- 
bgenital plate four lobed, median lobes 
shorter than lateral lobes; inner margins of 
lateral lobes bearing a short setal fringe. 
Sternum 9 with a broad transverse sclerite 
and a trilobed median setal patch; median 
lobe clothed with minute setae, lateral lobes 
with prominent setae (Fig. 7). 

Nymph. — Unknown . 

Etymology. — The name honors the Ara- 
wak people of Guyana and Suriname and is 
used as a noun in apposition. 

Types. — Holotype S and ? paratype 
from Suriname, Brokopondo District, 
Brownsberg Naturpark, Mazaroni Plateau, 
400-500 m, 16 Aug 1982, W. E. Steiner 
(USNM). 

Diagnosis. — This species does not ap- 
pear to be closely related to other regional 
Anacroneuria. The aedeagal shape and pale 
habitus are somewhat similar to A. isleta 
(Stark 1994) but that species has a trian- 
gular keel and a shorter and more rounded 
aedeagal tip. In dorsal aspect the aedeagus 
is similar to that of A. bari (Stark 1995), 
but that species does not have membranous 
ventral lobes on the aedeagus. The female 
is distinctive by virtue of the setal fringe on 
the lateral subgenital plate lobes (Fig. 7). 

Anacroneuria aroucana Kimmins 

Anacroneuria aroucana Kimmins, 1948: 
105. Holotype S , Arouca River, Trinidad. 
Anacroneuria aroucana: Stark, 1994:173. 

Material. — Venezuela: Sucre, Quebrada 
Zapateral, 1.5 km SE Las Piedras de Co- 
collar, 810 m, 9 Apr 1995, R. Holzenthal, 
O. S. Flint, 4 S (USNM, UCV). Sucre, Rio 
Cocollar, 1.5 km SE Las Piedras de Cocol- 
lar; 810 m, 7-8 Apr 1995, R. Holzenthal, 
O. S. Flint, 10 S (USNM). 

Comments. — These are the first mainland 
records for this species. 



Anacroneuria bari Stark 

Anacroneuria bari Stark, 1995:226. Holo- 
type S, El Tucuco, 45 km SW Mac- 
hiques, Zulia, Venezuela. 

Material. — Venezuela: Guarico, Parque 
Nacional Guatopo, Quebrada Guatopo, 0.5 
km NE La Colina, 600 m, 22 Jan 1994, R. 
Holzenthal, C. Cressa, Rincon, 3 S (UMSP, 
UCV). Trujillo, Quebrada Potrerito, 7.5 km 
NE Bocono, 1530 m, 29-30 Apr 1995, R. 
Holzenthal, C. Cressa, Gutic, 2 S (BPS). 

Comments. — This species was previously 
reported from the Venezuelan states of Bar- 
inas, Miranda, and Zulia (Stark 1995). 

Anacroneuria bifasciata (Pictet) 

Perla bifasciata Pictet, 1841:231. Holotype 

?, Moritz, Colombia. 
Anacroneuria bifasciata: Zwick, 1972: 

1154. 
Anacroneuria bifasciata: Stark, 1995:239. 

Material. — Venezuela: Distrito Federal, 
Rio Camuri Grande, 1 km S Camuri (nu- 
cleo USB), 30 m, 24 Jan 1994, R. Holzen- 
thal, C. Cressa, Rincon, 1 c?, 1 $ (BPS). 

Comments. — This species was previously 
reported from the Venezuelan states of Ar- 
agua, Distrito Federal, and Lara (Stark 
1995). 

Anacroneuria caraca Stark 

Anacroneuria caraca Stark, 1995:228. Ho- 
lotype (?, Rio Limon, Parque Nacional 
Henri Pittier, Aragua, Venezuela. 

Material. — Venezuela: Guarico, Parque 
Nacional Guatopo, Quebrada Guatopo, 0.5 
km NE La Colina, 600 m, 22 Jan 1994, R. 
Holzenthal, C. Cressa, Rincon, 1 S 
(UMSP). 

Comments. — This species was previously 
reported from the Venezuelan states of Ar- 
agua and Barinas (Stark 1995). 

Anacroneuria chaima, new species 

Adult habitus. — Head with a wide brown 
area forward of ocelli extending to M-line; 



74 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



lappets and anterior margin of head brown. 
Median pronotal stripe pale; broad lateral 
stripes dark brown (Fig. 12). Wing mem- 
brane and veins brown. 

Male. — Forewing length 8 mm. Hammer 
cylindrical, height greater than basal diam- 
eter (Fig. 13). Aedeagal apex simple, scoop 
shaped, slender and rounded at the tip; ven- 
tral aspect with a small circular pair of 
membranous lobes; dorsal aspect with a 
low wide keel; hooks slender (Figs. 14-16). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — The name honors the Chai- 
ma people of Venezuela and is used as a 
noun in apposition. 

Types. — Holotype S (pinned) from Ven- 
ezuela, Sucre, Rio CocoUar, 1.5 km SE Las 
Piedras de Cocollar, 810 m, 7-8 Apr 1995, 
R. Holzenthal, O. S. Flint (USNM). 

Diagnosis. — This species is similar to A. 
perija in general aedeagal structure but dif- 
fers in having small, circular membranous 
aedeagal lobes (Fig. 14) and in having the 
arms of the dorsal aedeagal keel divergent 
(Fig. 16). The dark color pattern of A. per- 
ija is also much more prominent, particu- 
larly over the occiput (Fig. 38). 

Anacroneuria chorrera Stark 

Anacroneuria chorrera Stark, 1995:230. 
Holotype S , La Chorrera Canyon, Meri- 
da, Venezuela. 

Material. — Venezuela: Merida, Rio La 
Gonzalez, between Merida and Jaji, 1870 
m, 25 Apr 1995, R. Holzenthal, C. Cressa, 
Gutic, 3 S (UCV). Merida, Rio Albarregas, 
1 km NW Universidad de los Andes, 1980 
m, 17 Jan 1994, R. Holzenthal, C. Cressa, 
Rincon, 3 S (UMSP). Merida, Parque Na- 
cional Sierra Nevada, Quebrada La Mucuy, 
7 km E Tabay, 2200 m, 18 Jan 1994, R. 
Holzenthal, C. Cressa, Rincon, 4 S (BPS). 
Trujillo, Quebrada Potrerito, 7.5 km NE 
Bocono, 1530 m, 20-30 Apr 1995, R. Hol- 
zenthal, C. Cressa, Gutic, 4 S (UMSP). 
Tachira, tributary to El Valle, 3.8 km SE El 
Zumbador, 2730 m, 21 Apr 1995, R. Hol- 



zenthal, C. Cressa, Gutic, 7 c^ (UMSP). 
Tachira, Quebrada Los Mirtos, 8 km S El 
Cobre, 2400 m, 22 Apr 1995, R. Holzen- 
thal, C. Cressa, Gutic, 1 S (UCV). 

Comments. — This species was previously 
reported from the Venezuelan states of Ar- 
agua, Distrito Federal, and Merida (Stark 
1995). 

Anacroneuria claudiae, new species 

Adult habitus. — Head yellow with dif- 
fuse brown pattern extending from ocelli to 
anterior margin but interrupted by three 
pale areas at M-line; lappets brown. Median 
pronotal stripe pale, lateral stripes dark, an- 
terolateral margins with pale spot (Fig. 17). 
Wing membrane transparent, veins brown. 

Male. — ^Forewing length 8 mm. Hammer 
thimble shaped, height greater than basal 
diameter (Fig. 18). Aedeagal apex simple, 
scoop shaped, slender and truncate at the 
tip; ventral aspect with a small pair of mem- 
branous lobes; dorsal aspect with a narrow 
keel, hooks slender (Figs. 19-21). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — The matronym honors 
Claudia Cressa of the Universidad Central 
de Venezuela, Caracas. 

Types. — Holotype S from Venezuela, 
Zulia, Rio Yasa, 3 km E Kasmera (Estacion 
Biologica), 150 m, 14 Jan 1994, R. Holzen- 
thal, C. Cressa, Rincon (USNM). Paratype 
S from Venezuela, Zulia, Parque Nacional 
Perija, Rio Negro, Toromo, 360 m, 15 Jan 
1994, R. Holzenthal, C. Cressa, Rincon 
(UCV). 

Diagnosis. — This species is similar to A. 
chiquita in general aedeagal structure but 
differs from that species in having small 
paired membranous ventral lobes (Fig. 17) 
and in having unmodified aedeagal hook 
apices (Fig. 17). 

Anacroneuria cruza Stark 

Anacroneuria cruza Stark, 1995:231. Ho- 
lotype S , Exp. Culebra, N Duida, Terri- 
torio Federal Amazonas, Venezuela. 



VOLUME 112, NUMBER 1 



75 




■\« 






mm/M 



V/.: 



1 -K' 



Vt^A 



mi 



■'.y.'.-i 



^■^/m 






r-i-fi 



13 



t:s 





Figs. 12-16. A. chaima structures. 12. Head and pronotum. 13. Male sternum 9. 14. Aedeagus, ventral. 15. 
Aedeagus, lateral. 16. Aedeagus, dorsal. Scales: 0.6 mm (12), 0.3 mm (13). 0.15 mm (14-16). 



76 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




17 



■ ■ .1 • *'■■ .-..•> 







18 



^"4 





Figs. 17-2L A. Claudia structures. 17. Head and pronotum. 18. Male sternum 9. 19. Aedeagus, ventral. 20. 
Aedeagus, lateral. 21. Aedeagus, dorsal. Scales: 0.6 mm (17), 0.3 mm (18), 0.15 mm (19-21). 



VOLUME 112, NUMBER 1 



77 



Material. — Guyana: Kanuku Mountains, 
Kumu River, 28-30 Apr 1995, O. S. Flint, 
2 6 (USNM). Kanuku Mountains, Moco 
Moco River, 29 Apr 1995, O. S. Flint, 1 S 
(USNM). Venezuela: Sucre, Rio Cocollar, 
1.5 km SE Las Piedras de Cocollar, 810 m, 
7-8 Apr 1995, R. Holzenthal, O. S. Flint, 
4 c^ (USNM, UCV). 

Comments. — This species was previously 
known from Territorio Federal Amazonas, 
Venezuela (Stark 1995). 

Anacroneuria karina, new species 

Adult habitus. — Head yellow with a dif- 
fuse brown spot forward of ocelli; lappets 
brown. Irregular midlateral pronotal bands 
dark brown, median field pale, grading to 
pale brown (Fig. 22). Wing membrane pale 
brown, veins brown, R and Sc dark brown. 

Male. — Forewing length 11 mm. Ham- 
mer thimble shaped, height greater than 
basal diameter (Fig. 23). Aedeagal apex 
simple, scoop shaped and broadly rounded; 
ventral membranous lobes narrow; dorsal 
keel well developed; hooks slender (Figs. 
24-26). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — The name honors the Kar- 
ina people of Venezuela and is used as a 
noun in apposition. 

Types. — Holotype S (USNM) and 7 S 
paratypes (USNM, UCV) from Venezuela, 
Sucre, Rio Cocollar, 1.5 km SE Las Piedras 
de Cocollar, 810 m, 7-8 Apr 1995, R. Hol- 
zenthal, O. S. Flint. 

Diagnosis. — Many species of Anacro- 
neuria share the color pattern of A. karina 
and the aedeagus also bears a general sim- 
ilarity to that of other species such as A. 
bari and A. arcuata (Stark 1995). Anacro- 
neuria karina, however, lacks the transverse 
dorsal aedeagal keel found in A. arcuata 
and the aedeagal apex is much longer and 
narrower in A. bari. 

Anacroneuria liana Stark 

Anacroneuria liana Stark, 1995:234. Ho- 
lotype (5, La Escalera, 108 km S Rio Cu- 
yuni, Bolivar, Venezuela. 



Material. — Guyana: Potaro River, Kaie- 
teur Falls, 1350', 21-23 Aug 1997, O. S. 
Flint, 9 c^, 58 $ (USNM). 

Comments. — This species was previously 
known from the holotype. The Guyana 
males agree with the holotype in all re- 
spects except in having falcate aedeagal 
hooks. The females collected with these 
males share the same color pattern (Fig. 63) 
and are associated on that basis. These fe- 
males have eggs with a terminal spine sim- 
ilar to those of A. blanca (Stark 1995) and 
they also are similar in color pattern, size 
and in having an intersegmental band of mi- 
crotrichia. 

Female. — Forewing length 13-14 mm. 
Subgenital plate four lobed, outer lobes 
slightly longer. Median sclerite of sternum 
9 densely hirsute, hairs slightly longer lat- 
erally. Intersegmental membrane covered 
with a dense microtrichia band (Fig. 64). 

Anacroneuria makushi, new species 

Adult habitus. — Head yellow; pronotum 
with diffuse midlateral brown bands (Fig. 
27). Wing membrane transparent, veins 
pale. 

Male. — Forewing length 9.5 mm. Ham- 
mer laterally compressed, height greater 
than basal diameter (Fig. 28). Aedeagal 
apex scoop shaped, tip blunt; ventral aspect 
with a pair of subapical membranous lobes; 
dorsal aspect with a long, low keel; dorsal 
sclerite basal to keel bearing an arcuate pro- 
cess; hooks irregularly scalloped on inner 
margin (Figs. 29-31). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — The name honors the Mak- 
ushi people of Guyana and is used as a 
noun in apposition. 

Types. — Holotype S (pinned) and 2 S 
paratypes from Guyana, Kanuku Moun- 
tains, Moco Moco River, 29 Apr 1995, O. 
S. Flint (USNM). 

Diagnosis. — A. makushi is similar to A. 
baniva in aedeagal structure (Stark 1995). 
The most conspicuous difference between 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 22-26. A. karina structures. 22. Head and pronotum. 23. Male sternum 9. 24. Aedeagus, ventral. 25. 
Aedeagus, lateral. 26. Aedeagus, dorsal. Scales: 0.6 mm (22), 0.3 mm (23), 0.15 mm (24-26). 



VOLUME 112, NUMBER 1 



79 




Figs. 27-31. A. makushi structures. 27. Head and pronotum. 28. Male sternum 9. 29. Aedeagus, ventral. 30. 
Aedeagus, lateral. 31. Aedeagus, dorsal. Scales: 0.6 mm (27), 0.3 mm (28), 0.15 mm (29-31). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



these species is the arcuate process on the 
dorsal aedeagal sclerite in A. makushi (Fig. 
31). 

Anacroneuria paleta Stark 

Anacroneuria paleta Stark, 1995:236. Ho- 
lotype S , 4 km S Santo Domingo, Mer- 
ida, Venezuela. 

Material. — Venezuela: Trujillo, Quebra- 
da Potrerito, 7.5 km NE Bocono, 1530 m, 
29-30 Apr 1995, R. Holzenthal, C. Cressa, 
Gutic, 31 S (UMSP, UCV, BPS). 

Comments. — This species was previously 
recorded from the Venezuelan states of Bar- 
inas and Merida (Stark 1995). 

Anacroneuria paria, new species 

Adult habitus. — Very diffuse brown pat- 
tern over ocelli and midlaterally on prono- 
tum (Fig. 32). Wing membrane pale with 
distinctive longitudinal brown band extend- 
ing from anal cell to tip; band narrow in 
proximal half, expanded at the cord, but in- 
terrupted by a large circular "window" be- 
yond the cord (Fig. 37). 

Male. — Fore wing length 10 mm. Ham- 
mer thimble shaped, height less than basal 
diameter (Fig. 33). Aedeagal apex with five 
dorsal lobes and a short narrow keel; ven- 
tral aspect of mesal lobe scoop shaped, 
without membranous lobes; hooks slender 
(Figs. 34-36). 

Female. — Forewing length 12 mm. Su- 
bgenital plate four lobed; lateral lobes larg- 
er than inner lobes, mesal notch deep and 
V-shaped. Transverse sclerite of sternum 9 
wide, sinuate and prominent, mesal sclerite 
T-shaped; lateral lobes of mesal sclerite 
with prominent setae, stalk covered with 
minute setae (Fig. 54). 

Nymph. — Unknown. 

Etymology. — The name is based on the 
Paria Peninsula and is used as a noun in 
apposition. 

Types. — Holotype S (pinned) and 9 ? 
paratypes from Venezuela, Sucre, Rio Co- 
collar, 1.5 km SE Las Piedras de Cocollar, 



810 m, 7-8 Apr 1995, R. Holzenthal, O. S. 
Flint (USNM). Additional paratypes; Ven- 
ezuela: Sucre, Parque Nacional Peninsula 
de Paria, Uquire, Rio La Vidua, 15 m, 30 
Mar-1 Apr 1995, R. Holzenthal, O. S. Flint, 
C. Cressa, 3 ? (UCV). Sucre, Peninsula de 
Paria, Puerto Viejo, "Rio el Pozo", 20 m, 
3 Apr 1995, R. Holzenthal, O. S. Flint, C. 
Cressa, 6 ? (USNM). 

Diagnosis. — This species is similar to A. 
bifasciata (Pictet) and A. vistosa Stark in 
displaying a distinctive wing pigmentation 
pattern, however A. paria is readily distin- 
guished from both species on the basis of 
color pattern, aedeagal shape and details of 
female sternum 9 (Stark 1995). Anacroneu- 
ria bifasciata adults have tiny ocelli, a 
prominent dark mesal pronotal band and the 
wing pigmentation is separated into three 
distinct transverse bands. The aedeagal 
apex of A. bifasciata also bears five dorsal 
lobes but the mesal lobe lacks a keel and 
the mesal sclerite of the female 9th sternum 
is triangular and evenly setose. The aedea- 
gal apex of A. vistosa is trilobed and the 
mesal sclerite of the female 9th sternum is 
evenly setose. 

Anacroneuria perija, new species 

Adult habitus. — Center of frons bright 
yellow, anterior head margin, posterolateral 
head margins, and ocellar area dark brown. 
Median pronotal stripe pale, lateral third 
brown (Fig. 38). Wing membrane brown, 
veins dark brown, Sc pale. 

Male. — Forewing length 9 mm. Hammer 
cylindrical, height greater than basal diam- 
eter (Fig. 39). Ventral aedeagal apex a short 
scoop covered with a pair of membranous 
lobes; dorsal aspect with a wide keel and a 
low U-shaped region; hooks slender (Figs. 
40-42). 

Female. — Unknown. 

Nymph. — Unknown . 

Etymology. — The name is based on the 
type locality and is used as a noun in ap- 
position. 

Types. — Holotype S from Venezuela, 



VOLUME 112, NUMBER 1 



81 




Figs. 32-37. A. paria structures. 32. Head and pronotum. 33. Male sternum 9. 34. Aedeagus. ventral. 35. 
Aedeagus, lateral. 36. Aedeagus, dorsal. 37. Forewing. Scales: 1.2 mm (37), 0.6 mm (32), 0.3 mm (33), 0.15 
mm (34-36). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




':-V. 



"H^ 






















39 





Figs. 38-42. A. perija structures. 38. Head and pronotum. 39. Male sternum 9. 40. Aedeagus, ventral. 41. 
Aedeagus, lateral. 42. Aedeagus, dorsal. Scales: 0.6 mm (38), 0.3 mm (39), 0.15 mm (40-42). 



VOLUME 112, NUMBER 1 



83 



Zulia, Parque Nacional Perija, Rio Negro, 
Toromo, 360 m, 15 Jan 1994, R. Holzen- 
thal, C. Cressa, Rincon (USNM). 

Diagnosis. — This species keys to A. ban- 
iva in Stark (1995) but the aedeagal apex is 
much shorter and narrower than in that spe- 
cies, and the dorsal keel and subapical U- 
shaped process of A. perija also will distin- 
guish the two. Although the condition of A. 
baniva type material left much of the color 
pattern obscured, it is clear that species 
lacks the distinctive head pattern of A. per- 
ija. 

Anacroneuria phantoma (Banks) 

Neoperla phantoma Banks, 1914:609. Ho- 
lotype S , Mallali, Guyana. 

Adult habitus. — Head yellow with dif- 
fuse yellow brown lappets. Pronotum with 
pale median stripe and broad, diffuse brown 
midlateral bands; lateral margins pale (Fig. 
43). Wing membrane and veins pale. 

Male. — Forewing length 9—10 mm. 
Hammer laterally compressed, height great- 
er than basal diameter (Fig. 44). Aedeagal 
apex bluntly pointed, shoulders undevel- 
oped. Dorsal keel long, ventral membra- 
nous lobes large. Hooks scalloped along in- 
ner margins (Figs. 45-47). 

Female. — Unknown. 

Nymph. — Unknown . 

Material. — Guyana: Mallali, Mar, H. S. 
Parish, 1 S (holotype) (MCZ). Potaro Riv- 
er, Kaieteur Falls, 1350', 21-23 Aug 1997, 
O. S. Flint, 4 S (USNM). 

Diagnosis. — This species is quite similar 
to and perhaps synonymous with A. baniva 
(Stark 1995). The subtle differences in ae- 
deagal structure include a small apical 
notch, decurved apex and low dorsal keel 
in A. baniva. The holotype of A. phantoma 
has a blunt aedeagal apex, straight in lateral 
aspect and a more pronounced dorsal keel 
(Figs. 45—47). Anacroneuria makushi rep- 
resents another member of this complex 
with a more robust aedeagal apex and a 
transverse arcuate keel (Figs. 29-31). 

Comments. — The type locality above 



Linden on the Demerara River is one of the 
H. S. Parish sites of 1912-13 (Adams, pers. 
comm.). 

Anacroneuria pictipes Klapalek 

Anacroneuria pictipes Klapalek, 1923:21. 
Holotype 9, Haut-Carsevenne, Guyana. 

Adult habitus. — (Modified from Klapa- 
lek, 1923) Body ochre yellow, pronotum 
darker brown along the posterior margins 
and near lateral margins (Fig. 48). Wing 
membrane and veins brown; R dark brown, 
costal border pale (Fig. 53). 

Male. — Forewing length 10 mm. Ham- 
mer laterally compressed at tip, height 
greater than basal diameter (Fig. 49). Ae- 
deagal apex truncate to emarginate, arising 
from low broadly rounded shoulders; dorsal 
keel triangular, hooks slender (Figs. 50-52). 

Female. — Forewing length 11 mm. Su- 
bgenital plate weakly four-lobed. Median 
field of sternum nine weakly sclerotized 
and sparsely hirsute. Posterior margin of 
nine without transverse sclerite (Fig. 55). 

Nymph. — Unknown. 

Material. — Guyana [French Guiana ?]: 
Haut-Carsevenne, 1878, F Geay, 1 $ (ho- 
lotype) (NMH). Guyana: Kanuku Moun- 
tains, Moca Moca River, 29 Apr 1995, O. 
S. Flint, 2 6 (USNM). Paramaketoi Village, 
2350', 24-25 Aug 1997, O. S. Hint, 3 S 
(USNM). 

Comments. — This association of males 
with the holotype female is based on the 
distinctive wing pigmentation pattern. Fe- 
males collected with the males at Moca 
Moca River represent another species, de- 
scribed below as "GU-1" which lacks this 
wing pattern. 

Anacroneuria timote, new species 

Adult habitus. — Head yellow with dif- 
fuse brown area forward of ocelli and M- 
line; lappets brown. Broad median pronotal 
stripe pale, irregular midlateral stripes 
brown, lateral margins pale (Fig. 56). Wing 
membrane transparent, C and Sc veins pale, 
R vein dark brown. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 43-47. A. phantoma structures. 43. Head and pronotum. 44. Male sternum 9. 45. Aedeagus, ventral. 
46. Aedeagus, lateral. 47. Aedeagus, dorsal. Scales: 0.6 mm (43), 0.3 mm (44), 0.15 mm (45-47). 



VOLUME 112, NUMBER 1 



85 




Figs. 48-53. A. pictipes structures. 48. Head and pronotum. 49. Male sternum 9.50. Aedeagus, ventral. 51. 
Aedeagus, lateral. 52. Aedeagus, dorsal. 53. Forewing. Scales: 1.2 mm (53), 0.6 mm (48), 0.3 mm (49), 0.15 
mm (50-52). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 







1 . r 



54 










Figs. 54-55. Anacroneuria female sterna 8 and 9. 54. A. paria. 55. A. pictipes. Scale: 0.3 mm. 



VOLUME 112, NUMBER 1 



87 








Figs. 56-60. A. timote structures. 56. Head and pronotum. 57. Male sternum 9. 58. Aedeagus, ventral. 59. 
Aedeagus, lateral. 60. Aedeagus, dorsal. Scales: 0.6 mm (56), 0.3 mm (57), 0.15 mm (58-60). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Male. — Fore wing length 17 nun. Ham- 
mer thimble shaped, height less than basal 
diameter (Fig. 57). Aedeagal apex simple, 
scoop shaped with broadly rounded tip; 
ventral aspect with an inconspicuous pair of 
membranous lobes; dorsal keel well devel- 
oped, hooks slender (Figs. 58-60). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — The name honors the Tim- 
ote people of Venezuela and is used as a 
noun in apposition. 

Types. — Holotype 6 (USNM) and 4 (? 
paratypes (UCV, UMSP) from Venezuela, 
Tachira, Quebrada Los Mirtos, 8 km S El 
Cobre, 2400 m, 22 Apr 1995, R. Holzen- 
thal, C. Cressa, Gutic. 

Diagnosis. — The head and pronotal pat- 
tern, size, and general aedeagal structure of 
this species is similar to A. shamatari (Stark 
1995). The most conspicuous difference in 
aedeagal structure involves the membra- 
nous area between the hooks. In A. sha- 
matari this area terminates well short of the 
aedeagal base whereas in A. timote it ex- 
tends almost to the aedeagal base (Fig. 58). 

Anacroneuria wapishana, new species 

Adult habitus. — Head dark brown except 
for anterior margin and callosities. Prono- 
tum dark brown except for V-shaped me- 
dian band and small anterolateral areas 
(Fig. 61). Fore and mid femora and tibiae 
dark brown; hind femora yellow in basal 
two thirds, dark brown apically. Wing 
membrane brown except for transparent 
window at cord; veins dark brown except 
for pale costal area; wing tips dark brown. 

Male. — Unknown. 

Fem^ale. — Forewing length 11 mm. Su- 
bgenital plate four lobed; outer lobes small, 
notches shallow; median notch deep. Mesal 
sclerite of sternum 9 with short broad me- 
dian field of fine setae, lateral fields with 
long thick bristles; transverse posterior 
sclerite indistinct (Fig. 62). 

Nymph. — Unknown. 

Etymology. — The name honors the Wap- 



ishana people of Guyana and is used as a 
noun in apposition. 

Types. — Holotype 2 from Guyana, Po- 
taro River, Kaieteur Falls, 1350', 21-23 
Aug 1997, O. S. Flint (USNM). 

Diagnosis. — The exceptional color pat- 
tern of the holotype is the basis for naming 
this species from a female. The species 
bears some similarity to A. dourada (Jewett 
1960) but is much smaller and has a more 
extensive area of dark pigment on the frons. 

Unassociated Females 
Anacroneuria GU-1 

Adult habitus. — Head yellow, without 
dark pattern. Pronotum with irregular 
brown midlateral bands and a broad yellow 
median band. Wing membrane transparent, 
veins pale. 

Female. — ^Forewing length 11 mm. Su- 
bgenital plate with four subequal lobes. Me- 
sal sclerite of sternum 9 with broad median 
field of short, fine setae, lateral field with 
longer thick bristles; posterior margin with 
a narrow U-shaped median sclerite (Fig. 
65). 

Material. — Guyana: Kanuku Mountains, 
Moco Moco River, 29 Apr 1995, 4 $, O. 
S. Flint (USNM). Kanuku Mountains, 
Kumu River and faUs, 28-30 Apr 1995, 1 
$, O. S. Flint (USNM). 

Anacroneuria GU-2 

Adult habitus. — Head yellow brown, lap- 
pets pale. Pronotum with diffuse brown 
midlateral bands. Wing membrane transpar- 
ent, veins pale. 

Female. — Forewing length 11 mm. Su- 
bgenital plate four lobed; lateral lobes wide 
and offset from narrow median lobes by 
shallow acute notches. Mesal sclerite of 
sternum 9 with median field covered with 
fine short setae which form a sagittate patch 
separated from lateral patches covered with 
longer bristles; posterior sclerite absent 
(Fig. 66). 

Material. — Guyana: Dubulay Ranch, Ar- 



VOLUME 112, NUMBER 1 



89 








61 



62 





\ ■',' 



.(-,■<• 



,*-■;■ 



■'■■'r-v. 



■'.IV- 



•/■-^■If 






<y.//> 



.[■•..•- 



M 









.l-..t. ^. 



/'tZ/t- 






^/, 






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:/"-•/• 



■) .(• 



■.r; 



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I:' 



^.4' 



^- • ^-z* M^. i: /■ I 



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,V 



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Figs. 61-64. Anacroneuria female structures. 61. A. wapishana head and pronotum. 62. A. wapishana sterna 
8 and 9. 63. A. liana head and pronotum. 64. A. liana sterna 8 and 9. Scales: 0.6 mm (61, 63), 0.3 mm (62, 
64). 



amatani Creek, 15-18 Apr 1995, 2 9, O. S. 
Flint (USNM). 

Anacroneuria GU-3 

Adult habitus. — Head yellow with dif- 
fuse brown pigment forward of ocelli; lap- 
pets brown. Pronotum with irregular diffuse 
brown spots scattered over midlateral area 
(Fig. 67). Tibiae banded, basal and apical 



bands brown, median band pale. Wing 
membrane transparent except for pale 
brown tips which accent an obscure win- 
dow at the cord; veins brown, R darker, cos- 
tal area pale. 

Female. — Fore wing length 10 mm. Su- 
bgenital plate four lobed; outer lobes small, 
notches shallow, median notch deep and U- 
shaped. Median sclerite of sternum 9 with 



90 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 










65 







66 



Figs. 65-66. Anacroneuria female sterna 8 and 9. 65. Anacroneuria GU-L 66. Anacroneuria GU-2. Scale: 
0.3 mm. 



VOLUME 112, NUMBER 1 



91 







m. 



i.v../.' 






rn 



■n 



m 



'(^M'^'M. 






.\^-'- 






Hiift 



■ifi:-i:-'!':^-;^Mi^ 



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T- 






i f 






^/■■' 



-rtr^Kfil'f 



■I; 



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4 



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'^(M 



Figs. 67-68. Anacroneuria GU-3 female structures. 67. Head and pronotum. 68. Sterna 8 and 9. Scales: 0.6 
(67), 0.3 (68). 



long narrow stem; stem and mesal field 
with short, fine setae, lateral areas with 
thicker setae. Transverse sclerite sinuate 
(Fig. 68). 

Material. — Guyana: Mazaruni-Potaro 
District, Takutu Mountains, 20 Dec 1983, P. 
J. Spangler, 1 9 (USNM). Same location, 
12 Dec 1983, P J. Spangler, R. A. Faitoute, 
R D. Perkins, 1 $ (USNM). 

Provisional Key for Male Anacroneuria 

from Venezuela, Guyana, and Suriname 

(Modified from Stark 1995) 

1 . Wings banded in amber and dark brown 

2 

Wings variable, but without bands ... 3 

2. Dark pigment separated into basal, me- 
dian and apical bands A. bifasciata 

Dark pigment forming a narrow lon- 
gitudinal band from base to cord (Fig. 
37) A. paria 

3. Forewing length greater than 13.5 mm 

4 

Forewing length less than 1 3 mm . . 11 

4. Hammer low, scarcely elevated above 

sternum A. chorrera 

Hammer length equal to apical diam- 
eter 5 

5. Hammer apex quadrate; aedeagal hook 
apices footlike A. cuadrada 



Hammer apex circular; aedeagal hooks 
without footlike apices 6 

6. Dorsum of aedeagus with a transverse, 

subapical arcuate lobe A. arcuata 

Dorsum of aedeagus without arcuate 
lobe 7 

7. Hammer tiny; aedeagal apex notched 

A. muesca 

Hammer thimble shaped; aedeagal 
apex rounded 8 

8. Aedeagal apex strongly trilobed; pale 
mesal pronotal band not extending lat- 
erally beyond ocelli A. fenestrata 

Aedeagal apex simple; pale mesal 
pronotal stripe wide, extending later- 
ally beyond ocelli 9 

9. Aedeagal apex abruptly narrowed at 

shoulder (Fig. 58) 10 

Aedeagal apex gradually tapered .... 

A. paleta 

10. Membranous area between hooks ex- 
tends almost to aedeagal base (Fig. 58) 

A. timote 

Membranous area between hooks ex- 
tends far short of aedeagal base 

A. shamatari 

11. Aedeagal apex simple 12 

Aedeagal apex multilobed 25 

12. Aedeagal hooks straight, daggerlike . . 

A. cniza 

Aedeagal hooks curved, scythelike ... 13 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



13. Apex of aedeagal hooks fingerlike, ae- 
deagal apex surmounting a distinct 

neck A. digitata 

Apex of aedeagal hooks not fingerlike, 
subapical area of aedeagus without 
neck 14 

14. Ventral aspect of aedeagus with a con- 
spicuous pair of membranous lobes . . 15 
Ventral aspect of aedeagus with mem- 
branous lobes absent or inconspicuous 

22 

15. Dorsal keel formed by a pair of widely 
spaced ridges (Fig. 16); head with a 
conspicuous dark area at ocelli (Fig. 

12) 16 

Ridges of dorsal keel closely spaced 
(Fig. 31); head with diffuse or pale 
ocellar pigment (Fig. 27) 17 

16. Keel arms divergent anteriorly (Fig. 
16); occipital area pale (Fig. 12) .... 
A. chaima 

Keel arms convergent anteriorly (Fig. 
42); occipital area dark (Fig. 38) .... 
A. perija 

17. Inner margins of aedeagal hooks scal- 
loped (Fig. 29) 18 

Inner margins of aedeagal hooks 
smoothly curved (Fig. 24) 20 

18. Aedeagus with a dorsomesal arcuate 

keel (Fig. 31) A. makushi 

Aedeagus without dorsomesal arcuate 
keel 19 

19. Aedeagal apex notched, decurved in 

lateral aspect A. baniva 

Aedeagal apex rounded, straight in lat- 
eral aspect (Figs. 45-46) A. phantoma 

20. Membranous ventral aedeagal lobes 
wider than long (Fig. 24); submarginal 
pronotal bands narrow (Fig. 22) 

A. karina 

Membranous ventral aedeagal lobes 
longer than wide (Fig. 19); submargin- 
al pronotal bands wide (Fig. 17) ... 21 

21. Aedeagal apex truncate (Fig. 19); ven- 
tral membranous aedeagal lobes small 

(Fig. 19) A. claudiae 

Aedeagal apex pointed (Fig. 9); ventral 
membranous aedeagal lobes large (Fig. 

9) A. arawak 

22. Aedeagal apex gradually narrowed to 

a point A. bari 

Aedeagal apex truncate or broadly 
rounded 23 



23. Aedeagus with a small ventral mem- 
branous lobe A. chiquita 

Aedeagus without ventral membranous 
lobe 24 

24. Projecting apex of aedeagus subequal 

to shoulder A. liana 

Projecting apex of aedeagus about 
twice as long as shoulder .... A. blanca 

25. Aedeagal apex with a dorsolateral pair 

of small hornlike processes 26 

Aedeagal apex without hornlike pro- 
cesses 27 

26. Hornlike aedeagal processes acute and 
conspicuous in ventral aspect 

A. caraca 

Hornlike aedeagal processes rounded 
and inconspicuous in ventral aspect . . 

A. aroucana 

27. Dorsal aedeagal keel V-shaped (Fig. 

52) 28 

Dorsal aedeagal keel not V-shaped ... 29 

28. R and basal half of Sc vein covered 
with dark brown pigment (Fig. 53) . . 

A. pictipes 

R and basal half of Sc without con- 
strasting pigment band A. achagua 

29. Ventral aspect of aedeagus with a large 

membranous lobe A. pequena 

Ventral aspect of aedeagus without 
membranous lobe 30 

30. Lateral aedeagal lobes scarcely pro- 
jecting A. menuda 

Lateral aedeagal lobes distinctly pro- 
jecting 31 

31. Lateral aedeagal lobes about as wide 

as median lobe 32 

Lateral aedeagal lobes about half as 
wide as median lobe A. sp VZ-10 

32. Wings with a large transparent spot be- 
yond cord, and an irregular transparent 

costal band A. vistosa 

Wings rather uniformly pigmented . . . 

A. pinza 

Acknowledgments 

I thank R. Holzenthal, University of Min- 
nesota, and O. S. Flint and N. Adams, Unit- 
ed States National Museum, for the loan of 
specimens. I also thank P. Zwick and cu- 
rators at the California Academy of Science 
and the Museum of Comparative Zoology 



VOLUME 112, NUMBER 1 



93 



for their help in arranging loans of type ma- 
terial. 

Literature Cited 

Banks, N. 1914. New neuropteroid insects, native and 
exotic. — Proceedings of the Academy of Natu- 
ral Science of Philadelphia 66:608-632. 

Jewett, S. G. 1960. Two new species oi Anacroneuria 
(Plecoptera) from Goias, Brazil. — Los Angeles 
County Museum Contributions in Science 36: 
1-4. 

Kimmins, D. E. 1948. A new species of Anacroneuria 
(Plecoptera, Perlidae) from Trinidad. — The Pro- 
ceedings of the Royal Entomological Society of 
London, Series B 17:105-106. 



Klapalek, F. 1923. Plecopteres nouveaux. — Annales de 
la Societe Entomologique de Belgique 63:21- 
29. 

Pictet, F. J. 1841. Histoire naturelle generale et parti- 
culiere des insectes Nevropteres. Famille des 
Perlides. 1. Partie: 1-423. 

Stark, B. P. 1994. Anacroneuria from Trinidad and To- 
bago (Plecoptera: Perlidae). — Aquatic Insects 
16:171-175. 

Stark, B. P. 1995. New species and records of Anacro- 
neuria (Klapalek) from Venezuela (Insecta, Ple- 
coptera, Perlidae). — Spixiana 18:211-249. 

Zwick, P. 1972. Die Plecopteren Pictets und Burmeis- 
ters, mit Angaben uber weitere Arten (Insec- 
ta). — Revue Suisse de Zoologie 78:1123-1194. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):94-96. 1999. 

A new species of Siamosquilla from Indonesia 
(Crustacea: Stomatopoda: Protosquillidae) 

Mark V. Erdmann and Raymond B. Manning 

(MVE) Department of Integrative Biology, University of California at Berkeley, Berkeley, 

California 94720, U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of 

Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. 

Abstract. — Siamosquilla sexava, the second species of the genus, is de- 
scribed from Indonesia. It can be distinguished from the type species, S. hyl- 
lebergi Naiyanetr, from Thailand, by the shorter median rostral spine, the larger 
ocular scales, the much broader telson, and differences in ornamentation of the 
fused sixth abdominal somite and telson. 



Among the stomatopods collected by one 
of us (M. V. E.) during a six-year field study 
in Indonesia was a minute protosquillid 
which proved to be the second known spe- 
cies of Siamosquilla Naiyanetr, 1989. It is 
described below. 

Abbreviations used in the account in- 
clude: LON, Lembaga Oseanologi Nasional 
(National Institute of Oceanoraphy), Jakar- 
ta, Indonesia; TL, total length, measured on 
the midline; USNM, National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C, U.S.A. 

The holotype and some paratypes have 
been deposited in the USNM; two paraty- 
pes are in the LON. 

Family Protosquillidae Manning, 1980 

Siamosquilla Naiyanetr, 1989 

Siamosquilla sexava, new species 

Fig. 1 

Material examined. — Indonesia: Moro- 
maho, Tukang Besi: 1$, TL 18 mm (ho- 
lotype, USNM 260927), 2??, TL 16-17 
mm (paratypes, LON). — Gili Meno, Lom- 
bok: 1(5, TL 17 mm, 1$, TL 18 mm (par- 
atypes, USNM 260928).— Melanguane, 
Sangihe-Talaud: 1$, TL 17 mm (paratype, 
USNM 260929).— Taupun, Togian Islands: 
M, TL 13 mm (paratype, USNM 260930). 

Diagnosis. — Size very small, TL less 



than 20 mm in adults. Cornea broadened, 
set obliquely on stalk. Ocular scales well 
developed, produced into triangular lobes 
laterally, extending nearly to lateral rostral 
spine in adults. 

Rostral plate sharply trispinous, median 
spine distinctly upturned distally, extending 
to base of corneas; ventral projection of ros- 
tral plate large, obtusely rounded ventrally. 
Lateral rostral spines extremely long, slen- 
der and recurved, length nearly two-thirds 
that of median spine. Basal part of rostral 
plate very thin. Anterior margins of lateral 
plates of carapace concave, anterolateral an- 
gles strongly produced to a sharp point, ex- 
tending anteriorly to base of rostral plate. 

Mandibular palp absent. Five epipods 
present. 

Raptorial claw with inflated part of outer 
margin of dactylus notched. Propodus with 
single movable spine proximally on inner 
margin. 

Anterior 4 abdominal somites smooth, 
unarmed, not carinate dorsally. Fifth ab- 
dominal somite smooth medially, with sin- 
gle low longitudinal carina laterally above 
lateral margin, separated from margin by a 
groove, armed with posterolateral spine. 
Sixth abdominal somite entirely fused with 
telson in adults, dorsal surface rough, with 
shallow, irregularly curved grooves, lacking 
carinae entirely. 



VOLUME 112, NUMBER 1 



95 




'■^V-^ A 



Fig. 1. Siamosquilla sexava, new species, female holotype, TL 18 mm. a. Anterior part of body, dorsal view 
(apex of left ocular scale damaged); b, Rostral plate, lateral view; c. Fused sixth abdominal somite and telson 
and left uropod, dorsal view. Scales = 1 mm. 



Telson much wider than long, dorsal sur- 
face rough, slightly inflated, with 3 very in- 
distinct bosses. Median boss outlined by a 
shallow groove in dorsal surface of telson; 
boss rounded anteriorly, converging poste- 
riorly; outline of submedian bosses even 
less distinct, extending posteriorly almost to 
posterior margin of telson. Median fissure 
completely fused, no longer visible. Three 
pairs of marginal teeth, submedians with 
moveable apices arising submarginally un- 
der submedian marginal projections. Inter- 
mediate teeth very low and rounded, almost 
indistinguishable in larger specimens. Lat- 
eral teeth more distinctly produced, with 
rounded apices. Separation between three 
pairs of marginal telson teeth shallow and 
rounded. Submedian teeth with 12-14 den- 
ticles on either side of midline, increasing 
noticeably in length laterally, intermediate 
teeth each with 2 fixed mesial denticles, lat- 
erals each with 1 denticle. Lateral margins 
of telson straight, rounded distally. 

Uropods stout, proximal segment of ex- 



opod with 9 short movable spines laterally, 
distalmost extending beyond midlength of 
distal segment, with fixed distal spine ven- 
trally. Inner margin of uropodal exopod se- 
tose; uropodal endopod with normal com- 
plement of setae. Inner spine of basal pro- 
longation of uropod much shorter than out- 
er. 

Size. — Males (n = 2), TL 13-17 mm. Fe- 
males (n = 5), TL 13-18 mm. 

Remarks. — This small species has a very 
distinctive rostral plate, with long, recurved 
lateral spines, and a telson that is most re- 
markable for its lack of distinguishing fea- 
tures; the sixth abdominal somite is fused 
to the telson, which has almost indiscern- 
ible bosses and a fused median fissure. Sia- 
mosquilla sexava differs from S. hyllebergi 
in numerous features: the median rostral 
spine is shorter, extending to, rather than 
beyond, the cornea and the lateral rostral 
spines are longer; the ocular scales are 
broader; in adults the anterolateral angles of 
the carapace are sharper and more pro- 



96 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



nounced; the dorsal sculpture of the fused 
sixth abdominal somite and telson is much 
more elaborate; and the telson is much 
broader. 

The fixed projections on the posterior 
margin of the telson are much more pro- 
nounced in S. hyllebergi than in S. sexava, 
and the submedian projections in the former 
species probably obscure the submarginal, 
movable submedian teeth of the telson, 
which are clearly visible in 5. sexava. 

In 5". sexava, the size of the ocular scales 
increases allometrically, and in very small 
specimens of S. sexava the scales resemble 
those of larger S. hyllebergii; they increase 
in width with increasing TL. 

Distribution. — Known only from eastern 
Indonesia, where it is relatively widespread. 
Recorded from five regions: Lombok, San- 
gihe-Talaud, Togian Islands, and Tukang 
Besi Archipelago. 

Etymology. — The species name is from 
the word sexava, which means "mantis" in 
Bahasa Talaud, the local language in one of 
the collection localities (Melanguane, San- 
gihe-Talaud). The local fishermen, school 
teachers and children there were most help- 
ful in assisting with collecting, and were 
fully cognizant of the differences between 
stomatopods and other decapod shrimp, 
comparing stomatopods to the "sexava" 
without any solicitation. Their assistance is 
most appreciated. 



Acknowledgements 

We thank the Indonesian Institute of Sci- 
ences for sponsoring Erdmann's project, 
and Kalam Sebayang and Ibu Dewi Soen- 
arijadi for their assistance with research vi- 
sas. Dr. Mohammad Kasim Moosa was the 
Indonesian sponsor of this research, and 
Erdmann gratefully acknowledges his men- 
torship on stomatopod morphology and his 
continual support of this project. We also 
thank Dr. Sukarno, Dr. Suharsono, and Dr. 
Anugerah Nontji of the Indonesian Institute 
of Sciences. 

Funding for Erdmann's research was pro- 
vided by grants from the UC Pacific Rim 
Research Program, the NSF International 
DDIG (#9503060 and #9704616) Program 
and a visiting scientist appointment from 
the Smithsonian Institution. Lilly King 
Manning prepared the figure. This is con- 
tribution no. 467 from the Smithsonian Ma- 
rine Station at Fort Pierce, Florida; the sup- 
port of that program for Manning's studies 
of stomatopod systematics is gratefully ac- 
knowledged. We gratefully acknowledge 
the reviews of Shane Ahyong, David K. 
Camp, and Roy K. Kropp as well as the 
cooperation of Rafael Lemaitre. 

Literature Cited 

Naiyanetr, P. 1989. Siamosquilla hyllebergi, a new ge- 
nus and new species of stomatopod from Thai- 
land. In E. A. Ferrero, ed., Biology of stomato- 
pods. Selected symposia and monographs, 
U.Z.I., 3:281-284. Muchi, Modena. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):97-105. 1999. 

The taxonomic status and zoogeography of Cambarus bartonii 
carinirostris Hay, 1914 (Crustacea: Decapoda: Cambaridae) 

Roger F. Thoma and Raymond E Jezerinact 
(RFT) Ohio Environmental Protection Agency, 2110 Aurora Rd., Twinsburg, Ohio 44087, U.S.A. 

Abstract. — Historically, Cambarus bartonii carinirostris was considered a 
subspecies of Cambarus bartonii. Recent studies indicate that C b. cariniros- 
tris is indistinguishable from the nominate species and should be relegated to 
synonymy with C bartonii. In other studies this assignment is not accepted 
and subspecific status has been maintained for C b. carinirostris when re- 
porting on crayfish closely related to C bartonii. Work in the area of northern 
Appalachian crayfishes has made it apparent to us that C. b. carinirostris should 
be elevated to full species status based on its unique meristic and morphometric 
characteristics. 



For nearly two decades we have studied 
the crayfishes of the northern Appalachians. 
Of the many questions associated with this 
area, the proper identity and distribution of 
C b. bartonii (Fabricius 1798) has been and 
continues to be a principal concern. Ort- 
mann's (1905, 1906, 1931) studies remain 
the foundation on which the distribution 
and systematics of northern Appalachian 
crayfish are presently understood. After 
Ortmann's death in 1933, Horton H. Hobbs, 
Jr., shifted the center of crayfish systematic 
studies to the southern Appalachians (and 
other southern areas) and the identity of C. 
b. bartonii and its subspecies remained un- 
resolved. In the early 1970s we became in- 
terested in the ecology of crayfish in our 
home state, Ohio. We were immediately 
confronted with the problem of C. b. bar- 
tonii and the subspecies associated with it, 
since both C b. carinirostris (Hay 1914) 
and C. b. cavatus (Hay 1902) are reported 
from Ohio (Hobbs 1974, 1989; Thoma & 
Jezerinac 1982). With the unresolved taxo- 
nomic problems and a domination by mem- 
bers of the subgenus Cambarus in Ohio, it 
became apparent that a taxonomic study 



t Deceased, 21 April 1996. 



was needed before the study of crayfish 
ecology could commence. 



Materials and Methods 

Specimens examined came from the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 
(USNM), and the Ohio State University 
Museum of Biological Diversity, Colum- 
bus, Ohio (OSU-MBD). The paratypes, 
housed at the Museum of Comparative Zo- 
ology (MCZ) were not examined. Field col- 
lections (now house at OSU-MBD) were 
made using a 1.3 X 2 m seine or by hand. 
A total of 95 specimens of C carinirostris 
(33 Form I males, 27 Form II males, and 
35 females) and 191 specimens of C b. 
bartonii (79 Form I males, 43 Form II 
males, and 69 females) were measured for 
this study. Measurements were made to the 
nearest 0.1 mm using a vernier caliper and 
followed Hobbs (1981) and Jezerinac 
(1985). Measurements of regenerated body 
parts were avoided. Analysis was per- 
formed using SYSTAT 5.2.1. Principal 
component analysis used only Form I male 
specimens. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Cambarus (Cambarus) carinirostris Hay, 
new status 

Cambarus bartonii carinirostris Hay, 1914: 
384. 

Cambarus bartonii montanus. — Faxon, 
1914:386 (in part). — Newcombe, 1929: 
286 (in part). 

Cambarus montanus montanus. — Ortmann, 
1931:106 (in part). 

Cambarus {Cambarus) bartoni cariniros- 
tris. — Ortmann, 1931:107. 

Cambarus {Cambarus) bartonii cariniros- 
tris.— Uohbs 1969: 109, fig. 19m; 1974: 
11, fig. 24; 1989: 13 fig. 30.— Thoma, 
1982:875. — Thoma & Jezerinac, 1982: 
136. — Jezerinac, 1983: 4. — Jezerinac & 
Thoma, 1984: 120 figs. 8-9. — Jezerinac 
& Stocker, 1989: 2. — Jezerinac & Stock- 
er, 1990: 1. — Jezerinac, Stocker & Tarter, 
1995: 76-83, fig. 35-38. 

Diagnosis. — Body pigmented. Carapace 
subcylindrical, slightly flattened dorsoven- 
trally. Eyes slightly reduced. Rostrum with 
parallel or slightly concave margins, mar- 
gins thickened, lacking marginal spines or 
tubercles, rostrum curved abruptly cephali- 
cally and terminating in upturned corneous 
tubercle; frequently with a median carina. 
Areola 3.4-12.0 times longer than wide 
(median = 5.7), comprising 25.2-40.4% of 
total length of carapace (median = 37.9%), 
bearing 3 to 4 punctations across narrowest 
part. Cervical spine absent or reduced to 
blunt tubercle. Cervical grove uninterrupt- 
ed. Suborbital angle acute. Postorbital ridge 
lacking cephalic spine or tubercle. Bran- 
chiostegal spine reduced to small knob. An- 
tennal scale approximately 1.5 times as 
long as broad, with mesial and lateral mar- 
gins subparallel near and at midlength; dis- 
tomesial margin strongly sloping. Basio- 
podite of antenna lacking spine. Ischiopod- 
ite of antenna with blunt spine. Chela 
smooth, robust, length 79.6-111.9% of total 
carapace length (median = 94.0%) in Form 
I males (71.8-88.4%, median = 78.8% in 
females), bering 1 row of 5 to 8 (median = 
7.0) adpressed squamous tubercles along 



mesial margin of palm and 5 to 7 puncta- 
tions dorsal to such tubercles (occasionally 
distal 1 to 4 punctations replaced by tuber- 
cles). Width of gape of fingers of Form I 
male 17.4-46.3% of palm length (median 
= 27.2%), less so in Form II males (median 
= 16.3%) and females (median = 17.2%). 
Lateral margin of fixed finger weakly cos- 
tate; moderately developed dorsomedial 
ridges on both fingers flanked by parallel 
rows of punctations; fixed finger impressed 
at dorsal and ventral lateral bases; dactyl 
0.9-2.5 (median = 2.1) times longer than 
mesial margin of palm; palm width 41.5- 
51.1% (median = 46.1%) of chela length; 
third or fourth tubercle of mesial margin of 
fixed finger enlarged; 1—2 tubercles usually 
present on mid-subpalmar surface; never 
with elongated setae at base of fixed finger. 
Dorsomesial margin of carpus of chela with 
1 distal spine and 1 proximal blunt tubercle; 
ventral surface with 1 or 2 conical tubercles 
at distal margin. Ventrolateral ridge of mer- 
us usually with 2-3 spines. Hook only on 
ischium of third pereiopod of male. Basal 
boss on coxa of fourth pereiopod well de- 
veloped. First pleopods of Form I male con- 
tiguous at base, with 2 short terminal ele- 
ments bent at approximately 90° to main 
shaft; corneous central projection truncated 
distally, bearing subapical notch; mesial 
process inflated, tapering distally; central 
projection of Form n male pleopods non- 
corneous, club-shaped. Females with an- 
nulus ventralis slightly embedded in ster- 
num, asymmetrical, subrhomboid, slightly 
movable, lacking cephalolateral promi- 
nence; first through fifth pleopods similar 
in shape. 

Color notes. — For the most part, this spe- 
cies is dorsally a uniform brown ranging 
from chestnut to tan with ventral surfaces 
fading to cream. Some populations have a 
greenish hue. The thickened rostral margins 
and postorbital ridges are a brownish red 
and the larger tubercles and spines orange. 
No banding or striping evident. 

Types. — "Type" and paratypes USNM 
23962 (1 Form I male, 7 Form II male, 15 



VOLUME 112, NUMBER 1 



99 



female); paratypes, MCZ 7399 (1 Form I 
male, 1 Form II male, 1 female). The Form 
I male housed at the USNM is herein des- 
ignated the lectotype of the species. 

Type locality. — Hay (1914:385) stated 
"Gandy Creek, Oceola, Randolph Co., W. 
Va." in his original description. A visit to 
this area and discussions with local resi- 
dents indicated Oceola (38°42'50"N, 
79°38'00"W) was the location of a now 
non-extant school house at the site of a his- 
toric lumber camp. The old school house 
was located near the Sinks of Gandy. We 
made a topotype collection from Gandy 
Creek upstream of County Road 40, just 
west of County Road 29/1 (38°43'22"N, 
79°37'38"W). This site is the first road 
crossing downstream of the mouth of the 
Sinks of Gandy. 

Range. — Found throughout the drainages 
of the Allegheny and Monongahela rivers 
in Pennsylvania, New York and West Vir- 
ginia; tributaries of the Ohio River up- 
stream of Sunfish Creek in Ohio and Fish 
Creek in West Virginia; southern tributaries 
of Lake Erie and Lake Ontario from the 
Grand River, Ohio, to the Gennesse River, 
New York; throughout the Greenbrier River 
and tributaries of the New River upstream 
of the Greenbrier; upper Elk River, West 
Virginia. 

Variation. — In the upper reaches of the 
Monongahela basin (Cheat and Tygart/ 
Buckhannon basins) C. carinirostris prob- 
ably retains its most plesiomorphic state. 
This is the only portion of the range in 
which no other stream dwelling forms of 
Cambarus are found. Here, C. carinirostris 
attains its greatest degree of sculpturing (in- 
cluding the carina) and approaches the body 
size of members of the subgenera Camba- 
rus and Puncticambarus that inhabit larger 
rivers. The plesiomorphic appearance is 
likely in part due to the larger sizes at- 
tained. In the remainder of the range, Cam- 
barus {Puncticambarus) robustus Girard, 
1852, or Cambarus (C) sciotensis Rhoades, 
1944, occupy the larger mainstem streams 
and C. carinirostris is confined to the 



smaller tributaries. In the Casselman River 
of the Youghiogheny River, Pennsylvania, 
we have seen specimens with body forms 
reminiscent of the subgenus Erebicambarus 
Hobbs, 1969, in that they display a more 
tubular, sausage-shaped carapace that is less 
dorsoventrally compressed. The chelae re- 
main decidedly within the range of C. car- 
inirostris, though they have the least 
amount of gape between the fingers. Spec- 
imens from the southernmost extent of the 
range display reduced inflation of the ros- 
tral margins and lack the 90° angle at the 
rostral tip. The rostra in this population 
most closely resemble the probable ple- 
siomorphic state. 

Occasionally, a second row of slightly 
produced tubercles can be found on the 
palm of the chela. This character does not 
exhibit a defined geographical pattern but 
appears in some individuals in most collec- 
tions. All other populations exhibit the nor- 
mal characteristics given in the Diagnosis. 

Size. — Mature specimens range from 26 
to 48.8 mm (median = 34.9 mm) total car- 
apace length (median: Form I males 36.9 
mm. Form II males = 31.9 mm, females = 
35.4 nrni). 

Life history notes. — Jezerinac et al. 
(1995) reported Form I males from late 
April through early September and oviger- 
ous females from July through mid August. 
This study found Form I males as late as 
October and ovigerous females in the re- 
ported range of dates. No information exists 
on longevity, growth rates, thermal prefer- 
ences or dietary habits. 

Habitat and ecology. — The normal hab- 
itat occupied by C. carinirostris is pools 
and riffles of high gradient first and second 
order streams. Populations can be found in 
intermittent streams but the abundance is 
greatly reduced. Some burrowing occurs, 
mostly sub-boulder, in mid-stream or on the 
edges of streams. This species is a second- 
ary burrower. Cambarus carinirostris is ca- 
pable of expanding its niche in the absence 
of other species of Cambarus that are pri- 
mary burrowers or mainstem inhabitants. 



100 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Often caves, if present, yield specimens, 
usually in the vicinity of the mouth. This 
species is sensitive to excess silt, bed load 
sediments, nutrient enrichment, acid mine 
impacts, and habitat alterations that reduce 
cobble and boulder abundance. 

Taxonomic status. — Numerous authors 
have commented on the distribution and 
taxonomic status of C. carinirostris. In his 
original description Hay (1914: 385) re- 
ported the species from Tygart Valley and 
Cheat River in Randolph County, West Vir- 
ginia, stating that C. carinirostris was "... 
a well marked subspecies ..." differing 
from the nominate species in that "... the 
carapace is a little more cylindrical, the ros- 
trum broader and flatter, and always fur- 
nished near the tip with a median longitu- 
dinal carina." Faxon (1914: 385), com- 
mented that the rostral carina was a rather 
elusive character and reported additional re- 
cords for this taxon from the Greenbrier 
River basin (West Virginia). The next report 
on C carinirostris was Ortmann (1931: 
139) in which he retained the subspecific 
status for C. carinirostris, and stated C. car- 
inirostris differs from C. b. bartonii only in 
the presence of the carina. After Ortmann's 
death, no further comments were published 
on the taxonomy of C. carinirostris until 
Hobbs (1972: 111) commented that it prob- 
ably should not be recognized. Bouchard 
(1976: 588) recognized no subspecies of C 
bartonii, reporting that no characters sepa- 
rate the currently recognized subspecies. He 
also noted the instability of the rostral char- 
acters. Thoma (1982), Thoma & Jezerinac 
(1982), Jezerinac (1983), and Jezerinac et 
al. (1995) continued to use subspecific sta- 
tus for C. carinirostris. 

We agree with Faxon (1914), Ortmann 
(1931), and Bouchard (1976) that the carina 
of C. carinirostris is an elusive trait, fre- 
quently absent or greatly reduced. In our 
study of the species we found the popula- 
tion in the vicinity of the type locality ex- 
hibited the carina most strongly and fre- 
quently, and in this respect one could con- 
clude that C. carinirostris is a local varia- 



tion. When we examined the bartonii 
complex for other traits, we noted clear dif- 
ferences in chela structure between popu- 
lations of the Atlantic and Mississippi 
drainages in Pennsylvania (especially in 
form I males). Thoma & Jezerinac (1982: 
137) reported that C. carinirostris could be 
distinguished from C. bartonii bartonii (Fa- 
bricius, 1798) by the thickened and fre- 
quently concave rostral margins; an abrupt- 
ly ending rostrum that forms a 90° angle 
with the base of the acumen; more strongly 
developed postorbital ridges; stronger de- 
velopment of the lateral impression and 
dorsal ridges of the chela; a sUght devel- 
opment of a second row of 2 or 3 tubercles 
on the mesial margin of the palm; reduced 
gape between the dactyl and propodus; and 
an enlarged third tubercle on the mesial 
margin of the opposable propodus. Contin- 
ued work throughout the ranges of the two 
forms (and the rest of the range of the sub- 
genus) revealed that the combination of 
thickened rostral margins, an enlarged third 
(or fourth) tubercle on the mesial margin of 
the opposable finger of the propodus, mod- 
erate development of the lateral impression 
and dorsal ridges of the chela, and less than 
2 full rows of palmar tubercles is sufficient 
to distinguish this species from the nomi- 
nate species and all other taxa presently 
known in the bartonii complex. The partial 
second row of palmar tubercles (not con- 
sistently present), 90° angled rostral mar- 
gins, strong postorbital ridges, and dactyl 
gape are useful in distinguishing this spe- 
cies individually from other species of the 
bartonii complex. Within the subgenus two 
forms (C b. cavatus Hay, 1902, and C. sci- 
otensis Rhoades, 1944) consistently display 
the development of a second row of tuber- 
cles on the mesial margin of the palm. 
Cambarus sciotensis displays a full com- 
plement of 5 or 6 tubercles in the second 
row. Thoma & Jezerinac (1982) reported 
that C. b. cavatus consistently had 3 to 5 
tubercles in the second row of palmar tu- 
bercles, noninflated rostral margins, and a 
narrower areola (the narrower areola may 



VOLUME 112, NUMBER 1 



101 



relate to the burrowing habits of C b. ca- 
vatus). 

Principal component analysis of morpho- 
metric data for C. carinirostris and C. b. 
bartonii (Fig. 1) illustrates slight differenc- 
es in body structure. The gape of the chela 
fingers, dactyl width, and areola width are 
the strongest loading variables (Table 1). 
Areola and dactyl width tend to be wider 
and chela gape narrower (Fig. 2) in C. car- 
inirostris. No single body proportions are 
sufficient to separate the two taxa on a con- 
sistent basis. The most reliable character 
traits (using Form I males only) for consis- 
tently separating the two taxa are develop- 
ment of an enlarged third tubercle on the 
inner margin of the opposable propodus, 
lateral impression of the chela, and dorsal 
ridges on the chela fingers of C. cariniros- 
tris (Fig. 2). The northern form of C. b. 
bartonii (Atlantic drainage from the Poto- 
mac basin northward) lacks these chela 
characters altogether while the southern 
complex of C. b. bartonii lacks the lateral 
impression and thickened rostral margins. 
Some southern representatives of C. b. bar- 
tonii have an enlarged third tubercle on the 
opposable dactyl. This is accompanied by 
an enlarged first and fourth (occasionally 
third) tubercle on the opposable finger of 
the propodus, a characteristic lacking in C. 
carinirostris. 

No intergrade populations have been 
found (using the characters we employed) 
between C. carinirostris and any other 
closely related taxon. For these reasons we 
elevate C. carinirostris to full species sta- 
tus. Important to note is that the above char- 
acteristics are most reliable in Form I 
males, preferably of a large size. 

Evolution, zoogeography and associated 
variation. — Cambarus carinirostris is pri- 
marily an inhabitant of the still extant por- 
tions of the preglacial Pittsburgh River 
(Leverett 1902, Tight 1903) and (in part) 
the New River basin. This distribution is a 
reflection of the evolutionary history of the 
species. Cambarus carinirostris was most 
likely derived from an ancestral stock of the 



bartonii complex that inhabited the New 
River (Kanawha basin). This stock was in- 
tern derived from populations inhabiting 
the upper Tennessee River basin, in partic- 
ular the Clinch and Powell systems. Hobbs 
(1969) concluded that the subgenus Cam- 
barus originated in the Tennessee basin and 
entered the Kanawha River basin from a 
northward migrating member of the '"extra- 
neus group." We originally believed the 
subgenus Cambarus originated in the Ka- 
nawha basin (Jezerinac et al. 1995), but the 
recent discovery of a previously unknown 
archetypal Cambarus species (a description 
of this species and a discussion of its evo- 
lutionary significance is in development) in 
an isolated portion of what was once the 
Tennessee basin and now part of the Cataw- 
ba River basin, confirms Hobbs' conclu- 
sion. 

Upon entering the New River basin 
through interdigitating headwater streams, 
probably in the Burk's Garden area of Vir- 
ginia, the ancestral C carinirostris spread 
steadily through the system. Cambarus car- 
inirostris probably was the first Cambarus 
to enter the upper New River system, thus 
having uninhibited dispersal opportunities. 
Access was gained to the preglacial Pitts- 
burgh River basin in the headwaters of the 
Greenbrier River, again before other mem- 
bers of the subgenus Cambarus. No further 
dispersal occurred until postglacial times 
when C. carinirostris followed the retreat- 
ing glaciers into the new Laurencian basin, 
where it established populations from the 
Gennesse River of New York in the east to 
the Grand River of Ohio in the west. Dis- 
persal on the western edge of the range was 
thwarted by the presence of crayfish popu- 
lations (presently assigned to C b. cavatus) 
that occupied the preglacial Teays River 
system (Jezerinac 1983). Cambarus b. ca- 
vatus most likely entered the Teays basin 
via the Big Sandy River. The Teays River 
population was a stream dwelling species of 
the subgenus Cambarus that apparently had 
ecological preferences similar to C. carini- 
rostris. In the Allegheny Plateau portion of 



102 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



3 


Factor 3 




: A 


2 


- 




• 


1 


- 





- 


1 


- 


2 


- 



C. carinirostris • # 



,oo 

O. Q 



ib oo 



C.b.bartonii 



_0 r t ■ » ■ I ■ ■ » ■ I ■ ■ ■ i 1 ■ ■ « « I I « ■ « I ■ ■ ' ■ I ■ ■ I ■ I 



till 



-4 -3 -2-10 1 

Factor 2 



7 
6 
5 
4 
3 
2 
1 




Gape (mm) 



III lll II ll 
CO 


. @ 8 • 


• 
•• 


- 


o • 




I—I 1 i 


c.b.bartonii rn n ^- 

. 1 . . . . 1 . . . . 1 . . . 


carinirostris 
1 1 1 1 1 1 1 



10 



20 30 40 

Chela Length (mm) 



50 



60 



Fig. I. A, Principal Component Analysis of Cambanis carinirostris (black circles, N = 33) and C. bartonii 
bartonii (open circles, N = 79) using Factors 2 and 3 (Table 1), Form I males only; B, Gape of chela fingers 
by length of chela for C. carinirostris (black circles, N = 33) and C. b. bartonii (open circles, N = 79), Form 
I males only. 



VOLUME 112, NUMBER 1 



103 



Table 1. — Factor loadings for Principal Component 
Analysis of morphometric data of Form I male speci- 
mens of C. b. bartonii and C. carinirostris. 



Measurement 


Factor 2 


Factor 3 


Chela length 


0.066 


-0.044 


Chela width 


0.017 


0.032 


Chela depth 


0.030 


0.037 


Gape of fingers 


0.227 


-0.326 


Palm length 


0.039 


-0.031 


Dactyl length 


0.088 


-0.042 


Dactyl width 


-0.095 


0.394 


Propodus finger length 


0.045 


-0.078 


Propodus finger width 


-0.076 


0.120 


Carapace length* 


0.040 


0.027 


Carapace width 


0.016 


0.035 


Areola length 


0.110 


0.023 


Areola width 


-0.667 


-0.189 



* Measurement includes rostrum. 

the New-Kanawha River basin and those 
streams draining to the Ohio River down- 
stream of the Sardis Col, C. b. cavatus pre- 
vailed. In the newly formed Ohio River, the 
two species are presently found on their re- 



spective preglacial sides of the Sardis Col, 
with neither having been able to advance 
into the other's range. To the north and to 
some degree to the east, the dispersal of C. 
carinirostris was inhibited by the presence 
of C b. bartonii. It appears that the post- 
glacial dispersal of C b. bartonii (in the 
Atlantic slope drainage) was much more 
rapid than that of C. carinirostris, with C. 
b. bartonii effectively excluding C carini- 
rostris in those areas. Few opportunities ex- 
isted for eastward dispersal of C. cariniros- 
tris in the Greenbrier, Monongahela, and 
Allegheny river basins because of physical 
barriers. Where potential stream piracies 
have been identified, no C. carinirostris 
could be found in Atlantic drainage 
streams. The ecological preferences of C 
carinirostris and C b. bartonii appear to be 
very similar. 

Taxonomic distinction. — No other spe- 
cies in the subgenus Cambarus possesses 
the character combination of thickened ros- 






Fig. 2. A, Dorsal view of pereiopod I chela of Cambarus carinirostris: 1 — enlarged third tubercle on mesial 
margin of immovable finger. 2 — lateral impression, 3 — dorsal ridge; B, Dorsal view of carapace of Cambarus 
carinirostris; 4 — rostral carina, 5 — thickened rostral margins; C, Dorsal view of pereiopod I chela of Cambarus 
b. bartonii. 



104 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tral margins, an enlarged third tubercle on 
the mesial margin of the opposable finger 
of the propodus, no enlarged tubercles on 
the mesial margin of the dactyl, moderate 
development of the lateral impression and 
dorsal ridges of the chela, and 1 row of pal- 
mar tubercles. 

Acknowledgments 

We would like to thank the many indi- 
viduals who helped us collect the crayfish 
used for this paper (including those who 
went before us). A special posthumous 
thanks to Dr. H. H. Hobbs, Jr and to the 
National Museum of Natural History, 
Smithsonian Institution, for access to the 
crayfish collections. We thank Dr. Joseph F. 
Fitzpatrick, Jr., of the University of South 
Alabama, Michael Bolton of the Ohio En- 
vironmental Protection Agency and two 
anonymous reviewers for evaluating the 
manuscript. 

Literature Cited 

Bouchard, R. W. 1976. Geography and ecology of 
crayfishes of the Cumberland Plateau and Cum- 
berland Mountains, Kentucky, Virginia, Tennes- 
see, Georgia and Alabama. Part II. The Genera 
Fallicambarus and Cambarus. Pp. 585—605 in 
J. W. Avault, Jr., ed.. Freshwater Crayfish, Lou- 
isiana State University Division of Continuing 
Education. 

Fabricius, J. C. 1798. Supplementum Entomologiae 
Systemicae. Hafniae: Proft et Storch, 527 pp. 

Faxon, W. 1914. Notes on the crayfish in the United 
States National Museum and the Museum of 
Comparative Zoology with descriptions of new 
species and subspecies to which is appended a 
catalogue of the known species and subspe- 
cies. — Memoirs of the Museum of Comparative 
Zoology, Harvard College 40(8):350-427. 

Girard, C. 1852. A review of the North American As- 
taci, with observations on their habits and geo- 
graphical distribution. — Proceeding of the 
Academy of Natural Sciences of Philadelphia 6: 
87-91. 

Hay, W. P. 1902. Observations on the crustacean fauna 
of Nickajack Cave, Tennessee, and vicinity. — 
Proceedings of the United States National Mu- 
seum, 25(1292):417-439. 

. 1914. ''Cambarus bartonii carinirostris 

Hay". Pp 384-385 in Walter Faxon, 1914. 



Notes on the crayfish in the United States Na- 
tional Museum and the Museum of Compara- 
tive Zoology with descriptions of new species 
and subspecies to which is appended a cata- 
logue of the known species and subspecies. — 
Memoirs of the Museum of Comparative Zo- 
ology, Harvard College 40(8):350-427. 

Hobbs, H. H., Jr. 1969. On the distribution and phy- 
logeny of the crayfish genus Cambarus. Pp 93- 
178 in The distributional history of the biota of 
the Southern Appalachians, Part I: Inverte- 
brates. P C. Holt, R. L. Hoffman, and C. W. 
Hart, Jr., eds., Virginia Polytechnic Institute and 
State University, Blacksburg, Virginia. Re- 
search Division Monograph 1. 295 pp. 

. 1972. Crayfishes (Astacidae) of North and 

Middle America, Identification Manual 9, Biota 
of freshwater ecosystems. United States Envi- 
ronmental Protection Agency, Water Pollution 
Research Control Series, 173 pp. 

. 1974. A checklist of the North and Middle 

American crayfishes (Decapoda: Astacidae and 
Cambaridae). — Smithsonian Contributions to 
Zoology 166, 161 pp. 

. 1981. The crayfishes of Georgia. — Smithson- 
ian Contributions to Zoology 318, 549 pp. 

. 1989. An illustrated checklist of the American 



crayfishes (Decapoda: Astacidae, Cambaridae, 
and Parastacidae). — Smithsonian Contributions 
to Zoology 480, 236 pp. 

Jezerinac, R. F 1983. Possible correlations of present 
distributions of Ohio crayfishes (Decapoda: 
Cambaridae) with Teays-Age drainages. [Ab- 
stract.] Teays-Age drainage effects on present 
distributional patterns of Ohio biota — an Ohio 
biogeography conference: Abstracts and Sup- 
plements. Ohio Biological Survey Informative 
Circular No. 11:4-5. 

. 1985. Morphological variations of Cambarus 

{Cambarus) bartonii cavatus (Decapoda: Cam- 
baridae) from Ohio, with a diagnosis of the 
Ohio form. — Ohio Journal of Science 85(3): 
131-134. 

, & G. W. Stocker. 1989. Distribution of the 

stream crayfishes of the genus Cambarus (De- 
capoda: Cambaridae) in West Virginia. — Ohio 
Journal of Science 89(2):2. 

, & . 1990. Distribution of crayfishes 

(Decapoda: Cambaridae) of West Virginia. Part 
II: The genera Cambarus and Fallicambarus. — 
Proceedings of the West Virginia Academy of 
Sciences 62(l):7-8. 

, & R. F Thoma, 1984. An illustrated key to 

the Ohio Cambarus and Fallicambarus (Deca- 
poda: Cambaridae) with comments and a new 
subspecies record. — Ohio Journal of Science 
84(3):120-125. 

, G. W Stocker, & D. C. Tarter. 1995. The cray- 



VOLUME 112, NUMBER 1 



105 



fishes (Decapoda: Cambaridae) of West Virgin- 
ia. — Ohio Biological Survey Bulletin New Se- 
ries 10(1), 193 pp. 

Leverett, F. 1902. Glacial formations and drainage fea- 
tures of the Erie and Ohio Basins.- — Mono- 
graphs of the United States Geological Survey, 
No. 91, 802 pp. 

Newcombe, C. L. 1929. The crayfishes of West Vir- 
ginia. — Ohio Journal of Science 29(6):267-288. 

Ortmann, A. E. 1905. The mutual affinities of the spe- 
cies of the genus Cambarus, and their dispersal 
over the United States. — Proceedings of the 
American Philosophical Society, 44(180):91- 
136. 

. 1906. The crawfishes of the state of Pennsyl- 
vania. — Memoirs of the Carnegie Museum. 
2(10):343-523. 

. 1931. Crawfishes of the southern Appala- 



chians and the Cumberland Plateau. — Annals of 
the Carnegie Museum. 20(2):61-160. 

Rhoades, R. E. 1944. Further studies on distribution 
and taxonomy of Ohio crayfishes and the de- 
scription of a new subspecies. Ohio Journal of 
Science 44(2):95-99. 

Tight, W G. 1903. Drainage modifications in south- 
eastern Ohio and adjacent parts of West Virgin- 
ia and Kentucky. United States Geological Sur- 
vey Professional Paper 13:1 1 1 pp. 

Thoma, R. F. 1982. The zoogeography and phylogeny 
of the subgenus Cambarus (Decapoda: Cam- 
baridae). [Abstract.] American Zoologist. 22(4): 
875. 

, & R. F Jezerinac. 1982. New distributional 

records of crayfish {Cambarus and Fallicam- 
barus) from Ohio, including a new subspecies 
record. Ohio Journal of Science. 82(3): 136-138. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):10^119. 1999. 

Two new species of Aegla Leach (Crustacea: 
Decapoda: Anomura: Aeglidae) from southern Chile 

Carlos G. Jara and Victor L. Palacios 

Institute de Zoologia "Dr. Ernst Kilian", Universidad Austral de Chile, Casilla 567, 

Valdivia, Chile 

Abstract. — Two new species of the genus Aegla Leach from southern Chile 
are described. Aegla cholchol, new species, a medium to large sized spinulated 
aeglid, has the River Chol-Chol (Cautin Province) as its type locality. It is 
probably more related to A. rostrata, from subandean lakes at the Tolten and 
Valdivia River basins than to A. bahamondei, from the Tucapel River Basin on 
the western slope of the Nahuelbuta Coastal Cordillera. The other, A. hueicol- 
lensis, new species, a small to medium sized non- spinulated aeglid, has the 
River Pichihueicolla (Valdivia Province), on the western slope of the Pelada 
Coastal Cordillera, as its type-locaUty. A. hueicollensis is similar to A. abtao 
with which it shares more morphological attributes than with other non-spi- 
nulated aeglids of the same geographic region (i.e., A. alacalufi, A. manni, and 
A. concepcionensis). 



Continental Chile is a long and narrow 
strip of land extending from Arica to Cape 
Horn, characterized by a series of climatic 
and biogeographical regions (see Di Castri 
et al. 1968). Three main topographical fea- 
tures, namely, the Andes Cordillera on the 
east, the Coastal Cordillera on the west, 
and the Central Valley between both rang- 
es dominate the landscape from north to 
south (B or gel 1983). The central southern 
part of Chile, between the cities of Con- 
cepcidn (36°55'S) and Puerto Montt 
(41°28'S), harbors the highest diversity of 
aeglids and parastacids in the country (Ba- 
hamonde & Lopez 1963). This diversity 
appears associated to the many rivers and 
lakes that constitute the drainage system of 
this area. 

Aeglids are distributed along a stretch of 
about 2000 kilometers, from the Choapa 
River (3r38'S) in the north down to Ma- 
dre de Dios Island (50°02'S) in the south 
(Bahamonde & Lopez 1963, Jara & Lopez 
1981). At least ten species and two sub- 
species of Aegla (A. concepcionensis 



Schmitt, 1942a, A. expansa Jara, 1992, A. 
pewenchae Jara, 1994, A. bahamondei 
Jara, 1982, A. spectabilis Jara, 1986, A. 
rostrata Jara, 1977, A. abtao Schmitt, 
1942a, A. d. denticulata Nicolet, 1845, A. 
d. lacustris Jara, 1989, A. manni Jara, 
1980, A. araucaniensis Jara, 1980, and A. 
alacalufi Jara & Lopez, 1981) have been 
described for the area between Concepcion 
and Puerto Montt, several associated to the 
Coastal Cordillera. In fact, three, A. con- 
cepcionensis, A. bahamondei, and A. man- 
ni, are restricted to small basins on its 
western slope. 

In this paper two new species of Aegla, 
both from localities in the Chilean Coastal 
Cordillera between Concepcion and Puerto 
Montt, are described. 

Specimens are deposited in the Crusta- 
cean Collection of the Instituto de Zoologia, 
Universidad Austral de Chile (IZUA C), 
Valdivia, Chile. The size of the specimens 
was recorded as carapace length (CL), i.e., 
distance between rostral apex and posterior 
margin of cephalothorax. 



VOLUME 112, NUMBER 1 



107 




Fig. 1. Aegla cholchol, new species, male allotype, Chol-Chol River, IZUA C-328-B, dorsal view. 



Aegla cholchol, new species 
Figs. 1, 2a-i 

Aegla bahamondei. — Jara, 1982:232 (in 
part), Fig. 1 (see Remarks). 

Type locality. — Chol-Chol River, 100 m 
downriver from bridge in the town of Chol- 
Chol, 38°36'40"S, 72°51'05"W, 29 km NW 
of Temuco, Province of Cautin, Chile. 

Type material. — Holotype: female (24.5 
mm CL), C. G. Jara coll., 22 Dec 1982, 
IZUA C-328-B. Allotype: male (22.4 mm 
CL). Paratypes: 9 females (17.5-30.3 mm 



LC), 6 males (14.9-27.2 mm LC), same 
data as holotype. 

Other material. — 3 females (21.5-23.5 
mm LC), Quepe River, 38°51'40"S, 
72°37'02"W, C. A. Viviani coll., 24 Feb 
1969, IZUA C-008-B. 3 males (24.3-26.1 
mm LC), 2 females (23.8-25.6 mm LC), 
Quepe River at Boroa, 39°49'06"S, 
72°53'02"W, C. G. Jara coll., 21 Dec 1982, 
IZUA C-297-B. 7 females (18.1-23.4 mm 
LC), Traiguen River, 38°14'45"S, 
72°39'40"W, C. G. Jara & R. Navarro coll., 
22 Dec 1982, IZUA C-316. 3 males (26.9- 



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Fig. 2. Aegla cholchol, new species, male allotype, Chol-Chol River, IZUA C-328-B; a, anterior part of 
cephalothorax, dorsal view; b, anterior part of cephalothorax, lateral view; c, third and fourth sterna; d, second 
abdominal epimeron; e, telson plate; f, ischium of left cheliped, ventral view; g, carpus of left cheliped, ventral 
view; h, left cheliped, dorsal view; i, merus of cheliped, lateral view. Scales: d = 1 mm; a, b, c, e, f, g, i = 5 
mm; h = 10 mm. 



VOLUME 112, NUMBER 1 



109 



18.9 mm LC), 5 females (26.4-15.5 mm 
LC), Colpi River near Galvarino, 
38°19'52"S, 72°47'10"W, C. G. Jara & R. 
Navarro coll., 22 Dec 1982, IZUA C-317- 
B. 2 males (19.3-29.3 mm LC), 6 females 
(23.6-26.0 mm LC), Quepe River, 
38°51'40"S, 72°37'02"W, C. G. Jara & R. 
Navarro coll., 21 Dec 1982, IZUA C-320- 

A. 8 males (13.6-30.8 mm LC), 9 females 
(14.3-21.8 mm LC), Lumaco River at Lu- 
maco, 38°09'46"S, 72°54'21"W, C. G. Jara 
& R. Navarro coll., 22 Dec 1982, IZUA C- 
321. 5 males (15.8-25.5 mm LC), 9 females 
(17.3-24.1 mm LC), Cautin River, Temuco, 
38°44'30"S, 72°35'00"W, C. G. Jara & R. 
Navarro coll., 23 Dec 1982, IZUA C-322- 

B. 8 males (11.7-21.9 mm LC), 8 females 
(17.8-24.7 mm LC), Chol-Chol River near 
Galvarino, 38°36'40"S, 72°51'05"W, C. G. 
Jara & R. Navarro coll., 22 Dec 1982, 
IZUA C-324-C. 12 males (12.4-27.3 mm 
LC), 9 females (12.5-21.2 mm LC), Lu- 
maco River at Pellahuen, 38°25'40"S, 
72°56'00"W, C. G. Jara & R. Navarro coll., 
22 Dec 1982, IZUA C-326-A. 1 juvenile 
(5.6 mm LC), Pichicautin River near Lu- 
maco, 38°09'25"S, 72°54'15"W, T. Gonser 
coll., 16 Jan 1983, IZUA C-424. 

Diagnosis. — Carapace slightly longer 
than wide, sparsely setose, moderately ex- 
panded at branchial level. Rostrum long, 
subtriangular, acute, styliform at distal half. 
Orbital spine prominent; extraorbital sinus 
wide, U-shaped. Anterolateral angle of first 
hepatic lobe spiniform. Anterior branchial 
margin subdenticulated; posterior finely 
serrated, dorsally upturned. Anterolateral 
angle of second abdominal epimeron spi- 
niform. Fourth thoracic sternum with blunt 
medial tubercle, occasionally with 1 or 2 
scales on frontal border. Telson divided. 
Adult males markedly heterochelous; fe- 
males weakly so. Chelae robust; propodus 
inflated though dorso-ventrally flattened; 
dactylar lobe blunt; palmar crest subrect- 
angular or arcuate, expanded, dentated; pal- 
mar lobe low, blunt, with scales only on 
frontal edge; medial scale largest; carpal 



lobe with prominent conical acute spine. 
Middorsal carpal ridge tubercles blunt. 

Description. — Carapace slightly longer 
than wide, somewhat setose especially on 
branchiostegal and pterygostomial areas. 
Abdominal terga and middorsal line of mer- 
us of second to fourth pereopods also se- 
tose. Precervical portion narrower than 
postcervical; apex of acute pyramidal bran- 
chial lobe separated from margin of third 
hepatic lobe by wide notch. Rostrum long, 
acute, narrow, styliform, slightly expanded 
at its proximal %\h but conical and some- 
what flattened on distal l/gth, gently re- 
curved. Rostral tip with acute scale. Rostral 
borders scarcely marginated, bearing mi- 
nute scales along %\h of rostrum length, na- 
ked distally. Rostral carina prominent and 
scaly on proximal half of rostrum length, 
absent on distal half; scales small, button- 
like, in 2 subparallel rows. Epigastric prom- 
inences forming low, smooth, arcuate ridge, 
broadest just behind deepest point of orbital 
sinus. Protogastric prominences forming 
low tuberculiform area bearing short 
oblique row of tiny stud-like scales. An- 
terolateral lobe of carapace and hepatic 
lobes well defined; former elongated as 
straight conical spine reaching up to middle 
of adjacent cornea. Gastric area broad, pos- 
terior more protuberant. Anterolateral angle 
of first hepatic lobe spiniform; spine point- 
ing straight forward. Branchial lobe pyra- 
midal, elongated, acutely tipped, well de- 
tached from margin of third hepatic lobe 
and from anterolateral angle of anterior 
branchial area; with single irregular row of 
acute scales on outer border. Branchial mar- 
gins moderately expanded; anterior subden- 
ticulated, posterior finely serrated; denticles 
on anterior margin short, ill-defined, except 
anteriormost which is as large as branchial 
lobe; each denticle carrying 3 to 4 tiny 
acute scales, mingled with short stiff setae. 
Dorsum of anterior branchial area flat to 
slightly concave; posterior branchial area 
and adjacent marginal cardial area dome 
shaped; free border of posterior branchial 
area recurved and upturned. Anterolateral 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



angle of second abdominal epimeron spi- 
niform; spine apex scarcely detached from 
and at level with posterior branchial mar- 
gin. Lateral angle of third to fifth abdominal 
epimera spiniform. Fourth thoracic sternum 
with blunt median tubercle, occasionally 
carrying 1 or 2 scales on frontal border. Tel- 
son longitudinally articulated. Chelae ro- 
bust; propodus inflated though dorso-ven- 
trally flattened. Dactylar lobe, at proximo- 
dorsal end of dactylus, blunt, with 1 to 4 
tiny scales in a row along apex. Palmar 
crest subrectangular or arcuate, moderately 
expanded, border coarsely dentated; teeth 
subtriangular, recurved, apices point front- 
dorsally, especially the most distal. Palmar 
lobe, just behind dactylus-propodus joint, 
blunt, low, with acute scales only on frontal 
edge, its medial scale generally largest; 
frontal border of palmar crest overhanging 
palmar lobe, separated by deep notch. Car- 
pal lobe prominent, massive, supporting ro- 
bust, conical, acute, somewhat recurved 
spine. Dorsointemal border of carpus, be- 
hind carpal lobe, with neat row of 2 to 3 
large conical acute spines, decreasing in 
size towards rear. Carpal ridge as arcuate 
row of blunt knob-like tubercles topped by 
minute scale and/or short setae. Carpus ven- 
tral face with robust conical spine. Merus 
of cheliped with row of blunt to spiniform 
tubercles, distalmost largest; dorsodistal an- 
gle of merus tuberculiform, bearing group 
of short stiff setae, and 2 to 4 scales of 
which median is largest. 

Color. — Live, with dorsum of carapace 
and abdomen olive green; sternal surface 
and ventral face of pereopods creamy 
white. Dorsum of pereopods with alternat- 
ing yellow green and dark green bars. Tip 
of largest spines pale yellow to creamy, 
darkening to green at their base. Joints of 
pereopods bright red, minutely spotted 
white. In alcohol, carapace and abdomen 
smoky tan to creamy white. 

Geographic range (Fig. 3). — Found on 
the Chol-Chol River basin, from Lumaco to 
Chol-Chol; also in the Cautin River, in front 
of Temuco, and in the Quepe River near 



Quepe; all these are tributaries of the Im- 
perial River basin which discharge into the 
Pacific Ocean, at Puerto Saavedra. 

Habitat. — Aegla cholchol is found in me- 
dium sized rivers (order 2 to 3, after Strah- 
ler, 1957) with coarse gravel and boulder 
substrates where water current velocity 
varies between 0.1 to 0.7 m/sec. As a con- 
sequence of intensive agricultural activity, 
the fluvial environment of some stretches of 
the Chol-Chol River basin have undergone 
severe alterations; the riparian vegetation, 
mostly non-native species, is in a hetero- 
geneous state of conservation. At these 
sites, specimens of A. cholchol, new spe- 
cies, were collected on coarse gravel to fine 
quartzitic micaceous sand, sharing the bio- 
tope with A. spectabilis. 

Etymology. — The specific name is de- 
rived from the name of the type locality, as 
a latinized noun used in apposition. It is 
treated as indeclinable, for the purposes of 
Article 31b of the International Code of 
Zoological Nomenclature (International 
Trust for Zoological Nomenclature 1985). 

Affinities. — Aegla cholchol, new species, 
A. rostrata, and A. bahamondei are similar 
in appearance but the new species is more 
similar to A. rostrata than to A. bahamon- 
dei. Common to the three species is a wide 
carapace with expanded branchial areas and 
denticulated or subdenticulated branchial 
borders; steep inward downward inclination 
of the branchiostegal surface; dorsum of an- 
terior branchial area subconcave; well de- 
fined branchial lobe; protuberant gastric and 
cardiac areas with elevated middorsal line 
but non-keeled as in A. denticulata; acute 
first hepatic angle and anterolateral angle of 
carapace; ample front and orbital sinuses 
limited by acute orbital spines, one on the 
plane of the orbital border and one just be- 
low the former, usually not visible from 
above; narrow, slightly troughed, elongated 
and conically tipped rostrum. In each of the 
three species, the appendages, especially 
the chelipeds, present a basic morphologi- 
cal pattern with proper modifications. The 
new species differs from A. rostrata in hav- 



VOLUME 112, NUMBER 1 



111 




••— International BORDER 

REOIONAL BORDER 



25 50 75 



100 Km 



Fig. 3. Map showing the geographical distribution of Aegla cholchol, new species. Asterisks indicate local- 
ities where specimens were collected. In some cases, asterisk indicates more than one sampling site. 



112 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ing: blunt tubercles on the carpal ridge; dor- 
sodistal angle of merus of chelipeds, sec- 
ond, and third pereopods blunt, or at most, 
with small round tubercle tipped by minute 
scales and/or setae; dactylar or palmar lobes 
not spiniform; edge of the anterior branchi- 
al area subdenticulated, teeth apressed non 
spine-like; fourth thoracic sternum with me- 
dian tubercle. Aegla cholchol, new species, 
differs from A. bahamondei by having: epi- 
branchial lobe monocuspidate; dactylar 
lobe low and blunt; protogastric promi- 
nences little protuberant; tubercle on fourth 
thoracic sternum acute but non- spiniform; 
posteroventral edge of merus of second to 
fourth pereopods smooth, non-denticulated; 
surface of carapace and chelae with few mi- 
nute scales. 

Remarks. — A. cholchol, new species, A. 
rostrata, and A. bahamondei share a series 
of complex characters. In particular, a ten- 
dency to have every protruding angle of the 
carapace, abdomen, and appendages, as a 
pointed conical tubercle or spine. However, 
the spines and spiniform tubercles are cer- 
tainly more developed in A. rostrata than 
in A. cholchol, new species, and A. baha- 
mondei. In the latter, the angles of the car- 
apace are acute but never spiniform. Jara 
(1986:40) argued that "the trend toward 
profuse spinulation seems correlated with 
living in lentic environments." In this con- 
text, both A. cholchol, new species, and A. 
bahamondei are restricted to rivers while A. 
rostrata is found mostly in lakes. Jara 
(1982:237) indicated that "some specimens 
. . . found in Quepe River near Temuco . . . 
could prove to be conspecific with ... [A. 
bahamondeiy\ On reexamination, those 
specimens proved to be A. cholchol, new 
species (IZUA C-008-B, formerly C-328). 

Aegla hueicollensis, new species 
Figs. 4, 5a-i 

Type locality. — Pichihueicolla River, 0.5 
km SE from its outlet into the Pacific Ocean 
(40°10'S, 73°40'W), on the western slope 



of Cordillera Pelada (Coastal Range), Prov- 
ince of Valdivia, Chile. 

Type series. — Holotype: male (16.5 mm 
CL), 26 Jan 1983, C. G. Jara coll., IZUA 
C-245. Allotype: female (16.5 mm CL). 
Paratypes: 7 males (10.9-19.7 mm LC), 3 
females (14.1-14.6 mm LC), 2 juveniles 
(7.3-7.7 mm LC), same data as holotype. 

Other material. — 3 males (10.7-14.1 
mm LC), 3 females (19.3-20.7 mm LC), 
Chiveria, Cordillera Pelada, 40°04'50"S, 
73°10'50"W, R. Formas coll., 14 Jan 1971, 
IZUA C-025. 5 females (12.1-18.1 mm 
LC), Hueicolla River, 40°08'30"S, 
73°37'42"W, R. Formas coll., 1974, IZUA 
C-026. 14 males (8.9-18.3 mm LC), 7 fe- 
males (1 1.7-16.4 mm LC), Amargos Creek, 
Corral, Valdivia, 39°52'24"S, 72°26'25"W, 
C. G. Jara coll., 19 Mar 1974, IZUA C-042. 
1 male (13.4 mm LC), 3 females (6.8-13.2 
mm LC), Trainel River, Lago Huillinco, 
Chiloe, H. J. Wetzlar coll, 07 Oct 1975, 
IZUA C-131. 20 males (10.1-24.8 mm 
LC), 11 females (7.8-2L3 mm LC), San 
Carlos Creek, San Carlos, Corral, Valdivia, 
39°51'40"S, 73°26'25"W, C. G. Jara coll, 20 
Mar 1976, IZUA C-175. 9 males (11.7- 
24.2 mm LC), 4 females (18.3-19.9 mm 
LC), Chiveria, Cordillera Pelada, 
40°04'20"S, 73°10'30"W, R. P Schlatter 
coll., 21 Jan 1977, IZUA C-184. 3 males, 
(12.3-18.3 mm LC), 9 females (10.9-17.6 
mm LC), Hueicolla River, 40°08'30"S, 
73°39'05"W, R. P Schlatter coll., 22 Jan 
1977, IZUA C-185. 18 males (10.5-16.8 
mm LC), 8 females (10.2-14.6 mm LC), 
Manzano River, Todos los Santos Lake, 
Llanquihue, 41°07'20"S, 72°24'50"W, G. 
Milhe coll., 26 Apr 1974, IZUA C-196. 3 
males (11.3-14.6 mm LC), 4 females 
(11.3-15.9 mm LC), Refugio River, Chiloe, 
R. P Schlatter coll., 11 Oct 1977, IZUA C- 
197. 2 males (8.3-23.6 mm LC), 1 female 
(26.4 mm LC), Roblental Creek, eastern 
slope of Cordillera Pelada, 40°16'S, 
73°12'W, C. G. Jara coll., 12 Feb 1978, 
IZUA C-206. 1 male (14.8 mm LC), 3 fe- 
males (15.7-21.4 mm LC), Pichihueicolla 
River, 40°09'40"S, 73°39'50"W, C. G. Jara 



VOLUME 112, NUMBER 1 



113 




mm 



Fig. 4. Aegla hueicollensis, new species, male holotype, Pichihueicolla River, IZUA C-245, dorsal view. 



coll., 26 Jan 1982, IZUA C-246. 27 males 
(4.5-15.9 mm LC), 22 females (7.9-17.6 
mm LC), Hueicolla River, 40°09'07"S, 
73°38'25"W, C. G. Jara coll., 27 Jan 1982, 
IZUA C-247. 5 males (10.3-19.7 mm LC), 
9 females (10.1-22.1 mm LC), Chaihmn 
River, 40°05'07"S, 73°2r51"W, C. G. Jara 
coll., 15 Feb 1984, IZUA C-363. 10 males 
(4.2-27.2 mm LC), 22 females (6.9-26.3 
mm LC), Collin River, 40°02'31"S, 
73°31'26"W, C. G. Jara coll., 15 Feb 1984, 



IZUA C-364. 6 males (7.0-17.2 mm LC), 
6 females (8.5-15.1 mm LC), Hueicolla 
River, 40°08'30"S, 73°31'38"W, H. F Jara 
coll, 07 Feb 1992, IZUA C-507. 6 males 
(9.6-16.7 mm LC), 1 female (13.9 mm 
LC), Hueicolla River, 40°08'27"S, 
73°30'39"W, H. E Jara coll, 08 Feb 1992, 
IZUA C-508. 

Diagnosis. — Carapace ovoidal, not ex- 
panded at branchial areas. Epibranchial 
lobe pyramidal, acute, borders scaly. Ros- 



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Fig. 5. Aegla hueicollensis, new species, male holotype Pichihueicolla River, IZUA C-245; a, anterior part 
of cephalothorax, dorsal view; b, anterior part of cephalothorax, lateral view; c, third and fourth sterna; d, second 
abdominal epimeron; e, telson plate; f, ischium of left cheliped, ventral view; g, carpus of left cheliped, ventral 
view; h, left cheliped, dorsal view; i, merus of cheliped, lateral view. Scales: c, d, f, i = 1 mm; a, b, e, g, h = 
5 mm. 



VOLUME 112, NUMBER 1 



115 



trum short, wide at base, neatly triangular, 
both sides of low-profiled carina barely 
troughed, conical at apex, acute. First he- 
patic lobe blunt. Orbital and extraorbital si- 
nus present. Branchial margin smooth, 
slightly nodulated. Second abdominal epi- 
meron acute but not spiniform. Telson di- 
vided. Fourth thoracic sternum smooth. 
Male chelae elongated, pyriform, inflated at 
propodus; palmar crest flabelliform, sub- 
rectangular, dentate; dactylar lobe subspi- 
niform; carpal lobe spiniform, apex mark- 
edly displaced towards distal end of article, 
frontal edge with row of tufts of short setae 
mingled with scales; tubercles of carpal 
ridge blunt, low, topped by transversal row 
of scales and setae. Tubercles on dorsal 
edge of merus of chelipeds blunt, tipped by 
2 to 4 scales roughly ordered in transversal 
oblique row. Ventroextemal border of is- 
chium, ventral face of carpus, and ventral 
borders of chelae with tufts of long setae in 
row. 

Description. — Carapace surface finely 
punctated, each pit with a minute scale and 
1 or 2 short stiff setae; highest density of 
setae on abdominal epimera. Precervical 
carapace almost as wide as postcervical. 
Epibranchial lobe pyramidal, acute, its edg- 
es lined with short acute stout scales; sim- 
ilar scales along outer borders of carapace 
except for posterior border and external half 
of orbital sinus. Rostrum short, wide at 
base, neatly triangular, barely troughed; 
low-profiled carina merges into rostrum 
body at proximal half; rostral apex conical, 
with stout acute apical scale encircled at 
base by 3 to 4 minor scales; dorsum of ca- 
rina with double row of tiny scales proxi- 
mally, and single unordered row distally; 
ventral carina not particularly prominent, its 
deepest point at base. Epigastric promi- 
nences nodular, blunt, topped with 3 to 4 
scales in transverse row; protogastric prom- 
inences barely discernible, marked by small 
field of button-like scales mingled with stiff 
setae. Anterolateral lobe of carapace later- 
ally expanded, flattened proximally, its ex- 
ternal border slightly sinuose; anterolateral 



angle of carapace short, acute, its apex 
reaching up to proximal third of adjacent 
cornea. Orbital sinus wide, delimited by an 
inconstantly present orbital spine; extraor- 
bital sinus variable in depth and width, gen- 
erally narrowly V-shaped. First hepatic lobe 
blunt, well separated from base of antero- 
lateral lobe; second and third lobes scarcely 
defined, separated by shallow notches in- 
terrupting marginal row of scales. Branchial 
margin narrow, smooth, slightly nodulated; 
posterior slightly upturned. Areola broad, 
convex, elevated over remaining cardial 
area. Thoracic sterna flat; fourth sternum 
with frontal fringe of long stiff setae; sel- 
dom with low, blunt, median tubercle, car- 
rying 1 or 2 scales. Anterolateral angle of 
second abdominal epimeron acute, dorsum 
subcarinate behind anterolateral angle; la- 
teroventral face of epimeron deeply con- 
cave, profusely covered by short stiff setae; 
free ventral border with dense row of long 
plumose setae, also present over posterior 
branchiostegal surface, edge of remaining 
abdominal epimera, and borders of uropods 
and telson. Telson longitudinally articulat- 
ed. 

Males heterochelous; left hand largest. 
Chelae robust but not particularly massive, 
subovoidal in outline, surface microtuber- 
culate; each tubercle carrying a minute 
acute scale and 2 short stiff microsetae, 
most evident on distal half of propodus and 
dactylus. Molar process well developed on 
cutting edge of fixed finger and dactylus of 
left chela but negligible or absent on right 
one. Dorsum of dactylus with robust spi- 
niform lobe on proximal end. Ventral face 
of chelae with prominent ridge along ven- 
troextemal border of propodus, marked by 
row of pits and tufts of long setae; pits more 
numerous along dactylus and fixed finger. 
Base of palmar crest on ventral side of che- 
lae parallel to short tuberculate ridge with 
large pits and tufts of setae. Central part of 
ventral face of chelae broadly convex, 
forming longitudinal ridge between center 
of propodus-carpus joint and distalmost 
dactylus-propodus joint, marked by large 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



pits and setae. Palmar crest prominent, 
thick, its border denticulated with scattered 
small acute scales between contiguous larg- 
er denticles. Distal end of palmar crest sep- 
arated from predactylar lobe of carpus by 
almost right-angled deep notch; predactylar 
lobe elevated, its free border with irregular 
row of short acute scales; largest scale at 
anterodorsal angle. Carpus robust, with two 
dorsal ridges; outermost lower, distinguish- 
able as slightly arcuate row of broad low 
tubercles; apex of tubercles with row of 3 
to 6 short stiff setae mingled with minute 
scales; innermost prominent, formed by 
row of blunt tubercles topped by tranverse 
row of 3 to 5 acute scales and setae. Carpal 
lobe spiniform, asymetrically subtriangular, 
apex leveled with frontal border of same 
lobe. Inner border of carpus with row of 3 
large spines proximally decreasing in size. 
Dorsum of anteriormost spine with 2 to 3 
scales or setae protruding from large pits in 
a row along spine axis. Same arrangement 
on minor spines or at least, 1 pit and setae 
present. Ventral surface of carpus with ro- 
bust conical spine; between it and inner 
border of carpus several thick, long, simple 
setae, uneven in size. Dorsal edge of merus 
with row of subacute tubercles tipped by 1 
or 2 scales, 3 or 4 on distalmost tubercle; 
frontal side of tubercles with tufts of long 
simple setae; frontodorsal angle blunt, 
thick, little prominent, with patch of short 
stiff setae and scales; ventral borders with 
row of tubercles tipped with scale and/or 
seta; distal angles as short conical spine. 
Dorsal edge of merus of second, third and 
fourth pereopods with fringe of long plu- 
mose setae covering minute spiniform tu- 
bercles; carpus, propodus and dactylus with 
fringe of short stiff simple setae. Postero- 
ventral edge of merus scabrous, subserrat- 
ed, with narrow band of setae and scales. 

Color. — Not observed live. In alcohol, 
carapace, abdomen, and appendages uni- 
formly smoky-tan to creamy white. 

Geographic range (Fig. 6). Small 
streams and creeks flowing from east to 
west on the western slope of Cordillera Pe- 



lada (Coastal Range between Corral to the 
north and the outlet of the Bueno River to 
the south). Specimens collected are from: 
Amargos Creek (Corral, Valdivia), Colun, 
Chaihuin, HueicoUa, and PichihueicoUa riv- 
ers. Also found in Manzano River, at Todos 
Los Santos Lake, in the Andean district of 
the Llanquihue province. 

Habitat. — Small rivers and fast flowing 
streams on moderate to steep slopes, cov- 
ered or surrounded by cold-temperate Val- 
divian rain forest. Specimens are found 
among pebbles and boulders on the stream 
bottom, at sites where a heterogeneous 
mass of vegetal detritus accumulates serv- 
ing as shelter for the crabs. 

Etymology. — The specific name derives 
from Hueicolla, a small town on the Pacific 
coast about 40 km to the south from Corral, 
where the rivers Hueicolla and Pichihuei- 
coUa discharge into the Pacific Ocean. 

Affinities. — Aegla hueicollensis, new spe- 
cies, shares a series of features with other 
species of Aegla. In general, the new spe- 
cies closely resembles A. abtao. Several 
morphological similarities suggest a close 
ancestral relationship between the two. A. 
hueicollensis, new species, also displays 
morphological similarity to A. alacalufi. 
Both share the almond-shaped outhne of 
the carapace, the short triangular acute ros- 
trum, and the pronounced distalward dis- 
placement of the apex of the carpal lobe. 
However, A. alacalufi has an undivided tel- 
son plate, and lacks minute scales on the 
carapace surface. The new species also 
shares features with A. manni and A. neu- 
quensis Schmitt, 1942b, such as: the irreg- 
ular row of scales and/or setae along the 
dorsal axis of the larger spines on the inner 
border of carpus of chelipeds and the trans- 
verse row of scales on the tubercles of the 
carpal ridge of chelae. The new species dif- 
fers from A. manni in having the rostral ca- 
rina and rostral margins scaly; from A. neu- 
quensis by having the anterolateral angle of 
second abdominal epimeron subspinose. 
The low, somewhat concave, profile of the 
rostral carina and the protuberant gastric 



VOLUME 112, NUMBER 1 



117 




Fig. 6. Map showing the geographical distribution of Aegla hueicollensis, new species. Asterisks indicate 
localities where specimens were collected. In some cases, asterisk indicates more than one sampling site. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



area of A. hueicollensis, new species, re- 
semble those of A. concepcionensis. These 
similarities led Schmitt (1942b:504) to re- 
port specimens of Aegla from Corral 
(northern limit of Cordillera Pelada) as A. 
concepcionensis. Similarly, Haig (1955: 
504) classified as A. concepcionensis spec- 
imens from Ancud (230 km to the south of 
Corral), Chiloe Island. To determine if these 
specimens are conspecific with A. hueicol- 
lensis, new species, they would have to be 
re-examined. 

Remarks. — The geographic range of A. 
hueicollensis, new species, includes a series 
of localities on the western slope of the 
Coastal Cordillera and one locality (Man- 
zano River) on the western slope of the An- 
des Cordillera, about 100 km southeast of 
the type locality. There is no hydrological 
connection between these areas at present. 
This fact leads to the assumption that, in 
the past, the geographic range of the new 
species must have been broader, enclosing 
localities on the intervening territory be- 
tween the Andean and the Coastal ranges. 
Connected to this is the fact that southern 
Chile, from paralell 38°S southward, was 
severely affected by Quaternary glaciations. 
lilies (1960) found evidence that during 
glaciations the ice-sheet front advanced 
west from the Andean highlands completely 
covering the Central Valley; the ice front 
pushed upon the eastern slope of the Coast- 
al Range but never surmounted it. The 
western slope of the Coastal Range which 
was never completely covered by ice (lilies 
1960) could have served as a refugial area 
for A. hueicollensis, new species. Its pres- 
ence in the small Manzano River basin, 
amid the Andean massif, presumes that 
somehow the site was unaffected by de- 
structive glacial effects, thus preserving the 
preexisting population. 

Acknowledgments 

The authors thank Drs. R. Formas, T 
Gonser, H. F. Jara, R. Navarro, and R. R 
Schlatter for the presentation of valuable 



material; Dr. Rafael Lemaitre and two 
anonymous reviewers for their efforts to 
improve the readability of the manuscript. 
The present paper was financed by grants 
91-0900 of the Fondo de Investigacion 
Cientifica y Tecnologica (Fondecyt, Chile) 
and S-91-4 of the Direccion de Investiga- 
cion y Desarrollo, Universidad Austral de 
Chile. 

Literature Cited 

Bahamonde, N., & M. T Lopez. 1963. Decapodos de 
aguas continentales en Chile. — Investigaciones 
Zoologicas Chilenas 10:123-149. 

Borgel, R. 1983. Geomorfologia. in Geografia de 
Chile, 2. Institute Geografico Militar, Santiago, 
Chile, 182 pp. 

Di Castri, F. 1968. Esquisse ecologique du Chili. Biol- 
ogie de TAmerique Australe. Pp. 7-82 in C. 
Delamare Deboutteville & E. Rapoport, eds.. 
Biologic de TAmerique Australe, 4. Editions du 
Centre National de la Recherche Scientifique, 
Paris, 685 pp. 

Haig, J. 1955. The Crustacea Anomura of Chile. — Re- 
ports of the Lund University Chile Expedition 
1948-49, 20. Lund Universitets Arsskrift. N.F 
Avd. 2, 5 1(12): 1-68. 

lilies, H. 1960. Geologic der Gegend von Valdivia 
(Chile). — Neues Jahrbuch fiir Geologic und Pa- 
lacntologie 111:30-110. (Translated into Span- 
ish by J. Walper. Universidad Austral de Chile, 
Valdivia, 1970, 50 pp.) 

International Trust for Zoological Nomenclature. 1985. 
International Code of Zoological Nomenclature, 
Third edition, London, 338 pp. 

Jara, C. 1977. Aegla rostrata, n. sp. (Decapoda, Ae- 
glidae), nuevo crustaceo dulceacuicola del Sur 
de Chile. — Studies on Neotropical Fauna and 
Environment 12:165-176. 

. 1980. Dos nuevas especies de Aegla Leach 

(Crustacea, Decapoda, Anomura) del sistema 
hidrografico del Rio Valdivia. — Anales del Mu- 
seo de Historia Natural de Valparaiso 13:255- 
266. 

. 1982. Aegla bahamondei, new species (Crus- 
tacea: Decapoda: Anomura) from the Coastal 
Mountain Range of Nahuelbuta, Chile. — Jour- 
nal of Crustacean Biology 2(2):232-238. 

. 1986. Aegla spectabilis, a new species of 

freshwater crab from the eastern slope of the 
Nahuelbuta Coastal Cordillera, Chile. — Pro- 
ceedings of the Biological Society of Washing- 
ton 99:34-41. 

. 1989. Aegla denticulata lacustris, new sub- 
species, from Lake Rupanco, Chile (Crustacea: 
Decapoda: Anomura: Aeglidae). — Proceedings 



VOLUME 112, NUMBER 1 



119 



of the Biological Society of Washington 102: 
385-393. 

— . 1992. Aegla expansa, new species (Crustacea: 
Decapoda: Anomura: Aeglidae), from the Low- 
er Bio-Bio river basin, Concepcion, Chile. — 
Gayana (Zoologia) 56(l-2):49-57. 

— . 1994. Aegla pewenchae, new species of Cen- 
tral Chilean freshwater decapod (Crustacea: An- 
omura: Aeglidae). — Proceedings of the Biolog- 
ical Society of Washington 107:325-339. 

-, & M. T. Lopez. 1981. A new species of fresh- 
water crab (Crustacea: Anomura: Aeglidae) 
from insular South Chile. — Proceedings of the 
Biological Society of Washington 94:88-93. 



Nicolet, H. 1849. Crustaceos. Pp. 113-318 in C. Gay, 
Historia Fisica y Politica de Chile, Zoologia 3, 
547 pp. 

Schmitt, W. L. 1942a. Two new species of Aegla from 
Chile. — Revista Chilena de Historia Natural 
44(1940):25-31, pi. 1. 

. 1942b. The species of Aegla, endemic South 

American freshwater crustaceans. — Proceedings 
of the United States National Museum 91:431- 
520. 

Strahler, A. N. 1957. Quantitative analysis of water- 
shed geomorphology. — Transactions of the 
American Geophysicists Union 38:13-920. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):120-132. 1999. 

The Hawaiian parthenopid crabs of the genera 

Garthambrus Ng, 1996, and Dairoides Stebbing, 1920 

(Crustacea: Decapoda: Brachyura) 

Peter K. L. Ng and S. H. Tan 

Department of Biological Sciences, National University of Singapore, Lower Kent Ridge Road, 

Singapore 119260, Republic of Singapore 

Abstract. — The taxonomy of four species of deep-water parthenopid crabs, 
Garthambrus stellata (Rathbun, 1906), G. lacunosa (Rathbun, 1906), G. com- 
planata (Rathbun, 1906) and Dairoides kusei (Sakai, 1938) from Hawaii is 
treated. Dairoides kusei is recorded from Hawaii for the first time. The iden- 
tities of G. stellata, G. lacunosa and G. complanata are clarified, with detailed 
comparisons and figures provided. Despite earlier reports, there are no inter- 
mediates between these three species. A key to the genus Garthambrus is 
provided. 



Rathbun (1906), in her treatment of the 
Hawaiian Brachyura, had originally recog- 
nized eight parthenopid species from these 
islands. One species, Harrovia truncata 
Rathbun, 1906, was transferred to a sepa- 
rate genus (Cyrtocarcinus Ng & Chia, 
1994) in the Xanthidae by Ng & Chia 
(1994). Another species, Parthenope (Pla- 
tylambrus) stellata Rathbun, 1906, was 
originally established with three subspecies, 
P. (P.) stellata stellata, P. (P.) stellata la- 
cunosa Rathbun, 1906, and P. (P.) stellata 
complanata Rathbun, 1906. Garth (1993) 
reappraised the validity of these three taxa 
and regarded them as distinct species. He 
briefly commented that the features used by 
Rathbun (1906) to separate the three vari- 
eties were good species characters. Al- 
though Garth examined the type series of 
all three taxa, he did not comment much 
about them or figure any of the types. This 
despite the fact that one taxon, P. (P. ) com- 
planata, had never been figured before. Nor 
did he comment on specimens which Rath- 
bun (1906) had regarded as being interme- 
diate between the three species. In describ- 
ing a new species (Parthenope cidaris) 
from Australia, Garth & Davie (1995) pro- 



vided photographs of the types of P. (P.) 
stellata, P. (P.) lacunosa and P. (P.) com- 
planata but did not make any additional 
comments other than again stating briefly 
that all three taxa were good species. 

Sakai (1938, 1976) had commented that 
P. (P.) lacunosa (as a subspecies) was a 
junior synonym of Lambrus (Parthenopo- 
ides) pteromerus Ortmann, 1893, a species 
which he transferred to the genus Tutankh- 
amen Rathbun, 1925. Garth (1993), how- 
ever, disagreed, commenting that P. (P.) la- 
cunosa lacked the lamellar ridges lining the 
afferent channels on the carapace found on 
Tutankhamen cristatipes (A. Milne Ed- 
wards, 1880), the type and only species of 
the genus. On the basis of Garth's (1993) 
redescription of P. (P.) stellata and his 
comments on P. (P.) lacunosa and P. (P.) 
complanata, Ng (1996) subsequently trans- 
ferred all three species to the genus Gar- 
thambrus Ng, 1996. 

Nine species of Parthenopidae are cur- 
rently known from the Hawaiian islands, 
Garthambrus stellata (Rathbun, 1906), G. 
lacunosa (Rathbun, 1906), G. complanata 
(Rathbun, 1906), Platylambrus nummifera 
(Rathbun, 1906), Rhinolambrus lamelligera 



VOLUME 112, NUMBER 1 



121 



(White, 1847), Aulacolambrus hoplonotus 
(Adams & White, 1848), Aulacolambrus 
whitei (A. Milne Edwards, 1878), Pseudo- 
lambrus calappoides (Adams & White, 
1848), and Daldorfia horrida (Linnaeus, 
1758). The generic classification used here 
follows that proposed by Ng & Rodriguez 
(1986) and Ng (1996). 

The present paper reports on the deep- 
water Parthenopidae from Hawaii primarily 
based on material deposited in the Bemice 
P. Bishop Museum, Honolulu; and National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. Garthambrus 
stellata, G. lacunosa and G. complanata are 
rediagnosed, and detailed figures of their di- 
agnostic characters provided for the first 
time. Using new characters identified in this 
study, the "intermediates" reported by 
Rathbun (1906) can easily be assigned to 
the three species. A key to the genus Gar- 
thambrus is provided. The unusual deep- 
water species, Dairoides kusei (Sakai, 
1938), previously known only from Japan, 
is also reported from Hawaii for the first 
time. 

The terms used here essentially follow 
those used by Garth (1958, 1993). The ab- 
breviations cw and cl are for the carapace 
width and length, and Gl and G2 are for 
the male first and second pleopods respec- 
tively. Specimens are deposited in the Bem- 
ice P. Bishop Museum, Honolulu (BPBM); 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM); Nationaal Naturhistorish Muse- 
um, Leiden (ex Rijksmuseum van Natuur- 
lijke Historic, RMNH); and the Zoological 
Reference Collection, National University 
of Singapore (ZRC). 

Systematic Account 
Genus Garthambrus Ng, 1996 

Garthambrus Ng, 1996: 156 (type species: 
Garthambrus posidon Ng, 1996, by orig- 
inal designation). 

Diagnosis. — Carapace subtriangular in 
shape, broader than long; angle between an- 



tero- and posterolateral margins strongly 
produced; dorsal surfaces granulose, spi- 
nose or rugose; progastric, mesogastric, 
metagastric, mesobranchial, metabranchial, 
cardiac and intestinal regions strongly in- 
flated; gastric and branchial regions sepa- 
rated by deep grooves. Median lobe of ros- 
trum prominent, sub-spatulate, deflexed 
downwards. Hepatic region and margin 
separated from anterolateral margin by 
cleft, notch or large tubercle. Posterolateral 
margin and metabranchial regions without 
long or prominent teeth or spines. Cheli- 
peds at least 2.5 times carapace length. Gl 
relatively stout, not armed with long spines 
or stiff hairs. Distal segment of G2 elon- 
gate, subequal to or distinctly longer than 
basal segment (modified from Ng 1996: 
156). 

Remarks. — Ng (1996) removed a number 
of species previously assigned to Parthen- 
ope, Platylambrus or Parthenope (Platy- 
lambrus) to a separate genus which he char- 
acterised by their triangular carapaces and 
long distal segment of the second male ple- 
opod. All eight species (see key) currently 
placed in Garthambrus are essentially deep- 
water species. 

One unusual species which requires men- 
tion is the Japanese species Parthenope 
pteromerus (Ortmann, 1893). It is very sim- 
ilar in appearance to G. lacunosa, which 
Sakai (1976) synonymised under P. pter- 
omerus. Sakai (1976) also transferred P. 
pteromerus to the genus Tutankhamen 
Rathbun, 1925, arguing that the afferent re- 
spiratory channels on its carapace are lined 
with lamellar ridges. Garth (1993) refuted 
this synonymy because G. lacunosa and all 
Garthambrus species do not have such la- 
mellar ridges. Subsequently, Ng (1996) 
suggested that P. pteromerus was not as- 
signable to Tutankhamen, and probably be- 
longed in a separate genus. While P. pter- 
omerus resembles many Garthambrus spe- 
cies superficially, the possession of lamellar 
ridges lining the afferent respiratory chan- 
nels on its carapace argues against its in- 
clusion in Garthambrus as presently de- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



fined. Parthetiope pteromerus is also a rel- 
atively shallow-water species compared to 
known species of Garthambrus (unpub- 
lished data). 

Key to Garthambrus species 

1 a. Carapace and legs covered with numer- 
ous long, sharp spines; margin of bran- 
chial region with deep, distinct cleft . . 
G. mironovoi 

lb. Carapace and legs not covered with nu- 
merous long, sharp spines; margin of 
branchial region entire or almost so . . 2 

2a. Granules on branchial, cardiac and gas- 
tric regions of carapace fused to various 
degrees forming distinct granular 
patches G. allisoni 

2b. Carapace dorsal surface almost smooth 
or if granules present, those on bran- 
chial, cardiac and gastric regions never 
distinctly fused 3 

3a. Margins of ambulatory meri distinctly 

cristate G. lacunosa 

3b. Margins of ambulatory meri not dis- 
tinctly cristate, but lined with spines, 
teeth or granules 4 

4a. Carapace and cheliped surfaces without 

pits or lacunae G. complanata 

4b. Carapace and cheliped surfaces with 

distinct pits and/or lacunae 5 

5a. Margins of ambulatory meri with 

prominent rounded granules 6 

5b. Margins of ambulatory meri with un- 
even granules, tubercles or small spines 7 

6a. Median protuberance of rostrum large, 
margins of ambulatory propodi and car- 
pi distinctly granulated G. poupini 

6b. Median protuberance of rostrum small, 
margins of ambulatory propodi and car- 
pi smooth, unarmed G. posidon 

7a. Dorsal surface of carapace densely 
granulated all over; metabranchial re- 
gions strongly swollen, appears peak- 
like from frontal view G. stellata 

7b. Dorsal surface of carapace with rugae 
and granules but never densely packed; 
metabranchial regions high but not very 
swollen, not peak-like from frontal 
view G. cidaris 



Garthambrus stellata (Rathbun, 1906) 
(Figs. lA-D, 6A-C) 

Parthenope (Platylambrus) stellata Rath- 
bun, 1906:884 (part), pi. 15 Figs. 1, 2.— 
Garth, 1993:786, Figs. 3, 4. 

Parthenope (Platylambrus) stellatus. — Se- 
rene, 1968:60. 

Parthenope stellata. — Garth & Davie, 
1995:225, Fig. 2B. 

Garthambrus stellata. — Ng, 1996:158. 

Material examined. — Holotype, 1 male, 
cw 48.6 mm, cl 32.8 mm (USNM 29839), 
south coast of Oahu Island, Hawaii, 435- 
461 m, station 3811, coll. Albatross Expe- 
dition, 27 Mar 1902. Paratype, 1 male, cw 
32.9 mm, cl 23.1 mm (USNM 29840), Ka- 
waihae, Hawaii Island, Hawaii, 20°01'45"N 
155°54'15"W, 269-362 m, station 4045, 8 
hemp tangles gear, coll. Albatross Expedi- 
tion, 11 Jul 1902. Others — 1 female 
(USNM 239148), Pearl Harbor, Mamala 
Bay, Oahu Island, Hawaii, disposal site, 
2ri6'48"N 157°56'30"W, 366 m, coll. RA^ 
Hurl, 30 Apr 1983. 1 male, cw 70.2 mm, 
cl 45.6 mm (ZRC 1997.441), Off Pearl Har- 
bor, Oahu Island, Hawaii, station 82-105, in 
dredge spoil site, on sediment bottom in vi- 
cinty of outcrop, 366 m, coll. RA^ Hurl, 
Makali'i Dive, D. M. Devaney & B. Bart- 
ko, 1 Sep 1982. 1 female (carapace only) 
(BPBM 1978.417), Off Makupuu Point, 
Molokai Channel, Hawaii, 12 km straight 
cast, 2r02'N 157°32'W, coll. Valkryrien, 
Capt. S. Rayner, 20 May 1967. 1 male, ca. 
cw 65.5 mm, cl 43.4 imn (left edge of car- 
apace broken) (BPBM 1976.259), Entrance 
to Pearl Harbor, Oahu Island, Hawaii, 2.5 
km off buoy 1,338 m, coll. in shrimp traps, 
Easy Rider, E. H. Chave, 27 Sep 1976. 2 
females, cw 21.6 mm, cl 16.5 mm, ca. cw 
16.0 mm, cl 12.5 mm (right side of cara- 
pace crushed) (BPBM 5508), Oahu Island, 
1.9 km off Kahala, Hawaii, 46 m, coll. 
Brock, 14 Apr 1949. 

Diagnosis. — Surface of carapace densely 
and granulated uniformly throughout, in- 
cluding grooves and depressions; metabran- 
chial regions strongly swollen, appearing 



VOLUME 112, NUMBER 1 



123 




Fig. 1. Garthambrus stellata (Rathbun, 1906). Holotype male (48.6 by 32.8 mm) (USNM 29839). A. Car- 
apace; B, Abdomen; C, Front; D, Left chela. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Garthambrus lacunosa (Rathbun, 1906). Holotype male (30.9 by 21.8 mm) (USNM 29842). A, 
Carapace; B, Abdomen. 




Fig. 3. Male telson and sixth abdominal segment. 
A, Garthambrus stellata, holotype male (USNM 
29839); B, Garthambrus lacunosa, holotype male 
(USNM 29842). C, Garthambrus lacunosa, male 
(USNM 29842b). 



peak-like in frontal view (Fig. IC), highest 
point with sharp granule; sub-branchial, 
suborbital, subhepatic and branchiostegal 
regions distinct, densely covered with dis- 
tinct small granules. Anterolateral margin 
with sharp, acutely triangular teeth, lateral 
margins of teeth with numerous accessory 
spinules and/or small, sharp granules. Che- 
Hped surfaces (especially merus, carpus, 
propodus and dactylus) with well devel- 
oped sharp spines, spinules and granules 
(Fig. ID); merus with oblique ridge of 
strong spines about % from proximal end; 



VOLUME 112, NUMBER 1 



125 




Fig. 4. Garthambrus complanata (Rathbun, 1906). Lectotype male (17.3 by 12.7 mm) (USNM 29845a). A, 
Carapace; B, Abdomen. 



surfaces never with pits or lacunae. Margins 
of ambulatory legs unevenly cristate to den- 
tate, with numerous accessory spinules; sur- 
faces densely covered with small granules, 
never with pits or lacunae. Anterior thoracic 



stemites densely covered with small gran- 
ules, without pits or lacunae (Fig. IB). Ab- 
dominal surfaces densely covered with 
small granules, never with pits or lacunae; 
lateral margins of male telson gently con- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



vex. Gl relatively slender, tip distinctly 
turned inwards towards median part of tho- 
racic sternum. 

Remarks. — Rathbun (1906) described 
Parthenope stellata on the basis of ten 
specimens from various parts of Hawaii. In 
her discusion on variation within the spe- 
cies, however, she noted that only two spec- 
imens conformed with P. stellata sensu 
stricto, and the other eight specimens ap- 
peared to belong to two other varieties or 
were intermediate in form (see discussion 
for next two species). One specimen 
(USNM 29839) was designated as the 
"type" {= holotype) of P. stellata. How- 
ever, Rathbun (1906:884), noted that the 
smaller specimen (USNM 29840) "... 
shows the tubercles and spines all sharp in- 
stead of blunt pointed, and lacks the hair 
near the ends of the legs." She regarded 
this smaller specimen as belonging to the 
"sharp-spined variety" of P. stellata sensu 
stricto, but did not apply any name. Ex- 
amination of a relatively good series of 
specimens has shown that differences noted 
by Rathbun (1906) between her two type 
specimens can easily be accounted for by 
infraspecific variation. Edmondson (1951: 
213) had reported two small specimens 
from shallow waters in Hawaii and Ng 
(1996:159) commented that "... his spec- 
imens are probably misidentified" as the 
carapace proportions were different, the 
rostrum was not trifid and the ambulatory 
dactylus was more styliform. We have re- 
examined these two specimens (BPBM 
5508), and they are clearly juvenile G. stel- 
lata. The differences in carapace propor- 
tions are almost certainly due to their small 
size and the ambulatory dactylus is actually 
not as styliform as depicted in Edmondson 's 
(1951:Fig. 18d) illustration. The bifid ros- 
trum shown for the specimen figured by 
Edmondson (1951:Fig. 18a) is such because 
the median lobe is broken. In the other 
specimen, the rostrum is clearly trifid. 

The relatively good series of specimens 
of various sizes and sexes shows that the 
characters identified here are consistent and 



are useful for defining this species. The Gl 
shows slight variation, in that the distal part 
may appear slightly more flared or slightly 
more bent inwards (towards the median part 
of the thoracic sternum). The density of the 
granules does not change much with 
growth. However, with an increase in size, 
the individual granules do become larger, 
more prominent and distinctly more stellate 
in structure. 

Garthambrus stellata is known from Ha- 
waii, Tuamotu and Taiwan (Garth 1992, 
Poupin 1996:32, Tan et al. 1999). In Ha- 
waii, the species is recorded up to depths 
of 461 m, although juveniles have been 
found in shallower waters of only about 46 
m. The record from Tuamotu is of an adult 
from about 300 m depth. The specimen fig- 
ured by Poupin (1996: pi. 15f) from Tua- 
motu is a dirty white overall and speckled 
with numerous fine brown spots, and the 
fingers are pigmented brown. 

Garthambrus lacunosa (Rathbun, 1906) 
(Figs. 2A-B, 3A-C, 6D-F) 

Parthenope (Platylambrus) stellata lacu- 
nosa Rathbun, 1906:884, pi. 15 Fig. 7. 

Parthenope (Platylambrus) lacunosa. — 
Garth, 1993:788. 

Parthenope lacunosa. — Garth & Davie, 
1995:226, Fig. 3A. 

Garthambrus lacunosa. — Ng, 1996:158 

Material examined. — Holotype, 1 male, 
cw 30.9 mm, cl 21.8 mm (USNM 29842), 
Kawaihae, west coast of Hawaii Island, Ha- 
waii, 269-362 m, 20°01'45"N 155°54' 
15"W, station 4045, 8 hemp tangles gear, 
coll. Albatross Expedition, 11 Jul 1902. 
Paratypes, 1 male, cw 30.4 mm, cl 31.6 mm 
(USNM 29843a), Maui Island, Lipoa Point, 
Pailolo Channel, Hawaii, 238-276 m, 
2r05'30"N 156°40'30"W, station 4100, 8 
hemp tangles gear, coll. Albatross Expedi- 
tion, 23 Jul 1902, (1 other paratype male 
specimen transferred to Stanford University 
according to labels, not examined). 1 fe- 
male (USNM 29844), northwest coast of 
Oahu Island, Hawaii, 282-357 m, coll. Al- 



VOLUME 112, NUMBER 1 



127 



batross Expedition. 1 female, cw 30.2 mm, 
cl 21.3 mm (USNM 29841), south coast of 
Molokai Island, Hawaii, off Lae-O-Ka Laau 
Lighthouse, 309-333 m, 64°00'N 
00°13'42"W, station 3835, coll. Albatross 
Expedition, 3 Apr 1902. Others — 1 male, 
cw 31.4 mm, cl 21.5 mm (USNM 29843b), 
Maui Island, Lipoa Point, Pailolo Channel, 
Hawaii, 238-276 m, 2r05'30"N 
156°40'30"W, station 4100, 8 hemp tangles 
gear, coll. Albatross Expedition, 23 Jul 
1902. 

Diagnosis. — Surface of carapace slightly 
rugose, with scattered to relatively numer- 
ous granules; metabranchial regions mod- 
erately raised but not strongly swollen, not 
peak-like from frontal view, highest point 
unarmed; sub-branchial, suborbital, subhe- 
patic and branchiostegal regions distinct, 
covered with numerous but scattered small 
granules. Anterolateral margin with lobi- 
form to truncate teeth, lateral margins of 
teeth lined with numerous accessory spi- 
nules or small, sharp granules especially 
along anterior margin. Outer surfaces of 
chelipeds (especially merus, carpus, pro- 
podus and dactylus) with well developed 
spines, spinules and granules; merus with 
oblique ridge of strong spines about V3 from 
proximal end; outer surfaces with numerous 
pits, often with well developed lacunae. 
Margins of ambulatory legs distinctly cris- 
tate, entire to uneven; surfaces almost 
smooth or with very small, scattered gran- 
ules, usually with distinct pits and/or lacu- 
nae. Anterior thoracic stemites with numer- 
ous scattered small granules and pits, often 
with lacunae. Abdominal surfaces covered 
with numerous scattered small granules and 
pits, often with lacunae; lateral margins of 
male telson concave. Gl relatively slender, 
tip turning slightly inwards towards median 
part of thoracic sternum. 

Remarks. — Rathbun (1906) regarded five 
specimens of the original nine specimens of 
Parthenope (Platylambrus) stellata belong- 
ing to a separate variety P. (P.) stellata la- 
cunosa. She has also mentioned that the dif- 
ferences of these five specimens are so dif- 



ferent from the type specimen of G. stel- 
lata, that they could be a different species 
altogether and she mentioned several char- 
acters: "The branchio-cardiac depression is 
deep, and another depression runs along the 
outer side of the branchial region, adjacent 
to the marginal teeth. The elevated part of 
this region has a row of large pits through 
its middle, and similar line of pits dividing 
the gastric region in three and roughening 
the chelipeds. The granules are in large part 
confluent and thus obliterated, especially on 
the higher parts of the carapace and the che- 
lipeds. The legs have smooth surfaces, thin 
cristate margins which are somewhat cre- 
nate or dentate in the merus and are desti- 
tute of long hair. Along with two of this 
variety from station 4100 is one which is 
intermediate between the typical {P. stellata 
stellata) and varietal form (P. stellata la- 
cunosa), the stellate granules being every- 
where fairly well shown, and also the lines 
of pits." (Rathbun, 1906:884). 

One of the original nine type specimens 
of Garthambrus stellata was considered as 
intermediate between P. stellata and P. la- 
cunosa. This specimen (USNM 29843b) is 
similar to P. stellata sensu stricto in having 
smaller granules overall, even on the cara- 
pace depressions and grooves, compared to 
typical P. lacunosa. The lacunae on the car- 
apace and chelipeds are also smaller and 
less developed. A close examination of this 
supposedly intermediate specimen, howev- 
er, shows that it only represents the extreme 
end of the variation in P. lacunosa. In the 
presence of lacunae on the carapace and 
chelipeds, truncate anterolateral teeth, dis- 
tinctly cristate ambulatory meri, low, non- 
peaked metabranchial regions which lack a 
sharp median granule and gently concave 
male telson (Fig. 3C), the specimen clearly 
represents P. lacunosa. Even though this 
specimen is more granulated than typical P. 
lacunosa, it is still much less granulated 
overall compared to any specimen of P. 
stellata sensu stricto we have examined, 
even specimens of similar sizes. 

Rathbun (1906) identified one specimen 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



(USNM 29842) as the "type" (= holotype) 
of Garthambrus lacunosa. The other four 
specimens listed by Rathbun (1906) are 
thus paratypes. The supposedly intermedi- 
ate specimen (USNM 29843b) was not re- 
garded as belonging to P. (P.) stellata la- 
cunosa by Rathbun (1906) when she orig- 
inally named the taxon, and thus, is not part 
of the type series. 

Garthambrus complanata (Rathbun, 1906) 
(Figs. 4A-B, 6G-I) 

Parthenope (Platylambrus) stellata com- 
planata Rathbun, 1906:884. 

Parthenope (Platylambrus) stellata com- 
planataus Serene, 1968:60. 

Parthenope {Platylambrus) complanata. — 
Garth, 1993:789. 

Parthenope complanata. — Garth & Davie, 
1995:226, Fig. 3B. 

Garthambrus complanata. — Ng, 1996:158. 

Material examined. — Lectotype, herein 
designated, 1 male, cw 17.3, cl 12.7 mm 
(USNM 29845a), Hanamaulu Bay, Kauai 
Island, Hawaii, 470-570 m, 22°01'30"N 
150°2riO"W, station 4132, 8 foot Blake 
Beam trawl gear, coll. Albatross Expedi- 
tion, 1 Aug 1902. Paralectotype, 1 male 
(USNM 29845b), same data as lectotype. 

Diagnosis. — Surface of carapace smooth 
to gently rugose, not granulated; metabran- 
chial regions raised but not strongly swol- 
len, not peak-like from frontal view, highest 
point unarmed; sub-branchial, suborbital, 
subhepatic and branchiostegal regions dis- 
tinct but almost smooth except for a few 
scattered granules. Anterolateral margin 
with liboform to truncate teeth, lateral mar- 
gins of teeth lined with scattered accessory 
spinules or small, sharp granules especially 
along anterior margin. Surface of chelipeds 
(especially merus, carpus, propodus and 
dactylus) with few, scattered granules, but 
with distinct simple spines; merus without 
distinct oblique ridge of spines; surfaces 
with few, scattered simple granules. Mar- 
gins of ambulatory legs smooth, not cristate 
but with well spaced spines and/or teeth; 



surfaces smooth, without lacunae. Anterior 
thoracic stemites almost smooth, without 
pits or lacunae. Abdominal surfaces almost 
smooth, without pits or lacunae; lateral 
margins of male telson concave. Gl rela- 
tively stout, straight. 

Remarks. — For her third variety of P. 
(P.) stellata, Rathbun (1906:884) noted that 
"Still a third type seems worthy of a dis- 
tinguishing name, P. (P.) stellata complan- 
ata. It differs from the type of P. (P.) stel- 
lata stellata in the surface of carapace and 
chelipeds being smooth to the naked eye, 
though under the lens finely punctate and 
roughened; the tubercle or spine at the inner 
third of the postero-lateral margin is repre- 
sented by a triangular nodule; tubercle at 
each end of posterior margin large and 
round; antero-lateral teeth broader and more 
dentiform than in other forms; no teeth nor 
spines at outer end of postero-lateral mar- 
gin, but a nodule on the dorsal surface at 
that point may represent them; marginal 
spines of chelipeds inclining to sharp; legs 
approaching the type in roughness; margins 
prominently spinate, without long hair." 
Most of these differences are valid, and G. 
complanata can easily be separated from 
congeners by the carapace sculpturation, ar- 
mature of the antero-lateral margin, absence 
of an oblique ridge on the relatively smooth 
merus of the cheliped, structure of the am- 
bulatory merus, and form of the thoracic 
stemites and abdominal surface. 

Garth & Davie (1995:Fig. 3B) provided 
a photograph of the "holotype" of this spe- 
cies, but no measurements were provided. 
However, Rathbun (1906) listed two male 
specimens (USNM 29845a) only as 
"types" without designating a holotype; 
thus, both are syntypes. Garth (1993) had 
earlier examined both specimens and listed 
them as syntypes. The larger specimen 
(17.3 by 12.7 mm) (USNM 29845a) in bet- 
ter condition, is here designated as the lec- 
totype. It is the same specimen figured by 
Garth & Davie (1995). 



VOLUME 112, NUMBER 1 



129 




Fig. 5. Dairoides kusei (Sakai, 1938). 1 male (52.2 by 47.3 mm) (ZRC 1997.447). 



Genus Dairoides Stebbing, 1920 

Dairoides Stebbing, 1920:233 (type spe- 
cies: Dairoides margaritus Stebbing, 
1920, by monotypy). 

Astherolambrus Sakai, 1938:341 (type spe- 
cies: Astherolambrus kusei Sakai, 1938, 
by monotypy). 

Remarks. — Sakai (1938) established 
Astherolambrus for a new species from Ja- 
pan, A. kusei. Sakai (1965) later synony- 
mised Astherolambrus under Dairoides 
Stebbing, 1920, which had been established 
for D. margaritus Stebbing, 1920, a species 
known only from South Africa. Takeda & 
Ananpongsuk (1991) subsequently de- 
scribed a third species, D. seafdeci, from 
the Andaman Sea in the Indian Ocean. 

Dairoides kusei (Sakai, 1938) 
(Figs. 5, 6 J-L) 

Astherolambrus kusei Sakai, 1938:341, pi. 
41 Figs. 5, 6. 



Dairoides kusei. — Sakai, 1965:99, pi. 45 
Fig. 2.— Sakai, 1976:288, pi. 97 Figs. 1, 

2. 

Material examined. — 1 male, 2 females 
(BPBM 1980.194), Oahu Island, off Bar- 
ber's Point, Hawaii, 117-128 m, trapped by 
gill nets laid overnight, coll. Teritu, T. 
Clarke, 19-20 Apr 1971. 1 male, cw 52.2 
mm, cl 47.3 mm (ZRC 1997.447), Hawaii, 
coll. E. Bilderback, 17 Jan 1979. 1 male 
(dried), cw 54.8 mm, cl 44.6 mm (RMNH), 
Wagu, Kii Peninsula, Mie Prefecture, Ja- 
pan, coll. N. Yamashita, 1978-79. 2 males, 
1 female (RMNH 32004), Wagu, Kii Pen- 
insula, Mie Prefecture, Japan, coll. N. Ya- 
mashita, 1978-79. 

Remarks. — Dairoides kusei had previ- 
ously been reported only from Japanese wa- 
ters. The specimens reported here agree 
well with the descriptions and figures of 
this species by Sakai (1938, 1965, 1976). 
We have also examined four specimens of 
D. kusei from Japan in the RMNH donated 



130 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Gls. A-C. Garthambrus stellata (Rathbun, 1906), holotype male (48.6 by 32.8 mm) (USNM 29839); 
D-F, G. laciinosa (Rathbun, 1906), holotype male (30.9 by 21.8 mm) (USNM 29842); G-I, G. complanata 
(Rathbun, 1906), lectotype male (17.3 by 12.7 mm) (USNM 29845a); J-L, Dairoides kusei (Sakai, 1938), 1 
male (52.2 by 47.3 mm) (ZRC 1997.447). 



VOLUME 112, NUMBER 1 



131 



by the late Tune Sakai. They agree with the 
specimens from Hawaii in all major as- 
pects, including the structures of their gon- 
opods. 

Acknowledgments 

The first author is most grateful to Lu 
Eldredge for his excellent hospitality during 
his visit to the BPBM. Thanks are also due 
to Rafael Lemaitre and Lipke Holthuis for 
their kind assistance in examining the spec- 
imens in the USNM and RMNH respec- 
tively. This study has been partially sup- 
ported by grant RP 3972371 of the National 
University of Singapore to the first author. 

Literature Cited 

Adams, A., & A. White. 1848. Crustacea. In A. Ad- 
ams, ed.. The zoology of the voyage of the 
H.M.S. Samarang; under the command of Cap- 
tain Sir Edward Belcher, C.B., ER.A.S., EG.S. 
during the Years 1843-1846. Viii + 66 pp., pis. 
1-13. Reeve, Benham & Reeve, London. [Pp. 
1-32 and pis. 1-6 were published in 1848; pp. 
i-viii, 33-66 and pis. 7-13 in 1849] 

Edmondson, C. H. 1951. Some central Pacific crusta- 
ceans. — Occasional Papers of the Bernice P. 
Bishop Museum, 20:183-243. 

Flipse, H. J. 1930. Oxyrhyncha: Parthenopidae. — Die 
Decapoda Brachyura der Siboga-Expedition, 
VL Siboga-Expeditie, 39c2(l 12):l-96. 

Garth, J. S. 1958. Brachyura of the Pacific Coast of 
America Oxyrhyncha. — Allan Hancock Pacific 
Expeditions, 21, part 1:1-499; part 2:677-854, 
pis. A-Z, 1-55. 

. 1993. Some deep-water Parthenopidae (Crus- 
tacea, Brachyura) from French Polynesia and 
nearby eastern Pacific Ridges and Seamounts. — 
Bulletin du Museum national d'Histoire natu- 
relle, Paris, 1992, (4)14(A3-4):781-795. 

, & P. J. F. Davie. 1995. A new species of Pa r- 

thenope (Crustacea: Decapoda: Brachyura) 
from deep-water off northern Queensland. — 
Memoirs of the Queensland Museum. 38(1): 
223-227. 

Linnaeus, C. 1758. Systema naturae per regna tria na- 
turae, secundum classes, ordines, genera, spe- 
cies cum characteribus, differentiis, synonymies 
locis. 10th Edition, 1:1-854, Holmiae. 

Milne Edwards, A. 1878, (in 1873-1881) Etudes sur 
les Xiphosures et les Crustaces de la region 
Mexicaine. In Mission scientifique au Mexique 
et dans TAmerique centrale, Recherches Zoo- 
logiques pour servir a I'histoire de la faune de 



I'Amerique centrale et du Mexique 5:1-368, 
pis. 1-61. 
-. 1880. Etudes preliminaires sur les Crustaces, 



lere partie. In Reports on the results of dredging 
under the supervision of Alexander Agassiz, in 
the Gulf of Mexico, and in the Caribbean Sea, 
1877, '78, '79, by the U.S. Coast Survey 
Steamer "Blake", Lieut. Commander C. D. 
Sigsbee, U.S.N., and Commander Bartlett, 
U.S.N., commanding. — Bulletin of the Museum 
of Comparative Zoology, Harvard College, 
8(l):l-68, pis. 1, 2. 

Ng, P. K. L. 1996. Garthambrus, a new genus of deep- 
water parthenopid crabs (Crustacea: Decapoda: 
Brachyura) from the Indo-Pacific, with descrip- 
tion of a new species from the Seychelles. — 
Zoologische Mededelingen, Leiden, 70(10): 
155-168. 

, & D. G. B. Chia. 1994. The genus Glypto- 

carcinus Takeda, 1973, with descriptions of a 
new subfamily, two new genera and two new 
species from New Caledonia (Crustacea: De- 
capoda: Brachyura: Xanthidae). — Raffles Bul- 
letin of Zoology, 42(3):701-730. 

, & G. Rodriguez. 1986. New records of Mim- 



ilambrus wileyi Williams, 1979 (Crustacea: De- 
capoda: Brachyura), with notes on the system- 
atics of the Mimilambridae Williams, 1979 and 
Parthenopoidea MacLeay, 1838 sensu Guinot, 
1978. — Proceedings of the Biological Society 
of Washington 99(1): 88-99. 

Ortmann, A. 1893. Die Decapoden-Krebse des Strass- 
burger Museums, mit besonderer Beriicksichti- 
gung der von Herm Dr. Doderlein bei Japan und 
bei den Liu-Kiu-Inseln gesammelten und zur 
Zeit im Strassburger Museum aufbewahrten 
Formen. VII. Theil. Abtheilung: Brachyura 
(Brachyura genuina Boas) II. Unterabtheilung 
Cancroidea, 2. Section: Cancrinea. 1. Gruppe 
Cyclometopa. — Zoologische Jahrbuchner, 7 
411-495, pi. 17. 

Poupin, J. 1996. Atlas des Crustaces Marins profonds 
de Polynesie Fran^aise. Recoltes du navire 
MARARA (1986/1996).— Service Mixte de 
Surveillance Radiologique et Biologique, Mon- 
tlhery, Cedex, France, pp. 1-59. pis. 1-20. 

Rathbun, M. J. 1906. The Brachyura and Macrura of 
the Hawaiian Islands. — United States Fisheries 
Commission Bulletin. 3[1903]:827-930, pis. 1- 
24. 

. 1925. The spider crabs of America. — Bulletin 

of the United States National Museum, 129:1- 
613, pis. 1-283. 

Sakai, T 1938. Brachygnatha, Oxyrhyncha. In Studies 
on the crabs of Japan 3:193-364, Figs. 1-55, 
pis. 20-41. 

. 1965. The Crabs of Sagami Bay collected by 

His Majesty the Emperor of Japan. Maruzen 



132 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Co., Tokyo, pp. 1-206. Figs. 1-27, pis. 1-100. 
1 map. 
-. 1976. Crabs of Japan and the adjacent seas. 



Kodansha Ltd., Tokyo. 3 vol. Volume 1 [En- 
glish text]: pp. 1-773, Figs. 1-379, maps 1-3; 
Volume 2 [Japanese text]: pp. 1-461, Figs. 1- 
2; Volume 3 [plates]: pp. 1-61, pis. 1-251. 
Serene. R. 1968. The Brachyura of the Indo-West Pa- 
cific region. Pp. 33-112 in Prodromus for a 
Check List of the Non-Planctonic Marine Fauna 
of South East Asia. — Singapore National Acad- 
emy of Science, Special publication number 1: 
1-120. 



Stebbing, T. R. R. 1920. South African Crustacea, Part 
X. — Annals of the South African Museum 
17(4):23 1-272, pi. 1. 

Takeda, M.. & S. Ananpongsuk 1991. A new deep-sea 
crab from the Andaman Sea off Thailand. — 
Bulletin of the National Science Museum, Se- 
ries A (Zoology) 17(2):93-100. 

Tan, S. H., J.-F Huang. & P K. L. Ng. 1999. Crabs of 
the Family Parthenopidae (Crustacea, Decapo- 
da, Brachyura) from Taiwan. — Zoological Stud- 
ies. Taipei (in press). 

White, A. 1847. Descriptions of new Crustacea from 
the eastern seas. — Proceedings of the Zoologi- 
cal Society of London 15:56-58. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):133-140. 1999. 

Reinstatement and further description of Eualus subtilis Carvacho & 

Olson, and comparison with E, lineatus Wicksten & Butler 

(Crustacea: Decapoda: Hippolytidae) 

Gregory C. Jensen and Rachel C. Johnson 

(GCJ) School of Fisheries, Box 357980, University of Washington, Seattle, Washington 98195, 
U.S.A.; (RCJ) National Marine Fisheries Service, Southwest Fisheries Center, 

Tiburon, California 94920, U.S.A. 

Abstract. — Eualus subtilis Carvacho & Olson 1984 is presently considered 
a synonym of E. lineatus Wicksten & Butler 1983. Comparisons of the two 
forms revealed differences in the armature of the antennular peduncle, stylo- 
cerite, and pereopods, as well as differences in color and maximum size, in- 
dicating that E. subtilis is a valid species. Furthermore, E. subtilis, unlike E. 
lineatus, exhibits marked sexual dimorphism, and characteristics of the previ- 
ously undescribed, diminutive males of this species are provided. 



The genus Eualus Thallwitz, 1891 is 
comprised of relatively small caridean 
shrimps that occur primarily in the higher 
latitudes. Butler (1980) illustrated a small, 
striped shrimp as E. herdmani (Walker, 
1898), and mentioned that it could be 
trawled in small numbers in Departure Bay, 
Canada. However, subsequent examination 
of the holotype and only known specimen 
of E. herdmani by Wicksten & Butler 
(1983) revealed that E. herdmani belonged 
in the genus Heptacarpus (Holmes, 1900) 
and the specimens illustrated by Butler 
(1980) were described as a new species, E. 
lineatus Wicksten & Butler, 1983. One of 
the characters distinguishing E. lineatus 
was the presence of three moderate dorso- 
distal spines on the first article of the an- 
tennular peduncle (hereafter referred to as 
"antennular spines"). 

A second description of a small eualid, E. 
subtilis Carvacho & Olson, 1984 was pub- 
lished based on a single specimen trawled 
off Baja California. As Carvacho & Olson 
did not mention E. lineatus, and still referred 
to *'£. herdmani'', they were evidently un- 
aware of Wicksten & Butler's (1983) paper. 
Eualus subtilis strongly resembled E. linea- 
tus in the size and armature of the rostrum. 



but had only a single, stout, dorsolateral an- 
termular spine. Wicksten (1988) considered 
E. subtilis to fall within the range of varia- 
tion of E. lineatus and thus a junior syno- 
nym, and used the record as a southern range 
extension for E. lineatus. 

Subtidal sampling in the Puget Sound re- 
gion revealed an extremely abundant small 
eualid that matched the description of E. 
subtilis. However, out of hundreds of spec- 
imens examined, all had a single antennular 
spine. Based on the length of the rostrum 
some of these specimens keyed out to E. 
pusiolus (Kr0yer, 1841), yet they bore sev- 
eral strong, distal spines on the merus of 
the walking legs while E. pusiolus has only 
a single spine (Squires 1990). Furthermore, 
this species exhibited marked sexual di- 
morphism, whereas there is no distinct dif- 
ferences between sexes in E. pusiolus 
(Greve 1963). In view of these observa- 
tions, a study was undertaken to determine 
if E. subtilis is distinct from E. lineatus, and 
provide information about the unusual, 
small males. 

Materials and Methods 

Due in part to the ambiguity of existing 
keys, museum specimens of these shrimps 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



have been variously cataloged under the 
names E. herdmani, E. pusiolus, or E. li- 
neatus. These specimens needed to be re- 
examined. Sixty-one specimens from the 
British Columbia Provincial Museum col- 
lection were examined, as was the holotype 
of E. lineatus (AHF 4129) deposited in the 
Natural History Museum of Los Angeles 
County and the paratype from the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM). The 
two paratype s reportedly deposited in the 
National Museum of Canada were not sent 
at the time E. lineatus was described and 
now cannot be located. Additional material 
was acquired from Friday Harbor Labora- 
tories of the University of Washington (43 
specimens), the California Academy of Sci- 
ences (8 specimens), and our own collec- 
tions ranging from the Pribilof Islands to 
Puget Sound (45 specimens). Twenty-two 
North Atlantic specimens of E. pusiolus 
from the Royal Norwegian Society of Sci- 
ences were examined for comparison. 

Live specimens having a single anten- 
nular spine were abundant in shell rubble 
and easily collected while diving by care- 
fully placing dead bivalve shells in a fine- 
meshed bag. Hand-operated suction devices 
were used to sample in rock crevices. 

Measurements were taken using an im- 
age analysis system (Optimus®) on a Wild 
MC3 dissecting microscope, with additional 
measurements taken using an ocular micro- 
meter. Carapace length (cl) was measured 
from the posterior margin of the orbit to the 
middorsal posterior margin of the carapace; 
rostrum length, from the same position on 
the orbit to the tip. Sex, rostral formula, 
number of distal spines on the basal article 
of the antennular peduncle, and relative 
lengths of the stylocerite and rostrum were 
also noted, as was the number of meral 
spines on pereopods 3-5. Drawings were 
made with the aid of a camera lucida on a 
Wild® M5 microscope. 

Results 

Of the 159 shrimp measured, only 17 
were found bearing multiple antennular 



spines, and these were compared with 93 
specimens that had only a single antennular 
spine and multiple spines on the merus of 
the third pereopod. The remaining 49 spec- 
imens were identified as E. pusiolus based 
on published descriptions and comparison 
with the specimens from Norway. 

The 17 specimens with multiple anten- 
nular spines reached substantially larger 
sizes than those bearing a single spine (Fig. 
la, b). Multiple antennular spines were also 
correlated with a reduced number of meral 
spines on the third pereopod. Those with 
multiple antennular spines typically had 
only one meral spine, although some spec- 
imens had a second, much reduced spine 
present on at least one side. Shrimp with a 
single antennular spine had significantly 
more meral spines (x" = 81.21; p ^ 0.001), 
the great majority bearing three strong mer- 
al spines and the remainder varying from 
two to five (Fig. 2). 

Those with multiple antennular spines 
also had a curved, dorsal tooth near the 
base of the stylocerite not previously noted 
in the literature (Fig. 3); that tooth was 
lacking in those with a single antennular 
spine. A blunt suborbital carapace spine 
was also present in those with multiple an- 
tennular spines, and missing in those with 
a single antennular spine and more than one 
spine on the merus. 

Males with a single antennular spine typ- 
ically had a very thin, bifid rostrum (Fig. 
4c); the largest male (2.1 mm cl) was small- 
er than any of the ovigerous females (2.6- 
3.8 mm). Unlike that of females, the pro- 
podus of pereopods 3-5 of males was dis- 
tinctly broadened distally and armed with 
two rows of spines forming a dense comb 
on the flexor margin, with spines increasing 
in length distally (Fig. 4a); the male dac- 
tylus was also armed with an unusual series 
of compound spines on the flexor margin 
(Fig. 4b). The appendix masculina was sub- 
equal in length to the appendix interna, and 
tipped with two long spinules and five short 
ones. Males comprised only 10% of the 
samples. 



VOLUME 112, NUMBER 1 



135 



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postorbital carapace length (mm) 



Fig. 1. Scatterplot of postorbital carapace length vs. proportional rostrum length for specimens ha%ing a 
single distodorsal spine on the basal article of the antennular peduncle (a), and specimens with multiple disto- 
dorsal spines on the basal article of the antennular peduncle (b). 



Only two males with multiple antennular 
spines were available for examination, but 
other than the presence of an appendix mas- 
culina they did not appear to differ from 
females with multiple spines. The appendix 
masculina in these specimens was slightly 
more than half the length of the appendix 
interna and tipped with eight long spinules, 
as previously described (Wicksten & Butler 
1983; Fig. 2d). 



The form with only a single antennular 
spine consistently displayed the color pat- 
tern described for E. lineatiis (Wicksten & 
Butler, 1983) as shown in Butler (1980: 
plate IC), having thin red diagonal lines on 
the carapace and first two abdominal seg- 
ments and red spotting on the remainder of 
the abdomen. In contrast, a live specimen 
of the form that has multiple antennal 
spines was boldly marked on both the car- 



136 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



80 n 



60- 



S 
I 40 



20- 






r 



i ■ 



number of 

antennular 

spines 

Q nrYultiple 
Q single 



12 3 4 5 
number of leg 3 merus spines 

Fig. 2. Bar graph showing number of lateral spines 
on the merus of the third pereopod for specimens with 
single (n = 85) or multiple (n = 12) distodorsal spines 
on the first article of the antennular peduncle. 



apace and abdomen with broad orange 
bands against a translucent background 
(Fig. 5). 

The holotype of E. lineatus has multiple 
antennular spines, a suborbital carapace 
spine, dorsal tooth on the stylocerite, and 
two spines on the merus of the third pereo- 
pod, whereas the USNM paratype of E. li- 
neatus has only a single antennular spine, 
no suborbital spine or stylocerite tooth, and 
three strong spines on the merus of the third 
pereopod. 

Discussion 

The combination of size, color and mor- 
phological differences clearly indicate that 
E. subtilis should be considered a valid spe- 
cies, distinctly separate from E. lineatus. 
Eualus subtilis lacks the multiple dorsodis- 
tal spines on the basal article of the anten- 
nular peduncle, the suborbital carapace 
spine, and the dorsal tooth on the stylocerite 
that are all present in E. lineatus. Further- 
more, the largest E. subtilis barely exceed 
half the length of E. lineatus, and E. subtilis 
exhibits marked sexual dimorphism while 
E. lineatus does not. The number of spines 



on the merus of the third walking leg is 
usually reliable for differentiating the two 
species, but there is some slight overlap, 
with E. subtilis varying from 2 to 5 (usually 
3) spines whereas E. lineatus varies from 
to 2 (usually 1). The type series for E. li- 
neatus consists of a mix of both of these 
species, but since the specimen designated 
as the holotype has multiple antennular 
spines and a spine on the stylocerite, this is 
the form that should retain the name E. li- 
neatus. 

Much of the confusion regarding these 
species is due to variability in the length of 
the rostrum of E. subtilis (Fig. 4c-f). Most 
keys continue to follow the pattern estab- 
lished by Rathbun (1904) of separating E. 
pusiolus and "£. herdmanV (= E. lineatus) 
solely on the basis of whether the rostrum 
overreaches the second article of the anten- 
nular peduncle. We found many cases 
where, depending on the length of the ros- 
trum, specimens of E. subtilis from the 
same haul had been cataloged as E. pusi- 
olus and E. herdmani or E. lineatus. 

The presence of multiple antennular 
spines makes E. lineatus very easy to dif- 
ferentiate from E. subtilis and E. pusiolus. 
The latter two species can be reliably sep- 
arated by the number of meral spines on the 
pereopods: E. subtilis has 2 to 5 strong, dis- 
tal spines on the merus of the third pereo- 
pod (and normally multiple spines on the 
fourth), whereas E. pusiolus has only a sin- 
gle spine on each of these pereopods. Eu- 
alus pusiolus has a small, rounded subor- 
bital carapace spine, while E. subtilis has 
none. In this respect E. subtilis resembles 
the South American species E. dozei (A. 
Milne Edwards, 1891), but this species also 
has only single meral spines (Holthuis 
1952). 

We found that some of the characters 
given by Carvacho & Olson (1984) to dif- 
ferentiate E. subtilis were not useful. The 
ventral spines on the abdomen (considered 
by these authors to be unique to this spe- 
cies) were present on all males and most 
non-ovigerous females of E. pusiolus and 



VOLUME 112, NUMBER 1 



137 




Fig. 3. Eualus lineatus Wicksten & Butler, 1983. Lateral view of anterior region of carapace, showing curved 
tooth on the base of the stylocerite and multiple spines on the basal article of the antennular peduncle. Scale 
bar is 1 mm. 



E. lineatus. Furthermore, the basicerite of 
E. pusiolus has only one (not two) lateral 
spines, so this character is not useful for 
separating this species from E. subtilis. 

Eualus subtilis is quite possibly the most 
abundant shrimp in Puget Sound, occurring 
subtidally on virtually any bottom type 
from mud to solid rock. It sometimes oc- 
curs in the low intertidal and has been col- 
lected in trawls to at least 74 m, and to date 
has been found from Barkley Sound, Brit- 
ish Columbia (this study) to Bahia de Todos 
Santos, Punta Banda, Baja California (Car- 
vacho & Olson 1984). Males of E. subtilis 
are fairly uncommon in collections, proba- 



bly due to their very diminutive size. Al- 
though males were always much smaller 
than ovigerous females, the presence of 
small females suggests that the species is 
not strictly protandric. The unusual modi- 
fications to the pereopods bear some resem- 
blance to those described by Bauer (1986) 
for another small hippolytid, Thor manningi 
Chace, 1972, a species that exhibits a novel 
reproductive strategy involving nearly 
equal proportions of protandric individuals 
and primary males. 

Less is known about the habits and hab- 
itat of the much rarer E. lineatus. Speci- 
mens have been collected at depths of 12- 



138 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



0.5 mm 

a 
0.2 mm 

b 
1.0 mm 

c d ef 




a 




Fig. 4. Eualus subtilis Carvacho & Olson, 1984. a. Third pereopod of male, lateral view; b, dactyl of third 
pereopod of male, lateral view; c, lateral view of male carapace; d, lateral view of female carapace; e, rostrum 
variation, female, lateral view; f, rostrum variation, female, lateral view. 



VOLUME 112, NUMBER 1 



139 




Fig. 5. Eualus lineatus Wicksten & Butler, 1984. Dorsal view of live specimen (male, 3.5 mm postorbital 
carapace length) showing pattern of broad orange bands. 



120 m from Juneau, Alaska to at least Santa 
Cruz Island, California. This species may 
associate with sponges since the holotype 
was collected from "mud and sponge bot- 
tom," and the trawl collection records for 
the specimens examined often indicated 
that the shrimp were removed from sponge 
cavities, or noted the presence of sponges 
in the haul. The live specimen collected by 
one of us (GCJ) was found in a suction 
sampler that had been used in both crevices 
and small sponges on a vertical rock face, 
but it is not known at which point in the 
dive it was captured. 

Given the confusion that has surrounded 
these species, any records should be con- 
sidered suspect until the specimens have 
been reexamined. It is likely that at least 
some of the depth and range records will 
be revised, and perhaps new information on 
the habitat of E. lineatus will come to light. 



Acknowledgments 

We are grateful for the help and support 
of the staff of the Shannon Point Marine 
Center, where part of this study was sup- 
ported by the National Science Foundation 
Research Experiences for Undergraduates 
program, and J. Orensanz for the loan of 
his camera lucida. The Royal Norwegian 
Society of Sciences, Trondheim, provided 
specimens of E. pusiolus; K. Sendall, K. 
Reid, R. Van Syoc, and J. W. Martin gra- 
ciously provided specimens from their re- 
spective museum collections, J. Price of the 
Canadian Museum of Nature went to great 
lengths to try to find the missing paratypes, 
and P. Jensen provided invaluable assis- 
tance in field sampling. 

Literature Cited 

Bauer, R. T. 1986. Sex change and life history pattern 
in the shrimp Thor manningi (Decapoda: Cari- 



140 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



dea): a novel case of partial protandric her- 
maphroditism. — Biological Bulletin 170:11-31. 

Butler, T. H. 1980. Shrimps of the Pacific coast of Can- 
ada. — Canadian Bulletin of Fisheries and 
Aquatic Sciences 202:1-280. 

Carvacho, A., & Y. R. Olson. 1984. Nuevos registros 
para la fauna carcinologica del noreste de Mex- 
ico y descripcion de una nueva especie: Eualus 
subtilis, n.sp. (Crustacea: Decapoda: Natan- 
tia). — The Southwestern Naturalist 29(1):59- 
71. 

Chace, F. A., Jr. 1972. The shrimps of the Smithsonian- 
Bredin Caribbean expeditions with a summary 
of the West Indian shallow-water species (Crus- 
tacea: Decapoda: Natantia). — Smithsonian Con- 
tributions to Zoology 98:1-179. 

Greve, L. 1963. The genera Spirontocaris, Lebbeus, 
Eualus and Thoralus in Norwegian waters 
(Crustacea, Decapoda). — Sarsia 11:29-42. 

Holmes, S. J. 1900. Synopsis of California stalk-eyed 
Crustacea — Occasional Papers of the California 
Academy of Sciences 7:1-262. 

Holthuis, L. B. 1952. The Crustacea Decapoda Macru- 
ra of Chile. — Reports of the University of Chile 
Expedition 1948-49, 5:1-110. 

Kr0yer, H. 1841. Udsigt over de nordiske Arter af 
Slaegten Hippolyte. — Naturhistorisk Tidsskrift 
3:570-579. 

Milne Edwards, A. 1891. Crustaces. Mission scienti- 



fique du Cap Horn 1882-1883, volume 6, Zo- 
ology, part 2F:l-54. 

Rathbun, M. J. 1904. Decapod crustaceans of the 
northwest coast of North America. — Harriman 
Alaska Expedition Series 10:1-210. 

Squires, H. J. 1990. Decapod Crustacea of the Atlantic 
coast of Canada. — Canadian Bulletin of Fish- 
eries and Aquatic Sciences 221:1-532. 

Thallwitz, J. 1891. Decapoden-Studien, ibessondere 
basirt auf A. B. Meyer's Sammlungen im Ostin- 
dischen Archipel, nebst einer Aufzahlung der 
Decapoden und Stomatopoden des Dresdener 
Museums. Abhandlungen und Berichte des 
Koniglichen Zoologischen und Anthropolo- 
gisch-Ethnographischen Museums zu Dresden 
1890-91(3): 1-55. 

Walker, A. O. 1898. Crustacea collected by W. A. 
Herdman in Puget Sound, Pacific Coast of 
North America, Sept. 1897. — Transactions of 
the Liverpool Biological Society 12:268-287. 

Wicksten, M. K. 1988. New records and range exten- 
sions of shrimps and crabs from California, 
U.S.A. and Baja California, Mexico. — Califor- 
nia Fish and Game 74(4): 23 6-248. 

, & T. H. Butler. 1983. Description of Eualus 

lineatus new species, with a redescription of 
Heptacarpus herdmani (Walker) (Caridea: Hip- 
polytidae). Proceedings of the Biological Soci- 
ety of Washington 96:1-6. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):141-144. 1999. 

Deilocerus captabilis, a new species of cyclodorippid crab from 
southeastern Brazil (Crustacea: Decapoda: Brachyura: 

Cyclodorippidae) 

Marcos Tavares 
Universidade Santa Ursula, ICBA, Rio de Janeiro 22231-040, Brasil 

Abstract. — Deilocerus captabilis, new species, the third western Atlantic 
species and seventh representative of the genus is herein described based on 
an adult female collected on calcareous algae nodules from 50 m depth, south- 
eastern Brazil. Deilocerus captabilis, new species, is the only species in the 
genus with three teeth on the lateral margin of the carapace. 

Re sumo. — Deilocerus captabilis nova especie, terceira a ocorrer no Atlantico 
sul ocidental das sete conhecidas para o genero, e descrita e ilustratada. A nova 
especie e baseada em uma femea adulta coligida a 50 m de profundidade em 
fundos de nodulos de algas calcareas. A nova especie se distingue facilmente 
das demais especies do genero por ser a unica a apresentar tres dentes na 
margem lateral da carapa^a. 



As part of an ongoing project (REVI- 
ZEE) of the Brazilian government's depart- 
ment "Ministerio do Meio Ambiente" to 
evaluate the country's marine living re- 
sources, a biological survey of the conti- 
nental shelf and slope from Salvador (12°S) 
to Cabo de Sao Tome (23°S) was conducted 
in 1997 onboard the "Astro Garoupa". 

The samples yielded an adult female of 
a new species of cyclodorippid crab, Deil- 
ocerus captabilis, described herein. 

The holotype is deposited in the Museu 
Nacional, Rio de Janeiro (MNRJ). Descrip- 
tive terminology follows Tavares (1991, 
1996). Abbreviations: Mxpl-3, first to third 
maxillipeds; P2-P5, second to fifth pereo- 
pods, PI, cheliped; cl, carapace length; cw, 
carapace width; mm, millimeters. 

Deilocerus captabilis, new species 
Fig. 1 

Material examined. — Brazil: Espirito 
Santo. "Astro Garoupa", REVIZEE Cen- 
tral II, St. 34C, 3 Nov 1997, 20°24'S, 
39049' W, 50 m: female holotype cl 1.7 mm, 
cw 2.0 mm (MNRJ 7303). 



Type locality. — Brazil: Espirito Santo 
(20°24'S, 39°49'W, 50 m). 

Description. — Carapace slightly broader 
than long. Dorsal surface ornamented with 
very fine scattered granules, denser near 
margins, except on smooth, shallow 
grooves defining gastric regions. Ventrolat- 
eral surfaces of carapace almost smooth; 
subhepatic region densely covered with 
rounded coarse granules, coarser on dorsal 
surface. Frontolateral tooth rounded, dense- 
ly covered with small granules; exorbital 
tooth blunt. Hepatic and anterolateral teeth 
short, blunt, covered with rounded granules; 
anterolateral tooth smallest. Laterobranchial 
tooth present as low lobe. Anterolateral 
margin (from exorbital tooth to branchial 
tooth) rounded, about as long as posterolat- 
eral margin (measured from branchial tooth 
to posterior margin). Posterolateral margin 
straight, well defined by row of small gran- 
ules. 

Ocular peduncle covered with small 
rounded tubercles, anterodistal tubercles 
more acute; cornea pigmented. Antenna 
very small, hidden in dorsal view; articles 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Deilocerus captabilis, new species, Espirito Santo, Brazil, 20°24'S, 39°49'W, 50 m: female holotype 
cl 1.7, cw 2.0 mm (MNRJ 7303). A, dorsal view of whole crab. B, right cheliped outer view. 



1-4 ornamented with small granules; sec- 
ond article slightly flattened; fifth article al- 
most smooth; flagellum obsolete. 

Endostomial channel visible dorsally be- 
tween frontolateral teeth. Third maxilliped 
with ischium and merus, each more than 2 



times as long as broad, with outer surfaces 
covered with rounded granules; palp with- 
out granules, articulated on inner surface of 
merus. 

Chelipeds densely covered with small tu- 
bercles, much less dense and smaller on in- 



VOLUME 112, NUMBER 1 



143 



ner surfaces; inner surfaces of merus, car- 
pus and chela forming concave surface fit- 
ting closely against walls of carapace; dor- 
sal and ventral margins of merus, carpus, 
palm and fingers well defined. Fingers ter- 
minating in sharp tips, cutting edge with 
few small acute teeth. Dactyl smaller than 
palm, set obliquely relative to palm axis. 
Fixed finger about 2 times broader proxi- 
mally than distally. Palm about 2 times lon- 
ger than broad, ornamented longitudinally 
with 2 rounded protuberances, proximal 
one largest. 

P2 longer than P3, otherwise similar. P2 
and P3 laterally flattened; both legs with 
propodus, carpus, and merus densely orna- 
mented with tubercles on dorsal and ventral 
margins, flanks almost smooth; dactyl cy- 
lindrical, with minute granules. 

P4 and P5 generally similar, subdorsal, 
subcheliform, much smaller than P2 and 
P3; P4 with ischium about 3 times longer 
than in P5; dactylus and propodus short, 
strongly curved; propodus twisted, dactylus 
flexing on its lateral surface. Ornamentation 
on propodi, carpi and meri less pronounced 
on P4-5 than on P2-3. 

Female abdomen with 6 segments dense- 
ly covered with small tubercles diminishing 
in size and density from pleotelson to first 
segment. Pleotelson as wide as fifth seg- 
ment, lateral margins broadly rounded. 

Distribution. — Known only from the 
type-locality Espirito Santo, Brazil 
(20°24'S, 39°49'W, 50 m). 

Etymology. — The specific name, capta- 
bilis (Latin, that can take), refers to the sub- 
cheliform P4 and P5. 

Remarks. — The genus Deilocerus Tava- 
res, 1993, is strictly American in distribu- 
tion. The genus is represented in the west- 
em Atlantic by two species, and in the east- 
em Pacific by four (Tavares 1993, 1996). 
Two groups of species are recognizable. 
The first group includes species with only 
one anterolateral tooth on the margin of the 
carapace: D. perpusillus (Rathbun, 1900) 
and D. analogus (Coelho, 1973), from the 
western Atlantic, and D. laminatus (Rath- 



bun, 1935) from the eastern Pacific. The 
second group is found only in the eastern 
Pacific and encompasses species with two 
teeth (one hepatic and one anterolateral), 
along the margin of the carapace: D. planus 
(Rathbun, 1900), D. decorus (Rathbun, 
1933) and D. hendrickxi Tavares, 1993. 
Deilocerus captabilis, new species, falls 
into the second group and is, therefore, the 
first western Atlantic representative of that 
group. Deilocerus captabilis, new species 
is, so far, the only species in the genus with 
three teeth on the lateral margin of the car- 
apace (hepatic, anterolateral, and latero- 
branchial): the first two teeth being well de- 
veloped, and the laterobranchial represented 
by a low lobe. Deilocerus captabilis, new 
species, can also be readily recognized from 
the remaining six species of the genus by 
the shape and ornamentation of the fronto- 
orbital margin of the carapace. 

Although no carrying behavior (Guinot 
et al. 1995) has been observed in Deiloce- 
rus captabilis, new species, it is possible 
that the crab can hold an object over its 
carapace using the last two pairs of legs. P4 
and P5 are subchelate, with the propodus 
and especially the dactyl strongly curved; 
the propodus is twisted so the dactyl closes 
on the lateral surface of the propodus. This 
structure allows a small object, such as a 
piece of shell, to be held over the carapace. 
Carrying behavior has been observed in 
other species of the genus, such as D. lam- 
inatus (see Garth 1946, Tavares 1994) and 
D. planus (see Schmitt 1921, Wicksten 
1982). 

Acknowledgments 

I am grateful to Daniele Guinot (Museum 
national d'Histoire naturelle, Paris), Rafael 
Lemaitre (Smithsonian Institution, Wash- 
ington, D.C.) and Peter K. L. Ng (National 
University of Singapore) for reading a draft 
of the manuscript and providing helpful 
suggestions, and to Maria Helena Pinheiro 
(Universidade Santa Ursula) for preparing 
drawings. This research has been funded by 



144 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Ministerio do Meio Ambiente, SECIRM, 
Programa REVIZEE, and Universidade 
Santa Ursula, Rio de Janeiro. Thanks are 
due also to CNPq for supporting studies on 
the systematic of decapod crustaceans in the 
form of ongoing grant 30.09.15/97-7. 

Literature Cited 

Coelho, P. A. 1973. Descri9ao preliminar de Clythro- 
cerus analogus, n. sp., do literal brasileiro 
(Crustacea, Decapoda, Dorippidae). — Ciencia e 
Cultura 25(6)(suppl.):343-344. 

Garth, J. S. 1946. Littoral brachyuran fauna of the Gal- 
apagos Archipelago. — Allan Hancock Pacific 
Expedition 5(10):i-iv + 341-600. 

Guinot, D., D. Doumenc, & C. C. Chintiroglou. 1995. 
A review of the carrying behaviour in Brachy- 
uran crabs, with additional information on the 
symbioses with sea anemones. — Raffles Bulle- 
tin of Zoology 43(2):377-416. 

Rathbun. M. J. 1900. Synopses of North American in- 
vertebrates. X. The Oxyrhynchous and Oxysto- 
matous crabs of North America. — American 
Naturalist 34:503-520. 

. 1933. Preliminary descriptions of nine new 

species of oxystomatous and allied crabs. — Pro- 
ceedings of the Biological Society of Washing- 
ton 46:183-186. 

. 1935. Preliminary descriptions of seven new 

species of oxystomatous and allied crabs. — Pro- 



ceedings of the Biological Society of Washing- 
ton 48:1-4. 

Schmitt, W L. 1921. The Marine Decapod Crustacea 
of California with special reference to the Deca- 
pod Crustacea collected by the United States 
bureau of Fisheries Steamer "Albatross" in 
connection with the Biological Survey of San 
Francisco Bay during the years 1912-1913. — 
University of California Publications in Zoolo- 
gy 23:1-470. 

Tavares, M. 1991. Revision preliminaire du genre Ty- 
molus Stimpson, avec la description de Tymolus 
brucei sp. nov. d'Australie occidentale (Crus- 
tacea, Brachyura, Cyclodorippoidea). — Bulletin 
du Museum national d'Histoire naturelle, Paris 
(4), 13, sect. A, (3-4):439-456. 

. 1993. Crustacea Decapoda: Les Cyclodorip- 

pidae et Cymonomidae de I'lndo-Ouest-Paci- 
fique a 1' exclusion du genre Cymonomus. In A. 
Crosnier, ed., Resultats des Campagnes MU- 
SORSTOM, 10. — Memoires du Museum na- 
tional d'Histoire naturelle, Paris 156:253-313, 
Fig. 1-20. 

. 1994. Description preliminaire de quatre nou- 

veaux genres et trois nouvelles especes de Cy- 
clodorippoidea americains. — Vie Milieu 43(2- 
3):137-144. 

. 1996. Revision systematique des Cyclodor- 



ippidae americains (Crustacea, Decapoda, 
Brachyura). — Bulletin du Museum national 
d'Histoire naturelle, Paris (4), 18, sect. A, (1- 
2):233-295. 
Wicksten, M. K. 1982. Behavior in Clythrocerus plan- 
us (Rathbun, 1900) (Brachyura, Dorippidae). — 
Crustaceana 43(3):306-307. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):145-163. 1999. 

The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian 
Islands, with description of a new species 

Christopher B. Boyko 

Department of Invertebrates, American Museum of Natural History, Central Park West at 79th 

St., New York, New York 10024, U.S.A. and Department of Biological Sciences, University of 

Rhode Island, Kingston, Rhode Island 02881, U.S.A. 

Abstract. — Albunea danai, a new species that has been confused with A. 
speciosa Dana, is described from Oahu, Hawaii. The new species actually 
resembles A. carabus (Linnaeus) from the Mediterranean and western Africa. 
The discovery of this new species brings the total number of Indo-West Pacific 
members of the genus to eight. Albunea speciosa is broadly distributed in the 
Indo-West Pacific and is not a Hawaiian endemic as previously believed. As 
the identities of the two Hawaiian species have been confused, A. speciosa is 
here redescribed from new material and a neotype is selected. Both A. danai 
and A. speciosa are discussed using newly defined morphological characters. 



Albunea speciosa Dana, 1852, is the only 
species of albuneid heretofore known from 
the Hawaiian Islands and has been consid- 
ered a Hawaiian endemic (Serene 1973). In 
the course of examining specimens of sand 
crabs for a worldwide revision of the family 
Albuneidae, material from Hawaii was 
found labeled ''Albunea speciosa'' and 
''Albunea symnista'' [sic] in the collections 
of the Western Australian Museum (WAM), 
but that was clearly not conspecific with A. 
speciosa. Later, additional Hawaiian speci- 
mens were obtained from the Bemice P. 
Bishop Museum that were labeled "Albu- 
nea thurstonV that clearly are not A. thur- 
stoni Henderson, 1893, but instead are the 
same taxon as the WAM material. Since 
these specimens cannot be placed in any 
known species of albuneid, they are de- 
scribed here as a new species. In addition, 
my examination of specimens that are re- 
ferable to A. speciosa revealed that this spe- 
cies is not endemic to Hawaii, but has a 
broad range in the Indo-West Pacific. Be- 
cause the faunal composition and biogeog- 
raphy of island groups has attracted much 
attention of late and the fact that more than 
one species has been repeatedly identified 



with A. speciosa, that species is redescribed 
from new material and a neotype is desig- 
nated herein in order to fix its identity. This 
description of the new species brings the 
total number of Indo-West Pacific species 
of Albunea to eight: A. symmysta (Linnae- 
us, 1758), A. speciosa, A. microps Miers, 
1878, A. thurstoni, A. elioti Benedict, 1904, 
A. steinitzi Holthuis, 1958, A. madagascar- 
iensis Thomassin, 1973, and A. danai new 
species. 

Materials, Methods, and Morphological 
Terminology 

Materials. — Specimens for this study 
came from the collections of the American 
Museum of Natural History, New York 
(AMNH), Bemice P. Bishop Museum, Hon- 
olulu, Hawaii (BPBM), California Acade- 
my of Sciences, Invertebrate Zoology, San 
Francisco (CASIZ), Musee Royal de 
I'Afrique Centrale, Tervuren, Belgium 
(MRAC), Queensland Museum (QM), Uni- 
versity Museum of Zoology, Cambridge 
(UMZC), National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. (USNM), Western Austrahan Muse- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



um, Perth (WAM), and Yale Peabody Mu- 
seum, New Haven, Connecticut (YPM). 

Methods. — Carapace length (CL), as 
measured from the midpoint of the anterior 
margin (including rostrum) to the midpoint 
of the posterior concavity, is provided as an 
indicator of specimen size. In the Hst of 
synonyms, asterisks refer to publications 
citing material examined during the present 
study. Absence of an asterisk in a specific 
entry does not imply that the identifications 
therein are in doubt, but only that it was not 
possible to examine the material cited in 
that publication. 

Illustrations were created using a modi- 
fied approach of Harvey & De Santo 
(1997). Specimen images were first cap- 
tured on a Macintosh® computer with a 
digital camera connected to a Wild M8 dis- 
secting microscope. These images were 
then prepared using the programs Adobe 
Photoshop® and Adobe Illustrator®. I at- 
tempted to record the position and size of 
setae in these drawings as accurately as 
possible, although for clarity of presenta- 
tion the plumules of plumose setae were not 
drawn. 

Morphological terminology. — During the 
course of this study and that of Boyko & 
Harvey (in press), several important diag- 
nostic morphological features were encoun- 
tered that have not been described previ- 
ously for albuneids. Although these, along 
with other features, are discussed in Boyko 
& Harvey (in press), they are also consid- 
ered here to faciUtate a clearer understand- 
ing of the descriptive terms used in this pa- 
per. 

The front of the carapace of albuneids 
bears a broad mat of very short, dense, sim- 
ple setae. This mat, hereafter called the se- 
tal field, varies in shape and extent across 
genera and species, but appears to be rela- 
tively invariant within species. The cara- 
pace also possesses numerous transverse, 
setose grooves. Although carapace grooves 
(CG) have been scarcely mentioned by pre- 
vious authors, 1 1 major grooves (numbered 
1-11, Fig. 1) have been identified which 



Setal field 




Fig. \. Diagrammatic albuneid carapace based on 
Albunea microps Miers, 1878, showing setal field and 
setose carapace grooves (CG 1-11) discussed in text. 



can be recognized across albuneid genera. 
Variability in the presence and the degree 
of fragmentation of specific grooves, in the 
anterior-posterior displacement of individ- 
ual fragments, and in the texture of the 
grooves (e.g., smooth, crenulate) tends to 
be conservative within species, and thus 
carapace grooves are useful in recognizing 
species. 

Because several of the specimens exam- 
ined during this study were incorrectly 
sexed, brief remarks on the determination 
of male and female identity in albuneids are 
presented here. As in most decapod crus- 
taceans, albuneid females have gonopores 
on the coxae of the third pereopods, where- 
as males have gonopores on the coxae of 
the fifth pereopods. However, in some al- 
buneid genera (e.g., Lepidopa Stimpson, 
1858 and some Albunea, including both 
species from Hawaii), males also have a 
small pore on the coxa of the third pereo- 
pod in a position analogous to that of the 
female gonopore. The precise nature and 
function of this pore is unknown. 

In albuneids, females have well devel- 
oped uniramous pleopods on abdominal so- 



VOLUME 112, NUMBER 1 



147 




Fig. 2. Pereopod II dactyl of Albunea sp., lateral 
view, showing terms used in species accounts for land- 
marks on pereopod dactyli. 



mites II- V. Male albuneids have tradition- 
ally been considered to lack pleopods (Ef- 
ford & Haig, 1968). However, rudimentary 
or small pleopods were found on abdominal 
somites II-V of male specimens with well 
developed gonopores on the fifth pereopods 
in species of several albuneid genera (e.g., 
Lepidopd). In some Albunea species (e.g., 
A. microps, A. speciosa), specimens with 
large pores on the fifth pereopods and with 
small pores on the coxae of the third pereo- 
pods, show no signs of pleopod develop- 
ment. In those species in which the male 
pore occurs, it is always smaller than gon- 
opores of same-sized females; likewise, the 
pleopods of females are always much more 
developed than those of males. Males are 
most reliably recognized by the presence of 
a gonopore on the fifth pereopod and the 
rudimentary degree of development of the 
pleopods or lack thereof. In small speci- 
mens, however, the presence or absence of 
the male gonopore is a more reliable indi- 
cator of sex than is pleopod development 
because both males and females may have 
small pleopod buds as juveniles. 

The shape of the dactylus of the pereo- 
pods, particularly the third pereopod, has 
been used to distinguish among species of 
albuneids. To facilitate the description of 
the complex shape of this segment, several 
terms are used to refer to important land- 
marks (Fig. 2). The "base" of the dactylus 
is the ventroproximal angle; the "heel" cor- 
responds to the dorsoproximal angle, which 



is often strongly produced. The dorsal mar- 
gin is almost always concave, sometimes 
smoothly so; in most species, however, the 
dorsal margin has a distinct angle, the apex 
of which is referred to as an "indent." The 
dactylus terminates in a "tip," which is 
somewhat rounded and lacking in a corne- 
ous nail. 

In some species of albuneids, certain seg- 
ments of the pereopods bear a large trans- 
parent, decalcified area, hereafter called the 
"window," that has not been previously re- 
ported in this family. This area, when pre- 
sent, is most prominent on the lateral sur- 
face of the merus, where it is comparable 
to the "leg membranes" of porcelain crabs 
(Porcellanidae) discussed in detail by Still- 
man & Somero (1996). These windows can 
also be found to a lesser degree on other 
pereopod segments, both laterally and me- 
sially. 

Superfamily Hippoidea Latreille, 1825 

Family Albuneidae Stimpson, 1858 

Albunea Weber, 1795 

Albunea speciosa Dana, 1852 

Figs. 3, 4 

Albunaea speciosa Dana, 1852: 405-406; 
1855: pi. 25, figs. 6a-f. — Stimpson, 
1858: 230 (list). 

Albunea speciosa. — Miers, 1878: 315 (list), 
331 (after Dana, 1852).— Ortmann, 1896: 
223 (key), 225 (hst), 239 (table).— Gor- 
don, 1938: 187 (list).— Edmondson, 
1946: 266.— Serene, 1973: 262-263, pi. 2. 

''1 Albunea speciosa''. — Borradaile, 1904: 
751*. 

Material examined. — Neotype: Hawaii, 
Oahu, 27 May 1938: 1 male, 9.0 mm CL 
(USNM 260868); same data as neotype: 2 
males, 9.5-10.4 mm CL, 1 female, 14 mm 
CL (USNM 287087); Kailua, Oahu, Mar 
1938: 1 male, 10.1 mm CL (BPBM 
SI 1781); Honolulu Harbor, Oahu, coll. E. 
M. Ehrhon, Dec 1916: 2 males, 6.4-9.9 mm 
CL, 1 female, 7.3 mm CL (CASIZ 109240); 
Halonu Blow Hole dive site, south shore, 
Oahu, coll. R. Holcom, 3 Aug 1997: 1 fe- 



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Fig. 3. Albunea speciosa Dana, 1852. A, male, 9.0 mm CL, USNM 260868, neotype; B-J, male, 10.4 mm 
CL, USNM 287087. A, carapace and eyes, dorsal view; B, eyes, dorsal view; C, left antennule, lateral view; D, 
left antenna, lateral view; E, left mandible; mesial view; F, right maxilla, lateral view; G, left maxillule, lateral 
view; H, left maxilliped I, lateral view; I, right maxilliped II, lateral view; J, left maxilliped III, lateral view. 
Scale = 1.6 mm (B, E, G), 2.2 mm (I), and 3.3 mm (A, C, D, F, H, J). 



VOLUME 112, NUMBER 1 



149 




Fig. 4. Albunea speciosa Dana, 1852. A-E, male, 10.4 mm CL, USNM 287087; F, male, 9.0 mm CL, USNM 
260868, neotype; G, female, 14 mm CL, USNM 287087. A, right pereopod I, lateral view; B, right pereopod 
II, lateral view; C, right pereopod III, lateral view; D, right pereopod IV, lateral view; E, abdominal somites I- 
VI, dorsal view; F, telson of male, dorsal view; G, telson of female, dorsal view. Scale = 3.0 mm (F, G), 4.0 
mm (A-D), and 4.2 mm (E). 



male, 6.6 mm CL, 1 broken unsexable/un- 
measurable specimen (QM W22284); Hal- 
onu Blow Hole dive site, south shore, 
Oahu, 12.2-13.7 m, coll. R. Holcom, 4 Apr 
1997: 2 males, 6.4-7.1 mm CL, 3 females, 
5.7-9.5 mm CL, 6 juveniles, 3.3-4.1 mm 
CL (QM W22285); Oahu, coll. R. Holcom, 
Apr 1997: 2 ovigerous females, 7.5 mm CL 
(QM W22286); "Hawaii," coll. C. M. 
Cook Jr., 1897: 1 female, 10 mm CL (YPM 
21133). 

Australia: Bay on north side of Point 



Cloates, lee side of reef. Western Australia, 
113°38'E, 22°4rS, depth 3.7 m, coll. WAM 
Ningaloo Expedition, 23 Aug 1968: 1 male, 
8.7 mm CL, 1 carapace, 9.1 mm CL (WAM 
489-97); southwest of Point Cloates, West- 
ern Australia, 113°39'30"E, 22°43'30"S, 
coll. WAM Ningaloo Expedition, 7 Sep 
1968: 1 ovigerous female, 9.7 mm CL 
(WAM 490-97). 

Seychelles: Mahe, coll. Mission Zoolo- 
gique MRAC-ULB, July-Sep 1966: 1 ovig- 
erous female, 9.9 nrni CL (MRAC 53.894). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Maldives: Hulule, Male Atoll, coll. J. S. 
Gardiner: 1 male, 8.4 mm CL (UMZC). 

Type locality. — Sandwich Islands (= Ha- 
waiian Islands) (Dana, 1852) herein re- 
stricted by neotype selection to Oahu, Ha- 
waii, USA, Pacific Ocean. 

Type material. — It is unclear exactly how 
many specimens of this species Dana 
(1852) had before him when writing the de- 
scription of this species, but the description 
suggests that there was only one. No type 
material of this species is extant in either 
USNM, the Museum of Comparative Zo- 
ology, Harvard University, or the British 
Museum (Natural History) (Evans 1967; 
Boyko, pers. obs.). Because these institu- 
tions are the only known repositories for 
Dana's extant type material (Evans 1967), 
the type material of A. speciosa must be 
considered lost. In light of the new infor- 
mation about the range of A. speciosa, and 
the discovery of a new species of albuneid 
in the Hawaiian Islands which has been re- 
peatedly misidentified as A. speciosa, it is 
appropriate to follow the suggestion of Se- 
rene (1973) and select a neotype for the 
species. A male, 9.0 mm CL (USNM 
260868) is herein designated as the neotype 
for Albunea speciosa. 

Diagnosis. — Carapace slightly longer 
than wide, covered with strongly setose 
grooves. Anterior margin with 13-17 teeth 
on either side of ocular sinus. Setal field 
with narrow lateral elements and concave 
anterior margin; posterior lateral elements 
extending to posterior lateral elements of 
CGI. CGI with separate posterior lateral el- 
ements but with anterior and posterior ele- 
ments united by posterior elements of setal 
field; CG4 with 2-3 short anteriorly dis- 
placed medial elements; CG5 entire, nearly 
reaching margins of CG6; CG6 and CG7 
separate; CG8 with 1-2 median elements 
separated from lateral elements; CGI 1 pres- 
ent. Rostrum reaching just beyond proximal 
margin of ocular plate. Ocular plate sub- 
quadrate. Ocular peduncles dorsoventrally 
flattened and elongate, rounded at tip, ap- 
proximate along miesial margin; lateral mar- 



gin concave; mesial margin straight. Cornea 
at tip of ocular peduncle. Antennule with 
48-53 flagellar exopod and 2 endopodal 
segments. Antenna with 5-6 flagellar seg- 
ments; acute spine on dorsolateral surface 
of peduncle segment I. Dactyli of pereo- 
pods II, III with heels low and smoothly 
rounded. Coxa of pereopod III of males 
with small male pore. Telson of male spat- 
ulate, laterally expanded, dorsoventrally 
flattened; produced slightly at tip. Telson of 
female flattened, rounded at tip. 

Redescription. — Carapace (Fig. 3a) 
slightly wider than long. Anterior margin 
concave on either side of ocular sinus, be- 
coming convex laterally with 10-12 large 
and 3-4 small spines along length. Rostrum 
a small acute tooth, reaching just beyond 
proximal margin of ocular plate. Ocular si- 
nus smoothly concave, unarmed. Frontal re- 
gion smooth; setal field broad posteriorly, 
narrowing anteriorly, with narrow anterior 
lateral elements and concave anterior mar- 
gin; posterior lateral elements reaching to 
posterior lateral elements of CGI. Medial 
portion of CGI parallel to anterior margin 
of carapace, sinuous, slightly crenulate, di- 
vided into medial fragment and curved pos- 
teriorly-displaced lateral elements, but with 
medial and lateral elements connected by 
posterior lateral elements of setal field. Me- 
sogastric region smooth; CG2 short, with 
1-2 elements; CG3 broken into 2 longer 
lateral elements and 1-3 short medial ele- 
ments; CG4 with 2-3 short medial elements 
displaced anteriorly, with gap at midline be- 
tween short elements. Hepatic region 
smooth with long setose groove at median 
of lateral margin. Epibranchial region ap- 
proximately triangular, smooth; posterolat- 
eral margin with 3 short rows of setae. Me- 
tagastric region smooth; CG5 ranging from 
entire to 4 elements, nearly reaching mar- 
gins of CG6. CG6 strongly crenulate, 
strongly anteriorly concave medially and 
sloping out to anteriorly convex lateral 
thirds. CG7 nearly straight relative to an- 
terior margin of carapace and separate from 
CG6. Cardiac region smooth; CG8 with 1- 



VOLUME 112, NUMBER 1 



151 



2 median elements separated from lateral 
elements. CG9 present as 2 lateral grooves 
with short gap at midline. CGIO present as 
2 curved lateral fragments, with gap be- 
tween fragments about half length of single 
fragment. CGll present. Branchial region 
with numerous short, transverse rows of se- 
tae. Posterior margin deeply and evenly 
convex, with submarginal groove reaching 
about half-way either side of posterior con- 
cavity. Branchiostegite with short anterior 
submarginal spine; anterior region with 
scattered short transverse lines ventral to 
linea anomurica; with many short rows of 
setae and sparsely covered with long plu- 
mose setae ventrally; posterior region mem- 
branous, with numerous irregular frag- 
ments, and sparsely covered with long plu- 
mose setae. 

Ocular plate (Fig. 3b) subquadrate, with 
broad median indentation; proximal ocular 
segments (Fig. 3b) reduced to small round- 
ed calcified area on either side of ocular 
plate. Ocular peduncles (Fig. 3b) elongate, 
with medially concave lateral margins, ta- 
pering to rounded distal comeae; mesial 
margins approximated along entire length; 
mesial and ventral margins of peduncle 
with sparse row of long plumose setae; tuft 
of plumose setae at proximal lateral ventral 
angle. 

Antennule (Fig. 3c) segment III narrow 
proximally, expanding distally to twice 
proximal width; with plumose setae on dor- 
sal and ventral margins; dorsal exopodal 
flagellum with 48-53 segments and long 
plumose setae on dorsal and ventral mar- 
gins; ventral endopodal flagellum short, 
with 2 segments and plumose setae on dor- 
sal and ventral margins. Segment II medi- 
ally inflated in dorsal view, with plumose 
setae on dorsal and ventral margins, and se- 
tae scattered on ventrolateral third of sur- 
face. Segment I wider than long, unarmed; 
dorsal third of lateral surface faintly rugose 
with long plumose setae; long plumose se- 
tae on dorsal and ventral margins. 

Antenna (Fig. 3d) segment V about 3 
times longer than wide, with long plumose 



setae on dorsal and ventral margins; flagel- 
lum 5-6 segmented, with long plumose se- 
tae on dorsal, ventral and distal margins. 
Segment IV expanded distally, with long 
plumose setae on dorsal, ventral and distal 
margins, and row of setae on dorsolateral 
margin. Segment III with long plumose se- 
tae on ventral margin. Segment II short, 
widening distally, with plumose setae on 
margins; antennal acicle long, thin, reach- 
ing to distal margin of segment IV, with 
long plumose setae on dorsal margin. Seg- 
ment I rounded proximally, flattened ven- 
trolaterally with long plumose setae on 
margins; lateral surface with acute spine 
dorsally, with low semicircular dorsolateral 
lobe ventrodistal to spine; segment with 
ventromesial antennal gland pore. 

Mandible (Fig. 3e) incisor process with 
1 tooth; cutting edge with 1 tooth. Molar 
process with 1-2 teeth. Palp 3-segmented, 
with plumose setae on margins and long, 
thick, simple setae arising from bend in sec- 
ond segment. 

Maxilla (Fig. 3f) exopod evenly rounded, 
with plumose setae along distal margin. 
Scaphognathite bluntly angled on posterior 
lobe, with plumose setae. Endopod and en- 
dites without distinctive characters. 

Maxillule (Fig. 3g) distal endite proxi- 
mally narrow, widening to inflated distal 
end, with thick simple setae on distal mar- 
gin. Proximal endite with thick simple setae 
on distal margin. Endopodal external lobe 
truncate distally and curled under; internal 
lobe reduced, with 3 thick setae at distola- 
teral margin. 

Maxilliped I (Fig. 3h) epipod with plu- 
mose setae on distal margin and distolateral 
surface. Endite tapered distally, subequal to 
first segment of exopod. Exopod with 2 
segments; proximal segment narrow, mar- 
gins parallel and with plumose setae; distal 
segment spatulate, about as long as wide, 
broadest medially, margins with long plu- 
mose setae. Endopod flattened and elon- 
gate, reaching to distal end of proximal ex- 
opodal segment; with plumose setae on 
margins. 



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Maxilliped II (Fig. 3i) dactylus evenly 
rounded, length equal to width, with thick 
simple setae distally. Propodus 1.5 times 
wider than long, with plumose setae on dor- 
sal margin and long simple setae on distal 
margin. Carpus not strongly produced dor- 
sodistally, about 2 times longer than wide, 
with long simple setae on dorsal margin. 
Merus about 3 times longer than wide, mar- 
gins parallel, with simple setae on ventro- 
lateral margin and plumose setae on dor- 
solateral margin. Basi-ischium incomplete- 
ly fused, with plumose setae on margins. 
Exopod V4 times longer than merus, with 
flagellum 1 -segmented. 

Maxilliped HI (Fig. 3j) dactylus evenly 
rounded; with long plumose setae on dorsal 
margin and lateral surface. Propodus with 
longitudinal median row of plumose setae 
on lateral surface; dorsal margin with plu- 
mose setae. Carpus slightly produced onto 
propodus; lateral surface with row of plu- 
mose setae ventromedially; plumose setae 
on dorsal margin. Merus unarmed, with 
plumose setae on all margins. Basi-ischium 
incompletely fused, without crista dentata. 
Exopod 2-segmented, proximal segment 
small; distal segment styliform, tapering, 
approximately Vs length of merus, with plu- 
mose setae scattered on surface; without 
flagellum. 

Pereopod I (Fig. 4a) subchelate. Dactylus 
curved and tapering; lateral and mesial sur- 
faces smooth; dorsal margin with long plu- 
mose and short simple setae; ventral margin 
with short simple setae. Propodus lateral 
surface with numerous short, transverse 
rows of setose rugae; dorsal margin un- 
armed; ventral margin produced distally 
into acute spine; cutting edge lacking teeth, 
lined with long plumose setae; dorsal mar- 
gin with long plumose setae, ventral margin 
with short simple setae. Carpus dorsodistal 
angle produced into strong corneous-tipped 
spine, dorsal margin with few large and 
small spines posteriorly along distal third; 
dorsal and distal margins with long plu- 
mose setae; lateral surface with small distal 
rugose area, with few transverse setose 



ridges on distal half of surface; mesial sur- 
face smooth with few scattered rows of 
long plumose setae, dorsal and ventral mar- 
gins with long plumose setae. Merus un- 
armed; lateral surface with scattered trans- 
verse rows of long plumose setae, dorsal, 
ventral and distal margins with long plu- 
mose setae; mesial side with few short rows 
of setae. Basi-ischium incompletely fused, 
unarmed. Coxa unarmed. 

Pereopods II-IV dactyli laterally com- 
pressed and dorsoventrally expanded. 

Pereopod II (Fig. 4b) dactylus smooth; 
with base to heel straight, heel smoothly 
rounded, heel to tip with rounded broad in- 
dent, tip acute, tip to base broadly convex; 
lateral surface smooth, with several small 
tufts of short setae in approximately straight 
line across medioproximal surface, several 
widely-spaced submarginal tufts of short 
setae dorsodistally; mesial surface smooth, 
ventral margin with long plumose setae, 
dorsal margin with short simple setae, with 
patch of long plumose setae at base. Pro- 
podus dorsal surface smooth, ventral mar- 
gin inflated and rounded; oblique row of 
long plumose setae on distal margin of lat- 
eral surface; distal and ventral margins with 
long plumose setae; dorsolateral surface a 
narrow, oblique, flattened shelf, with short 
setae on dorsal margin and long plumose 
setae on ventral margin; mesial surface with 
elevated, curved, setose ridge from ventral 
junction with dactylus almost to ventral 
proximal junction with carpus. Carpus 
slightly produced dorsodistally; lateral sur- 
face nearly smooth, with irregular broken 
row of rugae and submarginal elevated 
ridge ventrally, rugae and ridge with long 
plumose setae; dorsodistal projection with 
mat of short setae on lateral surface; mar- 
gins with long plumose setae; mesial sur- 
face smooth with long plumose setae in 
scattered patches on surface, and on mar- 
gins. Merus lateral surface with large de- 
calcified window and few scattered setae on 
surface and margins; mesial surface nearly 
smooth with few setae. Basi-ischium in- 



VOLUME 112, NUMBER 1 



153 



completely fused, unarmed. Coxa with 
small spine on anterior margin. 

Pereopod III (Fig. 4c) dactylus with base 
to heel straight, heel broadly rounded and 
low, heel to tip with broad evenly rounded 
indent, tip acute, tip to base smoothly con- 
vex to straight; lateral surface smooth, with 
several small tufts of short setae in roughly 
straight line across medioproximal surface, 
dorsodistal margin with tufts of short setae; 
ventromesial margin with long plumose se- 
tae, dorsal margin with short simple and 
plumose setae; mesial surface smooth, with 
plumose setae proximally at junction with 
propodus. Propodus not inflated dorsoven- 
trally; lateral surface smooth, with long plu- 
mose setae distally, with simple setae on 
dorsal margins, and long plumose setae on 
ventral margin; dorsolateral surface narrow, 
oblique, flattened; mesial surface with scat- 
tered long setae on and near distal margin. 
Carpus produced dorsodistally, exceeding 
proximal margin of propodus by about Va 
length of propodus, pointed but not acute; 
dorsolateral margin unarmed; lateral sur- 
face slightly rugose dorsodistally, with mat 
of short setae and 2 longer rows of setae 
ventrally; mesial surface smooth, with long 
plumose setae on margins and scattered on 
surface. Merus smooth with large decalci- 
fied window; dorsal and ventral margins 
unarmed, with long plumose setae; later- 
odistal margin with long plumose setae; 
mesial surface smooth. Basi-ischium in- 
completely fused and unarmed. Coxa un- 
armed. Female with large gonopore on an- 
terior mesial surface of coxa, surrounded 
with short plumose setae; male with small 
pore on coxa. 

Pereopod IV (Fig. 4d) dactylus with base 
to tip proximally convex becoming con- 
cave, heel and indent absent, tip acute, tip 
to base straight distally becoming convex 
proximally; lateral surface smooth, ventral 
margin with long plumose setae, dorsal 
margin with short simple setae; mesial sur- 
face with dorsal decalcified window, de- 
marcated ventrally by longitudinal elevated 
ridge bearing row of short setae; with setose 



punctae ventral to decalcified window. Pro- 
podus expanded dors ally and ventrally; 
ventral expansion exceeds ventral margin of 
dactylus, margin with long plumose setae; 
dorsal expansion with row of long plumose 
setae medially; lateral and mesial surfaces 
smooth. Carpus not produced dorsodistally; 
lateral and mesial surfaces smooth; dorsal 
margin with short simple and long plumose 
setae; ventral margin with short simple se- 
tae; mesial surface with decalcified win- 
dow. Merus lateral surface with scattered 
short transverse rows of setae, dorsal and 
ventrodistal margins with long plumose se- 
tae; mesial surface with large decalcified 
window proximo ventrally. Basi-ischium in- 
completely fused and unarmed. Coxa un- 
armed. 

Pereopod 5 reduced, slender, lacking dis- 
tinctive features. Coxa of male with large 
mesioproximal gonopore. 

Abdomen (Fig. 4e) somite I approxi- 
mately as long as wide, widest posteriorly; 
dorsal surface with anterior margin straight; 
posterior margin straight, with elevated 
submarginal row of short setae; with small 
transverse decalcified windows laterad to 
segment midline. Somite II dorsal surface 
with submarginal transverse ridge anterior- 
ly; with small transverse decalcified win- 
dows laterad to segment midline just ante- 
rior to submarginal ridge; with tuft of setae 
at posterolateral angle, extending onto pleu- 
ra posteromesially; posterior margin with 
indistinct punctate submarginal groove lat- 
erally; pleura expanded and directed slight- 
ly anteriorly; lateral margins rounded, an- 
terior and lateral margins with long plu- 
mose setae, posterior margin with short se- 
tae. Somite III similar to somite II, but 
narrower, shorter, and lacking anterior sub- 
marginal ridge; small tuft of short thick se- 
tae on posterolateral angle; pleura thinner 
and shorter than on somite II, directed an- 
terolaterally, with setae as in somite II; an- 
terolateral angle acute; dorsal surface 
obliquely flattened anterolaterally. Somite 
IV similar to somite III, but thinner and 
shorter; dorsal surface with thick setae pos- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



terolaterally; pleura thinner and shorter than 
on somite III, directed posterolaterally; dor- 
sal surface obliquely flattened anterolater- 
ally; margin with long plumose setae. So- 
mite V narrower than somite IV; lateral 
margins with short plumose setae; pleura 
absent. Somite VI subequal to somite V in 
width but longer; dorsal surface with short 
transverse rows of setae laterad to midline 
anteriorly; lateral margins with long plu- 
mose setae; pleura absent. 

Females with uniramous, paired pleopods 
on somites II- V; males lacking pleopods. 

Uropods lacking distinctive features. 

Telson of male (Fig. 4f) spatulate, later- 
ally expanded, with length subequal to 
width, produced into short rounded tip dis- 
tally; weakly calcified except for large tri- 
angular anterior plate; median longitudinal 
groove long, extending to distal end of cal- 
cified plate, fined with long thin simple se- 
tae; calcified plate slightly elevated medi- 
ally but without ridge. Telson of female 
(Fig. 4g) ovate, longer than wide, broadly 
triangular, dorsal surface smooth, with me- 
dian longitudinal groove anteriorly; with 
row of setose punctae lateral to midline 
from median of longitudinal groove to dis- 
tal end of groove; margins with long plu- 
mose setae. 

Coloration. — Off-white with whitish se- 
tae in fife and in preservative. 

Distribution. — Indo-West Pacific: Ha- 
waii; Western Austrafia; Seychelles; Mal- 
dives; 3.7-13.7 m. 

Remarks. — This species can easily be 
separated from all of the other Indo-West 
Pacific species of Albunea, except A. mad- 
agascariensis, by the concave shape of the 
lateral margins of the eyes. Although a di- 
rect comparison of A. speciosa with A. 
madagascariensis material is desirable, the 
types (and only known specimens) of A. 
madagascariensis apparently were not de- 
posited in the Museum National d'Histoire 
Naturelle (MNHN), contrary to Thomas sin 
(1973) (Nguyen, pers. coram.). Neverthe- 
less, based on Thomassin's (1973) descrip- 
tion and illustrations, the two species can 



tentatively be separated by several charac- 
ters. A. speciosa has a short rostrum that 
does not reach the distal margin of the oc- 
ular plate, a CG4 comprised of two medial 
elements, a more truncate heel of pereopod 
in, and a rounded distal tip on the telson 
of the male, while A. madagascariensis has 
a long rostrum that well exceeds the distal 
margin of the ocular plate, a CG4 of 1 me- 
dial element, a rounded heel of pereopod 
m, and a pointed distal tip on the telson of 
the male. Given a larger sample size, par- 
ticularly from the westernmost Indo-Pacific, 
all of these characters may prove to repre- 
sent only intraspecific variation of A. spe- 
ciosa, in which case A. madagascariensis 
would become a synonym of A. speciosa. 
Two of Thomassin's specimens exhibited a 
peculiar orange and brown banding pattern 
(Thomassin 1973: 268, pi. 1). No other spe- 
cies of Albunea have been reported with 
anything but almost uniform coloration, 
suggesting the possibifity that this may 
have been an artifact of the environment. 
More study is needed to determine whether 
this is similar to the type of environmen- 
tally-induced color changes observed in 
mole crabs of the genus Hippa Fabricius, 
1787 (e.g., Bauchau & Passelecq-Gerin 
1987). 

Little is known about the biology of this 
species other than the few records of ovig- 
erous females herein reported. Several A. 
speciosa specimens (QM W22285) were 
collected together with the holotype of Al- 
bunea danai new species, but it is unknown 
if the two species regularly coexist. 

Specimens of A. speciosa have been re- 
ported only three times in the literature 
(Dana 1852, Borradaile 1904, Serene 
1973). Examination of Borradaile's (1904) 
specimen from the Maldives confirms its 
identity as this species, making Borradaile's 
(1904) record the first from outside Hawaii, 
although his paper was overlooked by sub- 
sequent researchers. 

Because A. speciosa is now known to be 
a wide-ranging Indo-Pacific species, rather 
than a Hawaiian endemic, it would be use- 



VOLUME 112, NUMBER 1 



155 



ful to understand the origin and subsequent 
distribution of this species in order to better 
understand albuneid biogeography. The Ha- 
waiian specimens of A. speciosa are far re- 
moved from other populations of this spe- 
cies, a phenomenon that has been shown for 
other wide-ranging Hawaiian shallow-water 
fauna with western Indo-Pacific affinities 
(Newman 1986), and they probably reached 
the relatively young Hawaiian Islands by 
long distance dispersal. However, the clos- 
est locality where this species has been re- 
ported is not from the Philippines or nearby 
islands in the northern Indo-Pacific, but 
rather from Western Australia to the south. 
Unless the species is found on the east coast 
of Australia or in the island groups to the 
east, such as the New Hebrides or Society 
Islands, it is difficult to hypothesize the 
eastward dispersal route by which A. spe- 
ciosa reached the Hawaiian Islands. A 
northern or southern dispersal route across 
the Pacific appears equally likely, given the 
evidence currently available. 

Albunea danai, new species 
Figs. 5, 6 

Type material. — Holotype: Hawaii, Hal- 
onu Blow Hole dive site, south shore, 
Oahu, 12.2-13.7 m, coll. R. Holcom, 4 Apr 
1997: 1 male, 16.7 mm CL (QM W23105). 
Allotype: Kailua, Oahu, Mar 1938: 1 fe- 
male, 16.8 mm CL (BPBM SI 1782). Para- 
types: Waikiki, Oahu, 22.7 m, coll. Smith 
and Allen, 23 May 1948: 1 male, 11.6 mm 
CL (BPBM S5343); off Waikiki, Oahu, 6.1 
m, coll. Allen and Smith, 30 May 1948: 1 
female, 10.8 mm CL (AMNH 17716); Ka- 
hana Bay, Oahu, 7.6-9.1 m, coll. "Pele" 
Expedition, 25 Jul 1959: 1 male, 8.6 mm 
CL (BPBM S6775); off Sand Island, Oahu, 
4.8-7.6 m, coll. "Pele" Expedition, 17 July 
1959: 1 male, 10.4 mm CL (AMNH 
17717); Diamond Head, Oahu, 7.6-13.6 m, 
coll. "Pele" Expedition, 9 Sep 1959: 1 fe- 
male, 13.0 mm CL (BPBM S6777); [Ma- 
mala Bay], off Honolulu, Oahu, 27.4-40.2 
m, coll. T. Richert, Feb-Mar 1962: 1 male. 



16.4 mm CL, 1 female, 15.5 mm CL 
(WAM 481-97); [Mamala Bay], off Ewa 
Beach, near Pearl Harbor, Oahu, 27.4 m, 
coll. B. R. Wilson on RA^ "Pele," 5 Jul 
1964: 1 ovigerous female, 10.8 mm CL 
(WAM 143-70). 

Additional material examined (non 
type). — Hawaii: Kanoehe Bay, Oahu, 1924: 
1 male (poor condition), 4.0 mm CL 
(BPBM S7806). 

Type locality. — Halonu Blow Hole dive 
site, south shore, Oahu, Hawaii, USA, Pa- 
cific Ocean. 

Diagnosis. — Carapace slightly longer 
than wide, covered with lightly setose 
grooves. Anterior margin with 8-9 teeth. 
Setal field with narrow lateral elements and 
slightly concave anterior margin; posterior 
lateral elements not extending to posterior 
lateral elements of CGI. CGI with separate 
posterior lateral elements; CG4 with 2-4 
short medial elements; CG5 divided into 2 
lateral elements, not nearly reaching mar- 
gins of CG6; CG6 and CG7 separate, but 
almost approximate; CG8 with 1-2 poste- 
riorly displaced median elements separated 
from lateral elements; CGll absent. Ros- 
trum present, not reaching proximal margin 
of ocular plate. Ocular plate subquadrate. 
Ocular peduncles dorsoventrally flattened 
and elongate, pointed at tip, approximate 
along mesial margin; lateral margin convex; 
mesial margin straight proximally, convex 
distally. Cornea at lateral margin of tip. An- 
tennule with 87-92 flagellar exopodal and 
3-4 endopodal segments. Antenna with 7 
flagellar segments; acute spine on dorsolat- 
eral surface of peduncle segment I. Dactyli 
of pereopods II, III with heels low and 
smoothly rounded. Coxa of pereopod III of 
males with small male pore. Telson of male 
triangular, dorsoventrally flattened laterally 
and distally, inflated medially. Telson of fe- 
male flattened, rounded at tip. 

Description. — Carapace (Fig. 5a) slightly 
wider than long. Anterior margin slightly 
concave on either side of ocular sinus, be- 
coming convex laterally, with 8-9 large 
spines along length. Rostrum a small acute 



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Fig. 5. Albiinea danai, new species. A, male, 16.7 mm CL, QM W23105, holotype; B-F, H-J, ovig. female, 
10.8 mm CL, WAM 143-70; G, male 11.6 mm CL, BPBM 5343. A, carapace and eyes, dorsal view; B, eyes, 
dorsal view; C, left antennule, lateral view; D, right antenna, lateral view; E, left mandible, lateral view; F, left 
maxilla, lateral view; G, left maxillule, lateral view; H, left maxilliped I, lateral view; I, left maxilliped II, lateral 
view; J, left maxilliped III, lateral view. Scale = 1.2 mm (C, D, F, H, J), 1.6 mm (B, E, G), 2.2 mm (I), and 
6.7 mm (A). 



tooth, extending only half the distance be- 
tween distal margin of ocular sinus and oc- 
ular plate. Ocular sinus smoothly concave 
and unarmed. Frontal region smooth; setal 
field broad posteriorly, narrowing anterior- 
ly, with narrow anterior lateral elements and 
slightly concave anterior margin; posterior 



lateral elements thin and not reaching to 
posterior lateral elements of CGI. CGI me- 
dial portion parallel to anterior margin of 
carapace, faintly sinuous, strongly crenu- 
late, divided into medial fragment and 
curved, posteriorly displaced lateral ele- 
ments. Mesogastric region smooth; CG2 



VOLUME 112, NUMBER 1 



157 




Fig. 6. Albunea danai, new species. A, F, male, 16.7 mm CL, QM W23105, holotype; B-E, male 11.6 mm 
CL, BPBM 5343; G, female, 16.8 mm CL, BPBM SI 1782, allotype. A, left pereopod I, lateral view; B. right 
pereopod II, lateral view; C, right pereopod III, lateral view; D, right pereopod IV, lateral view; E, abdominal 
somites I-VI, dorsal view; F, telson of male, dorsal view; G, telson of female, dorsal view. Scale = 3.0 mm (F, 
G) and 4.4 mm (A-E). 



absent; CG3 broken into 6 short elements 
approximately equally spaced between pos- 
terior lateral elements of CGI; CG4 with 
2-4 short medial elements spaced approx- 
imately equally between longer lateral ele- 
ments of CG4. Hepatic region smooth with 
oblique setose groove at median of lateral 
margin. Epibranchial region roughly trian- 



gular, smooth; posterolateral margin with 2 
short rows of setae. Metagastric region 
smooth; CG5 divided into 2 short lateral el- 
ements. CG6 strongly crenulate, strongly 
anteriorly concave medially and sloping out 
to anteriorly convex lateral thirds, median 
and lateral thirds separated by short setae- 
free gap lateral to small depressions. CG7 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



oblique, almost reaching lateral margins of 
median segment of CG6. Cardiac region 
smooth; CG8 present as 2 very short later- 
omedial elements displaced posteriorly 
from longer lateral elements. CG9 present 
as 2 short lateral grooves with gap at mid- 
line. CO 10 present as 2 curved lateral frag- 
ments, with gap between fragments approx- 
imately equal to length of single fragment. 
CGll absent. Branchial region with nu- 
merous short, transverse rows of setae. Pos- 
terior margin deeply and evenly convex, 
with submarginal groove reaching about 
half-way either side of posterior concavity. 
Branchiostegite with short anterior submar- 
ginal spine; anterior region with scattered 
short transverse lines ventral to linea ano- 
murica; with many short rows of setae and 
sparsely covered with long plumose setae 
ventrally; posterior region membranous 
with numerous, irregular fragments and 
sparsely covered with long plumose setae. 

Ocular plate (Fig. 5b) subquadrate with 
narrow indentation; proximal ocular seg- 
ments (Fig. 5b) reduced to small rounded 
calcified area on either side of ocular plate. 
Ocular peduncle (Fig. 5b) elongate, with 
medially convex lateral margins, tapering to 
rounded distal cornea located in lateral 
notch; mesial margin approximate almost 
all of length; mesial and proximolateral 
margins of segment with sparse row of long 
plumose setae; tuft of plumose setae at 
proximolateral ventral angle. 

Antennule (Fig. 5c) segment III narrow 
proximally, expanding distally to twice 
proximal width; with plumose setae on dor- 
sal and ventral margins and sparsely scat- 
tered on lateral surface; dorsal exopodal fla- 
gellum with 87-92 segments and long plu- 
mose setae on dorsal and ventral margins; 
ventral endopodal flagellum short, with 3- 
4 segments and plumose setae on dorsal and 
ventral margins. Segment II medially in- 
flated in dorsal view, with plumose setae on 
dorsal and ventral margins and scattered se- 
tae on ventrolateral third of surface. Seg- 
ment I wider than long, unarmed; dorsal 
third of lateral surface rugose with long plu- 



mose setae; long plumose setae on dorsal 
and ventral margins. 

Antenna (Fig. 5d) segment V about 2 
times longer than wide, with long plumose 
setae on dorsal margin and scattered setae 
on distal half of lateral surface; flagellum 
7-segmented, with long plumose setae on 
dorsal, ventral and distal margins. Segment 
IV expanded distally with long plumose se- 
tae on dorsal, ventral and distal margins, 
and 2 rows of setae on dorsolateral surface. 
Segment III with long plumose setae on 
dorsal and ventral margin. Segment II short, 
widening distally, with plumose setae on 
margins and scattered on lateral surface; an- 
tennal acicle long, thin and exceeding distal 
margin of segment IV by Va the length of 
segment IV, with long plumose setae on 
dorsal margin. Segment I rounded proxi- 
mally, flattened ventrolaterally, with long 
plumose setae on margins; lateral surface 
with acute spine dorsally, with low semi- 
circular dorsolateral lobe ventrodistal to 
spine; segment with ventromesial antennal 
gland pore. 

Mandible (Fig. 5e) incisor process with 
1 tooth; cutting edge with 1 tooth. Molar 
process with 2-3 teeth. Palp 3-segmented, 
with plumose setae on margins and long, 
thick, simple setae arising from bend in sec- 
ond segment. 

Maxilla (Fig. 5f) exopod evenly rounded, 
with plumose setae along distal margin. 
Scaphognathite bluntly angled on posterior 
lobe, with plumose setae. Endopod and en- 
dites without distinctive characters. 

Maxillule (Fig. 5g) distal endite proxi- 
mally narrow, widening to inflated distal 
end, with thick simple setae on distal mar- 
gin. Proximal endite with thick simple setae 
on distal margin. Endopodal external lobe 
truncate distally, and curled under; internal 
lobe reduced, with 3 thick setae at distolat- 
eral margin. 

Maxilliped I (Fig. 5h) epipod with plu- 
mose setae on margins, distolateral surface 
and mesial surface (epipod shown curled in 
Fig. 5h). Endite tapered distally and sube- 
qual to first segment of exopod. Exopod 



VOLUME 112, NUMBER 1 



159 



with 2 segments; proximal segment narrow, 
margins parallel, margins with plumose se- 
tae; distal segment spatulate, about as long 
as wide, broadest medially, margins and 
mesioventral surface with long plumose se- 
tae. Endopod flattened and elongate, reach- 
ing to distal end of proximal exopodal seg- 
ment; plumose setae on margins and me- 
dian of lateral surface. 

Maxilliped II (Fig. 5i) dactylus evenly 
rounded, length equal to width, with thick 
simple setae distally and on distolateral sur- 
face. Propodus 2 times wider than long, 
slightly produced at dorsodistal angle, with 
plumose setae on dorsal margin and long 
simple setae on dorsodistal margin. Carpus 
not produced dorsodistally, about 2 times 
longer than wide; long simple setae on dor- 
sal and distal margins. Merus about 3 times 
longer than wide, margins parallel; with 
simple setae on ventrolateral margin and 
plumose setae on dorsolateral margin. Basi- 
ischium incompletely fused, plumose setae 
on margins. Exopod Vs times longer than 
merus, with flagellum 1 -segmented. 

Maxilliped III (Fig. 5j) dactylus with 
rounded tip; with long plumose setae on 
dorsal margin and lateral surface. Propodus 
with longitudinal median row of plumose 
setae on lateral surface; dorsal margin with 
plumose setae. Carpus slightly produced 
onto propodus; lateral surface with row of 
plumose setae ventromedially; plumose se- 
tae on dorsal margin. Merus unarmed, with 
plumose setae on dorsal and ventral mar- 
gins and scattered on lateral surface. Basi- 
ischium incompletely fused, with weak 
crista dentata of about 2 teeth. Exopod 2- 
segmented: proximal segment small; distal 
segment styliform, tapering, approximately 
Vz length of merus; with plumose setae on 
surface; without flagellum. 

Pereopod I (Fig. 6a) subchelate. Dactylus 
curved and tapering; lateral and mesial sur- 
faces smooth; dorsal margin with long plu- 
mose and short simple setae; ventral margin 
with short simple setae. Propodus lateral 
surface with numerous short, transverse 
rows of setose rugae; dorsal margin un- 



armed; ventral margin produced distally 
into acute spine; cutting edge lacking teeth, 
lined with long plumose setae; dorsal mar- 
gin with long plumose setae, ventral margin 
with short simple setae. Carpus with dor- 
sodistal angle produced into strong corne- 
ous-tipped spine; dorsal margin otherwise 
unarmed; dorsal and distal margins with 
long plumose setae; lateral surface with 
small distal rugose area, with few trans- 
verse setose ridges on distal half of surface; 
mesial surface smooth with few median 
rows of setae, margins with long plumose 
setae. Merus unarmed; lateral surface with 
scattered transverse rows of long plumose 
setae, margins with long plumose setae; 
mesial side with few short rows of setae. 
Basi-ischium incompletely fused, unarmed. 
Coxa unarmed. 

Pereopods II-IV with dactyli laterally 
compressed and dorsoventrally expanded. 

Pereopod II (Fig. 6b) dactylus smooth; 
base to heel straight, heel smoothly round- 
ed, heel to tip with wide acute indent, tip 
acute, tip to base broadly convex distally 
and slightly concave proximally; lateral sur- 
face smooth, with several small tufts of 
short setae in roughly straight line across 
medioproximal surface, several widely 
spaced submarginal tufts of short setae dor- 
sodistally; mesial surface smooth, ventral 
margin with long plumose setae, dorsal 
margin with short simple setae, with patch 
of long plumose setae at base. Propodus 
dorsal surface smooth, ventral margin in- 
flated and rounded; oblique row of long 
plumose setae on distal margin of lateral 
surface; distal and ventral margin with long 
plumose setae; dorsolateral surface a nar- 
row, oblique, flattened shelf, with short se- 
tae on dorsal margin and long plumose se- 
tae on ventral margin; mesial surface with 
elevated, curved setose ridge from ventral 
junction with dactylus almost to ventral 
proximal junction with carpus. Carpus pro- 
duced and gently rounded dorsodistally, 
dorsal margin unarmed; lateral surface 
smooth, with irregular broken row of rugae 
and submarginal elevated ridge ventrally, 



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rugae and ridge with long plumose setae; 
margins with long plumose setae; mesial 
surface smooth with long plumose setae in 
scattered patches on dorsal half of surface 
and on margins. Merus with large median 
decalcified window covering nearly all of 
lateral surface, with few scattered setae on 
surface and margins; mesial surface nearly 
smooth with few setae, with decalcified 
area on proximal Va near junction with basi- 
ischium. Basi-ischium incompletely fused 
and unarmed. Coxa with small spine on an- 
terior margin. 

Pereopod III (Fig. 6c) dactylus with base 
to heel straight, heel broadly rounded and 
slightly produced, heel to tip with broadly 
concave indent, tip acute, tip to base 
smoothly convex distally to straight proxi- 
mally; lateral surface smooth, with several 
small tufts of short setae in approximately 
straight line across medioproximal surface, 
dorsodistal margin with tufts of short setae; 
ventral margin with long plumose setae, 
dorsal margin with short simple and plu- 
mose setae; mesial surface smooth with 
plumose setae proximally at junction with 
propodus. Propodus not inflated dorsoven- 
trally; lateral surface smooth, with long plu- 
mose setae distally, with simple setae on 
dorsal margins; dorsolateral surface narrow, 
oblique, flattened; mesial surface with scat- 
tered long setae on and near distal margin. 
Carpus produced dorsodistally, exceeding 
proximal margin of propodus by about Vs 
length of propodus, rounded; dorsolateral 
margin unarmed; lateral surface slightly ru- 
gose dorsodistally, with mat of short setae 
and row of setae ventrally; mesial surface 
smooth, with long plumose setae on mar- 
gins and scattered on surface. Merus 
smooth, with large decalcified window cov- 
ering nearly half of lateral surface medially; 
dorsal and ventral margins unarmed, with 
long plumose setae; laterodistal margin 
with long plumose setae; mesial surface 
smooth. Basi-ischium incompletely fused 
and unarmed. Coxa with tubercle on ante- 
rior margin. Female with large gonopore on 
anterior mesial margin of coxa, surrounded 



with short plumose setae; male with small 
pore on coxa. 

Pereopod IV (Fig. 6d) dactylus with base 
to tip proximally convex becoming distally 
concave, heel and indent absent, tip acute, 
tip to base straight distally, becoming con- 
vex proximally; lateral surface smooth, 
ventral margin with long plumose setae, 
dorsal margin with short simple setae; me- 
sial surface with dorsal decalcified window, 
demarcated ventrally by longitudinal ele- 
vated ridge with row of short setae; with 
setose punctations ventral to decalcified 
window. Propodus expanded dorsally and 
ventrally; ventral expansion exceeds ventral 
margin of dactylus, margin with long plu- 
mose setae; dorsal expansion with row of 
long plumose setae medially; lateral and 
mesial surfaces smooth. Carpus not pro- 
duced dorsodistally; ventral Va of lateral sur- 
face and mesial surface smooth, dorsal Va of 
lateral surface with mat of short setae; me- 
sial surface with decalcified window; dorsal 
margin with short simple and long plumose 
setae; ventral margin with short simple se- 
tae. Merus lateral surface with scattered 
short transverse rows of setae, dorsal and 
ventrodistal margins with long plumose se- 
tae; mesial surface with large decalcified 
window proximo ventrally. Basi-ischium in- 
completely fused and unarmed. Coxa un- 
armed. 

Pereopod 5 reduced, slender, lacking dis- 
tinctive features. Coxa of male with large 
mesioproximal gonopore. 

Abdomen (Fig. 6e) somite I approxi- 
mately as long as wide, widest posteriorly; 
dorsal surface with anterior margin straight; 
posterior margin straight with elevated sub- 
marginal row of short setae; with small 
transverse decalcified windows laterad to 
segment midline. Somite II dorsal surface 
with submarginal transverse ridge anterior- 
ly; with small transverse decalcified win- 
dows laterad to segment midline just ante- 
rior to submarginal ridge; with tuft of setae 
at posterolateral angle, extending onto pleu- 
ra posteromesially; pleura expanded and di- 
rected slightly anteriorly; lateral margins 



VOLUME 112, NUMBER 1 



161 



rounded, anterior and lateral margins with 
long plumose setae, posterior margin with 
short setae. Somite III similar to somite II, 
but narrower, shorter, and lacking anterior 
submarginal ridge; small tuft of short thick 
setae on posterolateral angle; pleura thinner 
and shorter than on somite II, directed pos- 
terolaterally, with setae as in somite II; an- 
terolateral angle acute; dorsal surface 
obliquely flattened anterolaterally. Somite 
IV similar to somite III, but thinner and 
shorter; dorsal surface with few thick setae 
posterolaterally; pleura thinner and shorter 
than on somite III, directed posterolaterally; 
dorsal surface obliquely flattened anterolat- 
erally; margins with long plumose setae. 
Somite V subequal to somite IV; lateral 
margins with plumose setae; pleura absent. 
Somite VI subequal to somite V in length 
but wider; dorsal surface with short trans- 
verse rows of setae laterad to midline an- 
teriorly and posteriorly; lateral margins 
with long plumose setae; pleura absent. 

Females with uniramous, paired pleopods 
on somites II- V; males without pleopods. 

Uropods lacking distinctive features. 

Telson of male (Fig. 6f) triangular, slight- 
ly longer than wide, with smoothly rounded 
tip; proximal half heavily calcified, distal 
half weakly calcified except for large me- 
dian region; median longitudinal groove ex- 
tending to distal end of calcified area, line 
with long thin simple setae; junction of 
proximal and distal regions demarcated by 
strong line of long setae laterally; calcified 
plate slightly elevated medially but without 
ridge. Telson of female (Fig. 6g) ovate, lon- 
ger than wide, rounded distally; dorsal sur- 
face smooth, with median longitudinal 
groove anteriorly; with row of setose punc- 
tae lateral to midline from posterior end of 
longitudinal groove to % length of telson; 
margins with long plumose setae. 

Coloration. — In life, brownish with red- 
dish-brown setae. Preserved, uniform off- 
white to tan. 

Distribution. — Known only known from 
Oahu, Hawaii; 4.8-40.2 m. 

Etymology. — Named after James D. 



Dana (1813-1895), famed carcinologist and 
describer of A. speciosa, and many other 
species of Indo-Pacific Crustacea. Gender: 
masculine. 

Remarks. — This species is most similar 
to A. carabus (Linnaeus, 1758), from the 
Mediterranean and western Africa, in the 
shape of the dactyli of pereopods II-IV and 
telson morphology. Albunae carabus can be 
easily separated from A. danai, new spe- 
cies, by its CG8 of four medial elements, 
more strongly crenulated CGs, more pro- 
nounced heel on the dactyli of pereopods II 
and III, and a less inflated maxilliped III 
merus. A. danai new species can be distin- 
guished from other Indo-West Pacific spe- 
cies by the triangular shape of the telson of 
the male, the rounded dactyl of pereopod 
III, and setal patterns on the carapace of 
both sexes. All other species of Albunea 
from the Indo-West Pacific region with 
smoothly rounded heels on pereopod III 
(e.g., A. speciosa) also have much more 
strongly setose carapace grooves than A. 
danai, new species, and distinctive male 
telson morphologies. 

As previously indicated, the holotype 
was collected with specimens of A. specio- 
sa (QM W22285). 

This species, unlike A. speciosa, appears 
to be a true Hawaiian endemic based on all 
available data. However, this conclusion 
should be accepted cautiously, given the 
"endemic" label applied to A. speciosa pri- 
or to this study and that endemicity appears 
to be the exception, rather than the rule, in 
the Hawaiian biota (Newman 1986). As a 
whole, the Hawaiian albuneids seem to fit 
in the category of attenuated Indo-West Pa- 
cific fauna (Newman 1986), given the 
markedly greater diversity of albuneids (17 
species in six genera) throughout the rest of 
the Indo-West Pacific. 

Acknowledgments 

I thank R. Van Syoc (CASIZ), R. Jocque 
(MRAC), R. Symonds (UMZC), R. Lemai- 
tre (USNM), J. Griffith and M. Hewitt 



162 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



(WAM), and E. Lazo-Wasem (YPM) for 
making specimens available for study. 
Also, thanks to Nguyen Ngoc-Ho (MNHN) 
for trying to locate the types of A. mada- 
gascariensis. L. Eldredge loaned specimens 
from the collections of BPBM and gener- 
ously offered to deposit two paratypes of A. 
danai in the AMNH. Special thanks to P. 
Davie (QM) for allowing me to study the 
holotype specimen of A. danai. Color pho- 
tographs of both species were generously 
provided by R. Holcom of Hawaii. A. Har- 
vey (Georgia Southern University) was in- 
strumental in providing training in comput- 
er aided biological illustration, co-produced 
Figs. 1 and 2, and reviewed the manuscript. 
Two anonymous reviewers also contributed 
greatly to the final product. 

Literature Cited 

Bauchau, A. G., & E. Passelecq-Gerin. 1987. Morpho- 
logical color changes in anomuran decapods of 
the genus Hippa. — Indo-Malayan Zoology 4: 
135-144. 

Benedict, J. E. 1904. A new genus and two new spe- 
cies of crustaceans of the family Albuneidae 
from the Pacific Ocean; with remarks on the 
probable use of the antennule in Albunea and 
Lepidopa. — Proceedings of the United States 
National Museum 27(1367):621-625. 

Borradaile, L. A. 1904. Marine crustaceans. XIII. The 
Hippidea, Thalassinidea and Scyllaridea. In J. 
S. Gardiner, ed. The fauna and geography of the 
Maldive and Laccadive Archipelagoes 2:750- 
754, pi. 58. 

Boyko, C. B., & A. W. Harvey. 1999. Crustacea De- 
capoda: Albuneidae and Hippidae of the Indo- 
West Pacific region. In A. Crosnier, ed. Resul- 
tats des Campagnes MUSORSTOM, volume 
20. — Bulletin du Museum National d'Histoire 
Naturelle 180:377-404 (in press). 

Dana, J. D. 1852. Crustacea. U. S. Exploring Expedi- 
tion. During the years 1838, 1839, 1840, 1841, 
1842. Under the command of Charles Wilkes, 
U.S.N. 13(1). Philadelphia: C. Sherman. 685 
pp. 

. 1855. United States Exploring Expedition. 

During the years 1838, 1839, 1840, 1841, 1842. 
Under the command of Charles Wilkes, U.S.N. 
Atlas. Crustacea. Philadelphia: C. Sherman, 27 
pp., 96 pis. 

Edmondson, C. H. 1946. Reef and shore fauna of Ha- 
waii. — Bemice P. Bishop Special Publication 
22:381 pp. 



Efford, I. E., & J. Haig. 1968. Two new genera and 
three new species of crabs (Decapoda: Ano- 
mura: Albuneidae) from Australia. — Australian 
Journal of Zoology 16(6):897-914. 

Evans, A. C. 1967. Syntypes of Decapoda described 
by William Stimpson and James Dana in the 
collections of the British Museum (Natural His- 
tory). — Journal of Natural History 1:399-411. 

Fabricius, J. C. 1787. Mantissa insectorum sistens eor- 
um species nuper detectus adjectis characteribus 
genericis differentiis specificis, emendationibus, 
observationibus. Hafniae. 1:348 pp. 

Gordon, I. 1938. A comparison of the two genera Al- 
bunea and Lepidopa (Crustacea, Anomura), 
with description of a new species from Singa- 
pore. — Bulletin of the Raffles Museum 14:186- 
197. 

Harvey, A. W., & E. M. De Santo. 1997. A new spe- 
cies of Pachycheles from the Hawaiian Islands 
(Crustacea: Decapoda: Porcellanidae). — Pro- 
ceedings of the Biological Society of Washing- 
ton 110(l):65-68. 

Henderson, J. R. 1893. A contribution to Indian car- 
cinology. — Transactions of the Linnaean Soci- 
ety of London, 2nd ser. 5(10):325-458, pis. 36- 
40. 

Holthuis, L. B. 1958. Crustacea Decapoda from the 
northern Red Sea (Gulf of Aqaba and Sinai Pen- 
insula) II. Hippidea and Brachyura (Dromiacea, 
Oxystomata, and Grapsoidea). — Israel Sea Fish- 
eries Research Station Bulletin 1 7(9) :4 1-54. 

Latreille, P. A. 1825. Families naturelles du regne an- 
imal, exposees succinctement et dans un ordre 
analytique, avec I'indication de leurs genres. 
Paris: J.-B. Bailliere. 570 pp. 

Linnaeus, C. 1758. Systema namrae per regna tria na- 
turae, secundum classes, ordines, genera, spe- 
cies, cum characteribus, differentis, synonymis, 
Locis. 824 pp. 

Miers, E. J. 1878. Revision of the Hippidea. — Zoolog- 
ical Journal of the Linnean Society 14(76):312- 
336, pi. 5. 

Newman, W. A. 1986. Origin of the Hawaiian marine 
fauna: dispersal and vicariance as indicated by 
barnacles and other organisms. In R. H. Gore 
& K. L. Heck, eds. Crustacean Biogeography. 
Crustacean Issues 4. Rotterdam: A. A. Balkema. 
pp. 21-49. 

Ortmann, A. E. 1896. Die geographische verbreitung 
der decapodengruppe der Hippidea. — Zoologis- 
che Jahrbucher, Abteilung fiir Systematik, Geo- 
graphic und Biologic der Thiere 9(2):2 19-243. 

Serene, R. 1973. A new species of Decapoda Hippi- 
dea: Albunea mariellae nov. sp. from the Banda 
Sea. — Crustaceana 24(3):26 1-264, 2 pis. 

Stillman, J. H., & G. N. Somero. 1996. Adaptation to 
temperature stress and aerial exposure in con- 
generic species of intertidal porcelain crabs (ge- 



VOLUME 112, NUMBER 1 



163 



nus Petrolisthes): correlation of physiological 
biochemistry and morphology with vertical dis- 
tribution. — Journal of Experimental Biology 
199(8): 1845-1855. 
Stimpson, W. 1858. Prodromus descriptionis animal- 
ium evertebratum, quae in Expeditione ad 
Oceanum Pacificum Septentrionalem, a Repub- 
lica Federata missa, Cadwaladaro Ringgold et 
Johanne Rodgers Ducibus, observavit et des- 
cripsit. Pars VII. Crustacea Anomura. — Pro- 
ceedings of the Academy of Natural Sciences 
of Philadelphia 10:225-252. 



Thomassin, B. A. 1973. Albunea madagascariensis n. 
sp., nouvelle espece d'Hippidea (Decapoda, An- 
omura) des sables coralliens de la region de Tu- 
lear (S. W. de Madagascar). — Crustaceana 
24(3):265-274. 

Weber, E 1795. Nomenclator entomologicus secundum 
entomologiam systematicam ill. Fabricii adjec- 
tis speciebus recens detectis et varietatibus. Chi- 
lonii et Hamburgi: Carolum Ernstum Bohn. viii 
+ 172 pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):164-174. 1999. 

Two new species of Hansenium (Crustacea: Isopoda: Asellota) from 

Madang, Papua New Guinea 

Kathrin S. Bolstad and Brian Kensley 

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian 

Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — Two new species of Hansenium, H. tropex and H. thomasi, are 
described from Paddock Reef, Madang, Papua New Guinea. H. tropex is char- 
acterized by the possession of a broad oar-shaped lobe of the carpus of pereo- 
pod 1 in the male. H. thomasi is characterized by a narrow tapering mesially- 
directed lobe of the carpus of pereopod 1 of the male. The genus is redefined, 
with its chief character being the presence of a large carpal lobe on the first 
male pereopod. The eight stenetriid genera are compared on the basis of six 
characters. 



The shallow water marine crustaceans 
(with the exception of the Amphipoda) of 
the Papua New Guinea region are poorly 
known. The Isopoda have received scant at- 
tention (in all, only about 15 species have 
been recorded from the region.) Stebbing 
(1900), recorded three cirolanids, one cy- 
mothoid and one sphaeromatid from New 
Britain, New Guinea, Nobili (1905) de- 
scribed a corallanid and a bopyrid, while 
Nierstrasz (1931) lists three cymothoids, 
two cirolanids, three corallanids, and one 
sphaeromatid in his catalogue of isopod re- 
cords. Bruce (1982, 1993, 1994) has de- 
scribed cirolanids from the Madang area, 
while Keable (1997) added further cirolanid 
records; Jones et al. (1983) described three 
species of corallanids; Williams & Bunk- 
ley- Williams (1992) described two new 
species of cymothoids. No asellote isopods 
have been reported. The finding of two very 
distinctive stenetriids was, therefore, 
thought to be well worth recording. 

This paper was written while the authors 
participated in the Mentorship Program of 
the Thomas Jefferson High School for Sci- 
ence and Technology in Alexandria, Virgin- 
ia. 

The material described in this paper was 
collected by J. D. Thomas and J. Clark- 



Walker of the National Museum of Natural 
History, Smithsonian Institution. The col- 
lectors were carrying out a survey of the 
shallow water marine invertebrate fauna of 
Madang, Papua New Guinea, in coopera- 
tion with the Christensen Research Insti- 
tute. Jebb & Lowry (1995) provide a useful 
description of the habitats of Madang La- 
goon. 

Suborder Asellota 

Family Stenetriidae Hansen, 1905 

Genus Hansenium Serov & Wilson, 1995 

Hansenium Serov & Wilson, 1995: 72. 

Diagnosis. — Lateral tooth of cephalon 
moderately well developed, antennal tooth 
subequal to lateral, or rounded. Rostrum 
short, rectangular, anterior margin truncate. 
Eyes reniform, of 13-19 ommatidia. Pereo- 
pod 1 in male with carpus produced pos- 
terodistally into lobe; propodus longer than 
width of palm, latter often with few teeth 
close to articulation of dactylus. Pleopod 2 
in male with rounded protopodal lobe dis- 
tally, appendix masculina of endopod dis- 
tally somewhat broadened, truncate, often 
ringed by cuticular hairs. 

Type species. — Stenetrium hanseni No- 
bili, 1906. 



VOLUME 112, NUMBER 1 



165 



Remarks. — Serov & Wilson (1995), di- 
vided the genus Stenetrium into five sepa- 
rate genera, and reviewed the remaining 
three genera in the family Stenetriidae. 
They also listed the constituent species for 
each of the genera. Examination of the two 
present species, as well as undescribed ma- 
terial from the Indian Ocean has led us to 
revise the diagnosis of Hansenium, as given 
above. The presence of a variously devel- 
oped posterodistal lobe on the carpus of pe- 
reopod 1 in the male would seem to be the 
most characteristic synapomorphy of the 
genus, along with the presence of one or 
more teeth on the short propodal palm. The 
extreme development of this lobe into the 
expanded and flattened oar-like structure 
seen in H. tropex and H. wilsoni (Miiller, 
1991a) is approached in two undescribed 
species from the Indian Ocean. A review of 
the 1 3 species of Hansenium listed by Ser- 
ov & Wilson (1995) shows that only Ste- 
netrium bowmani Kensley, 1984, and S. gil- 
kertense Nordenstam, 1946 do not possess 
a carpal lobe on pereopod 1 in the male. 

Table 1 summarizes the six characters 
thought most significant in defining the 
eight genera of the Stenetriidae. 

Species included in Hansenium. 
Hansenium caicosense (Kensley & Heard, 

1991). Turks & Caicos Islands. 
Hansenium dodo (Miiller, 1991b). Reunion 

Island. 
Hansenium entale (Nordenstam, 1946). Gil- 
bert Islands. 
Hansenium hanseni (Nobili, 1906). Tua- 

motu Islands. 
Hansenium monodi (Nordenstam, 1946). 

Gulf of Suez; Seychelles. 
Hansenium spathulicarpus (Kensley, 1984). 

Belize. 
Hansenium stebbingi (Richardson, 1902) 

(= Stenetrium antillense Hansen, 1904, 

and = Stenetrium occidentale Hansen, 

1904). Bermuda; Belize; St. Thomas, 

West Indies. 
Hansenium thomas n. sp. Madang, Papua 

New Guinea. 



Hansenium tropex n. sp. Madang, Papua 

New Guinea. 
Hansenium wilsoni (Miiller, 1991a). Moo- 

rea. 

Hansenium tropex, new species 
Figs. 1-2 

Material examined. — Holotype, USNM 

253348, S tl 4.8 mm, Paratypes, USNM 

253349, 2 ovigerous 9 tl 3.5 mm (dam- 
aged), 5.0 mm. Paddock Reef, Madang, 
Papua New Guinea, coral rubble, 3-4 m, 
coll. J. D. Thomas, 14 Jan 1989. 

Description. — Body slender, about 4 
times as long as wide, cephalon about twice 
as wide as long. Rostrum short, rectangular, 
apically truncate, subequal in length to an- 
tennal teeth. Antennal teeth elongate, acute. 
Lateral teeth pronounced, subequal to an- 
tennal teeth. Frontal margin of cephalon be- 
low rostrum slightly convex. Eyes reniform, 
consisting of about 14 ommatidia. 

Antennule, flagellum of about 20 articles; 
1 aesthetasc on first article, 2 on second ar- 
ticle. Mandible with 4-cusped incisor; 3- 
cusped lacinia mobilis; left mandible with 
spine row of 4 comb setae; right mandible 
spine row with 7 comb setae and 3 simple 
setae; molar with 8 plumose setae below 
marginal serrations. Maxilla 1 inner lobe 
with 2 large fringed setae and 2 smaller 
simple setae; outer lobe with 8 fringed se- 
tae. Maxilla 2 outer lobe with 4 stout 
fringed setae and 1 simple seta; middle lobe 
with 5 stout fringed setae; inner lobe with 
8 simple setae and 4 stout plumose setae on 
mesial margin. Maxilliped palp with 3 
proximal articles wider than 2 distal arti- 
cles, latter longer than wide; endite broad 
with 6 broad flattened fan setae distally; 
mesial margin with 5 coupling hooks. 

Pereonites 1-4 decreasing in length pos- 
teriorly, with anterolateral margins acute; 
pereonites 5-7 increasing in length poste- 
riorly; pereonites 5-6 with rounded pos- 
terolateral margins; pereonite 7 with angu- 
lar posterolateral margins. Male pereopod 1 
elongate, slender; dactylus equal in length 



166 



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VOLUME 112, NUMBER 1 



167 




Fig. 1. Hansenium tropex, new species. A, (?, dorsal view; B, Pereopod 1 d; C, Pereopod 1 9 ; D. Pereopod 
2; E, Pereopod 7; F, Operculum 9; G, Pleopod 1 cJ; H, Pleopod 3. 



to propodal palm; short unguis extending 
beyond proximal spine of propodal palm; 
propodal palm serrate; carpus with large 
stout distally rounded densely setose lobe 



projecting anteroventrally, about 0.8 times 
length of propodus (Fig. IB). Female pe- 
reopod 1 significantly smaller than in male, 
with sparser setae; dactylus equal in length 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Hansenium tropex, new species. A, Right mandible; B, Left mandible; C, Mandibular palp; D, Maxilla 
1; E, Maxilliped; F, Maxilla 2; G, Antennule. 



VOLUME 112, NUMBER 1 



169 



to propodal palm, with 10 short simple se- 
tae on posterodistal margin; propodal palm 
with 6 short simple setae, terminating in 2 
long setae; carpal lobe much less prominent 
than in male. 

Pleon having 2 vestigial pleonites plus 
pleotelson; posterolateral spines pro- 
nounced; postanal margin nearly semicir- 
cular. Male pleopod 1 biramous with rami 
elongate; mesial margins of rami parallel; 
lateral margins rounded and sparsely setose. 
Female pleopod 2 1.4 times longer than 
wide, pentagonal, with outer margins 
sparsely setose. Pleopod 3 operculiform, 
with outer ramus broad, sparsely setose; in- 
ner ramus narrow and nearly triangular, 
with 4 terminal setae. 

Remarks. — Hansenium tropex bears the 
closest resemblance to H. wilsoni (Muller, 
1991a), but several features distinguish the 
two. In H. tropex, the antennal spines are 
clearly longer than the lateral spines, while 
in H. wilsoni, the antennal spines are re- 
duced, rounded, and shorter than the lateral 
spines. The anterior margin of the cephalon 
in H. tropex is convex, straight in H. wil- 
soni. In H. tropex the maxilliped has six 
flattened fan setae on the endite where H. 
wilsoni has three. In pereopod 1 of the male 
in H. tropex, the carpal process extends 
about Va the length of the propodus, the pro- 
podal palm terminates in a simple blunt 
seta, the merus is not produced and the is- 
chium is sparsely setose; in H. wilsoni, the 
carpal lobe extends beyond the propodus 
and the dactylus distally, the propodal palm 
has no articulated seta, the merus is slightly 
produced, and there is a dense cluster of 
setae on the ischium. Pleopod 1 in H. tro- 
pex females is nearly pentagonal, with setae 
at regular intervals on the lateral margins; 
in H. wilsoni, the female pleopod 1 oper- 
culum has only four setae and is more tri- 
angular in shape. Pleopod 1 in the male of 
H. tropex is nearly semicircular, tapering 
proximally and distally, while in H. wilsoni 
the male pleopod 1 only narrows distally. 

Etymology. — The specific name is de- 
rived from the Greek 'tropex', an oar, refers 



to the paddle-like carpal lobe of the male 
pereopod 1, and is used as a noun in ap- 
position. 

Hansenium thomasi, new species 
Figs. 3-5 

Material examined. — Holotype, USNM 

253350, S tl 4.3 mm, Paratypes, USNM 

253351, 3 (5, 8 ovigerous $ tl 3.4-4.5 mm, 
4 juveniles. Paddock Reef, Madang, Papua 
New Guinea, 1.5-4 m, coll. J. D. Thomas, 
14 Jan 1989. 

Diagnosis. — Body about 3.5 times longer 
than wide, width of cephalon about 1.5 
times length. Rostrum short, rectangular, 
anterior margin truncate; frontal margin of 
cephalon convex posterior to rostrum. An- 
tennal spines equal in length to rostrum; lat- 
eral spines subequal in length to antennal 
spines. Eyes reniform with about 19 om- 
matidia. 

Antennule of about 8 articles; 2 aesthe- 
tascs on terminal article; 1 aesthetasc on 
subterminal; 1 aesthetasc on following ar- 
ticle. Mandible with 4-cusped incisor and 
3-cusped lacinia mobilis; left mandible with 
spine row of 4 comb setae, left molar with 
8 fringed setae; right mandible with spine 
row of 7 comb setae, 2 plumose setae, and 
2 simple setae; right molar with 5 plumose 
setae. Maxilla 1 inner ramus with 3 stout 
plumose setae, 1 shorter simple seta; outer 
ramus with 5 stout comb setae and 5 stout 
simple setae. Maxilla 2 outer lobe with 2 
long fringed terminal setae, 2 shorter simple 
setae, mesial margin with 6 simple setae; 
middle lobe with 3 long fringed terminal 
setae, 1 simple terminal seta, 6 simple setae 
on mesial margin; inner lobe with 6 fringed 
setae and 6 simple setae. Maxillipedal en- 
dite broad; distal margin with 6 round flat- 
tened fan setae and 5 narrow fringed setae, 
with 1 short stout simple terminal seta; me- 
sial margin with 4 coupling hooks, 4 
fringed setae. 

Pereonites 1-3 increasing in length pos- 
teriorly; pereonite 4 longer than pereonite 
5; pereonite 6 longer than pereonite 7; cox- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 3. Hansenium thomasi, new species. A, S, dorsal view; B, Antennule; C, Left mandible; D, Right 
mandible; E, maxilla 2; F, Maxilla 1; G, Maxilliped. 



VOLUME 112, NUMBER 1 



171 




Fig. 4. Hansenium thomasi, new species. A, Pereopod 1 d; B, Pereopod 1 9 ; C, Pereopod 2; D, Pereopod 
7; E, Uropod; F, Pleopod 1 c?; G, Pleopod 3; H, Operculum 9; I, Pleopod 2 6. 



172 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 






/ 


%Mh ^ " -^p 




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f 


T> £71600 10KV X65'. 0* ■ ■ iUmrn 



Fig. 5. 
situ. 



Hansenium thomasi, new species. A, Cephalon in dorsal view; B, Left and right pereopod 1 c? in 



ae visible on pereonites 1 and 3-5; pereon- 
ites 1-2 with acute anterolateral angles; per- 
eonites 3-4 with concave lateral margins; 
pereonite 5 with flat lateral margins, round- 
ed posterolateral angle; pereonites 6—7 lat- 
erally broad and rounded. 

Male pereopod 1 dactylus elongate, nar- 
row with acute unguis, extending far be- 
yond propodal palm; propodal palm broad, 
with 3 sharp teeth, posterodistal angle 
greatly produced into narrow lobe termi- 
nating in simple stout seta; carpus with 
strongly produced posterodistal setose lobe; 
merus with small posterodistal projection 
and tufts of setae on postero- and antero- 
distal margins. Female pereopod 1 much 
smaller than in male; dactylus equal in 
length to broad propodal palm, with row of 
short setae along posterodistal margin; pro- 
podal palm broader than in male, with 7 
teeth; carpal process much smaller than in 
male; merus with setal tufts and low anter- 
odistal lobe. Pereopods 2-7 similar, carpus 
with 2 posterodistal setae; propodus with 
4-6 stout setae on posterior margin. 



Pleon with 2 vestigial pleonites plus 
pleotelson, latter with single produced pos- 
terolateral spine on each lateral margin; 
postanal region gently convex. Male pleo- 
pod 1 mesial margins finely setose, parallel; 
lateral margin convex with 6—7 setae dis- 
tally. Male pleopod 2 endopod simple, outer 
margin with about 9 setae; protopod distally 
subacute, exopod rounded. Female pleopod 
2 operculum subtriangular, 1.3 times longer 
than wide, sparsely setose. Pleopod 3 rami 
together sub-circular, with lateral margin of 
exopod setose; endopod nearly triangular 
with 6 distal spines. 

Remarks. — Hansenium thomasi bears 
considerable resemblance to H. entale 
(Nordenstam, 1946) from the Gilbert Is- 
lands in the Pacific. Comparison of the 
male pereopod 1, however, easily distin- 
guishes the two species. The dactylus of H. 
thomasi is less curved. There are three teeth 
on the propodal palm of H. thomasi, more 
apically acute than the four found on H. 
entale. The propodal teeth of H. thomasi are 
well separated from each other, whereas in 



VOLUME 112, NUMBER 1 



173 



H. entale their fused bases join the propodal 
palm. Hansenium thomasi has a large pos- 
terodistal propodal lobe, separate from the 
teeth, which terminates distally in a simple 
stout seta. Hansenium entale has a similar 
large toothlike process, but it is more di- 
rectly a part of the propodal palm, close to 
the other four teeth. Both H. thomasi and 
H. entale have produced carpi in the male 
pereopod 1, but the process is more setose 
and distally rounded in H. thomasi. The 
meral process in H. thomasi is much shorter 
than in H. entale, where it extends almost 
half the length of the carpus. 

The anterior margin of the cephalon in 
H. thomasi is gently convex, straight in H. 
entale. Hansenium entale has a character- 
istic pigment pattern, with a line across the 
anterior cephalon and a broad band between 
the eyes; H. thomasi apparently has no pig- 
mentation. In H. entale, the coxae are vis- 
ible on pereonites 3 and 4 only, while in S. 
thomasi they are visible on pereonites 3—5. 

Etymology. — The species is named for 
James Darwin Thomas, amphipod special- 
ist, who collected the specimens. 

Acknowledgments 

We thank Dr. Jim Thomas and Ms. Janice 
Clark- Walker for collecting the material de- 
scribed herein. The collectors' work was 
supported by funding from the National 
Geographic Society, the Smithsonian Insti- 
tution, and the Christensen Research Insti- 
tute of Madang, Papua New Guinea. We 
thank Dr. Niel Bruce, and two anonymous 
reviewers for their valuable comments on 
an earlier draft of this paper. We thank Ms. 
Marilyn Schotte who assisted with SEM 
preparation of specimens. 

Literature Cited 

Bruce. N. L. 1982. Records of isopod Crustacea (Cor- 
allanidae. Cirolanidae) from Papua New Guin- 
ea, with the description of a new species. — 
Journal of Crustacean Biology 2:612-618. 

. 1993. Two new genera of marine isopod crus- 
taceans (Cirolanidae) from Madang. Papua New 



Guinea. — Memoirs of the Queensland Museum 
31(1):1-15. 

. 1994. Cirolana and related marine isopod 

crustacean genera (family Cirolanidae) from the 
coral reefs of Madang, Papua New Guinea. — 
Cahiers de Biologic Marine 35:375-413. 

Hansen, H. J. 1904. On the morphology and classifi- 
cation of the Asellota group of crustaceans, with 
descriptions of the genus Stenetrium Hasw. and 
its species. — Proceedings of the Zoological So- 
ciety of London 19:302-331. 

. 1905. On the morphology and classification 

of the Asellota group of crustaceans with de- 
scriptions of the genus Stenetrium Hasw. and its 
species. — Proceedings of the Zoological Socie- 
ty of London 1904, (2 Suppl. n):302-331. 

Has well, W. A. 1881. On some new Australia marine 
Isopoda. Part I. — Proceedings of the Linnean 
Society of New South Wales 5:470-481. 

Jebb, M. H. P, & J. K. Lowry. 1995. Natural history 
of Madang Lagoon with an appendix to col- 
lecting localities. Pp. 1-24 in J. K. Lowry, ed.. 
The Amphipoda (Crustacea) of Madang La- 
goon, Papua New Guinea, Part 1. — Records of 
the Australian Museum, Supplement 22:1-174. 

Jones, D. A., J. D. Icely, & S. M. Cragg. 1983. Some 
corallanid isopods associated with wood from 
Papua New Guinea, including three new species 
(Isopoda: Corallanidae). — Journal of Natural 
History 17:837-847. 

Keable, S. J. 1997. The Cirolanidae (Crustacea: Iso- 
poda) of Darwin Harbour, Northern Territory, 
with additional records from northern Australia 
and Papua New Guinea. Pp. 245-278 in J. R. 
Hanley, G. Caswell, D. Megirian, and H. K. 
Larson, eds.. Proceedings of the Sixth Interna- 
tional Marine Biological Workshop. The marine 
flora and fauna of Darwin Harbour, Northern 
Territory, Australia. Museums and Art Galleries 
of the Northern Territory and the Australian 
Marine Sciences Association, Darwin. 

Kensley, B. 1984. The Atlantic Barrier Reef ecosystem 
at Carrie Bow Cay, Belize, III. New marine Is- 
opoda. — Smithsonian Contributions to Marine 
Sciences 24:1-81. 

, & R. Heard. 1991. Studies on the Crustacea 

of the Turks and Caicos Islands. British West 
Indies I. Four new marine isopod crustaceans 
from the vicinity of Pine Cay. — Gulf Research 
Reports 8(3):237-246. 

Miiller, H. G. 1991a. The marine isopod family Ste- 
netriidae from coral reefs at Bora Bora and 
Moorea. Society Islands, with descriptions of 
four new species (Crustacea). — Revue Suisse de 
Zoologie 1991. 98(l):70-76. 

. 1991b. Stenetriidae from coral reefs at Re- 
union Island, southern Indian Ocean. Descrip- 
tion of three new species (Crustacea: Isopoda: 



174 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Asellota). — Senckenbergiana Biologia 71(4/6): 
303-318. 

Nierstrasz, H. F 1931. Die Isopoden der Siboga-Ex- 
pedition III. Isopoda Genuina II. Flabellifera. — 
Siboga-Expeditie Monografie 32C: 121-232. 

Nobili, G. 1905. Decapodi e isopodi della Nuova 
Guinea tedesca, raccolti dal Sign. L. Biro. — An- 
nates Musei Nationalis Hungarici 3:480-507. 

. 1906. Diagnoses preliminaires de crustaces, 

decapodes et isopodes nouveaux recueillis par 
M. le Dr. G. Seurat aux iles Touamotu. — Bul- 
letin du Museum National d'Histoire Naturelle, 
Paris 1906, 5:256-270. 

Nordenstam, A. 1946. Marine Isopoda from Professor 
Dr. Sixten Bock's Pacific Expedition 1917- 
1918.— Arkiv for Zoologi 37A(7):26-29. 

Richardson, H. 1902. The marine and terrestrial iso- 
pods of the Bermudas, with descriptions of new 
or little known species. — Proceedings of the 
United States National Museum 11:277-310. 

Schultz, G. A. 1978. Protallocoxoidea new superfam- 
ily (Isopoda Asellota) with a description of Pro- 
tallocoxa weddellensis new genus, new species 



from the Antarctic Ocean. — Crustaceana 34(3): 
245-250. 

-. 1979. A new asellote (Stenetriidae) and two, 
one new, Anthuridea (Anthuridae) from Ber- 
muda (Crustacea, Isopoda). — Proceedings of 
the Biological Society of Washington 91:904- 
911. 
-. 1982. Species of Protallocoxoidea and Ste- 



netrioidea (Isopoda, Asellota) from the Antarc- 
tic and southern seas. Biology of Antarctic Seas 
10. — Antarctic Research Series, 32(2): 17-62. 

Serov, R A., & G. D. F Wilson. 1995. A review of the 
Stenetriidae (Crustacea: Isopoda: Asellota). — 
Records of the Australian Museum 47(1):39- 
82. 

Stebbing, T. R. R. 1900. On Crustacea brought by Dr. 
Willey from the South Seas. — Willey's Zoolog- 
ical Results 5:605-690. 

Williams, E. H. Jr., & L. Bunkley- Williams. 1992. 
Renocila loriae and R. richardsonae (Crustacea: 
Isopoda: Cymothoidae), external parasites of 
coral reef fishes from New Guinea and the Phil- 
ippines. — Proceedings of the Biological Society 
of Washington 105:299-309. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):175-180. 1999. 

Caecidotea simulator, a new subterranean isopod from the Ozark 
Springfield Plain (Crustacea: Isopoda: Asellidae) 

Julian J. Lewis 
217 W. Carter Avenue, Clarksville, Indiana 47129 U.S.A. 

Abstract. — With the description of Caecidotea simulator, a. phreatobitic iso- 
pod reported herein from Arkansas and Kansas, a total of six species of sub- 
terranean asellids is now known from the Springfield Plain section of the Ozark 
Plateau. The presence of C. macropropoda in Arkansas, a subterranean species 
previously known with certainty only from Oklahoma, is confirmed by two 
collections examined from northeastern Arkansas. 



The Springfield Plain is a region of gent- 
ly rolling karst landscape formed on flat- 
bedded limestones in the southwestern part 
of the Ozark Plateau. This area of the cen- 
tral United States includes parts of south- 
western Missouri, northwestern Arkansas, 
northeastern Oklahoma, and a tiny piece of 
southeastern Kansas. Previously reported 
from the Springfield Plain were the subter- 
ranean asellids Caecidotea ancyla (Flem- 
ing, 1972), C. antricola Creaser (1931), C 
macropropoda Chase & Blair (1937), C 
stiladactyla Mackin & Hubricht (1940), and 
C. steevesi (Fleming 1972). The type spec- 
imens of C. steevesi were examined and 
three species were found to be present: (1) 
C. steevesi from Carrico Cave, Dade Coun- 
ty, Missouri (type-locality), and Gitten 
Down Mountain Cave, Adair County, 
Oklahoma; (2) C antricola, also in the Car- 
rico Cave collection; and (3) the new spe- 
cies described below from Baxter Springs, 
Cherokee County, Kansas. 

Caecidotea simulator, new species 
Figs. 1, 2a, c-f, 3a, b 

Asellus steevesi Fleming 1972: 491-494 
[Baxter Springs, Kansas record]; 1973: 
295, 300 [in part]. 

Material examined. — Kansas: Cherokee 
County, Baxter Springs, seeps off 7th Av- 
enue, 12 Jun 1964, J. R. Holsinger, lOdd, 



13$ ?. — Arkansas: Washington County, O. 
A. Lasterling's well, 0.25 mile west High- 
way 71, Fayetteville, 22 Jul 1965, E. H. 
Schmitz, 15(5 (?, 11$$. 

An 11.0mm 6 from Baxter Springs, 
Kansas is designated as the holotype 
(USNM 216971), with paratypes from Bax- 
ter Springs (USNM 222477) and Laster- 
ling's well (USNM 216972). All of the ma- 
terial has been deposited in the collection 
of the National Museum of Natural History, 
Smithsonian Institution. 

Description. — Eyeless, unpigmented, 
longest male to 19mm, female to 13mm. 
Body slender, linear, about 6.5 X as long as 
wide. Margins of head, pereonites, and 
pleotelson moderately setose. Head about 
1.5 X as wide as long, anterior margin con- 
cave, postmandibular lobes moderately pro- 
duced. Pleotelson about 2X as long as wide, 
sides subparallel, caudomedial lobe slightly 
produced. 

Mandibles with 4-cuspate incisors and la- 
cinia mobilis, palp with rows of plumose 
setae on distal segments. Maxilla 1 with 5 
robust plumose setae on inner lobe, 13 
spines on outer lobe. Antenna 1 flagellum 
to 18 segments, esthete formula 3-0-1-0-1. 
Antenna 2, last segment of peduncle about 
1.2X length of preceding segment, flagel- 
lum of 19mm S with 168 segments. 

Male pereopod 1, propus 1.5X as long as 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 1. Caecidotea simulator, male from Baxter Springs, Cherokee Co., Kansas: (a) habitus, (b) distal seg- 
ments of antenna 1 flagellum, (c) right mandible, incisor, (d) left mandible, incisor and lacinia mobilis, (e) 
mandibular palp, (f) maxilla 1, outer lobe, (g) same, inner lobe, (h) pleopod 3, (i) pleopod 5, (j) pereopod 4. 



VOLUME 112, NUMBER 1 



177 




Fig. 2. Caecidotea simulator, (a, c-f) male from Baxter Springs, Cherokee Co., Kansas, and Caecidotea 
steevesi, (b, g) male from Carrico Cave, Dade Co., Missouri: (a) pleopod 1, (b) same, (c) gnathopod propus, 
palmar margin, (d) pleopod 2, (e) same, endopod tip, (f) same, caudal processes folded over anterior processes 
under coverslip, (g) pleopod 2, endopod tip. 



wide, palm with raised proximal spine, me- 
dian triangular process and slightly bicuspid 
distal process close together, sexual dimor- 
phism absent. Pereopods 2-7 with moderate 
setation as figured, sexual dimorphism for 
clasping slight with male pereopod 4 carpus 
2.6X as long as wide female 2.9X. 

Male pleopod 1, protopod about 0.6 X 
length of exopod, with 4 retinacula. Exopod 
about 1.6X as long as wide; lateral margin 



concave, distal margin with 6—8 long plu- 
mose setae. Pleopod 2, protopod with 3 me- 
sial setae. Exopod, proximal segment with 
about 5 lateral setae, distal segment with 
about 15 long plumose setae along margin. 
Endopod with distinct basal apophysis, en- 
dopod tip twisted in appearance, processes 
directed away from axis of endopod, can- 
nula tapering to a stylet, mesial process ta- 
pering and becoming digitiform, decurved 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Fourth pleopod exopods of related subterranean Caecidotea from the Springfield Plain: (a) C sim- 
ulator, Baxter Springs, Cherokee Co., Kansas, (b) C. simulator, Lasterling's well, Washington Co., Arkansas, 
(c) Caecidotea stiladactyla, small seeps 4.0 m. S. Boxley, Newton Co., Arkansas, (d) C. steevesi, Gittin Down 
Mountain Cave, Adair Co., Oklahoma, (e) C. steevesi, Carrico Cave, Dade Co., Missouri, (f) C. steevesi. War 
Eagle Cave, Madison Co., Arkansas. 



VOLUME 112, NUMBER 1 



179 



mesiad, caudal process heavily sclerotized, 
thickened laterally and mesially. Pleopod 3 
exopod, proximal segment about 9.7 X 
length of distal segment, with about 6 plu- 
mose setae on distal margin. Pleopod 4 with 
up to 18 proximolateral setae, two false su- 
tures present. Pleopod 5 with 2 sutures. 
Uropods of male about 2X length of pleo- 
telson, equal to length of pleotelson in fe- 
male. 

Etymology. — The noun "simulator", 
from the Latin meaning imitator or pretend- 
er, indicates the close resemblance of C. 
simulator to C. steevesL The vernacular 
name suggested for this species is the 
Springfield Plain groundwater isopod. 

Habitat and range. — Caecidotea simu- 
lator is known from the type-locality in 
southeastern Kansas and one locality in the 
adjacent comer of northwestern Arkansas. 
From its vermiform, eyeless, unpigmented 
appearance C. simulator is clearly an in- 
habitant of subterranean waters. Caecidotea 
simulator has not been found in caves and 
presumably lives in the saturated soil inter- 
stices that supply groundwater to the two 
sites from which it has been taken, a well 
and a seep spring. 

Relationships. — Caecidotea simulator 
and C. steevesi are very similar morpholog- 
ically. The gnathopods of both species are 
nearly identical. The apex of the first ple- 
opod exopod has elongate plumose setae in 
both species, a characteristic also shared 
with all species of the Hobbsi Group (Lew- 
is, 1982). In C simulator these setae occur 
all the way across the distal margin of the 
pleopod and are about 7-8 in number. In C 
steevesi the setae are located only on the 
mesial half of the distal margin and number 
4-5 (Fig. 2b). 

Due to torsion of the second pleopod en- 
dopod the taxonomically important tip pro- 
cesses are difficult to interpret and compare 
in C simulator and C. steevesi, a charac- 
teristic shared with C macropropoda and 
C. stiladactyla. The critical features are not 
easily seen without applying a coverslip, 
and the positions of the tip processes are 



quite distorted under the pressure of the 
glass (as the endopod twists). The endopod 
tips illustrated in Fig. 2 are the result of 
numerous attempts to gain the same per- 
spective for both C simulator and C. steev- 
esi. The endopods appear to be fundamen- 
tally similar, although side by side compar- 
ison (Fig. 2e-g) illustrates minor differenc- 
es of questionable importance. For 
example, the mesial process in C simulator 
tapers to a narrow, cylindrical digitiform 
process, while in C steevesi it is wider and 
obliquely truncate. 

The best way to separate C simulator 
from C. steevesi is by the structure of the 
fourth pleopod exopod (Fig. 3a— b, d-f), 
which has two false sutures in C. simulator 
(and C macropropoda, Lewis, 1982, Fig. 
3h), but only a single sigmoid suture in C. 
steevesi (and C stiladactyla. Fig. 3c). 

Caecidotea macropropoda Chase & Blair, 
1937 

Material examined. — Arkansas: Carroll 
County, White River at Beaver Down, G. 
C. Kephart, Jul 1978, ASS, 5$ $.— Wash- 
ington County, spring 2.2 miles north 
Dutch Mills, L. Hubricht, 21 May 1942, 
6S S , 5 $ $ ; seep, 1.5 miles north Winslow, 
L. Hubricht, 22 May 1940, 31c?(5, 7? ?. 

Remarks. — Lewis (1982) synonymized 
C ozarkana with C. macropropoda and re- 
described the species. Dearolf (1953) re- 
ported this species from two sites in Arkan- 
sas, but the validity of the records was un- 
known (Lewis, 1982). The records provided 
here substantiate those of Dearolf for the 
occurrence of C. macropropoda in Arkan- 
sas. The species is endemic to the Spring- 
field Plain, where it is found in cave 
streams, springs, and seeps. 

Acknowledgments 

The late Dr. Thomas E. Bowman provid- 
ed the loan of type-specimens of Caecido- 
tea steevesi from the collections of the 
Smithsonian Institution, read the manu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



script, and offered suggestions for its im- 
provement. 

Literature Cited 

Chase, H. D.. & A. P. Blair. 1937. Two new blind 
isopods from northeastern Oklahoma. — Ameri- 
can Midland Naturalist 18:220-224. 

Creaser, E. P. 1931. A new blind isopod of the genus 
Caecidotea, from a Missouri cave. — Occasional 
Papers of the Museum of Zoology, University 
of Michigan 222: 1-7. 

Dearolf, K. 1953. The invertebrates of 75 caves in the 
United States. — Pennsylvania Academy of Sci- 
ence 27:225-241. 



Fleming, L. E. 1972. Four new species of troglobitic 
asellids (Crustacea: Isopoda) from the United 
States. — Proceedings of the Biological Society 
of Washington 84(57):489-500. 

Lewis, J. J. 1982. A diagnosis of the Hobbsi Group, 
with descriptions of Caecidotea teresae, new 
species, and C. macropropoda Chase and Blair 
(Crustacea: Isopoda: Asellidae). — Proceedings 
of the Biological Society Washington 95:338- 
346. 

Mac kin, J. G., & L. Hubricht. 1940. Descriptions of 
seven new species of Caecidotea (Isopoda, 
Asellidae) from the central United States. — 
Transactions of the American Microscopical So- 
ciety 59:383-397. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):181-188. 1999. 

Collocherides brychius, a new species (Copepoda: Siphonostomatoida: 
Asterocheridae) from a deep-water hydrothermal site in the 

northeastern Pacific 

Arthur G. Humes 

Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 

02543, U.S.A. 

Abstract. — Collocherides brychius is described from a depth of 2253 m at 
a hydrothermal site on the Juan de Fuca Ridge in the northeastern Pacific. The 
species is close to C astroboae Stock 1971, but may be distinguished from 
that species by its relatively small size (length of female 0.57 mm) and by the 
inner terminal seta on the caudal ramus of the female being approximately as 
long as the caudal ramus, instead of more than 3 times its length. Although its 
three congeners are associated with ophiuroid echinoderms in shallow water, 
the new species was recovered free in microfaunal samples in the deep sea. 



The copepod genus Collocherides Stock 
1971 (Siphonostomatoida: Asterocheridae), 
contains three species. Collocherides astro- 
boae Stock 1971, lives in the stomach of 
the basket star Astroboa nuda (Lyman) at 
Eilat in the Gulf of Aqaba and at the Dahlak 
Archipelago (depth about 50 cm) in the Red 
Sea (Stock 1971). It has also been recov- 
ered from the stomach of Astroboa alba- 
trossi Doderlein in Indonesia (precise lo- 
cality unknown but probably Java Sea) 
(Stock 1971). Large numbers of C. astro- 
boae were found on A. nuda in 18 m at 
Nosy Be, northwestern Madagascar (Humes 
1973). Collocherides singularis Humes 
1986, lives with Astroboa nuda (depth 5 m) 
at Poelau Gomumu, Moluccas, 01°50'00"S, 
127°30'54"E (Humes 1986). Collocherides 
bleptus Humes 1993, occurs intertidally on 
the ophiuroid Macrophiothrix sp. at Nosy 
Be, northwestern Madagascar (Humes 
1993). 

Sixty-seven species of copepods have 
been recorded in deep water from hydro- 
thermal vents and cold seeps in the world's 
oceans (Humes & Segonzac 1998). These 
species consist for the most part of siphon- 
ostomatoids, with fewer calanoids, miso- 



phrioids, cyclopoids, poecilostomatoids, 
and harpacticoids. In this paper a new spe- 
cies of the siphonostomatoid genus Col- 
locherides is described from a deep-water 
hydrothermal site in the northwestern Pa- 
cific. With the addition of this new species, 
a new erebonasterid poecilostomatoid re- 
ported by Martinez Arbizu (1999), and a 
new aegisthid harpacticoid described by 
Conroy-Dalton & Huys (1999), the number 
of copepods known from deep-sea hydro- 
thermal vents and cold seeps rises to 70. 

Materials and Methods 

The copepods were collected at a low 
temperature vent, where the highest reading 
was 52° Celcius. The vent (Marker M) was 
situated on new lava flows, and is believed 
to be relatively young. (Vents at older ma- 
ture lavas are believed to support more het- 
erogeneous faunas, with more species, than 
young vents (Milligan & Tunnicliffe 
1994).) The diving submersible Alvin used 
its arm or "claw" to gather tube worms 
(Vestimentifera) and associated fauna and 
deposit them in a biobox mounted on a bas- 
ket. The sample was preserved for later 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



sorting, during which copepods were recov- 
ered (Tsurumi, pers. comm.). 

The copepods, which had been preserved 
in ethanol, were cleared and dissections 
made in lactic acid, using the wooden slide 
method described by Humes & Gooding 
(1964). All figures were drawn with the aid 
of a camera lucida. The letter after the ex- 
planation of each figure refers to the scale 
at which it was drawn. 

Siphonostomatoida Thorell, 1859 

Asterocheridae Giesbrecht, 1899 

Collocherides Stock, 1971 

Collocherides brychius, new species 

Figs. 1-3 

Type material. — 9 ? $, 3 S S, in 2253 
m, Juan de Fuca Ridge, Segment Cleft, 
North Field, Vent Marker M, northeastern 
Pacific, 44°58.97'N, 130°12.35'W, 28 Aug 
1990. Holotype 9 (USNM 243645), allo- 
type S (USNM 243646), and 6 paratypes 
(5 $ ?, 1 (?) (USNM 243647) deposited in 
the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
Remaining specimens (dissected) in the col- 
lection of the author. 

Female. — Body slender (Fig. la), not 
flexed between prosome and urosome (Fig. 
lb). Average length (not including setae on 
caudal rami) 0.57 mm (0.55-0.58 mm) and 
average width 0.18 nam (0.17-0.19 nam), 
based on 9 specimens. Dorso ventral thick- 
ness (at level of slight protuberance be- 
tween maxillipeds and first pair of legs) 127 
|xm. Epimera of metasomal somites round- 
ed (Fig. lb). Ratio of length to width of 
prosome 2.03:1. Ratio of length of prosome 
to that of urosome 1.31:1. 

Urosome with 5 somites. Somite bearing 
leg 5 (Fig. Ic) 47 X 83 jjim. Genital double- 
somite in dorsal view 81 ixm long and 81 
|jLm wide at widest part. In lateral view (Fig. 
Id), this double-somite with dorsal and ven- 
tral sides slightly rounded. Genital areas lo- 
cated laterally in anterior half of double- 
somite (Fig. Ic, d). Each genital area with 
small seta and minute spine (Fig. Id). Three 



postgenital somites from anterior to poste- 
rior 36 X 53, 31 X 40, and 23 X 36 fjim. 
Genital double-somite and 3 postgenital so- 
mites with pairs of small posterolateral 
spinelike processes or spinules (Fig. Id, e), 
those on anal somite prominent. 

Caudal ramus (Fig. le) moderately elon- 
gate, unomamented, length including ter- 
minal pointed process 44 iJim, length with- 
out process 35 ixm, width 16 iJim. Ratio of 
length (without process) to width 2.19:1, ra- 
tio including process 2.75:1. With 6 smooth 
setae, 1 long and 5 short, longest seta 52 
|jLm long, much longer than other setae and 
slightly swollen proximally. Long seta 
slightly longer than ramus, ratio 1.18:1. 

Body surface with few sensilla on uro- 
some, otherwise unomamented. 

Egg sac (Fig. Id), seen on only 1 female, 
large, oval, 179 X 99 jxm. 

Rostral area (Fig. If) triangular, not pro- 
truding. Antennule (Fig. 2a) 20-segmented, 
187 |jLm long, with aesthetasc 60 jJim long 
on segment 18. Formula for armature: 1, 2, 
2, 2, 2, 2, 2, 2, 6, 2, 2, 2, 2, 2, 2, 2, 2, 2+ 
1 aesthetasc, 2, and 9. All setae smooth. 
Antenna (Fig. 2b) 99 ixm long, including 
terminal spine 22 |xm. First segment (coxa) 
short and unarmed. Second segment (basis) 
with 1 minute seta (exopod). Third segment 
long and unarmed. Short fourth segment 
bearing 2 short setae and 1 long terminal 
spine, its truncated tip with few extremely 
minute setules. 

Oral cone (Fig. If) short, oval in ventral 
view, prominent in lateral view (Fig. lb). 

Mandible (Fig. 2c) with slender 2-jointed 
palp 60 |jLm long. Gnathobase (Fig. 2d) 50 
jjim long., with several blunt terminal teeth. 
Maxillule (Fig. 2e) with 1 seta on slender 
outer lobe and 4 setae on stout inner lobe. 
Maxilla (Fig. 2f) 2-segmented, first segment 
unarmed, second segment bearing recurved 
claw with truncated minutely spinulose tip. 
Maxilliped (Fig. 2g) 5-segmented. First 
segment with inner distal seta. Second seg- 
ment elongate (52 jjim long) with 1 seta on 
inner margin. Third and fourth segments 
short with 1 seta. Fifth segment elongate 



VOLUME 112, NUMBER 1 



183 




Fig. 1. Collocherides brychius, new species. Female, a, body, dorsal (scale A); b, body, lateral (A); c, 
urosome, dorsal (B); d, urosome, lateral (B); e, anal somite and caudal ramus, ventral (C); f, cephalosome, 
ventral (B). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Collocherides brychius, new species. Female, a, antennule, ventral (scale C); b, antenna, dorsal (C); 
c, mandible, ventral (C); d, gnathobase of mandible, flat view (D); e, maxillule, anterior (C); f, maxilla, posterior 
(C); g, maxilliped, posterior (C); h, leg 1 and intercoxal plate (E); i, leg 2 and intercoxal plate, posterior (E); j, 
leg 3 and intercoxal plate, anterior (E); k, leg 4 and intercoxal plate, anterior (E); 1, leg 5, ventral (C). 



VOLUME 112, NUMBER 1 



185 




Fig. 3. Collocherides brychius, new species. Male, a, body, dorsal (scale A); b, urosome, dorsal (B); c, 
antennule, ventral (C); d. leg 5, ventral (C); e, somites 1-3 of urosome, showing legs 5 and 6, ventral (E). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



with 1 distal seta. Claw (26 jjiin long) with 
truncated tip bearing minute spinules. 

Ventral area between maxillipeds and 
first pair of legs (Fig. If) slightly protuber- 
ant (Fig. lb). 

Legs 1-4 (Fig. 2h-k) biramous with 3- 
segmented rami. Formula for spines (Ro- 
man numerals) and setae (Arabic numerals) 
as follows: 



coxa 0-0 basis 1-0 expI-1; 



enpO-1; 
P2 coxa 0-0 basis 1-0 expI-1; 



expI-1; 
enpO-1; 



P3 coxa 0-0 basis 1-0 expI-1; 



enpO-1; 
P4 coxa 0-0 basis 1-' 



-0 expI-1; 
enp 0- 1 



l-l; 11,2,2 

0-2; 1.2,3 

I-l; 111,1,4 

0-2; 1,2,3 

I-l; 111,1,3 

0-2; 1,1,3 

I-l; 111,1,3 

0-2, 1,1,2 



Inner coxal seta absent in all 4 legs. Out- 
er margins of exopod segments of all 4 legs 
with very small spinules. 

Leg 5 (Fig. 21) 2-segmented. First seg- 
ment with outer seta and produced medially 
as broad triangular flap, pointed at tip, and 
having inner and outer setules. Second seg- 
ment 31 X 14 jjim, ratio 2.2:1, bearing 4 
setae and ornamented along inner margin 
with few setules. 

Leg 6 represented by seta and small spi- 
niform process on genital area (Fig. Id) 

Color of living specimens unknown. 

Male. — Body (Fig. 3a) elongate, slender, 
and, as in female, not flexed. Length (not 
including setae on caudal rami) 0.50 mm 
(0.50-0.51 mm) and greatest width 0.15 
mm (0.15-0.16 mm), based on 3 speci- 
mens. Greatest dorsoventral thickness at 
level of small ventral protuberance 96 ixm. 
Ratio of length to width of prosome 1.98: 
1 . Ratio of length of prosome to that of uro- 
some 1.36:1. 

Urosome (Fig. 3b) with 6 somites. So- 
mite bearing leg 5 32 X 70 |jim. Genital 
somite 60 X 83 ixm, with rounded lateral 
margins. Four postgenital somites from an- 
terior to posterior 34 X 49, 32 X 39, 26 X 
32, and 21 X 31 fxm. 

Caudal ramus similar to that of female. 



37 X 15 |jLm including terminal process. In- 
ner terminal seta 143 jxm long, relatively 
much longer than in female, 3.86 times lon- 
ger than ramus. 

Rostral area like that of female. Anten- 
nule (Fig. 3c) 18-segmented, geniculate. 
Aesthetasc on segment 17 59 jxm long. Ar- 
mature: 1, 2, 2, 2, 2, 2, 2, 2, 6, 2, 2, 2, 2, 
2, 2, 1, 1 + aesthetasc, and 9. Antenna as 
in female. 

Oral cone, mandible, maxillule, maxilla, 
maxilliped, and legs 1-4 like those of fe- 
male. 

Leg 5 (Fig. 3d) with second segment 20 
X 12 fxm, ratio 1.67:1, bearing 3 slender 
outer setae and 2 broad, hyaline, inner se- 
tae, 33 |xm, with blunt tips. 

Leg 6 (Fig. 3e) usual posteroventral flap 
on genital somite bearing 2 setae. 

Color of living specimens unknown. 

Etymology. — The specific name bry- 
chius, from Greek brychios, meaning from 
the depths of the sea, refers to the depth at 
which specimens of the new species was 
found. 

Remarks. — The female of Collocherides 
brychius may be differentiated from its 
three congeners as follows: 

From C. astroboae Stock 1971, in which 
the long terminal seta on the caudal ramus 
is approximately 3.6 times longer than the 
ramus, and the body length is 0.62 mm 
(measurements based on an average of five 
specimens from Astroboa nuda from the 
Dahlak Archipelago (Zoologisch Museum 
Amsterdam, cat. no. 101.090). 

From C. singularis Humes, 1986, in 
which the caudal ramus lacks a terminal 
process, the long terminal seta on the cau- 
dal ramus is approximately 1.8 times longer 
than the ramus, the second segment of the 
endopod of legs 1-3 has one inner seta, and 
the average body length is 0.64 mm. 

From C bleptus Humes, 1993, in which 
the ratio of the innermost terminal seta on 
the caudal ramus to the length of the ramus 
itself is approximately 2.51:1, the genital 
double-somite is 120 X 80 jjim, distinctly 
longer than wide, the average body length 



VOLUME 112, NUMBER 1 



187 



is 0.77 mm, and the ventral surfaces of the 
genital and postgenital somites have nu- 
merous small scalelike spines. 

In C brychius sexual dimorphism occurs 
in the length of the inner terminal seta on 
the caudal ramus, which in the female is 42 
|jLm, but in the male is 143 ixm and less 
swollen proximally than in the female. 

The four species of Collocherides may 
be further distinguished by the following 
key. 

Key to females of the genus Collocherides 

1. Genital double-somite distinctly longer 
than wide, ratio 1.4:1; body length 0.77 

mm (0.75-0.78 mm) C. bleptus 

Genital double-somite quadrate or near- 
ly so, ratio approximately 1:1; body 
length not exceeding 0.67 mm 2 

2. Long inner terminal seta on caudal ra- 
mus 3.6 times longer than ramus 

C astroboae 

Longest seta on caudal ramus 1.18 times 
(or less) longer than ramus 3 

3. Body length 0.57 mm (0.55-0.58 mm); 
legs 1-3 with second segment of endo- 

pod having 2 inner setae C brychius 

Body length 0.64 mm (0.63-0.65 mm); 
legs 1-3 with second segment of endo- 
pod with 1 inner seta C. singularis 

The ecological and host relationships 
within the genus Collocherides are poorly 
known. Collocherides astroboae, C singu- 
laris, and C. bleptus are associated with 
shallow-water ophiuroids, while C. bry- 
chius lives in deep— water hydrothermal 
sites and was recovered free in meiofaunal 
samples. However, much more information 
is needed to understand these relationships. 
The possibility that there may have been 
ophiuroids at the deep-sea vent site cannot 
be excluded. Ophiuroids, known from abys- 
sal depths (e.g., Lauermann and Kaufman 
1998), have been reported from deep-sea 
hydrothermal sites (Hessler and Smithey 
1983; Grassle 1986; Tunnicliffe 1991, 
1998; Segonzac 1992; Sibuet & Olu 1998). 
A new genus and species of ophiuroid, 
Ophioctenella acies, was described from 



the Mid- Atlantic Ridge by Tyler et al. 
(1995). However, no ophiuroids are yet 
known from Juan de Fuca vents, where the 
C brychius was found (Tunnicliffe, pers. 
comm.). 

Acknowledgments 

I thank Dr. Verena Tunnicliffe and Maia 
Tsurumi, Department of Biology, Univer- 
sity of Victoria, British Columbia, for send- 
ing the specimens of C. brychius to me for 
study. I also thank Dirk Platvoet, Zoolo- 
gisch Museum, Amsterdam, for arranging 
the loan of Stock's specimens of Astroboa 
nuda from Dahlak Archipelago. 

Literature Cited 

Conroy-Dalton, S., & R. Huys. 1999. A new genus of 
Aegisthidae (Copepoda: Harpacticoida) from 
hydrothermal vents on the Galapagos Rift. — 
Journal of Crustacean Biology (in press). 

Hessler, R. R., & W. M. Smithey, Jr. 1983. The distri- 
bution and community structure of megafauma 
at the Galapagos Rift hydrothermal vents. — In 
Hydrothermal processes and seafloor spreading 
centers. R A. Rona, K. Bostrom, L. Laubier, & 
K. L. Smith, Jr., eds., NATO conference series 
4:735-770. Plenum Press, New York. 

Humes, A. G. 1973. Cyclopoid copepods associated 
with the ophiuroid Astroboa nuda in Madagas- 
car. — Beaufortia 21:25-35. 

. 1986. Two new species of Copepoda associ- 
ated with the basket slar Astroboa nuda (Ophiu- 
roidea) in the Moluccas. — Zoologica Scripta 15: 
323-332. 

. 1993. Collocherides bleptus n. sp. (Copepoda: 

Siphonostomatoida) associated with an intertid- 
al ophiuroid in Madagascar. — Bijdragen tot de 
Dierkunde 63:121-127. 

, & R. U. Gooding. 1964. A method for study- 
ing the external anatomy of copepods. — Crus- 
taceana 6:238-240. 

, & M. Segonzac. 1998. Copepoda from deep- 



sea hydrothermal sites and cold seeps: descrip- 
tion of a new species of Aphotopontius from the 
East Pacific Rise and general distribution. — Ca- 
hiers de Biologic Marine 39:51-62. 

Lauermann, L. M. L., & R. S. Kaufman. 1998. Deep- 
sea epibenthic echinoderms and a temporally 
varying food supply: results from a one year 
time series in the N. E. Pacific. — Deep-Sea Re- 
search 1145:817-843. 

Martinez Arbizu, P. 1999. New Erebonasteridae (Co- 
pepoda) from Vilkitzky Strait in the Arctic and 



U^ 



II 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



from a Pacific hydrothermal vent site (northern 
Fiji Basin). — Journal of Crustacean Biology 19: 
93-105. 

Milligan, B. N., & V. Tunnicliffe. 1994. Vent and non- 
vent faunas of the Cleft segment, Juan de Fuca 
Ridge, and their relations to lava age. — Journal 
of Geophysical Research 99:4777-4786. 

Segonzac, M. 1992. Les peuplements associes a 
I'hydrothermalisme oceanique du Snake Pit 
(dorsale medio-atlantique; 23°N, 3480 m): com- 
position et microdistribution de la megafau- 
ne. — Comptes Rendus de I'Academie des Sci- 
ences Paris 314 (serie 3):593-600. 

Sibuet, M., & K. Olu. 1998. Biogeography, biodiver- 
sity and fluid dependence of deep-sea cold-seep 
communities at active and passive margins. — 
Deep-Sea Research II 45:517-567. 



Stock, J. H. 1971. Collocherides astroboae n. gen., n. 
sp., a siphonostome cyclopoid copepod living 
in the stomach of basket stars. — Bijdragen tot 
de Dierkunde 41:19-22. 

Tunnicliffe, V. 1991. The biology of hydrothermal 
vents: ecology and evolution. — Oceanography 
and Marine Biology 29:319-407. 

, A. G. Mc Arthur, & D. McHugh. 1998. A bio- 
geographical perspective of the deep-sea hydro- 
thermal vent fauna. — Advances in Marine Bi- 
ology 34:353-442. 

Tyler, R A., G. J. L. Paterson, M. Sibuet, A. Guille, B. 
J. Murton, & M. Segonzac. 1995. A new genus 
of ophiuroid (Echinodermata: Ophiuroidea) 
from hydrothermal mounds along the Mid-At- 
lantic Ridge. — Journal of the Marine Biological 
Association of the United Eangdom 75:977- 
986. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):189-198. 1999. 

Cymbasoma californiense, a new monstrilloid (Crustacea: Copepoda: 
Monstrilloida) from Baja California, Mexico 

E. Suarez-Morales and R. Palomares-Garcia 

(ES-M) El Colegio de la Frontera Sur (ECOSUR)-Unidad Chetumal. A.P. 424. Chetumal, 

Quintana Roo. 77000. Mexico; (RP-G) Centro Interdisciplinario de Ciencias Marinas 

(CICIMAR). COFAA. A.P 592. La Paz, B.C.S., Mexico 

Abstract. — Cymbasoma californiense is described from three adult females 
collected during three cruises carried out in an area adjacent to Bahia Mag- 
dalena, a large coastal system on the southwest coast of the Baja California 
Peninsula, and in the southern Gulf of California. The new species is very 
closely related to at least three other Cymbasoma (C. longispinosum (Bourne) 
and its allies), but can be distinguished by a combination of features including: 
the presence of cuticular striations on the forehead, cephalothorax, and genital 
somite; relatively long antennules; the structure of the fifth legs, and the relative 
length of the ovigerous spines. Cymbasoma californiense is the first species of 
this genus recorded in the Calif omian region and the Eastern Tropical Pacific. 



Monstrilloids are protelean parasitic co- 
pepods which are free-living only as first 
nauplii and adults (Raibaut & Trilles 1993). 
The first nauplius is the infective stage, but 
later stages complete their development as 
endoparasites of benthic organisms such as 
polychaetes and prosobranch molluscs. 
Adults are free-swimming, exclusively re- 
productive, and have no feeding append- 
ages (Isaac 1975; Davis 1984). 

The number of described species has 
been estimated as about 90 (Grygier 
1994a), and they are grouped into three rec- 
ognized genera (Monstrilla, Monstrillopsis, 
and Cymbasoma). The generic nomencla- 
ture of the group is still unclear, and efforts 
have been made to solve this problem, par- 
ticularly with regard to the genus Cymba- 
soma (Grygier 1994a, 1997). This genus 
contains now about 57 described species, 
with about half of them being described un- 
der the invalid genus name Thaumaleus 
(Razouls 1996). 

There are several (six) records of mon- 
strilloid copepods in the Northeastern Pa- 
cific region (Park 1967, Grygier 1995, Ra- 
zouls 1996). However, only three species of 



monstrilloid copepods have been recorded 
in lower latitudes, such as the California re- 
gion or the Eastern Tropical Pacific. All 
three regional records are of the genus 
Monstrilla. Monstrilla capitellicola Hart- 
man, 1961 was recorded at La Jolla Canyon 
as a parasite of a polychaete of the genus 
Capitella (Hartman 1961), and was de- 
scribed upon V stage copepodids. Monstril- 
la spinosa Park, 1967, originally described 
from the Strait of Georgia, was recorded by 
Suarez-Morales & Vasquez-Yeomans 
(1996) off Bahia de Todos Santos, Baja 
California, and M. gibbosa Suarez-Morales 
& Palomares-Garcia, 1995 was collected on 
the southeastern coast of the Baja Califor- 
nia Peninsula (Suarez-Morales & Paloma- 
res-Garcia 1995). 

As part of a survey of the main coastal 
systems along the southern portion of the 
Baja California Peninsula, plankton sam- 
ples were collected at Bahia de Magdalena, 
a large bay located on the western coast. 
Our taxonomic analysis of the copepods in 
the samples revealed the presence of several 
monstrilloids which turned out to belong to 
an undescribed species. The description of 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



the new species is presented herein follow- 
ing the standards set by Grygier & Ohtsuka 
(1995) for descriptions of monstrilloid co- 
pepods. 

Materials and Methods 

Plankton samples were collected during 
the BAMA8611, BAMA9710, and CER- 
RALVO9609 cruises. The first two were 
carried out in Bahia Magdalena (BAMA), 
a large coastal system on the southern por- 
tion of the western coast of the Baja Cali- 
fornia Peninsula, Mexico. The other cruise 
took place in the southern portion of the 
Gulf of California, along the southeastern 
coast of the Baja California. One female of 
the same previously undescribed species of 
monstrilloid copepod as captured during 
each of the three cruises: one at station K2 
(BAMA9710), on 27 October 1997, another 
at station R2 (BAMA9611), on 5 February 
1996, and the third was collected at station 
SIIG49 (CERRALVO9609), 20 September 
1996 (Fig. 1). Samples were collected dur- 
ing surface hauls with a standard plankton 
net (0.333 mm mesh-size). Zooplankton 
was fixed with a buffered formalin solution. 
Monstrilloid copepods were sorted and 
transferred to 70% ethanol. Observations 
were made under a Zeiss microscope and 
drawings were made with the aid of a cam- 
era lucida. Standard terminology for cope- 
pod morphology (Huys & Boxshall 1991) 
and for monstrilloid antennular armature 
(Grygier & Ohtsuka 1995) was followed. 

Order Monstrilloida 

Family Monstrillidae Dana, 1849 

Cymbasoma Thompson, 1888 

Cymbasoma califomiense, new species, 

Suarez-Morales 

Material examined. — Holotype, 2.1 mm, 
adult female, undissected, ethanol-pre- 
served. Sta. K2 (coordinates in decimal no- 
tation: 24.69°N, 112.07°W) BAMA9710 
cruise, Bahia Magdalena, South Baja Cali- 
fornia, Mexico, 27 Oct 1997. Vial deposited 
in the Collection of Crustacea, National 



Museum of Natural History, Smithsonian 
Institution, Washington D.C., under number 
USNM-261422. Paratype adult female, par- 
tially dissected, permanent slide, mounted 
in CMC. Sta. SIIG49 (24.27°N, 109.85°W) 
CERRALVO9609 cruise, Isla Cerralvo, 
Southern Gulf of California, Mexico, 20 
Sep 1996. Deposited in the NMNH, 
USNM-261422. Paratype aduh female, to- 
tal length: 2.03 mm, ethanol-preserved. Sta- 
tion R2 (24.41°N, 111.65°W), BAMA8611 
cruise, Bahia Magdalena, Baja California 
Sur, Mexico, 5 Feb 1986. Vial deposited in 
the Zooplankton Collection of El Colegio 
de la Frontera Sur Unidad Chetumal, Mex- 
ico, under number ECO-CH-Z-00369. 

Type locality. — Bahia Magdalena, south- 
em portion of the eastern coast of the Baja 
California Peninsula, Mexico. Water col- 
umn. 

Etymology. — The species is named after 
the Californian region in which this species 
was collected. 

Description of female. — Average body 
length of 3 individuals, 2.1 mm measured 
from forehead to posterior margin of anal 
somite. Cephalothorax (incorporating first 
pedigerous somite) accounting for almost 
65% of total body length (Fig. 2A, B). 
Ventral margin of anteriormost portion of 
cephalothorax slightly curved. Oral papilla 
as in Fig. 2A, lying midventrally 0.21 of 
way back along cephalothorax. Forehead 
with strong cuticular striations mainly be- 
tween antennule bases, and, more dorsally, 
a circular, spiral pattern with two anterior 
and two posterior whorls, present in all the 
individuals. Nauplius eye present, weakly 
developed, ocelli slightly pigmented with 
rounded shape (Fig. 3E). Pair of low 
bumps on ventral side of cephalothorax be- 
tween antennule bases and oral papilla, 
surrounded by circles with wrinkles ar- 
ranged in spiral-shaped pattern. Cuticular 
ornamentation also on lateral posterior 
margin of cephalothorax: clear vertical 
striations (Fig. 2B). Antennules four-seg- 
mented, armed with 0, I; 1,V; 2, I; 10, VII 
setae (Arabic numbers) and spines (Roman 



VOLUME 112, NUMBER 1 



191 



25.0 



24.9- 



24.8- 



24.7 



24.6 



24.5 



24.4 



24.3 



32' 



MEXICO 



PACIFIC OCEAN 




1 r 



go ^^40 ^^2° 110° 108° 



PACIFIC OCEAN 



112.2 112.1 112.0 111.9 111.8 111.7 111.6 111.5 



24.3- 



24.2 



24.1 



Cerralvo Island 




24.0 

110.10 110.00 109.90 109.80 

Fig. 1. Study area showing the sites in which specimens of the new species were collected. 



192 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. 
spine. 



Cymbasoma califorriiense n. sp.. holotype female. A. habitus, lateral. B. habitus lateral. C. ovigerous 



numbers) (Fig. 3B, C). Terminal two 
spines asymmetrical, forming chela-like 
structure. Last segment with one small 
aesthetasc on distal margin, large aesthe- 
tasc (4aes) on proximal outer margin. Most 
spines and setae of general pattern de- 



scribed by Grygier & Ohtsuka (1995) pre- 
sent (Fig. 3C). Only setae b5 and b6 miss- 
ing on last segment of right and left anten- 
nules, setae bl-4 unbranched. Ratio of 
lengths of antennule segments: 1 1.3 : 23.1 : 
14.3:51.3 = 100. Antennule slightly Ion- 



VOLUME 112, NUMBER 1 



193 




0.12 mm 



0.12 mm 



Fig. 3. Cymbasoma californiense n. sp., holotype female. A. genital complex and fifth legs, ventral. B right 
antennule, dorsal. C. left antennule, dorsal, with armature following the nomenclature of Grygier & Ohtsuka 
(1995). D. urosome and furcal rami, dorsal. E. forehead, dorsal. 



194 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ger than 17.5% of total body length, and 
25.5% of cephalothorax length. 

Incorporated first pedigerous somite and 
three free succeeding pedigerous somites 
(latter accounting for 22% of total length in 
dorsal view) each bearing a pair of bira- 
mous swimming legs (Fig. 4A-D). Endop- 
odites and exopodites of legs 1-4 triarticu- 
lated. Legs 1—4 decreasing in size posteri- 
orly. Armature of swimming legs as: 



legl 



basis endopodite exopodite 
1-0 0-l;0-l; 1,2,2 I-1;0-1;1, 1,3 



legs2-4 1-0 0-l;0-l; 1,2,2 I-1;0-1;I,2,3 

Basis separated from coxa posteriorly by 
diagonal articulation, with lateral hair-like 
seta on legs 1-4; seta on leg 3 at least 1.5 
times longer and noticeably thicker than the 
others, and plumose, while others are sim- 
ple. Inner seta on first exopodite segment of 
legs 1-4 plumose, long, reaching to distal 
margin of endopodite. Inner margin of first 
endopodal segments and outer margin of 
second and third exopodal segments of legs 
1-4 with short hairs. Outer exopodal apical 
seta of legs 1—4 with row of setules along 
inner side, but bearing row of small, closely 
spaced denticles along outer side. Most se- 
tae biserially plumose. 

Fifth legs joined medially; each with 
rounded endopodal lobe. Exopodal lobe 2 
times longer than wide, with two long, 
equal plumose setae, and one short inner, 
plumose seta, about 28% as long as the oth- 
er two (Fig. 3A). 

Urosome consisting of fifth pedigerous 
somite, genital double somite, one free ab- 
dominal (anal) somite, and furcal rami, al- 
together representing 14.3% of total body 
length. Genital double-somite slightly lon- 
ger than one-third total length of urosome 
(35%). Ratio of lengths of fifth pedigerous 
somite, genital double somite, and free ab- 
dominal somite: 37.5:35.5:26.6 = 100. 
Genital somite with strong transverse cutic- 
ular wrinkles on lateral margins and dorsal 
surface (Fig. 3D). Medial portion of genital 
double somite moderately swollen, bearing 



long, basally conjoined, ovigerous spines. 
Spines separate from level of posterior mar- 
gin of furcal rami. Spines slender at their 
base, swollen distally, in dorsal view, left 
one of holotype shorter than right one (Fig. 
4E). Spines about 1.4 times total body 
length (Fig. 2C). Caudal rami short, 1.2 
times longer than wide, moderately diver- 
gent, bearing three strong, terminal setae, as 
usual in genus. Middle seta slightly thicker 
than others (Fig. 3D). 

Male. — unknown. 

Remarks. — The new species from Baja 
California is assigned to the genus Cym- 
basoma by virtue of the presence of two 
urosomal somites in the female, with only 
one free somite between the genital double 
somite and the caudal rami (Isaac 1975). 

According to the most comprehensive 
key to the species of this genus (Isaac 
1975), the Calif omian specimens would be 
identified as females of C. longispinosum 
(Bourne, 1890). However, other species 
such as C. chelemense Suarez-Morales & 
Escamilla, 1997, recently described from 
the Gulf of Mexico, and C. morii Sekigu- 
chi, 1982, redescribed by Grygier (1994b) 
from Japan are closely related to and can 
also be confused with C. longispinosum. 
The female specimens of C. californiense 
share several features with C. longispinos- 
um, C. morri, and C. chelemense, such as 
the relatively long cephalothorax, the gen- 
eral structure of the fifth legs, with a round- 
ed endopodal lobe, the position of the oral 
papilla, the antennule/cephalothorax length 
ratio, the dorsal striations on the genital 
double segment, the swollen tips and the 
asymmetry of the ovigerous spines, and rel- 
atively long ovigerous spines conjoined at 
the base. However, the new species differ 
from these other species in several key 
structures generally regarded as important 
in monstrilloid taxonomy, such as the mor- 
phological details of the fifth legs, the struc- 
ture of the genital complex, and the body 
proportions. 

In the females of C. californiense the 
cephalothorax represents 65% of the total 



VOLUME 112, NUMBER 1 



195 





E 

E 

r^ 
o 



Fig. 4. Cymbasoma califomiense n. sp., holotype female. A. right leg 1, posterior. B. left leg 2, posterior. 
C. right leg 3, posterior. D. right leg 4, posterior. E. ovigerous spines, detail of distal portion. 



196 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



body length; the figure for C. chelemense is 
68%, while in C. longispinosum it is less 
than 65%, and in C. morii the reported 
range (Grygier 1994b) is 66-73%. In the 
new species the oral papilla is relatively 
much smaller and shows a different aspect 
with respect to those of C. chelemense and 
C. morii, but is similar to that of C. longis- 
pinosum. The strong cuticular protuberanc- 
es on the forehead of the new species are 
shared only by C. chelemense, but are dif- 
ferent; in the latter, the striations show a 
more complex pattern. The forehead is only 
slightly rugose in C. morii (see Grygier 
1994b). Cymbasoma calif omiense shows 
anteroventral knobs (nipples), which were 
described for C. morii, and C chelemense 
and are probably also present in C. longis- 
pinosum. However, in the new species these 
knobs are reduced, not protruding as far as 
those described for the other species. 

The fifth leg's endopodal lobe in Cym- 
basoma longispinosum has a clearly trian- 
gular-elongated shape (Giesbrecht 1892; 
Sars 1921; Isaac 1975), while the same 
structure is completely rounded in the new 
species, as it is in both C. morii (Grygier 
1994b) and C. chelemense (Suarez-Morales 
& Escamilla 1997). However, this structure 
is relatively smaller in C. morii than in the 
other two species. 

The fifth legs inner seta in C. calif or- 
niense is relatively short in comparison 
with those of the other species. It is about 
28% as long as the other two setae (3.5 
times shorter), while this figure is 62.5% in 
C. chelemense (1.6 times shorter), 40% (2.5 
times shorter) in C. morii, and 66% (1.5 
times shorter) in C. longispinosum. 

The armature of the antennules is similar 
in the four species. The new species shows 
the same antennular armament pattern as 
described in C. morii and C. chelemense, 
with only slight differences in the size and 
position of some of the smallest setae. The 
largest aesthetasc on the distal segment is 
relatively shorter in C. califomiense than it 
is in C. chelemense and in C. morii. Pro- 
portions of antennular segments are similar 



in these species, with the terminal segment 
representing more than 50% of the total an- 
tennular length. However, the antennules in 
the new species are relatively longer than 
in the other species compared herein, rep- 
resenting 25.5% of the cephalothorax 
length versus 21-22% in the other three 
species considered herein. 

In C. califomiense and C. chelemense, 
the genital double somite's posterior margin 
is about 40% wider than the anterior margin 
of the succeeding free (anal) somite. From 
Giesbrecht's (1892) illustrations, in C. lon- 
gispinosum the proportional value is less 
than half (16%) that measured for the new 
species; in C. morii the value is around 
30%, and the posterior margin of the genital 
double somite is slender and somewhat 
convex. This margin is slender in the other 
species and clearly irregular in the new spe- 
cies. 

The proportional length of the ovigerous 
spines differs among the four species. In C. 
longispinosum and C. chelemense the 
spines are only 14% longer than the body 
(Giesbrecht 1892; Sars 1921; Suarez-Mo- 
rales & Escamilla 1997). In C. morii they 
are much longer, about twice as long as the 
body, whereas in the new species they are 
42% longer than the body. All four species 
have a long common base for the two ovig- 
erous spines. 

Cymbasoma califomiense shows cuticu- 
lar wrinkles along the lateral margins of the 
fifth pedigerous and anal somites, and on 
the posterior lateral margins of the cepha- 
lothorax. These features are absent in the 
other species and could be considered as a 
key character to recognize this species. The 
total body length recorded for C. califor- 
niense (2.2 mm) is slightly under the range 
reported for C. longispinosum and C. che- 
lemense (2.3 to 3.16 mm) (Isaac 1975, Sua- 
rez-Morales & Escamilla 1997), but is 
slightly over the average length of C. morii 
(2.18 mm) (Grygier 1994b). 

There are several species of Cymbasoma 
with a long, slender cephalothorax: C. mo- 
rii (Sekiguchi, 1982), C. longispinosum 



VOLUME 112, NUMBER 1 



197 



Table 1 . — Comparison of features present in females of two groups of species of Cymbasoma with a relatively 
long cephalothorax. 



Species 


C. gigas 


C. morii 


C. chelemense 


C. longispinosum 


C californiense 


C. bowman i 


C reticulatum 


Lobes on fifth 

leg 
Inner lob 


2 


2 


2 


2 


2 


1 


1 


ovate 


rounded 


rounded 


subtriangular 


rounded 








Nipples 


7 


present 


present 


present 


reduced 


absent 


absent 


Ovig. spines 


7 


joined at 


joined at 


joined at base 


joined at 


separated 


separated at 






base 


base 




base 


at base 


base 


O.S. longer 


7 


2 times 


14% longer 


14% longer 


43% longer 


shorter 


shorter 


than body 
















Striation on 


7 


present 


present 


not described 


present 


absent 


absent 


gen. som. 
















Striation on 


7 


absent 


absent 


not described 


present 


absent 


absent 


anal som. 
















Striation on 


7 


present 


present 


not described 


present 


present 


absent 


forehead 




(light) 


(strong) 




(strong) 


(light) 





(Bourne, 1890), C. chelemense (Suarez- 
Morales & Escamilla), 1997, Cymbasoma 
gigas (A. Scott, 1909), C. reticulatum 
(Giesbrecht, 1892), and C. bowmani (Sua- 
rez-Morales & Gasca), 1998. As suggested 
by Suarez-Morales & Gasca (1998), these 
Cymbasoma species can be divided into 
two groups (see Table 1). In the first group, 
with a bilobed fifth leg, nipple-like ventral 
cephalic protuberances, striated genital so- 
mite, and basally fused genital spines 
which are longer (1.5-2 times) than the 
body, C. morii, C. longispinosum, C. che- 
lemensis, probably C. gigas, and now C 
californiense, could be included. The sec- 
ond group shares a single-lobed fifth leg, 
the absence of nipple-like ventral protu- 
berances, and basally separated genital 
spines that are shorter than the body (C 
reticulatum, C. bowmani). 

The lack of comparative biogeographical 
information on monstrilloids is related to 
several factors: the scarcity of the group in 
plankton collections, the unsolved taxono- 
my of the group, the uncertainty in recog- 
nizing the male and female of the same spe- 
cies from plankton samples, and the irreg- 
ular putative distributional patterns 
throughout the group (Isaac 1975). How- 
ever, it is relevant to mention the known 
geographical ranges of the four species 



compared herein. The Mediterranean, Great 
Britain, Mindanao, Portugal, Black Sea, 
Gulf of Suez, Vietnam, and the Arabian 
Gulf for C. longispinosum (Isaac 1975); the 
wide distribution reported for this species 
could be the result of overlooking closely 
related species. This has been suggested by 
Grygier & Ohtsuka (1995) for Monstrilla 
helgolandica Glaus (1863). Cymbasoma 
morii has been recorded from Japan, sup- 
posedly Vietnam, and probably India (Gry- 
gier 1994b); C chelemense from the south- 
em Gulf of Mexico (Suarez-Morales & Es- 
camilla 1997), and C. californiense from 
Baja California. Presumably, the geograph- 
ical isolation of the new species with re- 
spect to the others would favour the con- 
ception of a different taxon. 

This new species of Cymbasoma repre- 
sents the second record of the genus in the 
Northeastern Pacific region; the only pre- 
vious record was of C. rigidum Thompson, 
1888, near the Kodiak Island, Alaska 
(Threlkeld 1977). It is also the first record 
of the genus in the California region. 

Acknowledgments 

One specimen was obtained during the 
operation of the project "Potencial de la 
Isla Cerralvo, Baja California Sur como 



198 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



area de reserva pesquera". Comments of 
three anonymous reviewers improved an 
earlier version of this manuscript. 

Literature Cited 

Davis, C. C. 1984. Planktonic Copepoda (including 
Monstrilloida). Pp. 67-91. in K. A. Steidinger 
& L. M. Walter, eds.. Marine plankton life cycle 
strategies. C. R. C. Press. Florida. 

Giesbrecht, W. 1892. Systematik und Faunistik der pe- 
lagischen Copepoden des Golfes von Neapel 
und der angrenzenden Meeres-Abschnitte. — 
Fauna Flora Golf. Neapel 19:1-831 + Atlas 
(pis. 1-54). 

Grygier, M. J. 1994a [dated 1993]. Identity of Thau- 
matoessa (=Thaumaleus) typica Kr0yer, the 
first described monstrilloid copepod. — Sarsia 
78:235-242. 

. 1994b. Nomenclature, redescription, and new 

record from Okinawa of Cymbasoma morii Sek- 
iguchi, 1982 (Monstrilloida). — Hydrobiologia 
292/293:23-29. 

. 1995. Annotated chronological bibliography 

of Monstrilloida (Crustacea: Copepoda). — Gal- 
axea 12:1-82. 

. 1997. Monstrilla Dana, 1849 and Thaumaleus 

Kr0yer, 1849 (Crustacea, Copepoda): con- 
served. — Bulletin of Zoological Nomenclature 
54:131-132. 

, & S. Ohtsuka. 1995. SEM observations of the 



nauplius of Monstrilla hamatapex, new species, 
from Japan and an example of upgraded de- 
scriptive standards for monstrilloid copepods. — 
Journal of Crustacean Biology 15:703-719. 

Hartman, O. 1961. A new monstrillid copepod para- 
sitic in capitellid polychaetes in southern Cali- 
fornia. — Zoologischer Anzeiger 167:325-334. 

Huys, R., & G. A. Boxshall. 1991. Copepod evolu- 
tion. — Ray Society, London, England 159:1- 
468. 

Isaac, M. J. 1975. Copepoda. Sub-order: Monstrilloi- 



da. — Fiches d' Identification du Zooplankton 
144/145: 10 pp. 

Park, T. S. 1967. Two unreported species and one new 
species of Monstrilla (Copepoda: Monstrilloida) 
from the Strait of Georgia. — Transactions of the 
American Microscopical Society 86:144-152. 

Raibaut, A., & J. R Trilles. 1993. The sexuality of 
parasitic crustaceans. — Advances in Parasitolo- 
gy 32:367-444. 

Razouls, C. 1996. Diversite et repartition geographique 
chez les copepodes pelagiques.2. — Platycopioi- 
da, Misophrioida, Mormonilloida, Cyclopoida, 
Poecilostomatoida, Siphonostomatoida, Harpac- 
ticoida, Monstrilloida. — Annales de I'lnstitut 
Oceanographique 72:1-149. 

Sars, G. O. 1921. An account of the Crustacea of Nor- 
way with short descriptions and figures of all 
the species. VIII. Copepoda Monstrilloida & 
Notodelphyoida. Bergen Museum, Bergen, 91 
pp, pis. I-XXXVII. 

Sekiguchi, H. 1982. Monstrilloid copepods from Ago 
Bay, central Japan. — Proceedings of the Japa- 
nese Society of Systematic Zoology 22:24-34. 

Suarez-Morales, E., & J. B. Escamilla. 1997. An un- 
described monstrilloid copepod (Copepoda: 
Monstrilloida) from the northern Yucatan Pen- 
insula, Mexico. — Bulletin of Marine Science 
61:539-547. 

, & R. Gasca. 1998. Cymbasoma bowmani sp. 

no v., a new monstrilloid (Copepoda: Monstril- 
loida) from a Caribbean reef, with notes on spe- 
cies variation. — Journal of Marine Systems 15: 
433-439. 

, & R. Palomares-Garcia. 1995. A new species 

of Monstrilla (Copepoda: Monstrilloida) from a 
coastal system of the Baja California Peninsula, 
Mexico. — Journal of Plankton Research 17: 
745-752. 

, & R. Vasquez-Yeomans. 1996. On Monstrilla 



spinosa Park, 1967 (Copepoda, Monstrilloida) 
in the eastern Pacific. — Crustaceana 69:288- 
294. 
Threlkeld, S. T 1977. The distribution of five species 
of Monstrillidae (Copepoda, Monstrilloida) near 
Kodiak Island, Alaska. — Crustaceana 32:225- 
228. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):199-209. 1999. 

Early development of the deep-sea ampharetid 
(Polychaeta: Ampharetidae) Decemunciger apalea Zottoli 

Robert Zottoli 
Department of Biology, Fitchburg State College, Fitchburg, Massachusetts 01420, U.S.A. 

Abstract. — Early development of the opportunistic, deep-sea ampharetid 
polychaete Decemunciger apalea Zottoli is described from specimens removed 
from wood panels placed on the deep-sea floor by Turner (1973). Larvae with 
less than 8 setigers were not found. Larval spatulate notosetae and neuropodial 
uncini on segments 3—6 formed during early development are eventually lost 
by the 14-setiger stage making segment 7 the first uncinigerous thoracic seg- 
ment in juveniles and adults. Early development of this species is generally 
similar to that described for other shallow and deep-sea ampharetids. 



The deep-sea ampharetid polychaete De- 
cemunciger apalea Zottoli inhabits galleries 
in wood originally excavated by bivalve 
molluscs belonging to the genera Xylopha- 
ga and Xyloredo (Family Pholadidae, sub- 
family Xylophagainae) (Turner 1973, 1977; 
Zottoli 1982). The wood was placed by R. 
D. Turner, using the submersible DSRV Al- 
vin, at four experimental bottom stations in 
the North Atlantic to "test the hypothesis 
that wood is an important source of nutri- 
ents and contributes to diversity in the 
deep-sea" (Turner 1973, 1977). This study 
is based on a detailed examination of 393 
complete juvenile and adult specimens from 
wood panel N31 (DOS2) (Fig. 1). Early de- 
velopment and further role of this species 
in deep-sea ecosystems is discussed. 

Materials and Methods 

Three experimental islands, each with 12, 
one foot spruce cubes, were placed by 
Turner (1977), for a period of five years at 
the following locations: Deep Ocean Sta- 
tion 1 (DOS-1), 39°46'N, 70°41'W, 110 
miles south of Woods Hole, Mass., in 1830 
m; Deep Ocean Station 2 (DOS-2), 
38°18.4'N, 69°35.6'W, 190 miles southeast 
of Woods Hole, Mass., in 3506 m; Tongue 



of the Ocean, Bahama Islands (TOTO Tow- 
er 3), 24°53.2'N, 77°40.2'W, in 2066 m. 

Each experimental island is surrounded 
by wood panels (24" X 5" X 1") which are 
removed and replaced each time the islands 
are visited. Each panel is enclosed in a 
mesh bag when it is retrieved, to prevent 
loss of specimens. The mesh bags and their 
contents are then transferred to retrieval 
boxes carried on the DSRV Alvin basket. 
The contents of the bags may be preserved 
at the time the boxes are closed for return 
to the surface by puncturing plastic bags 
containing glutaraldehyde, previously 
placed in the retrieval boxes. In certain cas- 
es, the panels were preserved immediately 
upon reaching the surface. Wood panel N31 
(DOS 2) was submerged by Alvin on dive 
601, 5 Sep 1975, and removed on dive 777, 
3 Aug 1977 harboring 393 whole amphar- 
etids. Refer to Turner (1973, 1977) for a 
more detailed description of the methods 
used to place and retrieve the panels from 
experimental islands. 

Systematics 

Ampharetid polychaetes are for the most 
part widest anteriorly, tapering gradually to- 
wards the posterior end. The prostomium, a 
pre-segmental structure, is situated imme- 



200 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



c 
0) 



0-5.95 (Adults) 


^^^^^^^^^^^^^^^■89 


126 


178 


.45 (Juveniles) 




)-1. 25 (Larvae) 




^BH 


_ ( 1 1 1 1 j j 1 1 



20 40 60 80 100 120 

Number of individuals 



140 



160 



180 



200 



Fig. I. Frequency histogram for length from a collection of 393 Decemunciger apalea from wood panel 
N31 (DOS2). 



diately in front of the peristomium. Seg- 
ments 1 and 2 that are fused together follow 
behind the peristomium. Refer to Zottoli 
(1974) for additional information on the 
formation of these structures. The segmen- 
tal numbering system used in this paper is 
that of Malmgren (1865-1866) and Fauvel 
(1927) who recognized two segments in 
front of the paleal segment (segment 3). 
The thorax begins on segment 4. Segment 
3 in some species has one dorsolateral bun- 
dle of paleal setae, or notopod with a bun- 
dle of winged capillary setae, on each side. 
Segment 3 is achaetous in adult Decemun- 
ciger apalea; however, small notopods with 
larval, hooded, flared (spatulate) notosetae 
or winged capillary setae are present in ear- 
ly stages. Adult worms have 13 thoracic se- 
tigers, the last 10 of which are unciniger- 
ous, and 14 abdominal uncinigerous seg- 
ments (Zottoli 1982). In addition, 4 pairs of 
smooth branchiae are present on the dorsal 
surface of segments 3-5. Refer to Zottoli 
(1982) for a more complete description of 
adult characteristics. 



Larval and juvenile stages of D. apalea 
in this study are named according to the 
total number of segments bearing spatulate 
and/or capillary notosetae (setigers). Post- 
setigerous segments are those that follow 
these setigers and bear uncinigerous pin- 
nules (neuropodia with uncini). For adults 
the word setiger refers to any segment with 
setae of either the capillary or uncinate 
type. The number of thoracic segments with 
both capillary notosetae and neuropodial 
uncini is a diagnostic feature of juveniles 
and adults but not larvae since larvae gain 
and lose setae during development, and the 
thorax is not fully formed. 

Early Development 

The eight female D. apalea with mature 
eggs in their body cavities ranged from 2.5 
to 5.6 mm in length and from 0.5 to 0.7 
mm in maximum width, whereas the 13 
males with mature sperm ranged from 2.6 
to 5.95 mm in length and from 0.42 to 0.7 
mm in maximum width. Therefore, speci- 



VOLUME 112, NUMBER 1 



201 



mens with the full complement of adult se- 
tigers and ranging from 1.3 to 2.45 mm in 
length are considered juveniles. Worms less 
than 1.3 mm in length and lacking the full 
complement of adult setigers are considered 
larvae. On this basis, 89 adults, 126 juve- 
niles and 178 larvae were identified from 
the 393 whole specimens removed from 
Panel N31 (DOS 2) (Fig. 1). Sexually ma- 
ture specimens of this species previously 
studied (Zottoli 1982) ranged from 3.6 to 
6.3 mm in length and from 0.54 to 0.9 mm 
in maximum width. 

8-Setiger Stage. — The 8-setiger stage 
(Fig. 2) is named for its eight segments 
bearing notosetae. Larvae with less than 8 
setigers were not found. In the 8-setiger 
stage (Fig. 2), larval ciliary bands were not 
evident; a short, medial, ventrally ciliated 
tentacle protrudes from the mouth. A mus- 
cular ventral bulb is visible internally, just 
below the pharynx and above the lower lip. 
Pharynx, esophagus, stomach and intestine 
are visible through the transparent body 
wall. The digestive tract is filled with par- 
ticulate matter as well as occasional Xylo- 
phaga larvae, suggesting that they feed in 
the manner described for Hobsonia florida 
by Zottoli (1974): "larvae begin feeding on 
microscopic plant and animal material at 
about the two-setiger stage by forcing ma- 
terial from the mud surface into the diges- 
tive tract through the action of the ventrally 
located buccal mass and by cilia on the up- 
per lip." 

Prostomial and pygidial pigment spots 
are lacking. Fused segments 1 and 2 are 
situated immediately behind the prostomi- 
um. Segments 3-6 (setigers 1-4) bear 2 
small dorsolateral notopods each with 1 
spatulate (hooded, flared) seta (about 1 1 jjim 
long, 6 jjim maximum width) (Fig. 3A) and 
1-2 smooth, winged, bilimbate capillary se- 
tae (about 25 |xm long, 2 ijim wide basally) 
(Fig. 3A). The number of smooth, winged, 
bilimbate capillary setae per notopod in- 
creases as the larva grows from the 8- to 
the 14-setiger stage (Table 1). 

One uncinus (about 6 jJim long) is em- 




Fig. 2. Decemunciger apalea 8-setiger larva. Lat- 
eral view of entire worm, 0.6 mm long. 



bedded in the epidermis, below the noto- 
pod, on each side of segments 3-6 (setigers 
1-4). Each of the first 4 pairs of uncini have 
about 12 teeth above a basal prow (Fig. 



202 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Decemunciger apalea 8-setiger larva. A. Single spatulate seta and 2 smooth, winged, bilimbate 
capillary setae from the left notopod of the second setiger; B. Uncinus, frontal view, from the same setiger as 
above; C. Uncinus, frontal view, from the left uncinigerous pinnule of the 5th setiger. 



VOLUME 112, NUMBER 1 



203 



Table 1. — Number of smooth, winged, bilimbate capillary setae per notopodium on "setigerous" segments 
of the 8- to 14-setiger stages of Decemunciger apalea. Note the loss of 1 setiger between the 14b- and 13a- 
setiger stages. 





Setiger 










Setiger stage 










Segment 


8 


n 


12 


14a 


14b 


13a 


13b 


13c 


13d 


3 


1 


1 


1 


2 


1 


1 














4 


2 


2 


3 


3 


2 


3 


3 


3 


6 


4 


5 


3 


2 


3 


5 


3 


3 


3 


4 


6 


4 


6 


4 


2 


4 


5 


3 


3 


5 


6 


8 


6 


7 


5 


3 


3 


5 


4 


6 


6 


7 


8 


8 


8 


6 


2 


2 


3 


3 


6 


6 


6 


8 


8 


9 


7 


2 


2 


3 


3 


5 


6 


6 


9 


8 


10 


8 


1 


2 


2 


3 


4 


6 


6 


9 


9 


11 


9 




1 


2 


4 


4 


5 


6 


8 


9 


12 


10 




1 


2 


3 


4 


5 


5 


8 


9 


13 


11 




1 


2 


3 


4 


4 


5 


8 


8 


14 


12 






2 


3 


4 


4 


5 


8 


6 


15 


13 








3 


3 


4 


5 


7 


6 


16 


14 








2 


2 


3 


3 


7 


6 



(1 pair br.), 11 = 1.0 X 0.19 mm (1 pair br.), 12 = 0.9 X 0.18 nmi (2 pair br.), 14a 
r br.), 14b = 1.56 X 0.30 mm (3 pair br.), 13a = 1.75 X 0.30 mm (4 pair br.), 13b 
;^ K. ^ 1 ^^ - ^ <n ^ n ^n tA «o;. k. ^ i ^^ ^ 5.00 X 0.80 mm (4 pair br.), 14b 



8 = 0.6 X 0.19 mm , 

1.4 X 0.28 mm (3 pair br.), 14b . , , 

2.10 X 0.75 mm (4 pair br.), 13c = 3.50 X 0.50 mm (4 pair br.), 13d 
1.56 X 0.30 mm (3 pair br.), br = branchiae. 



3B); they disappear in later stages making 
segment 7 (setiger 5) the first uncinigerous 
thoracic segment in juveniles and adults. 
Segments 7-10 (setigers 5-8) bear 2 dorso- 
lateral notopods each with one or two 
smooth, winged, bilimbate capillary noto- 
setae (about 25 ixm long and 2 jjim wide 
basally) (Fig. 2; Table 1). Uncini (about 8 
jjim long) from segments 7-15 (setigers 5- 
8 and postsetigers 1—5), are borne on ven- 
trolateral extensions of the body (uncini- 
gerous pinnules), one pair per segment (Fig. 
2). Uncini have about 14 teeth above a bas- 
al prow (Fig. 3C). The numbers of uncini 
per uncinigerous pinnule on all setigers, ex- 
cept those of segments 3-6 (setigers 1—4) 
gradually increase from the 8- to the 14- 
setiger stage (Table 2). 

There are 5 post-setigerous segments 
(segments 1 1-15) bearing uncinigerous pin- 
nules but lacking notopodia and notosetae. 
The pygidium lies immediately behind seg- 
ment 15. One pair of branchiae is present 
on the dorsoposterior edge of segment 3. 
There is a narrow mid-dorsal gap between 
the branchium of each side. 



Larvae at this stage were found in mem- 
branous tubes covered by fine particulate 
matter. The membranous lining is most 
likely mucus secreted by the thick, glan- 
dular pads on the ventral surface of most 
thoracic setigerous segments. 

11 -Setiger Stage. — The 11 -setiger stage 
(Fig. 4), named for its eleven segments 
bearing notosetae, is similar to the 8-setiger 
stage except for an increase in number of 
segments and setae (Table 1), an increase in 
length of the median tentacle, addition of 2 
lateral tentacles, loss of larval spatulate se- 
tae on setiger 4, and loss of larval uncini 
on setigers 1 and 2 (Table 2). 

12-Setiger Stage. — By the 12-setiger 
stage, named for its twelve segments bear- 
ing notosetae, the larval uncini on setigers 
3 and 4 (Table 2), and the larval spatulate 
setae on setigers 1-3 disappear. There is an 
increase in the number of smooth, winged, 
bilimbate capillary notosetae per notopo- 
dium (Table 1) and in the total number of 
"setigerous" and "post-setigerous" seg- 
ments (Table 2). There is an increase in the 
length of the median tentacle, and a second 



204 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Number of uncini per uncinigerous pinnule on "setigerous" and "post-setigerous" segments of 
the 8- to 14-setiger stages of Decemunciger apalea. Note the loss of 1 setiger between the 14b- and 13a-setiger 
stages. Stage lengths are listed in Table 1. 















Setiger stage 










Segment 


Setiger 


8 


11 


12 


14a 


14b 


13a 


13b 


13c 


13d 


3 


1 


1 


























4 


2 


1 





1 




















5 


3 


1 


1 


1 




















6 


4 


1 


2 


1 




















7 


5 


3 


5 


7 


14 


11 


15 


25 


29 


30 


8 


6 


2 


3 


6 


11 


11 


14 


24 


29 


28 


9 


7 


1 


2 


4 


9 


9 


11 


18 


27 


26 


10 


8 


1 




3 


8 


8 


13 


18 


25 


21 


11 


9 


1 




3 


8 


6 


10 


16 


23 


20 


12 


10 


1 




2 


6 


5 


8 


15 


23 


19 


13 


11 


1 




1 


5 


5 


8 


14 


21 


18 


14 


12 


1 




1 


4 


3 


7 


14 


18 


13 


15 


13 


1 




1 


3 


3 


6 


14 


17 


10 


16 


14 
"Post setiger 


1 
' (Abdomen) 


1 


2 


2 


5 


14 


17 


















17 


15 






1 


2 


2 


3 


5 


16 


10 


18 


16 










2 


3 


5 


10 


10 


19 


17 










2 


3 


5 


10 


9 


20 


18 










2 


3 


5 


8 


9 


21 


19 










1 


3 


5 


7 


9 


22 


20 










1 


2 


5 


7 


6 


23 


21 










1 


2 


5 


7 


7 


24 


22 










1 


1 


5 


7 


5 


25 


23 












1 


5 


5 


4 


26 


24 












1 


4 


5 


4 


27 


25 
















4 


4 


28 


26 
















2 


3 


29 


27 
















3 





pair of branchiae appears on the dorsolat- 
eral, posterior surface between segments 4 
and 5. Later, a third pair of branchiae arises 
from the dorso-lateral surface of segment 5. 

14-Setiger Stage. — By the 14-setiger 
stage, all larval uncini on segments 3-6 
have disappeared (Table 2, 14a, b). There is 
a general increase in the number of noto- 
setae (Table 1) and uncini (Table 2). A 
fourth pair of branchiae arises from the an- 
terior, dorsolateral surface of segment 6. 
Notosetae are eventually lost on segment 3, 
thus creating the 13-setiger stage described 
below (Table 1). 

13 -Setiger Stage. — The early 13-setiger 
stage (Fig. 5) (Tables 1-2, 13a) has the full 
complement of adult thoracic setigerous 



segments of which 10 are uncinigerous. 
Uncinigerous thoracic segments begin on 
segment 7. Changes that occur between 
this stage and adult worms (Tables 1-2, 
13c, d) include an increase in number of 
capillary notosetae per notopod, an in- 
crease in number of uncini per unciniger- 
ous pinnule, an increase in number of teeth 
per uncinus to about 20, and an increase in 
number of uncinigerous abdominal seg- 
ments to 12-14. 

Uncini are first formed in the dorsal por- 
tion of each uncinigerous pinnule. They 
range in length from 6-8 ixm in an 8-se- 
tiger stage to 6-13 jjim in a 12- setiger stage 
and from 12-18 jjim in a 13-setiger stage. 
As new, longer uncini are created, older. 



VOLUME 112, NUMBER 1 



205 




Fig. 4. Decemunciger apalea 11-setiger larva. Lateral view of entire worm. 1.0 mm long. 



and shorter ones are pushed ventrally and 1988). Few of the originally formed uncini 

eventually re-absorbed. This process is remain in late juvenile and adult stages, 

similar to that in A. galapagensis (Zottoli suggesting an overall rapid growth rate for 

1983) and E. nebulosa (Bhaud & Gremare D. apalea. 



206 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 5. Decemunciger apalea 1 3-setiger juvenile. Lateral view of entire worm, 1.75 nun long. 



VOLUME 112, NUMBER 1 



207 



Discussion 

Larval ciliated bands, although not found 
in D. apalea, disappear at about the 6-se- 
tiger stage in H. florida (Hartman) (Zottoli 
191 A), the 7-setiger stage in Alkmaria rom- 
ijni Horst (Cazaux 1982), and were not 
found by Zottoli (1983) in later stages of 
Amphisamytha galapagensis. 

Spatulate setae similar to those described 
in D. apalea are found in notopods of the 
first 3 setigers of larval A. galapagensis 
ZottoU (ZottoU 1983), H. florida (Hartman) 
(Zottoli 1974), and Schistocomus sovjeticus 
Annekova (Okuda 1947) and in the noto- 
pods of the first 4 setigers of larval Am- 
pharete acutifrons (Grube) (Clavier 1984) 
and Melinna palmata (Grehan et al. 1991). 
What appear to be hooded, flared notosetae 
can be observed on Nyholm's (1950) pho- 
tograph of a 3-setiger Melinna cristata Sars. 
Curiously, none were reported by Cazaux 
(1982) for A. romijni. Russell (1987) de- 
scribed hooded, flared setae in notopodia of 
segments 3-6 from the paedomorphic am- 
pharetid Paedampharete acutiseries Rus- 
sell. He suggested that "the spatulate setae 
of H. florida and A. galapagensis may rep- 
resent a type of "spatulate" setae distinctly 
different from that of S. sovjeticus, A. acu- 
tifrons and P. acutiseries.'' A detailed ex- 
amination of spatulate setae from larval A. 
galapagensis, D. apalea, and H. florida 
shows a similar 3 -dimensional form to that 
described by Russell (1987). Notosetae, 
which are generally similar in appearance 
to ampharetid spatulate setae, have been de- 
scribed, respectively, by the bracketed au- 
thors on the first 6, 8 and 11 setigers of 
Eupolymnia nebulosa (Bhaud 1988 and 
Bhaud & Gremare 1988), Thelepus setosus 
(Duchene 1983), and Loima medusa (Wil- 
son 1928). Hooded, flared, notosetae even- 
tually disappear, with the possible exception 
of P. acutiseries, in all of the species dis- 
cussed above. 

The initial formation of spatulate setae, 
capillary notosetae and uncini and their 
subsequent loss on anterior segments here. 



are generally similar to that described for 
the ampharetids A. galapagensis (Zottoli 
1983), M. palmata (Grehan et al. 1991), H. 
florida (ZottoU 1974), and the terebeUids E. 
nebulosa (Bhaud & Gremare 1988), Loima 
conchilega (Kessler 1963), L. medusa (Wil- 
son 1928), and Nicolea zostericola (Eckel- 
barger 1974). 

Larval and adult uncini are similar in D. 
apalea, in contrast to A. galapagensis (Zot- 
toU 1983) and H. florida (ZottoU 1974) 
where multi-toothed larval uncini are re- 
placed in most segments by uncini with a 
single row of teeth. 

Branchial formation in D. apalea is sim- 
ilar to that of the ampharetids A. galapa- 
gensis (Zottoli 1983), A. romjini (Cazaux 
1982), H. florida (ZottoU 1974), and pos- 
sibly P. acutiseries (Russell 1987). 

Tubes, similar to those of D. apalea, 
were formed by 3-setiger, 1.75-2 day old 
H. florida larvae (ZottoU 1974), by 4-setig- 
er M palmata larvae (Grehan et al. 1991) 
and by 8-setiger, 20 day old Alkmaria rom- 
ijni larvae (Cazaux 1982), shortly after they 
had abandoned the interior of the maternal 
tube. Larvae of the terebeUids, E. nebulosa 
(Bhaud & Gremare 1988), L. conchilega 
(Kessler 1963), A^. zostericola (Eckelbarger 
1974), and T. setosus (Duchene 1983), 
formed similar tubes, respectively, at the 6-, 
2-, 14- and 4-setiger stages. Tube formation 
at such an early stage suggests that D. apa- 
lea larvae are benthic, remaining in the 
same general area as their parents. This 
does not preclude the possibility of larval 
transport from place to place by bottom cur- 
rents. Once larvae form their own tube, 
they most likely remain permanently af- 
fixed, reaching new feeding areas by tube 
elongation as described by Fauchald & Ju- 
mars (1979). Ecklebarger (1974) reported 
that juvenile and adult N. zostericola, 
forced from their tubes, moved by repeat- 
edly folding the abdomen upon itself and 
then straightening out. These actions lifted 
worms off the bottom and into the water 
column, transporting them short distances. 
The author observed similar movements in 



208 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



juvenile and adult H. florida forced from 
their tubes. Because of the general similar- 
ity in body shape and structure of amphar- 
etids and terebellids, it is assumed that 
most, including D. apalea, could move 
short distances in the manner described 
above if displaced from their tubes. 

Panel N31 (DOS 2) was submerged for 
about 2 years from 5 Sep 1975 to 3 Aug 
1977. There are more larvae and juveniles 
than adults (Fig. 1). Assuming that larvae 
and juveniles of D. apalea remain in the 
area where they were bred. Fig. 1 most like- 
ly reflects the distribution of the species as 
a whole. This assumption is supported by 
the fact that worms live in attached tubes 
in all but the earliest phases of their life and 
that their body shape is not conducive to 
sustained active locomotion in a pelagic en- 
vironment. The presence of numerous lar- 
vae and adults with gametes in their body 
cavities suggests that breeding took place 
shortly before the time of retrieval. The 
small number of large adults suggests a Hfe 
span of one year. This is also supported by 
the presence of gravid adults, juveniles and 
larvae in a panel (N68, DOSl) retrieved af- 
ter approximately one year. If the life span 
of D. apalea were more than one year, one 
would expect a proportionately greater 
number of adults. 

Acknowledgments 

Thanks are due to Dr. J. Fred Grassle, 
Charlene D. Long, the late Dr. Meredith L. 
Jones and Dr. Ruth D. Turner for making 
specimens available. The study, conducted 
by Turner (1977), was supported by the Of- 
fice of Naval Research (ONR Contract No. 
14_76-C-1281, NR 104-687 to Harvard 
University). 

Literature Cited 

Bhaud, M. 1988. Change in setal pattern during early 
development of Eupolymnia nebulosa (Poly- 
chaeta: Terebellidae) grown in simulated natural 
conditions. — Journal of the Marine Biological 
Association of the United Kingdom 68:677- 
687. 



, & A. Gremare. 1988. Larval development of 

the terebellid polychaete Eupolymnia nebulosa 
(Montagu) in the Mediterranean Sea. — Zoolo- 
gica Scripta 17(4):347-356. 

Cazaux, C. 1982. Developpement larvaire de 
I'Ampharetidae Lagunaire Alkmaria romijni 
Horst 1919. — Cahiers de Biologic Marine 23: 
143-157. 

Clavier, J. 1984. Description du cycle biologique d' 
Ampharete acutifrons (Grube, 1860) (Annelide 
Polychete). — Comptes Rendus Academic des 
Sciences, Paris, Serie III 299(3):59-62. 

Duchene, J-C. 1983. Developpement larvaire et fixa- 
tion chez Thelepus setosus (Annelide Polyche- 
te) a Kerguelen, Province Subantarctique. — Vie 
et Millieu 33(2):65-77. 

Eckelbarger, K. J. 1974. Population biology and larval 
development of the terebellid polychaete Nico- 
lea zostericola. — Marine Biology 27:101-113. 

Fauchald, K., & R A. Jumars. 1979. The diet of 
worms: A study of polychaete feeding guilds. — 
Oceanography and Marine Biology Annual Re- 
view 17:193-284. 

Fauvel, P. 1927. Polychetes Sedentaires. — ^Faune de 
France 16:1-494. 

Grehan, A., C. Retiere, & B. Keegan. 1991. Larval 
development in the ampharetid Melinna pal- 
mata Grube (Polychaeta). — Ophelia Supple- 
ment 5:321-332. 

Kessler, Von M. 1963. Die entwicklung von Lanice 
conchilega (Pallas) mit besonderer beriicksich- 
tigung der lebensweise. — Helgolander Wissen- 
schaftliche Meersuntersuchungen 8(4):425— 
476. 

Malmgren, A. J. 1865-1866. Nordiska Hafs-Annula- 
ter. — Ofversigt Svenska Vetenskaps Akade- 
miens Forhandhngar 22:181-192; 355-410. 

Nyholm K-G. 1950. Contributions to the life history 
of the ampharetid Melinna cristata. — Zoologis- 
ka Bidrag fran Uppsala 29:79-91. 

Okuda, S. 1947. On an ampharetid worm, Schistoco- 
mus sovjeticus Annekova, with some notes on 
it's larval development. — ^Journal of the Faculty 
of Science Hokkaido Imperial University, Series 
6 9:321-329. 

Russell, D. E. 1987. Paedampharete acutiseries, a new 
genus and species of Ampharetidae (Polychae- 
ta) from the North Atlantic Hebble area, exhib- 
iting progenesis and broad intraspecific varia- 
tion. — Bulletin of the Biological Society of 
Washington 7:140-151. 

Turner, R. D. 1973. Wood-boring bivalves, opportu- 
nistic species in the deep sea. — Science 180: 
1377-1379. 

. 1977. Wood, molluscs, and deep-sea food 

chains. — Bulletin of the American Malacologi- 
cal Union 1977:13-19. 

Wilson, D. P. 1928. The post-larval development of 



VOLUME 112, NUMBER 1 



209 



Loimia medusa Sav. — Journal of the Marine Bi- 
ological Association of the United Kingdom 15: 
129-147. 

Zottoli, R. A. 1974. Reproduction and larval devel- 
opment of the ampharetid polychaete Amphic- 
teis floridus. — Transactions of the American 
Microscopical Society 93:78-89. 

. 1982. Two new genera of deep-sea polychaete 

worms of the family Ampharetidae and the role 



of one species in deep-sea ecosystems. — Pro- 
ceedings of the Biological Society of Washing- 
ton 95:48-57. 

— . 1983. Amphisamytha galapagensis, a new 
species of ampharetid polychaete from the vi- 
cinity of abyssal hydrothermal vents in the Gal- 
apagos Rift, and the role of this species in Rift 
ecosystems. — Proceedings of the Biological So- 
ciety of Washington 96:379-391. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):210-215. 1999. 

A new species of Spiochaetopterus (Chaetopteridae: Polychaeta) from 
a cold-seep site off Hatsushima in Sagami Bay, central Japan 

Eijiroh Nishi, Tomoyuki Miura, and Michel Bhaud 

(EN) Natural History Museum and Institute, Chiba, Aoba 955-2, Chuo, Chiba 260-8682, Japan; 

(TM) United Graduate School of Agricultural Science, Kagoshima University, 1-21-24, Korimoto, 

Kagoshima, 890 Japan; (MB) Observatoire Oceanologique de Banyuls, Universite P. & M. Curie, 

(Paris VI) Laboratoire Arago, B.P 44, F-66651 Banyuls-sur-mer, France 

Abstract. — Spiochaetopterus sagamiensis, a new species (Polychaeta: Chae- 
topteridae), is described from two specimens collected from a cold-seep site 
off Hatsushima in Sagami Bay, central Pacific side of Japan. In this species, 
the large chaetae on chaetiger 4 (A4) are distinctive in having a triangular 
profile of the head, a nearly straight ventral edge of the inflated distal part, and 
two dorsolateral grooves on the shaft. The species has trilobed notopodia on 
the first segment of the median region (Bl), bilobed ones on the second (B2), 
and entire ones on the third (B3) and on all succeeding segments. Uncinal 
plates of neuropodium B2 have more than 30 teeth; the upper toothed edge is 
smoothly curved. Anterior eyespots are absent. Spiochaetopterus sagamiensis 
occurs in deep water (800-1 100 m depth) and is the first member of this family 
to be recorded in a deep-sea chemosynthetic community. 



Chaetopterid polychaetes commonly oc- 
cur on mud and sandy mud bottoms from 
the intertidal to shallow shelf waters. In 
Japanese waters, two species of Spiochae- 
topterus are known: S. okudai Gitay, 1969 
and S. costarum (Claparede, 1870) (Okuda 
1935, Gitay 1969, Nishi & Aral 1996). 
Chaetopterids are poorly known from deep- 
sea bottoms: only one species, the type spe- 
cies S. typicus Sars, 1856, has been record- 
ed from shallow water to a depth of 2700 
meters off W. Greenland (Kirkegaard 
1960), at depths of 20 to 1865 meters in 
Arctic Seas and the Atlantic Ocean (Fauvel 
1914), and at depths of 35 to 2030 meters 
in the North Pacific and associated water- 
bodies (Uschakov 1955). Levin et al. 
(1991) listed unidentified chaetopterid poly- 
chaetes present in the fauna of seamounts 
from the eastern Pacific Ocean at depths of 
1058 to 3353 meters. Other records from 
deep-sea bottoms are worthy of re-exami- 
nation (e.g., Levenstein 1961; Hartman 



1971; Hartman & Fauchald 1971). This pa- 
per describes a new species of Spiochae- 
topterus found in a deep sea cold-seep off 
Hatsushima in Sagami Bay (Miura 1988). 
It is the first chaetopterid polychaete found 
in a chemosynthetic community (cf. Des- 
bruyeres & Segonzac 1997) and the third 
species in the genus described from Japan. 

Materials and Methods 

During dive 115 of the Deep-sea Re- 
search Vehicle (DSRV) Shinkai 2000 (ob- 
server: K. Egawa) in Sagami Bay, two 
chaetopterid specimens were collected with 
three Calyptogena shells at depths of about 
800-1100 m (Sugiura & Egawa 1985). The 
specimens were fixed in 10% sea- water for- 
malin on the mother ship and later trans- 
ferred to 70% ethanol. The specimens were 
sent to the second author for further study 
of the faunal structure of the Hatsushima 
cold-seep site (Miura 1988). 



VOLUME 112, NUMBER 1 



211 



Parts of the specimens, such as segment 
7 of the anterior region and large (cutting) 
chaetae were removed dehydrated through 
an alcohol series, air-dried, and observed 
with scanning electron microscopy (Hitachi 
S-800). Type series were deposited in the 
Natural History Museum and Institute, Chi- 
ba (CBM-ZW). For the description, the ter- 
minology of Crossland (1904) and Bhaud 
et al. (1994) is followed. In these papers, 
three body regions are defined as region A 
(anterior), region B (middle) and region C 
(posterior); in each region, segments are 
numbered from anterior to posterior: for in- 
stance, the fourth segment of the anterior 
region is termed segment A4 or simply A4. 

Family Chaetopteridae Malmgren, 1867 

Genus Spiochaetopterus Sars, 1856 

Spiochaetopterus sagamiensis, new species 

Figs. 1-2 

Spiochaetopterus sp. — Miura, 1988 (dive 
numbers 177 and 115 inadvertently 
switched in table 2). 

Material examined. — Off Hatsushima, 
Sagami Bay, DSRV Shinkai 2000 Dive 
115, 5 Jun 1984, 35°01'N, 139°12'E, 800- 
1100 m, collected with Calyptogena shells, 
holotype (CBM-ZW-701), incomplete, with 
a fragment of tube and paratype (CBM-ZW- 
702). 

Diagnosis. — Spiochaetopterus of small 
size, eyes absent, with one pair of long 
palps. Large (cutting) chaetae of segment 
A4 with inflated head. Distal end of A4 
chaetae triangular ventrally; shaft without 
ventral groove, with two dorso-lateral 
grooves. Segment A7 with brownish ventral 
gland; A8 and A9 with whitish ventral 
glands observed in alcohol preserved spec- 
imens. Notopodia trilobed in segment Bl, 
bilobed in B2, entire in B3 and following 
segments. Neuropodia entire in Bl, bilobed 
in all other middle segments. Uncini with 
over 30, maximum 40 teeth on uncinal 
plates. Tube unbranched, smooth, not an- 
nulated, with serrated opening. 

Description. — Both holotype and para- 



type incomplete, lacking posterior region 
(region C). Holotype 18 mm long excluding 
palps, about 1 mm wide. Paratype 24 mm 
long excluding palps, 0.8-1.0 mm wide. 
Regions A and B (anterior and middle) 
creamy white, dorsal side of region B light 
brown in alcohol apart from glandular re- 
gions. 

Region A narrow, 3.5 mm long for nine 
segments in holotype; 9 mm in paratype. 
Prostomium ovoid. Peristomium horseshoe- 
shaped, plump with prostomium on dorsal 
side (Fig. IB). Eyespots absent. Prostomial 
antennae absent (Fig. IB, C). Paired palps 
long, grooved, 10 mm long in holotype (14 
mm in paratype), arising from posterior 
border of peristomium, near posterolateral 
border of prostomium (Fig. lA, B, C, E). 
Dorsal groove ciliated, extending from base 
of palpi in Al to A9 (Fig. IB, E). Ventrum 
of region A with a long slender plastron 
(ventral glandular area) (Fig. lA, C, E), 
with longitudinal white stripes, separated 
into five portions (Fig. IC); portion I, lon- 
ger than others, overlaying from peristomi- 
um to segment A2, white in colour, portion 
II from A3 to A4, narrower at A4, cream- 
colored; portion III from A5 through ante- 
rior half of A6, dark brown; portion IV 
from posterior half of A6 through A7, 
white; portion V from A8 to A9, white. 
Segments A1-A3 short, parapodia with a 
single row of 25-40 lanceolate chaetae; A4 
elongate, with two large (cutting) chaetae, 
and more than 20 lanceolate chaetae; A5- 
A9 longer and wider than anterior three 
segments, with single row of 35-40 lance- 
olate chaetae (Fig. 2E-H). 

Large (cutting) A4 chaetae obliquely tri- 
angular, occurring singly on both side (Fig. 
2A— F); overhang of ventral edge of head 
distinct but weak. Distal part 300 jjim long, 
230 jxm wide, with blunt distal tip and near- 
ly horizontal ventral edge 200 jxm wide; 
shaft 1300 jjim long, 10-15 fim wide in 
middle portion, asymmetrical, nearly oval 
in cross section, without ventral groove; 
neck between head and shaft as wide as 
shaft (Fig. 2A, B, E). 



212 



PROCEEDmCS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 1. Spiochaetopterus sagamiensis new species, drawn from holotype (A-E, J-M) and paratype (F-H). 
For all drawings (A to M) anterior part of the body is on the right. A. Whole body, lateral view. B. Anterior 
portion enlarged. C. Anterior portion including A, segment Bl and anterior B2, ventral view. D. Same, lateral 
view. E. Regions A and B, dorso-lateral view. E A8 to B2, lateral view. G. B3 neuropodium, lateral view. H. 
A6 to B8 showing rami of neuropodia, ventrolateral view. J. Segments A9 and Bl, dorsal view. K. Segment 
B2, dorsal view. L. Segment B3, dorsal view. M. Segment B4, dorsal view. Abbreviations. — I to V, indication 



VOLUME 112, NUMBER 1 



213 




Fig. 2. Spiochaetopterus sagamiensis new species. Scanning electron micrographs of chaetae and tube of 
paratype. A-D. Distal part of A4 large (cutting) chaeta, ventral view (A, B) and lateral view (C, D); scales equal 
200 jxm (A) and 50 ixm (B-D). E & F. Distal part of A4 large (cutting) chaeta, ventrolateral (E) and anterior 
dorsolateral view (F); unlabeled arrows point to dorsolateral groove on shaft; scale equal 100 |jim (E) and 50 
(xm (F). G & H. Chaetae from region A, close-up view of thoracic setae; scales equal 75 jxm (G) and 50 ixm 
(H). I. Uncini of B2 neuropodium; scale equals 10 |jLm. J. Part of tube, lower arrow shows undulation, upper 
one serration; scale equals 750 |jLm. 

<— 



of ventral parts of region A, i.e., separated portion of ventral plastron and posterior peristomium; al to a9. 
segments of region A; bl, first segment of region B; b2, b3, second or third segment of region B; n, notopodium; 
ne, neuropodium; p. palp; pe, peristomium; po, prostomium. All scales equal 0.5 mm. 



214 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Head of A4 chaetae inflated, obliquely 
truncate; upper tip of head tilted toward 
longitudinal axis of body; consequently ex- 
ternal lateral oblique edge longer than in- 
ternal edge. Horizontal edge smoothly cir- 
cular without ventral sinus. Dorsolateral 
grooves on shaft (Fig. 2E, F). Ventral edge 
of oblique plane perpendicular to axis of 
shaft. Collar decreasing regularly in diam- 
eter. 

Remaining anterior segments (A5-A9) 
large, inflated, shorter and wider ventrally; 
parapodia with 35-40 lanceolate chaetae in 
single rows; lanceolate chaetae asymmetri- 
cal, longitudinaUy folded, sUghtly brown- 
ish, with smooth surface and serrated edges 
(Fig. 2G, H). 

Region B longer than anterior region, 
with eight elongate glandular segments 
(seven in paratype); paddle and cupule ab- 
sent. Bl shorter than following segments 
(Fig. IE). Middle parapodia biramous, 
along posterior margins of segments (Fig. 
IE, E, H). Notopodia with one to three 
lobes; neuropodia bilobed except in seg- 
ment Bl, with inflated uncinal plates. No- 
topodia of segment Bl foliaceous, trilobed, 
medial lobes resulting from a subdivision of 
inner lobe; outer lobe cirriform (Fig. IE, J). 
Notopodia of B2 bilobed, with single, entire 
inner lobes (Fig. IE, K); those of segments 
B3 and following entire (Fig. IL, M). Neu- 
ropodia of segment Bl entire, with only 
lower lobe developed (Fig. IF); other neu- 
ropodia bilobed, with rounded upper lobes 
and elongate lower lobes (Fig. IG, H). On 
neuropodia of B3, teeth on uncinal plate di- 
rected backward on anterior smaller dorsal 
lobe; that on larger ventral lobe directed 
forward. 

Uncini bluntly triangular, with single row 
of 35 to 40 minute teeth {n = 6, average 
38.2 ± 1.94) (Fig. 21). Segment B4 of par- 
atype with more than 40 uncini on outer 
lobe and more than 15 on inner one. Outer 
serrated edge of uncinal plate smoothly 
curved. 

Posterior body region (region C) lacking 
in types. 



Both examined specimens are mature, 
with segments of region B distended; the 
notopodia appear as distended, inflated 
transverse bags. 

Tube is fragile, slender, straight, smooth, 
and not annulated. Collar folds occur at 
subequal intervals along tube length. Tube 
wall is thin, consisting of several layers of 
secretion, and regularly undulated external- 
ly (Fig. 2J). 

Etymology. — The species epithet is de- 
rived from the type locality, Sagami Bay. 

Remarks. — Among the 1 1 described spe- 
cies in Spiochaetoptenis, 5 species lack 
eyespots: S. sagamiensis, S. typicus Sars, S. 
bergensis Gitay, S. okudai Gitay and 5. 
monroi Gitay. These five species may be 
distinguished mainly on the shape of the 
ventral edge of the obHque section of A4 
setae: straight without protuberances in S. 
sagamiensis, circular in S. typicus, with 
three protuberances in S. bergensis (Bhaud, 
1998). In these three cases the ventral edge 
is horizontal, perfectly perpendicular to the 
shaft Spiochaetoptenis okudai and S. mon- 
roi were reexamined. The ventral edge of 
A4 chaetae is obHque relative to the lon- 
gitudinal axis of the shaft in both species. 
S. okudai lacks the sinus and S. monroi has 
a marked sinus. Secondarily, Spiochaetop- 
terus okudai and S. monroi have bilobed 
notopodia in region B (Gitay 1969); how- 
ever, most notopodia are entire in S. saga- 
miensis. From observations by the third au- 
thor of Atlantic-Mediterranean and Pacific 
species, smoothly curved serrated edges of 
uncinal plates of 5. sagamiensis may be re- 
garded as a discriminatory character. 

In conclusion, four diagnostic character- 
istics may be retained for this new species: 
eyes are absent; the morphology of A4 
chaetae with a triangular profile of the head, 
a nearly straight ventral edge of the inflated 
distal part, and two dorsolateral, and not 
ventral, grooves on the shaft; the notopodia 
on B3 have only one lobe; and, outer ser- 
rated edges of uncini plates smoothly 
curved. 



VOLUME 112, NUMBER 1 



215 



Acknowledgments 

The first author wishes to thank the staff 
of the marine laboratory of Natural History 
Museum and Institute, Chiba; a facility yet 
to be completed. This study was partly sup- 
ported by the grant to young scientists from 
the Ministry of Education, Sports and Cul- 
ture of Japan (to E. N., No. 09740645). The 
third author would like to thank the co-au- 
thors for carrying out this work. For him, 
it is an element of a larger program to ver- 
ify the reality of species with a cosmopol- 
itan distribution, to define the role of large- 
scale larval dissemination and to identify 
the spatial limits of successful recruitment. 
His participation was funded by the French 
National Programme on Determinism of 
Recruitment. We are grateful to Dr. Kristian 
Fauchald, Smithsonian Institution, for his 
useful suggestions on the manuscript. 

Literature Cited 

Bhaud, M. 1998. The species of the genus Spiochae- 
topterus (Polychaeta, Chaetopteridae) in the At- 
lantic-Mediterranean biogeographic area. — Sar- 
sia (in press). 

, M. C. Lastra, & M. E. Petersen. 1994. Re- 
description of Spiochaetopterus solitarius (Rio- 
ja, 1917), with notes on tube structure and com- 
ments on the generic status (Polychaeta; Chae- 
topteridae). — Ophelia 40:115-133. 

Crossland, C. 1904. The polychaeta of the Maldive 
Archipelago from the collection made by Stan- 
ley Gardiner in 1899. — Proceedings of the Zoo- 
logical Society of London 1:270-289. 

Desbruyeres, D., & M. Segonzac. 1997. Handbook of 
deep-sea hydrothermal vent fauna. Editions 
IFREMER, Brest, France, 279 pp. 

Fauvel, P. 1914. Annelides Polychetes non pelagiques 
provenant des campagnes de I'Hirondelle et de 
la Princess- Alice (1885-1910). — Resultats des 



Campagnes Scientifiques du Prince de Monaco 
46:1-432. 

Gitay, A, 1969. A contribution to the revision of Spi- 
ochaetopterus (Chaetopteriidae, Polychaeta). — 
Sarsia 37:9-20. 

Hartman, O. 1971. Abyssal polychaetous annelids 
from the Mozambique basin off Southeast Af- 
rica, with a compendium of abyssal polychae- 
tous annelids from world-wide areas. — Journal 
of Fisheries Research Board of Canada 28: 
1407-1428. 

, & K. Fauchald. 1971. Deep-water benthic 

polychaetes annelids off New England to Ber- 
muda and other North Atlantic areas, part 2. — 
Allan Hancock Foundations, Monograph Series 
6:1-327. 

Kirkegaard, J. B. 1960. Polychaeta and Pogonophora 
from the deepest part of the Skagerrak. — Vi- 
denskabelige Meddelelser fra Dansk naturhis- 
torisk Forening Kopenhaven 123:211-226. 

Levenstein, R. Y. 1961. Polychaeta from the Bering 
Sea. — Trudy Instalagione Okeanologi Akade- 
miia Nauk SSSR 46:147-178. (In Russian) 

Levin, L. A., L. D. McCann, & C. L. Thomas. 1991. 
The ecology of polychaetes on deep-seamounts 
in the Eastern Pacific Ocean. — Ophelia, Supple- 
ment 5:467-476. 

Miura, T. 1988. Parasitic animals collected in a Calyp- 
togena-dormndmi community developing off 
Hatsushima, The Sagami Bay. — JAMSTEC 
Deepsea Research 9 1988:239-244. (In Japa- 
nese with English abstract) 

Nishi, E., & Y. Aral. 1996. Chaetopterid polychaetes 
from Okinawa. — Publications of the Seto Ma- 
rine Biological Laboratory, Kyoto University 
37:51-61. 

Okuda, S. 1935. Chaetopteridae from Japanese waters. 
Journal of the Faculty of Science, Hokkaido 
University, sen 6, Zoology 4:87-102. 

Sugiura, A., & K. Egawa, 1985. Research of bottom 
fishing grounds in Sagami Bay. — Technical Re- 
ports of Japan Marine Science and Technology 
Center 1985:67-72. 

Ushakov, P. V. 1955. Polychaeta of the far eastern seas 
of the U.S.S.R. — Izdatel'stvo Akademii Nauk 
SSSR, Moskva-Leningrad, Translated from 
Russian and published by Israel Program for 
Scientific Translations, Jerusalem 1965, 430pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(1):216-219. 1999. 

Two new subtribes, Stokesiinae and Pacourininae, of the Vernonieae 

(Asteraceae) 

Harold Robinson 

Department of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560, U.S.A. 

Abstract. — New subtribes Stokesiinae and Pacourininae are established in 
the Vernonieae (Asteraceae) for the Southeastern United States genus Stoke sia 
and the South American genus Pacourina 



Revisions of the Vernonieae by the au- 
thor have until recently involved mostly 
Western Hemisphere members (Robinson 
1996). In the absence of studies of Old 
World members of the tribe, among which 
relatives might have been discovered, the 
monotypic Western Hemisphere genera Sto- 
kesia UHer. and Pacourina Aubl. were left 
unplaced as to subtribe. More recent studies 
of Old World Vernonieae by the author 
have made it clear that Stoke sia and Pa- 
courina are most closely related to other 
Western Hemisphere genera, but that they 
are deserving of separate subtribal status. 
The new subtribes are described below. 

The genus Stokesia has long been noted 
for its mostly liguliform corollas, resem- 
bling those of the tribe Lactuceae, Fitchia 
Hook.f. of the Heliantheae, and some Mu- 
tisieae such as Hyaloseris Griseb. (Espinar 
1973). The tribal position has sometimes 
been questioned, but placement has usually 
been in its correct position in the tribe Ver- 
nonieae (Bentham & Hooker 1873, Hoff- 
mann 1890-1894). The genus contains a 
single species, and has been promoted as a 
possible crop plant useful as a source for 
epoxy resins (Gunn & White 1974). The 
plant is also widely cultivated as an orna- 
mental. A drawing has been provided by 
Gunn and White (1974), and a color pho- 
tograph can be seen in Rickett (1967). The 
pollen of the genus has a rather weak per- 
forated tectum and a unique lophate pattern 
with trisected colpi meeting at the poles 



(Figs. 1-4). The genus has a chromosome 
number that differs from most other Ver- 
nonieae, especially other genera from the 
Western Hemisphere. A first count of n = 
9 (Jones, 1968) has been corrected by a se- 
ries of six subsequent counts of n = 7 
(Jones 1974). Almost all other New World 
Vernonieae have n = 16 or 17. Most Old 
World Vernonieae have n = 9 or 10. 

Stokesiinae H.Rob., subtribus nov. 

Type: Stokesia UHer., Sertum Angl. 27. 
1789. 

Plantae herbaceae perennes ad 0.5 m al- 
tae, sparce pilosae, pilis longis simplicibus 
non septatis. Folia plerumque rosulata al- 
tema base anguste petioliformia in nodis 
vaginata. Inflorescentiae pauce capitatae 
laxe cymosae. Capitula pedunculata; brac- 
teae involucri 40-50 in seriebus 3-4, brac- 
teae exteriores in appendicibus longe foli- 
iformes margine spinosae, bracteae inter- 
iores angustiores in apicibus setiferae. Flo- 
res 60-70 homogami; corollae azurae vel 
albae plerumque late liguliformes in limbis 
5-lobatae, corollae centrales ca. 3 actino- 
morphae; thecae antherarum base rotunda- 
tae, cellulis endothecialibus distincte linea- 
tis, lineis in partibus longitudinalibus cet- 
erum variabiliter arcuatis; appendices api- 
cales antherarum breves glabrae in 
parietibus cellularum tenues; basi stylorum 
non noduliferi; rami stylorum glandulo- 
punctati, papillis aciculiformibus argutis. 



VOLUME 112, NUMBER 1 



217 






p 


n 




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^^ogg 


) 


m 


W^k 


H 




f 


■p 


HIS 


^^H 


^^^^^^^v* ^^^ 


■mi^ 






^^^^H 




W 




^ 


■ 


E 


1 


4 




3 




^^H 




« 





Figs. 1-4. Stokesia laevis (Hill) Greene, Knobloch 1426 (US), Mississippi. 1. Polar view, line = 20 |jim. 2. 
Colpar view, line = 20 fxm. 3, 4. Detailed views of muri with baculae and perforated tectum. 3. line = 3.8 fjim. 
4. line = 1.76 |xm. 



Achenia 3-4-angulata plerumque prope bas- 
em glandulo-punctata, cellulis subsuperfi- 
cialibus porosis fibriformibus, raphidis sub- 
nullis minutis breviter oblongis; squamae 
pappi 4 aut 5 subulatae perfacile deciduae 
8-9 mm longae. Grana pollinis triporata, la- 
cunis colpi rhomboideis, muds minute 
crenulatis (Figs. 1-4). Numerus chromoso- 
matum n = 1 . 

The single species in the subtribe is Sto- 
kesia laevis (Hill) Greene, which is native 



to the Southeastern United States in south- 
em South Carolina, Georgia, Alabama and 
Mississippi, western Florida and eastern 
Louisiana. 

Pacourina is a singularly distinctive 
emergent aquatic plant of tropical America. 
The inflorescence, with heads sessile in a 
series of leaf axils, is reminiscent of the 
Lepidaploa Group in the subtribe Vemoni- 
nae. The sclerified apical anther appendage, 
however, is totally foreign to that group and 



218 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 5, 6. Pacourina edulis Aubl., Killip and Smith 14576 (US), Colombia. 5. Whole grain, line = 15 ixm. 
2. Closer view showing lacuna with pore and lack of micropunctations on muri, line = 6.7 ixm. 



is extreme for even the Piptocarphinae and 
Lychnophorinae. Triporate, psilolophate 
pollen is known otherwise in the tribe Ver- 
nonieae only in the Paleotropical subtribe 
Erlangeinae, but the pollen of Pacourina is 
larger than pollen in any members of that 
group, and the inflorescence and anther ap- 
pendages are totally different. Thus, the 
combination of characteristics precludes 
placement in any presently existing subtribe 
of the Vemonieae. Closest relationships of 
the new subtribe are not known, but they 
are presumed to be Neotropical. The dis- 
tinctive nature of the plant may derive to 
considerable extent from is aquatic special- 
ization. 

Pacourininae H.Rob., subtribus no v. 

Type: Pacourina Aubl., Hist. PI. Guiane 
2:800. 1775. 

Plantae camosae aquaticae, folia simpli- 
cia altema valde dentata. Inflorescentiae se- 
riate cymosae, bracteis foliiformibus; capit- 
ula sessilia axillaria solitaria late campan- 
ulata homogama; bracteae involucri ca. 50 
latae virides et margine albae; receptacula 



epaleacea; flores ca. 50 in capitulo; corollae 
purpureae, lobis distaliter valde scleroideis; 
thecae anterharum base dentate appendicu- 
latae; appendices apicales antherarum gla- 
brae valde scleroideae; basi stylorum leniter 
latiores, pilis stylorum acicularibus. Ach- 
enia 10-costata suberose corticata in sulcis 
idioblastifera; setae pappi breves multiser- 
iatae deciduae, squamellis persistentibus. 
Grana poUinis triporata psilolophata emi- 
cropunctata (Figs. 5, 6). 

The single species in the subtribe in Pa- 
courina edulis Aubl. of Central America 
and tropical South America. The species is 
well illustrated in Nash and Williams (1976, 
fig. 7, p. 461). 

Acknowledgments 

The SEM photographs were made by Su- 
sann Braden of the National Museum of 
Natural History SEM Laboratory using a 
Hitachi S-570. Prints were prepared by 
Sherry Pittam, previously of the Depart- 
ment of Botany. 



VOLUME 112, NUMBER 1 



219 



Literature Cited 

Bentham, G., & J. D. Hooker. 1873. Compositae. — 
Genera Plantarum. Vol. 2(1): 163-533. Reeve & 
Co., London. 

Espinar, L. A. 1973. Revision del genero Hyalosehs 
(Compositae). — Kurtziana 7:195-21 1. 

Gunn, C. R., & G. A. White. 1974. Stokesia laevis: 
taxonomy and economic value. — Economic 
Botany 28:130-135. 

Hoffmann, O. 1890-94. Compositae 4(5):87-387. In 
A. Engler & K. Prantl, (eds.). — Die Natiirlichen 
Pflanzenfamilien. Leipzig. 

Jones, S. B. 1968. Chromosome numbers in South- 
eastern United States Compositae, I. — Bulletin 
of the Torrey Botanical Club 95:393-395. 



. 1974. Vernonieae (Compositae) chromosome 

numbers. — Bulletin of the Torrey Botanical 
Club 101:31-34. 

Nash, D. L., & L. O. Williams. 1976. Flora of Gua- 
temala. — Fieldiana: Botany 24(12): 1-603. 

Rickett, H. W. 1967. Wild flowers of the United States, 
the southeastern states. 2(2):601, pi. 224. Mc- 
Graw-Hill, New York. 

Robinson, H. 1996. The status of generic and subtribal 
revisions in the Vernonieae. Pp 511-526 in D. 
J. N. Hind & H. J. Beentje, eds., Compositae: 
Systematics. — Proceedings of the International 
Compositae Conference, Kew, 1994. (D. J. N. 
Hind, Editor-in-Chief)- Vol. 1, Royal Botanic 
Gardens, Kew. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):220-247. 1999. 

Revisions in paleotropical Vernonieae (Asteraceae) 

Harold Robinson 

Department of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0166, U.S.A. 

Abstract. — The paleotropical subtribes Erlangeinae, Centrapalinae and Gym- 
nantheminae are described as new. Fourteen paleotropical genera are described, 
raised from lower rank, resurrected, enlarged or reduced with 110 new com- 
binations. Cabobanthus (2 spp.), Hilliardiella (8 spp.), Orbivestus (4 spp.), 
Vemoniastrum (8 spp.), Koyamasia (1 sp.), Brenandendron (3 spp.), Myan- 
maria (1 sp.) and Manyonia (1 sp.) are described as new. Oocephala (S. B. 
Jones) H. Rob. (2 spp.) and Lampropappus (O. Hoffm.) H. Rob. (3 spp.) are 
elevated to generic rank. Four genera are resurrected, the Asian Acilepis D. 
Don with 9 of 10 species newly combined, the African Bechium DC. (2 spp.), 
Centrapalus Cass. (9 spp.), Linzia Sch. Bip. ex Walp. with 5 of 7 species newly 
transferred, and Polydora Fenzl (8 spp.). One species is transferred to make a 
total of 27 in Distephanus, 4 species are transferred to total 7 in Cyanthillium, 
and 39 species are transferred to total 43 in Gymnanthemum. Lamprachaenium 
Benth. is synonymized with Phyllocephalum Blume with 1 species transferred. 



The present paper provides a limited re- 
organization of the Eastern Hemisphere 
Vernonieae for purposes of a projected ge- 
neric review of the tribe. The need to sum- 
marize the whole tribe presents a special 
problem. The half of the tribe in the West- 
em Hemisphere has been rather well delim- 
ited into workable and phyletically accept- 
able genera (Robinson 1996). This includes 
an accurate delimitation of the type genus 
Vemonia Schreb. that is primarily found in 
eastern North America. The Eastern Hemi- 
sphere members of the tribe have thus been 
left in a particularly untenable position, 
with an excessively paraphyletic core genus 
and mostly paraphyletic or polyphyletic 
segregates, none of which are congeneric 
Vemonia. In fact, none of the existing sub- 
tribal names in the Vernonieae are appli- 
cable to the large elements that are endemic 
to the Eastern Hemisphere. 

At the time of the summary of the New 
World Vernonieae (Robinson 1966), only a 
few Eastern Hemisphere elements have 
been revised by the author. The name Bac- 



charoides was resurrected for the members 
of the Stengelia Group as early as Robinson 
et al. (1980), and transfers were made into 
Baccharoides and Cyanthillium (Robinson, 
1990) mostly to accomodate species adven- 
tive in the Western Hemisphere. Distephan- 
us was resurrected and partially revised as 
a last step in the removal of some of its 
species from the tribe Senecioneae (Rob- 
inson & Kahn 1986). In the latter study the 
opportunity was taken to provide the proper 
combination for the type species of Gym- 
nanthemum Cass, that had originally been 
named as a Eupatorium. Nevertheless, no 
further work was planned in the Eastern 
Hemisphere members of the Vernonieae as 
recently as the time of the publication of 
the preliminary Western Hemisphere revi- 
sion in the Kew International Compositae 
Conference volume (Robinson 1996). 

The present study has built upon the ref- 
erence works of many other authors. Hum- 
bert (1960), Wild (1977, 1978), Wild & 
Pope (1978a, 1978b), Jeffrey (1988), which 
cover all the genera of the tribe in various 



VOLUME 112, NUMBER 1 



221 



parts of Africa, have proven most useful in 
the present study. The study of Jones 
(1981), restricted to Vemonia sensu lato, 
and Lisowski (1992) that excludes Vemon- 
ia, have been less useful. The works of 
Kirkman (1981), Pope (1983), Koyama 
(1984), Isawumi (1993, 1995), and Isawumi 
et al. (1996) have added important infor- 
mation. The present study has used the 
summary of secondary metabolite chemis- 
try of the Vemonieae by Bohlmann and 
Jakupovic (1990) and the various reports of 
chromosome numbers by Jones (1974, 
1979, 1982). DNA studies are as yet limited 
to those mentioned by Keeley (1994, 1995) 
and Kim et al. (1996). The final conclusions 
of the present study are, nevertheless, ulti- 
mately based on examination of specimens 
in the United States National Herbarium 
(US) and on some specimens kindly sent on 
loan by Kew (K), British Museum (BM), 
Bruxelles (BR) and Paris (P). 

Jones (1977, 1981) noted basic differenc- 
es between the Vemonieae in the Eastern 
and Western Hemispheres, and this has 
been generally accepted by others such as 
Jeffrey (1988). These differences have been 
reviewed to some extent by Isawumi 
(1995). The Hemispheric trends noticed by 
Jones (1977) are in the chromosome num- 
bers and chemistry. The Western Hemi- 
sphere species usually have a chromosome 
number of N = 17, and the Eastern Hemi- 
sphere species have mostly N = 9 or 10. 
The chemicals cited were terpenoids and 
flavonoids. Jones (1981) also stated that tri- 
porate pollen grains were known only from 
the Eastern Hemisphere. Since that time, 
some examples of triporate grains have 
been found in Latin America, Acilepidopsis 
(Robinson 1989), Mesanthophora (Robin- 
son 1992a), Pacourina (Robinson 1992b), 
but at least the first two of these are con- 
sidered to be related to Eastern Hemisphere 
members of the tribe. Robinson and Kahn 
(1986) and Isawumi et al. (1996) briefly 
noted the apparent restriction of glandular 
dots on the anthers and their appendages to 
New World members of the Vemonieae. In 



the present study, the raphids of the achene 
wall in the Eastern Hemisphere species 
have proven to be predominently elongate, 
with only a few entities such as Gutenber- 
gia having characteristically subquadrate or 
short raphids. Most groups of Western 
Hemisphere Vemonieae have characteristi- 
cally subquadrate raphids. In their chemical 
survey, Bohlmann and Jakupovic (1990) 
found the distinctive 5-alkyl coumarins 
only in Eastern Hemisphere genera and spe- 
cies, groups that are all placed in this study 
in the new subtribe Erlangeinae. 

Inflorescences with scorpioid or well de- 
veloped seriate cymes in the Vemonieae 
seem almost entirely restricted to the New 
World and New World genera like Stru- 
chium that have become Pantropical. A no- 
table exception, Manyonia, described below 
as a new genus from East Africa, is consid- 
ered to be a relative of New World genera, 
and it is described in the subtribe Vemon- 
iinae. 

Some doubts about the hemispheric dif- 
ferences were expressed by Keeley and 
Turner (1990). They mention some cases of 
Western Hemisphere ''Vernonia'' with 
chromosome numbers of N = 10 or 12, and 
they call attention to similarities between 
the groups referred to by the present author 
as Piptocarphinae in the Americas and the 
Gymnantheminae in the Eastern Hemi- 
sphere. The latter groups both tend to be 
woody, often have deciduous inner involu- 
cral bracts and usually have blunt-tipped 
sweeping hairs on their styles. Neverthe- 
less, the possible relationship is considered 
here to be above the generic and subtribal 
levels. The doubts of Keeley & Turner 
(1990) are justified to the extent that Ver- 
nonia cannot be simply divided into two 
genera, one for each Hemisphere. Many 
segregates are needed for each hemisphere. 

There is an important limitation to the 
use of pollen structure as a phyletic char- 
acteristic in the Vemonieae, a limitation al- 
ready seen in both Western and Eastern 
Hemisphere members of the tribe (Robin- 
son 1996). First, the simple non-lophate or 



222 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



sublophate pollen, Type A, often consid- 
ered primitive in the tribe, was not consid- 
ered primitive by the present author (Rob- 
inson 1996). Second, some pollen varia- 
tions are of more importance than others. 
Differences between various lophate pat- 
terns seem to correlate well with other char- 
acters, but all groups seem able to revert 
erratically to non-lophate or sublophate 
forms, the Type A of Jones (1981). The tru- 
ly triporate forms, with irregularly orga- 
nized polar lacunae, are evidently not close- 
ly related to other lophate types with dis- 
tinct colpi. However, they may be very 
closely related to colporate forms that are 
non-lophate. This does not mean that the 
difference between triporate and non-lophate 
tricolporate cannot be used as a key char- 
acter when the character happens to corre- 
late with other features, as in Cyanthillium. 

The present study has been restricted by 
available time and specimens. The new sub- 
tribes are kept to a minimum, although that 
minimum seems to fit the Paleotropical ma- 
terial rather well. The new genera and taxa 
elevated to generic rank are those that seem 
inescapable, and no attempt is made here to 
dispose of some additional distinctive ele- 
ments in the Eastern Hemisphere that do 
not fit in the genera presently recognized. 
Possibly, some smaller distinctive elements 
have been completely missed in the present 
survey. The number of species transferred 
into the various segregate genera is neces- 
sarily incomplete, but proper names are 
now available for many species that should 
not be retained in Vemonia. It is hoped that 
other authors will continue the process of 
making transfers. An attempt has been 
made to avoid overly broadening the con- 
cepts of the segregate genera. Even so, 
Gymnanthemwn has been interpreted to in- 
clude many elements that differ from the 
type by corolla lobe, style and achene char- 
acters. This has been done with the convic- 
tion that the broader concept of that genus 
will prevail. 

The synonymy given for most of the Af- 
rican species follows Jeffrey (1988). 



New Subtribes of the Eastern Hemisphere 
Vemonieae 

All presently named subtribes of the Ver- 
nonieae are based on primarily Western 
Hemisphere groups. None of these subtribal 
names apply to the primarily Eastern Hemi- 
sphere groups. To rectify this situation, the 
following three new subtribes are named. 

Erlangeinae H. Rob., subtribus no v. Type: 
Erlangea Sch. Bip., Flora 36:34. 1853. 
Type: E. plumosa Sch. Bip. 

Vemonia sect. Tephrodes DC. Prodr. 5: 

24. 1836. Type: Conyza cinerea L. 
Bechium DC, 5:70. 1836. Type: B. scap- 

iforme DC. 
Vemonia sect. Lepidella Oliver & Hiem, 

Fl. Trop. Afr. 3:267. 1877. Type: Ver- 

nonia petersii OHver & Hiem. 
Vemonia subg. Orbisvestus S. B. Jones, 

Rhodora 83:60. 1981. Type: Vemonia 

karaguensis Oliver & Hiem. 
Vemonia sect. Orbisvestus S. B. Jones, 

Rhodora 83:61. 1981. Type: as above. 
Vernonia subsect. Orbisvestus S. B. 

Jones, Rhodora 83:61. 1981. Type: as 

above. 
Vemonia subsect. HiUiardianae S. B. 

Jones, Rhodora 83:66. 1981. Type: 

Webbia oligocephala DC. 
Vemonia subsect. Tephrodes (DC.) S. B. 

Jones, Rhodora 83:70. 1981. 
Vemonia subsect. Lepidella (OUver & 

Hiem) S. B. Jones, Rhodora 83:72. 

1981. 
Vernonia subsect. Oocephalae S. B. 

Jones, Rhodora 83:72. 1981. Type: 

Vemonia oocephala Baker 
Vemonia subsect. Bechium (DC.) S. B. 

Jones, Rhodora 83:73. 1981. 

Plantae herbaceae annuae aut perennes 
vel frutescentes, pihs saepe symmetriciter 
T-formibus. Foha altema vel opposita vel 
temata pinnatinervata. Receptaculum epa- 
leaceum vel raro paleaceum. Flores 3 vel 
ca. 100 in capitulo; corollae lavandulae vel 
purpureae; thecae antherarum base non vel 
tenuiter caudatae; appendices apicales gla- 



VOLUME 112, NUMBER 1 



223 



brae in parietibus cellularum tenues raro 
tenuiter omatae; base stylorum noduliferi; 
rami stylorum aciculiformiter papillosi. 
Achenis 4-6 vel 8-10 costata; raphidis 
plerumque elongatis raro subquadratis; car- 
popodia anguste cylindrica; pappus longe 
setiformis vel abbreviatus vel coroniformis 
saepe facile deciduus. Grana pollinis non 
lophata et tricolporata vel lophata et tripor- 
ata in formibus lophatis lacunis polaribus 
irregularibter dispositis, tectis micropunc- 
tatis vel emicropunctatis. Numerus chro- 
mosomatum N = 9, 10, 20. 

The name is chosen to conform with the 
already established term "Erlangeoid" 
(Pope 1983). The subtribe is circumscribed 
to include all the Vemonieae presently 
known with triporate pollen (excluding Pa- 
courina Aubl., Robinson 1992b) or 5-alkyl 
coumarins (Bohlmann & Jakupovic 1990). 
The unquestioned core of the subtribe in- 
cludes genera with only 4-5 angled achenes 
and herbaceous habits, but, at present, ad- 
ditional forms with 10-ribbed achenes and 
woody habits are also included. All the in- 
cluded genera have acicular sweeping hairs 
of the style. The genera included are the 
New World Acilepidopsis H. Rob. (1989) 
and Mesanthophora H. Rob. (1992a), and 
Old V^orXd Acilepis D. Don, Ageratinastrum 
Mattf., Ambassa Steetz, Bechium DC, 
Bothriocline Oliv. ex Benth., Brachythrix 
Wild & G. V. Pope, Cyanthillium Blume, 
Decastylocarpus Humbert, Dewildemania 
O. Hoffm., Diaphractanthus Humbert, Er- 
langea Sch. Bip., Ethulia L. f., Gossweilera 
S. Moore, Gutenbergia Sch. Bip. ex Walp., 
Hystrichophora Mattf., lodocephalus Tho- 
rel ex Gagnep., Kinghamia C. Jeffrey, Lam- 
prachaenium Benth., Msuata O. Hoffm., 
Phyllocephalum Blume, Omphalopappus 
O. Hoffm., Rastrophyllum Wild & G. V. 
Pope, and six genera newly named or ele- 
vated below. 

Centrapalinae H. Rob., subtribus nov. 
Type: Centrapalus Cass., Diet. Sci. Nat. 
ed. 2, 7:382. 1817. Type: Centrapalus 
galamensis Cass. 



Vernonia subsect. Stengelia Sch. Bip. ex 
Walp., Repert. Bot. Syst. 2 [Suppl. 1]: 
946. 1843. Type: Vernonia adoensis 
Sch. Bip. ex Walp. 

Vernonia sect. Stengelia (Sch. Bip. ex 
Walp.) Benth. & Hook, f.. Gen. PI. 2: 
227. 1873. 

Vernonia subsect. Centrapalus (Cass.) S. 
B. Jones, Rhodora 83:69. 1981. 

Vernonia sect. Azureae S. B. Jones, Rho- 
dora 83:74. 1981. Type: Vernonia gla- 
bra Vatke. 

Plantae herbaceae perennes vel raro an- 
nuae, pilis simplicibus multiseptatis vel 
asymmetriciter T- formibus vel nullis. Folia 
altema pinnatinervata vel longitudinaliter 
nervata. Bracteae involucri apice saepe ap- 
pendiculatae vel plerumque herbaceae. Flo- 
res numerosi in capitulo; corolla purpurea 
vel azurea; thecae antherarum base rotun- 
datae; appendices apicales glabrae vel raro 
papilliferae in parietibus cellularum leniter 
incrassatae; base stylorum noduliferi vel 
non noduliferi; rami stylorum acute papil- 
liferi. Achenis 10-costata, raphidis elongatis 
vel interdum subquadratis; pappus pler- 
umque setiformis. Grana pollinis tricolpor- 
ata lophata vel non lophata micropunctata 
vel non micropunctata in formibus lophatis 
lacunis polaribus solitariis interdum prae- 
sentibus. Numerus chromosomatum N = 9, 
10. 

The subtribe is typified by Centrapalus 
Cass., and includes Adenoon Dalz., Aedesia 
O. Hoffm, Baccharoides Moench, Cam- 
chaya Gagnep., Lachnorhiza A. Rich., Lin- 
zia Sch. Bip. ex Walp., Muschleria S. 
Moore, Neurolakis Mattf. and Pleurocar- 
paea Benth. The sesquiterpene constituents 
include elemanolides (Bohlmann & Jaku- 
povic 1990). 

The genera of the subtribe are herba- 
ceous or weakly shrubby, and the sweeping 
hairs of the styles are acicular. The subtribe 
includes elements with the distinctive Lin- 
z/a-type pollen cited by Jeffrey (1988) such 
as Linzia and Aedesia and one with polar 
lacunae on its pollen grains and a lack of 



224 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



basal stylar nodes like Baccharoides (Isa- 
wumi 1993; Isawumi et al. 1996). 

Gymnantheminae H. Rob., subtribus nov. 
Type: Gymnanthemum Cass., Bull. Soc. 
Philom. Paris 1817:10. 1817. Type: Gym- 
nanthemum cupulare Cass. [= G. color- 
atum (Willd.) H. Rob. & B. Kahn]. 

Distephanus Cass., Bull. Soc. Philom. 

Paris 1817:151. 1817. Type: Diste- 
phanus populifolius (Lam.) Cass. 
Vernonia sect. Strobocalyx Blume ex 

DC, Prodr. 5:21. 1836. Type: Vernon- 
ia arborea Buch.-Ham. 
Gongrothamnus Steetz in Peters., Reise 

Mossamb. Bot. 336. 1864. Type: G. 

divaricatus Steetz in Peters 
Vernonia sect. Distephanus (Cass.) 

Benth. & Hook, f.. Gen. pi. 2:228. 

1873. 
Vernonia sect. Lampropappus O. Hoffm., 

Bol. Soc. Broter. 13:14. 1896. Type: 

Vernonia lampropappa O. Hoffm. 
Vernonia subsect. Strobocalyx (Bl. ex 

DC.) S. B. Jones, Rhodora 83:64. 

1981. 
Vernonia subsect. Gongrothamnus 

(Steetz) S. B. Jones, Rhodora 83:65. 

1981. 
Vernonia subsect. Pawekianae S. B. 

Jones, Rhodora 83:66. 1981. Type: 

Vernonia angulifolia DC. 
Vernonia subsect. Urceolatae S. B. 

Jones, Rhodora 83:67. 1981. Type: 

Vernonia sphaerocalyx O. Hoffm. 
Vernonia subsect. Turbinella S. B. Jones, 

Rhodora 83:67. 1981. Type: Vernonia 

lampropappa O. Hoffm. 
Vernonia subsect. Distephanus (Cass.) S. 

B. Jones, Rhodora 83:68. 1981. 
Vernonia subsect. Glutinosae S. B. Jones, 

Rhodora 83:73. 1981. Type: Vernonia 

glutinosa DC. 

Plantae fruticosae vel arborescentes vel 
scandentes, pilus simplicibus et arachnoid- 
eis vel L-formibus vel T-formibus. Folia al- 
tema pinnatinervata vel trinervata. Bracteae 
involucri 20-70 in seriebus 2-7 plerumque 
gradatae interiores interdum deciduae; re- 



ceptacula epaleacea vel paleacea. Flores 1- 
40 (-75) in capitulo; corollae lavandulae 
vel roseae vel flavae, lobis plerumque er- 
ectis et longe triangularibus rariter elongatis 
et arete revolutis; thecae antherarum base 
plerumque valde caudatae; appendices api- 
cales induratae et glabrae in parietibus cel- 
lularum aliquantum ornate incrassatae; basi 
stylorum non noduliferi vel noduliferi in- 
terdum abrupte noduliferi; rami stylorum 
plerumque obtuse papilliferi. Achenia 5 aut 
10-12-costata, raphidis plerumque elongatis 
interdum subquadratis vel nuUis, carpopo- 
dia lata; pappus plerumque setiformis, Gra- 
na pollinis tricolporata non lophata vel sub- 
lophata vel raro lophata in formibus lopha- 
tis lacunis polaribus solitariis nullis; tectis 
micropunctatis. Numerus chromosomatum 
N = 10, 2N = 30. 

The subtribe is typified by the genus 
Gymnanthemum Cass., but it also includes 
Distephanus Cass., Centauropsis Boj. in 
DC, Oliganthes Cass., and three genera 
named or elevated below. The sesquiter- 
pene lactone constituents include eleman- 
olides (Bohlmann & Jakupovic 1990). 

The subtribe includes all of the true large 
shrub and tree Vernonieae in the Eastern 
Hemisphere. The sweeping hairs of the 
styles often have rather blunt tips. Inner in- 
volucral bracts are persistent or deciduous, 
and a few species of Gymnanthemum and 
Brenandendron have long coiled corolla 
lobes. These characteristics are generally 
shared by the Gymnantheminae and the 
American Piptocarphinae, but deciduous 
bracts and reflexed corolla lobes are much 
more consistently present and blunt sweep- 
ing hair much less consistently present, in 
the Piptocarphinae. The Gymnantheminae 
lack stellate hairs like those common in the 
Piptocarphinae . 

New Genera and New Combinations of 

Eastern Hemisphere Vernonieae 

Subtribe Vemoniinae 

Manyonia H. Rob., gen. nov. (Vemoni- 
inae). 



VOLUME 112, NUMBER 1 



225 



Type: Vemonia peculiaris Verde. 

Plantae herbaceae perennes ad 1 m altae; 
caules brunnescentes striati sparce hispidu- 
li, pilis simplicibus multiseptatis uniseriatis, 
internodis 4-6 cm longis. Folia altema, pe- 
tiolis 8-10 mm longis; laminae membran- 
aceae ovatae 8-11 cm longae 3.2-5.0 cm 
latae base anguste cuneatae margine biser- 
ratae apice plusminusve acuminatae supra 
sparce pilosae subtus dense glandulo-punc- 
tatae, nervis secundariis utrinque 5 vel 6 su- 
pra et subtus puberulis. Inflorescentiae se- 
riate cymosae, ramis ad 14 cm longis, pe- 
dunculis plerumque brevibus 2-3 (-15) mm 
longis dense puberulis. Capitula campanu- 
lata 6 mm alta et lata; bracteae involucri ca. 
100; seriebus exterioribus 3-4 patentes 
longe subulatae 2.5 mm longae base dila- 
tatae 0.1—0.2 mm latae infeme margine 
puberulae; bracteae intermediae oblongae 4 
mm longae apice longe aristatae; bracteae 
interiores anguste oblongae 5 mm longae et 
2 mm latae apice acuminatae; receptaculum 
glabrum in diametro ca. 1.8 mm. Flores ca. 
35 in capitulo; corollae lilacinae ca. 5.5 mm 
longae, tubis angustis ca. 2.5 mm longis su- 
peme infundibularibus, faucibus ampliatis 
ca. 1 mm longis, lobis oblongis ca. 1.5 mm 
longis et 0.3 mm latis extus glanduliferis; 
thecae antherarum ca. 2 mm longae base 
non caudatae; appendices apicales ca. 0.4 
mm longae in parietibus cellularum tenues 
glabrae; basi stylorum disciformiter nodati; 
rami stylorum in papillis aciculiformibus 
obsiti. Achenia oblonga 1.5 mm longa 5- 
costata inter costam breviter setulifera et 
longitudinaliter multiseriate idioblastifera, 
costis glabris prominentibus et induratis, ra- 
phidis perdense dispositis subquadratis vel 
polygonatis; setae pappi ca. 20 barbellatae 
ca. 4 mm longae; squamulae exteriores ca. 
25 minute ciliatae. Grana pollinis in dia- 
metro ca. 35 |JL sublophata tricolporata. 

The genus Manyonia has been consid- 
ered by the author as a relative of New 
World Vernonieae since he first saw the il- 
lustration of the seriate cymes accompany- 
ing the original description (Verdcourt 



1956). This structure, otherwise lacking in 
native Old World Vernonieae, is confirmed 
in material borrowed from The Royal Bo- 
tanic Gardens, Kew. This relationship 
seems confirmed by a poorly researched 
characteristic, the type of ornamentation of 
the endothecial cells of the anther. The cells 
in Manyonia have rather strong thickenings, 
longitudinal in some cell rows, and in other 
rows, arching across the lower end. This is 
a pattern seen in many Western Hemisphere 
Vernonieae. Eastern Hemisphere Vernon- 
ieae have weaker ornate thickenings if any. 
Further examination of specimens of Man- 
yonia shows a strong resemblance to New 
World genera such as Heterocypsela H. 
Rob. and Dipterocypsela S. F. Blake, es- 
pecially in the wall of the achene with ex- 
tremely crowded subquadrate or polygonal 
raphids. The new genus differs from Het- 
erocypsela and Dipterocypsela in lacking 
the heteromorphic achenes and the glan- 
duliferous anther appendages of its Ameri- 
can relatives. The pollen of Manyonia is 
sublophate, more like Dipterocypsela and 
unlike the lophate form in Heterocypsela. 

The generic name is derived from the 
vernacular name Manyoni cited on Burtt 
5119 (K). The needed combination is as 
follows: 

Manyonia peculiaris (Verde.) H. Rob., 
comb. nov. basionym: Vemonia pecu- 
liaris Verde, Kew Bull. 1956:447. 1956. 
Tanzania. 

New Genera and New Combinations of 

Eastern Hemisphere Vernonieae 

Subtribe Erlangeinae H. Rob. 

Acilepis D. Don, Prodr. Fl. Nepal. 169. 
1825. Type: Acilepis squarrosa D. Don. 

Lysistemma Steetz in Peters, Reise Mos- 
samb. Bot. 340. 1864. Type: Lysistem- 
ma dendigulense (DC.) Steetz. 

Xipholepis Steetz in Peters, Reise Mos- 
samb. Bot. 344. 1864. Type: Xiphole- 
pis silhetensis Steetz. 

Vernonia sect. Xipholepis (Steetz) Benth. 
& Hook.f., Gen. pi. 2:229. 1873. 



226 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Erect perennial herbs; stems pentangular, 
with hairs multiseptate at base and often 
with long subfusiform apical cell. Leaves 
alternate, obovate to oblong-ovate. Inflores- 
cences of single heads, spiciform cymes, or 
corymbose cymes with few to many heads. 
Involucres funnelform to campanulate; bracts 
50-200 in 6-12 series, persistent, apiculate to 
subacute; receptacle epaleaceous. Heads with 
25-80 florets; corollas lavender, tubes slender 
below, funnelform above into throat, throat 
half or less as long as anther thecae, lobes 
long and narrow, with glandular dots; anther 
bases blunt, not tailed; apical anther append- 
ages glabrous, with thin- walled ceUs; style 
base with node, style branches with acicular 
sweeping hairs. Achenes 8-10 ribbed, setulae 
with one cell long, other cell short, raphids 
oblong with rhomboid tips; pappus whitish, 
both series rather easily deciduous, with 
many barbeUate inner setae, outer setae short- 
er, scarcely broader. PoUen triporate, lophate, 
nearly psilate, emicropunctate, with ca. 20 la- 
cunae. 

The Asiatic Acilepis is distinct in the 
rather simple stem hairs, the pedunculate or 
separated heads, the triporate pollen and in 
such details as the often totally deciduous 
pappus and the highly unequal cells of the 
setulae of the achene. Jeffrey (1988) rec- 
ognized the group for an AMcan species, 
Vemonia polysphaera, treated below as the 
new genus Cabobanthus. The latter has 
basal tubers, sessile clustered heads, a more 
persistent pappus and the cells of the setu- 
lae of equal length. 

The following ten species are recognized 
in the genus: 

Acilepis aspera (Buch.-Ham.) H.Rob., 
comb. nov. basionym: Vemonia aspera 
Buch.-Ham., Trans. Linn. Soc. London 
14:219. 1824. 

Eupatorium pyramidale D. Don, Prodr. 

Fl. Nepal. 170. 1825. 
Vemonia roxburgii Less., Linnaea 6:674. 

1831. 
Xipholepis aspera (Buch.-Ham.) Steetz in 



Peters, Reise Mossamb. Bot. 345. 
1864. 
Vemonia pyramidalis (D. Don) Mitra, 
Ind. For. 99:100. 1973. China, India, 
Myanmar, Nepal, Thailand. 

Acilepis clivorum (Hance) H. Rob., comb, 
nov. basionym: Vemonia clivorum 
Hance, J. Bot. 7:164. 1869. 

Aster coriaceiformis H. Lev. & Vaniot, Re- 
pert. Spec. Nov. Regni Veg. 8:358. 1910. 
China. 

Acilepis dalzelliana (J. R. Drumm. & 
Hutch.) H. Rob., comb. nov. basionym: 
Vemonia dalzelliana J. R. Drumm. & 
Hutch., Kew Bull. 1909:261. 1909. 
India. 

Acilepis dendigulensis (DC.) H. Rob., 
comb. nov. basionym: Decaneurum den- 
digulense DC. in Wight, Contr. Bot. Ind. 
7. 1834, not Vemonia dendigulensis DC. 

Lysistemma dendigulense (DC.) Steetz in 
Peters, Reise Mossamb. Bot. 341. 
1864. 

Vemonia indie a C. B. Clarke, Comp. Ind. 
16. 1876. Western peninsular India. 

Acilepis nantcianensis (Pamp.) H. Rob., 
comb. nov. basionym: Vemonia bractea- 
ta var. nantcianensis Pamp., Nouv. 
Giom. Bot. Ital., n.s. 18:98. 1911. 

Vemonia silhetensis var. nantcianensis 
(Pamp.) Hand.-Mazz., Symb. Sin. 7: 
1084. 1936. 

Vemonia nantcianensis (Pamp.) Hand.- 
Mazz., Noitsibl. Bot. Gart. Mus. Berl.- 
Dahl. 13:608. 1937. 
China. 

Acilepis saligna (DC.) H. Rob., comb. nov. 
basionym: Vemonia saligna DC, Prodr. 
5:33. 1836. 

Vemonia longicaulis DC, Prodr. 5:33. 
1836. 

Vemonia martinii Vaniot, Bull. Acad. In- 
tern. Geogr. Bot. 12:124. 1903. 

Vemonia sequinii Vaniot, Bull. Acad. In- 
tern. Geogr. Bot. 12:241. 1903. 
China, India, Myanmar. 



VOLUME 112, NUMBER 1 



227 



Acilepis scariosa (DC.) H. Rob., comb, 
nov. basionym: Decaneurum scariosum 
DC, Prodr. 7:264. 1838. 

Vemonia scariosa Am., Nova Acta Phys. 

Med. Acad. Caes. Leop. Carol. Nat. 

Cur. 18:346. 1836, horn, illeg., not V. 

scariosa Poir., 1808. 
Gymnanthemum scariosum (DC.) Sch. 

Bip. ex Walp., Rep. 2:949. 1843. 
Centratherum scariosum C. B. Clarke, 

Comp. Ind. 4. 1876. 
Vemonia lankana Grierson, Ceylon J. 

Sci., Biol. Sci. 10:43. 1972. 

Sri Lanka. 

Acilepis silhetensis (DC.) H. Rob., comb, 
nov. basionym: Decaneurum, silhetense 
DC, Prodr. 5:67. 1836. 

Eupatorium glabrum Heyne ex Wallich, 
Num. List. Dr. pi. 3283. 1831, nom. 
nud. 

Decaneurum glabrum DC, Prodr. 5:67. 
1836. 

Gymnanthemum glabrum (DC) Sch. Bip. 
ex Walp., Rep. 2:948. 1843. 

Gymnanthemum silhetense (DC) Sch. 
Bip. ex Walp., Rep. 2:948. 1843. 

Xipholepis silhetensis (DC) Steetz in Pe- 
ters, Reise Mossamb. Bot. 344. 1864. 

Vemonia bracteata Wall, ex C B. 
Clarke, Comp. Ind. 17. 1876. 

Vemonia silhetensis (DC) Hand.-Mazz., 
Symb. Sin. 7:1084. 1936. 
China, India, Thailand. 

Acilepis spirei (Gandog.) H. Rob., comb, 
nov. basionym: Vemonia spirei Gandog., 
Bull. Soc. Bot. France 54:194. 1907. 
China, Laos, Vietnam. 

Acilepis squarrosa D. Don, Prodr. Fl. Nepal 
169. 1825. 

Vemonia squarrosa (D. Don) Less., Lin- 

naea 6:627. 1831. 
Vemonia rigiophylla DC, Prodr. 5:15. 

1836. 
Vemonia teres Wall, ex DC, Prodr. 5:15. 

1836. 

China, India, Nepal, Sikkim, Thailand, 

Vietnam. 



Bechium DC, Prodr. 5:70. 1836. Type: Be- 
chium scapiforme DC. 

Erect or subhorizontally proliferating an- 
nual or biennial herbs, 1-4 dm high; hairs 
mostly of one long cell, appearing seri- 
ceous, red stipitate glands with multicellular 
tips on stems and bracts. Leaves alternate, 
rosulate or subrosulate, subsessile, blades 
oblong. Inflorescences scapiform with 1 to 
many corymbosely disposed heads. Heads 
shortly to longly pedunculate; involucral 
bracts ca. 30 in ca. 3 series, with red stip- 
itate glandular hairs; receptacle epalea- 
ceous. Florets 25-50; corollas reddish- vio- 
let, slender tubes funnelform above, throat 
very short, lobes with sparse biseriate non- 
glandular hairs, rarely with stipitate reddish 
gland at tip; anther bases rounded, without 
tails; apical appendage glabrous, with thin 
cell walls; style base with distinct wide 
node; style with sweeping hairs fat, pointed, 
few to many septate. Achenes 10-costate, 
with many unevenly pointed setulae, many 
idioblasts, raphids elongate with rhomboid 
tips; pappus with single series of easily de- 
ciduous to subpersistent bristles and short 
outer squamae, bristles narrowed below, 
some wider distally. Pollen tricolporate, 
non-lophate, echinate. 

The genus is recognized primarily on the 
basis of the herbaceous annual or biennial 
habit, elongate raphids of the achene wall, 
slender white pappus bristles, type A pol- 
len, and the rather broad, pointed sweeping 
hairs with few to many septations. The spe- 
cies placed in the genus here are two of the 
members of the Vemonieae in Madagascar 
that have reddish glands with multicellular 
caps on their peduncles and involucral 
bracts. Both species need new combinations 
since the oldest name for the type species 
has never been transferred to the genus. 
Bechium nudicaule (Less.) H. Rob., comb. 

nov. basionym: Vemonia nudicaulis 

Less., Linnaea 6:637. 1831. 

Bechium scapiforme DC, Prodr. 5:71. 

1836. 
Vemonia scapiformis (DC.) Drake, Bull. 



228 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Soc. Bot. France 46:244. 1889. 

Madagascar. 
Bechium rhodolepis (Baker) H. Rob., 
comb. nov. basionym: Vemonia rhodo- 
lepis Baker, J. Bot. 20:139. 1882. 

Vemonia purpureo-glandulosa Klatt, 
Bot. Jahrb. Syst. 12 (Beibl. 27):21. 
1890. 
Madagascar. 

Cabobanthus H. Rob., gen. nov. (Erlangei- 
nae). 
Type: Vemonia polysphaera Baker. 

Plantae herbaceae perennes base tuberosae. 
Caules subglabri aut tomentosi, pilis simpli- 
cibus uniseriatis multiseptatis. FoUa altema 
sessilia vel subsessilia. Capitula in axillares 
superiores ad 5 aggregata; involucra infun- 
dibularia, bracteis ca. 35 in seriebus ca. 5 
ovatis vel oblongis apiculatis; receptacula 
epaleacea. Flores ca. 10 in capitulo; corollae 
purpureae, tubis cyHndricibus supeme infun- 
dibularibus; faucibus quam theceis dimidiis 
brevioribus; thecae base breviter caudatae; 
appendices antherarum apicales in parietibus 
cellularibus tenues non glanduliferae; basi 
stylorum noduliferi; rami stylorum acicuUfor- 
miter papillosi. Achenia 8-10-costatae, setuHs 
in celluHs aequales, raphidis minutis anguste 
oblongis; setae pappi arete barbellatae. Grana 
pollinis triporata ca. 35 |xm lacunosa emicro- 
punctata. 

The genus is notable for the erect stems 
bearing axillary clusters of heads and the 
triporate, emicropunctate pollen grains. The 
following two species are recognized: 

Cabobanthus bullulatus (S. Moore) H. 
Rob., comb, nov., basionym: Vemonia 
bullulata S. Moore, J. Bot. 65, suppl. 2: 
44. 1927. 
Zambia. 

Cabobanthus polysphaerus (Baker) H. 
Rob., comb, nov., basionym: Vemonia 
polysphaera Baker, Kew Bull. 1898:148. 
1898. 

Vernonia humblei De Wild., Repert. 
Spec. Nov. Regni Veg. 13:207. 1914. 
Congo, Tanzania, Zambia. 



Cyanthillium Blume, Bidjr. 889. 1826. 
Type: Cyanthillium villosum Blume. 

Isonema Cass., Bull. Soc. Philom. Paris 
1817:152. 1817, hom. illeg., not Iso- 
nema R. Br., 1810. Type: Isonema 
ovata Cass. 

Vemonia sect. Tephrodes DC, Prodr. 5: 
24. 1836. Lectotype: Conyza cinerea 
Blume (Jones 1981). 

Cyanopsis Blume ex DC, Prodr. 5:69. 
1836, nom. illeg. et superfl., not Cass., 
1817. 

Claotrachelus ZoU. & Moritz ex ZoU., 
Natuur-Genrrsk. Arch. Ned. Indie 2: 
263, 565. 1845. Type: Claotrachelus 
rupestris ZoU. & Moritz ex ZoU. 

Seneciodes L. ex Post & O. Kuntze, Lex. 
Gen. Phan. 2:515. 1903. Type: Conyza 
cinerea L. 

Triplotaxis Hutch., Bull. Misc. Inform. 
1914:355. 1914. Lectotype: Herderia 
stellulifera Benth. in Hook.f. (Robin- 
son 1990). 

Vemonia subsect. Tephrodes (DC.) S. B. 
Jones, Rhodora 83:70. (1981). 

Annual or short-lived perennial herbs ca. 1 
m taU; stem hairs asymmetricaUy and sym- 
metricaUy T-shaped. Leaves alternate, nar- 
rowly petiolate, blades thinly papery. Inflo- 
rescences terminal, corymbose to pyramidal 
cymes. Heads pedunculate; involucral bracts 
papery, green with pale or purplish margins, 
ca. 30 in 3(-5) gradate series, persistent; re- 
ceptacle epaleaceous. Florets 15-94; coroUas 
bluish to lavender, funnelform with slender 
lower tubes, throat a third as long to nearly 
as long as lobes, lobes with simple hairs es- 
peciaUy near tips; anthers without tails; apical 
appendages glabrous, with thin ceU walls; 
style base with broad node; sweeping hairs 
acicular. Achenes 5-angled or ribbed, or te- 
rete, with idioblasts, sometimes with glands, 
raphids elongate; pappus with many long, 
fragUe, slender-tipped bristles or squameUae, 
persistent, with caUose ring in one species. 
PoUen triporate, echinolophate. N = 9, 11, 18 
(Jones 1979, 1982). 

The synonymy foUows that in Robinson 



VOLUME 112, NUMBER 1 



229 



(1990a) with the removal of Vemonia sub- 
sections Orbisvestus and Hilliardianae. At- 
tempts to interpret the genus more broadly to 
include Gutenbergia Sch.Bip. ex Walp. (Rob- 
inson, 1990b), are rejected here. At this time, 
other more closely related species, with 
shrubbier habits, non-lophate pollen, tailed 
anthers and T-shaped hairs or no hairs on the 
coroUa lobes, are placed in separate genera 
(see Hilliardiella and Orbivestus below). The 
species of Cyanthillium are all annuals or 
weak perennial subshrubs with triporate lo- 
phate pollen. The following seven species are 
presently placed in the genus. 

Cyanthillium albicans (DC. in Wight) H. 
Rob., comb. nov. basionym: Vemonia al- 
bicans DC. in Wight, Contrib. Bot. Ind. 
6. 1834. 
Western peninsular India. 

Cyanthillium cinereum (L.) H. Rob., Proc. 
Biol. Soc. Wash. 103:252. 1990. 

Conyza cinerea L., Sp. PI. 862. 1753. 
Vemonia cinerea (L.) Less., Linnaea 4: 

291. 1829. 
Seneciodes cinerea (L.) Post & O. Kuntze, 

Lex. Gen. Phan. 2:515. 1903. 

Throughout paleotropical region, 

widely adventive in Neotropics. 

Cyanthillium conyzoides (DC. in Wight) H. 
Rob., comb. nov. basionym: Vemonia 
conyzoides DC. in Wight, Contr. Bot. Ind. 
6. 1834. 
Western peninsular India. 

Cyanthillium hookerianum (Am.) H. Rob., 
comb. nov. basionym: Vemonia hooker- 
iana Am., Nov. Act. Nat. Cur. 18:346. 
1836. 
Sri Lanka. 

Cyanthillium patulum (Ait.) H. Rob., Proc. 
Biol. Soc. Wash. 103:252. 1990. 

The synonymy is as in Robinson (1990a) 
with the exclusion of Conyza chinensis 
Lam. and its combinations. 

Tropical Asia, Indonesia, Madagascar, 

adventive in West Indies. 

Cyanthillium stelluliferum (Benth.) H. 



Rob., Proc. Biol. Soc. Wash. 103:252. 
1990. 

Herderia stellulifera Benth. in Hook. f. 

& Benth., Niger FL, 425. 1849. 
Triplotaxis stellulifera (Benth.) Hutch., 

Bull. Misc. Inf. Kew 1914:356. 1914. 

West and central tropical Africa, 

Uganda to Angola. 

Cyanthillium vernonioides (Muschl.) H. 

Rob., comb. nov. basionym: Erlangea 

vernonioides Muschl., Bot. Jahrb. Syst. 

46:62. 1911, not Vemonia vernonioides 

(A. Gray) Bacigalupo, 1931. 
Vemonia me io Stephana C. Jeffrey, Kew 

Bull. 43:225. 1988. 

Hilliardiella H. Rob. gen. nov. (Erlangei- 
nae). Type: Vemonia pinifolia Less. 

Webbia DC, Prodr. 5:72. Oct. 1836, lec- 
totype Vemonia pinifolia Less., horn, 
illeg., not Webbia Spach, Jun 1836. 

Vernonia subsect. Hilliardianae S. B. 
Jones, Rhodora 83:66. 1981. Type: Ver- 
nonia oligocephala (DC.) Sch. Bip. 

Plantae herbaceae perennes ad 1 m altae; 
caules pilosi, pilis aequaliter T-formibus. 
Folia altema; laminae subtus saepe dense 
canescentiter pilosae. Inflorescentiae laxe 
vel subdense corymbose cymosae; capitula 
pedunculata; involucra campanulata; brac- 
teae 25-40 ca. 3-4-seriatae persistentes; re- 
ceptacula epaleacea. Flores 12-20 in capi- 
tulo; corollae purpureae extus pauce vel 
dense pilosae, pilis T-formibus leniter con- 
tortis; tubis supeme infundibularibus, fau- 
cibus brevibus, lobis linearibus; thecae base 
non vel breviter appendiculatae; appendices 
apicales antherarum glabrae in parietibus 
cellularum tenuis; basi stylorum noduliferi, 
papillae ramorum aciculiformes. Achenia 
4-5-costata dense setulifera et idioblasti- 
fera, raphidis elongatis; carpopodia anguste 
cylindrica; setae pappi albae barbatae ten- 
ues subpersistentes, seriebus exteriores 
breviter lanceolatis. Grana pollinis non lo- 
phata tricolporata echinata. Numerus chro- 
mosomatum N = 9, 10 (Jones 1942). 

Chemistry reported for the genus in- 



230 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



eludes germacranolides, hirsutanolides, 
quaianolides and bisabolene derivatives 
(Bohlmann & Jakupovic 1990). Most chro- 
mosome counts have reported N = 10. 

Hilliardiella is closely related to Cyan- 
thillium, but it seems consistently different 
by the more perennial habit, the non-lophate 
pollen, and especially the T-shaped hairs of 
the corolla. The group has been rather well 
defined at various levels by Candolle 
(1836) and Jones (1981). 

The name is derived from the subsectional 
name of Jones (1981), but the genus is de- 
scribed as new to avoid any complications 
from spelling. The name, as indicated by 
Jones (1981), honors Dr. OUve M. Hilliard, 
student of the Asteraceae of Natal. The fol- 
lowing eight species are credited to the genus. 

Hilliardiella aristata (DC.) H. Rob., comb, 
nov. basionym: Webbia aristata DC, 
Prodr. 5:73. 1836, not Vemonia aristata 
Less., 1829. 

Vemonia natalensis Sch. Bip. ex Walp., 
Rep. 2:947. 1843. 
South Africa. 

Hilliardiella calyculata (S. Moore) H. 
Rob., comb. nov. basionym: Vernonia 
calyculata S. Moore, J. Linn. Soc. Bot. 
35:316. 1902. 

Congo, Tanzania, Malawi, Mozambique, 
Zambia. 

Hilliardiella hirsuta (Sch. Bip. ex Walp.) 
H. Rob., comb. nov. basionym: Vernonia 
hirsuta Sch. Bip. ex Walp., Rep. 2:947. 
1843. 
South Africa. 

Hilliardiella leopoldii (Vatke) H. Rob., 
comb. nov. basionym: Vemonia leopoldii 
Vatke, Linnaea 39:478. 1878. 
Ethiopia. 

Hilliardiella nudicaulis (DC.) H. Rob., 
comb. nov. basionym: Webbia nudicaulis 
DC, Prodr. 5:73. 1836. 

Vemonia dregeana Sch. Bip. ex Walp., 
Rep. 2:947. 1843. 
South Africa. 

Hilliardiella oligocephala (DC.) H. Rob., 



comb. nov. basionym: Webbia oligoce- 
phala DC, Prodr. 5:73. 1836. 

Webbia elaegnoides DC, Prodr. 5:73. 

1836, not Vernonia elaegnoides 

H.B.K., 1818. 
Vemonia oligocephala (DC) Sch. Bip. 

ex Walp., Rep. 2:947. 1843. 
Vemonia krausii Sch. Bip. ex Walp., 

Rep. 2:947. 1843. 

Tanzania south to the Cape Province. 
Hilliardiella pinifolia (Less.) H. Rob., 
comb. nov. basionym: Vernonia pinifolia 
Less., Linnaea 4:257. 1829. 

Webbia pinifolia (Less.) DC, Prodr. 5:72. 
1836. 
South Africa. 

Hilliardiella smithiana (Less.) H. Rob., 
comb. nov. basionym: Vemonia smithi- 
ana Less., Linnaea 6:638. 1831. 
West and Central Tropical Africa. 

Orbivestus H. Rob., gen. nov. (Erlangei- 
nae). Type: Vemonia karaguensis Oliv. 
& Hiem. 

Vernonia subg. Orbisvestus S. B. Jones, 
Rhodora 83:60. 1981. Type: Vemonia 
karaguensis Oliv. & Hiem. 

Plantae suffiiitescentes vel frutescentes; 
caules pilosi, pilis T-formibus. Folia altema 
breviter petiolata; laminae sparse pilosae. In- 
florescentiae laxe corymbose cymosae. Ca- 
pitula pedunculata; involucra campanulata; 
bracteae 25-30 ca. 4-seriatae persistentes; re- 
ceptacula epaleacea. Flores 8-16 in capitulo; 
corollae purpureae extus non vel perpauce pi- 
losae, piUs raris sub-T-formibus, tubis super- 
ne leniter infundibularibus, faucibus et lobis 
subaequilongis, lobis longe triangularibus; 
thecae antherarum base breviter vel longe 
caudatae; appendices antherarum apicales 
glabrae in parietibus cellularum tenues; basi 
stylorum breviter et distincte noduliferi; pa- 
pillae ramorum aciculiformes. Achenia 4-5- 
costata setulifera multo idioblastifera, raphi- 
dis elongatis; carpopodia anguste cylindrica; 
setae pappi barbellatae facile deciduae, 
squamis exterioribus brevis persistentibus. 
Grana pollinis non lophata tricolporata echin- 



VOLUME 112, NUMBER 1 



231 



ata. Numerus chromosomatum N = 9, 20 
(Mangenot & Mangenot 1962, Mehra et al. 
1965, Jones 1982). 

The genus Orbivestus is distinct from the 
related Cyanthillium by the more shrubby 
habit, the lack of simple hairs on the corolla 
lobes, and the non-lophate pollen. It is dis- 
tinct from Hilliardiella by the longer co- 
rolla throats and more triangular lobes that 
bear few or no hairs. The few hairs seen on 
the corolla lobes in the type species are 
asymmetrically T-shaped, quite different 
from those in Hilliardiella. The genus has 
more prominent tails on the anthers than in 
either of the related genera. Chemical con- 
stituents include 5-alkyl coumarins, bisa- 
bolene derivatives and glaucolides (Bohl- 
mann & Jakupovic 1990). 

Although the subgeneric name by Jones 
(1981) is the inspiration for the generic 
name, the genus has been described as new 
to avoid the consistent extra "s" in the 
Jones spelling. Jones did not explain his 
spelling, and the reason for his choice is 
unknown. The spelling used here is that 
found in Jeffrey (1988) and Bohlmann & 
Jakupovic (1990). Still, it seems unneces- 
sary to completely abandon the name 
coined by Jones. 

The following four species are placed in 
the genus. 

Orbivestus cinerascens (Sch. Bip. in 
Schweinf.) H. Rob., comb. nov. basio- 
nym: Vemonia cinerascens Sch. Bip. in 
Schweinf., Beitr. Fl. Aeth. 162. 1897. 

Vemonia tephrodioides Chiov., Fl. So- 
mal. 2:255. 1932. 

Senegal east to western India, south to 
Angola and Botswana. 

Orbivestus homilanthus (S. Moore). H. 
Rob., comb. nov. basionym: Vernonia 
homilantha S. Moore, J. Bot. 41:138. 
1903. 

Vernonia sennii Chiov., Fl. Somal. 2:256. 
1932. 
Kenya, Somalia. 

Orbivestus karaguensis (Oliv. & Hiern) H. 



Rob., comb. nov. basionym: Vernonia 
karaguensis Oliv. & Hiern, Trans. Linn. 
Soc. London 29:91. 1873. 

Vernonia cistifolia O. Hoffm., Engl. 

Pflanzenw. Ost-Afr. C 404. 1895. 
Vemonia elliotii S. Moore, J. Linn. Soc. 

Bot. 35:315. 1902. 
Vemonia bothrioclinoides C. H. Wright, 

Bull. Misc. Inf. 1906:108. 1906. 
Vernonia porphyrolepis S. Moore, J. Bot. 

46:39. 1908. 
Vemonia campanea S. Moore, J. Bot. 54: 

251. 1916. 
Vemonia melanacrophylla Cufod., Nouv. 

Giom. Bot. Ital., n.s. 50:102. 1943. 

Sudan south to Mozambique and Zim- 
babwe, west to Nigeria. 

Orbivestus undulatus (Oliv. & Hiern) H. 
Rob., comb. nov. basionym: Vernonia 
undulata Oliv. & Hiern, Fl. Trop. Afr. 3: 
276. 1877. 

West and central tropical Africa, north to 
Sudan, south to Angola. 

Oocephala (S. B. Jones) H. Rob., stat. nov. 
(Erlangeinae). 

Vernonia subsect. Oocephalae S. B. 
Jones, Rhodora 83:72. 1981. 

Type species: Vernonia oocephala Baker. 

Low much-branched shrubs to 1 m high, 
stems with weakly L-shaped simple hairs and 
with multiseptate simple hairs. Leaves alter- 
nate, subsessile, linear to elliptical, sometimes 
serrate. Inflorescences corymbose cymes with 
usually shortly pedunculate heads or with 
heads sessile in apical clusters of leaves. In- 
volucre ovoid or cylindrical; bracts 20-40 in 
4-7 gradate series, ovate to oblong, ap- 
pressed; receptacle without pales. Florets ca. 
15 in a head; corollas white or lavender, tu- 
bular to narrowly funnelform, throat as long 
as lobes, tips without hairs or with few short 
biseriate hairs; anther bases rounded, apical 
appendages glabrous, with thin- walled cells; 
style base with indistinct ring; style branches 
with acicular sweeping hairs. Achenes weak- 
ly 8-ribbed, sericeous with many setulae, id- 
ioblasts numerous, raphids narrowly elon- 



232 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



gate; pappus biseriate, outer shorter and 
broader, inner setiform, subplumose, glabrous 
near base. Pollen triporate, lophate, minutely 
papillate on murae, emicropunctate or weakly 
micropunctate. 

The genus is distinquished by subplumose 
inner pappus, triporate pollen and stems with 
weakly L-shaped hairs. Vemonia sect. Ooce- 
phala is based on a single species with pe- 
dunculate ovoid heads containing few tubular 
florets. The additional species here referred 
to Oocephala was described by Jeffrey 
(1988) in his Group 3 subgroup A with Cen- 
trapalus, but the achene is most like Ooce- 
phala and Vemoniastrum, and a small corolla 
remnant shows triporate pollen. The species 
is not related to the Centraplinae, and is 
placed here on the basis of its subplumose 
pappus, in spite of considerable difference in 
general appearance. 

The genus contains the following two 
species. 

Oocephala agrianthoides (C. Jeffrey) H. 
Rob., comb. nov. basionym: Vernonia 
agrianthoides C. Jeffrey, Kew Bull. 43: 
227. 1988. 
Burundi, Congo, Tanzania. 

Oocephala stenocephala (Oliv.) H. Rob., 
comb. nov. basionym: Vemonia steno- 
cephala Oliv. in Hook., Ic. PL 14:35, t. 
1349A. 1881. 

Vernonia oocephala Baker, Bull. Misc. 

Inf. 1895:68. 1895. 
Vemonia luteoalbida De Wild., Repert. 

Spec. Nov. Regni Veg. 13:207. 1913. 

Nigeria east to Tanzania and south to 

Mozambique. 

Polydora Fenzl, Flora 27:312. 1844. Type 
species: Polydora stoechadifolia Fenzl = 
Webbia serratuloides DC. 

Crystallopollen Steetz in Peters, Reise 
Mossamb. Bot. 363. 1864. Type spe- 
cies: Crystallopollen angustifolium 
Steetz. 

Mostly annuals; stems with one-armed T- 
shaped hairs. Leaves alternate. Inflores- 
cence a lax thyrsoid panicle with corym- 



bosely cymose branches with pedunculate 
heads or with single terminal head. Invo- 
lucral bracts ca. 80 in ca. 7 series, often 
with widely scarious margins and awned 
often black tips, receptacles epaleaceous. 
Florets ca. 30 in a head; corollas whitish to 
purplish, tube longly funnelform, throat as 
long as the narrow glabrous lobes; anther 
bases plain, not tailed; apical appendage 
glabrous, with cell walls thin, sometimes 
weakly ornate; style base with distinct an- 
nular node; branches with acicular sweep- 
ing hairs. Achenes 5 or 8-10 ribbed, setu- 
liferous, raphids narrowly elongate; pappus 
with copious barbellate setae, greenish, yel- 
lowish or tawny, rarely white; outer pappus 
short, squamiform. Pollen triporate, scarce- 
ly echinolophate to psilolophate, with or 
without micropunctations. Chromosome 
number N = 9 (Jones 1979, 1982). Report- 
ed sesquiterpene lactones are germacranol- 
ides, hirsutanolides and furanhehangolides 
(Bohlmann & Jakupovic 1990). 

The genus is distinct in the annual habit, 
chromosome number of N = 9, the one- 
armed T-shaped hairs, untailed anthers and 
triporate pollen. 

Some of the species of the Vernonia 
chloropappa group are keyed by Pope 
(1986). 

The genus is here credited with the fol- 
lowing eight species. 

Polydora angustifolia (Steetz in Peters) H. 
Rob., comb. nov. basionym: Crystallo- 
pollen angustifolium Steetz in Peters, Re- 
ise Mossamb. Bot. 366. 1864, not Ver- 
nonia angustifolia Michx., 1803 or V. an- 
gustifolia D. Don ex Hook. & Am., 1835. 

Vernonia erinacea H. Wild, Kirkia 11:2. 
1978. 

Tanzania and Mozambique, east to 
Zambia and Zimbabwe. 

Polydora bainesii (Oliv. & Hiem) H. Rob., 
comb. nov. basionym: Vernonia bainesii 
Oliv. & Hiem in OUv., R. Trop. Afr. 3: 

272. 1877. 

Tanzania and Mozambique, east to Zam- 
bia and Zimbabwe. 



VOLUME 112, NUMBER 1 



233 



Poly dor a chloropappa (Baker) H. Rob., 
comb. nov. basionym: Vemonia chloro- 
pappa Baker, Bull. Misc. Inf. 1898:146. 
1898. 

Vemonia kassneri De Wild. & Muschl., 
Bull. Soc. Bot. Belg. 49:242. 1912. 

Vemonia smaragdopappa S. Moore, J. 
Linn. Soc. Bot. 47:284. 1925. 
Congo, Malawi, Tanzania, Zambia. 

Polydora jelfiae (S. Moore) H. Rob., comb, 
nov. basionym: Vemonia jelfiae S. 
Moore, J. Linn. Soc. Bot. 47:262. 1925. 
Angola, Burundi, Congo, Malawi, Tan- 
zania, Zambia, Zimbabwe. 

Polydora poskeana (Vatke & Hildebr.) H. 
Rob., comb. nov. basionym: Vemonia 
poskeana Vatke & Hildebr., Oesterr. Bot. 
Zeitschr. 25:324. 1875. 
Angola, Botswana, Nambia, Transvaal, 
Zimbabwe. 

Polydora serratuloides (DC.) H. Rob., 
comb. nov. basionym: Webbia serratu- 
loides DC, Prodr. 5:72. 1836, not Ver- 
nonia serratuloides H. B. K., 1818. 

Vemonia perrottetii Sch. Bip. ex Walp., 

Rep. 2:947. 1843. 
Polydora stoechadifolia Fenzl, Flora 27: 

312. 1844. 

West and central tropical Africa from 

Sudan and Ethiopia south to Angola 

and Zambia. 

Polydora steetziana (Oliv. & Hiem) H. 
Rob., comb. nov. basionym: Vemonia 
steetziana Oliv. & Hiem in Oliv., Fl. 
Trop Afr. 3:278. 1877. 
South Africa. 

Polydora sylvicola (G. V. Pope) H. Rob., 
comb. nov. basionym: Vemonia sylvicola 
G. V. Pope., Kew Bull. 41:395. 1986. 
Angola, Congo, Malawi, Mozambique, 
Tanzania, Zambia, Zimbabwe. 

Vernoniastrum H. Rob., gen. nov. (Erlan- 
geinae) Type: Crystallopollen latifolium 
Steetz in Peters 

Vemonia sect. Lepidella Oliv. & Hiern, 
Fl. Trop. Afr. 3:267. 1877. Type: Ver- 
nonia petersii Oliv. & Hiem, not Lep- 



idella Tiegh., 1912 or Lepidella E. J. 
Gilbert, 1925. 
Vemonia subsect. Lepidella (Oliv. & 
Hiem) S. B. Jones, Rhodora 83:72. 
1981. 

Plantae herbaceae perennes 0.3-1.0 m al- 
tae; caules pilosi, pilis simpUces in ceUuUs 
apicalibus elongatis base leniter inaequalibus. 
FoUa altema. Inflorescentiae uni- vel multo 
capitatae. Involucra campanulata; bracteae ca. 
50 in ca. 3 series gradatae persistentes; re- 
ceptacula epaleacea. Flores ca. 50 in capitulo; 
coroUae rubro-purpurascentes, tubis anguste 
infundibularibus, faucibus quoad lobis et the- 
cis brevioribus, lobis distaUter pilosis; thecae 
antherarum base acuminate vel acute cauda- 
tae; appendices antherarum apicales glabrae 
in parietibus ceUarum tenues; base stylorum 
noduHferi; rami stylorum aciculariter piHferi. 
Achenia 4-5-angulata, idioblastis saepe in 
seriebus transversahbus aggregatis, raphidis 
elongatis; setae pappi interiores margine 
dense barbellatae subpersistentes, squamae 
exteriores lateriores persistentes. Grana pol- 
Unis triporata lophata micropunctata vel em- 
icropunctata. Numerus chromosomatum N = 
10 (Jones 1979, 1982). 

Vernoniastrum seems closely related to 
the foregoing Polydora Fenzl, but differs by 
the perennial habit, the non-T-shaped hairs, 
the tailed anther bases and the chromosome 
number of N = 10. The core element of the 
genus also has the idioblasts of the achene 
in distinct transverse bands, a feature not 
seen outside of the genus Vernoniastrum. 

The genus is credited here with the fol- 
lowing eight species: 

Vernoniastrum aemulans (Vatke) H. Rob., 
comb. nov. basionym: Vemonia aemu- 
lans Vatke, Oesterr. Bot. Zeitschr. 27:195. 

1877. 

Kenya, Tanzania. 

Vernoniastrum ambiguum (Kotschy & 
Peyr.) H. Rob., comb. nov. basionym: 
Vemonia ambigua Kotschy & Peyr, PI. 
Tinn., 35, t. 17B. 1867. 
West tropical Africa to Sudan and Tan- 
zania and south to Angola. 



234 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Vernoniastrum latifolium (Steetz in Peters) 
H. Rob., comb. nov. basionym: Crystal- 
lopollen latifolium Steetz in Peters, Reise 
Mossamb. Bot. 364, t. 48a. 1864, not V. 
latifolia Lem., 1855. 

Vernonia petersii Oliv. & Hiem, Trans. 
Linn. Soc. London 29:90. 1873. 

Vernonia eriocephala Klatt, Bull. Herb. 
Boiss. 4:826. 1896. 
Angola and Congo east to Mozam- 
bique and Tanzania. 

Vernoniastrum musofense (S. Moore) H. 
Rob., comb. nov. basionym: Vernonia 
musofensis S. Moore, J. Bot. 56:206. 
1918. 

Vernonia lappoides O. Hoffm., Bol. Soc. 
Brot. 13:19. 1896, hom. illeg., not 
Baker, 1873. 

Vernonia miamensis S. Moore, J. Bot. 
64:304. 1926. 

Vernonia hoffinanniana Hutsch. & Dalz., 
Fl. W. Trop. Afr. 2:167. 1931, nom. 
nov. for V. lappoides O. Hoffm. 

Vernonia philipsoniana Lawalree, Expl. 
Hydrobiol. Lac. Tanganyika (1946— 
47) Res. Sc. 4, 2:59. 1955; nom. nov. 
superfl. for V. lappoides O. Hoffm. 
Tropical Africa from Nigeria to Tan- 
zania, south to Angola and Zimbabwe. 

Vernoniastrum nestor (S. Moore) H. Rob., 
comb. nov. basionym: Vernonia nestor S. 
Moore, J. Linn. Soc. Bot. 35:317. 1902. 

Vernonia chariensis O. Hoffm., Bull. 

Soc. Bot. France 55, mem. 8:40. 1908. 
Vernonia cannabina MuschL, Bot. Jahrb. 

Syst. 46:94. 1911. 

West Africa east to Tanzania south to 

Natal. 

Vernoniastrum ugandense (S. Moore) H. 
Rob., comb. nov. basionym: Vernonia 
ugandensis S. Moore, J. Linn. Soc. Bot. 
35:314. 1902. 

Vernonia caput-medusae S. Moore, J. 

Linn. Soc. Bot. 37:166. 1905. 
Vernonia fontinalis S. Moore, J. Bot. 52: 

90. 1914. 



Vernonia punctulata S. Moore, J. Linn. 

Soc. Bot. 47:262. 1925. 
Vernonia proclivicola S. Moore, J. Linn. 

Soc. Bot. 47:262. 1925. 
Vernonia mgetae Gilli, Ann. Naturhist. 

Mus. Wien 78:164. 1974. 

Congo, Burundi and Tanzania south to 

Angola, Zambia and Mozambique. 

Vernoniastrum uncinatum (Oliv. & Hiem 
ex Oliv.) H. Rob., comb. nov. basionym: 
Vernonia uncinata Oliv. & Hiem ex 
Oliv., Fl. Trop Afr. 3:277. 1877. 

Vernonia amplexicaulis Baker, Kew Bull. 
1895:216. 1895. 
Ethiopia, Kenya, Somalia. 

Vernoniastrum viatorum (S. Moore) H. 
Rob., comb. nov. basionym: Vernonia 
viatorum S. Moore, J. Linn. Soc. Bot. 35: 
315. 1912. 
Malawi. 

Koyamasia H. Rob., gen. nov. (Erlangei- 
nae). Type: Camchaya calcarea Kitam. 

Plantae herbaceae perennes ad 0.5 m al- 
tae pauce ramosae; caules pilis simplicibus 
multiseptatis obsiti. Foha altema anguste pe- 
tiolata. Inflorescentiae terminales et axiUares. 
Capitula longe pedunculata late campanulata; 
bracteae involucri ca. 90 ca. 4-5-seriatae per- 
sistentes in partibus majoribus herbaceae et 
reflexae; receptaculm glabrum. Flores ca. 90 
in capitulo; coroUae tenuiter camosae, tubis 
angustis, faucibus abrupte campanulatis, lobis 
quoad faucibus leniter longioribus non reflex- 
is extus glanduHferis; thecae antherarum val- 
de exertae saepe nigricans base rotundatae, 
ceUuHs endotheciahbus elongatis, in apicibus 
noduHferis; appendices apicales antherarum 
non longiores quam lateriores glabrae in par- 
ietibus ceUularum firmis; basi stylorum non 
noduliferi. Achenia 10-costata glabra; raphi- 
dis minutis anguste oblongis; setae pappi 
paucae breves facile deciduae. Grana poUinis 
triporata emicropunctata. Numerus chromo- 
somatum 2N = 54 (Koyama 1984). 

The genus Koyamasia is estabUshed for a 
single Southeast Asian species occurring in 
Umestone areas. It was originally described 



VOLUME 112, NUMBER 1 



235 



in Camchaya Gagnep., but was excluded 
from it by Koyama (1984). The species is 
similar to Camchaya in its geography and re- 
duced pappus, but it differs in the triporate 
rather than tricolporate lophate pollen and the 
simple rather than T-shaped hairs. The genus 
may be more closely related to Kinghamia C. 
Jeffrey, which contrary to its original descrip- 
tion, has triporate rather than tricolporate pol- 
len, and is not closely related to Linzia. The 
West African Kinghamia is similar, but is a 
less robust plant with narrower leaf blades 
and much smaller heads. Kinghamia may 
represent convergent evolution in many of its 
similarities, but it seems to differ from Koy- 
amasia by only some technical characteristics 
such as the reflexed mature corolla lobe tips, 
the presence of a basal stylar node, the pale 
and only partially exerted anthers, the longer 
anther appendages and the subquadrate en- 
dothecial cells without nodular thickenings. 

The single species is as follows: 
Koyamasia calcarea (Kitam.) H. Rob., 
comb. nov. basionym: Camchaya calca- 
rea Kitam., Acta Phytotax. Geobot. 23: 
71. 1968. 
Thailand. 
Phyllocephalum Blume, Bidjr. Fl. Ned. Ind. 
888. 1826. Type: Phyllocephalum frutes- 
cens Blume 

Decaneurum DC. ex Wight, Contr. Bot. 
Ind. 7-8. (1833), not Decaneurum 
DC., 1836. Type: Decaneurum reti- 
culatum DC. ex Wight [= Phylloce- 
phalum indicum (Less.) Kirkman] 

Rolfinkia Zenker, PI. Ind. 13. 1837. Type: 
Rolfinkia centaureoides Zenker [ = 
Phyllocephalum indicum (Less.) Kirk- 
man]. 

Lamprachaenium Benth, in Benth. & 
Hook, f.. Gen. pi. 2:225. (1873). Type: 
Decaneurum microcephalum Dalzell 

The herbaceous genera Phyllocephalum 
Blume of India and Indonesia, with three spe- 
cies (Kingham, 1981), and Lamprachaenium 
Benth. of India, with one species, have been 
examined and found to differ mostly by the 
costate achenes of the former versus the shiny 



dark ecostate achenes of the latter In both 
genera, the achenes are oblong and glabrous, 
somewhat obcompressed, abruptly rounded 
above to the narrow attachment of the corol- 
la, the pappus is of short highly deciduous 
setae and the raphids in the achene wall are 
subquadrate. Both genera have lophate tri- 
porate pollen with minutely spinulose mar- 
gins of the muri as seen in Kirkman (1981) 
and Robinson & Marticorena (1986). Both 
have foHose lower involucral bracts or foUose 
tips on the bracts. The genera are here con- 
sidered the same and the needed combination 
is as follows: 

Phyllocephalum microcephalum (Dalzell) 
H. Rob., comb. nov. basionym: Decaneu- 
rum microcephalum Dalzell, Hooker's J. 
Bot. Kew Gard. Misc. 3:231. 1851. India. 

New Genera and New Combinations of 
Eastern Hemisphere Vemonieae 
Subtribe Centrapalinae H. Rob. 

Centrapalus Cass., Bull. Soc. Philom. Paris 
1817:10. 1817. Type: Centrapalus gala- 
mensis Cass. 

Vernonella Sond., Linnaea 23:62. 1850. 
Type: Vernonella africana Sond. 

Annual or perennial, scapose or subsca- 
pose herbs, with or without brown woolly 
hair on crown of rootstock, anthesis often 
prior to leaf emergence; stem hairs simple, 
multiseptate. Leaves basal or cauline, alter- 
nate, sessile. Inflorescences terminal on 
stems and branches. Involucre hemispheri- 
cal; bracts 125-150 in ca. 5 gradate series, 
linear, green, often with small teeth on dis- 
tal margin; receptacle epaleaceous. Florets 
ca. 100 in a head; corollas light blue to blu- 
ish purple, tube funnelform above, throat 
nearly half as long as thecae, lobes some- 
times fringed with long papillae; anther 
base not tailed; apical anther appendage 
glabrous, often colored, with slightly thick- 
ened cell walls; style base with broad node. 
Achenes weakly 10-costate, setuliferous, 
raphids narrowly oblong; pappus setae 
long, subpersistent, outer series setiform. 



236 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



short. Pollen tricolporate, echinate, lophate 
or non-lophate, without muri projecting into 
colpi, micropunctate. Chromosome number 
N = 9 (Jones 1974, 1979, 1982). Sesqui- 
terpene lactones include elemanolides 
(Bohlmann & Jakupovic 1990). 

The name Centrapalus has been used 
most recently within the broad concept of 
Vemonia for a group of coarse annual or 
perennial African herbs. The genus is of 
some potential commercial importance for 
the extraction of epoxy resins (Ayorinde et 
al., 1990). The proper combination has not 
previously been provided for the type spe- 
cies, and an additional eight species are 
placed in the genus here. 

Centrapalus acrocephalus (Klatt) H. Rob., 
comb. nov. basionym: Vemonia acroce- 
phala Klatt, Ann. Hofmus. Wien 7:100. 
1897. 

Sierra Leone, Nigeria and Congo south 
to Angola and Zimbabwe. 

Centrapalus africanus (Sond.) H. Rob., 
comb. nov. basionym: Vemonella afri- 
cana Sond., Linnaea 23:62. 1850. 
South Africa. 

Centrapalus chthonocephalus (O. Hoffm.) 
H. Rob., comb. nov. basionym: Vemonia 
chthonocephala O. Hoffm., Bol. Soc. 
Brot. 13:17. 1896. 

Sierra Leone and Sudan south to Angola 
and Malawi. 

Centrapalus denudatus (Hutch. & Burtt) 
H. Rob., comb. nov. basionym: Vemonia 
denudata Hutch. & Burtt, Rev. Zool. Bot. 
Afr. 23:37. 1932. 
Congo, Tanzania, Zambia. 

Centrapalus kirkii (Oliv. & Hiem in Oliv.) 
H. Rob., comb. nov. basionym: Vemonia 
kirkii Oliv. & Hiem in Oliv., Fl. Trop. 
Afr. 3:274. 1877. 

Vemonia swynnertonii S. Moore, J. Linn. 
Soc. Bot. 40:107. 1911. 

Vemonia zambesiaca S. Moore, J. Bot. 
55:102. 1917. 

Tanzania south to Zambia and Moz- 
ambique. 



Centrapalus pauciflorus (Willd.) H. Rob., 
comb. nov. basionym: Conyza pauciflora 
Willd., Sp. PI. 3:1927. 1803. 

Centrapalus galamensis Cass., Diet. Sci. 
Nat., ed. 2, 7:383. 1817. 

Vemonia pauciflora (Willd.) Less., Lin- 
naea 4:292. 1829, not (Pursh.) Poir., 
1817. 

Vemonia afromontana R. E. Fr., Acta. 
Hort. Berg. 9:116. 1929. 

Vemonia zemyi Gilli, Ann. Naturhist. 
Mus. Wein 78:165. 1974. 

Vemonia petitiana A. Rich., Tent. Fl. 
Abyss. 1:373. 1848. 
Sudan, Ethiopia, Somalia, Kenya, 
Uganda, Tanzania, Malawi, Zambia, 
Mozambique and West Africa. 

Centrapalus praemorsus (Muschl.) H. 
Rob., comb. nov. basionym: Vemonia 
praemorsa Muschl., Bot. Jahrb. Syst. 46: 
68. 1911. 

Angola, Congo, Malawi, Tanzania, Zam- 
bia. 

Centrapalus purpureus (Sch. Bip. ex 
Walp.) H. Rob., comb. nov. basionym: 
Vemonia purpurea Sch. Bip. ex Walp., 
Rep. 2:946. 1843. 

Vemonia inulifolia Steud. ex Walp., Rep. 

2:946. 1843. 
Vemonia jaceoides A. Rich., Tent. Fl. 

Abyss. 1:376. 1848. 
Vemonia rigorata S. Moore, J. Bot. 41: 

155. 1903. 
Vemonia scabrida C. H. Wright, Bull. 

Misc. Inf. Kew 1906:21. 1906. 
Vemnonia duemmeri S. Moore, J. Bot. 

52:91. 1914. 
Vemonia pascuosa S. Moore, J. Linn. 

Soc. Bot. 47:163. 1925. 
Vemonia keniensis R. E. Fr., Acta Hort. 

Berg. 9:114. 1929. 

West, central and south tropical Africa 

north to Sudan and Ethiopia. 

Centrapalus subaphyllus (Baker) H. Rob., 
comb. nov. basionym: Vemonia suba- 
phylla Baker, Bull. Misc. Inf. Kew 1895: 
290. 1895. 



VOLUME 112, NUMBER 1 



237 



Nigeria, Cameroon, Congo, Tanzania, 
Angola, Malawi, Zambia. 
Linzia Sch. Bip. ex Walp., Rep. 2:948. 
1843. Type: Linzia vemonioides Sch. 
Bip. ex Walp. 

Vemonia sect. Azureae S. B. Jones, Rho- 
dora 83:74. 1981. Type: Linzia glabra 
Steetz in Peters 

Perennial herbs; stems with simple mul- 
tiseptate hairs. Leaves alternate, subsessile 
to short-petiolate. Inflorescences corymbi- 
form cymes or single heads with short to 
long pecuncles. Involucre funnelform to 
campanulate; bracts 50-150 in 5-6 series, 
often pectinate-denticulate along distal mar- 
gins, outer tips often elongate, green and 
recurved; receptacle epaleaceous. Florets 
ca. 20-50 in a head; corollas bluish, tube 
very long, funnelform near throat, throat 
very short, lobes apically stiffly hairy; an- 
ther base rounded; apical anther appendage 
glabrous, triangular, with thickened orna- 
mentation in center; style base with small 
annulus. Achenes strongly 10-costate, usu- 
ally with rows of idioblasts along sides of 
costae, setuliferous, raphids subquadrate to 
short-oblong; pappus of many somewhat 
persistent, long bristles, with short outer se- 
ries. Pollen tricolporate, lophate, with muri 
intruding into short colpi, single polar la- 
cunae often present, with or without micro- 
punctations. Chromosome number N = 10 
(Jones 1979, 1982). Sesquiterpene lactones 
include germacranolides and hirsutanolides 
(Bohlmann & Jalupovic 1990). 

Linzia has been recognized rather accu- 
rately in recent literature because of the rather 
characteristic bluish flowers (Jones 1981) or 
distinctive pollen (Jeffrey 1988). The species 
are related to Centrapalus, but they differ by 
the more perennial habit, the very short 
throats of the coroUas, the stronger ribs on 
the achenes, the idioblasts that are positioned 
along those ribs in most species and the chro- 
mosome number of N = 10. 

The following seven species are credited 
to the genus. 
Linzia gerberiformis (Oliv. & Hiem in 



Oliv.) H. Rob., comb. nov. basionym: 
Vernonia gerberiformis Oliv. & Hiern in 
Oliv., Fl. Trop. Afr. 3:285. 1877. 

Vernonia macrocyanus O. Hoffm., Bol. 
Soc. Brot. 13:20. 1896. 

Vernonia nandensis S. Moore, J. Linn. 
Soc. Bot. 35:323. 1902. 

Vernonia towaensis De Wild., Bull. Jard. 
Bot. Brux. 5:96. 1915. 
Angola, Burundi, Cameroon, Congo, 
Malawi, Nigeria, Sudan, Tanzania, 
Uganda, Zambia, Zimbabwe. 

Linzia glabra Steetz in Peters, Reise Mos- 
samb. Bot. 353. 1864. 

Vemonia glabra (Steetz) Vatke, Oesterr. 
Bot. Zeitschr. 27:194. 1877. 

Vernonia obconica Oliv. & Hiem in 
Oliv., Fl. Trop. Afr. 3:286. 1877. 

Vemonia pogosperma Klatt, Ann. K.K. 
Naturhist. Hofmus. 7:99. 1892. 

Vemonia hindei S. Moore, J. Bot. 41: 
155. 1903. 

Vemonia piovanii Chiov., Race. Bot. 
Miss. Cons. Kenya 61. 1935. 

Vemonia roseopapposa Gilli, Ann. Na- 
turhist. Mus. Wien 78:165. 1974. 
Congo, Burundi, Kenya, Tanzania, 
south to Angola, Namibia, Mozam- 
bique, Transvaal, Swaziland, Natal, 
Madagascar. 

Linzia infundibulariformis (Oliv. & Hiem 
in Oliv.) H. Rob., comb. nov. basionym: 
Vernonia infundibulariformis Oliv. & 
Hiem in Oliv., H. Trop. Afr. 3:285. 1877. 

Vernonia saussureoides Hutch., Bull. 
Misc. Inf. Kew 1921:378. 1921. 
Burundi, Congo, Cameroon, Nigeria, 
Sudan, Tanzania, Uganda. 

Linzia ituriensis (Muschl.) H. Rob., comb, 
nov. basionym: Vernonia ituriensis 
Muschl., Wiss. Ergebn. Deutsch. Zent.- 
Afr. Exped. 1907-8 (2):364. 1911. 

Vernonia hi I Hi Hutch. & Dalz., Fl. W 
Trop. Afr. ed. 1, 2:165, 168. 1931. 

Vernonia muhiensis Kalanda, Bull. Jard. 
Bot. Nat. Belg. 52:125. 1982. 



238 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Burundi, Cameroon, Congo, Ethiopia, 
Nigeria, Rwanda, Sudan, Tanzania. 

Linzia melleri (Oliv. & Hiem in Oliv.) H. 
Rob., comb. nov. basionym: Vernonia 
melleri Oliv. & Hiem in Oliv., Fl. Trop. 
Afr. 3:282. 1877. 

Vernonia superba O. Hoffm., Engl. 
Pflanzenw. Ost-Afr. C. 406. 1895. 

Vernonia scabrifolia O. Hoffm., Bot. 
Jahrb. Syst. 30:424. 1901, nom. illeg., 
not Hieron., 1897. 

Vernonia paludigena S. Moore, J. Bot. 
52:91. 1914. 

Vernonia vanmeelii Lawalree, Expl. Hy- 
drobiol. Lac Tanganyika. Res. Sc. 6, 2: 
59. 1955, nom. nov. for V. scabrifolia 
O. Hoffm. 

Angola, Congo, Malawi, Mozam- 
bique, Rwanda, Tanzania, Zambia, 
Zimbabwe. 

Linzia vemonioides Sch. Bip. ex Walp., 
Rep. 2:948. 1843. 

Vernonia quartiniana A. Rich., Tent. Fl. 

Abyss. 1:379. 1848. 
Vernonia congolensis De Wild. & 

Muschl., Bull. Soc. Bot. Belg. 49:237. 

1912. 
Vernonia vernonioides (Sch. Bip. ex 

Walp.) Cufod., Bull. Jard. Bot. Brux. 

36, suppl.:1078. 1966, hom. illeg., not 

(A. Gray) Bacigal, 1931. 

Burundi, Congo, Ethiopia, Tanzania. 

Linzia usafuensis (O. Hoffm.) H. Rob., 
comb. nov. basionym: Vernonia usafuen- 
sis O. Hoffm., Bot. Jahrb. Syst. 30:425. 
1901. 

Vernonia candelabricephala Gilli, Ann. 
Namrhist. Mus. Wien 78:161. 1974. 
Tanzania. 

New Genera and New Combinations of 

Eastern Hemisphere Vemonieae 

Subtribe Gymnantheminae H. Rob. 

Distephanus Cass., Bull. Soc. Philom. Paris 
1817. 151. 1817. Type: Conyza populi- 
folia Lam. 



The genus characterization, synonymy, 
and many of the species are given by Rob- 
inson & Kahn (1986), who stressed the yel- 
low flowers and trinervate leaves as impor- 
tant generic characteristics. The type spe- 
cies has been redescribed and illustrated 
particularly well by Hind (1996), and it is 
suggested in Scott (1993) and Hind (1996) 
that the number of species in the genus 
should be nearer to 40 instead of the 26 
transferred by Robinson & Kahn (1986). 
During the present study only the following 
single additional species is transferred. 

Distephanus henryi (Dunn.) H. Rob., 
comb. nov. basionym: Vernonia henryi 
Dunn., J. Linn. Soc. Bot. 35:500. 1903. 
China. 

Gymnanthemum Cass., Bull. Soc. Philom. 
Paris 1817:10. 1817. Type: G. cupulare 
Cass. = Baccharis senegalensis = Gym- 
nanthemum coloratum (Willd.) H. Rob. 
& BKahn 

Bracheilema R. Br. ex Salt., Abyss. Ap- 
pend. 65. 1814, nom. nud. 

Decaneurum DC, Arch. Bot. (Paris) 2: 
516. 1833, nom. superfl. Type: as in 
Gymnanthemum. 

Monosis DC. in Wight, Contrib. Bot. Ind. 
5. 1834. Type: Monosis wightiana DC. 
in Wight [= Vernonia arborea Buch.- 
Ham.]. 

Vernonia sect. Strobocalyx Blume ex 
DC, Prodr. 5:21. 1836. Type: Vernon- 
ia arborea Buch.-Ham. 

Plectreca Rafin., Fl. Tellur. 4:119. 1838 
("1836"). Type: Staehelina corymbo- 
sa Thunb. 

Keringa Rafin., Sylva Tellur. 144. 1838. 
Type: Vernonia amygdalina Del. 

Cheliusia Sch. Bip., Flora 24, 1. Intell. 
26. 1841. Type: Cheliusia abyssinica 
Sch. Bip. = Gymnanthemum amyg- 
dalinum (Del.) Sch. Bip. ex Walp. 

Strobocalyx (Bl. ex DC.) Spach, Hist. 
Nat. Veg. Phan. 10:39. 1841. 

Punduana Steetz in Peters, Reise Mos- 
samb. Bot. 345. 1864. Type: P. vol- 
kameriifolia (DC) Steetz 



VOLUME 112, NUMBER 1 



239 



Vernonia subsect. Urceolata S. B. Jones, 
Rhodora 83:67. 1981. Type: Vernonia 
sphaerocalyx O. Hoffm. 

Shrubs or trees, moderately to densely 
branching; hairs often forming felt, with 
large often contorted cap cells basally or 
nearly basally attached. Leaves alternate, 
with short or winged petioles to rather long- 
petiolate. Inflorescences terminal on stems 
and branches, densely corymbiform. Heads 
with involucres campanulate to ovoid; 
bracts appressed, coriaceous to subcoria- 
ceous, 25-35 in 4-5 series, inner bracts per- 
sistent or easily deciduous; receptacle epa- 
leaceous. Florets 1-50; corollas white to vi- 
olet; tube cylindrical, throat longer than the- 
cae or very deeply cut, lobes with glands 
or spicules; anther base broadly tailed, often 
long; apical anther appendages glabrous, 
with rather thick- walled cells; style with or 
without node. Achenes 5-10-costate, with 
or without setulae or uniseriate hairs, ra- 
phids short to elongate or lacking; pappus 
of many rather persistent capillary bristles, 
often broad-tipped, with short outer squa- 
mellae. Pollen grains tricolporate, non-lo- 
phate or rarely lophate, echinate, micro- 
punctate. Chromosome numbers N = 7?, 
10, 20, 2N = 30 (Jones 1979, 1982). Ses- 
quiterpene lactones include elemanolides 
(Bohlmann & Jakupovic 1990). 

A broad interpretation of the Gymnanthe- 
mum is accepted here. The type species lacks 
a basal node on the style, but other closely 
related species have such nodes. The number 
of ribs on the achene varies from 5 to 10 
within many subgroups. At least some of the 
species with persistent inner involucral bracts 
seem very closely related to species in which 
they are deciduous. The leaves also seem to 
vary in size, texmre and margin. The corolla 
throat is usually long, but a number of species 
in Madagascar have sinuses cut nearly to the 
base of the throat. The corolla lobes usually 
show a characteristic long-triangular erect 
form, but a few species have long narrow 
lobes that are rolled back with age. The 
sweeping hairs are usually blunt, but those of 



G. amygdalinum are more pointed. The pol- 
len is almost always non-lophate or Type A, 
but rarely, as in the type of Punduana, Ver- 
nonia volkameriaefolia, the pollen is lophate. 
The genus is hmited within the subtribe in 
the present paper by only the most obvious 
differences in inflorescence shape, involucral 
bracts, leaf nervation, corolla symmetry and 
color, and lack of pales on the receptacle. 

The genus is credited here with the fol- 
lowing 43 species: 

Gymnanthemum amygdalinum (Del.) Sch. 
Bip. ex Walp., Rep. 2:948. 1843. 

Vernonia amygdalina Del., Cent. PI. Afr. 

Voy. Meroe41. 1826. 
Gymnanthemum abyssinicum Sch. Bip. 

ex Walp., Rep. 2:948. 1843. 
Vernonia vogeliana Benth. in Hook., Ni- 
ger Fl. 427. 1849. 
Vernonia condensata Baker., J. Bot. 8: 

202. 1875. 
Vernonia eritreana Klatt, Bull. Herb. 

Boiss. 4:826. 1896. 
Vernonia randii S. Moore, J. Bot. 37:369. 

1899. 
Vernonia giorgii De Wild., Bull. Jard. 

Bot. Brux. 5:92. 1915. 
Vernonia bahiensis Toledo, Arq. Bot. Es- 

tado Sao Paulo, n.s. 1:52. 1939. 
Vemonanthura condensata (Baker) H. 

Rob., Phytologia 73:69. 1992. 

Yemen and Ethiopia, South Uganda, 

Kenya and Tanzania, Brazil, widely 

cultivated. 

Gymnanthemum anceps (C. B. Clarke ex 
Hook.f.) H. Rob., comb. nov. basionym: 
Vernonia anceps C. B. Clarke ex Hook, 
f., Fl. Brit. Ind. 3:233. 1881. 
Sri Lanka. 

Gymnanthemum andersonii (C. B. Clarke) 
H. Rob., comb. nov. basionym: Vernonia 
andersonii C. B. Clarke, Comp. Ind. 27. 
1852. 

Vernonia chevalieri Gagnap., Bull. Mus. 
Hist. Nat. Paris 25:488. 1919, horn, il- 
leg., not O. Hoffm., 1908. 
Assam, Myanmar, China. 



240 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Gymnanthemum andrangovalense (Hum- 
bert) H. Rob., comb. nov. basionym: Ver- 
nonia andrangovalensis Humbert, Notul. 
Syst. (Paris) 13:311. 1948. 
Madagascar. 

Gymnanthemum antanalus (Humbert) H. 
Rob., comb. nov. basionym: Vernonia 
antanala Humbert, Notul. Syst. (Paris) 
13:314. 1948, in part., emend., Humbert, 
Not. Syst. (Paris). 15:361. 1959. 
Madagascar. 

Gymnanthemum appendiculatum (Less.) 
H. Rob., comb. nov. basionym: Vernonia 
appendiculata Less., Linnaea 6:636. 
1831. 
Madagascar. 

Gymnanthemum arboreum (Buch.-Ham.) 
H. Rob., comb. nov. basionym: Vernonia 
arborea Buch.-Ham., Trans. Linn. Soc. 
London 14:218. 1824. 

Monosis wightiana DC. in Wight, Con- 

trib. Bot. Ind. 5. 1834, not Vernonia 

wightiana Am. 
Vernonia celebrica DC, Prodr. 5:21. 

1836. 
Vernonia javanica DC, Prodr. 5:22. 

1836. 
Vernonia blumeana DC, Prodr. 5:22. 

1836. 
Strobocalyx arborea (Buch.-Ham.) Sch. 

Bip., Jahres. Pollichia 18-19:171. 

1861. 
Vernonia monosis Benth. ex C B. 

Clarke, Comp. Ind. 24. 1852, nom. 

nov. for Monosis wightiana DC in 

Wight, 1834. 
Vernonia vaniotii Levi., Repert. Spec. 

Nov. Regni Veg. 12:531. 1913. 

SE. Asia, Indonesia. 

Gymnanthemum baronii (Baker) H. Rob., 
comb. nov. basionym: Vernonia baronii 
Baker, J. Linn. Soc. Bot. 20:173. 1885. 

Vernonia campenoni Drake, Bull. Soc. 
Bot. France 46:241. 1899. 
Madagascar. 

Gymnanthemum bellinghamii (S. Moore) 
H. Rob., comb. nov. basionym: Vernonia 



bellinghamii S. Moore, J. Bot. 38:155. 
May 1900. 

Vernonia goetzeana O. Hoffm., Bot. 
Jahrb. Syst. 28:503. July 1900. 

Vernonia elizabethvilleana De Wild., Re- 
pert. Spec. Nov. Regni Veg. 13:20. 
1941. 

Congo and Tanzania south to Moz- 
ambique. 

Gymnanthemum bockianum (Diels) H. 
Rob., comb. nov. basionym: Vernonia 
bockiana Diels, Bot. Jahrb. Syst. 29:608. 
1900. 

Pluchea rubicunda Schneid. in Sarg., pi. 
Wils. 3:418. 1916. China. 

Gymnanthemum chapelieri (Drake) H. 
Rob., comb. nov. basionym: Vernonia 
chapelieri Drake, Bull. Soc. Bot. France 
46:234. 1899. 
Madagascar. 

Gymnanthemum coloratum (Willd.) H. 
Rob. & B. Kahn, Proc. Biol. Soc. Wash. 
99:501. 1986. 

Eupatorium coloratum Willd., Sp. PI. 3: 

1768. 1803. 
Baccharis senegalensis Pers., Syn. PI. 2: 

424. 1807. 
Gymnanthemum cupulare Cass., Diet. Sc. 

Nat., ed. 2, 20: 109. 1821. 
Vernonia senegalensis (Pers.) Less., Lin- 
naea 4:265. 1829. 
Decaneurum grande DC, Prodr. 5:67. 

1836. 
Decaneurum senegalense (Pers.) DC, 

Prodr. 5:68. 1836. 
Gymnanthemum grande (DC) Sch. Bip. 

ex Walp., Rep. 2:948. 1843. 
Gymnanthemum senegalense (Pers.) Sch. 

Bip. ex Walp., Rep. 2:948. 1843. 
Gymnanthemum quercifolium Steetz in 

Peters, Reise Mossamb. Bot. 334. 

1864. 
Vernonia oxyura O. Hoffm., Engl. Pflan- 

zenw. Ost.-Afr. C 403. 1895. 
Vernonia polyura O. Hoffm., Bot. Jahrb. 

Syst. 30:422. 1901. 



VOLUME 112, NUMBER 1 



241 



Vernonia cirrhifera S. Moore, J. Linn. 

Soc. Bot. 35:320. 1902. 
Vernonia longipetiolata Muschl., Bot. 

Jahrb. Syst. 46:74. 1911. 
Vernonia aldabrensis Hemsl., J. Bot. 54, 

suppl. 2:20. 1916. 
Vernonia grandis (DC.) Humb., Fl. Mad- 

ag. 189:44. 1960. 

West tropical Africa east to Kenya, 

south to Mozambique and Madagascar. 

Gymnanthemum corymbosum (Thunb.) H. 
Rob., comb. nov. basionym: Staehelina 
corymbosa Thunb., Prodr. pi. Cap. 2:143. 
1800. 

Vernonia corymbosa (Thunb.) Less., Lin- 

naea 6:647. 1831. 
Plectreca corymbosa (Thunb.) Rafin., Fl. 

Tellur. 4:119. 1838 ("1836"). 

South Africa. 

Gymnanthemum coursii (Humbert) H. 
Rob., comb. nov. basionym: Vernonia 
coursii Humbert, Notul. Syst. (Paris) 13: 
310. 1948. 
Madagascar. 

Gymnanthemum crataegifolium (Hutch.) 
H. Rob., comb. nov. basionym: Vernonia 
crataegifolia Hutch., Bull. Misc. Inf. 
Kew 7:330. 1912. 
South Africa. 

Gymnanthemum cumingianum (Benth. in 
Hook.f.) H. Rob., comb. nov. basionym: 
Vernonia cumingiana Benth., Hooker's J. 
Bot. Kew Gard. Misc. 4:232. 1852. 
China, Philippines. 

Gymnantheum cylindriceps (C. B. Clarke) 
H. Rob., comb. nov. basionym: Vernonia 
cylindriceps C. B. Clarke, J. Linn. Soc. 
Bot. 25:35. 1880. 
India, Nepal. 

Gymnanthemum dissolutum (Baker) H. 
Rob., comb. nov. basionym: Vernonia 
dissoluta Baker, J. Linn. Soc. Bot. 20: 

174. 1883. 

Vernonia sparsiflora Baker, J. Linn. Soc. 

Bot. 20:172. 1883. 
Vernonia capreaefolia Baker, J. Linn. 



Soc. Bot. 22:487. 1887. 
Madagascar. 

Gymnanthemum esculentum (Hemsl. ex F 
B. Forbes & Hemsl.) H. Rob., comb. nov. 
basionym: Vernonia esculenta Hemsl. ex 
F B. Forbes & Hemsl., J. Linn. Soc. Bot. 
23:401. 1888. 

Vernonia papillosa Franch., J. Bot. 10: 

368. 1896. 
Vernonia arbor Levi., Repert. Spec. Nov. 

Regni Veg. 11:304. 1912. 

China. 

Gymnanthemum exsertiflorum (Baker) H. 
Rob., comb. nov. basionym: Vernonia 
exsertiflora Baker, Bull. Misc. Inf. Kew 
1898:147. July 1898. 

Vernonia kreismannii Welw. ex Hiem, 
Cat. Afr. PI. Welw. 1, 3:517. Dec 
1898. 

Vernonia sphaerocalyx O. Hoffm., Bot. 
Jahrb. Syst. 30:423. 1901. 
Angola and Congo to Malawi and Tan- 
zania. 

Gymnanthemum exsertum (Baker) H. 
Rob., comb. nov. basionym: Vernonia 
exserta Baker, J. Linn. Soc. Bot. 22:488. 
1887. 

Vernonia grisea Baker, J. Linn. Soc. Bot. 

22:488. 1887, hom. illeg., not Baker, 

1873. 
Vernonia trichodesma Baker, J. Linn. 

Soc. Bot. 25:325. 1890. 
Vernonia lantziana Drake, Bull. Soc. 

Bot. France 46:235. 1899. 

Madagascar. 

Gymnanthemum glaberrimum (Welw. ex 
O. Hoffm.) H. Rob., comb. nov. basio- 
nym: Vernonia glaberrima Welw. ex O. 
Hoffm., Bol. Soc. Brot. 13:15. Sept 1896. 

Vernonia hensii Klatt, Bull. Herb. Boiss. 

4:828. Dec. 1896. 
Vernonia mashonica N. E. Br., Bull. 

Misc. Inf. Kew 1906:108. 1906. 

West, central and south tropical Africa 

north to Sudan. 

Gymnanthemum hildebrandtii (Vatke) H. 



242 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Rob., comb. nov. basionym: Vemonia 
hildebrandtii Vatke, Osterr. Bot. Zeit. 25: 
323. 1875. 

Vemonia taylorii S. Moore, J. Bot. 38: 

154. 1900. 

Kenya, Somalia, Tanzania. 

Gymnanthemum humblotii (Drake) H. 
Rob., comb. nov. basionym: Vernonia 
humblotii Drake, Bull. Soc. Bot. France 
46:235. 1899. 

Vemonia beforonensis Humbert, Notul. 
Syst. (Paris) 13:320. 1948. 
Madagascar. 

Gymnanthemum louvelii (Humbert) H. 
Rob., comb. nov. basionym: Vemonia 
louvelii Humbert, Notul. Syst. (Paris) 15: 
247. 1958. 
Madagascar. 

Gymnanthemum mespilifolium (Less.) H. 
Rob., comb. nov. basionym: Vemonia 
mespilifolia Less., Linnaea 6:641. 1831. 
South Africa. 

Gymnanthemum myrianthum (Hook.f.) H. 
Rob., comb. nov. basionym: Vemonia 
myriantha Hook.f., J. Linn. Soc. Bot. 7: 
198. 1864. 

Vemonia podocoma Sch. Bip. ex Vatke, 

Linnaea 39:476. 1875. 
Vemonia subuligera O. Hoffm., Engl., 

Pflanzenw. Ost.-Afr. C. 403. 1895. 
Vemonia stipulacea Klatt, Bull. Herb. 

Boiss. 4:457. 1896. 
Vemonia lujae De Wild., PL Nov. Herb. 

Hort. Then. 2:119, t. 96. 1900. 
Vemonia ampla O. Hoffm., Bot. Jahrb. 

Syst. 30:423. 1901. 
Vemonia myrianthoides Muschl., Bot. 

Jahrb. Syst. 46:84. 1911. 
Vemonia uhligii Muschl., Bot. Jahrb. 

Syst. 46:79. 1911. 
Vemonia oliveriana Pichi-Serm., Webbia 

7:345. 1950, nom. illegit. superfl. for 

V. podocoma Sch. Bip. ex Vatke. 
Vernonia chlarugii Pich-Serm., Miss. 

Stud. Lago Tana 7, Ricerche Bot. 1: 

155, t. 30. 1951. 

West Africa from Guinea and Sierra 



Leone to Cameroon, Congo, Sudan, 
Ethiopia, Kenya, Tanzania, Uganda to 
south tropical Africa. 

Gymnanthemum obovatum Gaudich., Voy. 
Uran. Bot. 471. 1830. 

Vemonia cuneata Less., Linnaea 6:644. 

1831. 
Vemonia vagans DC, Prodr. 5:32. 1836. 
Vernonia scandens DC, Prodr. 5:32. 

1836. 
Decaneurum obovatum (Gaudich.) DC, 

Prodr. 5:67. 1836. 
Gymnanthemum scandens (DC.) Steetz in 

Peters, Reise Mossamb. Bot. 338. 

1864. 
Gymnanthemum vagans (DC.) Steetz in 

Peters, Reise Mossamb. Bot. 338. 

1864. 
Gymnanthemum ajfine Steetz in Peters, 

Reise Mossamb. Bot. 338. 1864. 
Vernonia obovata (Gaudich.) Boerl., 

Handl. Fl. Ned. Indie 2:1. 1891, not 

Vemonia obovata Less., 1829. 

India, Myanmar, Indonesia. 

Gymnanthemum pectiniforme (DC. in 
Wight) H. Rob., comb. nov. basionym: 
Vemonia pectiniformis DC. in Wight, 
Contr. Bot. Ind. 2. 1834. 

Vemonia puncticulata DC, 7:264. 1838. 
Lysistemma pectiniforme (DC. in Wight) 
Steetz in Peters, Reise Mossamb. Bot. 
343. 1864. 
India, Sri Lanka. 

Gymnanthemum pectorale (Baker) H. 
Rob., comb. nov. basionym: Vemonia 
pectoralis Baker, J. Bot. 20:139. 1882. 
Madagascar. 

Gymnanthemum pleistanthum (Humbert) 
H. Rob., comb. nov. basionym: Vernonia 
secundifolia subsp. pleistantha Humbert, 
Bull. Soc. Bot. France 94:380. 1947. 
Madagascar. 

Gymnanthemum rueppellii (Sch. Bip. ex 
Walp.) H. Rob., comb. nov. basionym: 
Vemonia rueppellii Sch. Bip. ex Walp., 
Rep. 2:946. 1864. 

Vernonia francavillana Oliv. & Hiem in 



VOLUME 112, NUMBER 1 



243 



Oliv., Fl Trop. Afr. 3:296. 1877. 
Ethiopia. 

Gymnanthemum secundifolium (Boj. ex 
DC.) H. Rob., comb. nov. basionym: Ver- 
nonia secundifolia Boj. ex DC, Prodr. 5: 
22. 1836. 

Vemonia quadriflora Baker, J. Linn. Soc. 
Bot. 20:173. 1884. 
Madagascar. 

Gymnanthemum solanifolium (Benth.) H. 
Rob., comb. nov. basionym: Vemonia so- 
lanifolia Benth., Hooker's J. Bot. Kew 
Card. Misc. 1:486. 1842. 
China, Myanmar, Thailand, Vietnam. 

Gymnanthemum subcrassulescens (Hum- 
bert) H. Rob., comb. nov. basionym: Ver- 
nonia subcrassulescens Humbert, Notul. 
Syst. (Paris) 13:309. 1948. 
Madagascar. 

Gymnanthemum theophrastifolium 

(Schweinf. ex Oliv. & Hiem) H. Rob., 
comb. nov. basionym: Vemonia theo- 
phrastifolia Schweinf. ex Oliv. & Hiern, 
in Oliv., Fl. Trop. Afr. 3:294. 1877. 

Vemonia myriocephala A. Rich., Tent. 

Fl. Abyss. 1:374. 1848, hom. illeg., 

not DC, 1836. 
Cacalia richardiana O. Kuntze, Rev. 

Gen. PI. 2:969. 1891, nom. nov. for V. 

myriocephala A. Rich. 
Vemonia seretii De Wild., Ann. Mus. 

Congo V, 11:207. 1907. 
Vernonia macrophylla Chiov., Ann. Bot. 

Roma 9:70. 1911. 
Vernonia richardiana (O. Kuntze) Pichi- 

Serm., Webbia 7:340. 1950. 

Congo and Nigeria east to Uganda, 

Kenya and Ethiopia. 

Gymnanthemum thomsonianum (Oliv. & 
Hiem in Oliv.) H. Rob., comb. nov. bas- 
ionym: Vernonia thomsoniana Oliv. & 
Hiem in Oliv., Trans. Linn. Soc. London 
29:91. 1873. 

Vemonia living stoniana Oliv. & Hiem in 
Oliv., Fl. Trop. Afr. 3:295. 1877. 

Vernonia cruda Klatt, Bull. Herb. Boiss. 
4:456. 1896. 



Vemonia densicapitulata De Wild., Bull. 
Jard. Bot. Bmx. 5:92. 1915. 
West, central and south tropical Africa. 

Gymnanthemum urticifolium (A. Rich.) H. 
Rob., comb. nov. basionym: Vemonia ur- 
ticifolia A. Rich., Tent. Fl. Abyss. 1:378. 
1848. 

Vernonia antinoriana Avetta, Nouv. 

Giom. Bot. Ital. 21:348. 1889. 
Vernonia mellifera Muschl., Bot. Jahrb. 

Syst. 46:90. 1911. 

Congo, Ethiopia, Kenya. 

Gymnanthemum vidalii (G. Merr.) H. Rob., 
comb. nov. basionym: Vemonia vidalii 
G. Merr., Philipp. Is., Bur. Gov. Lab. 
Bull. 6:6. 1904 ("1903"). 
Philippines. 

Gymnanthemum volkameriifolium (DC.) 
H. Rob., comb. nov. basionym: Vernonia 
volkameriaefolia DC, Prodr. 5:32. 1836. 

Vemonia acuminata DC, Prodr. 5:32. 

1836. 
Vemonia punduana DC, Prodr. 5:32. 

1836. 
Vemonia esqirolii Levi., Repert. Spec. 

Nov. Regni Veg. 11:304. 1912. 

China, India, Myanmar. 
Gymnanthemum wightianum (Arn.) H. 
Rob., comb. nov. basionym: Vemonia 
wightiana Am., Pugill. pi. Ind. Or. 27. 
1836. 
Sri Lanka. 

Gymnanthemum zanzibarense (Less.) H. 
Rob., comb. nov. basionym: Vernonia 
zanzibarensis Less., Linnaea 6:637. 
1831. 
Kenya, Tanzania. 

Gymnanthemum zeylanicum (L.) H. Rob., 
comb. nov. basionym: Eupatorium zey- 
lanicum L., Sp. pi. 837. 1753. 

Vemonia zeylanica (L.) Less., Linnaea 4: 
344. 1829. 
Sri Lanka. 

A number of additional species have 
been placed in the genus that are exclud- 
ed here. These include G. congestum 
Cass., = Critoniopsis triflosculosa (H. B. 



244 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



K.) H. Rob. (Robinson 1993), G. frutes- 
cens (Bl.) Sch. Bip. ex Walp., G. molle 
(DC.) Sch. Bip. ex Walp., G. phyllolaen- 
um (DC.) Sch. Bip. ex Walp. and G. re- 
ticulatum (Wight) Sch. Bip. ex Walp. 
which belong to Phyllocephalum (Kirk- 
man 1981). 

Brenandendron H. Rob., gen. nov. (Gym- 
nantheminae). 

Type: Vemonia titanophylla Brenan 

Plantae arborescentes ad 8 m altae me- 
diocriter ramosae; caules velutini, pilis fu- 
siformibus vel leniter asymmetricis. Folia 
altema petiolata vel subsessilia, laminis 
magnis interdum lobatis. Inflorescentiae 
frondifoimes in ramulis racemosae vel spi- 
ciformes. Capitula sessilia vel breviter pe- 
dunculata; involucra campanulata; bracteae 
involucri usque ad 110 in seriebus 7—9 ap- 
pressae interiores leniter deciduae subcori- 
aceae vel coriaceae; receptacula epaleacea. 
Flores 30-50 in capitulo; coroUae pler- 
umque tubiformes in faucibus elongatae, 
lobis erectis vel interdum recur vatis; thecae 
antherarum base longe obtuse caudatae; ap- 
pendices apicales in parietibus cellularum 
firmae; basi stylorum annulate noduliferi. 
Achenia 8— 10-costata glabra idioblastifera, 
raphidis rhomboideis breviter vel mediocri- 
ter elongatis; setae pappi interiores subplan- 
ae subpersistentes exteriores squamiformes 
vel nullae. Grana pollinis tricolporata 
echinata non lophata. Numerus chromoso- 
matum N = 9 Mangenot & Mangenot 
(1957). 

Germacranolides have been reported 
from the genus (Bohlmann and Jakupovic 
1990). 

Brenandendron is closely related to Gym- 
nanthemum, but it differs by the distinctive 
frondiform inflorescence. Brenan (1953) pro- 
vided a key to the three species. In addition 
to the differences given by Brenan, the three 
species differ in their coroUa lobes. Brenan- 
dendron donianum has erect lobes much 
shorter than the corolla throat, B. frondosum 
has narrow erect lobes, and B. titanophyllum 
has strongly recurved narrow lobes. 



The three species are as follows: 

Brenandendron donianum (DC.) H. Rob., 
comb. nov. basionym: Vemonia doniana 
DC, Prodr. 5:23. 1836. 

Vemonia conferta Benth. in Hook., Niger 
Fl. 427. 1849. 

West and central tropical Africa from An- 
gola to Liberia and Sudan. 

Brenandendron frondosum (Oliv. & Hiem 
in Oliv.) H. Rob., comb. nov. basionym: 
Vemonia frondosa Oliv. & Hiem in 
Oliv., Fl. Trop. Afr. 3:294. 1877. 
West Africa from Guinea to Cameroon. 

Brenandendron titanophyllum (Brenan) H. 
Rob., comb. nov. basionym: Vemonia ti- 
tanophylla Brenan, Kew Bull. 1953:116. 
1953. 
Cameroon, Congo, Equatorial Guinea. 

Myanmaria H. Rob., gen. nov. (Gymnan- 
theminae). 

Type: Vemonia calycina DC. 

Plantae frutescentes ca. 1 m altae medi- 
ocriter ramosae; caules puberuli, pilis sim- 
plicibus uniseriatis multiseptatis. Folia al- 
tema breviter petiolata, laminis oblongo- 
o vatis apice obtusis. Inflorescentiae cor- 
ymbiformae. Capitula in ramis terminaUa 
pedunculata; bracteae involucri exteriores 
ca. 20 ca. 2-3-seriatae late ovatae virides 
foliiformes; bracteae interiores lineares acu- 
tae; receptacula epaleacea. Flores 35-50 in 
capitulo; coroUae puberulae, tubis elonga- 
tis, faucibus quoad lobis longioribus, lobis 
2-3 -plo longioribus quam latioribus extus 
glanduliferis et pauce breviter pilosulis; the- 
cae antherarum base distincte late caudatae; 
appendices apicales antherarum distincte 
induratae; basi stylorum non noduliferi; 
rami stylorum obtuse papilliferi. Achenia 
10-costata idioblastifera in costis setulifera, 
raphidis subquadratis; setae pappi barbel- 
latae numerosae 2-3-seriatae persistentes 
exteriores breviores vel nullae. Grana pol- 
linia ca. 50 jjum in diametro tricolporata 
echinata lophata micropunctata, lacunis po- 
laribus triplicibus intercolpatis. 

Myanmaria is endemic to Myanmar. The 
genus is easily distinguished by the ca. 20 



VOLUME 112, NUMBER 1 



245 



large, broad, green outer involucral bracts 
that completely conceal the inner bracts. 
The species is often described as herba- 
ceous, and it apparently commonly turns 
black when dry. 

The single known species is as follows: 

Myanmaria calycina (DC.) H. Rob., comb, 
nov. basionym: Vemonia calycina DC, 
Prodr. 5:60. 1836. 
Myanmar. 

Lampropappus (O. Hoffm.) H. Rob., stat. 
nov. (Gymnantheminae). Vemonia section 
Lampropappus O. Hoffm., Bol. Soc. Bro- 
ter 13:14. 1896. Type: Vemonia lampro- 
pappus O. Hoffm. Vemonia subsect. Tur- 
binella S. B. Jones, Rhodora 83:67. 1981. 
Type: Vemonia lampropappus O. Hoffm. 

Subshrubs to 0.6 m high; stems costate, 
tomentose, hairs with basally or slightly ex- 
centrically attached contorted capcells. 
Leaves alternate, shortly petiolate to more 
or less amplexicaul, blades densely tomen- 
tose abaxially. Inflorescence densely cor- 
ymbose, with many heads. Heads hemi- 
spherical to turbinate; involucral bracts ca. 
30, ca. 4-5-seriate, strongly gradate, outer 
bracts ovate, apices acute to shortly apicu- 
late; receptacle epaleaceous. Flowers ca. 20 
in a head; corollas white with purple tips to 
pale purple, actinomorphic or slightly zy- 
gomorphic, with lobe or sinus centered on 
outer side, longest lobe centered toward in- 
side. Bases of anther thecae rounded, with- 
out tails; apical appendage with somewhat 
thickened and ornamented cell walls; base 
of style with broadened node; sweeping 
hairs of style branches with obtuse tips. 
Achenes 5 -costate, densely villous with un- 
iseriate hairs, with few septae near base, 
with few idioblasts; raphids subquadrate to 
short-oblong; pappus setae persistent, 3-se- 
riate, broad and flat, densely scabrid on 
margins smooth on surfaces, outer seg- 
ments shorter and narrower. Pollen grains 
tricolporate, non-lophate, echinate. 

Lampropappus, as a section, was origi- 
nally compared with the American Stilp- 
nopappus Mart, ex DC. and the Eastern 



Hemisphere Vemonia sect. Stengelia { = 
Baccharoides Moench) by Hoffmann 
(1896) because of the broad flat pappus seg- 
ments, but was distinguished at least from 
the Vernonia sect. Stengelia by the exap- 
pendiculate involucral bracts. Lampropappus 
does not have the subulate tips on the pappus 
elements seen in Stilpnopappus. The African 
genus is further distinguished by the uniser- 
iate rather than biseriate hairs densely cov- 
ering the achene and by the non-lophate pol- 
len. One species of the present genus, L tur- 
binella, proves to be distinct in the sUghtly 
zygomorphic corollas that are rotated or re- 
supinate by 1/10 so that a sinus is centered 
to the outside and a lobe centered to the in- 
side. This is not seen in the other two species 
of the genus. This feature occurs elsewhere 
in the Vemonieae in the American genus 
Mattfeldanthus H. Rob. & R. M. King 
(1979), which also has zygomorphic coroUas. 
The latter genus is related to Lepidaploa 
(Cass.) Cass, with lophate pollen and seriate- 
cymose inflorescences. Such resupination of 
the corolla also seems to occur in the Gor- 
teriinae of the Arctoteae. 

The following three species are placed in 
the genus: 

Lampropappus eremanthifolia (O. Hoffm.) 
H. Rob., comb. nov. basionym: Vernonia 
eremanthifolia O. Hoffm., Bol. Soc. Bro- 
ter. 13:15. 1896. 
Angola. 
Lampropappus hoffmannii H. Rob., nom. 
nov. basionym: Vemonia lampropappus 
O. Hoffm., Bol. Soc. Broter. 13:14. 1896. 
Angola. 
Lampropappus turbinellus (S. Moore) H. 
Rob., comb. nov. basionym: Vernonia 
turbinella S. Moore, J. Linn. Soc. Bot. 
47:266. 1925. 

Congo, Malawi, Zambia. Zygomorphy of 
the corollas is not mentioned in the orig- 
inal description of the species (Moore 
1925). 

Acknowledgments 

The present study has been aided by loans 
of material kindly sent by The Royal Botanic 



246 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Garden, Kew; British Museum (Natural His- 
tory); Jardin Botanique de I'Etat, Bruxelles; 
and Museum National d'Histoire Naturelle, 
Laboratoire de Phanerogamie, Paris. Aspects 
of this study have been assisted by Margorie 
Knowles of the Department of Botany, 
Smithsonian Institution. John Pruski of the 
Smithsonian Department of Botany and Mi- 
chael O. Dillon of the Field Museum of Nat- 
ural History are thanked for critical reviews 
of the manuscript. 

Literature Cited 

Ayorinde, E O., B. D. Butler, & M. T. Clayton. 1990. 
Vemonia galamensis: a rich source of epoxy 
acid. — Journal of the American Oil Chemistry 
Society 67:844-845. 

Bohlmann, F, & J. Jakupovic. 1990. Progress in the 
chemistry of the Vernonieae (Compositae).- 
Plant Systematics and Evolution [Suppl. 4]:3- 
43. 

Brenan, J. P. M. 1953. Tropical African plants: XXIII. 
Compositae. — Kew Bulletin 1953:116-117. 

Candolle, A. P., de. 1836. Vemoniaceae in Prodromus 
Systematis Universalis regni vegetabilis 5:9- 
103. Treuttel et Wiirtz, Paris. 

Hind, D. J. N. 1996. 302 Distephanus populifolius. 
Curtis's Botanical Magazine 13:191-194. 

Hoffmann, O. 1896. Compostas da Africa Portugue- 
za. — Boletim da Sociedade Broteriana 13:11- 
35. 

Humbert, H. 1960. 189^ Famille-Composee In Flore 
de Madagascar et de Comores (plantes vascu- 
laires), 1:1-338. 

Isawumi, M. A. 1993. New combinations in Bacchar- 
oides Moench (Vernonieae; Compositae) in 
West Africa. — Feddes Repertorium 104:309- 
326. 

. 1995. 5. Notes on Vemonia (Vernonieae: 

Compositae) in West Africa. Pp. 51-106 in D. 
J. N. Hind, C. Jeffrey, & G. V Pope, eds.. Ad- 
vances in Compositae Systematics. Royal Bo- 
tanical Garden, Kew l:i-xiv, 1-784. 

, G. El-Ghazaly, & B. Nordenstam. 1996. Pol- 
len morphology, floral microcharacters and tax- 
onomy of the genus Baccharoides Moench 
(Vernonieae: Asteraceae). — Grana 35:205-230. 

Jeffrey, C. 1988. The Vernonieae in east tropical Af- 
rica, notes on Compositae: V — Kew Bulletin 
43:195-277. 

Jones, S. B., Jr. 1974. Vernonieae (Compositae) chro- 
mosome numbers. — Bulletin of the Torrey Bo- 
tanical Club 101:31-34. 

. 1977. Vernonieae — systematic review. Pp. 

503-521 in V. H. Heywood, J. B. Harbome & 



B. L. Turner, eds.. The Biology and Chemistry 
of the Compositae. Academic Press, London, 
New York. l:i-xiv, 1-619. 

. 1979. Chromosome numbers of Vernonieae 

(Compositae). — Bulletin of the Torrey Botani- 
cal Club 106:79-84. 

. 1981. Synoptic classification and pollen mor- 
phology of Vemonia (Compositae: Vernonieae) 
in the Old World).— Rhodora 83:59-75. 

. 1982. Documented chromosome numbers. 

Compositae. — Taxon 31:126-127. 

Keeley, S. C. 1994. Survey of the Vernonieae — A trib- 
al review [Abstract], p. 26 in D. J. N. Hind (co- 
ordinator), Compositae: Systematics Biology 
and Utilization, Paper and Poster Abstracts, i- 
XX, 1-151. 

. 1995. Exploration of ndhF sequence data in 

subtribal placements within the Vernonieae (As- 
teraceae) [Abstract]. — American Journal of Bot- 
any 82(6) [Suppl.]: 139. 

, & B. L. Turner. 1990. A preliminary cladistic 



analysis of the genus Vemonia (Vernonieae: 
Asteraceae). [Abstract]. — Plant Systematics and 
Evolution [Suppl. 4]:45-66. 

Kim, Hyi-Gyung, S. C. Keeley, & R. K. Jansen. 1996. 
Phylogenetic position of the Hawaiian endemic 
Hesperomannia (Mutisieae) based in ndhF se- 
quence data [Abstract]. — American Journal of 
Botany 83(6) [Suppl.]: 167. 

Kirkman, L. K. 1981. Taxonomic revision of Centrath- 
erum and Phyllocephalum (Compositae: Ver- 
nonieae). — Rhodora 83:1-24. 

Koyama, H. 1984. Taxonomic studies in the Compos- 
itae of Thailand 3. — Acta Phytotaxonomica et 
Geobotanica 35:49-58. 

Lisowski, S. 1992. Les Vernonieae (Asteraceae) dans 
la flore d'Afrique centrale (excl. le genre Ver- 
nonia). Fragmenta Floristica et Geobotanica 37: 
275-369. 

Mangenot, S., & G. Mangenot. 1957. Nombres chro- 
mosomiques nouveaux chez diverses Dicotyle- 
dones et Monocotyledones d'Afrique Occiden- 
tale. — Bulletin du Jardin Botanique de I'Etat 
27:637-654. 

, & . 1962. Enquete sur les nombres 

chromosomiques dans une collection de especes 
tropicales. — Revue de Cytologic et de Biolo- 
giae Vegetales 25:411-447. 

Mehra, P N., B. S. Gill, J. K. Mehta, & S. S. Sidhu. 
1965. Cytological investigations on the Indian 
Compositae. I. North Indian taxa. — Caryologia 
18:35-68. 

Moore, S. 1925. A third contribution to the Composite 
flora of Africa.-Joumal of the Linnean Society 
of Botany 47:257-284. 

Pope, G. V. 1983. Cypselas and trichomes as a source 
of taxonomic characters in the Erlangeoid gen- 
era.— Kirkia 12:203-231. 



VOLUME 112, NUMBER 1 



247 



. 1986. Vemonia chloropappa (Compositae) 

and related species in tropical Africa. — Kew 
Bulletin 41:393-397. 

Robinson, H. 1989. Acilepidopsis, a new genus of Ver- 
nonieae from South America (Asteraceae). — 
Phytologia 67:289-292. 

. 1990a. Six new combinations in Baccharoides 

Moench and Cyanthillium Blume (Vernonieae: 
Asteraceae). — Proceedings of the Biological 
Society of Washington 103:248-252. 

. 1990b. New combinations in the Asteraceae 

(Vernonieae, Heliantheae, Mutisieae). — Phyto- 
logia 69:105-107. 

. 1992a. Mesanthophora, a new genus of Ver- 
nonieae (Asteraceae) from Paraguay. — Novon 
2:169-172. 

. 1992b. The Asteraceae of the Guianas, III: 

Vernonieae and restoration of the genus Xiph- 
ochaeta. — Rhodora 94:348-361. 

. 1993. A review of the genus Critoniopsis in 

Central and South America (Vernonieae: Aster- 
aceae). — Proceedings of the Biological Society 
of Washington 106:606-627. 

. 1996. 33. The status of generic and subtribal 

revisions in the Vernonieae. Pp. 511-526 in D. 
J. N. Hind and H. J. Beentje, eds., Compositae 
Systematics. Proceedings of the International 
Compositae Conference, Kew, 1994. D. J. N. 
Hind, editor in chief. Royal Botanical Garden, 
Kew l:i-xiv, 1-784. 

, & B. Kahn. 1986. Trinervate leaves, yellow 

flowers, tailed anthers, and pollen variation in 



Distephanus Cassini (Vernonieae: Astera- 
ceae). — Proceedings of the Biological Society 
of Washington 99:493-501. 

-, & R. M. King. 1979. Mattfeldanthus miiti- 
sioides gen. et spec, no v. (Asteraceae: Vernon- 
ieae) from Bahia, Brazil. — Willdenowia 9:9-12. 

-, & C. Marticorena. 1986. A palynological 
study of the Liabeae (Asteraceae). — Smithson- 
ian Contributions to Botany 64:i-iv, 1-50. 

-, E Bohlmann, & R. M. King. 1980. Chemo- 



systematic notes on the Asteraceae. III. Natural 
subdivisions of the Vernonieae. — Phytologia 
46:421-436. 

Scott, A. J. 1993. Distephanus. in D. J. N. Hind, C. 
Jeffrey & A. J. Scott. Compositae. Pp. 19-21 
in J. Bosser et al., eds., Flore des Mascareignes. 
fasc. 109. Composees: Sugar Industry Research 
Institute (Mauritius), ORSTOM (Paris) and 
Royal Botanic Gardens, Kew. 

Verdcourt, B. 1956. Notes from the East African Her- 
barium: IV— Kew Bulletin 1956:445-454. 

Wild, H. 1977. The Compositae of the Flora Zambes- 
iaca area, 7 — Vernonieae (excluding Vemonia 
Schreb.).— Kirkia 10:339-384. 

. 1978. The Compositae of the Flora Zambes- 

iaca area 8 — Vernonieae {Vemonia). — Kirkia 
11:31-127. 

, & G. V. Pope. 1978a. Brachythrix: a new ge- 
nus of the tribe Vernonieae (Compositae) from 
South Central Africa. — Kirkia 11:25-30. 

, & . 1978b. The Compositae of the Flo- 
ra Zambesiaca area, 9-Vemonieae {Brachythrix 
and Hoehnelia). — Kirkia 11:133-142. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(l):249-250. 1999. 



INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 

Applications published in the Bulletin of Zoological Nomenclature 

The following Applications were published on 30 September 1998 in Vol. 55, 
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of 
these applications is invited for publication in the Bulletin and should be sent to the 
Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road, 
London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 

Case No. 

3050 Pachylops Fieber, 1858 (Insecta, Heteroptera): proposed designation of Cap- 

sus chloropterus Kirschbaum, 1856 (currently Orthotylus virescens 
(Douglas & Scott, 1865)) as the type species. 
3087 Hydrobia Hartmann, 1821 and Cyclostoma acutum Drapamaud, 1805 (cur- 
rently Hydrobia acuta; Mollusca, Gastropoda): proposed conserva- 
tion by replacement of the lectotype of H. acuta with a neotype; 
Ventrosia Radoman, 1977: proposed designation of Turbo ventrosus 
Montagu, 1803 as the type species; and HYDROBIINA Mulsant, 
1844 (Insecta, Coleoptera): proposed emendation of spelling to HY- 
DROBIUSINA, so removing the homonymy with HYDROBIIDAE 
Troschel, 1857 (Mollusca). 

3051 Scarus chrysopterus Bloch & Schneider, 1801 (currently Sparisoma chrysop- 

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neotype. 
1647 Cacatua Vieillot, 1817 and CACATUINAE Gray, 1840 (Aves, Psittacifor- 

mes): proposed conservation. 
3004 LoRisiDAE Gray, 1821 and Galagidae Gray, 1825 (Mammalia, Primates): 

proposed conservation as the correct original spellings. 



250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Opinions published in the Bulletin of Zoological Nomenclature 

The following Opinions were published on 30 September 1998 in Vol. 55, Part 3 
of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob- 
tained free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History 
Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 

Opinion No. 

1902. Anomalina d'Orbigny, 1826 (Foraminif erida) : Anomalina ariminensis 

d'Orbigny in Fomasini, 1902 designated as the type species. 

1903. Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): conserved as the correct 

original spelling, and corrections made to the entries relating to Um- 
bellularia Lamarck, 1801 on the Official Lists and Indexes of Names 
in Zoology. 

1904. Aporcelaimus Thome & Swanger, 1936 (Nematoda): Dorylaimus superbus 

de Man, 1880 designated as the type species. 

1905. S.D. Kaicher (1973-1992), Card Catalogue of World-Wide Shells: not sup- 

pressed for nomenclatural purposes. 

1906. Euchroeus Latreille, 1809 (Insecta, Hymenoptera): conserved; Chrysis pur- 

purata Fabricius, 1787 (currently Euchroeus purpuratus): specific 
name conserved; and Chrysis gloriosa Fabricius, 1793: specific 
name suppressed. 

1907. Nothosaurus Miinster, 1834 (Reptiha, Sauropterygia): given precedence over 

Conchiosaurus Meyer, [1833] 

1908. Hemidactylus gamotii Dumeril & Bibron, 1836 (Reptiha, Squamata): spe- 

cific name conserved. 

1909. Holotropis herminieri Dumeril & Bibron, 1837 (currently Leiocephalus her- 

minieri), Proctotretus bibronii T Bell, 1842 (currently Liolaemus 
bibronii) (Reptiha, Squamata): specific names conserved, and Lio- 
laemus bellii Gray, 1845 placed on the Official List. 



VOLUME 112, NUMBER 1 251 



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CONTENTS 

Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on the types and type lo- 
cality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and description of a new species 

i of Eleutherodactylus from Mato Grosso, Brazil (Amphibia: Anura: Leptodactylidae) 

% W. Ronald Heyer and Anna M. Muiioz 1 

A' new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Leptodactylidae) with 

comments on the zoogeography of the Brazilian campos rupestres W. Ronald Heyer 19 

.^A;new species of cardinalfish (Perciformes: Apogonidae) from the Bay of Bengal, Indian Ocean 
s: Thomas H. Fraser 40 

.i^ new species of deep-water skate, Rajella eisenhardti, (Chondrichthyes: Rajidae) from the 

,' Galapagos Islands Douglas J. Long and John E. McCosker 45 

A new genus of the subfamily Cubacubaninae (Insecta: Zygentoma: Nicoletiidae) from a Mex- 
ican cave Luis Espinasa 52 

Two new species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from Mexican 

caves, with redescription of the genus Luis Espinasa 59 

Anacroneuria from northeastern South America (Insecta: Plecoptera: Perlidae) Bill P. Stark 70 

A new species of Siamosquilla from Indonesia (Crustacea: Stomatopoda: Protosquillidae) 

Mark V. Erdmann and Raymond B. Manning 94 

The taxonomic status and zoogeography of Cambarus bartonii carinirostris Hay, 1914 (Crus- 
tacea: Decapoda: Cambaridae) Roger F. Thoma and Raymond F. Jezerinac 97 

Two new species of Aegla Leach (Crustacea: Decapoda: Anomura: Aeglidae) from southern 

Chile Carlos G. Jara and Victor L. Palacios 106 

The Hawaiian parthenopid crabs of the genera Garthambrus Ng, 1966, and Dairoides Stebbing, 

1920 (Crustacea: Decapoda: Brachyura) Peter K. L. Ng and S. H. Tan 120 

Reinstatement and further description of Eualus subtilis Carvacho & Olson, and comparison 
with E. lineatus Wicksten & Butler (Crustacea: Decapoda: Hippolytidae) 

Gregory C. Jensen and Rachel C. Johnson 133 

Deilocerus captabilis, a new species of cyclodorippid crab from southeastern Brazil (Crustacea: 

Decapoda: Brachyura: Cyclodorippidae) Marcos Tavares 141 

The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian Islands, with description 

of a new species Christopher B. Boyko 145 

Two new species of Hansenium (Crustacea: Isopoda: Asellota) from Madang, Papua New 

Guinea Kathrin S. Bolstad and Brian Kensley 164 

Caecidotea simulator, a new subterranean isopod from the Ozark Springfield Plain (Crustacea: 

Isopoda: Asellidae) Julian J. Lewis 175 

Collocherides brychius, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from a 

deep-water hydrothermal site in the northeastern Pacific Arthur G. Humes 181 

Cymbasoma californiense, a new monstrilloid (Crustacea: Copepoda: Monstrilloida) from Baja 

California, Mexico E. Suarez-Morales and R. Palomares-Garcia 189 

Early development of the deep-sea ampharetid (Polychaeta: Ampharetidae) Decemunciger apa- 

lea Zottoli Robert Zottoli 199 

A new species of Spiochaetopterus (Chaetopteridae: Polychaeta) from a cold-seep site off 
Hatsushima in Sagami Bay, central Japan 

Eijiroh Nishi, Tomoyuki Miura, and Michel Bhaud 210 

Two new subtribes, Stokesiinae and Pacourininae, of the Vemonieae (Asteraceae) 

Harold Robinson 216 

Revisions in paleotropical Vemonieae (Asteraceae) Harold Robinson 220 

International Commission on Zoological Nomenclature 249 

Announcement 25 1 



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VOLUME 112 
15 JUNE 1999 



NUMBER 2 



ISSN 0006-324X 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):253-302. 1999. 



A revision of the Antarctic genus Chlanidota 
(Gastropoda: Neogastropoda: Buccinulidae) 

M. G. Harasewych and Yuri I. Kantor 

(MGH) Department of Invertebrate Zoology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.; 

(YIK) Severtzov Institute, Russian Academy of Sciences, Leninski prospect 33, 

Moscow 117071, Russia 



Abstract. — The genus Chlanidota is revised to contain two subgenera, 
Chlanidota sensu stricto and Chlanidota {Pfejferia). Chlanidota s.s. has a broad 
distribution throughout the Antarctic region, and contains five species: Chlan- 
idota {Chlanidota) vestita (Martens, 1881), C (C) densesculpta (Martens, 
1885), C (C) paucispiralis Powell, 1951, C (C.) pilosa Powell, 1951, and C 
(C) signeyana Powell, 1951. The subgenus Pfejferia, which differs from 
Chlanidota primarily in the morphology of its operculum, is endemic to South 
Georgia Island, and is known from three species: Chlanidota {Pfejferid) chor- 
data (Strebel, 1908), C {P.) palliata (Strebel, 1908), C. (P.) invenusta, new 
species. With its bulliform shell lacking a siphonal canal, the monotypic, cir- 
cum- Antarctic genus Neobuccinum is conjectured to be the sister taxon of 
Chlanidota, but is readily distinguished by its smoother, higher-spired shell, 
radula in which the central cusp of the lateral teeth is medially placed, and a 
penis lacking a papilla. 



The genus Chlanidota Martens, 1878, is 
one of the more diverse and wide-ranging 
members of the antiboreal buccinoidean ra- 
diation comprising the family Buccinulidae. 
As is true of nearly all buccinoidean higher 
taxa, this genus has been defined primarily 
on the basis of shell morphology, occasion- 
ally supplemented with observations of the 
radula and operculum. Consequently, nei- 
ther the limits of this genus nor its relation- 
ships to other buccinoidean taxa have been 
well established. 

Martens (1878) first proposed Chlanidota 
as a monotypic subgenus of Cominella, but 
later (Martens 1881) transferred the subge- 
nus to Buccinum Linne, 1758. Watson 
(1886) included the type species of Chlan- 
idota in the genus Neobuccinum Smith, 
1877. Thiele (1912) erected the monotypic 
Ficulina as a subgenus of Cominella. Later, 
he discovered this taxon to be a junior syn- 



onym of Ficulina Gray, 1867, and proposed 
Notoficula Thiele, 1917, as a new name. 
Still later, Thiele (1929:315) treated Chlan- 
idota as a genus, and included Notoficula, 
Pfejferia Strebel, 1908, and Bathydomus 
Thiele, 1912, as subgenera. Powell (1951) 
elevated each of these subgenera to generic 
status, but regarded them to be closely re- 
lated. Oliver (1983) showed Notoficula to 
be a neotenous eratoid rather than a bucci- 
noidean. 

Chlanidota has been treated most recent- 
ly by Dell (1990), who included 13 species 
(Table 1), some tentatively, and noted that 
this genus has undergone an explosive ra- 
diation in Antarctic waters. He commented 
that the distributions, both geographic and 
bathymetric, "show puzzling patterns, es- 
pecially through the Scotia Arc and adja- 
cent regions of the Antarctic continent." 
The closely related taxon Pfejferia (see Ta- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Listing of species attributed to Chlanidota and Pfejferia and their present taxonomic placement. 
Taxa listed in bold are valid species referable to these subgenera of Chlanidota. 



Chlanidota bisculpta Dell, 1990 Not a Chlanidota. Belongs in an as yet undescribed genus. 

Chlanidota {Chlanidota) densesculpta (Martens, 1885) 



Chlanidota elongata (Lamy, 1910) 



Junior homonym of Cominella elongata Dunker, 1857; Chlani- 
dota lamyi Dell, 1990 is a replacement name. 
Not a Chlanidota. Belongs in an as yet undescribed genus. 
Belongs in the conoidean genus Belaturricula. 
Synonym of Chlanidota signeyana Powell, 1951 
Synonym of Chlanidota signeyana Powell, 1951 



Chlanidota eltanini Dell, 1990 
Chlanidota gaini Lamy, 1910 
Chlanidota lamyi Dell, 1990 
Chlanidota cf. lamyi Dell, 1990 

Chlanidota (Chlanidota) paucispiralis Powell, 1951 
Chlanidota {Chlanidota) pilosa Powell, 1951 

Chlanidota polysperia Dell, 1990 Not a Chlanidota. Belongs in an as yet undescribed genus. 

Chlanidota pyriformis Dell, 1990 Synonym of Chlanidota signeyana Powell, 1951 

Chlanidota smithi Powell, 1958 Synonym of Neobuccinum eatoni (Smith, 1875) 

Chlanidota (Chlanidota) signeyana Powell, 1951 
Chlanidota (Chlanidota) vestita (Martens, 1881) 
Chlanidota (Pfefferia) chordata (Strebel, 1908) 

Pfejferia cingulata Strebel, 1908 Synonym of Chlanidota (Pfefferia) palliata (Strebel, 1908) 

Pfefferia elata Strebel, 1908 Synonym of Chlanidota (Pfefferia) palliata (Strebel, 1908) 

Chlanidota (Pfefferia) palliata (Strebel, 1908) 

Chlanidota (Pfefferia) invenusta, new species. 



ble 1), endemic to South Georgia Island, 
was last reviewed by Powell (1951), who 
selected a type species and added several 
new records for two of the four species de- 
scribed by Strebel (1908). 

The present revision of the genus Chlan- 
idota, which includes the subgenus Pfeffer- 
ia, is based on a study of the large collec- 
tions of these taxa sampled under the aus- 
pices of the United States Antarctic Pro- 
gram (USAP), as well as the type material 
of all species described or subsequently as- 
signed to these taxa. The objectives of this 
study are: to assess the relationships among 
nominal taxa of Chlanidota and Pfefferia 
based on anatomical as well as concholog- 
ical characters; to discern biogeographic 
patterns in the distribution of these taxa; 
and to begin to build a framework of ana- 
tomical data that can be used to resolve 
phylogenetic relationships of and among 
the buccinoidean taxa of the Southern 
Hemisphere. 



Materials and Methods 

This report is based primarily on the ex- 
tensive collections of Chlanidota and Pfef- 
feria (including dry shells and alcohol pre- 
served anatomical material) collected by the 
United States Antarctic Program (USAP) 
and housed at the National Museum of 
Natural History, Smithsonian Institution 
(USNM). Additional material, including 
type specimens housed in the following in- 
stitutions, were examined: BMNH — The 
Natural History Museum, London; MNHN — 
Museum national d'Histoire naturelle, Par- 
is; MNH-U — Museum fiir Naturkunde der 
Humboldt-Universitat zu Berlin; SAM — 
The South Australian Museum, Adelaide; 
SMNH — Swedish Museum of Natural His- 
tory, Stockholm; ZMH — Zoologisches In- 
stitut und Zoologisches Museum der Uni- 
versitat Hamburg. 

In the material examined sections, "spec- 
imen" denotes alcohol preserved material. 



VOLUME 1 12, NUMBER 2 



255 





B 



Fig. 1. Measurements of shell and operculum features. AL, aperture length; BWL, body whorl length; OL. 
operculum length; PE, posterior edge; SL, shell length; SW, shell width; 6, deflection of aperture from shell 



axis. 



while "shell" refers to records containing 
only the empty shell. 

Shell and operculum measurements were 
obtained for representative specimens of 
each species (n = 10, when available), as 
detailed in Figure 1. Shell ultrastructure 
was observed along freshly fractured sur- 
faces parallel to the growing edge of the 
shell. Anatomical descriptions are based on 
gross dissections of preserved material. As 
the apex of the shell was strongly eroded in 
all specimens of Chlanidota, the aperture 



length (AL) rather than the shell length was 
used as the reference when reporting the 
relative size of anatomical features. Radulae 
were removed by dissection, cleaned using 
diluted bleach (NaClO), coated with carbon 
and gold, and examined using a Hitachi 
S570 Scanning Electron Microscope 
(SEM). 

Images were digitized using a Leaf Lu- 
mina Digital Scanning Camera. Optical and 
SEM images were processed using Pho- 
toshop Version 4.01 (Adobe). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Systematics 

Order Neogastropoda Wenz, 1938 

Superfamily Buccinoidea 

Rafinesque, 1815 

The buccinoidean whelks comprise one 
of the few uncontroversial, monophyletic 
groups within the order Neogastropoda. 
While buccinoideans are readily identified 
on the basis of shell, radular, and alimentary 
system morphology (e.g., weakly sculp- 
tured shells lacking spines or columellar 
folds, lateral teeth with >2 cusps, absence 
of accessory salivary glands and rectal 
gland), there is no consensus as to the rank 
or inter-relationships of the group as a 
whole, or its constituent higher taxa (e.g., 
Thiele 1929; Wenz 1938; Bouchet & Waren 
1985; 1986; Ponder & Waren 1988; Ponder 
& Lindberg 1996; Kantor ,1996). As a re- 
sult, most current classifications are of a ty- 
pological rather than phylogenetic nature. 

Based primarily on shell, radular, and 
opercular characters, Powell (1929) divided 
the Buccinoidea into the families Buccini- 
dae, Neptuneidae, Buccinulidae, Cominel- 
lidae and Fasciolariidae. In the same year, 
Thiele (1929) published an alternative and 
more widely accepted higher classification 
of the Buccinoidea, comprising the families 
Columbellidae, Buccinidae, Melongenidae, 
Nassariidae, and Fasciolariidae. Powell 
(1951) subsequently revised his classifica- 
tion, regarding the Southern Hemisphere 
Buccinulidae (now including the subfamily 
Cominellinae) to be more closely related to 
the boreal family Neptuneidae than to Buc- 
cinidae. Ponder (1971) reviewed the New 
Zealand species of Buccinulum and con- 
cluded that anatomical differences did not 
justify their segregation from Buccinidae. 
Ponder (1974) and subsequently Ponder & 
Waren (1988) reduced the Buccinoidea to a 
family within Muricoidea, and also reduced 
in rank or synonymized most of the previ- 
ously recognized families and subfamilies. 
Powell (1976) partially incorporated these 
demotions in taxonomic rank into revisions 
of his earlier work (e.g., Powell 1961), but 



(Powell 1979:192) clearly continued to re- 
gard the Southern Hemisphere Buccinuli- 
nae to be distantly related to the Northern 
Hemisphere Buccininae. Other authors have 
revised the higher systematics of the buc- 
cinoidean fauna of the Northern Hemi- 
sphere (Habe & Sato 1973) without treating 
austral representatives. 

Pending a phylogenetic revision of the 
higher taxa within Buccinoidea, we provi- 
sionally retain the taxon Buccinulidae and 
its subdivisions, as understood by Powell 
(1951), to encompass the austral radiation 
of buccinoidean taxa. 

Family Buccinulidae Finlay, 1928 

Finlay (1928:250) proposed the family 
Buccinulidae (with subfamilies Buccinuli- 
nae and Siphonaliinae) and enumerated the 
included genera, but provided no diagnosis 
or differentiating characters. Powell (1929: 
57) suggested that Finlay's work was based 
on protoconch and teleoconch characters, 
and provided a modified classification 
"based primarily on the dentition." Powell 
(1951:151) further refined his concept of 
Buccinulidae, noting that the chief charac- 
teristics of the group were tricuspid rachi- 
dian teeth and an operculum with a terminal 
or subterminal nucleus. He subdivided the 
family into three subfamilies, the Cominel- 
linae, Buccinulinae and Prosiphiinae based 
on the number of cusps on the lateral teeth. 

Subfamily Buccinulinae Finlay, 1928 

As refined by Powell (1951:151) the 
Buccinulinae are characterized by radulae 
with tricuspid rachidian teeth and tricuspid 
lateral teeth. The subfamily ranges from 
Antarctica to New Zealand, Australia, and 
along the eastern Pacific coast as far north 
as California. 

Genus Chlanidota Martens, 1878 

Powell speculated that Chlanidota, Pfef- 
feria and Neobuccinum may represent a 
transitional stage between the short-ca- 



VOLUME 112, NUMBER 2 



257 



nailed Cominellinae and the long-canalled 
Buccinulinae, the latter including the Ant- 
arctic genera Probuccinum, Cavineptunea, 
and Bathydomus . He included Chlanidota, 
Pfefferia, and Neobuccinum within Buccin- 
ulinae primarily on the basis of their tricus- 
pid lateral teeth, but noted that in these taxa 
the intermediate cusp was likely the result 
of a bifurcation of the inner cusp. 

Subgenus Chlanidota Martens, 1878 

Cominella {Chlanidota) Martens, 1878: 
[Type (by monotypy): Cominella (Chlan- 
idota) vestita Martens, 1878]. 

Chlanidota Thiele, 1929:314-315; Powell, 
1951:139; Dell, 1990:177. 

Description — Shell medium-sized for 
family, reaching 25.5 to 42.6 mm, depend- 
ing on species. Shell usually thin, fragile, 
ovate to elongate in outline, spire short to 
very short. Shell covered with thin to thick, 
smooth or hirsute periostracum. Spiral 
sculpture of either thin to very thin threads 
or of prominent cords. Aperture high, oval. 
Operculum very small (0.18-0.37 AL), 
ovate, coiled, with nucleus terminal, rotated 
to left, nearly perpendicular to long axis of 
operculum. Siphonal notch broad, dorsally 
recurved, siphonal fasciole with ridges 
along margins. Shell composed of 3 crys- 
talline layers, outermost layer prismatic, 
middle layer of crossed-lamellar crystals 
with crystal faces comarginal (oriented par- 
allel to growing edge), innermost layer of 
crossed lamellar crystals radial (oriented 
perpendicular to growing edge). 

Digestive system generally typical of 
buccinids. Proboscis of medium length or 
long. Radular ribbon long, triserial. Rachi- 
dian tooth with arched base, straight sides, 
3 large, robust cusps, usually of equal 
length. Lateral teeth with 3 cusps, outer 
cusp longest, middle cusp shortest, closely 
adjacent to innermost cusp. Salivary glands 
small, fused or separate. Valve of Leiblein 
large, well defined, lacking ciliary cone. 
Gland of Leiblein long, tubular, convoluted 
anteriorly, straightening and tapering pos- 



teriorly. Oesophagus wide, muscular, with 
posterior crop-like enlargement lined with 
tall longitudinal folds. Stomach broadly U- 
shaped, without caecum. 

Penis long, dorsoventrally flattened, with 
long, cylindrical terminal papilla, surround- 
ed by deep circular fold at its base. 

Remarks. — This subgenus is endemic to 
the Antarctic region. The highest diversity 
is in the Weddell Quadrant, especially in the 
vicinity of South Georgia Island. There are 
no records from the Ross Quadrant. Only 
Chlanidota vestita, the type species, ex- 
tends northward beyond the Antarctic Con- 
vergence. The bathymetric range of Chlan- 
idota sensu stricto is from 3 to 1116 m. 

Chlanidota {Chlanidota) vestita 

(Martens, 1878) 

Figs. 2-3; Table 2 

Cominella {Chlanidota) vestita Martens, 
1878:23; Martens, 1904:63; Thiele, 1904: 
168, Taf. DC, fig. 56 (radula); Lamy, 
1911:pl. I, fig. 5. 

Buccinum {Chlanidota) vestitum Martens, 
1881:43-44, Taf. 9, Fig. 3a-c. 

Chlanidota vestita Tryon, 1881:201, pi. 79, 
fig. 391; Smith, 1902:203; Powell, 1951 
139; Carcelles, 1953:191, pi. 3, fig. 58 
Powell, 1957:133; Powell, 1960:150 
Dell, 1964:288; Gaillard, 1971:100; Can- 
tera & Amaud, 1984:68; Dell, 1990:184, 
fig. 311. 

Neobuccinum vestitum Watson, 1886:216. 

Description. — Shell (Fig. 2) small for ge- 
nus (to 29 mm), thin, translucent, but 
strong, ovate in outline, with low, rounded 
spire. Protoconch unknown, upper teleo- 
conch whorls heavily eroded. Teleoconch of 
up to 3V2 evenly-rounded, convex whorls, 
deeply eroded where periostracum damaged 
or missing. Suture strongly impressed, 
forming extremely narrow channel between 
adapical spiral cord and previous whorl. 
Spiral sculpture of prominent but unequal 
cords, intervening spaces 2-4 times as 
broad as cords. Lectotype with 23 cords on 
body whorl, 8 on penultimate whorl. Para- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Chlanidota (Chlanidota) vestita (Martens, 1878). A-C. Lectotype, MNH-U 2561 3-a (here designated). 
D-F. Paralectotype, MNH-U 25613-b. Both from Kerguelen Island. Scale bar = 1 cm. 



lectotype with 22 cords on body whorl, 7 
on penultimate whorl. Numerous fine, sin- 
uated spiral threads between cords. Axial 
sculpture limited to growth lines, producing 
finely cancellate surface sculpture. Aperture 
broadly ovate, deflected from shell axis by 
14-18°. Outer lip thin, evenly rounded, 
simple. Columella —Vi AL, weakly con- 
cave, with strong siphonal fold. Parietal re- 
gion, siphonal fasciole overlain by thin, 
grayish callus. Siphonal notch broad, dor- 
sally recurved, with straight columellar, 
rounded apertural margins forming borders 
of fasciole. Ridge margin of fasciole runs 
from apertural margin of siphonal notch. 
Shell color chalky white, aperture weakly 
glazed. Periostracum thin, straw-colored. 



smooth, glossy. Operculum, as illustrated 
by Dell (1990:fig. 311), very small (0.28 
AL), ovate. Radula (Thiele 1904:fig. 56) 
with tricuspid rachidian and lateral teeth, as 
described for other species of Chlanidota 
illustrated herein. 

Type locality. — Kerguelen Island. 

Type material. — Two lots of syntypes, 
MNH-U 25613 (2 shells), MNH-U 25626 
(3 shells). One specimen from MNH-U 
25613, corresponding in size to the original 
description, is here designated as the lec- 
totype (Fig. 2A-C). The remaining four 
specimens are paralectotypes. 

Material examined. — The lectotype 
(MNH-U 2561 3-a) and one paralectotype 
(MNH-U 25613-b). 



VOLUME 112, NUMBER 2 



259 




150 



150^ 



to 



50 



lom 



150 



200 



"10 



a; 
O ~ 

50 



100 



150 



200 



B 



Fig. 3. Geographic distribution and bathymetric ranges of Chlanidota {Chlanidota) vestita (Martens. 1878), 
O = type locality. ▲ = published records, and Chlanidota {Chlanidota) pilosa (Powell, 1951). ^ = type locality. 
A. Bathymetric range of C. (C.) vestita (Martens, 1878). B. Bathymetric range of C. {C.) pilosa (Powell, 1951). 
Antarctic quadrants follow terminology of Markham (1912). Dashed line indicates Antarctic Convergence. 



Table 2. — Chlanidota {Chlanidota) vestita (Mar- 
tens, 1878). Shell measurements of lectotype and par- 
alectotype 1. Linear measurements in mm. 







Paralec- 


Character 


Lectotype 


totype 1 


Shell Length (SL) 


22.5 


19.4 


Body Whorl Length (BWL) 


20.1 


16.8 


Aperture Length (AL) 


17.1 


13.3 


Shell Width (SW) 


17.1 


14.5 


BWL/SL 


0.89 


0.87 


AL/SL 


0.76 


0.69 


SW/SL 


0.76 


0.75 


No. spiral cords on: 






Penultimate whorl 


8 


7 


Body whorl 


23 


22 



Published records. — H.M.S. Challenger. 
Sta. 149, Accessible Bay, Kerguelen, 
49°08'S, 70°12'E, in 20-25 fm volcanic 
mud; Sta. 149B, off Royal Sound, Kergue- 
len, 49°28'S, 70°30'W, 25 fm, volcanic 
mud; Sta. 149D, off Royal Sound, Kergue- 
len, 49°28'S, 70°13'W, 28 fm, volcanic 
mud; Sta. 151, off Heard Island, 
52°59'30"S, TS^SS'SO'^W, 75 fm, volcanic 
mud (Watson, 1886). British, Australian 
and New Zealand Antarctic Research Ex- 
pedition (BANZARE): Sta. 12, Off Grave 
Island, Island Harbor, Kerguelen, 5 m in red 
and brown algae; Sta. 58, Hydrography 
Channel, SE of Green Island, Kerguelen, in 
50 m (Powell 1951). RA^ Southern Cross: 
Cape Adare, Ross Sea, 43-47 m, BMNH 
(Smith 1902:203, see Dell 1990:fig. 311). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 3. — Chlanidota (Chlonidota) densesculpta (Martens, 1885). Measurements of shell characters. Linear 
measurements in mm (n = 10, holotype not included). 



Character 


Mean 


CT 


Range 


Holotype 


Shell Length (SL) 


34.3 


2.22 


29.5-36.6 


31.2 


Body Whorl Length (BWL) 


29.8 


1.90 


25.9-32.3 


27.8 


Aperture Length (AL) 


24.4 


2.18 


20.6-27.8 


24.4 


Shell Width (SW) 


23.2 


2.20 


18.7-27.2 


24.5 


Operculum Length (OL) 


6.2 


0.87 


4.4-7.5 


— 


BWL/SL 


0.87 


0.01 


0.85-0.88 


0.89 


AL/SL 


0.71 


0.03 


0.67-0.75 


0.78 


SW/SL 


0.68 


0.04 


0.63-0.74 


0.79 


OL/AL 


0.25 


0.04 


0.18-0.31 


— 



Distribution'. — Kerguelen, the Crouzets, 
Heard Island and the Ross Sea, in 5-150 m 
(Fig. 3). 

Remarks. — Chlanidota vestita is readily 
distinguished from all congeners by its 
small, broadly shouldered shell, stepped 
spire, high body whorl, and moderately 
strong spiral cords. 

In addition to the Kerguelen and Crozet 
Plateaus, records include a single report 
from Cape Adare on the Antarctic continent 
(Smith 1902). Dell (1990:184, fig. 311) ex- 
amined and figured the Cape Adare speci- 
men [BMNH], and confirmed it to be C. 
vestita, and not C smithi as reported by 
Powell (1958). Dell (1964:288) reported C. 
vestita from Heard Island. 

Chlanidota (Chlanidota) densesculpta 

(Martens, 1885) 

Figs. 4-7; Table 3 

Cominella (Chlanidota) densesculpta Mar- 
tens, 1885:91; Martens & Pfeffer, 1886: 
71, pi. 1, fig. 3a-f. 

Chlanidota densesculpta Thiele, 1912:263; 
David, 1934:128; Powell, 1951:140, pi. 
8, figs. 31-33; Carcelles, 1953:192, pi. 3, 
fig. 60; Powell, 1960:150; Dell, 1990 
(partim):183, fig. 305, non fig. 306. 

Description. — Shell (Fig. 4) large for ge- 
nus (to 40.3 mm), thin, translucent, fragile, 
ovate in outHne, with low, rounded spire. 
Protoconch unknown, early whorls heavily 
eroded. Teleoconch of up to 6 evenly- 
rounded convex whorls, deeply eroded 



where periostracum damaged or missing. 
Suture impressed, obscured by thick perios- 
tracum. Spiral sculpture of numerous ex- 
tremely fine spiral threads, broader than the 
spaces between them. Spiral threads sinu- 
ated, equal in strength, closely spaced (4-5 
per mm), —40 on penultimate whorl, >80 
on body whorl. Axial sculpture limited to 
growth lines, producing finely cancellate 
surface sculpture appearing in places more 
like a rectangular array of fine pits. Aper- 
ture broadly ovate, deflected from shell axis 
by 9-13°. Outer Hp thin, evenly rounded, 
slightly reflected. Columella <% AL, weak- 
ly concave, with strong siphonal fold. Cal- 
lus consisting of thin, silver-edged glaze 
overlying parietal region, siphonal fasciole. 
Siphonal notch broad, dorsally recurved, 
with straight columellar and rounded aper- 
tural margins that form borders of fasciole. 
Ridge margin of fasciole runs from aper- 
tural margin of siphonal notch, may be ev- 
ident beneath columellar callus in some 
specimens. Shell color chalky white, aper- 
ture weakly glazed. Periostracum (Fig. 6C) 
thick, orange-tan, hirsute. Hair-like projec- 
tions emanating from edges of lamellae at 
intersections of spiral threads with axial 
growth lines. Operculum (Fig. 4G-I, K) 
small (0.18-0.31 AL), broadly ovate, weak- 
ly coiled, with terminal nucleus (usually 
abraded) rotated nearly perpendicular to 
long axis. Attachment area nearly circular, 
spanning ^4 of inner surface, posterior, left 
margins thickened, glazed, abraded. 



VOLUME 112, NUMBER 2 



261 




Fig. 4. Chlanidota {Chlanidota) densesculpta (Martens, 1875). A-C. holotype, MNH-U 37478. South Geor- 
gia Island. D-E, H, J. RA^ Islas Orcadas, Sta. 20, Off South Georgia Island, 54°00'06"S, 37°40'36"W. in 68- 
80 m, USNM 906152. Periostracum of specimen in figure J removed with bleach to reveal spiral sculpture. F, 
G, I. RA^ Eltanin, Sta. 1533, South Georgia Island, 54°00'S, 37°27'W, in 3-6 m. F USNM 870722, G, I. USNM 
896097. K. Operculum of specimen in fig. D. Scale bar = 1 cm for A-J, 2 mm for K. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



rhd sg gL 



rhd 




Fig. 5. Chlanidota (Chlanidota) densesculpta (Martens, 1875). RA^ Islas Orcadas, Sta. 19, Off South Geor- 
gia Island, 54°01'42"S, 37°40'00"W, in 46-70 m, USNM 901651. A. Left, and B. right lateral views of 6 animal 
removed from shell. C. Ventral and D. left lateral views of anterior alimentary system. E. Posterior portion of 
the proboscis, with salivary glands removed to show the valve of Leiblein. F. Dorsal, and G. ventral views of 
stomach. H. Stomach, opened mid-dorsally. I. Penis. Scale bars = 5 mm. Abbreviations: ao, anterior aorta; ct, 
ctenidium; ctz, compacting zone of the stomach; ddg, duct of the digestive gland; dg, digestive gland; gL, gland 



VOLUME 112, NUMBER 2 



263 



Ultrastructure. — (Fig. 6D) Outermost 
layer prismatic (—90 fxm), middle layer 
comarginal crossed-lamellar (—140 ixm), 
inner layer radial crossed-lamellar (—32 
fxm). 

External anatomy. — (Fig. 5A-B). Soft 
tissues comprise approximately IVi whorls. 
Mantle cavity spans just under Vi whorl, 
kidney Va whorl, digestive gland and gonad 
VA whorls. Columellar muscle short, broad, 
attached to shell at rear of mantle cavity. 
Foot large, broadly rectangular (LAV ===^ 1- 
1.3). Body base color yellowish tan, dorsal 
surfaces of the head, tentacles, siphon, foot 
mottled with dark grayish black. Head 
small, with broad, tapering tentacles, black 
eyes. 

Mantle cavity. — Mantle edge smooth. Si- 
phon of medium length (— Vs AL), muscular, 
extending substantially beyond mantle 
edge. Osphradium greenish, bipectinate, 
with narrow axis, spanning about Vi mantle 
cavity length. Ctenidium large, wide, span- 
ning about ^A mantle cavity length. Hypo- 
branchial gland formed of few, distinct, 
widely spaced folds. 

Alimentary system. — Proboscis smooth, 
unpigmented, of moderate length when re- 
tracted (—0.6 AL), but long in semiprotract- 
ed position (—1.1 AL; Fig. 5C, D). Probos- 
cis sheath very thin-walled, translucent. 
Mouth opening forming triangular slit. Buc- 
cal mass muscular, large, filling retracted 
proboscis. Odontophoral cartilages paired, 
fused anteriorly, spanning Vs of buccal 
mass. Radular ribbon long, 10.9-13.2 mm 
(0.47-0.52 AL), =-540 fxm wide («0.023 
AL), triserial (Fig. 6A, B), consisting of 
75-80 rows of teeth, posteriormost 5-7 
teeth nascent. Rachidian teeth with 3 cusps 
on central portion of broad, anteriorly 
arched basal plate. Central cusp slightly 



longer, narrower than lateral cusps. Lateral 
teeth with 3 cusps. Outer cusp stouter, near- 
ly twice as long as inner cusp. Intermediate 
cusp slightly shorter than, immediately ad- 
jacent to inner cusp. Salivary glands small 
(Fig. 5C, D; sg), fused, dorsal to nerve ring, 
right salivary gland totally covering valve 
of Leiblein. Salivary ducts pass loosely 
along both sides of esophagus, becoming 
embedded in esophageal walls near rear of 
buccal mass. Valve of Leiblein well de- 
fined, large (Fig. 5E; vL), pyriform, without 
ciliary cone. Gland of Leiblein convoluted 
anteriorly, tapering posteriorly (Fig. 5C, D; 
gL), opening without constriction to mid- 
esophagus, just anterior to, left of crop (Fig. 
5C; poe). Esophagus wide, muscular, en- 
larging posteriorly (Fig. 5C; poe) to form a 
crop-like structure lined with tall longitu- 
dinal folds. Stomach U-shaped, without 
caecum (Fig. 5F, H). Paired ducts of diges- 
tive gland closely spaced (Fig. 5H; ddg). 
Typhlosoles present (Fig. 5H; tph), poorly 
defined. Digestive gland clearly divided 
into 2 lobes. Zone of compaction well pro- 
nounced (Fig. 5H; ctz). Rectum terminates 
with well-defined anal papilla. 

Female reproductive system. — Typically 
buccinoidean. Oviduct opens into medium- 
sized albumen gland. Ingesting gland sin- 
gle. Capsule gland large, occupies ^Vi of 
mantle cavity. Bursa copulatrix present, 
simple, hemispherical. 

Male reproductive system. — Seminal 
vesicle of medium size, highly coiled. Pros- 
tate narrow, running along posterior half of 
mantle cavity wall. Penis dorsoventrally 
compressed, long (0.8 X mantle cavity 
length), with smooth walls, long, cylindri- 
cal papilla surrounded by deep circular fold 
around its base (Fig. 51). 

Type locality. — South Georgia Island. 



of Leiblein; h, heart; ht, cephalic tentacles; ig, intestinal groove; mo, mouth; nep. nephridium; nr. circumoeso- 
phageal nerve ring; oe, oesophagus; op, operculum; os, osphradium; p, penis; poe. posterior oesophagus; pr, 
proboscis; prr, proboscis retractors; rhd, proboscis sheath; s, siphon; sg. salivary gland; st, stomach; t, testes; 
vL, valve of Leiblein. 



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Fig. 6. Chlanidota {Chlanidota) densesculpta (Martens, 1875). A. Dorsal, and B. left lateral (45°) views of 
the central portion of the radular ribbon taken from animal in Fig. 5. C-D. RA^ Eltanin, Sta. 1533, South Georgia 
Island, 54°00'S, 37°27'W, in 3-6 m, USNM 896097. C. Periostracum. D. Shell ultrastructure, fracture surface 
parallel to growing edge. Scale bar = 200 [xm for A-C, 100 jxm for D. 



Type material. — Holotype, MNH-U 

37478 (fig. 4A-C). 

Material examined. — Holotype. RA^ El- 
tanin: Sta. 1533, South Georgia Island, 
54°00'S, 37°27'W, in 3-6 m, 7 Feb 1966, 1 
shell, USNM 870722, 6 specimens, USNM 
896097. RA^ Mas Orcadas: Sta. 18, Off 
South Georgia Island, 54°02'30"S, 
37°39'36'W, in 60-71 m, 15 May 1975, 4 
specimens, USNM 901650; Sta. 19, Off 
South Georgia Island, 54°01'42"S, 
37°40'00"W, in 46-70 m, 15 May 1975, 10 
specimens, USNM 901651 (anatomical de- 



scriptions are based on specimens from this 
lot); Sta. 20, Off South Georgia Island, 
54°00'06"S, 37°40'36"W, in 68-80 m, 15 
May 1975, 7 specimens, USNM 901652; 
Sta. 21, Off South Georgia Island, 
53°57'30"S, 37°20'42"W, in 27-40 m, 16 
May 1975, 1 specimen, USNM 901653; 
Sta. 95, Off South Georgia Island, 
54°11'48"S, 37°41'06"W, in 68-80 m, 9 Jun 
1975, 3 specimens, USNM 901654. RA^ 
Prof. Siedlecki, Sta. 81, Off South Georgia 
Island, 54°43'S, 35°13'W, in 300-306 m, 11 
Dec 1986, 1 broken shell, USNM 897523. 



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265 




to 



-4-3 

C ~ 
50 

100 


1 


150 
200 

300 

"1 







No. of spec.x 5 

Fig. 7. Geographic distribution and bathymetric range of Chlanidota {Chlanidota) densesculpta (Martens, 
1875). The type locality is "South Georgia Island." 



South Georgia, Bay of Islands, in 7.5 m, 3 
specimens, USNM 252874. 

Published records. — RA^ Discovery Ih 
Sta. 45, 2.7 miles S 85°E of Jason Light, 
South Georgia, in 238-270 m; Sta. 141, 
East Cumberland Bay, South Georgia, 200 
yards from shore under Mt. Duse, in 17-27 
m; Sta. 145, Stromness Harbour, South 
Georgia, between Grass Island and Tons- 
berg Point, in 26-35 m; Sta. 1941, Leith 
Harbour, South Georgia, in 55-22 m; Sta. 
WS 62, Wilson Harbour, South Georgia, in 
26-83 m; Sta. MS 6, East Cumberland Bay, 
y^ mile south of Hope Point to Wa cables S 
X E of King Edward Point Light, South 
Georgia, in 24-30 m; Sta. MS 10, East 
Cumberland Bay, Va mile south of Hope 
Point to Va mile south of Government Flag- 
staff, South Georgia, in 26-18 m (Powell 
1951). SW of Snow Hill Island, 64°36'S, 
57°42'W, in 125 m (Strebel 1908). Kergue- 
len (22-345 m) and Crozet (22-505 m) Is- 
lands (Cantera & Amaud 1984). 

Distribution. — All of the specimens that 
we examined were from South Georgia Is- 



land. There have been several published re- 
ports of C. densesculpta occurring at other 
localities. According to Powell (1951), 
Martens (1903) record of C. densesculpta 
from Bouvet Island is Notoficula bouveti 
(Thiele, 1912). Strebel (1908:33) neither 
discussed nor figured the single specimen 
of C. densesculpta he reported from off the 
Antarctic Peninsula. Similarly, the recent 
reports of this species from the Kerguelen 
(22-345 m) and Crozet (22-505 m) Islands 
(Cantera & Amaud 1984:68) lack descrip- 
tions or illustrations. Pending confirmation 
of the identification of these records, we 
provisionally regard Chlanidota dense- 
sculpta to be endemic to South Georgia Is- 
land (Fig. 7). Live-collected specimens 
were all taken at depths ranging from 3 to 
80 m. The only record outside this range is 
a single dead and broken specimen (RA^ 
Prof. Siedlecki Sta. 81) taken in 300-306 
m. Dell (1990) misidentified two lots of the 
bathyal species Chlanidota (Pfefferia) in- 
venusta (described below) as C. (C.) den- 
sesculpta, which led to his incorrect report 



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Table 4. — Chlanidota {Chlanidota) pilosa Powell, 
1951. Shell measurements of holotype. Linear mea- 
surements in mm. 



Character 


Holotype 


Shell Length (SL) 


25.3 


Body Whorl Length (BWL) 


23.3 


Aperture Length (AL) 


21.1 


Shell Width (SW) 


20.0 


BWL/SL 


0.92 


AL/SL 


0.83 


SW/SL 


0.79 


No. spiral cords on: 




Penultimate whorl 


21 


Body whorl 


57 



of a very broad bathymetric range (0-1400 
m) for the latter species. 

Remarks. — This species has a striking 
conchological resemblance to Chlanidota 
(Pfefferia) palliata Strebel, which is also 
endemic to South Georgia Island, but can 
be discriminated most easily on the basis of 
its small operculum (18-0.31 AL, vs. 0.67- 
0.83 AL in C. palliata), which lacks the 
"feathered" posterior edge diagnostic of 



the subgenus Pfefferia. Chlanidota dense- 
sculpta is readily distinguished from other 
species of Chlanidota s.s. by its extremely 
fine and indistinct spiral sculpture as well 
as by the presence of a thick, finely hirsute 
periostracum. 

Skeletal ossicles of holothurians were 
found in the rectum of one of the specimens 
dissected. 

Chlanidota {Chlanidota) pilosa 

Powell, 1951 

Figs. 3, 8; Table 4 

Chlanidota pilosa Powell, 1951:139, 194, 
fig. L73, pi. 8, figs. 29-30; Kaicher 1990: 
5807. 

Description. — Shell (Fig. 8) small for ge- 
nus (to 25.5 nun), very thin, translucent, 
fragile, globose, with very low, rounded 
spire. Protoconch unknown, early whorls 
heavily eroded. Teleoconch of evenly 
rounded, highly convex whorls. Spiral 
sculpture of numerous fine spiral threads, 
broader than intervening spaces. Spiral 
threads sinuated, equal in strength, closely 




Fig. 8. Chlanidota {Chlanidota) pilosa Powell, 1951. Holotype, BMNH 1961500. RA^ Discovery II, Sta. 
456, 1 mile east of Bouvet Island, in 40-45 m. Scale bar = 1 cm. 



VOLUME 112, NUMBER 2 



267 



Table 5. — Chlanidota (Chlanidota) signeyana (Powell, 1951). Measurements of shell characters. Linear mea- 
surements in mm (n = 10, holotype not included). 



Character 


Mean 


CT 


Range 


Holotype 

of C. 
signeyana 


Holotype 

of C. 
pyriformis 


Holotype 

of C. 

elongata 

(from Lamy. 

1911) 


Shell Length (SL) 


29.8 


6.7 


21.4-42.2 


37.4 


28.6 


30.0 


Body Whorl Length (BWL) 


24.5 


5.1 


17.8-33.4 


31.9 


24.4 


26.5 


Aperture Length (AL) 


19.3 


3.5 


14.1-25.7 


25.3 


20.1 


20.7 


Shell Width (SW) 


18.2 


3.2 


13.5-22.8 


23.6 


18.6 


16.9 


Operculum Length (OL) 


5.9 


1.5 


4.5-9.4 


6.4 


— 


— 


BWL/SL 


0.83 


0.02 


0.79-0.86 


0.86 


0.85 


0.87 


AL/SL 


0.65 


0.04 


0.58-0.70 


0.68 


0.70 


0.69 


SW/SL 


0.61 


0.06 


0.54-0.71 


0.64 


0.65 


0.56 


OL/AL 


0.30 


0.04 


0.25-0.37 


0.27 


— 


— 


No. spiral cords on: 














Penultimate whorl 


10.1 


2.2 


7-14 


11 


7 


7 


Body whorl 


25.0 


6.2 


15-35 


26 


21 


21 



Spaced (2.4-3 per mm), 21 on penultimate 
whorl, 57 on body whorl. Axial sculpture 
limited to growth lines. Aperture broadly 
ovate, deflected from shell axis by 15°. Out- 
er Up very thin, evenly rounded, slightly re- 
flected. Columella <V2 AL, weakly con- 
cave, with strong siphonal fold. Callus con- 
sisting of a thin glaze overlying parietal re- 
gion and siphonal fasciole. Siphonal notch 
broad, slightly dorsally recurved, with 
straight columellar, rounded apertural mar- 
gins that mark borders of fasciole without 
forming raised ridge margins. Shell color 
white, aperture weakly glazed. Periostrac- 
um thin, light yellowish-brown, hirsute. 
Operculum small (0.3 AL), irregularly 
ovate. Radula is similar to that C (C.) dense- 
sculpta except rachidian cusps are closer to- 
gether, more broadly triangular, with the 
central cusp broader and longer than flank- 
ing cusps (Dell 1951:fig. L73). 

Type locality. — 1 mile east of Bouvet Is- 
land, in 40-45 m (RA^ Discovery II, Sta. 
456). 

Type material. — Holotype (live collect- 
ed) BMNH 1961500 (Fig. 8), and paratype 
(empty shell). 

Material examined. — The type material. 

Distribution. — Known only from Bouvet 
Island. 



Remarks. — This species is thus far 
known only from the type material. Chlan- 
idota pilosa is similar to low-spired, rotund 
specimens of C. densesculpta (e.g.. Fig. 
4E), but differs in having fewer and more 
pronounced spiral riblets on the penultimate 
and body whorls, and by having a fasciole 
that lacks a ridge along its margin. These 
two species are also widely separated in 
their distribution. Chlanidota pilosa differs 
from C vestita in having a lower spire, as 
well as more numerous and finer spiral 
threads. 

Chlanidota (Chlanidota) signeyana 

Powell, 1951 

Figs. 9-11; Table 5 

Chlanidota signeyana Powell, 1951:141, 
fig. F, 194, fig L74, 196, fig. N129, pi. 8, 
figs. 34-35; Carcelles, 1953:191, pi. 3, 
fig. 59; Powell, 1960:150; Kaicher, 1990: 
5801; Dell, 1990:177, fig. 307. 

Cominella (Chlanidota) vestita var. elon- 
gata Lamy, 1910:319; Lamy, 1911:6, pi. 
1 fig. 6. 

Chlanidota elongata Powell, 1951:140, 
194, fig. L76; Carcelles, 1953:191, pi. 3, 
fig. 56; Powell, 1960:150; Cemohorsky, 
1977:110; Horikoshi et al., 1979: 22, fig. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 9. Chlanidota (Chlanidota) signeyana Powell, 1951. A-C. Holotype, Chlanidota signeyana, BMNH 
1961512, RA^ Discovery //, Sta. 167, Off Signy Island, South Orkney Islands, 60°50'30"S, 46°15'00"W, in 244- 
344 m. D. Holotype, Cominella {Chlanidota) vestita var. elongata Lamy, 1910 (from Lamy, 1911), King George 
Island, South Shetland Islands, in 420 m. E, F Holotype, Chlanidota pyriformis Dell, 1990, USNM 613075, 



VOLUME 112, NUMBER 2 



269 



8; Hain, 1989:71, Taf. V/4, Taf. XXIII/1; 

Numanami, 1996: 160-162, figs. 106- 

107. 
Chlanidota lamyi Dell, 1990:182, fig. 310, 

new name for Cominella {Chlanidota) 

elongata Lamy, 1910, not Cominella 

elongata Dunker, 1857. 
Chlanidota pyriformis Dell, 1990:182, fig. 

309. 
Chlanidota cf. lamyi Dell, 1990:182, fig. 

315. 

Description. — Shell (Fig. 9) large for ge- 
nus (to 42.6 nun), thin, fragile, elongate to 
ovate in outline, with medium to high, 
rounded to turreted spire. Protoconch un- 
known, early whorls heavily eroded. Teleo- 
conch of up to 5+ evenly-rounded convex 
whorls, deeply eroded where periostracum 
damaged or missing. Suture deeply im- 
pressed, sharply abutting. Spiral sculpture 
typically of distinct, sharp, narrow cords 
(7-14 on penultimate whorl, 15-35 on body 
whorl) equal or subequal in prominence, 
that may be reduced or nearly absent in 
some specimens (Fig. 9F, K). In specimens 
with pronounced cords (Fig. 9H-I), includ- 
ing the holotype (Fig. 9A— C), 4—5 adapical 
cords on body and penultimate whorls usu- 
ally more widely spaced, giving the illusion 
that they are more pronounced. Spaces be- 
tween cords equal to cord width on early 
whorls, at least twice cord width on body 
whorl. Axial sculpture limited to fine, 
weakly recurved, prosocline growth lines. 
Aperture elongate to broadly ovate, deflect- 
ed from shell axis by 14-17°. Outer lip thin, 
evenly rounded, fragile. Columella 1/3-1/2 
AL, weakly convex, with strong siphonal 
fold. Callus of thin, silver-edged or thick, 
brownish glaze overlying parietal region, 
siphonal fasciole. Siphonal notch broad. 



dorsally recurved, with straight columellar 
and rounded apertural margins that form 
borders of fasciole. Ridge margin of fasci- 
ole extends from apertural margin of si- 
phonal notch. Shell color chalky-white to 
brownish or orange-tan. Aperture weakly 
glazed. Periostracum (Fig. lOE) thin to 
moderately thick, orange-tan to greenish- 
tan, smooth in weakly sculptured speci- 
mens, forming low, axial lamellae with 
short, hairlike projections on intersections 
with spiral cords of strongly sculptured 
specimens. Operculum (Fig. 9L) small 
(0.25-0.37 AL), broadly ovate, weakly 
coiled, with terminal nucleus rotated nearly 
perpendicular to long axis. Operculum may 
be thin, yellowish, translucent, but usually 
opaque, brownish to dark brown. Attach- 
ment area oval, spanning about % of inner 
surface, posterior and left margins thick- 
ened, abraded. 

Ultrastructure. — (Fig. lOF) Outermost 
layer prismatic (—38 ixm), middle layer 
comarginal crossed-lamellar (—60 |xm), in- 
ner layer radial crossed-lamellar (—20 fxm). 

Anatomy. — Gross anatomical features of 
C. signeyana very similar to those of C 
densesculpta, but left and right salivary 
glands of C. signeyana separate, not fused. 
Radular ribbon long, 9.6-12.2 mm (0.48- 
0.55 AL), 790-910 ixm wide (0.04 AL), 
triserial (Fig. lOA-D), consisting of 65-75 
rows of teeth, posteriormost 5-6 rows na- 
scent. Rachidian teeth with arched base, 
straight lateral sides, 3 large, robust cusps 
of equal length. Lateral teeth with 3 cusps, 
outer cusp longest, intermediate cusp short- 
est, situated close to inner cusp. 

Type locality. — [Chlanidota signeyana] 
Off Signy Island, South Orkneys, 
60°50'30"S, 46°15'00'W, in 244-344 m 



R/W Wesnvind, Sta. W-10, off South Shetland Islands, 63°00'S, 60°32'W, in 159 m. G. ¥UV Eltanin, Sta. 997. 
Gibbs Island. Bransfield Strait. South Shetland Islands, in 769 m, USNM 881971. H-1. RA' Eltanin. Sta. 426. 
Bransfield Strait. South Shetland Islands, in 809-1116 m, USNM 886204. J-K. R/V Islas Orcadas. Sta. 55. 
Saunders Island, South Sandwich Islands, in 64-88 m, USNM 901664. L. Operculum of specimen in figs. J, K. 
Scale bar = 1 cm for A-K, 1 mm for L. 



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Fig. 10. Chlanidota {Chlanidota) signeyana Powell, 1951. A. Dorsal and B. left lateral (45°) views of the 
central portion of the radular ribbon taken from the animal in Fig. 9J, K. C. Dorsal and D. left lateral (45°) 
views of the central portion of the radular ribbon taken from the animal in fig. 9H, I. E. Periostracum and F. 
shell ultrastructure of the same specimen. Scale bars = 200 ixm for A-D, 500 fxm for E, 50 ixm for F 



VOLUME 112, NUMBER 2 



271 




lOOl 



500l 



lOOOl 



No. of spec.x 5 



Fig. 11. Geographic distribution and bathymetric range of Chlanidota (Chlanidota) signeyana Powell, 1951. 
O = type locality, Chlanidota signeyana; + = type locality, Chlanidota lamyi; ♦ = type locality, Chlanidota 
pyriformis. # = examined material; D = published records. 



(RA^ Discovery II, Sta. 167); [Cominella 
(Chlanidota) vestita var. elongata Lamy, 
1910] King George Island, South Shet- 
lands, in 420 m; {^Chlanidota pyriformis 
Dell, 1990] Off South Shetland Islands, 
63°00'S, 60°32'W, in 26 m (RA^ Westwind: 
Sta. W.IO). 

Type material. — [Chlanidota signeyana^ 
Holotype, BMNH 1961512 (fig. 9A-C), 
and 2 paratypes; [Cominella (Chlanidota) 
vestita var. elongata Lamy, 1910] The ho- 
lotype (Fig. 9D) was not found in MNHN; 
[Chlanidota pyriformis Dell, 1990] Holo- 
type, USNM 613075 (Fig. 9E-F) paratypes 
1-6, USNM 860181, Paratype 7, MF 
47204. 

Material examined. — RA^ Eltanin: Sta. 
426, South Shetland Islands, Bransfield 
Strait, 62°27'S, 57°58'W, in 809-1116 m, 5 
Jan 1963, 3 shell fragments, USNM 



870290, 7 shells or fragments, USNM 

870291, 4 specimens, USNM 886204; Sta. 
428, South Shetland Islands, Bransfield 
Strait, 62°4rS, 57°51'W, in 662-1120 m, 5 
Jan 1963, 1 shell fragment, USNM 870293; 
Sta. 432, South Shetland Islands, 62°52'S, 
59°27'W, in 884-935 m, 7 Jan 1963, frag- 
ments of 2 shells, USNM 870303; Sta. 439, 
Antarctic Peninsula, 63°51'S, 62°38'W, in 
128-165 m, 9 Jan 1963, 1 shell, USNM 
870313, 1 specimen, USNM 881919; Sta. 
444, South Shetland Islands, 62°56'S, 
62°02'W, in 750-732 m, 11 Jan 1963, 1 
specimen, USNM 881923; Sta. 538, South 
Orkney Islands, Bransfield Strait, 60°30'S, 
47°34'W, in 616-662 m, 6 Mar 1963, frag- 
ments of 2 shells, USNM 870330; Sta. 993, 
South Shetland Islands, Elephant Island, 
6r25'S, 56°30'W, in 300 m, 13 Mar 1964, 
1 specimen, USNM 881970; Sta. 997, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



South Shetland Islands, Bransfield Strait, 
Gibbs Island, 61°44.18'S, 55°56.06'W, in 
769 m, 14 Mar 1964, 1 specimen, USNM 
881971; Sta. 1079, Scotia Ridge, 61°26'S, 
4r55'W, in 593-598 m, 13 Apr 1964, frag- 
ments of 3 shells, USNM 870627, 2 spec- 
imens, USNM 881983; Sta. 1084, Scotia 
Ridge, 60°22'S, 46°50'W, in 298-403 m, 15 
Apr 1964, 3 shells, USNM 870641, 7 shells 
and fragments, USNM 870642, 1 specimen, 
USNM 881988; Sta. 1997, Ross Sea, 
72°00'S, 172°28'E, in 530-549 m, 2 shells, 
USNM 898205; Sta. 2124, Ross Sea, 
71°38'S, 172°00'E, in 606-622 m, 1 shell, 
USNM 898202; Sta. 2127, Ross Sea, 
71°23'S, 17r36'E, in 515-521 m, 1 shell, 
USNM 898040. 

RA^ Eastwind: Sta. EW66-009, Palmer 
Peninsula 62°43.rS, 62°17.5'W, in 561 m, 
31 Jan 1966, 1 specimen, USNM 678473; 
EW66-012, Palmer Peninsula 63°23'S, 
60°51'W, in 405 m, 3 Feb 1966, 2 speci- 
mens, USNM 678475; EW66-021, off 
South Orkney Islands, 60°21'S, 45°55'W, in 
102 m, 9 Feb 1966, 1 specimen, USNM 
678396; EW66-022, off South Orkney Is- 
lands, 60°26.5'S, 45°53.3'W, in 168 m, 9 
Feb 1966, 1 specimen, USNM 678476; 
EW66-029, off South Orkney Islands, 
6r06'S, 44°57'W, in 290 m, 15 Feb 1966, 
1 specimen, USNM 678398; EW66-036, 
off Elephant Island, 6ri6'S, 54°45'W, in 
300 m, 17 Feb 1966, 2 shells, USNM 
678480; off beach wrack, Collins Pt., De- 
ception Island, 4 Jan 1966, 4 specimens, 
USNM 678378. 

RA^ Hero: Cruise 691, Sta. 23, off Ant- 
arctic Peninsula, Palmer Archipelago, Bra- 
bant Island, 64°12.06'S, 62°39.36'W, in 93- 
95 m, 9 Feb 1969, 1 specimen, USNM 
896260; Cruise 691, Sta. 31, South Shet- 
land Islands, Deception Island, 62°58.25'S, 
60°45.40'W, in 37-51 m, 13 Feb 1969, 7 
specimens, USNM 897557; Cruise 721, 
Sta. 751, off Antarctic Peninsula, 
64°46'28"S, 64°04'20"W, in 33 m, 31 Dec 
1971, 1 shell, USNM 901659; Cruise 721, 
Sta. 765, off Antarctic Peninsula, 64°47.3'S 
64°07.4'W, in 55 m, 3 Jan 1972, 1 shell. 



USNM 901671; Cruise 721, Sta. 1058, off 
Antartic Peninsula, 62°19.0'S, 59°11.4'W, 
in 44 m, 19 Dec 1971, 1 shell, USNM 
901657; Cruise 731, Sta. 1806, 64°46'31"S, 
64°04'52"W, in 47-75 m, 18 Feb 1973, 1 
specimen, USNM 901686; Cruise 824, Sta. 
32-1, Antarctic Peninsula, Anversa Island, 
64°37'S, 62°50.48'W, in 640-670 m, 23 
Mar 1982, 1 shell, USNM 881583; Cruise 
824, Sta. 7-1, Antarctic Peninsula, Adelaide 
Island, 66°40.40'S, 67°31.23'W, in 510-730 
m, 17 Mar 1982, 1 specimen, USNM 
896309; Sta. 32-1, Antarctic Peninsula, 
64°37'S, 62°50.80'W, in 640-670 m, 25 
Mar 1982, 1 shell, USNM 901658. 

RA^ Islas Orcadas: Sta. 39, South Sand- 
wich Islands, 57°01.2'S, 26°44.3'W, in 97- 
100 m, 23 May 1975, 1 shell, USNM 
901663; Sta. 40, South Sandwich Islands, 
Kadlemas Island, 57°06.8'S, 26°43.36'W, in 
15-33 m, 23 May 1975, 1 specimen, 
USNM 901660; Sta. 42, South Sandwich 
Islands, Kadlemas Island, 57°06.8'S, 
26°43.6'W, in 22-44 m, 24 May 1975, 12 
specimens, USNM 901669; Sta. 46, South 
Sandwich Islands, Kadlemas Island, 
57°06.2'S, 26°44.5'W, in 26-60 m, 25 May 
1975, 2 specimens, USNM 901665; Sta. 48, 
South Sandwich Islands, Kadlemas Island, 
57°06.3'S, 26°44.2'W, in 27-62 m, 23 May 
1975, 4 specimens, USNM 901670; Sta. 55, 
South Sandwich Islands, Saunders Island, 
57°47.12'S, 26°22.30'W, in 64-88 m, 27 
May 1975, 1 specimen, USNM 901664; 
Sta. 57, South Sandwich Islands, 27 May 
1975, 5 specimens, USNM 901656; Sta. 80, 
South Sandwich Islands, Zavodovski Is- 
land, 56°20.0'S, 27°38.8'W, in 351-393 m, 
4 Jun 1975, 13 specimens, USNM 901667; 
Sta. 110, Inaccessible Island, 60°28.rS, 
46°27.2'W, in 115-132 m, 16 Feb 1976, 2 
specimens, USNM 901666; Sta. 115, Inac- 
cessible Island, 60°32.4'S, 47°22.7'W, in 
567-671 m, 17 Feb 1976, 2 shells, USNM 
901655; Sta. 118, off South Orkney Islands, 
62°01.5'S, 43°06.2'W, in 759-857 m, 20 
Feb 1976, 1 specimen, USNM 901661; Sta. 
121, off South Orkney Islands, 6r47.00'S, 



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273 



43°40.00'W, in 616-642 m, 21 Feb 1976, 1 
specimen, USNM 901662. 

RA^ Westwind: Sta. W-10, off South 
Shetland Islands, 63°00'S, 60°32'W, in 159 
m, 26 Jan 1958, 6 shells, paratypes of C 
pyriformis Dell, 1991. USNM 860181, 4 
shells, USNM 890897. 

RA^ Polar Duke: off Palmer Peninsula, 
65°36'S, 67°2rW, in 200 m, 6 Sep 1985, 1 
shell, USNM 846179. 

RA^ Deepfreeze TV\ Sta. TD2-ED 14, off 
Cape Norwegia, Weddel Sea, 7r50'S, 
15°50'W, 1028-1122 m, 18 Jan 1959, 1 
shell and fragments, USNM 638862. 

Published records.-KIW Discovery II, 
Sta. 162, Off Signy Island, South Orkneys 
60°48'00"S, 46°08'00"W, in 320 m; Sta. 
170, off Cape Bowles, Clarence Is., 
6r25'30"S, 60°28'00"W, in 342 m; Sta. 
175, Bransfield Strait, South Shetlands, 
63°17'20"S, 59°48'15"W, in 200 m; Sta. 
1952, between Penguin Island and Lion's 
Rump, King George I, South Shetlands, in 
367-383 m; Sta. 1957, off south side of 
Clarence Island, South Shetlands, in 785- 
810 m (Powell, 1951). RA^ Eltanin: Sta. 
426, off South Shetland Islands, 62°27'S, 
57°58'W, in 1116-809 m; Sta. 439, west of 
Antarctic Peninsula, 63°51'S, 62°38'W, in 
128-165 m; Sta. 1084, north of South Ork- 
ney Islands, 60°22'S, 46°50'W, in 298-403 
m, RA^ Hero: Sta. 23, Antarctic Peninsula, 
64°12.1'S, 62°39.6'W, in 93-95 m; Arthur 
Harbor, Anvers Island, Antarctic Peninsula, 
30 m (Dell, 1990:180). PS ANT III/3: Sta. 
345, Weddel Sea, 73°27'S, 21°37'W, in 617 
m (Hain, 1989). Syowa Sta., Enderby Land, 
in 98 m (Horikoshi et al., 1979). JARE Sta. 

9, Breid Bay, Queen Maud Land, 
70°13.7'S, 24°25.7'E, in 276-289 m, Sta. 

10, 25 (Numanami, 1996); NZOI: Sta. 
A463, 72°20'S, 174°50'E, in 460 m; Sta. 
A464, 72°20'S, 174°00'E, in 376 m (Dell, 
1990, as C cf. lamyi). 

Distribution, — South Orkneys, South 
Shetlands, Antarctic Peninsula, and Queen 
Maud Land, and the eastern margins of the 
Ross Sea. Living specimens were taken in 
30-1116 m (Fig. 11). 



We include records for this species re- 
ported from the area of Queen Maud Land 
(as C. elongata). Figured specimens from 
this region (Numanami, 1996:fig. 106A-B, 
E-F). appear to have weaker, more numer- 
ous spiral cords. One specimen from Syowa 
station has rachidian teeth with narrower 
cusps and additional serrations, features that 
had not previously been observed in spec- 
imens of C signeyana. Powell (1951:142, 
fig. L78) reported and illustrated similar ad- 
ditional serrations in a specimen of Chlan- 
idota {Pfejferid) elata. 

We have examined and also include re- 
cords from the eastern margin of the Ross 
Sea, which had been listed as Chlanidota 
cf. lamyi by Dell (1990:310). Dell (1990) 
noted that these broken and worn speci- 
mens closely resemble C signeyana (as C 
lamyi) from the South Shetland Islands, but 
differed only in having more numerous spi- 
ral cords on their body whorl. 

Remarks. — Lamy (1910) was the first to 
recognize this species, proposing the taxon 
Cominella {Chlanidota) vestita var. elon- 
gata, without realizing that the name was 
preoccupied by Cominella elongata Dun- 
ker, 1857. Powell (1951:141) considered 
Chlanidota elongata to be restricted to the 
South Shetland Islands and erected the tax- 
on C. signeyana based on material from the 
South Orkney Islands. He recognized that 
C signeyana was closely related to C. elon- 
gata, but distinguished it as being "con- 
stantly broader and of ovate rather than cy- 
lindrical outline." Cemohorsky (1977:110) 
noted that Cominella (Chlanidota) vestita 
var. elongata Lamy, 1910 was preoccupied, 
but questioned whether this taxon was dis- 
tinct from C. vestita. After examining a 
broader range of specimens than were 
available to Powell, Dell (1990:180) noted 
that both ovate and cylindrical phenotypes 
co-occurred in South Shetland and South 
Orkney samples, but that specimens could 
be sorted into one or the other phenotype 
on the basis of the ratio of shell width (SW) 
to shell height (SH). He regarded Chlani- 
dota signeyana to be limited to the ovate 



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Table 6. — Chlanidota (Chlanidota) paucispiralis Powell, 1951. Measurements of shell characters. Linear mea- 
surements in mm {n = 10, holotype not included). 



Character 


Mean 


a 


Range 


Holotype 


Shell Length (SL) 


29.8 


3.0 


21.9-31.7 


21.9 


Body Whorl Length (BWL) 


25.2 


2.4 


19.2-27.2 


19.2 


Aperture Length (AL) 


19.9 


1.8 


15.8-21.9 


15.8 


Shell Width (SW) 


19.5 


2.1 


14.4-21.7 


14.4 


Operculum length (OL) 


4.5 


0.7 


4.0-5.0 


4.0 


BWL/SL 


0.85 


0.01 


0.84-0.88 


0.88 


AL/SL 


0.67 


0.03 


0.63-0.72 


0.72 


SW/SL 


0.65 


0.02 


0.62-0.68 


0.66 


OL/AL 


0.27 


0.02 


0.25-0.28 


0.25 


No. spiral cords on: 










Penultimate whorl 


4.4 


0.5 


4-5 


4 


Body whorl 


10.6 


1.6 


8-13 


10 



phenotype with SW/SH ratios ranging from 
0.60 to 0.66 and proposed the new name 
Chlanidota lamyi (for Cominella elongata 
Lamy, 1910, non Dunker, 1857) for the 
elongate phenotype, with SW/SH ratios be- 
tween 0.55 and 0.60. He did, however, rec- 
ognize the possibility that "these two forms 
are but extremes of the same species." 

Dell (1990) also described C. pyriformis 
based on a single lot (11 specimens) col- 
lected off the South Shetland Islands, which 
he regarded to be a morphologically con- 
sistent shallow-water form alHed to both C. 
signeyana and C. lamyi, but distinguished 
by its tapering, conical spire. 

We examined and measured a large series 
of specimens from throughout the ranges of 
the three nominal species, but were unable 
to differentiate between them in any repro- 
ducible manner. Larger samples from sev- 
eral stations (e.g., USNM 901669, USNM 
897557, USNM 860181) revealed that each 
population spans the range of morphologies 
of at least two, and sometimes all three of 
these nominal species. Chlanidota signey- 
ana is highly variable in shell outline and 
spiral sculpture, and encompasses the phe- 
notypes of Chlanidota lamyi and C pyri- 
formis. 

There does appear to be a clinal gradient 
in shell shape, with northern specimens 
tending to be more inflated, while popula- 
tions from the southern portion of the spe- 



cies' range, including the Antarctic main- 
land, have a higher proportion of narrower 
and more cylindrical shells. Numanami 
(1996:146) recognized a similar gradient in 
shell morphology in the taxon Neobuccin- 
um eatoni, noting that specimens from the 
Kerguelen and South Shetland Islands tend- 
ed to be more elongated, while the samples 
from Enderby Land tended to be more in- 
flated and shorter spired. However, his large 
sample (n = 98) from Enderby Land 
showed a wide variability in shell form, and 
included examples from the entire range of 
variation for the species. 

Chlanidota signeyana is most similar to 
C. (C.) paucispiralis Powell, 1951, but dif- 
fers in having more numerous and weaker 
spiral cords. Moreover, C. (C.) signeyana 
has not been recorded off South Georgia, 
while C. (C.) paucispiralis is endemic to 
South Georgia Island. 

Chlanidota (Chlanidota) paucispiralis 

Powell, 1951 

Figs. 12, 13, 14, 15; Table 6 

Chlanidota paucispiralis Powell, 1951:141, 
194, fig. L77, pi. 8, figs. 36-37; Carcel- 
les, 1953:191, pi. 3, fig. 57; Powell, 
1960:150; Kaicher, 1990:5806; Dell, 
1990:183-4, fig. 308. 

Description — Shell (Fig. 12) large for 
genus (to 40.3 mm), thin, strong, ovate to 



VOLUME 112, NUMBER 2 



275 



elongate-ovate in outline, with moderately 
high, conical spire. Protoconch unknown, 
early whorls heavily eroded. Teleoconch of 
up to SVi evenly-rounded convex whorls. 
Suture strongly impressed, forming very 
narrow channel between adapical spiral 
cord and previous whorl. Spiral sculpture of 
narrow, sharp, widely spaced cords, 4-5 on 
penultimate whorl, 8-13 on body whorl. 
Space between cords wide, evenly concave, 
with 0-8 very fine spiral threads. Two 
adapical cords form narrowly tabulate 
shoulder. Axial sculpture of fine, closely 
spaced growth lines. Aperture ovate, de- 
flected from shell axis by 9-15°. Outer lip 
thin, evenly rounded, with serrated edge 
formed by spiral cords. Columella <V2 AL, 
weakly concave, with pronounced siphonal 
fold. Parietal callus narrow, thick, overlying 
spiral cords. Siphonal notch dorsally re- 
curved, with straight columellar, rounded 
apertural margins that define limits of fas- 
ciole. Apertural margin of siphonal notch 
dorsally deflected, giving rise to low, sharp 
keel that forms "ridge margin" of fasciole. 
Shell color chalky white, aperture thinly 
glazed, occasionally with margins of glaze 
discolored to gray or golden tan. Perios- 
tracum (Fig. 13C, D) thin, smooth, yellow- 
ish-tan, sometimes consisting of closely ad- 
jacent, blade-like lamellae, which overlay 
spiral sculpture without giving rise to hair- 
like projections. Operculum (Fig. 12J) 
small (0.25-0.28 X AL), broadly ovate, 
weakly coiled, with laterally terminal nu- 
cleus (usually abraded) rotated nearly per- 
pendicular to long axis. Attachment area 
nearly circular, spanning % of inner surface, 
posterior and left margins thickened, 
glazed. 

Ultrastructure. — (Fig. 13D) Outermost 
layer prismatic (—90 ixm), middle layer 
comarginal crossed-lamellar (—235 (xm), 
inner layer radial crossed-lamellar (—55 
|xm). 

Anatomical features of C (C.) paucispir- 
alis are very similar to those of C. (C.) den- 
sesculpta. Only a single female specimen 



(SL = 27.7 mm) was available for dissec- 
tion. 

External anatomy. — Soft tissues com- 
prise —3^/^ whorls. Mantle cavity just over 
Vi whorl, kidney Va whorl, digestive gland 
IVa whorls. Columellar muscle short, broad, 
attaching to shell at rear of mantle cavity. 
Foot small, rectangular (LAV — 1.4), with 
long (0.2 X foot length) medial, ventral 
pedal gland. Base color yellowish tan, mot- 
tled with patches of grayish black on dorsal 
surfaces of siphon, tentacles, foot. Head 
small, with thin cylindrical tentacles, large 
black eyes. Siphon long, free, muscular. 

Alimentary system. — Radular ribbon 
« 9.0 mm long (0.51 AL), 830 jxm wide 
(0.05 AL), triserial (Fig. 13A-B), of -=70 
rows of teeth, of which 5-7 are nascent. 
Rachidian teeth with arched base, straight 
lateral sides, 3 large, robust cusps, central 
cusp slightly longer that lateral cusps. Lat- 
eral teeth with 3 cusps, outer cusp longest 
and, intermediate cusp shortest, adjacent to 
inner cusp. Anterior foregut very similar to 
that of its congeners. Stomach (Fig. 14) dif- 
fers in having very short caecum, which 
may be homologous to caecum (posterior 
mixing area) of Buccinidae. 

Type locality. — South Georgia, 53°52'30"S, 
36°08'00'W, in 160 m (RA^ Discovery 11, 
Sta. 159). 

Type material. — Holotype, BMNH 
1961513 (fig. 12A-C), and 2 paratypes, 
BMNH 1961513. 

Material examined. — Holotype. RA^ Is- 
las Orcadas: Sta. 8, off South Georgia Is- 
land, 53°35'48"S, 37°35'12"W, in 254-366 
m, 11 May 1975, 1 shell, USNM 881743; 
Sta. 10, off South Georgia Island, 
53°47'48"S, 37°26'42"W, in 165-234 m, 12 
May 1975, 1 shell, USNM 881745; Sta. 12, 
off South Georgia Island, 53°38'12"S, 
37°54'42"W, in 130-137 m, 13 May 1975, 
4 shells, USNM 881708; Sta. 13, off South 
Georgia Island, 53°44'12"S, 37°59'30'W, in 
128-137 m, 13 May 1975, 5 shells + frag- 
ments, USNM 901672; Sta. 14, off South 
Georgia Island, 53°4r48"S, 37°57'12'W, in 
144-150 m, 14 May 1975, 1 shell, USNM 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 12. Chlanidota (Chlanidota) paucispiralis Powell, 1951. A-C. Holotype, BMNH 1961513, RA^ Dis- 
covery II, Sta. 159, South Georgia, 53°52'30"S, 36°08'00"W, in 160 m. D, E. RA^ Prof. Siedlacki, Sta. 105, 
South Georgia, 53°40'S, 36°48'W, in 161-192 m, USNM 897515. F RA^ Prof. Siedlacki, Sta. 20, 53°58'S, 
38°42'W, in 189-200 m, USNM 897583. G. RA^ Prof Siedlacki, Sta. 16, Off South Georgia Island, 53°44'S, 



VOLUME 112, NUMBER 2 



277 




Fig. 13. Chlanidota (Chlanidota) paucispiralis Powell, 1951. A. Dorsal and B. left lateral (45°) views of 
the central portion of the radular ribbon taken from animal in fig. 12H, I. C. Periostracum and D. shell ultra- 
structure. RA^ Prof. Siedlacki, Sta. 24, 54°05'S, 38°25'W, in 197-207 m, USNM 897531. Scale bars = 200 |xm. 



881716; Sta. 15, off South Georgia Island, 
53°37'42"S, 38°04'00"W, in 128-137 m, 14 
May 1975, 1 shell, USNM 901673; Sta. 16, 
off South Georgia Island, 53°38'12"S, 
38°01'06"W, in 130-134 m, 14 May 1975, 
1 shell, USNM 901675; Sta. 17, off South 
Georgia Island, 53°36'00"S, 38°03'00'W, in 
122-124 m, 14 May 1975, 1 specimen + 2 
shells, USNM 901674 (anatomy studied); 
Sta. 30, off South Georgia Island, 



53°50'36"S, 36°18'36'W, in 185-205 m, 19 
May 1975, 2 shells, USNM 887839. 

RA^ Prof. Siedlecki: off South Georgia 
Island, Sta. 16, 53°44'S, 39°22'W, in 304- 
342 m, 02 Dec 1986, 4 shells, USNM 
897553; Sta. 20, 53°58'S, 38°42'W, in 189- 
200 m, 02 Dec 1986, 2 shells, USNM 
897583; Sta. 24, 54°05'S, 38°25'W, in 197- 
207 m, 03 Dec 1986, 2 shells, USNM 
897531; Sta. 28, 54°30'S, 38°56'W, in 220- 



<— 



39°22'W, in 304-342 m, USNM 897553. H, 1. RA^ Islas Orcadas, Sta. 17, Off South Georgia Island, 53°36'00"S, 
38°03'00"W, in 122-124 m, USNM 901674, (anatomical data based on this E specimen). J. operculum of 
specimen in figs. H, 1. Scale bar = 1 cm for A-1, 2 mm for J. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



poe 




Fig. 14. Chlanidota {Chlanidotd) paucispiralis Powell, 1951. Stomach of specimen in fig. 12H I. A. Dorsal 
view of the stomach. B. Stomach opened mid-ventrally. Scale bars = 2 mm. Abbreviations: ctz, compacting 
zone of the stomach; dc, duct pouch; ddg, duct of the digestive gland; ig, intestinal groove; pm, posterior mixing 
area, or caecum; poe, posterior oesophagus; sa, sorting area; tph, typhlosoles. 



232 m, 03 Dec 1986, 1 shell, USNM 
897484; Sta. 87, 54°18'S, 35°37'W, in 238- 
247 m, 11 Dec 1986, 2 shells, USNM 
897537; Sta. 101, 53°47'S, 36°34'W, in 
263-277 m, 1 shell, USNM 897457; Sta. 
105, 53°40'S, 36°48'W, in 161-192 m, 14 
Dec 1986, 1 shell, USNM 897515; Sta. 
116, 53°43'S, 38°36'W, in 260-306 m, 16 
Dec 1986, 1 shell, USNM 896993; Sta. 
121, 53°57'S, 38°10'W, in 90-100 m, 16 
Dec 1986, 1 shell, USNM 896991. 

RA^ Eltanin: Sta. 671, off South Georgia 
Island, 54°41'S, 38°38'W, in 220-320 m, 23 
Aug 1963, 1 shell, USNM 870378; Sta. 
678, off South Georgia Island, 54°49'S, 
38°01'W, in 732-814 m, 24 Aug 1963, 1 



juvenile specimen, USNM 881942; Sta. 
1535, off South Georgia Island, 53°5rS, 
37°38'W, in 97-101 m, 7 Feb 1966, 10 
shells + fragments, USNM 898666. 

Distribution. — South Georgia Island, in 
97-814 m (Fig. 15). Live specimens were 
collected in 122-814 m. 

Remarks. — Unlike most of its congeners, 
this geographically restricted species dis- 
plays remarkably little variation in shell 
morphology. It is readily distinguished from 
all other Chlanidota sensu stricto by its 
prominent, broadly spaced, spiral cords. It 
most closely resembles Chlanidota {Pfejfer- 
ia) chordata (Strebel, 1908), from which it 
is readily distinguished by its small, smooth 



VOLUME 112, NUMBER 2 



279 




100 
150 
2001 



300 



400 



500 




a 



700 



No. of spec.x 5 

Fig. 15. Geographic distribution and bathymetric range of Chlanidota (Chlanidota) paucispiralis Powell, 
1951. O = type locality; # = examined material. 



operculum. Chlanidota {Pfejferia) chordata 
(Strebel, 1908) has a thicker shell and a 
thicker, hirsute periostracum. 

Subgenus Chlanidota {Pfefferia) 
Strebel, 1908 

Pfefferia Strebel, 1908:33-4; Powell, 1951: 

142. 
Chlanidota (Pfefferia) Thiele, 1929:315. 

Type species. — Pfefferia palliata Strebel, 
1908, by subsequent designation, Powell, 
1951:142. 

Description. — Shell medium sized for 
family, maximum adult size 34.5 to 38.3 
mm, depending on species. Shell usually 
moderately thick, ovate to elongate in out- 
line, with short spire. Shell covered with 
thick periostracum, either smooth or with 
fine axial lamellae. Spiral sculpture of thin 
threads or prominent cords. Aperture high, 
oval. Operculum (Fig. 16) large (0.48-0.83 
AL), leaf-shaped, tapered above, below, 
with terminal nucleus rotated nearly per- 
pendicular to long axis of operculum. Pos- 
terior edge of operculum with tall ridge of 



feathered lamellae, broadest just above mid- 
length. Siphonal notch broad, dorsally re- 
curved, siphonal fasciole with ridged mar- 
gins. Shell ultrastructure and gross anatomy 
of mantle cavity, alimentary and male re- 
productive systems as in Chlanidota sensu 
stricto. 

Remarks. — The subgenus Chlanidota 
(Pfefferia) is endemic to South Georgia Is- 
land. In his description of Pfefferia, Strebel 
(1908) noted many similarities with Chlan- 
idota, especially in the morphology and 
sculpture of the shell and periostracum. He 
distinguished the two as genera on the basis 
of their opercula, that of Chlanidota being 
small, triangular, with rounded corners, and 
with its apex and nucleus along the poste- 
rior margin. The operculum of Pfefferia is 
much larger, elongated, tapering above and 
below, and has a characteristic, "feathered" 
posterior edge. Strebel further commented 
that low and high-spired forms (without in- 
termediates) were encountered at each of 
the two stations from which Pfefferia had 
been collected. Each station also contained 
strongly and weakly sculpture forms. While 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 16. Chlanidota {Pfejferia) palliata (Strebel, 1908). A. Operculum showing terminal nucleus rotated 
nearly perpendicular to long axis of operculum. Lines C-C and F-F' correspond to the sections shown by Figs. 
C and F B. Left lateral (45°) view of operculum. C. Section trough flat, central portion of operculum (along the 
line C-C on Fig. A), nucleus at the bottom. D, E. Successive layers are closely appressed, adherent in this 



VOLUME 112, NUMBER 2 



281 



expressing uncertainty as to whether these 
differences warranted specific status, Stre- 
bel nevertheless described four species, 
commenting that additional material would 
allow these questions to be better answered. 
Examination and dissection of additional 
material of three species of Chlanidota 
(Pfefferia) revealed that Chlanidota (Pfef- 
feria) cannot be distinguished from Chlan- 
idota sensu stricto on the basis of gross 
anatomy. Of special interest is the variable 
operculum of the newly described C (P.) 
invenusta, which ranges from the typical 
Chlanidota (Pfejferia) operculum to one 
very similar to, but larger than, that of 
Chlanidota (Chlanidota), and includes in- 
termediate morphologies. Due to the pau- 
city of distinguishing features, we consider 
Chlanidota (Pfefferia) to warrant only sub- 
generic distinction from Chlanidota sensu 
stricto. 

Chlanidota (Pfefferia) palliata 

(Strebel, 1908) 
Figs. 17, 18, 19, 20; Table 7 

Pfefferia palliata Strebel, 1908:34-5, pi. 3, 
fig. 39a-f; Thiele, 1912:pl. 16, fig. 20 
(radula); Powell, 1951:143; Carcelles, 
1953: 193. 

Pfefferia elata Strebel, 1908:35, pi. 3, fig. 
40; Powell, 1951:142, 194, fig. L78 (rad- 
ula), 196, fig. N 128 (operculum). 

Pfefferia cingulata Strebel, 1908:36, pi. 3, 
fig. 42a-c; Powell, 1951:142-3, 194, fig. 
L79 (radula); Carcelles, 1953: 193. 

Description. — Shell (Fig. 18) large for 
genus (to 34.5 mm, Powell, 1951:142), 
thick, solid, ovate-rounded. Protoconch, 
early teleoconch whorls eroded in all spec- 
imens. Teleoconch with up to 4 evenly in- 
flated whorls. Shoulder not pronounced. 
Suture deep, adpressed, narrowly chan- 



neled. Spiral sculpture of alternating broad, 
low cords (10-17 on body whorl, 3-5 on 
penultimate whorl) and fine threads (20-24 
per 5 mm), spanning entire shell surface. 
Cords more clearly visible on young spec- 
imens with thinner periostracum. Axial 
sculpture limited to indistinct growth lines. 
Aperture tall, ovate, deflected from the shell 
axis by 16-18°. Siphonal canal is not pro- 
nounced. Outer lip evenly rounded, thick, 
weakly reflected in adults, very thin in ju- 
venile and some adult specimens. Columel- 
la <V2 AL, weakly concave, with fine pus- 
tules. Siphonal fold strong. Callus thick, 
clearly demarcated, narrowly overlying pa- 
rietal region, siphonal fasciole in mature 
specimens. Siphonal notch broad, dorsally 
reflected, columellar margin straight, aper- 
tural margin rounded, reflected, forming 
pronounced ridge margin along adapical 
edge of fasciole. Shell color grayish white, 
translucent. Periostracum very thick, tightly 
adherent to shell surface, with evenly 
spaced, axially reflected lamellae with 
fringed edges evident on early whorls and 
juvenile specimens (Fig. 19C). Operculum 
(Fig. 16A, D) large, leaf-shaped, with api- 
cal nucleus, massive, raised, lamellose bor- 
der along posterior margin, 0.67-0.83 AL. 
Attachment area spans nearly entire inner 
surface, posterior, left margins thickened, 
glazed. 

Ultrastructure. — (fig. 19E) Outermost 
layer prismatic (—50 ixm), middle layer 
comarginal crossed-lamellar (—175 ixm), 
inner layer radial crossed-lamellar (—95 
fxm). Inner surface of the outer aperture lip 
with numerous, fine spherules (Fig. 17D). 

External anatomy. — Body of IVi whorls 
(Fig. 18A, B), mantle cavity spanning — V2 
whorl, kidney Vs whorl, digestive VA whorl. 
Foot of preserved specimens short (LAV 
— 1), with rounded posterior edge. Head 



^- 



region of the operculum. F. Section through "feathered" portion of operculum (along the line F-F' on Fig. A), 
nucleus at the bottom. G, H. Successive layers are unattached along apical portion of posterior edge of oper- 
culum. 1. new lamellae are deposited along the entire length of the operculum. 



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Fig. 17. Chlanidota (Pfejfeira) palliata (Strebel, 1908). A-C. Lectotype, SMNH (here designated), Sveska 
Siidpolarexp., Sta. 22, South Georgia, 54°17'S, 36°28'W, in 75 m. D. Operculum of lectotype, enlarged. 



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283 



Table 7. — Chlanidota {Pfejferia) palliata (Strebel, 1908). Measurements of shell characters. Linear measure- 
ments in mm (n = 10, type material not included). 











Lectotype 
of C. 


Lectotype 
of C 


Holotype of 
C. elata 


Character 


Mean 


(T 


Range 


palliata 


cingidata 


(broken) 


Shell Length (SL) 


27.1 


3.6 


18.1-30.0 


32.8 


25.9 


-37 


Body Whorl Length (BWL) 


24.6 


3.4 


16.4-27.1 


29.2 


23.7 


-32 


Aperture Length (AL) 


19.3 


2.0 


14.4-20.4 


23.5 


19.8 


-23 


Shell Width (SW) 


19.4 


2.3 


14.0-21.2 


23.4 


19.4 


-22 


Operculum Length (OL) 


14.1 


2.0 


9.6-16.7 


— 


15.4 


— 


BWL/SL 


0.91 


0.02 


0.86-0.95 


0.89 


0.92 


-0.86 


AL/SL 


0.72 


0.04 


0.67-0.80 


0.72 


0.76 


-0.62 


SW/SL 


0.72 


0.03 


0.68-0.78 


0.71 


0.75 


-0.60 


OL/AL 


0.73 


0.06 


0.67-0.83 


— 


0.78 


— 



narrow, with stout, conical tentacles, eyes 
on small lobes. Body lacks pigmentation. 

Mantle cavity. — Mantle edge smooth. Si- 
phon medium to long, muscular, extending 
substantially beyond mantle edge (~% AL). 
Osphradium dark greenish-brown, bipectin- 
ate, spans ~^/^ mantle length. Hypobranchi- 
al gland of few, distinct, widely spaced 
folds. Ctenidium large, wide, spans ~^/4 
mantle length, lamellae tallest in posterior 
Va of ctenidium, becoming shorter anterior- 

ly. 

Alimentary system. — Proboscis (Fig. 
19C, D, pr) thick (L/D « 3.5), of moderate 
length (0.9 AL). Mouth opening, triangular 
slit. Buccal mass muscular, large, nearly 
equal to retracted proboscis in length. 
Odontophoral cartilages paired, fused an- 
teriorly, spanning ~% of proboscis length. 
Radular ribbon (Fig. 18A, B) of moderate 
length, 10.6 mm (0.66 AL), —760 jjim wide 
(0.05 AL), triserial, consisting of 60-65 
rows of teeth, posteriormost 5 nascent. 
Rachidian teeth with arched base, nearly 
straight lateral sides, 3 large, robust cusps 
of equal length. Central cusp longer than 
flanking cusps in one specimen (Fig. 18 A, 



B). Lateral teeth with 3 cusps, outer cusp 
longest, intermediate cusp shortest, adjacent 
to inner cusp. Intermediate cusp nearly 
fused to inner cusp in one specimen (fig. 
ISA, arrow). Salivary glands small, fused 
(Fig. 19G), situated above nerve ring. Right 
salivary gland completely covering valve of 
Leiblein. Salivary ducts run loosely along 
both sides of oesophagus, entering esopha- 
geal wall near posteriormost portion of re- 
tracted proboscis. Valve of Leiblein (Fig. 
18E, vL) well defined, large, without cili- 
ated cone. Gland of Leiblein (Fig. 18C-E, 
gL) long, tubular, coiled anteriorly, tapering 
posteriorly. Oesophagus wide, muscular, 
expanding posteriorly to form crop (Fig. 
181, poe) Uned with tall longitudinal folds. 
Stomach (Fig. 18H, I) U-shaped, without 
caecum. Ducts of digestive gland paired, 
closely spaced, transverse fold slightly 
raised. Typhlosoles present, poorly defined. 
Rectum terminating with well defined anal 
papilla. 

Female reproductive system. — Typically 
buccinoidean. Oviduct opens into medium- 
sized albumen gland. Ingesting gland sin- 
gle. Capsule gland large, occupies ^Vi of 



<— 

E. Paralectotype, SMNH, from the type locality. F-H. Lectotype of Pfejferia cingulata (Strebel. 1908) (here 
designated), Sveska Sudpolarexp., Sta. 34, Cumberland Bay, South Georgia, 54°irS, 36°18'W, in 252-310 m. 
I. Holotype of Pfejferia elata (broken specimen), same station as types of C. (P.) palliata. J, K, L. both from 
RA^ Islas Orcadas, Sta. 33, 54°30.7'S, 35°35.9'W, in 261-267 m, USNM 901676. Scale bar = 1 cm for A-C. 
E-L, 5 mm for D. 



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Fig. 18. Chlanidota {Pfejferia) palliata (Strebel, 1908). A. Dorsal, and B. left lateral (45°) views of the 
central portion of the radular ribbon taken from animal in Fig. 17 J. C. Periostracum. D. Edge of outer lip. E, F. 
Shell ultrastructure. E. Fracture surface parallel to outer lip. F. Fracture surface perpendicular to outer lip. Scale 
bars = 200 ixm for A, B, 100 ixm for C-F 



VOLUME 112, NUMBER 2 



285 



mantle cavity. Bursa copulatrix present, 
simple, hemispherical. 

Male reproductive system. — Very similar 
to that of C. (C.) dense sculpta, penis has 
the same size, overall shape, terminal pa- 
pilla. 

Type locality. — {Pfejferia palliata & P. 
elata] South Georgia, 54°17'S, 36°28'W, in 
75 m (Sveska Siidpolarexp., Sta. 22); [Pfef- 
feria cingulata] Cumberland Bay, South 
Georgia, 54°irS, 36°18'W, in 252-310 m. 
(Sveska Siidpolarexp., Sta. 34). 

Type material. — [Pfefferia palliata] Lec- 
totype (here designated) SMNH (fig. 17A- 
C), and 2 paralectotypes SMNH (fig. 17E); 
[Pfefferia elata] Holotype, SMNH 3661 
(Fig. 171), broken specimen; [Pfefferia cin- 
gulata] Lectotype (here designated) (Fig. 
16F-H) and juvenile paralectotype, SMNH. 

Material examined. — Type material. RA^ 
Islas Orcadas: Sta. 31, 19 May 1975, 
54°05.36'S, 36°30.48'W, 130-143 m, 1 
shell, USNM 887863; Sta. 32, 19 May 
1975, 54°21.36'S, 35°58.42'W, 141-164 m, 
1 shell, USNM 887872; 9 shells, USNM 
887867; Sta. 33, 19 May 1975, 54°30.7'S, 
35°35.9'W, 261-267 m, 27 specimens, 
USNM 901676. 

Published records. — RA^ Discovery II, 
Sta. 30, West Cumberland Bay, South 
Georgia, 2.8 miles S, 24°W of Jason Light 
[16 Mar 1926], 251 m (Powell, 1951:142). 

Distribution. — Known only from off the 
northeastern coast of South Georgia (Fig. 
19). Live material was collected between 75 
and 310 m. 

Remarks. — The type specimens of 
Chlanidota {Pfefferia) palliata and C. (P.) 
elata were taken from the same dredge 
haul. Strebel (1908) distinguished these 
taxa primarily on the difference in their 
spire height and the strength of their spiral 
sculpture, but questioned whether these dif- 
ferences merited specific recognition. We 
were able to examine a larger sample than 
was available to Strebel (USNM 901676, n 
= 27) and found it to contain specimens 
spanning a morphological continuum be- 
tween these two forms. The type series of 



C. (P.) cingulata was collected very near 
the type locality of C. (P.) palliata, but at 
slightly greater depth. The specimen select- 
ed as lectotype of C. (P.) cingulata is, in 
our opinion, conspecific with the specimen 
selected as lectotype of C. (P.) palliata, as 
both fall within the range of morphological 
variation found in a single population 
(USNM 901676). Since Powell (1951) des- 
ignated P. palliata to be the type species of 
Pfefferia, we retain this name for this spe- 
cies, and synonymize the remaining nomi- 
na. 

Powell (1951:142) noted that the only 
radula he examined was abnormal, with two 
small, "incipient" cusps flanking the nor- 
mally tricuspid rachidian teeth. The radulae 
we examined were typical of Chlanidota. 

Juvenile specimens are bicolored, with 
the periostracum being dark chestnut brown 
above the periphery, and a pale, olive green 
below the periphery. The periostracum of 
adult specimens is thicker and uniformly 
chestnut brown in color. Conchologically 
this species is similar to C. (C) dense- 
sculpta, but may be readily distinguished on 
by the presence of weakly raised spiral 
cords, as well as by its distinctive opercu- 
lum. 

Chlanidota {Pfefferia) chordata 

(Strebel, 1908) 

Figs. 21, 22, 23; Table 8 

Pfefferia chordata Strebel, 1908:36-7, pi. 
3, fig. 41a-c; Powell, 1951:143; Carcel- 
les, 1953:193. 

Description. — Shell (Fig. 20) large for 
genus (to 35.3 mm), thick, solid, ovate- 
rounded, highly variable. Protoconch, early 
teleoconch whorls eroded in all specimens. 
Teleoconch with up to 4+ evenly rounded, 
very convex whorls. Shoulder not pro- 
nounced. Body whorl comprises 0.6-0.89 
of the total shell length. Suture strongly im- 
pressed, forming narrow channel between 
adapical spiral cord and previous whorl. 
Spiral sculpture variable, spiral cords (7-26 
on body whorl, 4-11 on penultimate whorl) 



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Fig. 19. Anatomy of Chlanidota {Pfejferia) palliata (Strebel, 1908). Specimen in fig. 17 J. A. Right and B. 
left lateral views of animal removed from shell (SL = 29.7 mm). C. Ventral, D. left lateral, and E. right lateral 
views of anterior alimentary system (salivary glands removed to show valve of Leiblein in E). F. Juncture of 
gland of Leiblein and esophagus. G. Ventral view of fused salivary glands. H. Dorsal view of stomach. I. 
Stomach, opened mid-dorsally. Scale bars = 5 mm for A-E, H-I; 2 mm for F, G. Abbreviations: ao, anterior 



VOLUME 112, NUMBER 2 



287 



41'W 40° 



56" 




South Georgia Island 







Q " 
50 



100 



150 



200 




300 



No. of spec.x 3 



Fig. 20. Geographic distribution and bathymetric range of Chlanidota (Pfefferia) palliata (Strebel, 1908). 
O = type locality; # = examined material. 



low, weak, and smooth to very tall and 
sharply-defined, forming corresponding spi- 
ral grooves on inner surface of outer Up. 
Cords of weakly sculptured specimens (Fig. 
20L, M) closely spaced, as wide or wider 
than intervening spaces. Cords of moder- 
ately sculptured specimens (e.g.. Fig. 20 A- 
C, F-G) Vi to Vs width of intervening spac- 
es. Cords of strongly sculptured specimens 
(Fig. 20D-E, K-J) more than Vi the width 
of intervening spaces. Adapical cord of 
strongly corded specimens may form nar- 
rowly concave subsutural rim. Fine spiral 
threads may be present on cords and in in- 
terspaces. Axial sculpture limited to weakly 
defined growth lines. Aperture tall (0.60- 
0.84 SL), ovate, deflected from shell axis 
by 16—19°. Outer lip thin to moderately 
thick, evenly rounded or scalloped, fragile 
to strong, sometimes slightly reflected. Col- 



umella less than half aperture length, weak- 
ly convex or straight, with strong siphonal 
fold. Callus usually thick, narrow, overlying 
parietal region, siphonal fasciole, grayish to 
brownish in color. Siphonal notch broad, 
dorsally recurved, with straight columellar 
and rounded apertural margins that form 
borders of fasciole. Ridge margin of fasci- 
ole pronounced, originating at apertural 
margin of siphonal notch, sometimes pro- 
truding through callus. Shell color chalky- 
white. Periostracum (Fig. 21) yellow-or- 
ange, olive to dark brown, nearly black, 
moderately to extremely thick, tightly ad- 
herent to the shell surface. Periostracum 
surface smooth or of closely adjacent, 
blade-like lamellae that overlay spiral 
sculpture without giving rise to hair-like 
projections, or hairy, producing one or sev- 
eral hairs at intersections of lamellae with 



aorta; ct, ctenidium; ddg, duct of the digestive gland: dg, digestive gland; dgL, duct of gland of Leiblein; gL, 
gland of Leiblein; h, heart; ht, cephalic tentacles; ig, intestinal groove; nep, nephridium; nr. circumoesophageal 
nerve ring; oe, oesophagus; op, operculum; os, osphradium; ov, ovary; poe, posterior oesophagus; pr, proboscis; 
prr, proboscis retractors; rhd, proboscis sheath; s, siphon; sd, salivary duct; sg, salivary gland; st, stomach; vL, 
valve of Leiblein. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 8. — Chlanidota (Pfefferia) chordata (Strebel, 1908). Measurements of shell characters. Linear mea- 
surements in mm {n = 10, including holotype). 



Character 


Mean 


a 


Range 


Holotype 


Shell Length (SL) 


27.9 


3.5 


23.2-34.4 


34.7 


Body Whorl Length (BWL) 


21.9 


3.7 


16.7-29.7 


29.0 


Aperture Length (AL) 


19.1 


2.9 


15.3-23.2 


2L7 


Shell Width (SW) 


21.1 


3.4 


17.2-26.7 


23.8 


Operculum Length (OL) 


12.0 


3.3 


7.7-16.2 


— 


BWL/SL 


0.79 


0.12 


0.60-0.89 


0.83 


AL/SL 


0.68 


0.05 


0.60-0.74 


0.62 


SW/SL 


0.75 


0.06 


0.68-0.86 


0.69 


OL/AL 


0.61 


0.09 


0.46-0.73 


— 


No. spiral cords on: 










Penultimate whorl 


5.7 


2.4 


4-11 


5 


Body whorl 


13 


6.3 


7-26 


12 



spiral cords. Operculum large (0.46-0.73 
AL), massive, typical of genus. 

Ultrastructure. — (Fig. 22C, E) Outer- 
most prismatic layer comprising spiral 
cords, greatly variable in thickness, ranging 
from 825 jxm in strongly corded specimens 
(Fig. C) to 58 |jLm in lightly corded shells 
(Fig. E). Middle, comarginal layer ranges 
from 60 to 210 |xm, inner, radial layer 
reaches —60 jjim in thickness. 

Anatomy. — Gross anatomy of C. (P.) 
chordata very similar to that of C. (P.) pal- 
liata. Radular ribbon long (0.66 AL), —720 
|jLm wide (0.036 AL), triserial (Fig. 22A- 
B), of —65 rows of teeth, of which 5 are 
nascent. Rachidian teeth with deeply arched 
base, nearly straight lateral sides, 3 large, 
robust cusps of equal-length. Lateral teeth 
with 3 cusps, outermost cusp largest, inter- 
mediate cusp thinnest and shortest, adjacent 
to inner cusp. Stomach U-shaped without 
obvious caecum, too poorly preserved to re- 
veal internal details. 

Type locality. — Cumberland Bay, South 
Georgia, 54°11'S, 36°18'W, in 252-310 m 
(Sveska Siidpolarexp., Sta. 34). 

Type material. — Holotype, SMNH 3660 

Material examined. — Holotype. RA^ Is- 
las Orcadas: South Georgia: Sta. 9, 
53°43'30"S, 37°30'06"W, 271-313 m, 12 
May 1975, 2 specimens, USNM 901677; 
Sta. 18, 54°02'30"S, 37°39'36'W, 60-71 m, 
15 May 1975, 3 specimens, USNM 901678; 



Sta. 19, 54°01'42"S, 37°40'00"W, 46-70 m, 
15 May 1975, 3 specimens + 1 shell, 
USNM 901679; Sta. 20, 54°00'06"S, 
37°40'36"W, 68-80 m, 15 May 1975, 4 
specimens, USNM 901680; Sta. 22, 
54°02'48"S, 37°23'42"W, 66-75 m, 16 May 
1975, 3 specimens + 6 shells, USNM 
881724; Sta. 25, 53°5r02"S, 36°49'03"W, 
199-247 m, 17 May 1975, 1 shell, USNM 
901681; Sta. 26, 53°43'06"S, 36°49'18"W, 
183-192 m, 17 May 1975, 3 specimens + 
3 shells, USNM 881756; Sta. 88, 
54°3r45"S, 36°48'42"W, 150-154 m, 7 Jun 
1975, 2 specimens + 1 shell, USNM 
896287; Sta. 89, 54°44'12"S, 37°11'12"W, 
225-265 m, 7 Jun 1975, 6 specimens + 7 
shells, USNM 881760; Sta. 90, 54°50'48"S, 
37°23'48"W, 223-227 m, 7 Jun 1975, 8 
shells, USNM 881762; Sta. 101, 
54°14'10"S, 37°54'20'W, 164-183 m, 10 
Jun 1975, 1 shell, USNM 901682. RA^ 
Prof. Siedlecki: South Georgia: Sta. 20, 
53°58'S, 38°42'W, 189-200 m, 2 Dec 1986, 
1 shell, USNM 901683; Sta. 37, 54°18'S, 
37°54'W, 158-194 m, 5 Dec 1986, 1 shell, 
USNM 897573; Sta. 83, 54°39'S, 35°49'W, 
98-127 m, 11 Dec 1986, 2 specimens + 4 
shells, USNM 897534; Sta. 84, 54°28'S, 
35°39'W, 231-249 m, 11 Dec 1986, 1 spec- 
imen, USNM 897536; Sta. 88, 54°15'S, 
35°51'W, 232-254 m, 11 Dec 1986, 2 
shells, USNM 897506; Sta. 90, 54°10'S, 
35°15'W, 242-262 m, 12 Dec 1986, 1 shell. 



VOLUME 112, NUMBER 2 



289 



USNM 896998; Sta. 91, 54°09'S, 35°55'W, 
218-227 m, 12 Dec 1986, 3 specimens, 
USNM 897450; Sta. 101, 53°47'S, 
36°34'W, 263-277 m, 2 shells, USNM 
897457; Sta. 105, 53°40'S, 36°48'W, 161- 
192 m, 14 Dec 1986, 2 specimens + 6 
shells, USNM 897515; Sta. 106, 53°44'S, 
36°51'W, 178-201 m, 14 Dec 1986, 5 
shells, USNM 896985. 

Distribution. — Off Southern Georgia 
(Fig. 22). Living specimens collected at 
depths from 46 to 313 m. 

Remarks. — Chlanidota (Pfefferia) chor- 
data is highly variable in shell morphology, 
with the extremities of the morphological 
range appearing very different (compare, 
e.g., Fig. 2 IK, J to L, M). Incrementally 
transitional forms ranging from nearly 
smooth to highly corded individuals were 
present in our samples (e.g.. Fig. 
21M«-^L<->F^^A^^D^^H<-^J). Specimens 
from depths (<100 m) (e.g., Fig. 21L, M) 
tend to have smaller shells with more nu- 
merous and more densely spaced spiral 
cords than specimens from deeper water 
(>150 m) (e.g.. Fig. 21D, H, J). Interme- 
diate specimens between these extremes oc- 
cur within single dredge hauls at both shal- 
low and deep stations. Specimens with in- 
termediate shell morphology might be con- 
fused with Chlanidota {Pfejferia) palliata 
(compare Fig. 21F-G with Fig. 17F-G). 

Chlanidota {Pfejferia) invenusta, 

new species 

Figs. 24-26; Table 9 

Chlanidota densesculpta (partim) — Dell, 
1990:183, fig. 306. 

Description. — Shell (Fig. 23) thick, sol- 
id, ovate-rounded. Protoconch, Wi upper te- 
leoconch whorls eroded. Teleoconch with 
~4 strongly convex whorls. Shoulder 
rounded. Suture adpressed, deep, wide, 
nearly channeled. Subsutural rim well pro- 
nounced, slightly convex. Spiral sculpture 
of weak cords (25-40 on body whorl, 5-14 
on penultimate whorl) constant in width 
adapically, of variable width and promi- 



nence below shoulder. Axial sculpture lim- 
ited to fine, often indistinct growth lines. 
Aperture of moderate height (0.60-0.72 
SL), elongate-ovate, deflected from the 
shell axis by 17-21°. Outer lip thin to thick, 
evenly rounded, weakly reflected in thick- 
lipped adults. Outer lip thin and evenly 
rounded, slightly deflected. Columella short 
{<V2 AL), straight, with strong, long si- 
phonal fold, siphonal region slightly elon- 
gated, inflected in some specimens. Callus 
narrow, thin to thick overlying parietal re- 
gion, siphonal fasciole, white when thin, 
greenish-gray at margins when thick. Si- 
phonal notch broad, dorsally recurved, with 
straight columellar, rounded apertural mar- 
gins forming borders of fasciole. Apertural 
margin of siphonal notch gives rise to weak 
ridge margin. Shell color grayish white. 
Periostracum (Fig. 24D) light-olive, mod- 
erately thick, tightly adherent to shell sur- 
face, forming very low, densely spaced la- 
mellae along the growth lines. Operculum 
(Fig. 23D, H, J, L) large (0.47-0.62 AL), 
oval, light to dark brown, apical end round- 
ed to weakly tapering, nucleus, terminal, ro- 
tated nearly perpendicular to long axis of 
operculum, posterior edge smooth (Fig. 
23d, J), or with weakly (Fig. 23H) to 
strongly (Fig. 23L) feathered lamellae. 

Ultrastructure. — (Fig. 25C) Outer pris- 
matic layer comprises spiral cords, variable 
in thickness, ranging from —80 jxm to 
— 140 jjim, depending on strength of cord. 
Middle layer comarginal (—120 jxm). Inner 
layer radial, reaching —50 ixm in thickness, 
is the last layer to be deposited, may not be 
evident in immature specimens (Fig. 25 C). 

Anatomy. — The gross anatomy of para- 
type 1 was examined, and found to be sim- 
ilar in all details to that of C {P.) palliata 
with the exception that the animal lacked 
eyes. Radular ribbon (Fig. 24A-B) long, 
13.6 mm (0.54 AL), --760 |xm wide (0.030 
AL), triserial, consisting of «80 rows, 6 of 
which are nascent. Rachidian teeth with 
arched base, straight lateral sides, 3 large, 
robust cusps, central cusp slightly larger 
that flanking cusps. Small intermediate ser- 



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Fig. 21. Chlanidota {Pfejferia) chordata (Strebel, 1908). A-C. Holotype, SMNH 3660, Sveska Siidpolarexp., 
Sta. 34, Cumberland Bay, South Georgia, 54°11'S, 36°18'W, in 252-310 m. D, E. RA^ Islas Orcadas, Sta. 89, 
South Georgia, 54°44'12"S, 37°11'12"W, in 225-265 m, USNM 881760. F, G. RA^ Islas Orcadas, Sta. 9, South 
Georgia, 53°43'30"S, 37°30'06"W, in 271-313 m, USNM 901677. H, I. R/V Prof. Siedlecki, Sta. 105, South 



VOLUME 112, NUMBER 2 



291 



ration present between central cusp and 
right flanking cusp of one specimen. Lateral 
teeth with 3 cusps, as in genus. 

Type locality. — South Georgia Island, 
53°31'12"S, 37°50'54"W, in 1267-1599 m 
(RA^ Islas Orcadas, Sta. 7). 

Type material. — Holotype, USNM 
881782, paratypes 1-9, USNM 880280, all 
from the type locality. 

Material examined. — South Georgia: 
RA^ Islas Orcadas, Sta. 7, 53°3ri2"S, 
37°50'54"W, 1267-1599 m, 11 May 1975, 
Holotype, USNM 881782, paratypes 1-9, 
USNM 880280, 17 shells, USNM 901684; 
STa. 27, 53°34.9'S, 36°47.8'W, 448-872 m, 
17 May 1975, 1 specimen, USNM 901685; 
RA^ Eltanin: Sta. 731, 53°35'S, 36°28'W, 
796-824 m, 11 May 1975, 13 specimens, 
USNM 896048; Sta. 734, 53°23'S, 
37°irW, 1299-1400 m, 11 May 1975, 4 
shells, USNM 870389, 8 specimens, 
USNM 896049. 

Distribution. — Known only from the 
northwestern coast of South Georgia (Fig. 
25). Living material was collected at depths 
ranging from 448 to 1599 m. 

Etymology. — invenustus (Lat.) — unat- 
tractive. 

Remarks. — Chlanidota (Pfefferia) inven- 
usta may be distinguished from the other 
species in the subgenus Pfefferia by its 
more inflated shell, with a stepped rather 
than conical spire, more numerous and finer 
spiral cords, a broader, more rounded ap- 
erture, and an operculum in which the 
feathering along the posterior edge is much 
narrower to entirely absent. Some speci- 
mens superficially resemble Chlanidota 
densesculpta, but may be distinguished on 
the basis of their larger operculum, thicker 
shell, more rounded and stepped spires, and 
proportionally smaller, more rounded, and 
more deflected apertures. 



The nine paratypes collected with the ho- 
lotype exhibit some variation in shell out- 
line and the number of spiral cords. Smaller 
specimens tend to have more rounded 
shells, while the number of spiral cords 
does not correlate with shell size. Opercula 
of the smallest specimens are yellow and 
translucent, and clearly show that the ter- 
minal nucleus is coiled, as in Neobuccinum. 

Species excluded from 
Chlanidota sensu lato 

Chlanidota smithi Powell, 1958 
Fig. 27 

Chlanidota smithi Powell, 1958:192, pi. 3, 
fig. 3; Dell, 1990:177. 

Type locality. — Off Enderby Land, 
65°50'S, 54°23'E, in 220 m (BANZARE, 
Sta. 42). 

Material examined. — Holotype, SAM 
D 15471 (Fig. 27). 

Published records. — BANZARE, Sta. 
41, Off Enderby Land, 65°48'S, 53°16'E, 
193 m (Powell, 1958:192). 

Remarks. — Powell (1958) placed this 
species in a group with Chlanidota dense s- 
culpta, C. vestita, and C pilosa, but distin- 
guished it from these taxa on the basis of 
its "disproportionally large," bulbous pro- 
toconch and the almost smooth surface on 
all whorls. Powell (1958) conjectured that 
Smith's (1902) record of C vestita from 
Cape Adare, 24-26 fm [45-48 m] might 
represent C smithi. However, Dell (1990: 
fig. 311) illustrated the Cape Adare speci- 
men and showed it to represent C vestita. 
Thus, records of C. smithi are restricted to 
Enderby Land. This species differs from 
typical Chlanidota in number of characters, 
most notably its very large size (SL = 51.5 
mm) and very large protoconch. Compari- 



<— 



Georgia, 53°40'S, 36°48'W, in 161-192 m, USNM 897515. J, K. RA^ Islas Orcadas, Sta. 26. South Georgia. 
53°43'06"S, 36°49'18"W, in 183-192 m, USNM 881756. L, M. Both from RA^ Islas Orcadas. Sta. 19. Off South 
Georgia Island, 54°0r42"S, 37°40'00"W, in 46-70 m, USNM 901679. Scale bar = 1 cm. 



292 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 22. Chlanidota {Pfejferia) chordata (Strebel, 1908). A. Dorsal and B. left lateral (45°) views of the 
central portion of the radular ribbon taken from specimen in fig. 21F-G. C. Shell ultrastructure and D. perios- 
tracum of the strongly corded shell. RA^ Prof. Siedlecki, Sta. 83, South Georgia, 54°39'S, 35°49'W, in 98-127 
m, USNM 897534. E. Shell ultrastructure and F periostracum of the weakly corded shell (fig. 20M). Scale bars 
= 200 |xm for A, B, E, F, 500 |jLm for C, 1000 \x.m for D. 



VOLUME 112, NUMBER 2 



293 



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son of the holotype of C smithi (Fig. 27) 
with the specimens of Neobuccinum eatoni 
Smith, 1875, clearly reveal a striking con- 
chological similarity between these species. 
Numanami (1996:146) noted that the ma- 
jority of specimens of A^. eatoni taken off 
Syowa Station (Enderby Land) are obese, 
with a large, well inflated body whorl. 
Comparison of Numanami's (1996:fig. 
94A, B, D-G) illustrations of Neobuccinum 
eatoni with the holotype of Chlanidota smi- 
thi leave no doubt that these two taxa are 
conspecific. 

Chlanidota gaini (Lamy, 1910) 

Sipho gaini Lamy, 1910:319; Lamy, 1911: 

7, pi. 1, figs. 7-8. 
Prosipho? gaini Thiele, 1912:262. 
?Chlanidota gaini Powell, 1951:142. 
Chlanidota gaini Carcelles, 1953:191; Dell, 

1990:177. 

Type locality. — Off King George Island, 
South Shetlands, in 420 m. 

Material examined. — Holotype, NMNH. 

Remarks. — ''Sipho'' gaini was provi- 
sionally attributed to Chlanidota by Powell 
(1951) and by Carcelles (1953). Dell (1990) 
considered Chlanidota gaini to be a species 
of uncertain affinity, known only from its 
holotype. 

Examination of the holotype, which had 
not been illustrated since its description, 
clearly indicates that this species is refer- 
able to the family Conidae (sensu Taylor et 
al., 1993), and is, in fact, the senior syno- 
nym of Belaturricula antarctica (Dell, 
1990). The composition and relationships 
of the genus Belaturricula are the subject 
of a separate report (Kantor & Harasewych 
1999). Hedley (1916) was the first to sur- 
mise the conoidean affinities of Sipho gaini, 
suggesting that it was closely related to 
Pontiothauma ergata Hedley, 1916. 

Chlanidota eltanini Dell, 1990 

Chlanidota eltanini Dell, 1990:184-5, figs. 
290, 292, 297, 314. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




No. of spec.x 4 



Fig. 23. Geographic distribution and bathymetric range of Chlanidota (Pfefferia) chordata (Strebel, 1908). 
O = type locality; # = examined material. 



Type locality. — East of Falklands Islands 
(Islas Malvinas), 51°58'S, 56°38'W, depth 
845-646 m (RA^ Eltanin, Sta. 558). 

Type material. — Holotype, USNM 
860124, 2 paratypes USNM 860125, 1 par- 
atype NMNZ MF.56613, from the type lo- 
cality. KIW Eltanin: Sta. 557, East of Falk- 
land Islands, 5r56'S, 56°39'W, 855-866 m, 
2 paratypes USNM 860126; Sta. 1521, 
South Atlantic Ocean, 54°09'S, 52°08'W, 
419-483 m, 2 paratypes USNM 860127, 1 
paratype NMNZ MF56614. 

Remarks. — ^Dell (1990) described Chlan- 
idota eltanini, C. bisculpta and C polys- 
peira from the Magellanic Province. While 
these three species are likely congeneric, 
they differ significantly from species as- 
signed to the subgenera Chlanidota and 
Pfefferia. Conchologically, these species 
are readily distinguished by their small (15- 
16 mm), slender, thick shell, very thin, 
smooth periostracum, and large (0.6 AL), 
coiled operculum. We were able to examine 
the anatomy of Chlanidota bisculpta, as 
well as of a closely related but as yet un- 
described species, and found differences in 



radular morphology, stomach shape, size of 
gland of Leiblein, and penis morphology. 
We provisionally consider these species to 
be more closely related to Neobuccinum, 
and exclude them from Chlanidota. 

A revision of this group will be pub- 
lished separately. 

Chlanidota bisculpta Dell, 1990 

Chlanidota bisculpta Dell, 1990:185, figs. 
291, 294, 295, 312. 

Type locality. — ^Burdwood Bank, 53°08'S, 
59023 'W, in 578-567 m (RA^ Eltanin, Sta. 
340). 

Type material. — Holotype, USNM 
860128; R/V Eltanin: Sta. 557, East of 
Falkland Islands (Islas Malvinas), 51°56'S, 
56°39'W, 855-866 m, 6 paratypes USNM 
860129, 1 paratype NMNZ MF56615; Sta. 
740, off Cape Horn, 56°06'S, 66°19'W, 
384-494 m, 2 paratypes USNM 860130, 1 
paratype NMNZ MF56616. 

Remarks. — See remarks under Chlani- 
dota eltanini Dell, 1990. 



VOLUME 112, NUMBER 2 



295 




Fig. 24. Chlanidota (Pfefferia) invenusta new species. A-C. holotype, USNM 881782, RA^ Islas Orcadas, 
Sta. 7, South Georgia, 53°31'12"S, 37°50'54"W, in 1267-1599 m. D. Operculum of holotype. enlarged. E. 
Paratype 1, (anatomy studied), and F. paratype 5, USNM 880280, both from type locality. G, H. Specimen with 
narrowly rimmed operculum. I, J, M. Specimens with smooth operculum. K, L. Specimen with broadly rimmed 
operculum, all from RA^ Eltanin, Sta. 734. South Georgia. 53°23'S, 37°irw, in 1299-1400 m. USNM 896049. 
Scale bar = 1 cm for shells. 5 mm for opercula. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 25. Chlanidota (Pfejferia) invenusta, new species. A. Dorsal and B. left lateral (45°) views of the central 
portion of the radular ribbon of paratype 1 (Fig. 24E). C. Shell ultrastructure, and D. periostracum of the paratype 
5 (Fig. 24F). Scale bar = 200 |xm for A, B, 100 (xm for C, 500 ixm for D. 



Chlanidota polyspeira Dell, 1990 

Chlanidota poly speira Dell, 1990:186, figs. 
292, 293, 313. 

Type locality. — Patagonian Shelf, 
54°04'S, 63°35'W, depth 247-293 m (RA^ 
Eltanin Sta. 369). 

Type material — ^Holotype, USNM 860131, 
3 paratypes, USNM 860132, 1 paratype 
NMNZ MF.56617, all from the type local- 
ity. 

Remarks. — See remarks to Chlanidota 
eltanini Dell, 1990. 



Discussion 

Buccinoidean classification has tradition- 
ally been based on a combination of shell, 
opercular, and radular characters, and these 
structures are well documented for the ma- 
jority of Antarctic genera (e.g., Thiele 
1904, Powell 1951, Hain 1989, Dell 1990, 
Numanami 1996). Ponder (1974) reported 
that most organ systems were weakly dif- 
ferentiated among buccinoidean higher 
taxa, and that it was difficult to identify an- 
atomical features that could be used to seg- 



VOLUME 112, NUMBER 2 



297 




No. of spec. X 5 



Fig. 26. Geographic distribution and bathymetric range of Chlanidota {Pfejferid) invenusta, new species. O 
= type locality, # = examined material. 

regate them reliably. More recently, Hara- gland, that vary among buccinoideans and 

sewych (1990) identified several characters, may be phylogenetically informative. While 

including the morphology of the gland of there have been a substantial number of an- 

Leiblein and the presence of an ingesting atomical investigations of boreal (e.g., Da- 




Fig. 27. Chlanidota smithi Powell, 1958. Holotype, SAM D15471, BANZARE Sta. 42, Off Enderby Land, 
65°50'S, 54°23'E, in 220 m. Scale bar = 1 cm. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



kin 1912; Lus 1978, 1981, 1989; Kantor 
1988, 1990), tropical (e.g., Marcus & Mar- 
cus 1962, 1964) and deep-sea (e.g.. Ponder 
1968, Harasewych 1990) buccinoidean 
taxa, the anatomy of Antarctic representa- 
tives has not previously been studied in any 
detail. 

It is not surprising that Chlanidota shares 
many anatomical features with the boreal 
family Buccinidae, among them similar ar- 
rangements of the mantle cavity organs, re- 
productive system, and many characters of 
the digestive system. Characters such as a 
long, thick proboscis and fused salivary 
glands occur in Chlanidota as well as in the 
buccinid taxa Habevolutopsius (Kantor 
1990) and Ancistrolepidinae (Kantor 1988). 
The stomach of Chlanidota, however, dif- 
fers from that of most Buccinidae in being 
broadly U-shaped and in lacking a caecum. 
The caecum, or posterior mixing area, is 
usually well developed in boreal Buccini- 
dae, and is sometimes very large (e.g., in 
Volutopsius, subfamily Volutopsiinae), 
making the stomach appear sac-like. The 
stomach of Southern Hemisphere taxa, such 
as Penion adustus (PhiUppi, 1845) (Ponder 
1973) and Ratifusus mestayerae (Iredale 
1915) (Ponder 1968), which had been as- 
signed to Buccinulidae by Powell (1961), 
also lack a caecum. 

The northern Atlantic buccinid Colus 
gracilis (da Costa 1778) is unusual in lack- 
ing a gastric caecum (Smith 1967). How- 
ever, its stomach differs from the stomach 
of Chlanidota in having a cuticularized 
shield and a distinct sphincter between the 
oesophagus and stomach. The stomach ep- 
ithelium of Colus gracilis is glandular and 
darker than the esophageal epithelium, 
while in Chlanidota the epitheUal Hning of 
the esophagus is dark and that of the stom- 
ach is Ughter. 

Unlike all other Chlanidota and Pfejfer- 
ia, the stomach of Chlanidota (Chlanidota) 
paucispiralis has a small enlargement (Fig. 
14, pm) corresponding in position to the 
caecum of Neptune a (Smith 1967), Buccin- 
um (Medinskaya 1993) and other boreal 



buccinids. It is still not clear, whether the 
presence of a gastric caecum is an advanced 
or plesiomorphic character, as the structure 
occurs in many neogastropod families (e.g., 
Buccinidae, Volutomitridae, Muricidae, Mi- 
tridae) but is absent in others (e.g., Col- 
umbelUdae, Volutidae, Cancellariidae and 
all Conoidea). 

The posterior oesophagus of Chlanidota 
is greatly enlarged, thick-walled, and Hned 
with tall epitheUal folds (e.g. Fig. 5H), 
which are darker than the rest of esophageal 
epitheUum. The transition between the pos- 
terior oesophagus and the stomach is clear- 
ly evident on dissection, as it is marked by 
a color change of the epithelium near the 
posterior duct of the digestive gland. 

The morphology of the radula and major 
organ systems is remarkably invariant in all 
of the species of Chlanidota and Pfefferia 
that we examined. The only exceptions 
were the presence of a small caecum in C 
(C.) paucispiralis and of separate rather 
than fused salivary glands in C (C.) sig- 
neyana. PreHminary dissections of other 
Antarctic buccinulids generally regarded as 
being closely related to Chlanidota (e.g., 
Neobuccinum eatoni, ''Chlanidota'' bis- 
culpta, see above) revealed significant dif- 
ferences in their alimentary and male repro- 
ductive systems, providing support for re- 
ducing Pfefferia to a subgenus of Chlani- 
dota. 

The morphology of the operculum of 
Chlanidota (Pfefferia) is both striking and 
unique within Neogastropoda. The mecha- 
nism by which the characteristic "feath- 
ered" margin is formed along the posterior 
edge of the operculum is readily inferred. 
Newly secreted layers of conchioUn are 
broader adapically than abapically, and are 
attached to the posterior edge of the oper- 
culum only along their proximal margins. 
While operculum morphology is constant in 
C (P.) palliata and C (P.) chordata, it 
varies considerably in the newly described 
C. (P.) invenusta. Juveniles of C. (P.) in- 
venusta have an operculum with a "feath- 
ered" margin. The width of this margin de- 



VOLUME 112, NUMBER 2 



299 



creases with increasing shell size, the mar- 
gin disappearing entirely in large specimens 
(see Fig. 24L^H^J^D). The size of the 
operculum of C. (P.) invenusta (0.47—0.62 
AL) is larger than in any species of Chlan- 
idota (Chlanidota) (0.18-0.37 AL), com- 
parable to that of C. (P.) chordata (0.46— 
0.73 AL), but smaller than that of C. (P.) 
elata (0.68-0.78 AL). The large operculum 
of Chlanidota {Pfejferid) allows it to seal 
the aperture tightly when the animal with- 
draws into its shell (e.g., Figs. 17F, 21H), 
while its "feathered" posterior margin pro- 
vides a flexible, tight seal against the outer 
lip. In contrast, the small size of the oper- 
culum in Chlanidota s.s. precludes its util- 
ity for sealing the aperture. The decrease in 
relative size and "degree of feathering" of 
the operculum in C (P.) invenusta with in- 
creasing shell size, suggests that the ability 
to seal the aperture tightly diminishes in im- 
portance as the animal grows. 

While the anatomy and radular morphol- 
ogy are largely invariant in Chlanidota and 
Pfefferia, shell shape and the strength and 
number of spiral cords and threads vary 
considerably. The limited number of spec- 
imens available to previous workers has, in 
a few instances, led to the description of 
species based on differences that now ap- 
pear to fall within a broad continuum of 
morphological variability revealed by larger 
sample sizes spanning wider geographical 
areas. 

As restricted in this paper, the genus 
Chlanidota is confined to the Antarctic re- 
gion, with only the type species ranging 
slightly beyond the Antarctic Convergence. 
The present revision reduces the number of 
recognized species within Chlanidota. Nev- 
ertheless, it remains the second most di- 
verse buccinoidean genus (following Pro- 
sipho) in Antarctic waters. 

Of the eight recognized species in the ge- 
nus Chlanidota, only C (C) signeyana has 
a broad circum-Antarctic distribution (al- 
though records from the poorly sampled 
Ross Quadrant are lacking). Except for a 
single record of C (C) vestita from Cape 



Adare, all remaining species of Chlanidota 
are restricted to subAntarctic islands, with 
two species of Chlanidota sensu stricto, and 
all three species of Pfefferia endemic to 
South Georgia. Of the species inhabiting 
subAntarctic islands, most are sublittoral, 
and only Chlanidota {Pfefferia^ invenusta is 
restricted to bathyal depths. In contrast, the 
circum-Antarctic C signeyana has a broad 
bathymetric range (30-1116 m). 

The protoconchs of all specimens of 
Chlanidota were severely eroded, hamper- 
ing inference of developmental mode. 
However, most polar species, including 
buccinoideans, develop directly without a 
pelagic larval stage (Thorson 1946). Thus, 
it seems likely that the island populations 
of Chlanidota are vicariant isolates from a 
more broadly ranging species. 

Acknowledgments 

We wish to express our thanks to Dr. An- 
ders Waren for loan of the Strebel type ma- 
terial, to Dr. Philippe Bouchet for access to 
Lamy's types, and to Kathie Way and Joan 
Pickering for the loan of Powell's types at 
BMNH. Dr. W Ziedler South AustraUan 
Museum made available the type specimen 
of C smithi. We are grateful to Ellen Strong 
for translation of the relevant portions of 
Strebel (1908), and to Dr. Guido Pastorino 
for his assistance at various stages of man- 
uscript preparation, especially with digital 
photography. We also thank Susann Braden 
for her assistance with scanning electron 
microscopy. 

The junior author gratefully acknowledges 
Mrs. Mabs Mango for her kind hospitahty 
during the preparation of this manuscript. 

This research was supported by a grant 
from the NSF-USAP United States Antarc- 
tic Program [Contract No. OPP-9509761]. 

Literature Cited 

Bouchet, P., & A. Waren. 1985. Revision of the North- 
East Atlantic bathyal and abyssal Neogastro- 
poda. excl. Turridae.^Bollettino Malacologico, 
Supplement 1:121-296. 



300 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



. 1986. Taxonomical notes on tropical deep wa- 
ter Buccinidae with descriptions of new taxa. — 
Memoires du Museum national d'Histoire na- 
turelle, serie A, Zoologie 133:457-499. 

Cantera, J. R., & R M. Amaud. 1984. Les Gastero- 
podes prosobranches des lies Kerguelen et 
Crozet (sud de 1' Ocean Indien) comparaison 
ecologique et particularites biologique. — 
CNFRA publications no. 56:5-169. 

Carcelles, A. 1953. Catalogo de la malacofauna Ant- 
arctica Argentina. — Anales del Museo Nahuel 
Huapi 3:150-250. 

Cemohorsky, W. O. 1977. The taxonomy of some 
southern ocean Mollusca (Gastropoda) mainly 
Antarctic and sub Antarctic. — Records of the 
Auckland Institute and Museum 14:105-119. 

Dakin, W. J. 1912. Buccinum (The Whelk). — Proceed- 
ings of the Liverpool Biological Society 20: 
253-367. 

David, L. 1934. Zoologische Ergebnisse der Reisen 
von Dr Kohl-Larsen nach den subantarktischen 
Inseln bei Neuseeland und nach Siidgeorgien. — 
Senckenbergiana 16:126-137. 

Dell, R. K. 1964. Marine Mollusca from Macquarie 
and Heard Islands. — Records of the Dominion 
Museum 4(2):267-301. 

. 1990. Antarctic Mollusca. — Bulletin of the 

Royal Society of New Zealand 27:1—311. 

Finlay, H. J. 1928. The Recent Mollusca of the Chat- 
ham Islands. — Transactions of the New Zealand 
Institute 59:232-286, pis. 38-43. 

Gaillard, J. M. 1971. Mollusques, Premier Partie, Am- 
phineures et Gasteropodes Prosobranches. In- 
vertebres de I'infralittoral rocheux dans 
I'archipel de Kergielen. — CNFRA (Comite Na- 
tional Fran^ais des Recherches Antarctiques) 
30:89-111. 

Habe, T, & J. Sato. 1973. A classification of the family 
Buccinidae from the North Pacific. — Proceed- 
ings of the Japanese Society of the Systematic 
Zoology 8:1-8, pis. 1, 2. 

Hain, S. G. 1989. Beitrage zur Biologic der beschalten 
Molluscen (Kl. Gastropoda und Bivalvia) des 
Weddellmeeres, Antarktis. — Dissertation zur 
Erlangung des Grades eines Doktors der Natur- 
wissenschaften, Bremen: 3-298. 

Harasewych, M. G. 1990. Studies on bathyal and abys- 
sal Buccinidae: 1. Metula fusiformis Clench and 
Aguayo, 1941.— The Nautilus 104(4): 120-130. 

Hedley, C. 1916. Mollusca. — Australian Antarctic ex- 
pedition 1911-1914 Scientific Reports. C. — Zo- 
ology and Botany 4(1): 1-80. 

Horikoshi, M., T Hoshiai, & Y. Naito. 1979. Nanky- 
okukai no teisei dobutsu no bunrui seitaigaku- 
teki kenkyu (Systematic and ecological studies 
on Antarctic benthos). Showa 54-nendo Kyodo 
Kenkyu Hokokusho (Progress Report of Joint 



Research for 1979), Tokyo, National Institute of 
Polar Research: 20-26. 

Kaicher, S. D. 1990. Buccinidae, Pack IV. Card cata- 
logue of world-wide shells 57:5792-5897. 

Kantor, Yu. I. 1988. Gastropods of the subgenus An- 
cistrolepis (Clinopegma) (Gastropoda, Buccin- 
idae) of the Okhotsk Sea. — Zoological Journal 
67(8): 1126-1 140. (In Russian). 

. 1990. Gastropods of the subfamily Volutop- 

siinae (Buccinidae) of the World Ocean. — Mos- 
cow, Nauka, 178 pp. (In Russian). 

. 1996. Phylogeny and relationships of Neogas- 

tropoda. Pp. 221-230 in J. D. Taylor ed., Origin 
and evolutionary radiation of Mollusca, Cente- 
nary Symposium of the Malacological Society of 
London. Oxford University Press, 392 pp. 

, & M. G. Harasewych. 1999. Rediscovery of 



the Antarctic species Sipho gaini Lamy, 1910 
(Gastropoda: Neogastropoda) with remarks on 
its taxonomic position. Antarctic Research (in 
press). 

Lamy, E. 1910. Mission dans I'Antarctique dirige par 
M. le Dr Charcot (1908-1910): collections re- 
cueillis par M. le Dr J. Liouville: Gastropodes 
prosobranches et scaphopodes. — Buletin du 
Museum d'Histoire naturelle 16:318-324. 

. 1911. Gastropodes Prosobranches, Scapho- 
podes et Pelecypodes. Deuxieme Expedition an- 
tarctique fran9aise (1908-1910) — Science Na- 
turelles: documents scientifiques: 1-32. 

Lus, V. Ya. 1978. A new genus and species of buccin- 
ids (Mollusca: Prosobranchia: Buccinidae) from 
lower abyss of Izu-Bonin Trench in the Pacif- 
ic. — Trudy Instituta Okeanologii AN SSSR 
(Proceedings of the Institute of Oceanology of 
the USSR Academy of Sciences) 114:147-157 
(in Russian). 

. 1981. On the abyssal species Sipho (Siphon- 

orbis) danielsseni (Friele) and Mohnia mohni 
(Friele) (Gastropoda: Buccinidae). — Trudy In- 
stituta Okeanologii AN SSSR (Proceedings of 
the Institute of Oceanology of the USSR Acad- 
emy of Sciences) 115:126-140 (in Russian). 

. 1989. More deep-sea gastropods from the 



Buccinacea (Neogastropoda: Buccinacea 
s.l.). — Trudy Instituta Okeanologii AN SSSR 
(Proceedings of the Institute of Oceanology of 
the USSR Academy of Sciences) 123:151-165 
(in Russian). 

Marcus, E., & E. Marcus. 1962. On Leucozonia nas- 
sa. — Boletim da Faculdade de Filosofia, Cien- 
cias e Letras, Universidade de Sao Paulo 232, 
Zoologia 24:11-30. 

, & . 1964. Studies on Columbellidae. — 

Boletim da Faculdade de Filosofia, Ciencias e 
Letras, Universidade de Sao Paulo, Zoologia 24 
(261):335-402. 

Markham, C. R. 1912. Review of the results of twenty 



VOLUME 112, NUMBER 2 



301 



years of Antarctic work originated by the Royal 
Geographical Society. — The Geographical Jour- 
nal 39:575-580. 

Martens, E. 1878. Einige Conchylien aus den kalteren 
Meeresgegenden der sudlichen Eidhalfte. — Sit- 
zungsbericht der Gesellschaft naturfirschender 
Freude zu Berlin 1878:20-26. 

. 1881. Conchologische Mittheilungen als Fort- 

setzung der Novitates conchologicae 1:1-101. 

. 1885. Vorlaufige Mitteilungen iiber die Mol- 

lusken-Fauna von Slid Georgien. — Sitzungsber- 
icht der Gesellschaft naturfirschender Freude zu 
Berlin 1885:89-94. 

. 1904. Die beschalten Gastropoden der 

deutschen Tiefsee-Expedition 1898-1899. A. 
Systematisch-geographischer Teil. — Wissen- 
schaftliche Ergebnisse der deutschen Tiefsee- 
Expedition auf dem Dampfer "Valdivia" 7:1- 
146, pis. 1-5. 

, & G. Pfeffer. 1886. Die Mollusken-Fauna von 



of Mollusca, Centenary Symposium of the Mal- 
acological Society of London. Oxford Univer- 
sity Press, 392 pp. 
-, & A. Waren. 1988. Appendix. Classification 



Siid-Georgien nach den Ausbeute der Deutsch- 
en Station. — Jahrbuch der Hamburgischen Wis- 
senschaftlichen Anstalten 3:65-135. 

Medinskaya, A. I. 1993. Anatomy if the stomach of 
some Neogastropoda from the offshore zone of 
the Japan Sea. — Ruthenica, Russian Malacolog- 
ical Journal 3(1): 17-24. 

Numanami, H. 1996. Taxonomic study on Antarctic 
gastropods, collected by Japanese Antarctic re- 
search expeditions. — Memoirs of National In- 
stitute of Polar Research, Series E, Biology and 
Medical Science, 39:1-244. 

Oliver, P. G. 1983. Notoficula Thiele, a neotenous ge- 
nus of eratoid gastropod from Antarctica. — 
British Antarctic Survey Bulletin 61:1-6. 

Ponder, W. F. 1968. Anatomical notes on two species 
of the Colubrariidae (Mollusca, Prosobran- 
chia). — Transactions of the Royal Society of 
New Zealand. Zoology 10(24):217-223. 

. 1971. A review of the New Zealand recent 

and fossil species of Buccinulum Deshayes 
(Mollusca: Neogastropoda: Buccinidae). — Jour- 
nal of the Royal Society of New Zealand 1(3/ 
4):23 1-283. 

. 1973. A review of the Australian species of 

Penion Fischer (Neogastropoda: Buccinidae). — 
Journal of the Malacological Society of Austra- 
lia 2:401-428. 

. 1974. The origin and evolution of Neogastro- 
poda. — Malacologia 12:295-338. 

. 1982. A new genus and species of Buccinidae 

(Mollusca: Gastropoda) from the Continental 
Slope of eastern and southern Australia. — Jour- 
nal of the Malacological Society of Australia 
5(3-4):201-207. 

, & D. R. Lindberg. 1996. Gastropod phylog- 

eny-challenges for the 90's. Pp. 135-154 in J. 
D. Taylor, ed.. Origin and evolutionary radiation 



of the Caenogastropoda and Heterostropha — a 
list of their family-group names and higher taxa. 
Pp. 288-328 in W. F Ponder, ed., Prosobranch 
Phylogeny. Malacological Review, Supplement 
4, 346 pp. 

Powell, A. W. B. 1929. The Recent and Tertiary spe- 
cies of the genus Buccinulum in New Zealand, 
with a review of related genera and families. — 
Transactions of the New Zealand Institute 60: 
57-101, pis. 1-4. 

. 1951. Antarctic and sub Antarctic Mollusca: 

Pelecypoda and Gastropoda. — Discovery Re- 
ports 26:47-196, pis. 5-10. 

. 1957. Mollusca of Kerguelen and Macquarie 

Islands. B. A. N. Z. — Antarctic Research Ex- 
pedition, Reports, Series B 6(7): 107-150. 

. 1958. Mollusca from the Victoria-Ross Quan- 

drants of Antarctica. B. A. N. Z. — Antarctic Re- 
search Expedition, Reports, Series B 6(9): 166- 
215. 

. 1960. Antarctic and sub Antarctic Mollusca. — 

Records of the Auckland Institute and Museum 
5:117-193. 

. 1961. Shells of New Zealand, 4th edition. 

Whitcombe & Tombs, Ltd., Christchurch, 203 
pp. 

. 1976. Shells of New Zealand, 5th Revised 

edition. Whitcombe & Tombs, Ltd., Christ- 
church, 154 pp. 

. 1979. New Zealand Mollusca, Marine, Land 



and Freshwater Shells. Collins Ltd., Auckland, 
500 pp. 

Smith, E. A. 1902. VII. Mollusca. — Report on the col- 
lections of natural history made in the Antarctic 
regions during the voyage of the "Southern 
Cross", London:201-213. 

Smith, E. H. 1967. The Neogastropod stomach, with 
notes on the digestive diverticula and intes- 
tine. — Transactions of the Royal Society of Ed- 
inburgh, 67(2):23-42. 

Strebel, H. 1908. Die Gastropoden. — Wissenschaftli- 
che Ergebnisse der Schwedischen Siidpolar-Ex- 
pedition (1901-1903) 6(1):1-112. pis. 1-6. 

Taylor, J. D., Yu. I. Kantor, & A. V. Sysoev. 1993. 
Foregut anatomy, feeding mechanisms, relation- 
ships and classification of Conoidea ( = Toxo- 
glossa) (Gastropoda). — Bulletin of the natural 
History Museum. London (Zoology) 59:125- 
169. 

Thiele, J. 1904. B. Anatomisch-systematische Unter- 
suchungen einiger Gastropoden. — Wissenschaf- 
tliche Ergebnisse der deutschen Tiefsee-Expe- 
dition auf dem Dampfer "Valdivia" 7:147-180, 
pis. 6-9. 



302 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



-. 1912. Die antarktischen Schecken und 
Muscheln. — Deutschen Siidopolar-Expedition 
1901-1903 13:183-285. 

-. 1929. Handbuch der systematischen Weich- 



tierkunde 1(1): 1-376. Gustav Fischer, Jena. 
Thorson, G. 1946. Reproduction and larval develop- 
ment of Danish marine bottom invertebrates. — 
Meddelelser Era Komissionen for Danmarks 
Fiskeri-Og Havunders0gelser, Serie: Plankton 
4(l):l-523. 



Tryon, G. W. 1881. Manual of Conchology, III. Tri- 
tonidae, Fusidae, Buccinidae:l-310. 87 pis. 

Watson, R. B. 1886. Report on the Scaphopoda and 
Gastropoda. — Report on the Scientific Results 
of the Voyage of H. M. S. Challenger during 
the years 1873-1876. Zoology 15:i-vi, 1-756, 
pis. 1-50. 

Wenz, W 1938-1944. Gastropoda. Teil 1, Allgemeiner 
Teil und Prosobranchia. — Handbuch der Palao- 
zoologie 6:1-1639. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):303-312. 1999. 

Hydroid and medusa stages of the new species Ectopleura obypa 
(Cnidaria: Hydrozoa: Tubulariidae) from Brazil 

Alvaro E. Migotto and Antonio C. Marques 

(AEM) Centre de Biologia Marinha, Universidade de Sao Paulo, Caixa Postal 83, 

11600-970 Sao Sebastiao, SP, Brazil; 

(ACM) Departamento de Biologia, Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto, 

Universidade de Sao Paulo, Av. Bandeirantes 3900, 14040-901 Ribeirao Preto, SP, Brazil 

Abstract. — Ectopleura obypa, a new species referable to the family Tubu- 
lariidae, is described from the southeastern coast of Brazil. Specimens were 
collected on experimental panels and other artificial substrates at depths from 
about 1 to 15 m. The hydroid stage is a remarkable green color when blasto- 
styles are developed. A free medusa, with 2 opposed capitate marginal tentacles 
and 8 meridional tracks of nematocysts on the exumbrella, is present in the 
life cycle. 



During a study of hydroid recruitment on 
experimental plates in waters of southeast- 
em Brazil we discovered a species of Ec- 
topleura, not yet described, remarkable for 
its bright green color. Several other speci- 
mens of the species were found at various 
times later on other artificial substrates in- 
cluding nylon ropes, buoys, and iron pipes. 
Specimens kept in the laboratory and cul- 
tured under controlled conditions liberated 
free medusae. 

The genus Ectopleura L. Agassiz, 1862, 
a basal clade of the family Tubulariidae, has 
20 valid species divided in two phyloge- 
netic groups distinguishable basically by 
their cnidomes and stolon growth pattern 
(Petersen 1990:160). There is much confu- 
sion between the genera Tubularia and Ec- 
topleura, the first having sessile gonophores 
and the latter traditionally considered as 
having free medusae (e.g., Millard 1975: 
31-32; Bouillon 1985:112). However, Pe- 
tersen (1979:120; 1990:160) redefined the 
tubulariid genera based on features of the 
hydroid stage. His classification of tubular- 
iid polyps included: the solitary Zyzzyzus, 
with a thin perisarc and tuber-like attach- 
ment of hydrocaulus; the solitary Tubularia 
with firm perisarc and producing either eu- 



medusoid or cryptomedusoid gonophores 
of the symmetrical type; the solitary Hy- 
bocodon, producing fixed asymmetrical 
gonophores; the colonial or solitary Ecto- 
pleura with firm perisarc and producing ei- 
ther free medusae or fixed gonophores; the 
solitary or colonial Ralpharia with firm 
perisarc and producing reduced medusae 
with an internal raised collar around bell 
opening; and the solitary Bouillonia with 
barrel- shaped hydranth, firm perisarc and 
reduced medusae. 

Material and Methods 

Hydroids of Ectopleura obypa, new spe- 
cies, were collected on a number of sub- 
strates (experimental plates, ropes, buoys, 
iron pipes) on the coast of Sao Sebastiao, 
Sao Paulo State, Brazil. Type material, 
studied here, was collected by skin and 
SCUBA diving from a light buoy in the Sao 
Sebastiao Channel. Part of the material was 
anesthetized in a 1:1 solution of 7.5% 
MgCl2 solution and seawater, and preserved 
in 4% formaldehyde solution in seawater; 
part was kept alive in the laboratory. Live 
hydranths were examined under a stereo- 
microscope, and newly liberated medusae 



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were transferred to finger bowls and kept in 
constant temperature chambers at 19, 24 
and 26°C with a 12 h light/ 12 h dark pho- 
toperiod. Water in the bowls was changed 
daily, shortly after medusae were measured 
and fed with Anemia nauplii. 

Stems for scanning electron microscopy 
(SEM) were post-fixed in 1% OSO4, dehy- 
drated in a graded series of ethanol, dried 
in a critical-point drier, and sputter-coated 
with gold. Nematocyst types and their dis- 
tributions were determined using a light mi- 
croscope with interference-contrast. Nema- 
tocyst nomenclature used here is that of 
Mariscal (1974) and Millard (1975). Only 
capsules of undischarged nematocysts were 
measured. Abbreviations are: AM (collec- 
tion of A. Migotto); MZUSP (Museu de 
Zoologia da Universidade de Sao Paulo, 
Sao Paulo); USNM (United States National 
Museum, Smithsonian Institution, Washing- 
ton D.C.). 

Results and Discussion 

Genus Ectopleura L. Agassiz, 1862 

Ectopleura obypa, new species 

Figs. 1-3; Tables 1-2 

Material. — Holotype, 1 hydroid colony, 
2.0 cm high, Sao Sebastiao Channel, Sao 
Sebastiao, Sao Paulo State, Brazil 
(23°49.86'S, 045°25.28'W), 4 Jan 1996, 5 
m, colony with gonophores and several hy- 
dranths growing epibiotically on barnacles 
and mussels, on light buoy, collected by 
hand, coll. A. E. Migotto (MZUSP- 12.8 13); 
paratypes (all from same locality as holo- 
type), medusae liberated in the laboratory 6 
Jan 1996 from hydroid colony collected 4 
Jan 1996, 6 medusae ca. 2 h old (AM1325); 
2 medusae, 6 d old (MZUSP- 12.8 14); 4 
medusae, 7 d old (AM1327); 26 medusae, 
34 d old (AM1328); one hydroid colony 35 
mm high, on mussels and ascidian, with 
gonophores, 16 Jan 1996, 2 m, coll. A. C. 
Morandini (AM 13 23); one hydroid colony 
25 mm high, on mussel, with gonophores, 
17 Jun 1996, 2-3 m, coll. A. E. Migotto 
(AM1324; USNM 99449); 84 medusae ca. 



3 h old, liberated in the laboratory on the 
same day of collection, from colony col- 
lected 17 Jun 1996 (AM1329). 

Description. — colony up to 35 mm high, 
arising from branched stolons. Hydrocaulus 
unbranched, increasing in width from base 
to distal end, covered with a smooth light 
brown perisarc without annuli. Neck region 
long (Figs. IC, 3 A), flexible and contrac- 
tile, covered by a filmy perisarc; extended 
neck up to 3.5 times longer than contracted; 
in live specimens, neck entirely white or 
light pink, with white pigments concentrat- 
ed in 2 narrow longitudinal bands. Hy- 
dranth vasiform, pink-white and transpar- 
ent, with one whorl of 16-30 aboral tenta- 
cles and one whorl of 14-29 oral tentacles 
(Figs. lA, 2A, 3A). Aboral tentacles long, 
filiform, laterally flattened (Fig. lA), with 
nematocysts concentrated along adoral sur- 
face, and on tip. Oral tentacles short, cir- 
cular in cross section, basal part adnate to 
hypostome (Fig. ID), with nematocysts 
concentrated in a terminal capitulum and 1- 
3 irregular swellings on the adoral surface 
of the tentacle (Fig. IB, D); free part of oral 
tentacles slightly fused at base (Fig. IB). 
Medusa buds on 4-14 (usually 7-10) blas- 
tostyles arising above aboral tentacles; old- 
er medusa buds at ends of blastostyles (Fig. 
3B). Blastostyles yellow-green, composed 
of a main stem on which medusae arise ei- 
ther directly or from irregular branches; 
main stem may bifurcate. In large speci- 
mens the developed blastostyles hang be- 
tween aboral tentacles. 

Newly-Uberated medusa with thin umbrel- 
la, dome-shaped and without a distinct apical 
projection but with a small apical canal, with 
8 slightly raised exumbrellar, meridional 
nematocyst tracks; with 4 marginal bulbs, 2 
of which with opposed tentacles; marginal 
tentacles with a terminal knob and 1-3 ab- 
axial nematocyst clusters (Fig. 2B); abaxial 
nematocyst cluster grows and involves the 
tentacle (Figs. 2D, 3C). Manubrium tubular, 
without Ups, with nematocysts around mouth, 
and about % the length of beU cavity. Apical 



VOLUME 112, NUMBER 2 



305 




Fig. 1. Scanning electron micrographs of Ectopleura obypa, new species. A, fronto-lateral view of hydranth, 
note incipient blastostyles just above aboral tentacles; B, oral view of hydranth; C, neck region; the arrow points 
to the groove where filmy perisarc is secreted; D, lateral view of hypostome and oral tentacles. Scale bar. A. B 
= 100 |jim; C, D = 50 ixm. 



canal short. Color: umbrella transparent; 
subumbrella green when illuminated side- 
ways; manubrium pink- white; marginal bulbs 
milky- white with small red dots. 

The types and measurements of nema- 
tocysts of hydranth and medusa are listed 
below. Morphological character variation of 
many colonies and newly released medusae 
are in Tables 1 and 2, respectively. 



Nematocysts. — (in jxm) (Fig. 3D) 

Hydranth 

Oral tentacle: 

Stenotele - 13.5 - 14.0 X 12.0 - 

13.0 (abundant) 
Stenotele - 6.0 - 8.5 X 6.0 - 7.5 

(abundant) 
Microbasic eurytele - 9.0 — 10.0 X 
4.0 - 4.5 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Photographs of live specimens of Ectopleiira obypa, new species. A, lateral view of a hydranth with 
incipient blastost\les: B. lateral view of a newly-liberated medusa: C. lateral view of a 6-day-old medusa: D. 
detail of a tentacle of a 7-day-old medusa. Scale bar, A = 1.0 mm: B, D = 100 |xm: C = 200 |xm. 



VOLUME 112, NUMBER 2 



307 




Fig. 3. Ectopleura obypa, new species. A, outline diagram of hydranth, traced from a whole mount prepa- 
ration; the upper arrow points to the region where the film perisarc is secreted; the lower arrow indicates the 
beginning of the firm perisarc; B, lateral view of a blastostyle with several well developed medusa buds; C. 
sequence in the development of a medusa tentacle; C4 is the oldest stage, just before liberation from blastostyle; 
D. nematocysts; Dl. stenotele; D2, basitrichous isorhiza; D3, microbasic mastigophore; D4, desmonome. Scale 
bar, A = 0.5 mm; B = 100 |xm; C = 50 \xm: D = 10 fxm. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Morphological character variation of hydroids of Ectopleura obypa, new species. 

Measurements 
[average ± SD (range)] 



Hydrocauli 
Length (mm) 
Distal diameter (mm) 
Proximal diameter (mm) 

Hydranth 

Total length (mm) 

Length from the base of aboral tentacles to the apex of hypostome (mm) 

Diameter at the level of aboral tentacles (mm) 

Tentacles 
Aboral 

Number 
Length (mm) 

Oral 

Number 
Length (mm) 

Gonosome 

Number 
Length (mm) 



18.5 ± 6.9 (0.87-30.3) 
0.20 ± 0.03 (0.13-0.26) 
0.34 ± 0.07 (0.22-0.60) 

1.85 ± 0.55 (1.18-3.30) 
1.41 ± 0.42 (0.88-2.70) 
1.09 ± 0.30 (0.62-1.80) 



23.8 ± 2.9 (16-30) 
2.50 ± 0.59 (0.91-3.90) 

20.8 ± 3.25 (14-29) 
0.79 ±0.16 (0.42-1.20) 

8.55 ± 2.10 (4-14) 
0.91 ± 0.55 (0.15-2.10) 



Aboral tentacle: 

Stenotele - 13.5 - 14.0 X 12.0 - 

13.0 (rare) 
Stenotele - 6.0 - 8.5 X 6.0 - 7.5 

(abundant) 
Microbasic eurytele — 9.0 — 10.0 X 

4.0 — 4.5 (abundant; more at tip) 
Desmoneme - 4.0 - 5.0 X 3.5 - 4.0 
(abundant) 
Newly released medusa 

Basitrichous isorhiza — 8.0 — 10.5 

8.0 - 9.0 
Stenotele - 8.0 - 8.5 X 7.0 - 8.0 
Microbasic eurytele — 9.0 — 10.5 

4.0 - 4.5 
Desmoneme — 6.0 X 4.0 
Etymology. — The species name is de- 
rived from the word "obypa", in the lan- 
guage of the Brazilian Tupi natives, mean- 
ing green, the color of the blastostyles. It is 
pronounced ' ' 6-bi-pa' ' . 

Development of medusa. — ^Medusae kept 
in the laboratory, despite several different 
combinations of temperature, salinity, type 
of food and volume of culturing water, grew 
slowly and did not change significantly in 



X 



X 



morphology. Medusae usually did not cap- 
ture food by themselves. Ingestion occurred 
only when the offered naupUus was touched 
to the manubrium; we never saw the trans- 
fer process of the prey from the tentacles to 
the mouth of the medusa. From the several 
cultures started only one batch had medusae 
with gonads, although even these were not 
completely developed (medusae 25 days 
old, from colony collected 17 Jun 1996, 
kept at 19°C). We are aware that these un- 
successful efforts probably indicate the cul- 
tivated medusae do not represent the spe- 
cies under natural conditions, but we decid- 
ed to include our observations on growth 
rate and morphology. Developing gonads 
were first observed in 7 to 9 d old medusae. 
These medusae were 1380-1440 jxm high, 
1050-1150 fxm in maximum diameter and 
had tentacles with 6 knobs, attaining ca. 
800 [xm in length (Fig. 2C, D). They Uved 
16 more days, maintaining approximately 
the same size and general features except 
for a slight growth of the gonads, which 
bulged a little around the middle of the ma- 
nubrium. Tentacles developed up to 7 nem- 



VOLUME 112, NUMBER 2 



309 



Table 2. — Morphological character variation of newly-released medusae of Ectopleura obypa, new species, 
from three different colonies. 



[average ± SD (range)] 



Height (|xm) 

Maximum diameter (ixm) 
Diameter at base (ixm) 
Diameter of aperture (ixm) 
Length of manubrium (ixm) 



594.5 ± 101.9 (460-780) 

516.0 ± 82.1 (396-660) 

344.5 ± 69.6 (240-440) 

120 

510.2 ± 141.7 (276-780) 



atocyst knobs. The number of tentacles did 
not change in any specimens during 
growth; nor was there any indication that 
the two other perradial bulbs would give 
rise to tentacles. 

Remarks. — This description of Ectopleu- 
ra obypa, new species, especially color, size 
and general shape, is based mainly on live 
material. Colors in fixed material disap- 
peared. Some anesthetized and fixed hy- 
dranths, although well preserved, did not 
keep their natural shape: hydranths from 
AM 1324 are inflated and most have the up- 
per part of the neck dilated. Fixed medusae 
usually have longer manubria than live 
specimens, and perradial and interradial 
grooves appear in the umbrella (as also ob- 
served for E. sacculifera, see Brinckmann- 
Voss 1970:28). 

The long, flexible and contractile neck 
enables the hydranth to bend in any direc- 
tion. In still water under laboratory condi- 
tions, the hydranths performed regular cir- 
cular movements interspersed with strong 
contractions of the neck. We could not as- 
certain the number of ridges in the endo- 
derm of hydrocaulus, but we suppose the 
two bands of white pigments present along 
the neck indicate the existence of two of 
these ridges in E. obypa. The blastostyles 
were so vivid a green, even early in devel- 
opment, that when fully developed the 
whole hydranth appeared green to the na- 
ked eye. The medusa started pulsating 
hours before liberation, already capable of 
catching and ingesting food, and the ma- 
nubrium moved peristaltically and vermic- 
ularly. 



Besides E. obypa, new species, two other 
species of Ectopleura were recorded from 
the region of Sao Sebastiao: Ectopleura du- 
mortieri (Van Beneden, 1844) and Ecto- 
pleura warreni (Ewer, 1953) (see Migotto 
& Silveira 1987:100-103, Migotto 1996: 
24-25). The polyp of Ectopleura dumorti- 
eri is solitary, and has free medusae with 4 
marginal tentacles; E. warreni has fixed 
gonophores. 

Ectopleura obypa is referred to a large 
group of species diagnosed by the presence 
of free medusae with 8 meridional nema- 
tocyst tracks in the umbrella, issuing in 
pairs from tentacle bulbs (Schuchert 1996: 
107). Complementary characters are: even- 
ly rounded umbrella; four radial canals and 
tentacle bulbs; manubrium short, not ex- 
tending beyond bell margin; medusa tenta- 
cles moniliform or with abaxial nematocyst 
clusters (Schuchert 1996:107). The cni- 
dome of species having free medusae is 
characterized by microbasic euryteles (be- 
sides other types such as stenoteles and des- 
monemes), a type of nematocyst not present 
in the species of Ectopleura with fixed gon- 
ophores (see Petersen 1990:160; note that 
the author interchanged the distribution of 
anisorhiza and microbasic euryteles in fig- 
ure 19, characters 2 and 6). 

Among species of Ectopleura with free 
medusae, two subgroups are clearly recog- 
nized on the basis of the number of mar- 
ginal tentacles in the medusa: 2 tentacles 
and 4 tentacles. 

Species described as having 4 perradial 
tentacles are E. americana Petersen, 1990, 
E. dumortieri (Van Beneden, 1844), E. be- 



310 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






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VOLUME 112, NUMBER 2 



311 



thae (Warren 1908) and E. indica Petersen, 
1990. 

Besides E. obypa there are six species of 
Ectopleura having medusae with 2 opposed 
tentacles: Ectopleura wrighti Petersen, 
1979, E. pacifica Thomely, 1900, E. mayeri 
Petersen, 1990, E. minerva Mayer, 1900, E. 
sacculifera Kramp, 1957, and£^. xiamenesis 
Zhang & Lin, 1984 (see Table 3). The last 
three of these are known only from the me- 
dusa stage. Ectopleura sacculifera and E. 
xiamenensis do not have an apical canal, 
contrasting in this with E. minerva and E. 
obypa. Moreover, Ectopleura sacculifera is 
clearly distinct from the rest by having a 
thick umbrella and pendent gonads on the 
manubrium wall (Kramp 1957:7, plate 3, 
figs. 1-3). Neither an apical canal nor an 
apical projection were mentioned by 
Brinckmann-Voss (1970:25-27) in the de- 
scription of young and adult medusae of E. 
wrighti (as E. larynx), and we suppose they 
are not present in this species. Nematocyst 
clusters of the marginal tentacles in medu- 
sae of E. obypa are clearly diffrent from 
those of E. minerva Mayer, 1900 (Mayer 
1900, fig. 125) and E. sacculifera (cf. 
Brinckmann-Voss 1970:28, fig. 29.1), 
which are exclusively abaxial and do not 
involve the tentacle as in E. obypa (exclu- 
sively abaxial clusters of nematocysts are 
also present in an unidentified species of 
Ectopleura with a two-tentacled medusa de- 
scribed by Schuchert 1996:112, fig. 67). 

It is difficult to link the species known 
only by the medusoid stage with polypoid 
stages already described in the literature, 
due to the lack of life cycle studies. Except 
for E. wrighti (see Brinckmann-Voss 1970: 
25, as E. larynx) and E. obypa, data on me- 
dusa stages are few and based on immature 
specimens still attached to blastostyles. Ec- 
topleura wrighti is the only species in 
which medusae obtained from hydroid col- 
onies were raised through maturity (Brinck- 
mann-Voss 1970:25). Petersen (1990:166) 
described Ectopleura mayeri (=E. pacifica 
of Calder 1988:53-55) as a "new species", 
even though he acknowledged the possibil- 



ity of its being the hydroid stage of E. mi- 
nerva Mayer, 1900, because the medusa of 
the first "has not been reared to a stage 
where it can be identified with certainty". 
Hydroid stages of other species with two- 
tentacled medusae have many similar fea- 
tures: general size, number of oral and ab- 
oral tentacles, and number of blastostyles. 
They all have 2 endodermal ridges in the 
hydrocaulus, but this character is shared 
with other species having different numbers 
of tentacles. The aboral tentacles of E. oby- 
pa and E. pacifica are slightly adnate to the 
hypostome, in which they differ from E. 
wrighti and E. mayeri. Ectopleura obypa 
differs from E. pacifica in the morphology 
of its oral tentacles (see Table 3), and is 
unique in having flattened aboral tentacles. 

Acknowledgements 

We thank the staff of the Laboratorio de 
Microscopia Eletronica and of the Depar- 
tamento de Zoologia, Instituto de Biocien- 
cias, USP, for assistance with scanning elec- 
tron microscopy. We also thank the two re- 
viewers (Dr. Dale Calder and Dr. Paul F. S. 
Cornelius) whose comments improved the 
manuscript. This work was supported by 
the Fundagao de Amparo a Pesquisa do Es- 
tado de Sao Paulo (FAPESP grants n° 95/ 
3022-5; 96/10544-0 and 97/04572-4). 

Literature Cited 

Agassiz, L. 1862. Contribution to the natural history 
of the United States of America, vol IV. Little, 
Brown and Company, Boston, 380 pp. 

Bouillon, J. 1985. Essai de classification des Hydro- 
polypes — Hydromeduses (Hydrozoa-Cnidar- 
ia). — Indo-Malayan Zoology 2(l):29-243. 

Brinckmann-Voss, A. 1970. Anthomedusae/Athecata 
(Hydrozoa, Cnidaria) of the Mediterranean, part 
I. Capitata. — Fauna e Flora del Golfo di Napoli 
39:1-96. 

Calder, D. R. 1988. Shallow water hydroids of Ber- 
muda. — The Athecatae. — Royal Ontario Muse- 
um, Life Science Contributions 148:1-107. 

Ewer, D. W. 1953. On a new tubularian hydroid from 
Natal. — Annals of the Natal Museum 12(3): 
351-357. 

Kramp. R L. 1957. Hydromedusae from the Discovery 
collections. — Discovery Reports 29:1-128. 



312 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Mariscal, R. N. 1974. Nematocysts. Pp. 129-178 in L. 
Muscatine & H. M. Lenhoff, eds., Coelenterate 
Biology. Reviews and new perspectives. Aca- 
demic Press, New York. 501 pp. 

Mayer. A. G. 1900. Some medusae from the Tortugas, 
Florida. — Bulletin of the Museum of Compar- 
ative Zoology at Harvard College 37:13-82. 

Migotto, A. E. 1996. Benthic shallow-water hydroids 
(Cnidaria, Hydrozoa) of the coast of Sao Se- 
bastiao, SP, Brazil, including a checklist of Bra- 
zilian hydroids. — Zoologische Verhandelingen 
306:1-125. 

Migotto, A. E., & E L. da Silveira. 1987. Hidroides 
(Cnidaria, Hydrozoa) do litoral sudeste e sul do 
Brasil: Halocordylidae, Tubulariidae e Cory- 
morphidae. — Iheringia, serie Zoologia 66:95— 
115. 

Millard, N. A. H. 1975. Monograph on the Hydroida 
of southern Africa. — Annals of the South Afri- 
can Museum 68:1—513. 

Petersen, K. W. 1979. Development of coloniality in 
Hydrozoa. Pp. 105-139 in G. Larwood & B. 



Rosen, eds.. Biology and systematics of colo- 
nial organisms. The Systematics Association, 
Academic Press, New York, 589 pp. 

. 1990. Evolution and taxonomy in capitate hy- 
droids and medusae. — Zoological Journal of the 
Linnean Society 100:101-231. 

Schuchert, P. 1996. The marine fauna of New Zealand: 
athecate hydroids and their medusae. — New Ze- 
land Oceanographic Institute Memoir 106:1- 
159. 

Thomely, L. R. 1900. The hydroid zoophytes collected 
by Dr. Willey in the southern seas. — A. Willey's 
Zoological Results 4:451-457. 

Van Beneden, P. J. 1844. Recherches sur 
I'embryogenie des tubulaires, et I'histoire na- 
turelle es differents genres de cette famille qui 
habitent la Cote d'Ostende. — Nouveaux Me- 
moires de TAcademie Royale des Sciences, des 
Lettres et des Beaux Arts de Belgique 36:1— 
207. 

Warren, E. 1908. On a collection of hydroids, mostly 
from the Natal coast. — Annals of the Natal 
Government Museum 1:269-355. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):313-318. 1999. 

Striatodoma dorothea (Cheilostomatida: Tessaradomidae), a new 
genus and species of bryozoan from deep water off California 

Judith E. Winston and Stace E. Beaulieu 

(JEW) Virginia Museum of Natural History, Martinsville, Virginia 24112, U.S.A.; 

(SEB) Marine Biology Research Division, Scripps Institution of Oceanography, 

La Jolla, California 92093-0202, U.S.A. 

(Current address) Applied Ocean Physics and Engineering Department, 

Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, U.S.A. 

Abstract — Striatodoma dorothea, a new genus and species of cheilostomate 
bryozoan, is described from material found attached to hexactinellid sponges 
and pogonophoran tubes at an abyssal station (4100 m depth) off central Cal- 
ifornia. Members of this new genus can be distinguished from other members 
of the family Tessaradomidae by the presence of biserial, rather than quadris- 
erial branches, and a peristomial sinus, rather than an enclosed spiramen. Two 
other Pacific species, Diplonotos striatum Canu & Bassler, 1930, and Tessar- 
adoma bifax Cheetham, 1972, are transferred to Striatodoma. 



Although bryozoans have been identified 
from deep-sea stations down to 8300 m, re- 
views by Schopf (1969) and Hay ward 
(1981) have shown that only a tiny portion 
of the deep ocean floor has been sampled 
for the group. Nothing is known of the 
deep-sea bryozoan fauna of the eastern Pa- 
cific with the exception of three species 
found at two stations in the eastern Pacific 
between Acapulco and Panama during the 
Galathea Expedition (Hayward 1981). Re- 
cently, as part of a study by Beaulieu 
(1998) of hard substrate epifauna at abyssal 
depths off California, an attempt was made 
to identify all taxa attached to biogenic 
structures that protruded from the soft sed- 
iment of the sea floor. Individual stalks of 
the hexactinellid sponge Hyalonema sp. and 
individual tubes of the pogonophoran Uni- 
brachium sp. were sampled in tube cores 
using the submersible Alvin at 4100 m 
depth. Two of the approximately 140 spe- 
cies found attached to the sponge stalks and 
pogonophoran tubes were bryozoans. One 
is a ctenostome, Arachnidium hippothooi- 
des Hincks, 1862. The other is an undes- 
cribed genus and species of deep-sea chei- 



lostome which we name and describe be- 
low. 

Tessaradomidae Jullien, 1903 
Striatodoma, new genus 

Diagnosis. — Tessaradomidae character- 
ized by subcylindrical, proximally thick- 
ened branches with two series of zooids, 
longitudinally striated calcification, a spi- 
ramen in close association with zooid peri- 
stome, rows of marginal pores, some of 
them replaced by oval avicularia, and a 
subglobular imperforate ovicell. Striatodo- 
ma differs from Tessaradoma in possessing 
branches made up of two, rather than four 
series of zooids, and in confluence of the 
spiramen with a peristomial sinus. 

Type species. — Striatodoma dorothea, 
new species, by present designation. 

Additional species of Striatodoma. — Tes- 
saradoma bifax Cheetham, 1972 and Di- 
plonotos striatum Canu & Bassler, 1930. 

Etymology. — The first part of the genus 
name is from the Latin, striatus = fur- 
rowed, channeled, descriptive of the striated 
appearance of colony walls. The second 



314 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



part, doma, is derived from the Greek 
8a)fxa, 8a)|jLaTo^, house, roof, to parallel the 
other genus name in the family, Tessara- 
doma. Gender neuter. 

Remarks. — ^During the last hundred years 
seven Uving and fossil species of deep wa- 
ter cheilostomes have been described and 
placed in the family Tessaradomidae Jul- 
lien, 1903, and genus Tessaradoma Nor- 
man 1869, type species Onchopora borealis 
Busk, 1860. Jullien defined the family as 
having erect or encrusting colonies and zo- 
oids, a tubular peristome with a tubuHform 
spiramen opening into its base, and with a 
small, spherical, imperforate ovicell, also 
opening into the peristome above the zo- 
oidal operculum. The genus Tessaradoma 
has a colony form consisting of erect, un- 
jointed cyHndrical branches arising from an 
encrusting base, zooids with a projecting 
peristome and prominent spiramen, imper- 
forate ovicells obscured by increasing cal- 
cification, and adventitious avicularia (Hay- 
ward & Ryland 1979). The type species, 
Tessaradoma boreale, has a Recent distri- 
bution in both the North and South Atlantic 
and is also known from Neogene fossil lo- 
caUties in western Europe and the Mediter- 
ranean (Cheetham 1972, Lagaaij & Cook 
1973). Its colonies are erect, rigid, and 
spreading, with quadriserial branches con- 
sisting of elongate oval zooids, arranged 
back to back in alternating pairs. The zooid 
primary orifice is concealed by a tubular 
peristome with a spiramen tube near its 
base, its opening projecting from the zooid 
frontal surface about one third of the way 
down the frontal waU. Wall calcification is 
granular, becoming striated around the con- 
spicuous marginal pores. Oval adventitious 
avicularia are also developed in the mar- 
gins, particularly lateral to the spiramen. 
The ovicell is small, smooth, imperforate, 
subspherical (slightly broader than long), 
and though conspicuous in young zooids 
becomes increasingly immersed in calcifi- 
cation as zooids age (Hayward & Ryland 
1979). 

Hayward (1981) described Tessaradoma 



brevissima from the Tasman Sea, but Gor- 
don (1989), who recorded T. brevissima 
from additional deep water localities off 
New Zealand, placed the species in the ge- 
nus Galeopsis (family CeUeporidae) on the 
basis of the following characters: paired av- 
icularia in close association with peristome 
and sinus, and an ovicell with a central tab- 
ulate or fenestrate area and labellum. Two 
South African species, Tessaradoma bispi- 
ramina and Tessaradoma circella, de- 
scribed by Hayward & Cook (1979), and 
the Indonesian species Tessaradoma bipa- 
tens, described by Harmer (1957) also seem 
to belong in this group. 

In contrast, the new Pacific species de- 
scribed below, as weU as two other Pacific 
species, the Eocene Tessaradoma bifax 
from the smaU western Pacific Island of 
Tonga (Cheetham 1972), and the Recent 
Tessaradoma striatum from the Galapagos 
(Canu & Bassler 1930), differ from them in 
the furrowed appearance of frontal wall cal- 
cification, the subspherical imperforate ovi- 
cell and the association of the spiramen 
with the peristomial sinus. These Pacific 
species appear to be related to Tessarado- 
ma, but their shared features indicate they 
should be grouped with the new species de- 
scribed below in a new genus, Striatodoma. 
The Eocene Tessaradoma bifax described 
by Cheetham (1972) from Tonga, is very 
similar in morphology to Striatodoma do- 
rothea, with biserial branches, tubular peri- 
stome, and with lateral pores and avicularia 
making a sinuate double trail along the 
sides of branches. In this species, however, 
the sinus (at least in the fragmentary ma- 
terial available) does not become calcified 
into a tube. In addition, its oviceU is not as 
prominent as that of S. dorothea, appearing 
only as a sUght enlargement of the peri- 
stome of the maternal zooid, and a swelling 
of the frontal shield of the distal zooid. The 
Recent Diplonotos striatum collected from 
the Galapagos at 1251 m depth (Canu & 
Bassler 1930) also belongs in this group. 
Based on his studies of syntype material, 
Cheetham (1972) transferred that species to 



VOLUME 112, NUMBER 2 



315 




Fig. 1. Striatodoma dorothea new species. All illustrations from portions of holotype colony. CASIZ 1 13579, 
attached to pogonophoran tube collected 4100 m depth onAlvin Dive 2828 at 34°42'N, 123°00'W, off California. 
(A) Largest colony fragment showing colony form and branching pattern [scale = 2 cm]. (B) Zooid morphology 
[from left to right] autozooids in side and front view; ovicelled zooids in front and side view [scale bar = 0.6 
mm]. (C) Closeup of peristome, showing spiramen sinus [scale bar =150 ixm]. (D) Closeup of mature ovicelled 
zooids, showing constricted peristome and completely enclosed sinus [scale bar = 0.270 mm]. (E) Closeup of 
lateral avicularium, showing diagonally tilted rostrum and calcified pivotal hinges [scale bar =100 jxm]. 



Tessaradoma on the basis of its similarity 
to T. bifax, remarking that, contrary to 
Canu and Bassler's original description, the 
primary orifice does lie at the base of a 
peristomial shaft, but that shaft is almost 
completely immersed in the thickened fron- 
tal shield. Its chief difference from the other 
two species lies in the extreme development 
of frontal wall calcification, in which stri- 
ation becomes rugosity and external zooid 
boundaries are lost. The positions of lateral 
avicularia and pores may still be faintly dis- 
cerned, however, and match the pattern in 
the other two species. Like S. dorothea, the 



sinus in S. striatum may become completely 
enclosed to form a tube. 

Striatodoma dorothea new species 
Fig. 1 

Holotype. — California Academy of Sci- 
ences, CASIZ 113579, attached to pogo- 
nophoran tube collected as stalk no. 3 
(Beaulieu 1998) at 34°42'N, 123°00'W, 
4100 m depth, 17 Sep 1994. 

Paratypes. — Virginia Museum of Natural 
History 567; from stalk 4, —4100 m, 
PULSE 22 Cruise, Chief Scientist Kenneth 



316 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Measurements (in mm) of holotype and paratype specimens of Striatodoma dorothea. 



Character 




Range 


Mean 


SD 


n 


Zooid Length 




0.364-0.501 


0.444 


0.038 


12 


Zooid Width 




0.191-0.273 


0.231 


0.027 


12 


Orifice Length (autozooid) 




0.073-0.100 


0.085 


0.009 


12 


Orifice Width (autozooid) 




0.091-0.109 


0.102 


0.008 


12 


Orifice Length (ovicelled zooid) 


0.073 


— 


— 


2 


Orifice Width (ovicelled zooid) 


0.100 


— 


— 


2 


Ovicell Length 




0.109-0.155 


0.134 


0.016 


6 


Ovicell Width 




0.182-0.210 


0.196 


0.011 


6 


Avicularia Length 




0.036-0.055 


0.044 


0.007 


11 


Avicularia Width 




0.027-0.046 


0.036 


0.004 


11 


Branch Width (Distal end) 




0.291-0.337 


0.301 


0.025 


12 


Branch Width (Basal end) 




0.364-0.728 


0.558 


0.103 


9 



L. Smith, Jr., Scripps Institution of Ocean- 
ography, Alvin Dive 2828, 17 Sep 1994. 
Water temperature 1.2°C, 34°42'N, 
123°00'W; CASIZ 113580, stalk 37, 
34°56'N, 123°07'W, 4100 m, PULSE 24 
FVGR, 16 Feb 1995; CASIZ 113581, stalk 
33, 34°42'N, 123°00'W, 4100 m, PULSE 
25, Dive 2920, 30 Apr 1995; CASIZ 
113582, stalk 12, 34°42'N, 123°00'W, 4100 
m, PULSE 22, Dive 2834, 23 Sep 1994; 
collector for all, Stace Beaulieu. 

Etymology. — dorothea, the Latinized 
spelling of Dorothy, used as a noun in ap- 
position. The species is named in honor of 
Dorothy E Soule, in recognition of her el- 
egant studies of Pacific bryozoans and of 
her active stewardship of California marine 
environments. 

Diagnosis. — Characterized by subcy- 
Hndrical branches made up of two series of 
rectangular zooids, with longitudinally stri- 
ated calcification, rows of marginal pores, 
with occasional pores replaced by oval av- 
icularia, short peristome with spiramen en- 
closed in proximal peristomial sinus, and 
relatively prominent ovicell. 

Description. — Colony erect, rigidly cal- 
cified, unjointed, broadly branching in a 
planar fashion, up to several cm in size, the 
two biggest branch fragments of the largest 
specimen found (the holotype), measuring 
3 cm h X 7 cm w and 5 cm h X 5 cm w, 
respectively (Fig. lA). Attached to stalks of 
deep water glass sponges and pogonopho- 



ran tubes by an encrusting base. Zooids 
elongate, rectangular, growing back to back 
in two alternating longitudinal series (Fig. 
IB). Frontal wall convex, with faint longi- 
tudinal striations, most of its surface im- 
perforate, but with a row of small oval 
pores just inside zooid lateral margins. Pri- 
mary orifice transversely oval, surrounded 
by a short peristome (Fig. IC). No out- 
wardly visible spiramen; instead, the peri- 
stome of young zooids has a proximal si- 
nus, which is encircled by calcification as 
the zooid ages, becoming increasingly tu- 
bular and projecting. Ovicells are smoothly 
calcified and globular, reaching the height 
of the peristome opening, which becomes 
slightly narrowed in fertile zooids (Fig. 
ID). About one pore per zooid is replaced 
by an oval adventitious avicularium, about 
50 |ULm in length. In side view the sinuate 
double track of pores and avicularia is dis- 
tinctive (Fig. IE, B). In basal regions of the 
colony zooid openings are calcified over 
and branches become thickened. Some of 
the avicularia and pores are calcified over 
also. Branch thickness (the depth of two 
back to back zooids) averages 0.301 mm in 
zooids near the growing edge of branches 
and 0.558 mm in zooids near the colony 
base. Zooid measurements are summarized 
in Table 1. 

Distribution and ecology. — The species 
was observed and/or collected at 4060- 
4100 m depths off California, between lat- 



VOLUME 112, NUMBER 2 



317 



itudes 34°38' and 34°56'N and longitudes 
122°59' and 123°15'W. General area: 220 
km west of Point Conception, CA. Beaulieu 
(1998) collected a total of 35 tube core 
samples at the abyssal station and found 
seven colonies of S. dorothea (one colony 
per substrate; Table 2). The species was at- 
tached to pogonophoran tubes, the basal 
spicules of Hyalonema sp., and to other or- 
ganisms that were attached to the host sub- 
strate. The branches of S. dorothea also 
provided substrate on which other species 
attached (Table 2). 

In order to determine the abundance of 
S. dorothea at the abyssal station, all colo- 
nies large enough to identify with certainty 
were enumerated in photographic transects 
of the sea floor (procedures described in 
Beaulieu 1998). We photographed approx- 
imately 9 km of the sea floor (total of seven 
transects) in which we encountered 55 col- 
onies of S. dorothea. Of these, 53 were at- 
tached to dead Hyalonema spicules, one to 
a pogonophoran tube, and one to an un- 
identified structure. The encrusting bases of 
most of the colonies were attached at the 
middle of the host substrate, elevating the 
colonies about 10 cm above the sea floor. 
Density estimates (no. colonies per unit 
area of sea floor) were calculated using the 
computer program DISTANCE (Laake et 
al. 1994). Mean density estimates for the 
individual transects ranged between 3 and 
8 colonies per 1000 m^. Only about 2% of 
the biogenic structures enumerated in pho- 
tographic transects appeared to be colo- 
nized by S. dorothea. However, 20% of the 
structures collected from the sea floor had 
S. dorothea attached; therefore, the density 
estimates from the photographic transects 
may be an order of magnitude low. 

Discussion. — Tessaradomids belong to 
the fauna of the outer continental shelf and 
slope. The 4100 m depth recorded for S. 
dorothea in this study is the deepest re- 
corded, but it is not much deeper than the 
3700 m recorded for T. boreale. However, 
T. boreale is found in much shallower wa- 
ter in the Arctic (70 m depth), and Cheet- 



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318 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ham (1972) suggested that distribution of 
such deep-sea species may be temperature 
controlled, limited to water temperatures 
between 2° and 13° C. This paper and Beau- 
lieu (1998) provide a glimpse of how erect 
bryozoans with an attached base may sur- 
vive on the muddy deep sea floor. But as 
yet, these species are known only from a 
few broken fragments, collected at widely 
scattered localities by various expeditions. 
Better understanding of their ecology as 
well as clarification of their relationships 
with shallow water species must await the 
kinds of collections that can provide mate- 
rial adequate for the detailed anatomic 
study necessary for phylogenetic analysis. 

Acknowledgments 

We would like to thank K. Smith and the 
pilots of Alvin for making the collections 
and in situ observations possible. S. Beau- 
lieu was supported by a National Defense 
and Engineering Grant and by NSF grant 
OCE92- 17334 to K. Smith. We are also in- 
debted to Dr. W C. Banta (American Uni- 
versity) and Dr. A. H. Cheetham (National 
Museum of Natural History) for their help- 
ful reviews of the manuscript. 

Literature Cited 

Beaulieu, S. E. 1998. The ecology of glass sponge 
communities in the abyssal N.E. Pacific. Un- 
published Ph.D. thesis. University of California, 
San Diego, 206 pp. 

Busk, G. 1860. Zoophytology. Shetland Polyzoa col- 
lected by Mr. Barlee. — Quarterly Journal of Mi- 
croscopical Science 8:143-145. 

Canu, R., & R. S. Basslen 1930. The Bryozoan fauna 



of the Galapagos Islands. — Proceedings of the 
U.S. National Museum 76, Art. 13:1-78. 

Cheetham, A. H. 1972. Cheilostome Bryozoa of Late 
Eocene Age from Eua, Tonga. — U.S. Geologi- 
cal Survey Professional Paper 640-E:l-26. 

Gordon, D. P. 1989. The marine fauna of New Zea- 
land: Bryozoa: Gymnolaemata (Cheilostomida 
Ascophorina) from the western South Island 
continental shelf and slope. — New Zealand 
Oceanographic Institute Memoir 97:1-158. 

Harmer, S. F. 1957. The Polyzoa of the Siboga Expe- 
dition, part 4, Cheilostomata Ascophora II. — 
Leyden, Siboga Expedition Reports, 28d:641- 
1147. 

Hay ward, P. J. 1981. The Cheilostomata (Bryozoa) of 
the deep sea. — Galathea Report 15:21-68. 

, & R L. Cook 1979. The South African Mu- 

seumis Meiring Naude Cruises, part 9. Bry- 
ozoa. — Annals of the South African Museum 
79:43-130. 

, & J. S. Ryland. 1979, British Ascophoran 

Bryozoans. London, Academic Press, 312 pp. 

Hincks, T 1862. A catalogue of the zoophytes of south 
Devon and south Cornwall. — Annals and Mag- 
azine of Natural History, series 3, 9:467-475. 

Jullien, J. 1903. Bryozoaires provenant des campagnes 
de I'Hirondelle (1886-88). — Resultats des cam- 
pagnes scientifiques accompli par le Prince Al- 
bert I, 23:1-120. 

Laake, J. S., S. T Buckland, D. R. Anderson, & K. P 
Bumham. 1994. DISTANCE User's Guide V2.1 
Colorado Cooperative Fish and Wildlife Re- 
search Unit, Colorado State University, Fort 
Collins, Colorado, 84 pp. 

Lagaaij, R., & P L. Cook. 1973. Some Tertiary to Re- 
cent Bryozoa. Pp. 489-498 in A. Hallam, ed.. 
Atlas of Palaeobiogeography. Elsevier Scientific 
Publishing Company, Amsterdam. 

Norman, A. M. 1869. Last report on dredging among 
the Shetland Islands. — Report of the British As- 
sociation for the Advancement of Science for 
1868:303-312. 

Schopf, T J. M. 1969. Geographic and depth distri- 
bution of the phylum Ectoprocta from 200 to 
6000 m. — Proceedings of the American Philo- 
sophical Society 113:464-474. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):319-326. 1999. 

Two new species of Dentatisyllis and Branchiosyllis 
(Polychaeta: Syllidae: Syllinae) from Venezuela 

Guillermo San Martin and David Bone 

(GSM) Laboratorio de Biologia Marina e Inveitebrados, Departamento de Biologia, 

Unidad de Zoologia, Universidad Autonoma de Madrid, Canto Blanco, 

E-28049 Madrid, Spain, e-mail: guillermo.sanmartin@uam.es; 

(DB) Departamento de Biologia de Organismos, Instituto de Tecnologia y Ciencias Marinas, 

Universidad Simon Bolivar, Caracas, 1080 Venezuela, A. P. 89000 

Abstract. — A new species of the genus Dentatisyllis Perkins, 1981 and an- 
other new species of the genus Branchiosyllis Ehlers, 1887 are described: Den- 
tatisyllis morrocoyensis and Branchiosyllis lorenae. The specimens of both new 
species were collected during a study of the polychaetes from Thalassia tes- 
tudinum beds in Morrocoy Park, Venezuela. One specimen of B. lorenae was 
previously collected and reported as Branchiosyllis sp., from Cuba by San 
Martin (1991), so this species could be distributed throughout the Caribbean 
area. Dentatisyllis morrocoyensis is distinguished from all other species of the 
genus by the high number and shape of teeth in the trepan and shape of the 
blades of the compound setae, unique in the genus. Branchiosyllis lorenae is 
very similar to B. exilis but differs by having, on the anterior and midbody 
parapodia, 1-2 compound setae, the bidentate blades of which are slightly 
curved and bear a distal tooth somewhat shorter than the proximal one. This 
latter characteristic is unique to the genus. 



During a study on the ecology of the 
polychaetes inhabiting Thalassia testudin- 
um beds in the Morrocoy Park, Venezuela, 
specimens of two undescribed species of 
Syllidae were collected. A series on the tax- 
onomy and ecology of several families of 
polychaetes from this area and habitat has 
been recently begun (Bone & Vieitez 
1999). This paper deals with the description 
of the two new species of Syllidae: Den- 
tatisyllis morrocoyensis and Branchiosyllis 
lorenae. The study of the syllids from Mor- 
rocoy Park has been supported by the 
Agreement between the Universities Simon 
Bolivar (Venezuela) and Autonoma de Ma- 
drid (Spain). 

Materials and Methods 

All samples were collected in shallow 
Thalassia testudinum seagrass beds in less 



than 0.5 m depth. The samples were taken 
using a 38 cm- corer which was pushed 25 
cm into the sediment. Sediment samples 
were preserved in 10% buffered formalin 
and washed through a 1 mm mesh sieve. 
All organisms were hand-picked under a 
magnifying-lens from the remaining mate- 
rial and separated for taxonomic identifi- 
cations. Measurements are referred to the 
holotype or largest specimen studied; width 
is measured across the proventriculus and 
excludes cirri, parapodia, and setae. Obser- 
vations, drawings, and measurements were 
made using a microscope with interference 
contrast optics. Drawings were made with 
the aid of a drawing tube. The SEM micro- 
graphs were taken at the SIDI (Servicio In- 
terdepartamental de Investigacion) of the 
University Autonoma of Madrid. Types are 
deposited in the Museo Nacional de Cien- 
cias Naturales de Madrid, Spain. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Results and Discussion 

Family Syllidae Grube, 1850 

Subfamily Syllinae Grube, 1850 

Genus Dentatisyllis Perkins, 1981 

Dentatisyllis morrocoyensis, new species 

Fig. 1 

Material examined. — Morrocoy Park 
(Venezuela), Thalassia testudinum beds, 
Holotype. 

Description. — Body small, thin, cylindri- 
cal, without color marking, incomplete, 
about 4.2 mm long, 0.32 mm wide, 30 se- 
tigers. Prostomium semicircular; four small 
eyes in open trapezoidal arrangement and 
two anterior eyespots. Only one lateral, bro- 
ken antenna present on this specimen, with 
10 articles, originating in front of anterior 
eyes. Palps slightly longer than prostomi- 
um, fused at bases. Tentacular segment dis- 
tinct, somewhat shorter than following seg- 
ments; dorsal tentacular cirri with about 13 
articles; ventral tentacular cirri somewhat 
shorter, with about 8 articles. Dorsal cirri of 
first setiger long, with about 23 articles; re- 
maining dorsal cirri alternating long and 
short; long dorsal cirri similar in length to 
body width, with about 17-18 articles, short 
dorsal cirri shorter than body width, with 
about 10 articles (Fig. lA). Parapodia elon- 
gate, each with distal anterior lobe, some- 
what shorter than distal posterior lobe; ven- 
tral cirri long, digitiform, extending past 
tips of parapodial lobes, distally broad (Fig. 
ID). Compound setae all heterogomph fal- 
cigers, similar throughout; about 8-10 setae 
on each parapodium. Blades bidentate, with 
both teeth very close, similar in length. 
Marked dorso-ventral gradation in shape 
and length of blades; blades of dorsal-most 
compound setae long, about 38 |ULm, with 
short and coarse spines on margin; remain- 
ing blades provided with long, coarse, up- 
wards dressed, 3-4 spines on bases, espe- 
cially on medium size ones, and shorter and 
thinner spines distally; blades of ventral- 
most compound setae about 20 |xm length 
(Fig. IF). Simple dorsal and ventral setae 
not seen. Parapodia each with two slender 



aciculae, ending with fine tips (Fig. IE). 
Pharynx long, everted on this specimen 
(Fig. lA); anterior margin (Fig. lA, B) sur- 
rounded by a trepan of about 50 curved, 
hooked teeth (Fig. IB, C), and a crown of 
about 20 soft papillae; midorsal pharyngeal 
tooth rhomboidal, small, located subtermin- 
ally to anterior margin. Proventriculus 
shorter than pharynx, through 4 segments, 
with about 27 muscle cell rows. 

Remarks. — The genus Dentatisyllis was 
erected by Perkins (1981) for species hav- 
ing a cylindrical body, pharyngeal tooth 
and a trepan on the anterior margin of the 
pharynx; the genus has been recently re- 
vised by Ding et al. (1998) who provided a 
diagnosis and a key to all the known spe- 
cies of the genus. Dentatisyllis morrocoy- 
ensis n. sp., is the only species of the genus 
provided with a very high number of mar- 
ginal teeth on the trepan; all other species 
have about 10, whereas D. morrocoyensis 
has about 50. The blades of the compound 
setae and the aciculae of D. morrocoyensis 
n. sp. are very similar to those of Opistho- 
syllis longidentata San Martin, 1991, but 
the pharyngeal armature is completely dif- 
ferent (San Martin 1991). 

Etymology. — The species is named after 
the type locality, Morrocoy Park (Venezue- 
la). 

Genus Branchiosyllis Ehlers, 1887 

Branchiosyllis lorenae, new species 

Figs. 2, 3, 4 

Branchiosyllis sp. San Martin (1991):234, 
fig. lO-S. 

Material examined. — Morrocoy Park 
(Venezuela), Thalassia testudinum beds, 
Holotype and 27 paratypes. Additional ma- 
terial: 2 specimens used for SEM. 

Description. — Body long, cylindrical 
(Figs. 2A, 4A), holotype incomplete speci- 
men, 5.3 mm long, 0.4 mm wide, 54 setig- 
ers; longest complete paratype 7.2 mm 
long, 0.48 mm wide, 50 setigers, a few 
somewhat longer, incomplete paratypes. 
Most anterior segments without color mark- 



VOLUME 112, NUMBER 2 



321 




Fig. 1. Dentatisyllis morrocoyensis, n. sp. Holotype. A, anterior end. dorsal view; B. ventral view of the 
anterior end of the pharynx; C, detail of the teeth of the trepan; D. midbody parapodium, anterior view: E, 
aciculae, from midbody; F, compound setae, midbody. Scale. — A:0.1 1 mm. B, D:65 |xm. C:48 ixm. E. F:20 |j.m. 



ing, anterior and midbody segments provid- dorsal cirri with dark spots (Fig. 2A). Pro- 

ed dorsally each with three ovate dark stomium oval, wider than long; four eyes in 

spots, sometimes forming nearly a row; very open trapezoidal arrangement, nearly 

some articles of antennae, tentacular and on line, and two small anterior eyespots. 



322 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




D 



Fig. 2. Branchiosyllis lorenae, n. sp. Holotype. A. anterior end, dorsal view: B. parapodial lobe, dorsal 
view. C, aciculae. anterior parapodium: D. aciculum. posterior parapodium. Scale. — A:0.11 mm. B:48 p.m. 
C. D:20 [xm. 



Median antenna originating between pos- 
terior pair of eyes, somewhat longer than 
prostomium and palps together, with 14 ar- 
ticles; lateral antennae shorter than median 
antenna, originating between anterior pair 
of eyes and eyespots. with 12-13 articles. 
Palps broad, longer than prostomium, fused 
at bases. Tentacular segment reduced, cov- 
ered dorsally by prostomium and setiger 1: 
dorsal tentacular cirri longer than median 



antenna, with about 20 articles; ventral ten- 
tacular cirri similar in length to lateral an- 
tennae, with about 13 articles. Dorsal cirri 
of setiger 1 very long, with about 30-33 
articles; remaining dorsal cirri alternating 
long, somewhat longer than body width, 
with 25-30 articles, and short cirri, shorter 
than body width, with about 16-18 articles. 
Parapodial lobes conical, provided with two 
distal, triangular lobes (branchiae), anterior 



VOLUME 112, NUMBER 2 



323 




Fig. 3. Branchiosyllis lorenae. n. sp. Compound setae: A. dorsal from anterior parapodium; B, median and 
ventral from anterior parapodium. C. from anterior midbody: D. from posterior midbody; E. from posterior 
parapodium; H claw-shaped setae, from posterior-most parapodia. Scale. — 20 |jLm. 



lobe somewhat shorter than posterior lobe 
(Figs. 2B, 4E). Ventral cirri digitiform, 
elongate, reaching distal level of parapodial 
lobes. Anterior parapodia each with 1-2, 
sometimes 3, dorsal, compound setae pro- 
vided with long, somewhat curved, biden- 



tate blades, 50-55 ixm, distal tooth short 
and small proximal tooth somewhat larger, 
more prominent (Figs. 3A, 4C); spines on 
margin short, several rows of spines ob- 
served by SEM, slightly longer than those 
of the rest of blade, partially covering prox- 



324 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 4. Branchiosyllis lorenae, n. sp. SEM. A, anterior end, dorsal view; B, anterior rim of pharynx; C, long 
bladed, dorsal compound seta, anterior parapodium; D, detail of the same, showing several rows of spines on 
distal part. E, midbody posterior parapodium; F, dorsal compound seta from midbody; G, ventral compound 
setae from midbody; H, claw-shaped compound setae, from posterior- most parapodia. 



VOLUME 112, NUMBER 2 



325 



imal tooth (Fig. 4D), 7-10 median and ven- 
tral compound setae with blades curved, bi- 
dentate, both teeth similar on longer blades, 
proximal tooth shorter than distal one on 
shorter blades; gradation in length, 32 ixm 
above, 24 |jLm below (Fig. 3B). Progres- 
sively decreasing number of compound se- 
tae on each parapodium, blades shorter, 
lacking compound setae with long blades; 
midbody parapodium with a few bidentate 
blades and several unidentate, smooth or 
slightly bidentate, hooked blades (Figs. 3D, 
4F, G); posterior parapodia each with a few 
modified compound setae with blades 
turned 180°, smooth, strongly hooked, 
claw-shaped (Fig. 4H); posterior-most para- 
podia only with 2-3 claw-shaped com- 
pound setae (Fig. 3F). Shafts of compound 
setae distally provided with spines, which 
are increasingly more numerous and longer 
in dorsal and anterior setae. Solitary dorsal 
and ventral simple setae absent. Anterior 
parapodia each with 3-4 aciculae, dimin- 
ishing progressively to only 1-2 on poste- 
rior parapodia; aciculae with curved tip 
(Fig. 2C, D). Pharynx broad, through 5 seg- 
ments; anterior margin provided with a 
crown of ten soft papillae and one conical 
middorsal tooth (Figs. 2A, 4B). Proventric- 
ulus somewhat longer than pharynx, 
through 6 segments, with about 30 muscle 
cell rows. Pygidium small, with two long, 
articulated anal cirri. 

Remarks. — Branchiosyllis is a genus 
with only five described species: B. oculata 
Ehlers, 1887 (Ehlers 1887), distributed in 
the Caribbean Sea; B. exilis (Gravier, 1900) 
(Gravier 1900, Westheide 1974, San Martin 
1984), a circumtropical species, also pre- 
sent in temperate waters; B. pacifica Rioja, 
1941 (Rioja 1941), from the Pacific coasts 
of Mexico; B. diazi Rioja, 1958 (Rioja 
1958), from the Caribbean coast of Mexico; 
and B. abranchiata Hartmann-Schroder, 
1965 (Hartmann-Schroder 1965) from Sa- 
moa. A table summarizing the main char- 
acters of all these species is provided in 
Hartmann-Schroder (1978) and San Martin 
(1984). Branchiosyllis lorenae, n. sp. is eas- 



ily distinguished from B. oculata, B. paci- 
fica and B. abranchiata because these three 
species have only claw- shaped seta, and 
lack unmodified ones. Branchiosyllis lor- 
enae differs from B. diazi in lacking a dor- 
sal branchia on each parapodium and in the 
shape of anterior compound setae. The most 
similar species is B. exilis; however, B. lor- 
enae differs in having compound setae on 
anterior parapodia with long, curved, 
blades, with the distal tooth small, shorter 
than the proximal tooth; this kind of blade 
is very unusual and B. lorenae, n. sp., is the 
only member of this genus with this kind 
of seta. 

Etymology. — The species is named in 
honor of Lorena Galindo, who collaborated 
in the collection of the samples and sepa- 
ration of specimens. 

Literature Cited 

Bone, D., & J. M. Vieitez, 1999. Spionidae from Mor- 
rocoy Park (Venezuela). — Bulletin of Marine 
Science (in press). 

Ding, Z., E Licher, & W. Westheide. 1998. New and 
newly assigned species of the genus Dentati- 
syllis (Polychaeta, Syllidae, Syllinae), with 
comments on the reproduction, together with a 
key and a synoptic table of all species of the 
genus. — Sarsia 83:29-43. 

Ehlers, E. 1887. Report on the annelids of the dredging 
expedition of the U. S. coast survey steamer 
"Blake". — Memoires of the Museum of Com- 
parative Zoology of Harvard 15:335 pp. 

Gravier, C. 1900. Contribution a I'etude des Annelides 
Polychetes de la Mer Rouge. — Nouvelles Arch- 
ives du Museum d'Histoire naturelle de Paris 2: 
137-282. 

Hartmann-Schroder, G. 1965. Zur Kenntnis der Euli- 
toralen Polychaetenfauna von Hawaii, Palmira 
und Samoa. — Abhandlungen und Verhandlun- 
gen des Naturwissensschaftlichen Vereins in 
Hamburg 9:81-161. 

. 1978. Einige Sylliden-Art (Polychaeta) von 

Hawaii und aus dem Karibischen Meer. — Mit- 
teilungen aus dem Hamburgischen Zoologisch- 
en Museum und Institut 75:49-61. 

Perkins, T. H. 1981. Syllidae (Polychaeta), principally 
from Florida, with descriptions of a new genus 
and twenty-one new species. — Proceedings of 
the Biological Society of Washington 93:1080- 
1172. 

Rioja, E. 1941. Estudios Anelidologicos. III. Datos para 
el conocimiento de la fauna de poliquetos de las 



326 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



costas del Pacifico de Mexico. — Anales del Insti- 
tute de Biologia de Mexico 12:669-746. 
-. 1958. Estudios Anelidologicos. XXII. Datos 



para el conocimiento de la fauna de Anelidos 
Poliquetos de las costas orientales de Mexico. — 
Anales del Institute de Biologia de Mexico 29 
(1/2):219-301. 
San Martin, G. 1984. Estudio biogeografico, faunistico 
y sistematico de los Poliquetos de la familia Sfl- 



idos (Syllidae: Polychaeta) en Baleares. — Tesis 
Doctoral. Ediciones de la Universidad Complu- 
tense de Madrid, n° 187:529 pp. 

. 1991. Syllinae (Polychaeta: Syllidae) from 

Cuba and the Gulf of Mexico. — Bulletin of Ma- 
rine Science 48:227—235. 

Westheide, W. 1974. Interstitielle Fauna von Galapa- 
gos. XI. Pisionidae, Hesionidae, Pilargidae, Syl- 
lidae. — Mikrofauna Meeresbodens 44:195-338. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

I12(2):327-337. 1999. 

Parvidrilus strayeri^ a new genus and species, an enigmatic interstitial 
clitellate from underground waters in Alabama 

Christer Erseus 

Department of Invertebrate Zoology, Swedish Museum of Natural History, 
Box 50007, SE-I04 05 Stockholm, Sweden 

Abstract. — An exceedingly small oligochaete, Parvidrilus strayeri, a new 
genus and species, is described from a streambed in northern Alabama, U.S.A. 
The species is up to 1.4 mm long (with up to 33 segments) and has several 
unusual features. In ovigerous specimens, a clitellum is developed as a pair of 
lateral rows of a few, large, swollen epidermal cells in segments (IX-) X-XII 
(-XIII, -XIV). Chaetae are absent from segment II. Otherwise, in the anterior 
part of the body, there are long hair chaetae as well as small crotchet chaetae 
in both dorsal and ventral bundles. Furthermore, the chaetal bundles are situated 
posteriorly in each segment. It is unclear whether gonads are paired or un- 
paired; testes (testis?) are in segment XI, ovaries (ovary?) in XII. The other 
reproductive organs include a U-shaped muscular 'genital body' in segment 
XII and a V-shaped 'copulatory organ' in segments XII-XIII; the exact nature 
and function of these structures are unknown. The genus is proposed to be 
classified as the single member of Parvidrilidae, new family. It appears to be 
most closely related to two, largely Southern Hemisphere, aquatic clitellate 
families, Capilloventridae and Phreodrilidae. A possible relationship to the 
monotypic, South American taxon, Narapidae, is also discussed. 



In the course of a study of the under- 
ground fauna of a crystal clear, spring fed 
stream in Alabama (Stray er et al. 1995), 
specimens of an exceedingly small, undes- 
cribed, oligochaete were encountered. Dr. 
David L. Strayer (Institute of Ecosystem 
Studies, Millbrook, New York, U.S.A.) 
placed the material at the present author's 
disposal. It is described as the type species 
of a new genus, Parvidrilus, in the present 
paper. The phylogenetic affinities of the 
new species, which appears to represent a 
new higher level taxon of the Clitellata, are 
also discussed. 

Material and Methods 

The worms were collected at a single site 
in Hendrick Mill Branch, northern Ala- 
bama, in October 1990 (Strayer et al. 1995). 
They were fixed in buffered formalin. 



stained with Rose Bengal, sorted under a 
dissecting microscope, and stored in 70% 
ethanol. The material was then sent to the 
present author, who stained several individ- 
uals in alcoholic paracarmine and mounted 
them whole in Canada balsam on micro- 
scope slides. Most of the measurements in 
the description refer to this whole-mounted 
material, examined under a light micro- 
scope. Due to the small dimensions, how- 
ever, the details of the chaetae could not be 
observed even when using a high-resolution 
lOOX oil immersion lens. Therefore, exter- 
nal traits were also investigated by means 
of scanning electron microscopy. Since the 
animals were too small for the perforated 
containers available for critical point dry- 
ing, they were simply dehydrated in 99% 
ethanol, transferred to butyl acetate, and air- 
dried. This is a rough method, but thanks 
to their sturdy cuticle and small size the 



328 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



worms maintained the body shape accept- 
ably, at least in a comparison with individ- 
uals prepared for light microscopy. Three 
specimens were mounted on double-sided 
tape, coated with gold and examined with 
SEM. 

The holotype and some paratypes of the 
new species, Parvidriliis strayeri, are de- 
posited in the National Museum of Natural 
History (USNM), Smithsonian Institution, 
Washington, D.C., additional paratypes in 
the Swedish Museum of Natural History 
(SMNH), Stockholm. 

ParvidriUdae, new family 

Etymology. — ^Family name (-idae) based 
on Parvidrilus, new genus (type genus). 

Diagnosis. — Until additional species are 
found, the new family is characterized by 
the features of the type species of the type 
genus. They can be summarized as follows: 

Meiofaunal freshwater oligochaetes, 
about 1 mm long. Chaetal bundles situated 
posterioriy in each segment, but completely 
absent from segment 11. In anterior part of 
body, both dorsal and ventral bundles with 
very long hair chaetae as well as short 
crotchet chaetae; dorsal crotchets single- 
pointed, ventral ones bifid. In posterior part 
of body, dorsal bundles similar to anterior 
dorsal ones, ventral bundles with bifid 
crotchets only. In ovigerous specimens, cH- 
tellum developed as a pair of lateral rows 
of large, swollen, transparent, cells in seg- 
ments (IX-) X-Xn (-Xm, -XIV). Ahmen- 
tary canal simple, without diverticula. It is 
unclear whether gonads are paired or un- 
paired; testes (testis?) in segment XI, ova- 
ries (ovary?) in XII. Reproductive organs 
complex, including U-shaped muscular 
'genital body' in XII and V-shaped 'copu- 
latory organ' in Xn-Xm. 

Parvidrilus, new genus 

Etymology^ — A combination of parvus 
(Latin for 'smaU') and drilus (Greek for 
'worm'). 

Diagnosis. — As for family. 



Type species. — Parvidrilus strayeri, new 
species. 

Other species. — None. 

Parvidrilus strayeri, new species 
Figs 1—2 

Ohgochaeta n. sp.: Stray er et al. 1995:506. 

Holotype.— VS'^M 185769, whole- 
mounted specimen. 

Type locality. — ^A hyporheic site in a sand- 
gravel bar, in Hendrick Mill Branch, a spring- 
fed stream about 20 km NE of Pinson, 
Blount Co., Alabama, U.S.A., 33°52'12''N, 
86°33'57'W, 4 Oct 1990, coll. D. L. Strayer. 
At this site, the stream is 3—6 m wide and 
runs through a hardwood forest in a lime- 
stone terrain. The bottom varies from sand 
to exposed bedrock, with a predominance 
of coarse sand, angular gravel and small 
(10-15 cm) stones. Generally the alluvium 
is thin, but a large deposit of sand and grav- 
el is held in place by a dam about 3 m high. 
At the time of collection, the emergent part 
of this deposit was 3-4 m wide, about 30 
m long, and reached about 0.5 m above the 
stream water level. Because the dam leaked 
from the base, water flowed down into and 
through the bar. Subsamples were taken 
from 16 wells in the bar, using a Bou- 
Rouch pump (Bou 1974), all wells yielded 
water freely. Dissolved oxygen was 0-8.0 
ppm, with most readings 2-6 ppm. Other 
chemical characteristics (means of several 
subsamples): Ca 22.2 ppm; Mg 11.6 ppm; 
K 0.5 ppm; Na 0.7 ppm; NO3 0.9 ppm; SO4 
1.8 ppm; CI 1.2 ppm; Dissolved Organic 
Carbon (DOC) 0.5 ppm (Strayer et al. 1995; 
Strayer, pers. comm.). 

Paratypes.— US^M 185770-185776- 
000000, seven whole-mounted specimens; 
SMNH Type coU. 5085-5092, eight whole- 
mounted specimens; SMNH Type coll. 
5093, three specimens mounted on a SEM 
stub; all from type locaHty. 

Etymology. — Named for Dr. David L. 
Strayer, who collected the material and was 
the first to reaUze that the species was a 



VOLUME 112, NUMBER 2 



329 




Fig. 1. Parvidrilus strayeri, new genus and species; SEM micrographs. A, postclitellar part of body, showing 
dorsal hair chaetae (he) ventral bifid chaetae (in bottom part of figure), and intersegmental fiirrows (if); note the 
posterior location of bundles in the segments; B, anterior end of body showing mouth (m), achaetous segment 11 (II), 
and dorsal (top of figure) and ventral chaetae (middle and bottom of figure) of segments III-V; C, bundle of ventral 
chaetae of segment EX, showing hair chaeta (he) and bifid crotchets (be); D, bundle of dorsal chaetae of segment X, 
showing hair chaeta (he), and single-pointed crotchets (sc); note also thin outer part of hair chaetae of another segment; 
E, bundle of ventral chaetae of postclitellar segment, showing two bifid crotchets; F, ventral view of genital bursa 
(bu) in segment XII, showing lateral bulbous swellings (bs), posterior triangular lappet (tl). and ventral chaetae (vc). 



330 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Parxidrilus strayeri, new genus and species. A, schematical, horizontal view of segments XII and 
Xm, showing general outline of 'genital body' and 'copulatory organ'; B, somewhat horizontal view of segments 
Xl-Xni of one paratype: C, somewhat lateral view of segments XI-XIII of another paratvpe. Abbreviations: bs, 
bulbous swelling (of body wall); bu. location of genital bursa (cf. Fig. IF); c. chaeta; cc. clitellar cell; co. 
copulatory organ; gb, genital body; ov, ovary; pe, penis/pseudopenis; ps, penial/pseudopenial sac; sf, sperm 
funnel, t. testis. 



VOLUME 1 12, NUMBER 2 



331 



new taxon of considerable systematic inter- 
est. 

Description. — Length (five complete 
specimens) 0.7-1.4 mm, 18-33 segments. 
Width at genital region 0.03-0.10 mm. Pro- 
stomium rounded, somewhat narrower than 
and well set off from peristomium; prosto- 
mial epidermis with numerous (stained) cell 
nuclei, probably indicating a glandular or 
sensory function. Posterior part of body 
(Fig. lA) more clearly annulated (i.e., with 
distinct intersegmental furrows; Fig. lA, if) 
than anterior part. Pygidium generally de- 
marcated, somewhat narrower than last 
(true) segment. Cuticle smooth, but foreign 
particles often adhering to it here and there 
along body of worm. Clitellum poorly de- 
veloped in most individuals; but in oviger- 
ous specimens, a row of large, swollen, 
transparent, epidermal cells (Fig. 2B, cc) 
present on each side of worm in segments 
(IX-) X-XII (-Xni, -XIV). 

Chaetal bundles situated posteriorly in 
each segment (Fig. lA), but lacking com- 
pletely in segment II (Fig. IB). Anterior 
dorsal bundles each with one to two (oc- 
casionally three) long, whip-Uke, hair chae- 
tae (Fig. ID, he), alternating with two or 
three (occasionally four) single-pointed, 
smaller crotchet chaetae (Fig. ID, sc); hairs 
occasionally up to about 200 |xm long (ex- 
act length difficult to measure due to strong 
curvature of hairs and poorly resolved view 
of tips), entaUy about 1 jjim wide, progres- 
sively tapering and only about 0. 1 ixm wide 
at ectal end; crotchets 15—25 ixm long, 
about 0.5-0.6 (xm thick, with somewhat 
blunt tips. Postclitellar dorsal bundles each 
with one or no hair chaeta (Fig. lA, he) and 
two single-pointed crotchets. In anterior 
segments (at least in III-VII, but generally 
also including a few additional pre-genital 
segments, and sometimes even as far back 
as in XV), ventral bundles each with one 
hair chaeta (Fig. IC, he), and three (occa- 
sionally four) bifid crotchets (Fig. IC, be); 
ventral hairs slightly shorter and thinner 
than corresponding dorsal hairs; bifids 
about 20-25 |jLm long, about 0.9-1.4 |ULm 



thick, with upper tooth thinner and shorter 
than lower tooth. Within these bundles, up- 
permost (most lateral) chaeta bifid, fol- 
lowed by a hair, and these two chaetae 
clearly separated from two, more ventral, 
bifids (Fig. IC). Postclitellar ventral bun- 
dles with two to three bifid crotchets (Fig. 
IE), similar to anterior ventral bifids. Mod- 
ified genital chaetae absent. 

Ventrally in segment XII, body wall 
forming a conspicuous, generally somewhat 
"X" -shaped genital bursa (Fig. IF); im- 
pression of an X facilitated by a pair of 
large, bulbous, swellings (Fig. IF, bs), one 
at each side of bursal opening, and an in- 
distinct, mid-ventral, triangular lappet (Fig. 
IF, tl), located immediately posterior to bur- 
sal opening. 

Spermathecal openings not observed. 

Brain located within, and with front end 
reaching anterior coelomic lining of, peris- 
tomium. Nerve cord ventral, of normal mi- 
crodrile oligochaete appearance. Alimenta- 
ry system of normal microdrile oligochaete 
type too: simple pharynx with dorsal pha- 
ryngeal bulb, somewhat sinuous esophagus 
with a few pharyngeal (?) gland cells scat- 
tered in segments V-VIII, followed by a 
narrow, simple intestine; no particular en- 
largements or diverticula on gut. Large, dif- 
fuse cells of chloragogen tissue present 
along most of gut. Free coelomocytes not 
observed. 

In three specimens, a small lump of cells 
in anterior end of coelom of segment XI 
probably an (unpaired?) testis (Fig. 2C, t?); 
in another worm, a single sperm bundle 
present in coelom of this segment; other- 
wise male gonads or developing spermato- 
zoa not observed. In nine specimens, an 
ovary (apparently developed on one side 
only) observed in anterior part of segment 
XII (Fig. 2B, C, ov); three of these worms 
ovigerous, i.e., with a large mature Qgg fill- 
ing whole width of coelom, and extending 
through one or two segments, in region of 
XIII-XV In two specimens (only), a sperm 
funnel discernible at anterior side of septum 



332 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



between XI and XII (Fig. 2B, C, sf), but its 
continuation into a vas deferens not seen. 

Other internal genitalia (in XII-XIII) 
complex (Fig. 2), but due to quality of fix- 
ation and small dimensions difficult to in- 
terpret using conventional oligochaete ter- 
minology. Two conspicuous structures pres- 
ent, here referred to as the 'genital body' 
and the 'copulatory organ', respectively. 

Genital body (Fig. 2A-C, gb) a "U"- 
shaped structure incorporated in ventral 
body wall in segment XII; the two arms 
forcing body to bulge considerably, corre- 
sponding to two bulbous swellings at lateral 
sides of genital bursa (Fig. IF, bs). Arms of 
genital body more or less oval, 20-35 ixm 
long, 15-23 |xm wide, oriented along long 
axis of worm, and united anteriorly. At ear- 
ly developmental stages, lobes appear to 
contain a lumen. At full maturity, wall of 
genital body thick, heavily muscular and 
folded in complex manner (Fig. 2B, C, gb), 
with indistinct lumen (lumina?); sometimes 
densely packed cilia (see Fig. 2B) and/or 
spermatozoa (?) visible inside. At late stag- 
es of development, lobes tend to break up 
into a number of secondary, pear-to- spin- 
dle- shaped, packages of muscle fibers (as in 
Fig. 2B). Function of genital body un- 
known; possibly having something to do 
with storage and/or ejaculation of sperm, ei- 
ther before (an atrium?) or after copulation 
(a spermatheca?). Whether genital body has 
any permanent or temporary opening to the 
exterior or any connection with the sperm 
funnels or copulatory organ unknown. 

Copulatory organ (Fig. 2A, co) bilobed, 
somewhat "V" -shaped, consisting of a pair 
of straight or somewhat curved sacs (Fig. 
2B, C, ps), conrununicating with each other 
mid-ventrally in most posterior part of seg- 
ment XIL At point of union of the two sacs, 
copulatory organ appears anchored in the 
ventral body wall, in a position correspond- 
ing to apex of the mid-ventral, triangular, 
lappet located in posterior wall of genital 
bursa (Fig. IF, tl; see above). Inner ends of 
sacs extending obliquely and freely back- 
wards into coelom of XIII. Each sac 25-60 



jjim long, 10-21 ixm wide, with thick, mus- 
cular, wall, and containing a coiled, slender 
tube. In one damaged specimen, tubes 
squashed out from sacs, appearing as long 
penes (Fig. 2C, pe), but it appears as if tips 
of these are normally located inside inner 
ends of sacs rather than at mid-ventral at- 
tachment point in body wall. It is thus un- 
known whether tubes are protrusible (i.e., 
penes) or eversible structures (i.e., pseudo- 
penes). 

Remarks. — Although there is a possibil- 
ity that the genital body in segment XII is 
used for storing sperm from a concopulant 
(see above), typical spermathecae were not 
observed. 

Several points in this description need to 
be clarified by future studies. Nevertheless, 
Parvidrilus strayeri is easily separated from 
all other known oligochaete taxa. See Dis- 
cussion below. 

Distribution and habitat. — Known only 
from the type locality in Alabama, USA. 
Interstitial groundwater. 

Discussion 

Parvidrilus strayeri is clearly a clitellate. 
It appears to be hermaphroditic (the geni- 
talia of all specimens look the same) with 
gonads in specific segments, and it has an 
eversible, thickened, pharynx roof, a brain 
located behind the prostomium, and large 
glandular epidermal cells in the genital re- 
gion suggesting the existence of a clitellum 
(see Purschke et al. 1993). In a traditional 
sense, the worm is a member of the Oli- 
gochaeta; but there is now increasing evi- 
dence that this taxon is paraphyletic unless 
leeches and leech-like groups are included 
(Purschke et al. 1993, Brinkhurst 1994, Sid- 
dall & Burreson 1996, Ferraguti & Erseus 
1999). A formal phylogenetic analysis to 
establish the more specific systematic po- 
sition of Parvidrilus within the Clitellata, is 
not meaningful until some uncertainties in 
the description have been clarified. As there 
is yet no strong indication as to which cli- 
tellate subgroup the new species belongs. 



VOLUME 112, NUMBER 2 



333 



outgroups for such an analysis would need 
to be selected from outside the Clitellata, 
implying that most ingroup and outgroup 
character states would not be comparable. 

In some ways, however, the new species 
is unusual and unique (see Table I). First, it 
is one of the smallest clitellates known. 
With a body length of about 1 mm it resem- 
bles only the smallest Chaetogaster spp. 
(Naididae) (see Sperber 1948), an undes- 
cribed freshwater species tentatively as- 
signed to Capilloventer (Capilloventridae) 
from Western Australia (Pinder & Brink- 
hurst 1997b); and a miniature terrestrial 
species of Enchytraeidae from Italy, Mar- 
ionina eleonorae Rota, 1995. Some marine 
oligochaetes are just slightly longer. Ex- 
amples within the 1.5-2.5 mm range are nu- 
merous species of the subfamily Phallodri- 
linae (Tubificidae; see, e.g., Erseus 1980, 
1989, 1990, 1992), and Randiella caribaea 
Erseus & Strehlow, 1986 and R. minuta Er- 
seus & Strehlow, 1986 (Randiellidae; see 
Erseus & Strehlow 1986, Erseus 1997). To 
my knowledge, there is no freshwater tu- 
bificid that is even close to the 0.7-1.4 mm 
range that characterizes Parvidrilus. 

Second, in Parvidrilus strayeri, the four 
chaetal bundles of each segment are located 
in a more posterior position within the seg- 
ment than in other oligochaetes. Third, the 
clitellar cells are large in relation to the 
body diameter, few in number, and restrict- 
ed to two lateral rows, one at each side, 
through a few segments of the worm. These 
features are likely to be autapomorphies of 
Parvidrilus or of a group including as yet 
unknown taxa. 

Although complete male ducts have not 
been observed in Parvidrilus, the position 
of the genital bursa indicates that male 
pores (or an unpaired male pore?) are pres- 
ent in segment XII. In this respect the new 
taxon bears resemblance to four other mi- 
crodrile families, Enchytraeidae, Propappi- 
dae, Capilloventridae and Phreodrilidae, 
and a few representatives of the Lumbri- 
culidae. Capilloventridae and Phreodrilidae 
are regarded as endemic to the Southern 



Hemisphere (although a few phreodrilids 
occur north of the equator), but a further 
comparison with them is pertinent here as 
both (see Harman & Loden 1984; Erseus 
1993; Pinder & Brinkhurst 1997a, 1997b) 
are characterized by absence of chaetae in 
segment II (for phreodrilids, at least with 
regard to the dorsal chaetae; Pinder & 
Brinkhurst 1997a), and presence of hair 
chaetae in the other segments, i.e., two 
striking similarities with the chaetal pattern 
in Parvidrilus (Table I). 

The additional feature of ventral hair 
chaetae is, within the CHtellata, shared only 
by Parvidrilus and Capilloventridae and 
may be homologous. However, Capilloven- 
tridae, as currently defined (Pinder & 
Brinkhurst 1997b), is characterized by 
modified hair-like genital chaetae, sperma- 
thecae in segment Vn, and a pair of blind 
ventral sacs (salivary glands?) opening into 
the mouth cavity (the two first features also 
known from the Randiellidae; see Erseus 
and Strehlow 1986). As none of these is 
present in Parvidrilus, inclusion of the new 
genus in Capilloventridae would imply re- 
ductions of all three traits, which is a less 
parsimonious hypothesis than to regard 
Parvidrilus as a taxon outside Capilloven- 
tridae. Moreover, in Capilloventridae but 
not in Parvidrilus, the chaetal bundles of 
each side are located close together and 
widely separated from those of the other 
side, somewhat like the distribution of 
chaetae on parapodia of polychaetes (Er- 
seus 1993). This could be a plesiomorphic 
condition reflecting ancestry of the Clitel- 
lata among the polychaetes (cf. Westheide 
1997), and in which case the Capilloventri- 
dae could be the most ancestral group of all 
Clitellata; alternatively, it is an additional 
autapomorphy of Capilloventridae. 

The Phreodrilidae comprises a total of 
about 60 species with great morphological 
variation with regard to chaetae as well as 
genital organs (Pinder & Brinkhurst 1997a). 
In phreodrilids, the dorsal chaetal bundles 
frequently consist of long hair-like chaetae 
accompanied by short, single-pointed, "lat- 



334 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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VOLUME 112, NUMBER 2 



335 



eral support chaetae", an arrangement sim- 
ilar to that found in Parvidrilus strayeri 
(but also similar to the dorsal 'hairs and 
needles' in species of Naididae and Opis- 
tocystidae). The crucial point would then be 
to what extent the curious genital body and 
copulatory organ of the new species are ho- 
mologous to phreodrilid genital structures. 
They show some superficial resemblance to 
the atria and eversible pseudopenes, respec- 
tively, of the species of Phreodrilus Bed- 
dard (see Pinder & Brinkhurst 1997a), al- 
though in Parvidrilus the two kinds of 
structures do not appear to be continuous 
with each other as the atria and pseudope- 
nes are in Phreodrilus. Moreover, the 
phreodrilid atria are filiform invaginations, 
lying freely in the coleomic cavity, whereas 
the Parvidrilus genital body is in very close 
contact with (merely as a pair of swellings 
of) the body wall. There are also other dif- 
ferences between Phreodrilidae and Parvi- 
drilus (Table I). In phreodrilids, the general 
body size is one order of magnitude larger 
than that of Parvidrilus, the ventral chaetae 
are consistently paired, there is a thin cli- 
tellum of normal microdrile type and it is 
generally restricted to segments XII-XIII, 
there are spermathecae in segment XIII, and 
if hairs are present, the lateral support chae- 
tae do not usually project from the chaetal 
sacs. Many of these features are shared with 
other aquatic clitellate taxa, but the lateral 
support chaetae, the position of the sper- 
mathecae (in segment XIII), and the well 
developed tubular atria mentioned above, 
are likely to be autapomorphies supporting 
monophyly of the Phreodrilidae. Thus, it 
does not appear appropriate to place Par- 
vidrilus in the Phreodrilidae. 

Narapa bonettoi Righi & Varela, 1983, 
the single representative of the family Nar- 
apidae and known only from the Parana 
River in Argentina (Righi & Varela 1983, 
Brinkhurst & Marchese 1989), is another 
aquatic oligochaete taxon with an unclear 
systematic position. It shows some curious 
resemblance to Parvidrilus. Although lack- 
ing chaetae completely and having its gen- 



ital system in a more anterior position (Ta- 
ble I), Narapa has a male gonoduct (in seg- 
ment VI) (Righi & Varela 1983:figs 4-6) 
with an outline comparable to one possible 
interpretation of the reproductive system in 
Parvidrilus. In Narapa, the sperm funnel is 
followed by a short vas deferens (not yet 
observed in Parvidrilus), leading to a ven- 
tral glandular, tubular atrium (possibly cor- 
responding to a lateral arm of the 'genital 
body' in Parvidrilus), followed by a pos- 
teriorly bent-over penial sac with a winding 
lumen (possibly corresponding to a copu- 
latory sac in Parvidrilus). It is also note- 
worthy that the ovary (in VII) is reported 
to be unpaired in Narapa', in none of the 
specimens of Parvidrilus strayeri was more 
than one ovary observed. It is an open 
question whether these similarities are syn- 
apomorphic or convergent, but Narapidae 
and Parvidrilidae are both monotypic and 
each possesses its own autapomorphies (Ta- 
ble I). As currently defined, they can, there- 
fore, at most be regarded as sister taxa. 

However, relevant comparisons with oth- 
er aquatic microdrile taxa have still not 
been exhausted. For instance, it could be 
suggested that the U-shaped genital body of 
Parvidrilus strayeri is a derivative of a pair 
of atria of the kind found in the family Tub- 
ificidae, and that its position in segment XII 
is merely an autapomorphic rearward shift 
of a tubificid male system. Following this 
line of reasoning, the copulatory organ of 
Parvidrilus (Fig. 2C) could be interpreted 
as an indication of a close relationship with, 
e.g., Teneridrilus flexus Erseus & Hiltunen, 
1990 (in Erseus et al. 1990). The latter is a 
North American Great Lakes freshwater tu- 
bificid (reported also by Stacey & Hubley 
1994), with small, convoluted, tubular pe- 
nial organs contained within muscular sacs 
(Erseus et al. 1990:fig. 2D, E). However, 
these similarities are probably coincident 
and convergent. As noted above, the chaetal 
pattern and the segmental position of the 
genitalia strongly suggest that Parvidrilus is 
more closely related to the Capilloventridae 
and Phreodrilidae than to the Tubificidae 



336 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



(Table I). Furthermore, the tubificid atria 
extend freely into the coelom, rather than 
being incorporated in the body wall as is 
the case with the genital body of Parvidri- 
lus. 

To conclude, it seems justified to estab- 
lish a higher level taxon, Parvidrilidae, new 
family, for Parvidrilus on the basis of the 
available morphological evidence. This 
classification is not likely to render any of 
the other clitellate families paraphyletic. 
Several features of Parvidrilus/^2cr\idu\i- 
dae are probably autapomorphic: the ex- 
treme miniaturization, the chaetal bundles 
situated posteriorly in each segment, the cli- 
tellum modified into two lateral rows of a 
few large glandular cells, and the unique, 
complex, genital organs surrounding a con- 
spicuous mid- ventral bursa in segment XII. 

In its area of distribution, Parvidrilus is 
part of a particular association of inverte- 
brates ('stygobionts') adapted to life in in- 
terstitial groundwater, an association which 
also includes the polychaete Troglochaetus 
sp., aeolosomatids (Annelida, Aphanoneu- 
ra), smaller lumbriculid (e.g., Stylodrilus 
wahkeenensis Rodriguez & Coates, 1996) 
and naidid oligochaetes, bathynellacean 
crustaceans, microcerberid isopods and nu- 
merous benthic cyclopoid copepods (Rod- 
riguez & Coates 1996, Strayer et al. 1995, 
Strayer & Reid in preparation, Reid et al. 
in preparation). Although still greatly over- 
looked, this rich stygobiont fauna appears 
to characterize a vast area of unglaciated 
ancient terrain in eastern North America. 
Some authors (e.g., Wagele et al. 1995, 
Reid 1998) have already pointed out that 
many endemic interstitial microcrustaceans 
of this region appear to represent an old 
continental fauna, with closely related, but 
anatomically distinct Eurasian counterparts. 
It is possible that Parvidrilus is a northern 
representative of a group of aquatic cHtel- 
lates, which also contains the Southern 
Hemisphere families, Capilloventridae, 
Phreodrilidae and, possibly also, Narapidae. 

Parvidrilus was common at Hendrick 
MiU (D. L. Strayer, pers. comm.) and is 



probably not a rare animal, once the right 
habitats are searched. 

Parvidrilus strayeri deserves further at- 
tention. New material needs to be studied 
by methods that would enable a detailed 
scrutiny of the true nature of its miniature 
genital organs, which may throw additional 
light on its systematic position. Molecular 
systematic analyses of this enigmatic taxon 
may contribute towards the same end. 

Acknowledgments 

I am indebted to Dr. David L. Strayer, for 
generously placing the material at my dis- 
posal, and for sharing published and un- 
published information; to Dr. Anders Waren 
(SMNH) for invaluable assistance with 
scanning electron microscopy; to Ms. Bar- 
bro Lofnertz (University of Goteborg), Mrs. 
Anna Hedstrom and Ms. Christine Hammar 
(both SMNH), for technical assistance; to 
Dr. Ralph O. Brinkhurst (Hermitage, TN, 
USA) and Dr. Kathryn A. Coates (Bermuda 
Biological Station for Research, Inc), for 
constructive criticism of the text; and the 
Swedish Natural Science Research Council, 
for financial support. 

Literature Cited 

Bou, C. 1974. Les methodes de recolte dans les eaux 
souterraines interstitielles. — Annals de Speleo- 
logie 29:611-619. 

Brinkhurst, R. O. 1994. Evolutionary relationships 
within the Clitellata: an update. — Megadrilogi- 
ca 5:109-116. 

, & M. Marchese. 1989. Guide to the fresh- 
water aquatic Oligochaeta of South and Central 
America. — Coleccion CLIMAX No. 6. Asocia- 
cion Ciencias Naturales del Litoral, Santo 
Tome, Argentina, 179 pp. 

Erseus, C. 1980. Taxonomic studies on the marine gen- 
era Aktedrilus Knollner and Bacescuella Hrabe 
(Oligochaeta, Tubificidae), with description of 
seven new species. — Zoologica Scripta 9:97— 
111. 

. 1989. Marine Tubificidae (Oligochaeta) of the 

Arabian Gulf coast of Saudi Arabia, part 5. — 
Fauna of Saudi Arabia 10:11-19. 

. 1990. Marine Oligochaeta of Hong Kong. Pp. 

259-335 in B. Morton, ed.. Proceedings of the 
Second International Marine Biological Work- 



VOLUME 112, NUMBER 2 



337 



shop: The marine flora and fauna of Hong Kong 
and Southern China, Hong Kong, 1986, vol. 1. 
Hong Kong University Press, Hong Kong, 448 
pp. 

— . 1992. A generic revision of the PhallodriUnae 
(Oligochaeta, Tubificidae). — Zoologica Scripta 
21:5-48. 

— . 1993. Taxonomy of Capilloventer (Capillov- 
entridae), a Httle-known group of aquatic Oli- 
gochaeta, with descriptions of two new spe- 
cies. — Journal of Natural History 27:1029- 
1040. 

-. 1997. A record of Randiella from New Cal- 
edonia, the first known occurrence of the marine 
interstitial family Randiellidae (Annelida; Oli- 
gochaeta) in the South Pacific Ocean. — Journal 
of Natural History 31:1745-1750. 

- J. K. Hiltunen, R. O. Brinkhurst, & D. W. 
Schloesser. 1990. Redefinition of Teneridrilus 
Holmquist (Oligochaeta: Tubificidae), with de- 
scription of two new species from North Amer- 
ica. — Proceedings of the Biological Society of 
Washington 103:839-846. 

-, & D. R. Strehlow. 1986. Four new species of 



marine Oligochaeta representing a new fami- 
ly. — Zoologica Scripta 15:53-60. 

Ferraguti, M., & C. Erseus. 1999. Sperm types and 
their use for a phylogenetic analysis of aquatic 
clitellates. — Hydrobiologia (in press). 

Harman, W. J., & M. S. Loden. 1984. Capilloventer 
atlanticus gen. et sp. n., a member of a new 
family of marine Oligochaeta from Brazil. — 
Hydrobiologia 115:51-54. 

Pinder, A. M., & R. O. Brinkhurst. 1997a. Review of 
the Phreodrilidae (Annelida: Oligochaeta: Tub- 
ificida) of Australia. — Invertebrate Taxonomy 
11:443-523. 

. 1997b. The family Capilloventridae (Anneli- 
da, Clitellata) in Australia, with descriptions of 
two new species of Capilloventer. — Zoologica 
Scripta 26:255-265. 

Purschke, G., W. Westheide, D. Rodhe, & R. O. Brink- 
hurst. 1993. Morphological reinvestigation and 



phylogenetic relationship of Acanthobdella pe- 
ledina (Annelida, Clitellata). — Zoomorphology 
113:91-101. 

Reid, J. W. 1998. How "cosmopolitan" are the con- 
tinental cyclopoid copepods? Comparison of the 
North American and Eurasian faunas, with de- 
scription of Acanthocyclops parasensitivus n. 
sp. from the U.S.A. — Zoologischer Anzeiger 
236:109-118. 

Righi, G., & M. E. Varela. 1983. Narapa bonettoi, gen. 
nov. sp. nov. (Oligochaeta, Narapidae, fam. 
nov.) de agua doce da Argentina. — Re vista de 
la Asociacion de Ciencias Naturales del Litoral 
14:7-15. 

Rodriguez, P., & K. A. Coates. 1996. A new American 
Stylodrilus species (Lumbriculidae, Oligochae- 
ta). — Canadian Journal of Zoology 74:92-96. 

Rota, E. 1995. Italian Enchytraeidae (Oligochaeta). 
I. — Bolletino di Zoologia 62:183-231. 

Siddall, M. E., & E. M. Burreson. 1996. Leeches (Oli- 
gochaeta?: Euhirudinea), their phylogeny and 
the evolution of life-history strategies. — Hydro- 
biologia 334:277-285. 

Sperber, C. 1948. A taxonomical study of the Naidi- 
dae. — ^Zoologiska Bidrag frn Uppsala 28:1-282. 

Stacey, D. F, & B. R. Hubley. 1994. New records of 
a sludge worm Teneridrilus flexus Erseus & 
Hiltunen (Oligochaeta: Tubificidae) from Lake 
Huron. — Canadian Field. Naturalist 108:182- 
185. 

Stray er, D. L., S. E. May, P Nielsen, W. Wollheim, & 
S. Hausam. 1995. An endemic groundwater fau- 
na in unglaciated eastern North America. — Ca- 
nadian Journal of Zoology 73:502-508. 

Wagele, J. W., N. J. Voelz, & J. V. McArthur 1995. 
Older than the Atlantic Ocean: discovery of a 
fresh-water Microcerberus (Isopoda) in North 
America and erection of Coxicerberus, new ge- 
nus. — Journal of Crustacean Biology 15:733- 
745. 

Westheide, W 1997. The direction of evolution within 
the Polychaeta. — Journal of Natural History 3 1 : 
1-15. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):338-35L 1999. 

On the entocytherid ostracods of the Brazos River basin and 
adjacent coastal region of Texas 

Daniel J. Peters and Jean E. Pugh 

(DJP) New Horizons Governor's School for Science & Technology, 520 Butler Farm Road, 

Hampton, Virginia 23666, U.S.A.; 
(JEF) Christopher Newport University, 6640 Fields Landing Road, Hayes, Virginia 23072, U.S.A. 

Abstract. — New records of entocytherid ostracods infesting crayfishes are 
recorded from the Brazos River basin in southeastern Texas, which extend the 
ranges of Ankylocythere ancyla, A. sinuosa, Entocythere harrisi, E. reddelli, 
and Uncinocythere simondsi. A review of the variations in the copulatory com- 
plex of A. ancyla is provided and the synonymy of A. tiphophila with A. 
sinuosa is proposed. 



Upon the completion of his study of the 
crayfishes of the Brazos River basin and 
nearby areas, Douglas W. Albaugh (1973, 
1975; Albaugh & Black 1973) sent sedi- 
ments from the containers in which his 
specimens were preserved to the Smithson- 
ian Institution. This report is based on the 
ostracods contained in these samples which 
were made available to us through the kind- 
ness of the late Horton H. Hobbs, Jr. 

Five species of entocytherid ostracods 
were retrieved from 118 samples collected 
between 21 February and 8 March 1973. 
Hosts, which were identified by Dr. Al- 
baugh (the subgeneric assignments were 
provided by Horton H. Hobbs, Jr.), include 
the following; Cambarellus {Dirigicamba- 
rus) shufeldtii (Faxon), C. (Pandicambarus) 
ninae Hobbs, C. {P.) puer Hobbs, Fallicam- 
barus (Creasehnus) fodiens (Cottle), Pro- 
cambarus (Capillicambarus) brazoriensis 
Albaugh, P. (C.) hinei (Ortmann), P. (C) 
incilis Penn, P. (Girardiella) simulans 
(Faxon), P. (Ortmannicus) a. acutus (Gi- 
rard), P. (O.) texanus Hobbs and P. (Sca- 
pulicambarus) clarkii (Girard). Insofar as 
we have been able to determine, all of the 
collections were made from open water; no 
crayfishes were taken from burrows. The 
ostracods, locality data, host and entocy- 
therid identifications are deposited at the 



National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 

Ankylocythere ancyla Crawford 
Figs, la-h, 2a-h, 4 

Ankylocythere ancyla Crawford, 1965; 148, 
149, 152, 153, figs. 1-3, 6, 7 [Type lo- 
cality; "... in the city limits of Greens- 
boro, Guilford County, North Carolina." 
Types; holotype, allotype, morphotype, 
and dissected male paratype USNM; par- 
atypes in the collection of E. A. Craw- 
ford, Jr., and USNM. Host; Cambarus la- 
timanus (LeConte) { = Cambarus (De- 
pressicambarus) catagius Hobbs & Per- 
kins, 1967)]. 

Ankylocythere species g Hobbs III, 1969; 
32-34, figs. 4d-g, 1. 

Ankylocythere species h Hobbs III, 1969; 
34-36, figs. 4h-k, 1. 

Except for the omission of the references 
to Hobbs III (1969), Andolshek & Hobbs 
(1986; 10) included a compilation of all ref- 
erences in the literature to this ostracod. 

Diagnosis of Texas material. — Shell 
length of male 336 to 378 (avg. 357) ixm; 
shell height 196 to 224 (avg. 218) ixm. Pen- 
iferum truncate to tapering with acute an- 
tero- and posteroventral angles. Clasping 
apparatus L-shaped with vertical ramus Ion- 



VOLUME 112, NUMBER 2 



339 




Fig. I. Variation in penifera, lateral view, of Ankylocy the re ancyla in Brazos River basin (a, h — Refugio 
Co.; b, f — Washington Co.; c, g — Fort Bend Co.; d — Austin Co.; e — Brazoria Co.); scale 0.02 mm. 



340 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Variation in clasping apparatus, lateral view, of Ankylocythere ancyla in Brazos River basin (a, 
h — Refugio Co.; b, e, g — Washington Co.; d — Fort Bend Co.; f — Austin Co.; c — Brazoria Co.); same scale 
as Fig. 1. 



ger than horizontal ramus; latter with tooth 
on preaxial border near midlength, and 
postaxial border with long, curved talon 
reaching to midway between preaxial tooth 
and apex of ramus; extreme apical part of 
talon sometimes strongly curved mesially. 
Apex of clasping apparatus with 2 denti- 
cles. 

Range.— Ando\s\iQ\L & Hobbs (1986) 
quoted the range of the species as cited by 
Hobbs & Peters (1977) as extending ". . . 



along the Atlantic and Gulf slopes from the 
Mobile River drainage in Alabama and 
Mississippi northeastward to the Potomac 
drainage in Virginia and in the New River 
Basin of North Carolina." The locality re- 
cords established herein extend the west- 
ward limits some 750 kilometers to the 
Brazos basin in Texas. 

Southeastern Texas records (Fig. 4). — 
Nineteen localities from the following 
counties: Austin (2), Brazoria (2), Brazos 



VOLUME 112, NUMBER 2 



341 



(2), Calhoun (3), Fort Bend (2), Matagorda 
(1), Refugio (6), and Washington (1). 

Remarks. — In his study of the entocy- 
therids infesting burrowing crayfishes in the 
coastal plain between extreme eastern Texas 
and the Apalachicola Basin in Alabama and 
Florida, Hobbs III (1969) recognized 12 
species belonging to the genus Ankylocy- 
there. Specimens of two of these, A. species 
"g" and "h" were lent to us by Dr. Hobbs, 
and, after comparing them with represen- 
tatives of A. ancyla from throughout its 
range, we are convinced that they are re- 
ferable to this species. Thus there are few 
gaps in its known range. Andolshek & 
Hobbs (1986:14) reviewed available data 
on the size of the shell of this ostracod and 
found that the smallest individuals occurred 
in southeastern Georgia and the largest in 
Virginia. Those in North Carolina were, on 
the average, intermediate in size. The range 
in size of the material from along the coasts 
of Florida, Alabama, Mississippi, Louisi- 
ana, and Texas reported herein falls within 
that cited for the species in southeastern 
Georgia (length, 315 to 399 luim; height 175 
to 245 |jLm). Thus it appears that the shells 
of southern populations of A. ancyla are 
smaller than those occurring in the more 
northern parts of the range. 

A cursory examination of the ventral part 
of the peniferum would suggest that a di- 
morphic condition exists in this appendage, 
one in which the anteroventral extremity is 
produced in a subspiculiform prominence 
(Fig. Id), and in the other, subtruncate (Fig. 
le). That the difference is more in the angle 
from which the penifera are viewed rather 
than due to morphological variation be- 
comes apparent when those of a number of 
specimens are compared. In this ostracod, 
the basic structure of the ventral extremity 
of the peniferum is more clearly observed 
in specimens from the eastern part of the 
range where a broad concavity exists be- 
tween the acute cephloventral and rounded 
posteroventral extremities (See fig. 4a in 
Andolshek & Hobbs (1986)). In specimens 
from the Brazos region, the anteroventral 



angulate extension often appears to be more 
strongly produced, and its base to bear a 
thickened, sclerotized prominence, which 
when viewed at some angles, seems to pro- 
ject posteriorly or posteroventrally; also the 
anteroventral apex of the posteroventral 
prominence is procurved, diminishing the 
maximum diameter of the concavity, and is 
frequently rather strongly sclerotized. (For 
variations compare Fig. lb, c, e, g, h). Thus, 
whereas the ventral extremity of the peni- 
fera of the eastern and western members of 
the species appear to be markedly different, 
the contrast is less marked than seems ap- 
parent when only a superficial comparison 
is made. 

Among other variations noted in the cop- 
ulatory complex of this ostracod are the 
thickness of the junction of the horizontal 
and vertical rami of the clasping apparatus 
and the curvature of the vertical ramus. In 
specimens from Washington County, TX, 
males were found that possess a sinuous 
vertical ramus (Fig. 2e), and in the area of 
the junction of the rami there occurs a con- 
spicuous thickening (Fig. 2e). Even though 
different, this variance must be considered 
to be within the range of variation in the 
species. For example, in one specimen one 
of the pair of clasping apparati exhibits 
such a thickening and the other resembles 
the more frequently observed apparatus 
(Fig. 2f, g). Considerable variation occurs 
in the curvature of the vertical ramus and 
in that of the talon (Fig. 2a-h). 

Hosts. — In the Brazos Basin, A. ancyla 
is known to infest F. (C.) fodiens, C. (P.) 
ninae, and P. (S.) clarkii, and has been re- 
trieved from collections containing all of 
the crayfishes known to occur in the area 
except C. (P.) texanus, P. (C.) hinei, and P. 
(C.) brazoriensis; rarely, however, was it 
found in collections containing representa- 
tives of C (P.) puer, C. (D.) shufeldtii, and 
P. (C.) incilis. 

Entocytherid associates. — In the 19 lo- 
calities in which this ostracod was found, it 
was the only one infesting the crayfishes in 
nine sites. Its most frequent associate (in 9 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



localities) was A. sinuosa, and only rarely 
was it found with E. reddelli (2), and U. 
simondsi (1). 

Ankylocy there sinuosa (Rioja) 
Figs. 3, 5 

Entocythere cambaria. — Hobbs, 1941:4 [in 
part]. 

Entocythere (Cytherites) heterodonta sin- 
uosa Rioja, 1942a:203, 204, figs. 5, 6 
[Type locality: La Cueva Chica, San Luis 
Potosi, Mexico. Types: not extant. Host: 
Procambarus acutus cuevachicae Hobbs 
1941.]; 1953:287. 

Entocythere (Cytherites) sinuosa. — Rioja, 
1942b:688, 689, 695, 696, fig. 20; 1943a: 
564; 1943b:576. 

Entocythere sinuosa. — Hoff, 1944:330, 
332, 356.— Rioja, 1949:321, 322 [in 
part], figs. 13, 14; 1951:170; 1953:291, 
292.— Tressler, 1954:138; 1959:731, fig. 
28.190.— Hobbs, 1957:431.— Crawford, 
1959:173, 178. 

Ankylocythere sinuosa. — Hart, 1962:127; 
1964:246.— Crawford, 1965: 149.— Red- 
dell, 1965:156; 1970:395; 1971:18; 1981: 
82.— Hobbs, 1966:70, fig. 18; 1971:34- 
35, fig. 22.— Ferguson, 1968:501.— 
Hobbs & Walton, 1968:246.— Baker, 
1969:293.— Reddell & Mitchell, 1969:6; 
1971:142.— Young, 1971:399-409.— 
Hart & Hart, 1974:1, 2, 14, 21, 22, 29- 
31, 34, pi. 3: figs. 11-13, pi. 41.— Hobbs 
III, 1969:5, 14, 20-22, 27, 30, 32-35, 39, 
41, 43, 46, 55, 65, 66, 71, 74, 78, 79, fig. 
5a-k; 1975:281, 290; 1978:506; 1982: 
2.— Hobbs & Peters, 1977:13; 1991:66, 
67.— Hobbs & McClure, 1983:773. 

Entocythere tiphophila Crawford, 1959: 
150, 151, 173-178, 180, 181, figs. 31-37 
[Type locality: roadside ditch 9.1 miles 
(14.6 km) SE of University of South Car- 
olina stadium, Richland County, South 
Carolina, on St. Rte. 48. Types: USNM. 
Hosts: Fallicambarus (C.) uhleri and 
Procambarus (F.) troglodytes]. — Hart, 
1962:123, 128. 

Ankylocythere tiphophila. — Hart, 1962:128; 



1964:245.— Crawford, 1965:149.— 
Hobbs, 1966:71, fig. 16. — Ferguson, 
1968:501.— Peters, 1974:74; 1975:iii, 5- 
8, 10, 13, 20, 22, 23, 27, 45, figs. 2c, 6k, 
14.— Hart & Hart, 1974:15, 20, 33, pi. 4: 
figs. 6-8, pi. 45.— Hobbs & Peters, 1977: 
iii, 3-7, 9, 12, 16, 19-22, 28, 40, 41, 43, 
46, 49-54, 70, fig. 5, map 5.— Hobbs, 
1981:140, 499, 501.— Hobbs & McClure, 
1983:773. 
Ankylocythere tiphophyla. — Hobbs, Holt, & 
Walton, 1967:77 [erroneous spelling]. 

Diagnosis of Texas material. — Shell 
length of male 329-378 (avg. 350) iJim; 
shell height 168-210 (avg. 191) fxm. Peni- 
ferum varying from deeply cleft to truncate 
with tapering acute anteroventrally project- 
ing prominence. Clasping apparatus L- 
shaped with vertical ramus longer that hor- 
izontal ramus; latter with truncate, almost 
straight, anteroventrally projecting talon sit- 
uated slightly proximal to midway between 
preaxial tooth and apex of ramus; apex of 
apparatus with 2 denticles. 

Range. — On the Gulf of Mexico versant, 
from the Cordillera volcanica Transversal 
along the Gulf and Atlantic (lower pied- 
mont and coastal plain) slope to the York 
River Basin in Virginia. Hart and Hart 
(1974:33) also reported it from two locali- 
ties in Ohio, records that should be con- 
firmed. 

Southeastern Texas records (Fig. 5). — 
This ostracod is the most widespread of the 
entocytherids within the study area, occur- 
ring in 94 of the 118 localities represented 
among the collections examined. 

Hosts. — In southeastern Texas this ostra- 
cod was associated with three crayfishes: P. 
(O.) a. acutus, P. (S.) clarkii and P. (G.) 
simulans, but has been found in collections 
containing specimens of all of the other 
species in the area. In the collections from 
94 localities where the ostracod was found, 
P. (O.) a. acutus was a potential host in 60 
of them, P. (5.) clarkii in 52, P. (G.) si- 
mulans in 31, P. (C) incilis in 17, and F. 
(C.)fodiens in 14. All of the other crayfish- 



VOLUME 112, NUMBER 2 



343 




Fig. 3. Variation in male copulatory complex, lateral view, of Ankylocy the re siniiosa (a — Tamaulipas. Ace- 
quia, Mexico; b — Burleson Co, TX; c — Robertson Co., Texas) and Ankylocythere tiphophila (d — Dorchester 
Co., South Carolina; e— Newport News, Virginia); same scale as Fig. 1. 



es were present in fewer than 10 of the col- 
lections. 

Entocytherid associates. — In the 94 lo- 
calities where A. sinuosa was found, it was 
the only ostracod infesting the crayfish(es) 
in 57 of them. In 26 localities it shared the 



host(s) with E. reddelli, in eight with A. an- 
cyla, and two each with E. harrisi and U. 
simondsi. 

Remarks. — There is nothing remarkable 
concerning the size of the animals or in the 
structures employed in distinguishing this 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




97" 96° 

Fig. 4. Distribution of Ankylocythere ancyla and Uncinocythere simondsi in the Brazos River drainage. 



ostracod from its congeners. The length of in Mexico, cited similar ranges in size: 0.34 

the shells of males ranges from 329 to 378 to 0.37 mm and 0.19 to 0.22 mm, respec- 

|jLm and the height from 168 to 210 iJim. tively. 
Hobbs (1971), in reporting on this ostracod Perhaps because of the apparent discon- 



VOLUME 112, NUMBER 2 



345 



y\ 




97" 96° 

Fig. 5. Distribution of Ankylocythere sinuosa in the Brazos River drainage. 



tinuity between the ranges of A. sinuosa of the latter, there seemed to be no reason 

and A. tiphophila and the seemingly con- to suspect that they represented two forms 

sistent presence of a cleft peniferum in the of a single species. With the acquisition of 

range of the former and its absence in that collections almost merging the ranges of 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



the two species, we were prompted to com- 
pare specimens from Mexico to Virginia 
(Fig. 3) and found that in the Gulf Coastal 
area forms with a cleft peniferum, the only 
feature that has served consistently to sep- 
arate the two (Fig. 3a, c), occur in localities 
in which some of the males were lacking a 
cleft (Fig. 3b, d, e). Moreover, in specimens 
from Bleckley County, Georgia (see An- 
dolshek & Hobbs, 1986:fig. 12a), the pen- 
iferum is distinctly excavate, tending to- 
ward the cleft condition. In view of the dis- 
covery of an almost continuous range and 
no character that can be relied upon invari- 
ably to separate the two, we propose that 
A. tiphophila be considered a synonym of 
Rioja's Entocy there sinuosa. 

Reinforcing this proposal is the seeming- 
ly subparallel clinal distribution with re- 
spect to the size of the shell. Andolshek & 
Hobbs (1986:25) reviewed in tabular form 
the shell size reported for A. tiphophila re- 
vealing that the largest specimens occur in 
North Carolina (410 |jLm) and Virginia (390 
(xm), smaller ones in South Carolina (346 
fxm), and the smallest in southeastern Geor- 
gia (321 |jLm). Perhaps significant are the 
sizes reported by Hobbs III (1969:74) for 
A. sinuosa occurring from eastern Texas to 
the panhandle of Florida. The mean shell 
length is 323 fxm as compared with one of 
321 |xm in the material from southeastern 
Georgia. The cline, however, seems to be 
reversed between eastern Texas and Mexi- 
co, for the mean shell length for specimens 
from Mexico was reported by Hobbs (1971: 
36) to be 350 ixm. 

Entocythere harrisi Peters 
Fig. 6 

Entocythere harrisi Peters, 1975:32-33, 
figs. 5a, 6e, f, 7a [Type locality: Rocky 
Creek 4.3 mi (6.9 km) E of U.S. Hwy 29 
on U.S. Hwy 60, Amherst County, Vir- 
ginia. Types: holotype and allotype, 
USNM; paratypes, USNM, H.H. Hobbs 
III, and DIP. Hosts: Cambarus (C.) bar- 
tonii bartonii (Fabricius), C. (Hiaticam- 



barus) longulus (Giraid), and C (P.) ac- 
uminatus Faxon]. — Hobbs & Peters, 
1977:iv, 5, 9, 12-14, 21, 29, 33, 36, 41, 
45, 47, 51, 52, 54, 55, 60, 61, 64, fig. 25; 
1982:314; 1989:328.— Andolshek & 
Hobbs, 1986:30. [The references cited 
here constitute a complete bibliography 
for the species.] 

Diagnosis. — Shell length of male 441- 
570 (avg. 477) jjim; shell height 210-300 
(avg. 244) fjim. Peniferum truncate distally. 
Clasping apparatus "with postaxial border 
[slightly] bowed into heellike prominence 
at junction of horizontal and vertical rami, 
junction thickened; mesial surface of area 
of junction without flange; horizontal ramus 
without oblique ridges on mesial surface" 
(Hobbs & Peters 1977:51). 

Range. — The most recent summary of 
the range of this ostracod was that of Hobbs 
& Peters (1977:52) who based their records, 
except for the type locality, on female spec- 
imens. In the present study we have become 
convinced that the characters they used for 
distinguishing between the females of this 
species and those of their E. intemotalis 
(=E. elliptica Hoff, 1944) are not reliable. 
This conclusion is based upon the obser- 
vation that in two collections of E. harrisi 
(one from Pike County, Arkansas, and an- 
other from Angelina County, Texas) con- 
taining several male and females, none of 
the latter possess the type of genital appa- 
ratus similar to those that were identified 
with that species in North Carolina and Vir- 
ginia. Instead, the genital apparatus of the 
Arkansas and Texas females are indistin- 
guishable from those of E. elliptica and E. 
reddelli. Thus we believe this ostracod is 
represented in collections only by the ho- 
lotype (from Amherst County, Virginia) 
and the specimens cited herein from the 
Brazos River basin, Texas. 

Remarks. — With the new records cited 
herein for the males of this species, we are 
inclined to propose a clinal distribution in 
size with respect to shell height. As the 
largest member is reported from Virginia 



VOLUME 112, NUMBER 2 



347 



y\ 




97 96° 

Fig. 6. Distribution of Entocythere harrisi and Entocythere reddelli in the Brazos River drainage. 



(300 fxm), smaller members occur from However, when the shell length is com- 

Pike Co., Arkansas (265 |xm) and the Braz- pared, the largest member remains in Vir- 

os River drainage (256 ixm), with the small- ginia (570 |jLm) and the smallest (456 iJim) 

est (224 |jLm) appearing in Angelina, Texas, in Angelina, Texas, whereas members from 



348 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



the Brazos River basin (490 fxm) and Ar- 
kansas (476 |jLm) appear reversed. This may 
be due to the small number of specimens 
available for examination. 

Southeastern Texas records (Fig. 6). — A 
male from each of the following localities: 
4.0 mi (6.4 km) E of Marlin, Falls County, 
and McLaughlin Creek, 12.5 mi (20 km) E 
of Cameron on U.S. 90 at Branchville, Mil- 
an County. 

Hosts. — In both localities this ostracod 
was found on P. (G.) simulans and P. (O.) 
a. acutus. 

Entocytherid associates. — A. sinuosa 
was present in the two Brazos localities. 

Entocythere reddelli Hobbs & Walton 
Fig. 6 

Entocythere reddelli Hobbs & Walton, 
1968:243-246, fig. 2a-d [Type locahty: 
Golden Fawn Cave, 8 mi (12.8 km) NNE 
of Boerne, Kendall County, Texas. 
Types: holotype, allotype, and paratypes, 
USNM; paratypes. Academy of Natural 
Sciences of Philadelphia and H.H. Hobbs 
m. Host: P. (5.) clarkii (Girard)]. — Red- 
dell & Mitchell, 1969:6.— Reddell, 1970: 
395.— Hobbs, 1971:40.— Hart & Hart, 
1974:83, 92-93, pi. XXVE: figs. 11-15, 
pi. LE.- Hobbs m, 1975:281.— Hobbs 
& Peters, 1977:iv, 5-7, 9-10, 12, 14, 29, 
36, 41, 43, 45, 47, 52, 54, 55, 73; 1982: 
313.— Andolshek & Hobbs, 1986:30. 
[These references constitute what we be- 
lieve to be a complete bibliography for 
the species.] 

Diagnosis. — Shell length of male 546— 
581 (avg. 564) ixm; shell height 280-294 
(avg. 287) |jLm. Peniferum subtruncate ven- 
trally and possessing rounded antero- and 
postero ventral extremities. "Clasping ap- 
paratus with postaxial border at junction of 
horizontal and vertical rami produced in 
heellike prominence and junction thick- 
ened; mesial surface of area of junction 
with angular flange, apex of angle reaching 
level proximal to proximal tooth on preax- 
ial margin of horizontal ramus, horizontal 



ramus lacking long oblique ridge extending 
across mesial surface. Female genital ap- 
paratus composed of ventrally directed sub- 
spiculiform projection arising from bipartite 
base, latter embedded in amorphous mass" 
(Hobbs & Peters 1977:55), second antenna 
with appendix at base of terminal claws 
conspicuously enlarged, pectinate, bearing 
2 or 3 broad teeth or as many as 6 finely 
divided denticles. 

Range. — In addition to its presence in 
Kendall County, Texas, Hart & Hart (1974: 
93) reported its occurrence in Greene Coun- 
ty, Arkansas, and Sumner County, Kansas. 
Hobbs & Peters (1977:55) also reported it 
from the Catawba, French Broad, and Hi- 
wassee basins in North Carolina. Twenty- 
five localities in the Brazos River basin of 
southeastern Texas are cited here. 

Southeastern Texas records (Fig. 6). — 
Twenty-five localities from the following 
counties: Brazoria (8), Brazos (1), Burleson 
(1), Fort Bend (7), Matagorda (4), Milan 
(1), Robertson (2), and Victoria (1). 

Remarks. — Except for the distribution of 
this ostracod in the Brazos River basin, its 
range is poorly known. The total absence of 
records in what appears to be the central 
part of its range is disturbing, but there is 
little reason to suspect that the type locality 
lies on its western limit, and if and where 
its range and that of E. cambaria Marshall 
and E. illinoisensis Hoff meet or intersect 
to the north have not been determined. 

Hosts. — In the Brazos basin, E. reddelli 
was found infesting P. (O.) a. acutus and 
P. (S.) clarkii. It also occurred in collections 
containing one or both of these crayfishes 
along with one or more of the following: 
C. (D.) shufeldtii, C. (P.) ninae, C. (P.) 
puer, C. (P.) texanus, F. (C.) fodiens, P. 
(C) brazoriensis, P. (C.) incilis, and P. (G.) 
simulans. 

Entocytherid associates. — In all 25 lo- 
cahties in which this ostracod was found, it 
was associated with A. sinuosa, and in one 
of them A. ancyla (hosts: P. (O.) a. acutus 
and P. (S.) clarkii) was also present; in an- 



VOLUME 112, NUMBER 2 



349 



Other of these locahties U. simondsi was in- 
festing the same two host species. 

Uncinocythere simondsi Hobbs & Walton 
Fig. 4 

Entocythere simondsi Hobbs & Wahon, 
1960:17, 20-21, figs. 1-10 [Type locali- 
ty: Dunn Creek, 1.9 mi (3 km) west of 
Fighting Town Creek on Hell's Hollow 
Road, Fannin County, Georgia. Hosts: C. 
(C) bartonii and C sp. {=C. (D.) lati- 
manus)]. 

Uncinocythere simondsi. — Hart, 1962:138. 
[A complete bibliography for the species 
is presented by Andolshek & Hobbs, 
1986:39.] 

Diagnosis. — Shell length of male 329- 
343 (avg. 336) ixm; shell height 154-196 
(avg. 175) |JLm. Copulatory complex of 
peniferum terminating distally in bifid 
tip. Clasping apparatus L- shaped with 
preaxial border bearing 3 distinct teeth; 
postaxial border entire, lacking any ex- 
crescence; distal extremity with 3 denti- 
cles. 

Range. — From Illinois, Kentucky, and 
North Carolina southward to Brazoria and 
Washington counties, Texas, and northern 
Florida, previously known no farther west 
than Mississippi (Hart & Hart 1974; Hobbs 
& Peters 1977, 1982; Andolshek & Hobbs 
1986). 

Southeastern Texas records (Fig. 4). — 
This ostracod was found in the following 
localities: 2 mi (3.2 km) N, 2 mi (3.2 km) 
E of Brenham on St Rte 90, Washington 
County; and 1.25 mi (2 km) E of Rosharon 
on Farm Rd 1462, Brazoria County. 

Hosts. — Procambarus (5.) clarkii was 
one of the hosts in both of the Brazos col- 
lections in which this ostracod was found. 
In one of them, P. (O.) a. acutus was also 
present, and in the other, P. (G.) simulans 
was in the container from which the ostra- 
cods were removed. 

Entocytherid associates. — In both of the 
localities in which this ostracod was found, 
A. sinuosa and E. reddelli also were pre- 



sent. In the collection from Washington 
County, A. ancyla also was found with 
them. 

Acknowledgments 

We extend our thanks to the late H. H. 
Hobbs, Jr. who encouraged us in this en- 
deavor and provided the materials, guid- 
ance, and insight for this study. We are also 
grateful to Horton Hobbs III for his criti- 
cism of the manuscript and to Brian Ken- 
sley, Ray Manning, and Karen Reed for 
their assistance. 

Literature Cited 

Albaugh, D. W. 1973. Life Histories of the Crayfishes 
Procambarus acutus and Procambarus hinei in 
Texas. Unpublished Ph. D. dissertation, Texas 
A & M University, College Station, 135 pp. 

. 1975. A new crawfish of the genus Procam- 
barus, subgenus Capillicambarus, from Tex- 
as. — Tulane Studies in Zoology and Botany 19: 
1-7. 

, & J. B. Black. 1973. A new crawfish of the 



genus Cambarellus from Texas, with new Texas 
distributional records for the genus (Decapoda, 
Astacidae). — Southwestern Naturalist 18:177- 
185. 

Andolshek, M. D., & H. H. Hobbs, Jr. 1986. The en- 
tocytherid ostracod fauna of southeastern Geor- 
gia. — Smithsonian Contributions to Zoology 
424, 43 pp. 

Baker, J. H. 1969. On the relationship of Ankylocy the re 
sinuosa (Rioja, 1942) (Ostracoda, Entocytheri- 
dae) to the crayfish Procambarus simulans sim- 
ulans (Faxon, 1884). — Transactions of the 
American Microscopical Society 88:293-294. 

Crawford, E. A., Jr. 1959. Five new ostracods of the 
genus Entocythere (Ostracoda, Cytheridae) 
from South Carolina. — University of South Car- 
olina Publications, Series III, Biology 2:149- 
189. 

. 1965. Three new species of epizoic ostracods 

(Ostracoda, Entocytheridae) from North and 
South Carolina. — American Midland Naturalist 
74:148-154. 

Ferguson, E. 1968. Recently described species and dis- 
tributional records for North American fresh- 
water Ostracoda.— American Midland Natural- 
ist 79:499-506. 

Hart, D. G., & C. W. Hart, Jr 1974. The ostracod fam- 
ily Entocytheridae. — Academy of Natural Sci- 
ences of Philadelphia Monograph 18:239 pp. 

Hart, C. W., Jr. 1962. A Revision of the ostracods of 



350 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



the family Entocytheridae. — Proceedings of the 
Academy of Natural Sciences of Philadelphia 
114:121-147. 
-. 1964. Two new entocytherid ostracods from 



of the Academy of Natural Sciences of Phila- 
delphia 120:237-252. 
-, P C. Holt, & Margaret Walton. 1967. The 



the vicinity of Washington, D.C. — Proceedings 
of the Biological Society of Washington 77: 
243-246. 

Hobbs, H. H., Jr. 1941. A new crayfish from San Luis 
Potosi, Mexico (Decapoda, Astacidae). — Zool- 
ogica. Scientific Contributions of the New York 
Zoological Society 26:1-4. 

. 1957. Observaciones acerca de las especies 

del genero Entocythere (Crustaceos, Ostraco- 
dos) de Cuba. — Anales del Instituto de Biolo- 
gia, Universidad Nacional Autonoma de Mexi- 
co 27:431-436. 

. 1966. An illustrated key to the species of the 

genus Ankylocythere with a description of a 
new species from Louisiana (Ostracoda, Ento- 
cytheridae). — Proceedings of the Louisiana 
Academy of Sciences 29:67-75. 

. 1971. The entocytherid ostracods of Mexico 

and Cuba. — Smithsonian Contributions to Zo- 
ology 81, 55 pp. 

. 1981. The crayfishes of Georgia. — Smithson- 
ian Contributions to Zoology 318, 559 pp. 

, & A. C. McClure. 1983. On a small collection 

of entocytherid ostracods with the descriptions 
of three new species. — Proceedings of the Bi- 
ological Society of Washington 96:770—779. 

, & F. O. Perkins. 1967. A new burrowing cray- 
fish from North Carolina (Decapoda, Astaci- 
dae). — Proceedings of the Biological Society of 
Washington 80:141-146. 

, & D. J. Peters. 1977. The entocytherid ostra- 
cods of North Carolina. — Smithsonian Contri- 
butions to Zoology 247, 73 pp. 

, & . 1982. The entocytherid ostracod 



fauna of northern Georgia. — Proceedings of the 
Biological Society of Washington 95:297-318. 

-, & . 1989. New records of entocytherid 

ostracods infesting burrowing crayfishes, with 
the description of a new species, Ascetocythere 
stockeri. — Proceedings Biological Society of 
Washington 102:324-330. 

-, & . 1991. Additional records of ento- 
cytherid ostracods infesting burrowing crayfish- 
es, with descriptions of five new species. — Pro- 
ceedings of the Biological Society of Washing- 
ton 104:64-75. 

— , & M. Walton. 1960. Three new ostracods of 
the genus Entocythere from the Hiwassee drain- 
age system in Georgia and Tennessee. — Journal 
of the Tennessee Academy of Science 35:17- 
23. 

— , & . 1968. New entocytherid ostracods 

from the southern United States. — Proceedings 



crayfishes and their epizootic ostracod and bran- 
chiobdellid associates of the Mountain Lake, 
Virginia, region. — Proceedings of the United 
States National Museum 123 (3602): 1-84, 22 
figs. 

Hobbs III, H. H. 1969. Studies in ecological and host 
specificity in entocytherid ostracods (Ostracoda: 
Entocytheridae). Unpublished M.S. thesis, Mis- 
sissippi State University, State College, 93 pp. 

. 1975. Distribution of Indiana cavemicolous 

crayfishes and their ecto-commensal ostra- 
cods. — International Journal of Speleology 7: 
273-302. 

. 1978. New species of ostracods from the Gulf 

coastal plain (Ostracoda: Entocytheridae). — 
Transactions of the American Microscopical So- 
ciety 97:503-511. 

. 1982. Host-commensal relationships of cray- 



fishes and ostracods of the gulf coast plain, 
U.S.A. — Ohio Journal of Science 82:2. 

Hoff, C. C. 1944. New American species of the ostra- 
cod genus Entocythere. — American Midland 
NaturaUst 32:327-357. 

Peters, D. J. 1974. The ecological and geographical 
distribution of some Virginia ostracods. — As- 
sociation of Southeastern Biologists Bulletin, 
21:74. 

. 1975. The entocytherid ostracod fauna of the 

James and York River basins with a description 
of a new member of the genus Entocythere. — 
Virginia Polytechnic Institute and University, 
Research Division Bulletin 93, 50 pp. 

Reddell, J. R. 1965. A checklist of the cave fauna of 
Texas. I. The Invertebrata (exclusive of Insec- 
ta). — Texas Journal of Science 17:143-187. 

. 1970. A checklist of the cave fauna of Texas. 

rV. Additional records of Invertebrata (exclusive 
of Insecta). — Texas Journal of Science 21:389— 
415. 

. 1971. A preliminary bibliography of Mexican 

cave biology with a checklist of published rec- 
ords. — Association for Mexican Cave Studies 
Bulletin 3:1-184. 

. 1981. A review of the cavemicole fauna of 

Mexico, Guatemala, and Belize. — Texas Me- 
morial Museum Bulletin 27:1-327. 

, & R. W. Mitchell. 1969. A checklist and an- 
notated bibliography of the subterranean aqua- 
ticfauna of Texas. — Special Report of the Inter- 
national Center for Arid and Semi-arid Land 
Studies 24:1-48. 

, & . 1971. Studies on the cavemicole 



fauna of Mexico. — Association for Mexican 
Cave Studies, Bulletin 4: 1-239. 
Rioja, E. 1942a. Descripcion de una especie y una sub- 



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351 



especie nueva del genero Entocythere Marshall, 
procedentes de La Cueva Chica. — Ciencia 
(Mexico) 3:201-204. 

-. 1942b. Estudios carcinologicos. XIII. Consi- 
deraciones y datos cerca del genero Entocythere 
(Crust. Ostracodos) y algunas de sus especies, 
con descripcion de una neuva. — Anales del In- 
stitute de Biologia, Mexico 13:685-697, 21 
figs. 

-. 1943a. Estudios carcinologicos. XIV. Nuevos 
datos acerca de los Entocythere (Crust. Ostra- 
codos) de Mexico. — Anales del Institute de 
Biologia, Universidad Nacional Autonoma de 
Mexico 14:553-566. 

-. 1943b. Estudios carcinologicos. XV. Polimor- 
fismo femenino en los ostracodos del genero 
Entocythere. — Anales del Institute de Biologia, 
Universidad Nacional Autonoma de Mexico 14: 
567-585. 

-. 1949. Estudios carcinologicos. XXI. Contri- 
bucion al conocimiento de las especies del Ge- 
nero Entocythere de Mexico. — Anales del In- 



stitute de Biologia, Universidad National Au- 
tonoma de Mexico 20:315-329. 

-. 1951. Estudios carcinologicos. XXV. El hal- 
lazge del genero Sphaeromicola en America 
(Ostracodos, Citeridos) y descripcion de una 
nueva especie. — Anales del Institute de Biolo- 
gia, Universidad Nacional Autonoma de Mexi- 
co 22:169-179. 

-. 1953. Los crustaceos cavemicolas de Mexico. 



IV. Centenario de la Universidad de Mexico 
(1851-1951). Memoria del Congreso Cientifico 
Mexicano. — Ciencias Biologicas 7:285-298. 

Tressler, W. L. 1954. Fresh-water Ostracoda from Tex- 
as and Mexico. — Journal of the Washington 
Academy of Sciences 44:138-149. 

. 1959. Ostracada. Pp. 657-734 in W. T. Ed- 

mondson, ed. Ward, H. B. and Whipple's Fresh- 
water Biology, 2nd edition, John Wiley and 
Sons, New York, 1248 pp. 

Young, W 1971. Ecological studies of the Entocyther- 
idae (Ostracoda). — American Midland Natural- 
ist 85:399-409. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):352-36L 1999. 

A new genus, Neodoxomysis 

(Crustacea: Mysidacea: Mysidae: Leptomysini), with 

description of two new species 

Masaaki Murano 

Institute of Environmental Ecology, METOCEAN Co. Ltd., 1334-5 Riemon, 
Ooigawa-cho, Shida-gun, Shizuoka 421-0212, Japan 

Abstract. — A new genus, Neodoxomysis, is established for two new species, 
N. elongata and N. sahulensis, collected from Sahul Shelf. The new genus is 
closely related to Doxomysis, but is different from the latter genus in having 
only a single strong, modified seta on the exopod of the fourth male pleopod, 
as compared with two in Doxomysis. Doxomysis littoralis Tattersall, 1922, is 
transferred to the new genus. 



A single male specimen identified by li 
(1964) as Doxomysis littoralis Tattersall, 
1922, is quite different from the other Dox- 
omysis species in having a single strong 
modified seta on the exopod of the fourth 
male pleopod. The exopod in all other Dox- 
omysis species is armed with two strong 
modified setae. 

During a cruise to southeastern Asian 
seas (KH-72-1) by the RA^ Hakuho Maru 
of the Ocean Research Institute, University 
of Tokyo, two undescribed species, which 
were similar to D. littoralis with respect to 
the morphology of the fourth male pleopod, 
were collected. This morphological char- 
acter is distinct these species from those of 
the genus Doxomysis. Therefore, a new ge- 
nus is estabhshed to receive these three spe- 
cies, and two new species are described. 
The type specimens are deposited in the 
National Science Museum, Tokyo (NSMT). 

Neodoxomysis, new genus 

Diagnosis. — Carapace produced anteri- 
orly into triangular rostral plate with round- 
ed apex. Antermal scale lanceolate with 
rounded apex, armed with setae on whole 
margins. Eye functionally normally devel- 
oped, without papilliform process on eye- 
stalk. Maxilla with second segment of en- 



dopod wider than long, expanded distally, 
with about 10 strong spines on distal mar- 
gin. Labrum with rounded frontal margin. 
Endopods of third to eighth thoracic limbs 
with propodus divided into 2 subsegments. 
Endopod of uropod with row of numerous 
spines along inner margin. Telson with lat- 
eral margin armed with spines throughout, 
apical cleft deep, armed with marginal spi- 
nules and pair of plumose setae arising 
from anterior end. Exopod of fourth pleo- 
pod of male modified, longer than endopod, 
ultimate segment small, with 1 or 2 short 
simple setae, penultimate segment with sin- 
gle strong seta. 

Type species. — Neodoxomysis elongata, 
new genus, new species. 

Etymology. — Derived from its relation- 
ship to Doxomysis. It is feminine in gender. 

Remarks. — The new genus Neodoxomy- 
sis closely resembles Doxomysis Hansen, 
1912, except for the fourth male pleopod. 
In the new genus, the exopod of the fourth 
male pleopod is armed with only a single, 
strong, modified seta arising from the pen- 
ultimate segment, while in Doxomysis as 
well as eight related genera in the tribe Lep- 
tomysini (Talbot 1997), the exopod is 
armed with two strong modified setae, one 
arising from the penultimate segment and 
the other from the antepenultimate segment. 



VOLUME 112, NUMBER 2 



353 



Among 11 Doxomysis species in which 
the exopod of the fourth male pleopod has 
been described, only D. littoralis bears a 
single strong modified seta on the exopod. 
This species was established on specimens 
from the Andaman Islands by Tattersall 
(1922); later the male was described from 
the South China Sea by li (1964). Accord- 
ingly, D. littoralis should be transferred to 
the new genus. The new genus Neodoxo- 
mysis comprises a total of three species. 

Neodoxomysis elongata, new species 
Figs. 1, 2 

Type series. — Holotype (NSMT-Cr 
12487), adult male (5.3 mm), allotype 
(NSMT-Cr 12488), adult female with em- 
bryos (4.9 mm), paratypes (NSMT-Cr 
12489), 6 adult males (6.1-6.8 mm) and 3 
adult females (4.5-5.0 mm); Sahul Shelf, 
12°37.3'S, 124°33.9'E to 12°36.0'S, 
124°36.4'E; 74-78 m; 25-26 June 1972; 
plankton net installed in mouth of 3-m 
beam trawl. 

Other material. — 8 adult males, 3 adult 
females, 15 immature males and 19 imma- 
ture females; collection data same as type 
series. 

Description. — Body somewhat slender. 
Carapace produced anteriorly into triangu- 
lar rostral plate with rounded apex extend- 
ing beyond basal margin of antennular pe- 
duncles, leaving whole eyes uncovered 
(Fig. lA, B); anterolateral comer rounded; 
posterior margin slightly emarginate, leav- 
ing last thoracic somite exposed. 

Eye developed, relatively large; cornea 
occupying more than half of whole organ, 
spherical, wider than eyestalk; eyestaUc his- 
pid on anterior and posterior surfaces, with- 
out papilliform process (Fig. lA, B). 

Antennular peduncle of male more robust 
than that of female, first segment with sev- 
eral setae at outer distal comer and 1 seta 
at inner distal comer, second segment short, 
narrower than preceding one; third segment 



as long as first, wider than preceding two 
segments, with 1 straight and 4 short 
curved setae at inner distal comer; proces- 
sus masculinus large, hirsute (Fig. lA). An- 
tennular peduncle of female slender; first 
segment as long as succeeding 2 segments 
together; second segment short, narrow, 
with 1 seta at inner distal comer; third seg- 
ment with 1 seta at distal third of inner mar- 
gin and 5 setae on inner distal margin (Fig. 
IB). 

Antennal scale extending beyond distal 
margin of third segment of antennular pe- 
duncle for about Vs of its length (Fig. lA, 
B), lanceolate, 6.3 times as long as broad, 
outer margin slightly concave, distal seg- 
ment ViQ of length of scale (Fig. IC). An- 
tennal peduncle short, reaching proximal 
third of antennal scale, 3-segmented, third 
segment longest; sympod with thorn at out- 
er distal comer (Fig. IC). 

Mandible with well developed mastica- 
tory edge. Mandibular palp 3-jointed, sec- 
ond segment elongated oval, third segment 
0.52 of second in length, outer margin with 
2 series of setae, proximal setae longer, 
barbed on proximal half, distal setae barbed 
on whole length (Fig. ID). Maxillule: inner 
lobe with 9 setae on inner margin, 5 setae 
on outer margin, 3 setae on ventral surface 
and 3 stout and 1 slender setae on distal 
margin; outer lobe with about 10 spines on 
apical margin and 3 setae on ventral surface 
(Fig. IE). Maxilla: second segment of en- 
dopod expanded distally, wider than long, 
distal margin slightly convex, 2.5 times as 
wide as at base, armed with 10 stout spines, 
which are rounded and flattened at tip, out- 
ermost spine longer than others, inner 6 
spines, especially 2 innermost ones armed 
with short setae on margins; exopod rather 
rectangular in shape, extending beyond dis- 
tal margin of first segment of endopod (Fig. 
IF). Labrum with frontal margin rounded 
(Fig. IG). 

Endopod of first thoracic limb robust, 
dactylus wider than long, with strong ter- 
minal claw (Fig. IH). Endopod of second 



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Fig. L Neodoxomysis elongata, new species; A, C-J: holotype; B: allotype. A, anterior end of adult male; 
B, anterior end of adult female; C, antenna; D, mandible and mandibular palp; E, maxillule; F, maxilla; G, 
labrum; H, endopod of first thoracic limb; I, endopod of second thoracic limb; J, sixth thoracic limb. 



VOLUME 112, NUMBER 2 



355 




Fig. 2. Neodoxomysis elongata, new species; A-F: holotype; G: allotype. A. first male pleopod; B, fourth 
male pleopod; C, distal part of fourth male pleopod; D, fifth male pleopod; E. endopod of uropod; F uropod 
and telson; G, distal part of telson. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



thoracic limb slender, basis with inner lobe 
developed, ischium with long setae on inner 
margin, merus as long as carpopropodus 
and dactylus combined, dactylus with ter- 
minal claw and 6 barbed setae (Fig. 11). 

Endopod of sixth thoracic limb slender, 
carpus separated obliquely from 2-subseg- 
mented propodus, distal propodal subseg- 
ment longer than proximal, equal to carpus 
in length; dactylus small, with slender ter- 
minal claw (Fig. IJ). Exopods of thoracic 
limbs with basal plate armed with small 
acute process at outer distal comer, flagel- 
liform part 8 -segmented in first and eight 
limbs, 9-segmented in second to seventh 
limbs (Fig. IJ). 

Abdomen consisting of 6 somites, first 5 
somites subequal, sixth somite 1.7 times 
longer than fifth. 

Male pleopods well developed, natatory. 
First pair with 7 -segmented exopod and un- 
segmented endopod (Fig. 2A). Second and 
third pairs with 7 -segmented exopod and 6- 
segmented endopod extending to distal 
margin of sixth segment of exopod. Fourth 
pair: exopod 1.6 times longer than endopod, 
11 -segmented, seventh and eighth segments 
unarmed with setae, ninth segment with 1 
short seta at outer distal end, tenth segment 
with 2 setae on distal end, longer one ex- 
tremely long, 0.4 of exopod in length, 
straight, feathered in distal half, shorter one 
simple, 0.2 as long as longer one, terminal 
segment very small, V3 of preceding one in 
length, with 1 short seta; endopod 6- seg- 
mented, reaching distal end of sixth seg- 
ment of exopod, without modified setae 
(Fig. 2B, C). Fifth pair: exopod 7-segment- 
ed, longer than endopod, endopod 6-seg- 
mented, with triangular lobe tipped with 
seta on outer margin of first segment (Fig. 
2D). 

Exopod of uropod slender, long, slightly 
curved outwardly, extending beyond distal 
end of telson for its distal half (Fig. 2F). 
Endopod of uropod extending beyond distal 
end of telson for distal third, armed along 



inner margin from statocyst region to near 
apex with about 42 spines which are made 
up with longer and shorter ones arranged 
alternately, except several spines in distal 
and proximal portions (Fig. 2E, F). 

Telson slightly longer than last abdomi- 
nal somite, 1.7 times as long as broad at 
base, abruptly narrowing near base, paral- 
lel-sided in middle part, then gradually nar- 
rowing towards distal end, cleft at apex for 
V3 of telson length; lateral margin armed 
along whole length with about 16 spines, 
sparsely in proximal third and densely in 
distal %; each apex of distal lobes rather 
truncate, with 3 somewhat obtuse spines, 
outermost one longest in male and middle 
one in female; cleft with small notch at an- 
terior end, each side with about 16-18 spi- 
nules along whole length except for poste- 
rior short distance naked, pair of plumose 
setae arising from anterior end of cleft (Fig. 
2F, G). 

Etymology. — The name elongata refers 
to the slender body. 

Remarks. — Neodoxomysis elongata is 
considerably different from the other two 
species of the genus, N. sahurensis and N. 
littoralis, in the following aspects: The an- 
tennal scale is 6.3 times as long as broad in 
this species, while it is about 5 times as 
long in the other two species; the exopod 
of the fourth male pleopod is 11 -jointed in 
this species as compared with 7 in the other 
two species; and, spines on the second en- 
dopod segment of the maxilla are rounded 
and flattened distally in this species, while 
these are sharply or obtusely pointed in the 
other two species. 

Neodoxomysis sahulensis, new species 
Figs. 3, 4 

Type series. — Holotype (NSMT-Cr 
12490), adult male (5.0 nam); allotype 
(NSMT-Cr 12491), adult female (4.0 mm); 
paratypes (NSMT-Cr 12492), 2 adult fe- 
males (3.7, 4.0 mm), 1 adult male (divided 
into two parts); Sahul Shelf, 12°17.3'S, 



VOLUME 112, NUMBER 2 



357 




Fig. 3. Neodoxomysis sahulensis, new species; A, C-G: holotype; B: allotype. A, anterior end of adult male; 
B, anterior end of adult female; C, antenna; D, mandible and mandibular palp; E. maxillule; F, maxilla; G, 
labrum; H, endopod of first thoracic limb; I, second thoracic limb. 



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Fig. 4. Neodoxomysis sahulensis, new species; A: allotype; B— F: holotype. A, endopod of one of posterior 
thoracic limbs; B, endopod of eighth thoracic limb; C, fourth male pleopod; D, distal part of exopod of fourth 
male pleopod; E, fifth male pleopod; F, uropod and telson. 



VOLUME 112, NUMBER 2 



359 



129°40.9'E to 12°17.2'S, 129°41.8'E; 49- 
52 m; 24 June 1972; plankton net installed 
in mouth of 3-m beam trawl. 

Description. — Carapace with anterior 
margin produced in triangular rostrum with 
narrowly rounded apex and almost straight 
lateral margins, leaving whole eyes exposed 
(Fig. 3A, B); anterolateral comer rounded; 
posterior margin somewhat emarginate, 
leaving last thoracic somite uncovered. 

Eye moderately developed; cornea oc- 
cupying about half of whole eye, slightly 
wider than stalk; stalk hispid on anterior 
and posterior surfaces (Fig. 3A, B). 

Antennular peduncle of male more robust 
than that of female, first segment as long as 
third, with anterolateral comer slightly pro- 
duced anteriorly, armed with several setae, 
second segment short and narrow, third seg- 
ment wider than preceding segments, with 
1 short seta at distal third of inner margin 
and 2 setae at anteromedian comer; proces- 
sus masculinus large (Fig. 3A). In female, 
first segment longer than and as wide as 
third, second segment with 1 seta at anter- 
omedian comer, third segment with 1 seta 
at distal third of inner margin and 4 setae 
at anteromedian comer (Fig. 3B). 

Antennal scale setose on whole margins, 
overreaching distal end of antennular pe- 
duncle for Vz of its length (Fig. 3 A, B), 
more than 5 times as long as broad, outer 
margin nearly straight, inner margin convex 
in proximal half, distal segment V^ of length 
of scale (Fig. 3C). Antennal peduncle short, 
barely extending to middle of antennal 
scale, composed of 3 segments, distal 2 
equal in length, longer than proximal one; 
antennal sympod with long, acute process 
at outer distal comer (Fig. 3C). 

Mandible with well developed masticatory 
edge; mandibular palp with second segment 
elongated oval, third segment 0.6 as long as 
second, outer margin with 2 series of setae, 
proximal setae longer, barbed on proximal 
half, distal one barbed along whole length 
(Fig. 3D). Maxillule: inner lobe with 7 setae 
on inner margin, 4 setae on outer margin and 



3 stout and 1 slender setae on apical margin; 
outer lobe with about 10 spines on apical 
margin, about 10 spinules on middle part of 
outer margin and 3 setae on ventral surface 
(Fig. 3E). Maxilla: distal segment of endopod 
expanded distally, wider than long, distal 
margin sUghtly arched, 2.5 times as wide as 
at base, armed with 7 strong spines of which 
outer one is longer than others; exopod ex- 
tending to distal end of first endopod segment 
(Fig. 3H). Labrum with frontal margin round- 
ed (Fig. 3G). 

First thoracic endopod robust, ischium 
wider than long, with developed inner lobe, 
merus relatively slender, dactylus wider 
than long, with stout, straight claw (Fig. 
3H). Second thoracic endopod rather slen- 
der, basis with developed inner lobe, ischi- 
um with about 8 long setae on inner margin, 
merus as long as carpopropodus and dac- 
tylus together, dactylus longer than wide, 
with 1 barbed claw and 6 barbed setae (Fig. 
31). Third to eighth thoracic endopods be- 
coming more slender towards posterior 
pairs, carpus articulated obliquely with pro- 
podus but in eighth limb articulation is 
nearly transverse, propodus divided into 2 
subsegments, in eight limb distal subseg- 
ment 1.6 times longer than proximal (Fig. 
4A, B). Exopods with flagelliform part 9- 
segmented in middle pairs; basal plate with 
outer distal comer pointed (Fig. 31). 

Abdomen with sixth somite longest, 1.3 
times as long as fifth. 

Male pleopods well developed, bira- 
mous. First pleopod with 7-segmented ex- 
opod and unsegmented endopod. Second 
and third pleopods with 7-segmented exo- 
pod longer than 6-segmented endopod. Ex- 
opod of fourth pleopod modified, 7-seg- 
mented, nearly 1.5 times longer than en- 
dopod, terminal segment very small, VS of 
preceding segment in length, armed with 1 
tiny seta at distal end, penultimate segment 
with 1 strong spinose seta which is nearly 

4 times longer than segment supporting it, 
antepenultimate segment armed with 1 
short feeble seta at distal end of outer mar- 
gin; endopod 6-segmented, normal, without 



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Table 1. — Morphological differences between Neodoxomysis sahulensis and A^. littoralis. 



N. sahulensis 



N. littoralis 



Second segment of endopod 

of maxilla 
Exopod of 4th pleopod of male 
Endopod of 4th pleopod of 

male 
Endopod of 5th pleopod of 

male 
Telson 



With 7 acute, simple spines 

7-jointed, with 1 seta at distal end 
Extending to middle of 4th segment 

of exopod 
Extending to distal end of 5th 

segment of exopod 
1.5 times as long as broad, lateral 

margin with 16—17 spines; cleft Va 

as long as telson, with 11-12 

spines on each side 



With 9 slender, bluntly pointed 

spines 
7-jointed, with 2 setae at distal end 
Extending to distal end of 5th 

segment of exopod 
Extending to distal end of 6th 

segment of exopod 
1.75 times as long as broad, lateral 

margin with 16 spines; cleft % as 

long as telson, with 16—19 spines 

on each side 



modified setae (Fig. 4C, D). Fifth pleopod 
with exopod 7-segmented, 1.2 times longer 
than 6-segmented endopod; endopod with 
first segment with digitiform projection 
tipped with 1 seta on outer margin in ad- 
dition to usual side lobe (Fig. 4E). 

Endopod of uropod extending beyond 
distal end of telson for V3 of its length, inner 
margin from statocyst region to near apex 
with 34 spines, which are composed of lon- 
ger and shorter spines arranged alternately 
except for several spines on proximal and 
distal portions, distal 4 spines long and 
acute, other spines obtuse (Fig. 4F). Exopod 
of uropod overreaching endopod for V^ of 
its length (Fig. 4F). 

Telson as long as last abdominal somite, 
1.5 times as long as broad at base, deeply 
cleft at apex; lateral margin furnished with 
16-17 rather long spines along whole 
length, distal 10 spines gradually increasing 
in length towards apex; each apex of distal 
lobes armed with 3 spines, outer one of 
which is slightly longer than others; apical 
cleft V4 as long as telson, furnished with 11- 
12 spinules on each side and pair of plu- 
mose setae arising from anterior end (Fig. 
4F). Spination on telson in female similar 
to that of male. 

Etymology. — The name sahulensis refers 
to the locality in which the specimens were 
collected. 

Remarks. — Neodoxomysis sahulensis is 
closely similar to N. littoralis (Tattersall, 



1922) in many respects, but distinguished 
from that species as shown in Table 1. 

In the new species the outer lobe of the 
maxillule is furnished with a row of spi- 
nules on the middle part of the outer mar- 
gin. Such a character has been observed in 
Doxomysis quadrispinosa by Pillai (1973) 
and D. australiensis by Tattersall (1940). 

Literature Cited 

Hansen, H. J. 1912. Report on the scientific results of 
the expedition to the eastern tropical Pacific, in 
charge of Alexander Agassiz, by the U.S. Fish 
Commission Steamer "Albatross", from Au- 
gust, 1899, to March, 1900, Commander Jeffer- 
son F. Moser, U. S. N., commanding. XVI. Re- 
ports on the scientific results of the expedition 
to the eastern tropical Pacific, in charge of Al- 
exander Agassiz, by the U.S. Fish Commission 
Steamer "Albatross", from October, 1904 to 
March, 1905. Lieut. -Commander L. M. Garrett, 
U. S. N., commanding. XXVII. The Schizopo- 
da. — Memoirs of the Museum of Comparative 
Zoology at Harvard College 35:175-296, pis. 
1-12. 

li, N. 1964. Fauna Japonica, Mysidae (Crustacea). Bio- 
geographical Society of Japan, Tokyo, 610 pp. 

Pillai, N. K. 1973. Mysidacea of the Indian Ocean. 
Papers on the zooplankton collection of the 
IIOE.-Handbook to the International Zooplank- 
ton Collections 4:1-125. 

Talbot, M. S. 1997. Doxomysis acanthina, a new lep- 
tomysinid (Crustacea: Mysidacea) from the 
northern Great Barrier Reef, Australia, with ex- 
tensions to the known distributions of D. aus- 
traliensis W. M. Tattersall, 1940 and D. spinata 
Murano, 1990, and a key to the genus Doxo- 



VOLUME 112, NUMBER 2 361 

my sis. — Proceedings of the Biological Society . 1940. Report on a small collection of Mysi- 

of Washington 110:426-438. dacea from the coastal waters of New South 

Tattersall, W. M. 1922. Indian Mysidacea. — Records Wales. — Records of the Australian Museum, 

of the Indian Museum 24:445-504. 20:327-340. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):362-367. 1999. 

First record of the family Gynodiastylidae Stebbing, 1912 

(Crustacea: Malacostraca: Cumacea) from Antarctic waters with the 

description of Gynodiastylis jazdzewskii, a new species 

Magdalena Blazewicz and Richard W. Heard 

(MB) University of Lodz, Department of Invertebrate Zoology and Hydrobiology, 

Laboratory of Polar Biology and Oceanobiology, 90-237 Lodz, S. Banacha str. 12/16, Poland; 

(RWH) University of Southern Mississippi Institute of Marine Sciences, 

Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 39566-7000 

Abstract. — An adult male of an undescribed gynodiastylid cumacean was 
collected in a Menzies trawl sample taken at a depth of between 388 and 399 
m in the Ross Sea (76°01.5'-76°01.0'S, 179°49.9'-179°52.3'E) during February 
1972. The specimen, which had lost its third maxillipeds and the last four 
articles of the first pereopods, is placed in the genus Gynodiastylis Caiman, 
1911. Gynodiastylis jazdzewskii, new species, the first member of its family 
reported from Antarctic waters, can be distinguished from the 59 previously 
described species of Gynodiastylis and members of the other five gynodiastylid 
genera by a combination of characters, including the finer, more irregularly 
wavy, longitudinal ridges, and the length and setation of the telson, on the 
carapace, the shape and terminal setation of the telson, the length of telson in 
relation to that of the last abdominal somite and uropodal peduncles, and the 
setation and relative length of the uropodal rami. 



An examination of the Cumacea collect- 
ed by the R/V Eltanin in the Ross Sea, re- 
vealed a single adult male specimen of an 
undescribed gynodiastyhd. The specimen 
was damaged (third maxillipeds and the 
four distal articles of the first pereopods 
missing), but can be reliably assigned to the 
genus Gynodiastylis Cahnan, 1911 and rep- 
resents the first record of the family Gyno- 
diastyhdae fi-om the Antarctic region. 

Gynodiastylis jazdzewskii, new species 
Figs. 1-2 

Material examined. — Holotype: adult 
male (USNM 243765); Ross Sea, between 
76°01.5'S-179°49.9'E and 76°01.0'S- 
179°52.3'E; depth 388-399 m; 08 February 
1972; Menzies trawl; RA^ Eltanin, Cruise 
51, Sta. 5761; Coll., Smithsonian Oceano- 
graphic Sorting Center. 

Diagnosis. — GynodiastyHdae. Adult male. 



Carapace with numerous (approximately 
17) fine, irregular, wavy, longitudinal ridg- 
es, many coalescing (especially anteriorly). 
Exopods on pereopods 1-4. Telson Ungui- 
form, distinctly longer than sixth abdominal 
somite, extending to distal % of uropodal 
peduncle, armed with 2 distinct, closely set, 
terminal spine-setae. Uropods relatively 
long, attenuated; peduncle with 4 spine-se- 
tae along iimer distal margin; endopod and 
exopod with 2 articles, endopod (excluding 
terminal setae) slightly longer than exopod, 
exopod with 2 long terminal setae, iimer- 
most about as long as rest of exopod, outer 
seta about % length of inner one. 

Description. — adult male holotype (Fig. 
1), carapace length 0.9 mm, total length 3.2 
mm. Carapace nearly Vs of total length; hav- 
ing numerous (approximately 17) fine, ir- 
regular, wavy, longitudinal ridges (or stria- 
tions), many coalescing anteriorly; ocular 
lobe well-developed, eyes obscure, unpig- 



VOLUME 112, NUMBER 2 



363 




Fig. 1. Gynodiastylis jazdzewskii, n. sp. Lateral view of male holotype. Scale = 0.2 mm. 



merited; pseudorostrum strongly developed, 
acutely pointed, slightly decurved, extend- 
ing well beyond ocular lobe. Antennal 
notch broad, not defined ventrally by sharp 
tooth. Thoracic somites 3-5 as illustrated 
(Fig. 1). Abdomen (Fig. 1) subequal in 
length to carapace; somites of similar 
length. Telson (Fig. 2J) linguiform, distinct- 
ly longer than sixth abdominal (telsonic) 
somite, extending to distal ^4 of uropodal 
peduncle, armed with 2 distinct, closely set, 
terminal spine-setae. First and second an- 
tennae (not dissected), appearing similar to 
those of other described members of genus. 
Mandible (Fig. 2A): with well developed 
incisor and molar. First maxilla (Fig. 2B): 



outer endite with 14 terminal or subtermi- 
nal, stout, spine-setae. Second maxilla (fig. 
2E): characterized by well-developed, stout 
comb setae. Maxilliped 1 (Fig. 2D): bran- 
chial lobe (not illustrated) lacking branchial 
processes; siphon long, twisted distally, ter- 
minating in bent acute tip; endopod with 
well-developed specialized setae and spine- 
setae on inner plate, coxa, and propodus 
(Fig. 2D, enlargements). Second maxilliped 
(Fig. 2C): carpus nearly twice as long as 
propodus; propodus more than twice as 
long as dactyl; dactyl terminating in 3 well 
developed, simple spine-setae. Third max- 
illipeds: lost. Pereopods 1-4: bearing exo- 
pods, exopods decreasing in size posteriorly 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Gynodiastylis jazdzewskii, n. sp. A, left mandible: incisor (on left), all spine teeth, except distal most 
broken off; molar (on right); B, first maxilla; C, second maxilliped (terminal articles); D, first maxilliped (ex- 
cluding damaged branchial lobe) showing enlargement of setal types; E, second maxillia F, second pereopod 
(exopod not shown); G, third pereopod; H, fourth pereopod (exopod not shown); I, fifth pereopod; J, sixth 
abdominal (telsonic) somite, uropods, and telson. Scale a = 0. 1 mm for F, G, H, I; Scale b = 0.3 mm for A, 
B, C and 0.1 mm for J. 



VOLUME 112, NUMBER 2 



365 



(Fig. 1). First pereopod (Fig. 1) damaged, 
only basis present; basis strongly devel- 
oped, extending anteriorly past midlength 
of carapace, nearly as long as entire second 
pereopod. Second pereopod (Fig. 2F), basis 
subequal to combined length of other arti- 
cles; ischium distinguishable; carpus longer 
than merus; propodus subequal in length to 
dactyl. Third pereopod (Fig. 2G), merus 
well developed, subequal in length to basis; 
carpus approximately as long as propodus, 
nearly as wide as long with 3 well devel- 
oped distal setae, 1 extending to tip of dac- 
tyl and 2 extending well beyond dactyl; 
dactyl about as long as propodus. Fourth 
pereopod (Fig. 2H) similar to third, but hav- 
ing more slender carpus. Fifth pereopod 
(Fig. 21), basis relatively short, subequal in 
length to combined lengths of ischium, 
merus, and carpus; merus broad, slightly 
longer than carpus; carpus longer than pro- 
podus; propodus with long, stout, distal seta 
extending well past dactyl; dactyl shorter 
than propodus. Uropods (Fig. 2J): relatively 
long, approximately equal to combined 
length of abdominal somites 5 & 6, atten- 
uated; peduncle with 4 spine- setae along in- 
ner distal margin; inner ramus with 2 arti- 
cles, longer than outer ramus, proximal ar- 
ticle with inner margin bearing 3 spine- se- 
tae, distal article longer than proximal, 
inner margin with 3 spine-setae, 2 terminal 
setae present, outer seta distinctly longer 
than distal article and over 4 times longer 
than inner seta; outer ramus with 2 articles, 
distal article with one relatively short sub- 
terminal seta on inner margin and 2 long 
terminal setae, innermost approximately as 
long as rest of exopod, outer seta about Vi, 
length of inner. 

Etymology. — This species is named in 
honor of Professor Krzysztof Jazdzewski 
(University of Lodz, Poland) in recognition 
of his many significant contributions to 
Antarctic biological research. 

Comparisons. — Gynodiastylis jazdzew- 
skii can be distinguished from the previ- 
ously described species of the genus by the 
fine, irregularly wavy, longitudinal ridges 



on the carapace, the shape and terminal se- 
tation of the telson, the relative length of 
the telson in relation to that of the last ab- 
dominal somite and the uropodal peduncles; 
and by the setation and relative length of 
the uropodal rami. 

Of the known species of Gynodiastylis, 
four (G. costata Caiman, 1911; G. turgida 
Hale, 1928; G. lata Hale, 1946; G. lineata 
Day, 1980) have carapaces with longitudi- 
nal ridges or striations. Of these, G. ja- 
zdzewskii appears to be most closely related 
to G. costata, the only other member of the 
genus having numerous, fine, irregular stri- 
ations on its carapace. Gynodiastylis cos- 
tata was originally described from Gulf of 
Siam at depths ranging from 9 to 37 m. It 
was later reported from "night surface 
plankton" in Japanese waters by Gamo 
(1968). 

Gynodiastylis jazdzewskii is readily dis- 
tinguishable from the comparable males of 
G. costata sensu Caiman (1911) and Gamo 
(1962) by having: (1) a longer and more 
acutely tipped rostrum, (2) a distinctly lon- 
ger telson (approximately Va the length of 
the uropodal peduncle), and (3) two long 
terminal setae on the tip of uropodal exo- 
pod. Based on Caiman's (1911) and Gamo's 
(1962) descriptions and illustrations (plate 
36, figs. 1-10 and fig. 40, respectively) the 
telson of male G. costata is half or less the 
length of the uropodal peduncle. There is 
only a single long inner terminal seta on the 
uropodal exopod of G. costata, the outer 
terminal seta being weakly developed (less 
than y^ the length of the inner). 

The variously modified peduncular arti- 
cles of the first antennae of Allodiastylis 
Hale, 1936 and Sheardia Hale, 1946 distin- 
guish G. jazdzewskii from the species of 
these two genera. The presence of uropodal 
endopods with two articles distinguishes G. 
jazdzewskii from the species of the genus 
Dicoides Hale, 1946, which are character- 
ized by uropodal endopods with three arti- 
cles. The members of the remaining two 
gynodiastylid genera, Halina Day, 1980, 
and Zimmeriana Hale, 1946, all lack the 



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fine, longitudinal, carapace ridges that char- 
acterize G. jazdzewskii. 

Discussion. — In her revision and rein- 
statement of the family Gynodiastylidae, 
Day (1980) recognized six genera, Gyno- 
diastylis, Allodiastylis, Dicoides, Haliana, 
Sheardia, and Zimmeriana. Based on the 
summary works by Day (1980) and Baces- 
cu (1992), records for all previously de- 
scribed gynodiastylids appear to be con- 
fined to the Indo-West Pacific region be- 
tween 40°N (Japan) and 43°S (New Zea- 
land). 

Because present day gynodiastylids have 
predominantly tropical and temperate dis- 
tributions, G. jazdzewskii might represent a 
reUc species whose ancestral stock was 
present in cold Antarctic waters before the 
break up of Pangaea over 180 million yr 
ago. Based on its similarity to other warmer 
water species like G. costata, G. jazdzews- 
kii could have split off from the same stock 
that gave rise to the rich present day gy- 
nodiastylid fauna. Following this reasoning, 
present day gynodiastylids may have 
evolved from a small group of cold water, 
Antarctic progenitors and then underwent 
rapid speciation and dispersal as Australia 
and South Africa slowly drifted northward 
into the warmer latitudes. There, however, 
are some problems with this hypothesis. For 
example, if the origin of gynodiastylids was 
the cold water of the Antarctic Ocean, their 
apparent absence from continental waters of 
South America is puzzling. Recently, how- 
ever, Les Watling (pers. comm) informed us 
that he has collected gynodiastyUds from 
waters off Chile. 

There is the additional possibility that, 
due to their small size and specialized hab- 
itat requirements, the lack of previous rec- 
ords for the family in Antarctic waters also 
may be due to an artifact of sampling. How- 
ever, if populations of gynodiastylids are 
common in the shelf waters of Antarctica, 
at least some additional specimens should 
have been collected in studies conducted in 
the waters adjacent to the Antarctic Penin- 
sula. There has been considerable benthic 



sampling in this region, and the cumaceans 
from the resulting collections have been 
identified (Lowry 1975, Richardson & 
Hedgpeth 1977, Ledoyer 1993, Blazewicz 
& Jazdzewski 1995, R. Heard, personal ob- 
servations). 

A third plausible option is that gynodias- 
tylids originated in the warm temperate wa- 
ters of the Indo-Pacific (Day 1980). If so G. 
jazdzewskii might simply be a pioneer spe- 
cies, representing a relatively recent and 
isolated dispersal event for the family in 
Antarctic waters. 

The occurrence of G. jazdzewskii at a 
depth between 388 and 399 m represents 
the second deepest record for the genus and 
family. A southern African species, G. pro- 
fondus Day, 1980, was reported from 
depths of 68 to 680 m by Day (1980). 

The description of any new species, 
based on a single damaged specimen, is 
tenuous. Notwithstanding, G. jazdzewskii is 
readily distinguishable from all previously 
described gynodiastylids and represents an 
important zoogeographic record for the 
family in Antarctic waters. 

Acknowledgments 

We are grateful to Dr. G. Hendler and 
Ms. B. Landrum for making material and 
station data from the RA^ Eltanin available. 
The senior author thanks Dr. Robert T. van 
AUer (former Director of the Gulf Coast 
Research Laboratory, Ocean Springs, MS) 
and Prof. Krzysztof Jazdzewski (Director of 
the Laboratory of Polar Biology & Ocean- 
obiology. University of Lodz, Poland) for 
their help in arranging financial assistance 
to work in the United States. Ms. Sara 
LeCroy graciously helped with the prepa- 
ration and reading of the manuscript. This 
research was supported in part by the U.S. 
Antarctic Research Program of the National 
Science Foundation, through a contract 
with the Smithsonian Oceanographic Sort- 
ing Center (Grant DPP-8214878). 



VOLUME 112, NUMBER 2 



367 



Literature Cited 

Bacescu, M. 1992. Cumacea II (Fam. Nannastacidae, 
Diastylidae, Pseudocumatidae, Gynodiastylidae, 
et Ceratocumatidae) — Crustaceorum Catalogus, 
Pars 8, H.-E. Gruner and L. B. Holthuis, eds., 
SPD Academic Publishing, The Hague, pp. i- 
iv, 175-468. 

Blazewicz, M., & K. Jazdzewski. 1995. Cumacea 
(Crustacea, Malacostraca) of Admiralty Bay 
(King George Island, South Shetlands): a pre- 
liminary note. — Polish Polar Research 16(1—2): 
71-85. 

Caiman, W. T. 1911. On new or rare Crustacea of the 
order Cumacea from the collection of the Co- 
penhagen Museum. II. The families Nannasta- 
cidae and Diastylidae. — Transactions of the 
Zoological Society of London 18:341-398. 

Day, J. 1980. South African Cumacea. Part 4. Families 
Gynodiastylidae and Diastylidae. — Annals of 
the South African Museum 82(6): 187-292. 

Gamo, S. 1962. On the Cumacean Crustacea from Tan- 
abe Bay, Kii Peninsula. — Publications of the 
Seto Marine biological Laboratory 10(2): 10- 
66. 

. 1968. Studies on the Cumacea (Crustacea, 

Malacostraca) of Japan, part III. — Publications 



of the Seto Marine biological Laboratory 16(3): 
147-192. 

Hale, H. M. 1928. Australian Cumacea. — Transactions 
of the Royal Society of South Australia 52:31- 
52. 

. 1936. Cumacea from a South Australian 

reef. — Records of the South Australian Museum 
5(4): 404-438. 

. 1945. Australian Cumacea. No. 11. The fam- 
ily Diastylidae, part 1. — Transactions of the 
Royal Society of South Australia 69:173-211. 

. 1946. Australian Cumacea. No. 12. The fam- 



ily Diastylidae, part 2 Gynodiastylis and related 
genera. — Records of the South Australian Mu- 
seum 8:357-444. 

Lowry, J. K. 1975. Soft bottom macrobenthic com- 
munity of Arthur Harbor, Antarctica. — Antarc- 
tic Research Series 23:1-19. 

Richardson, M., & J. W. Hedgpeth 1977. Antarctic 
soft-bottom macrobenthic community adapta- 
tions to cold stable, highly productive, glacially 
affected environment, pp. 181-196 In G. A. 
Liano, ed.. Adaptations within Antarctic eco- 
system. — Proceedings SCAR Symposium on 
Antarctic Biology, Smithsonian Institution, 
Washington. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

112(2):368-380. 1999. 

A new genus and species of sphaeromatid isopod (Crustacea) from 

Atlantic Costa Rica 

Regina Wetzer and Niel L. Bruce 

(RW) Department of Biological Sciences, University of South Carolina, 

Columbia, South Carolina 29208 U.S.A.; rwetzer@sc.edu; 

(NLB) 138 Carmody Road, St. Lucia, Queensland 4067, Australia; nbruce@globec.com.au 

Abstract. — Beatricesphaera ruthae, a new genus and species of sphaero- 
matid isopod inhabiting shallow water along the Atlantic Costa Rican coast is 
described. This small species is placed into a new genus as the combination 
of characters which describe it precludes inclusion into any known sphaero- 
matid genus. It is best recognized by these character states: short epistome, 
flattened antenna 1 peduncle articles 1 and 2, robust setose pereopods, lamellar 
uropods, simple pleopods 4 and 5, short and unfused penes and an elongate 
basally attached appendix masculina. It is not readily allied to any genus or 
group of genera within the Sphaeromatidae. 



Costa Rican shores boast every major 
tropical coastal marine habitat, from rocky 
shores and sandy beaches, to mangrove la- 
goons and coral reefs. In a joint project 
between Universidad de Costa Rica and 
the Los Angeles County Natural History 
Museum, the Cahuita Reef (9°18'N, 
82°7'W) at Parque Nacional on the Carib- 
bean coast of Costa Rica was surveyed and 
monitored between April 1986 and August 
1987 for habitat degradation resulting from 
upland deforestation. This national park in- 
cludes a small 240 hectare barrier reef 
(Wellington 1974; Risk et al. 1980; Cortes 
et al. 1984; Guzman & Cortes 1984a, 
1984b; Murillo & Cortes 1984; Cortes & 
Risk 1985). 

Knowledge of the Pacific Costa Rican 
isopod fauna rests with Brusca & Iverson's 
1985 summary. The greater Caribbean re- 
gion is covered by the publications of Kens- 
ley (1984), the field guide of Kensley & 
Schotte (1989) and miscellaneous other 
contributions, notably by Miiller (e.g., 
1993a, 1993b). This new genus and species 
of marine sphaeromatid isopod brings the 
total number of sphaeromatid genera known 
from the Caribbean to 17. 



Bruce (1993, 1995, 1997) discusses some 
of the taxonomic difficulties of the Sphae- 
romatidae (e.g., high levels of homoplasy 
and the certain lack of monophyly of many 
larger genera) when compared to other fla- 
belliferan families. The Sphaeromatidae are 
remarkable for their morphological diver- 
sity and are highly speciose. To date almost 
100 genera have been described (Bruce 
1995), and the latest count reports over 600 
described species (see Kensley & Schotte, 
USNM Isopod List on the WWW at: (URL: 
http://nmnhwww.si.edu/gopher-menus/Iso- 
pods.html)). For the Caribbean, Kensley & 
Schotte (1989) reported 32 species in 13 
genera, and White (1996 unpublished) re- 
ports 7 described sphaeromatids from the 
Cahuita reef and two undescribed species. 

Methods and terminology. — Coral rubble 
and algal samples from the intertidal to 12 m 
were collected and rinsed in fresh water and 
sieved through a 240 jxm mesh screen. All 
material was fixed and preserved in 95% 
ethyl alcohol. Material was examined with 
dissecting, compound, and scarming elec- 
tron microscopes. Appendages were drawn 
with the aid of a camera lucida. Setal and 
spine nomenclature as well as the nomen- 



VOLUME 112, NUMBER 2 



369 



clature of appendage orientation follows 
Brusca et al. 1995. Abbreviations used are: 
LACM — Los Angeles County Museum of 
Natural History, Los Angeles, California; 
MOV — Museum of Victoria, Melbourne, 
Australia; SDNHM — San Diego Natural 
History Museum, San Diego, California; 
UCR — Universidad de Costa Rica, San 
Jose, Costa Rica; USNM — National Muse- 
um of Natural History, Smithsonian Insti- 
tution, Washington, D.C.; PMS, plumose 
marginal setae; SEM, scanning electron mi- 
croscope. 

Systematics 

Order Isopoda Latreille, 1817 

Suborder Flabellifera Sars, 1882 

Family Sphaeromatidae 

Milne Edwards, 1840 

Beatricesphaera, new genus 

Type species. — Beatricesphaera ruthae, 
new species, here designated. 

Diagnosis of male. — Body vaulted, about 
twice as long as wide, widest at pereonites 
5 and 6; body with translucent, flexible 
membrane-like covering; body margins 
with setose fringe, membrana cingula (see 
Buss & Iverson 1981). Cephalon lacking 
rostral point, lateral margins not expanded; 
eyes lateral, distinctly faceted. Pereonites 
2-6 of about equal length, pereonites 1 and 
7 longest; pereonite 7 not extending to lat- 
eral margins, laterally overlapped by per- 
eonite 6; coxae without coxal keys (see 
Bruce 1994a, Figs. 2G, 22D), sutures ab- 
sent. Pleon consisting of 1 segment. Pleo- 
telson anterolateral margins extending to 
full body width, posterior margin narrowed, 
with distinct dorsal foramen, posterior mar- 
gins of which contact posteriorly. 

Antenna 1 peduncle articles 1 and 2 
strongly flattened, anteriorly expanded; ar- 
ticle 2 with anterodistal lobe; anterior mar- 
gins of article 1 and 2 contiguous; article 3 
slender, not expanded; flagellum short, 
slightly longer than article 3. Antenna 2 pe- 
duncle article 1 shortest, article 5 longest. 



articles 2-4 progressively increasing in 
length. 

Epistome short, not laterally narrowed or 
constricted, with blunt median point; not 
separating antennule bases; lateral margins 
narrow, encompassing labrum. Right man- 
dible incisor blade-like, narrow, unicuspid; 
without spine row and lacinia mobilis; left 
mandible incisor similar to right, but with 
lacinia mobilis and spine row of 2 spines; 
both mandibles with robust molar provided 
with proximal marginal teeth, gnathal sur- 
face smooth. Maxilla 1 outer lobe with 10 
spines, outer 5 smooth, inner 5 weakly ser- 
rate, inner lobe with 4 feathered spines and 
2 short simple spines. Maxilla 2 with 3 sub- 
equal lobes, provided with setae. Maxilli- 
ped palp articles 2 and 3 with short distinct 
distomedial lobes, article 4 and 5 quadrate; 
endite flat, distal margin subtruncate, with 
acute and blunt spines. 

Pereopods robust, ambulatory, accessory 
unguis of dactylus simple. Pereopod 2 
markedly slender in comparison to pereo- 
pod 1, pereopods 3-7 more robust than 1 
and 2, posterior margins of pereopods with 
fringe of simple and plumose setae on ven- 
tral margins of merus, ischium, and pro- 
podus. 

Paired short flattened penes present on 
posterior of stemite 7, not extending to ple- 
opod rami. 

Pleopods 1-3 rami with PMS, 4-5 with- 
out PMS. Pleopod 1 rami almost subequal 
in length, neither indurate nor operculate. 
Pleopod 2 endopod about 1.5 times as long 
as exopod; appendix masculina on basal 
lobe, nearly twice (1.8) as long as endopod, 
apically acute. Pleopod 3 endopod without 
suture; pleopods 4 and 5 without ridges or 
folds; pleopod 5 exopod with 3 scale patch- 
es. Uropods lamellar, extending just beyond 
posterior of pleotelson, not meeting at mid- 
line; exopod prominent, about half as long 
as endopod. 

Female. — Mouthparts not metamor- 
phosed; pleotelson posterior margin entire, 
without exit channel; oostegites absent; 



370 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



eggs apparently held within pereon; other- 
wise similar to male. 

Remarks. — This small species of sphae- 
romatid presents a combination of charac- 
ters that prevents its inclusion in any known 
sphaeromatid genus, and furthermore, can- 
not readily be allied to any group of genera 
within the Sphaeromatidae. We therefore 
regarded it as incertae sedis. Superficially it 
is perhaps most similar to the ''Cassidini- 
dea group" discussed by Bruce (1994b), 
but it lacks the large dorsally visible epi- 
stome, fused penes, laterally extended ceph- 
alon, and also has short rather than elongate 
pleopod 1 rami. In common with that 
group, it has an elongate appendix mascu- 
lina which is set on a basal lobe, the mar- 
supium is composed of anterior and poste- 
rior pockets (see Harrison 1984, Kensley & 
Schotte 1989: 205 for terminology), the 
pleon consists of a single segment entirely 
without sutures, and pleopods 4 and 5 are 
similar to those of Cassidinidea (all of these 
characters mentioned might be considered 
possible apomorphies). The significance of 
the shared characters is uncertain as all of 
these states are known to occur in other 
sphaeromatid genera. In common with the 
'' Leptosphaeroma group" (sensu Bruce 
1994b) are the short epistome and short pe- 
nes, but the genera of this group are oth- 
erwise distinct. What is remarkable, if the 
affinities of this new genus do lie with Cas- 
sidinidea and related genera, is that this is 
the first instance of a perforate pleotelson 
in that group of genera. 

There is, as mentioned earlier, a superfi- 
cial resemblance to the genus Cassidinidea 
Hansen, 1905 (reviewed in Bruce 1994b). 
That genus is immediately distinguished by 
the dorsally visible epistome, narrow anten- 
nule peduncle articles, expanded lateral 
margins of the head, and fusion of the pe- 
nial processes (Kensley & Schotte's 1989 
statement that the penes are "elongate, sep- 
arate" is in error, applying only to ''Cas- 
sidinidea'' mosaica Kensley & Schotte, 
1987, a species incorrectly placed in the ge- 
nus). In addition both male and female Cas- 



sidinidea have the pleotelson posterior mar- 
gin entire, whereas it is perforate in the 
male of Beatricesphaera new genus. A 
"gel-layer" occurs in Cassidinella (Bruce 
1994a) as well as other genera, but seems 
easily lost, and is usually not recorded, so 
its significance is entirely uncertain. It was 
not observed in Beatricesphaera. 

Lack of thickened ridges or folds on ple- 
opods 4 and 5 has repeatedly been shown 
to be a character of dubious phylogenetic 
significance (most recently Bruce 1995, 
1997) that can vary within genera, and the 
flattened antennule peduncle article(s) can 
also vary within genera (e.g., Cassidinella, 
Bruce 1994a). Another character state that 
is not widely distributed in the family is the 
marked slendemess and elongation of the 
second leg in comparison to the first. This 
is also known to occur in some genera and 
species of the Ischyromene-gxow^ (Bruce 
1995) as well as some species of Dyname- 
nella (e.g., D. nuevitas Kensley et al. 1997). 
Lamellar uropods, which occur in most 
genera formerly placed in the subfamily 
Cassidininae, are also to be found in other 
genera such as Exosphaeroma for example. 
Without a full phylogenetic analysis of the 
genera with the associated identification of 
informative characters and resolution of the 
evidently high level of homoplasy in the 
family (Bruce 1993, 1995, 1997), problems 
such as the one that this genus presents will 
continue to plague taxonomists working on 
the Sphaeromatidae. 

Of the other sphaeromatid genera known 
from the Caribbean region, none is partic- 
ularly similar, and the genus can best be 
identified by the characters of short epi- 
stome, flattened antenna 1 peduncle articles 
1 and 2, robust setose pereopods, lamellar 
uropods, simple pleopods 4 and 5, short and 
unfused penes and an elongate basally at- 
tached appendix masculina. Females can be 
identified by the antenna 1, epistome and 
pereopod characters. The prominent bosses 
on pereonite 7, while not included as a ge- 
neric character, further serve to identify the 
genus and species. 



VOLUME 112, NUMBER 2 



371 



Etymology. — During her tenure at the 
SDNHM, RW had the great pleasure of 
working with and benefiting from two tre- 
mendously wise women (Beatrice Koep- 
nick and Ruth Kantor), whose efficacy, in- 
sight, and hard-earned wisdom left an in- 
delible impression on all they met. It is the 
energy that these two women gave to so 
many scientific projects that this new genus 
and species honors. The new genus honors 
Bea who passed away April 1995 and un- 
fortunately before this manuscript was com- 
pleted. The generic name is feminine, as the 
second element, sphaera, is a feminine Lat- 
in noun. 

Beatricesphaera ruthae, new species 
Figs. 1-6 

Material examined. — Male holotype 
(LACM 86-196.4), Atlantic, Costa Rica, Li- 
mon Province, Parque Nacional Cahuita, 
Punta Cahuita Reef, 9°44.18'N, 82°48.7'W, 
inner reef flat, brown algae wash, 1-3 m 
depth. 29 Oct 1986. Coll. R. C. Brusca and 
P M. Delaney, LACM 86-196. Ovigerous 
female allotype (LACM 86-202.3) Parque 
Nacional Cahuita, Punta Cahuita Reef, 
9°40.3'N, 82°45'W, outer reef crest, algae 
and algae holdfast washes, 1-8 m depth. 30 
Oct 1986. Coll. R. C. Brusca and P M. De- 
laney, LACM 86-202. Paratypes (2 mature 
males, 2 juvenile males) (LACM 86-196.7), 
Parque Nacional Cahuita, Punta Cahuita 
Reef, 9°44.18'N, 82°48.7'W, inner reef flat, 
brown algae wash, 1-3 m depth. 29 Oct 
1986. Coll. R. C. Brusca and P M. Delaney, 
LACM 86-196. Paratypes (1 male, 1 fe- 
male, 1 juvenile) (LACM 86-202.4), Parque 
Nacional Cahuita, Punta Cahuita Reef, 
9°40.3'N, 82°45'W, outer reef crest, algae 
and algae holdfast washes, 1-8 m depth. 30 
Oct 1986. Coll. R. C. Brusca and P M. De- 
laney, LACM 86-202. Paratypes (2 males, 
3 females) (LACM 87-3.2), Parque Nacion- 
al Cahuita, Punta Vargas, 9°44.18'N, 
82°48.7'W, inner reef lagoon, algae washes, 
1-3 m depth. 8 Jan 1987. Coll. R. C. Brusca 
and R. Wetzer, LACM 87-3. Paratype (1 



male, dissected and figured appendages) 
(LACM 86-122.4), Limon Province, north 
of Puerto Viejo, 9°40.8'N, 82°45.53'W, cor- 
alline algae rock wash, offshore algae ridge, 
1-3 m depth, CRA-86-13. 27 Jul 1986. 
Coll. G. L. Hendler, LACM 86-122. Para- 
types (5 males, 5 females) (USNM 
286886), Parque Nacional Cahuita, Punta 
Vargas, 9°44.18'N, 82°48.7'W, inner reef la- 
goon, algae washes, 1-3 m depth. 8 Jan 
1987. Coll. R. C. Brusca and R. Wetzer, 
LACM 87-3. Paratypes (4 males, 4 fe- 
males) (MOV J45491 through J45498), 
Parque Nacional Cahuita, Punta Cahuita 
Reef, 9°40.3'N, 82°45'W, outer reef crest, 
algae and algae holdfast washes, 1-8 m 
depth. 30 Oct 1986. Coll. R. C. Brusca and 
P M. Delaney, LACM 86-202. Paratypes (2 
males, 3 females) (UCR-2248), Parque Na- 
cional Cahuita, Punta Cahuita Reef, 
9°40.3'N, 82°45'W, outer reef crest, algae 
and algae holdfast washes, 1-8 m depth. 30 
Oct 1986. Coll. R. C. Brusca and P M. De- 
laney, LACM 86-202. 

Other material examined: Atlantic, Costa 
Rica, specimens: Limon Province, Parque 
Nacional Cahuita, Punta Cahuita Reef, 
9°44.35'N, 82°48.7'W, approximately 500 m 
from shore, algae and coral rubble washes, 
3-4 m depth. 3 Apr 1986. Coll. R. C. Brus- 
ca and P M. Delaney, LACM 86-98, 2 
mancas. Parque Nacional Cahuita, Punta 
Cahuita Reef, 9°44.35'N, 82°48.7'W, ap- 
proximately 500 m from shore, brown algae 
washes, 3-4 m depth. 4 Apr 1986. Coll. R. 
C. Brusca, P. M. Delaney, and R. Wetzer, 
LACM 86-100, 9 specimens. Limon Prov- 
ince, north of Puerto Viejo, 9°40.8'N, 
82°45.53'W, coralline algae rock wash, off- 
shore algae ridge, 1-3 m depth, CRA-86- 
13. 27 Jul 1986. Coll. G. L. Hendler, 
LACM 86-122, 24 specimens. Limon Prov- 
ince, north of Puerto Viejo, 9°40.8'N, 
82°45.53'W, outer reef crest, coralline algae 
on Sargassum reef, 6—11 m depth. 27 Jul 
1986. Coll. R. Wetzer, LACM 86-128, 2 
specimens each attached to a walking leg 
of brachyuran crab: Acanthonyx petiverii. 
One specimen attached to left 3rd walking 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



leg; second specimen attached to left 2nd 
walking leg. Limon Province, Parque Na- 
cional Cahuita, Punta Cahuita Reef, 
9°44.18'N, 82°48.7'W, inner reef flat, 
brown algae wash, 1-3 m depth. 29 Oct 

1986. Coll. R. C. Brusca and P M. Delaney, 
LACM 86-196, 40+ specimens. Limon 
Province, Parque Nacional Cahuita, Punta 
Cahuita Reef, 9°40.3'N, 82°45'W, outer reef 
crest, algae and algae holdfast washes, 1-8 m 
depth. 30 Oct 1986. Coll. R. C. Brusca and 
R M. Delaney, LACM 86-202, 64+ speci- 
mens. Limon Province, Parque Nacional 
Cahuita, Punta Vargas, 9°44.18'N, 
82°48.7'W, inner reef lagoon, algae washes, 
1-3 m depth. 8 Jan 1987. Coll. R. C. Brusca 
and R. Wetzer, LACM 87-3, 70+ speci- 
mens. Limon Province, Parque Nacional 
Cahuita, Punta Vargas, 9°44.18'N, 82°48.7'W, 
inner reef lagoon, coral rubble washes, 1- 
12 m depth. 8 Jan 1987. Coll. R. C. Brusca 
and R. Wetzer, LACM 87-4, 2 females. Li- 
mon Province, Parque Nacional Cahuita, 
Punta Vargas, 9°44.18'N, 82°48.7'W, inner 
reef lagoon, rock, coral rubble, and algae 
washes, 1-13 m depth. 8 Jan 1987. Coll. R. 
C. Brusca and R. Wetzer, LACM 87-5, 2 
mancas. Limon Province, Parque Nacional 
Cahuita, Punta Vargas, 9°44.5'N, 82°48.37'W, 
inner reef lagoon, skimming Thallassia bed 
off point, intertidal to 1 m depth. 8 Jan 

1987. Coll. R. C. Brusca and R. Wetzer, 
LACM 87-7, 1 male. Limon Province, 
Parque Nacional Cahuita, Punta Vargas, 
9°44.18'N, 82°48.7'W, inner reef lagoon, 
broken coral and rubble washes, 1-3 m 
depth. 8 Jan 1987. Coll. R. C. Brusca and 
R. Wetzer. LACM 87-8, 1 molt. Limon 
Province, Puerto Viejo, 9°40.3'N, 82°45'W, 
forereef crest and along sand bottom junc- 
tion, algae and gorgonian washes, 1-6 m 
depth. 9 Jan 1987. Coll. R. C. Brusca and 
R. Wetzer, LACM 87-9, 34 specimens. Li- 
mon Province, Parque Nacional Cahuita, 
Punta Cahuita Reef, 9°44.37'N, 82°48.7'W, 
rock, coral, and algae holdfast washes, 0.9- 
3 m depth. 19 Aug 1987. Coll. R. C. Brusca 
and R. Wetzer, LACM 87-158, 21 speci- 
mens. Limon Province, Parque Nacional 



Cahuita, Punta Cahuita Reef, 9°44.52'N, 
82°48.37'W, rock, coral, and algae holdfast 
washes, 0.5-10 m depth. 20 Aug 1987. 
Coll. R. C. Brusca and R. Wetzer, LACM 
87-165, 64 specimens. 

Description. — Adult male: Body strongly 
vaulted, limpet-like, about 2.3 times as long 
as wide (Figs. lA, B, 2A). Antennular pe- 
duncle articles, lateral body margins, uro- 
pods, and pleotelson with conspicuous setal 
fringe, membrana cingula (Figs. lA, B, C, 
3A). Frontal margin of cephalon without 
rostral process (Fig. lA). Dorsum of per- 
eonite 7 elevated into large dome-like con- 
ical tubercle, with lateral notches on pos- 
terior margin. Single free pleonite also with 
tubercle, somewhat smaller than tubercle on 
pereonite 7; with lateral projections on pos- 
terior margin. Pleotelson acute; pleotelsonic 
foramen dorsally directed, triangular, entire- 
ly enclosed (Figs. lA, B, 2B). A thin mem- 
brane visible with SEM overlie the dorsal 
cuticle of entire body including antennular 
peduncle articles and appears finely granu- 
lar with small, numerous, evenly spaced 
pits; each pit with 1 to 3 simple or palmate 
setae (Fig. 2A, B, C). Coxal sutures indis- 
tinct dorsally and ventrally. Eyes round, 
well developed, slightly elevated, with fac- 
ets and pigmentation, deeply immersed in 
pereonite 1. Pereonite 1 longest, pereonites 
2-6 subequal, pereonite 7 longer than pre- 
ceding pereonites. First pleonite subequal in 
length to pereonites 2-6 (Figs. lA, B, 2A). 

Antennae 1 short, extending to posterior 
margin of pereonite 2 (Fig. 3A). Peduncular 
articles 1-2 shovel- shaped, dorsoventrally 
compressed and greatly expanded. Article 1 
rectangular, almost twice as long as wide. 
Last (third) peduncular article narrow, not 
expanded; about twice as long as wide. Fla- 
gellum 6-articulate, article 1 short, article 2 
longer, article 3 longest, terminal articles 
gradually tapering distally. One long, "ar- 
ticulated" aesthetasc on lateral, distal fla- 
gellar articles 2, 3, and 4 (Fig. 3 A, B). 

Antennae 2 extending to anterior margin 
of pereonite 3 ; usually not visible in dorsal 
aspect. Peduncle composed of 4 articles, 1 



VOLUME 112, NUMBER 2 



373 




Fig. 1. Beatricesphaera ruthae, new species. A, dorsal view, adult male, holotype LACM 86-196; B, lateral 
view, adult male, holotype LACM 86-196; C, uropod (right), adult male, paratype LACM 86-122. 



and 2 subequal in length, article 3 1.6 times 
as long as 2 and somewhat expanded, arti- 
cle 4 slender and about twice as long as 
article 1. Simple and palmate setae present 
on peduncular articles as figured. Flagellum 
8-articulate, each article with 2-6 simple 
setae on margin (Fig. 3C). 

Frontal lamina wide, crescent shaped, en- 
compassing clypeus and part of labrum. La- 
brum with dense short simple setae on me- 
dial margin, setae become less dense and 
longer on lateral margins (Figs. 3D, 4E, F). 
Clypeus and labrum fused, together form- 
ing a subovate structure. Mandibular inci- 
sors distally narrow, flattened, unicuspid; 
lacinia mobilis composed of 2 large bifid 



spines; spine row composed of 2 comb 
spines; molar process with proximal mar- 
ginal teeth; gnathal surface almost smooth; 
palp 3-articulate, article 2 with 3 stout bi- 
serrate setae, article 3 with 6 biserrate ro- 
bust setae (Figs. 3E, 4A, B, C). Maxilla 1 
outer lobe nearly twice as wide as inner 
lobe, with 5 flattened, smooth robust spines 
laterally, 4-5 large, serrate spines medially; 
inner lobe with 4 slender, plumose robust 
setae (Fig. 3F). Maxilla 2 with 3 subequal 
lobes; outer lobe with 4 long robust setae; 
middle lobe with 4 long robust setae; inner 
lobe with simple, plumose and comb setae, 
as figured; numerous short, simple setae 
dispersed along medial margin; endite basis 



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Fig. 2. Beatricesphaera ruthae, new species. Scanning electron micrographs. A, adult male LACM 86-122, 
lateral view; B, adult male LACM 86-122, pleotelson; C, male LACM 87-7 palmate setae on pleotelson, note 
torn "cuticle;" i.e., membrana cingula, scale bar = 20 fxm; D, gravid female LACM 86-122 ventral brood pouch; 
E, adult male LACM 86-7 penes, scale bar = 200 |xm. 



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375 




Fig. 3. Beatricesphaera ruthae, new species. Adult male paratype LACM 86-122. A, antenna 1 (right); B. 
flagellular articles antenna 1 (right); C, antenna 2 (right); D, frontal lamina, clypeus, labrum; E, mandible (left); 
F, maxilla 1 (right); G, maxilla 2 (right); H, maxilliped (right). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

4 




Fig. 4. Beatricesphaera ruthae, new species. Scanning electron micrographs. A, male LACM 87-7, mouth- 
field with right and left molar processes, left incisor, lacinia mobilis, and spine row, scale bar =100 |xm; B, 
same as A, scale bar = 50 |xm; C, male LACM 86-122, mouthfield, frontal view, scale bar =100 (xm; D, male 
LACM 87-7, palmate setae on pleotelson, membrana cingula torn, discs are diatom symbionts, scale bar = 20 
|xm; E, male LACM 87-7, peduncular articles of antennules, frontal lamina, scale bar = 200 [xm; F, male LACM 
87-3, mouthfield with clypeus, labrum, left and right maxillipeds, scale bar = 200 ixm. 



VOLUME 112, NUMBER