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Full text of "Proceedings of the Biological Society of Washington"

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PROCEEDINGS 

of the 

Biological Society of 
Washington 



VOLUME 95 
1982 



Vol. 95(1) published 13 April 1982 Vol. 95(2) published 1 1 August 1982 

Vol. 95(3) pubhshed 5 October 1982 Vol. 95(4) published 20 December 1982 



WASHINGTON 
PRINTED FOR THE SOCIETY 



EDITOR 
Brian Kensley 



ASSOCIATE EDITORS 

Classical Languages Invertebrates 

George C. Steyskal Thomas E. Bowman 

Plants Vertebrates 

David B. Lellinger Richard Banks 

Insects 
Robert D. Gordon 



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 1981-1982 



OFFICERS 

President 
RAYMOND B. MANNING 

Vice President 
PAUL J. SPANGLER 

Secretary 
MICHAEL A. BOGAN 

Treasurer 
LESLIE W. KNAPP 



COUNCIL 

Elected Members 
FREDERICK M. BAYER KRISTIAN FAUCHALD 

ISABEL C. CANET DAVID L. PAWSON 

AUSTIN B. WILLIAMS 



TABLE OF CONTENTS 



Volume 95 

Adis, Joachim, and Richard C. Froeschner. Notes on distribution of some Latin Amer- 
ican cotton-stainers {Dysdercus: Pyrrhocoridae: Hemiptera) and remarks on the biol- 
ogy of Dysdercus urbahni Schmidt 371-376 

Adkison, Daniel L. Description of Dactylokepon sulcipes n. sp. (Crustacea: Isopoda: 

Bopyridae) and notes on D. caribaeus 702-708 

Adkison, Daniel L., Richard W. Heard, and Guy T. Clark. Description of the male and 

notes on the female Argeiopsis inhacae (Crustacea: Isopoda: Bopyridae) 334-337 

Baker, H. R. A note on the genitalia of Potamothrix hammoniensis (Oligochaeta: Tubi- 
ficidae) 563-566 

Barnard, J. Laurens, and Margaret M. Drummond. Redescription of Exoediceros fossor 
(Stimpson, 1856) an Australian marine fossorial amphipod, the type-genus of the new 
family Exoedicerotidae 610-620 

Barnard, J. Laurens, and Gordon S. Karaman. Classificatory revisions in gammaridean 
Amphipoda (Crustacea), Part 2. 167-187 

Bayer, Frederick M. Some new and old species of the primnoid genus Callogorgia 
Gray, with a revalidation of the related genus Fanellia Gray (Coelenterata: An- 
thozoa) 116-160 

Bowman, Thomas E., and Richard Franz. Anopsilana crenata, a new troglobitic ciro- 

lanid isopod from Grand Cayman Island, Caribbean Sea 522-529 

Brooks, Daniel R., and Janine N. Caira. Atrophecaecum lobacetabulare, n. sp. (Di- 
genea: Cryptogonimidae: Acanthostominae) with discussion of the generic status of 
Paracanthostomum Fischthal and Kuntz, 1965, and Ateuchocephala Coil and Kuntz, 
1960 223-231 

Brown, Walter C, and Angel C. Alcala. A new cave Platymantis (Amphibia: Ranidae) 

from the Philippine Islands 386-391 

Bruce, Niel L., and Thomas E. Bowman. The status of Cirolana parva Hansen, 1890 

(Crustacea: Isopoda: Cirolanidae) with notes on its distribution 325-333 

Chernoff, Barry, and Robert Rush Miller. Mexican freshwater silversides (Pisces:Ath- 

erinidae) of the genus Archomenidia, with the description of a new species 428^39 

Child, C. Allan. Pycnogonida of the western Pacific islands I. The Marshall 
Islands 270-281 

Child, C. Allan, and Koichiro Nakamura. A gynandromorph of the Japanese pycnog- 

onid Anoplodactylus gestiens (Ortmann) 292-296 

Collette, Bruce B. South American freshwater needlefishes of the genus Potamorrha- 
phis (Beloniformes: Belonidae) 714-747 

Cressey, Roger F. A new genus of bomolochid copepods from Indo-West Pacific nem- 
ipterid fishes 495-504 

Crews, Celinda R., and Artie L. Metcalf. A new species of oreohelicid land snail from 

the San Agustin plains, New Mexico 256-264 

Cutler, Edward B., and Norma J. Cutler. A revision of the genus Siphonosoma 

(Sipuncula) 748-762 

Dawson, C. E. Review of the genus Micrognathus Duncker (Pisces: Syngnathidae), 

with description of M. natans n. sp. 657-687 

Dawson, C. E., and C. J. M. Glover. Hypselognathus horridus, a new species of 

pipefish (Syngnathidae) from South Australia — 403-407 

Desbruyeres, Daniel, and Lucien Laubier. Paralvinella grasslei, new genus, new species 
of Alvinellinae (Polychaeta: Ampharetidae) from the Galapagos Rift geothermal 
vents • 484-494 

Downey, Maureen E. Evoplosoma virgo, a new goniasterid starfish (Echinodermata: 

Asteroidea) from the Gulf of Mexico 111-111 

Ewing, R. Michael. A partial revision of the genus Notomastus (Polychaeta: Capitel- 

lidae) with a description of a new species from the Gulf of Mexico 232-237 

iv 



Fauchald, Kristian. Two new species of Onuphis (Onuphidae: Polychaeta) from Uru- 
guay 203-209 

Fauchald, Kristian. Some species of Onuphis (Polychaeta: Onuphidae) from the Atlan- 
tic Ocean 238-250 

Fauchald, Kristian. A eunicid polychaete from a white smoker 781-787 

Fauchald, Kristian. Description of Mooreonuphisjonesi, a new species of onuphid poly- 
chaete from shallow water in Bermuda, with comments on variability and population 
ecology 807-825 

Flint, Oliver S., Jr., and Joaquin Bueno-Soria. Studies on Neotropical caddisflies, XXXII: 
the immature stages of Macronema variipenne Flint & Bueno, with the division of 
Macronema by the resurrection of Macrostemum (Trichoptera: Hydropsychi- 
dae) 358-370 

Formas, J. R., and Alberto Veloso. Taxonomy of Bufo venustus Philippi, 1899 (Anura: 
Leptodactylidae) from central Chile 688-693 

Formas, J. R., and M. Ines Vera. The status of two Chilean frogs of the genus Eup- 
sophus (Anura: LeptodactyHdae) 594-601 

Fredette, Thomas J. Evidence of ontogenetic setal changes in Heteromastus filiformis 
(Polychaeta: Capitellidae) 194-197 

Garcia, Renato G., and Raymond B. Manning. Four new species of stomatopod crus- 
taceans from the Philippines 537-544 

Garrido, Orlando H., and Albert Schwartz. A new species of Sphaerodactylus (Reptiha: 

Sauria: Gekkonidae) from eastern Cuba 392-397 

George, Robert Y., and Noel A. Hinton. A new species of deep-sea isopod, Storthyn- 
gura myriamae, from the Walvis Ridge off South Africa 93-98 

Gleye, Linda G. Two new species of leptomysinid mysids (Crustacea: Mysidacea) from 
Southern California 3 19-324 

Green, Karen D. Uncispionidae, a new polychaete family (Annelida) 530-536 

Harasewych, M. G. Pterynotus xenos, a new species of muricid from off northern 
Jamaica (Mollusca: Gastropoda) 639-641 

Harding, Keith A. Courtship display in a Bornean frog 621-624 

Heaney, Lawrence R., and Gary S. Morgan. A new species of gymnure, Podogymnum 

(Mammalia: Erinaceidae) from Dinagat Island, Philippines 13-26 

Hershkovitz, Philip. Subspecies and geographic distribution of Black-mantle Tamarins 

Saguinus nigricollis Spix (Primates: Callitrichidae) 647-656 

Heyer, W. Ronald. Two new species of the frog genus Hylodes from Caparao, Minas 

Gerais, Brazil (Amphibia: Leptodactylidae) 377-385 

Heyer, W. Ronald, Charles H. Daugherty, and Linda R. Maxson. Systematic resolution 

of the genera of the Crinia complex (Amphibia: Anura: Myobatrachidae) 423-427 

Hobbs, Horton H., Jr., and Daniel J. Peters. The entocytherid ostracod fauna of north- 
ern Georgia 297-3 18 

Hobbs, Horton H., Jr., and Henry W. Robison. A new crayfish of the genus Procam- 
barus from southwestern Arkansas 545-553 

Holt, Perry C. A new species of the genus Cambarincola (Clitellata: Branchiobdellidae) 
from Illinois with remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the 
status of Sathodrilus Holt, 1968 251-255 

Huddleston, Richard W. Comments on the nomenclatural status of the families Cau- 

casellidae and Favusellidae (Foraminiferida) 637-638 

Huddleston, Richard W., and Drew Haman. Nomenclatural status of the foraminiferal 

genus Cubanella Saidova, 1981 114-115 

Huddleston, Richard W., and Drew Haman. Jascottella, nom. nov. for Mamilla Scott, 

1974 (Microproblematica) non Fabricius, 1823 (Mollusca) 421 

Hutchings, P. A., and C. J. Glasby. Two new species of Ceratonereis (Polychaeta: 

Nereididae) from estuarine areas of New South Wales, Australia 515-521 

Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XIII. Phagocata 
hamptonae, new species, from Nevada 161-166 

Kenk, Roman, and Anne M. Hampton. Freshwater triclads (Turbellaria) of North 
America. XIV. Poly cells monticola, new species, from the Sierra Nevada range in 
California 567-570 



V 



Kensley, Brian, and Gary C. B. Poore. Anthurids from the Houtman Abrolhos Islands, 

Western Australia (Crustacea: Isopoda: Anthuridae) 625-636 

Kornicker, Louis S. Alternochelata lizardensis, a new species of myodocopine ostra- 

code from the Great Barrier Reef of Australia (Rutidermatidae) 793-806 

Kudenov, Jerry D. Redescription of the major spines of Polydora ligni Webster 
(Polychaeta: Spionidae) 571-574 

Kudenov, Jerry D., and John H. Dorsey. Astreptosyllis acrassiseta, a new genus and 

species of the subfamily Eusyllinae (Polychaeta: Sylhdae) from AustraUa 575-578 

Kyte, Michael A. Ophiacantha abyssa, new species, and Ophiophthalmus displasia 
(Clark), a suggested new combination in the ophiuroid family Ophiacanthidae (Echi- 
nodermata: Ophiuroidea) from off Oregon, U.S.A. 505-508 

Lewis, Julian J. A diagnosis of the Hobbsi group, with descriptions of Caecidotea 
teresae, n. sp., and C macropropoda Chase and Blair (Crustacea: Isopoda: Aselli- 
dae) 338-346 

Louton, Jerry A. A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from 

the western Highland Rim in Tennessee 198-202 

Lynch, John D., and Pedro M. Ruiz-Carranza. A new genus and species of poison-dart 
frog (Amphibia: Dendrobatidae) from the Andes of northern Colombia 557-562 

Manning, Raymond B., and Ch. Lewinsohn. Rissoides, ^. new genus of stomatopod 

crustacean from the east Atlantic and South Africa 352-353 

Manning, Raymond B., and Marjorie L. Reaka. Gonodactylus insularis, a new sto- 
matopod crustacean from Enewetak Atoll, Pacific Ocean 347-351 

Marshall, Harold G. Phytoplankton distribution along the eastern coast of the USA. 
IV. Shelf waters between Cape Lookout, North CaroUna, and Cape Canaveral, 
Florida 99-113 

Martin, Joel W., and Lawrence G. Abele. Naushonia panamensis , new species (De- 
capoda: Thalassinidea: Laomediidae) from the Pacific coast of Panama, with notes on 
the genus 478-483 

McEachran, John D., and Janice D. Fechhelm. A new species of skate from Western 
Australia with comments on the status of Pavoraja Whitley, 1939 (Chondrichthyes: 
Rajiformes 1-12 

McEachran, John D. , and Janice D. Fechhelm. A new species of skate from the western 
Indian Ocean, with comments on the status of Raja (Okamejei) (Elasmobranchii: 
Rajiformes) 440-450 

McKaye, Kenneth R., and Catherine MacKenzie. Cyrtocara liemi, a previously un- 
described paedophagous cichlid fish (Teleostei: Cichhdae) from Lake Malawi, 
Africa 398-402 

McKenzie, K. G. Description of a new cypridopsine genus (Crustacea: Ostracoda) from 

Campbell Island, with a key to the Cypridopsinae 766-771 

Mendez, G. Matilde, and Mary K. Wicksten. Notalpheus imarpe: a new genus and 

species of snapping shrimp from western South America (Decapoda: Alpheidae) 709-713 

Murdy, Edward O., and John D. McEachran. Istigobius hoesi, a new gobiid fish from 
Australia (Perciformes: Gobiidae) 642-646 

Nakamura, Izumi. Lateral line of Diplospinus multistriatus (Teleostei: GempyH- 
dae) 408-411 

Nakamura, Koichiro, and C. Allan Child. Three new species of Pycnogonida from Sa- 

gami Bay, Japan 282-291 

Opell, Brent D. A new Uloborus Latreille species from Argentina (Arachnida: Araneae: 

Uloboridae) . 554-556 

Parenti, Lynne R. Relationships of the African killifish genus Foerschichthys (Teleostei: 

Cyprinodontiformes: Aplocheilidae) 45 1-457 

Petuch, Edward J. Paraprovincialism: remnants of paleoprovincial boundaries in Recent 

marine moUuscan provinces 774-780 

Rausch, V. R., and R. L. Rausch. The karyotype of the Eurasian flying squirrel, Ptero- 
mys volans (L.), with a consideration of karyotypic and other distinctions in Glaucomys 
spp. (Rodentia: Sciuridae) 58-66 

Rohr, David M., and Richard W. Huddleston. Yochelsoniella, nom. nov., a new name 

for Ellisella Rohr, 1980 (Gastropoda) non Gray, 1858 (Coelenterata) 269 

vi 



Rosewater, Joseph. A new species of the genus Echininus (Mollusca: Littorinidae: 

Echinininae) with a review of the subfamily 67-80 

Schultz, George A. Amerigoniscus malheurensis, new species, from a cave in western 

Orgeon (Crustacea: Isopoda: Trichoniscidae) 89-92 

Shelly, Roland M. A new xystodesmid milliped genus and three new species from the 

eastern Blue Ridge Mountains of North CaroHna (Polydesmida) 458-477 

Stauffer, Jay R., Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jen- 
kins. Checklist of the fishes of the central and northern Applachian Mountains 27-47 

Sterrer, Wolfgang, and Thomas M. Iliffe. Mesonerilla prospera, a new archiannelid 

from marine caves in Bermuda 509-514 

Thomas, Richard. A new dwarf Sphaerodactylus from Haiti (Lacertilia: Gekkoni- 

dae) 81-88 

Thompson, Fred G. A new species of Euglandina from Peru (Gastropoda: Pulmonata: 

Spiraxidae) 763-765 

Thompson, Fred G., and Jane E. Deisler. A new tree snail, genus Drymaeus (Buli- 

mulidae) from southeastern Peru 265-268 

Uebelacker, Joan M. Review of some little known species of syllids (Annehda: Poly- 

chaeta) described from the Gulf of Mexico and Caribbean by Hermann Augener in 

1924 583-593 

Vari, Richard P. Hemiodopsis ocellata, a new hemiodontid characoid fish (Pisces: Char- 

acoidea) from western Surinam 188-193 

Vari, Richard P. Curimatopsis myersi, a new curimatid characiform fish (Pisces: Char- 

aciformes) from Paraguay 788-792 

Vecchione, Michael. Morphology and development of planktonic Lolliguncula brevis 

(Cephalopoda: Myopsida) 602-609 

Wainright, Sam C, and Thomas H. Perkins. Gymnodorvillea floridana, a new genus 

and species of Dorvilleidae (Polychaeta) from southeastern Florida 694-701 

Wicksten, Mary K. New records of pinnotherid crabs from the Gulf of California 

(Brachyura: Pinnotheridae) 354-357 

Wicksten, Mary K. Pinnixa costaricana, a new species of crab from Central America 

(Brachyura: Pinnotheridae) 579-582 

Zibrowius, Helmut, and Stephen D. Cairns. Remarks on the stylasterine fauna of the 

West Indies, with a description of Stylaster antillarum, a new species from the Lesser 

Antilles (Cnidaria: Hydrozoa: Stylasterina) 210-221 

Zottoli, Robert. Two new genera of deep-sea polychaete worms of the family Amphar- 

etidae and the role of one species in deep-sea ecosystems 48-57 

Zusi, Richard L., and Gregory Dean Bentz. Variation in a muscle in hummingbirds and 

swifts and its systematic implications 412-420 



Vll 



INDEX TO NEW TAXA 



Volume 95 

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

COELENTERATA 

Hydrozoa 

Stylaster antillarum 213 

Anthozoa 

Callogorgia chariessa 128 

Fanellia corymbosa 154 

PLATYHELMINTHES 

Trematoda 

Atrophecaecum lobacetabulare 223 

Turbellaria 

Phagocata hamptonae 161 

Polycelis monticola 567 

ANNELIDA 

Archiannelida 

Mesonerilla /jroj'perfl 509 

Branchiobdellida 

Cambarincola illinoisensis 25 1 

Polychaeta 

Astreptosyllis 575 

acrassiseta 576 

Ceratonereis limnetica 515 

turveyi 519 

Decemunciger 49 

apalea 49 

Endecamera 50 

palea 52 

Eunice pulvinopalpata 78 1 

Gymnodorvillea 695 

floridana 695 

Haplosyllis floridana n.c. 584 

Moor tonwphis j one si 807 

Notomastus daueri 234 

Onuphis (Nothria) australatlantica 238 

heterodentata 241 

lithobiformis 243 

(Onuphis) declivorum 245 

difficilis 203 

orensanzi 205 

texana 247 

Paralvinella 485 

grasslei 485 

UNCISPIONIDAE 530 

via 



Uncispio 534 

hartmanae 534 

ARTHROPODA 

Crustacea 

Alternochelata lizardensis 793 

Amerigoniscus malheurensis 89 

Anopsilana crenata 522 

Apanthura zeewykae 625 

Aurohornellia 171 

Austrocypridopsis 766 

tenilli 768 

Caecidotea teresae 339 

Ciorida nazasaensis 538 

Cubanomysis mysteriosa 319 

Dactylokepon sulcipes 703 

Eisothistos bataviae 628 

EXOEDICEROTIDAE 610 

Feriharpinia 182 

Gonodactylus insulans 347 

Y{dipiosqm\\2i philippinensis 547 

Holocolax 495 

longisetus 495 

russelli 502 

Holsingerius 180 

Indocalliope 182 

Levisquilla armata 540 

Lupimaera 174 

lupana n.c. 174 

Maleriopa 176 

Mysidopsis cathengelae 321 

Nasageneia 170 

N2iUihom2L panamensis 478 

Notalpheus imarpe 709 

Oratosquilla microps 542 

Panathura haddae 634 

PARACALLIOPIIDAE 181 

Paramoera stephenseni n.c. 170 

Pinnixa costaricana 579 

Procambarus (Girardiella) parasimulans 545 

Relictomoera 168 

Rissoides 352 

Sternomoera 169 

Storthyngura myriamae 93 

Tegano 176 

Texiweckeliopsis 179 

Torridoharpinia 184 

Insecta 

Ophiogomphus bouchardi 198 

Myriapoda 

Prionogonus 460 

divaricatus 467 

haerens 463 

thrinax 470 

ix 



Pycnogonida 

Ammothella stauromata 271 

Anoplodactylus marshallensis 274 

perforatus 289 

shimodaensis 285 

Ascorhynchus utinomii 283 

Nymphon micronesicum 277 

Arachnida 

Uloborus elongatus 554 

MOLLUSCA 

Drymaeus aurantiostomus 265 

Echininus viviparus 69 

Euglandina haasi 763 

Oreohelix litoralis 256 

Pterynotus (Pterynotus) xenos 639 

Yochelsoniella 269 

ECHINODERMATA 

Evoplosoma virgo 772 

Ophiacantha abyssa 505 

Ophiophthalmus diplasia n.c. 508 

CHORDATA 

Pisces 

Archomenidia marvelae 430 

Curimatopsis inyersi 788 

Cyrotocara liemi 398 

Hemiodopsis ocellata 188 

Hypselognathus horridus 403 

Istigobius hoesi 643 

Micrognathus natans 682 

Pavoraja alleni 8 

Raja (Okamejei) heemstrai 441 

Amphibia 

Atopophrynus 557 

syntomopus 557 

Hylodes babax 380 

vanzolinii 382 

Platymantis spelaeus 386 

Reptilia 

Sphaerodactylus celicara 392 

omoglaux 81 

Mammalia 

Podogymnura aureospinula ^ 14 

Saguinus nigricollis hernandezi 649 

MICROPROBLEMATICA 

Jascottella 421 



(ISSN 0006-324X) 



Proceedings 

of the 

BIOLOQICAL SOCIETY 

of 

WASHINQTON 



/ 



-^^ONMA/ 



utim^"^: 



""-.'i^ 



Volume 95 13 April 1982 Number 1 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

1981-1982 

Officers 



President: Raymond B. Manning 
Vice President: Paul J. Spangler 



Frederick M. Bayer 
Kristian Fauchald 



Elected Council 



Austin B. Williams 



Secretary: Michael A. Bogan 
Treasurer: Leslie W. Knapp 

Isabel C. Canet 
David L. Pawson 



Custodian of Publications: Michael J. Sweeney 



PROCEEDINGS 



Editor: Brian Kensley 



Co-editor: Stephen D, Cairns 



Associate Editors 

Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman 

Plants: David B. Lellinger Vertebrates: Richard Banks 

Insects: Robert D. Gordon 



Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 
Annual dues of $10.00 include subscription to the Proceedings of the Biological Society of 
Washington. Library subscriptions to the Proceedings are: $18.00 within the U.S.A., $23.00 
elsewhere. 

The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. 
Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued 
sporadically) are available. Correspondence dealing with membership and subscriptions should 
be sent to The Treasurer, Biological Society of Washington, National Museum of Natural 
History, Smithsonian Instutution, Washington, D.C. 20560. 

Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological 
Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- 
ington, D.C. 20560. 

Known office of publication: National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560. 

Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 
Second class postage paid at Washington, D.C, and additional mailing office. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 1-12 



A NEW SPECIES OF SKATE FROM WESTERN 

AUSTRALIA WITH COMMENTS ON THE 

STATUS OF PAVORAJA WHITLEY, 1939 

(CHONDRICHTHYES: RAJIFORMES) 

John D. McEachran and Janice D. Fechhelm 



Abstract. — The genus Pavoraja is resurrected for the Australian skates Raja 
nitida and Pavoraja alleni, n. sp., which differ from all recognized genera of 
skates in clasper and rostral structure, and by their combination of derived char- 
acter states. Pavoraja alleni is described and diagnostic characters are given for 
Pavoraja and P. nitida. 



Raja nitida, which occurs off Tasmania, Victoria, and southern New South 
Wales (Whitley 1940), was placed in a new genus Pavoraja, along with another 
eastern Australian skate, Raja polyommata Ogilby, 1916 by Whitley (1939). Whit- 
ley (1939) only briefly described Pavoraja and gave no characteristics that dis- 
tinguish it from other skate genera. The following year Whitley (1940) offered 
little to augment his description, but stated that P. nitida resembles the South 
American skate, Malacorhina scobina {=Psammobatis scobina Philippi, 1857), 
that the type locality of P. nitida may be South America rather than Australia, 
and that the Australian specimen of P. nitida may represent a new species or 
subspecies. More recent authors (Fowler 1941, Bigelow and Schroeder 1953) 
considered Pavoraja synonymous with the genus Raja, because of Whitley's lack 
of a generic diagnosis. 

During investigations of the skate fauna of Western Australia a new species 
was discovered which closely resembles Raja nitida Giinther, 1880. Herein we 
describe the taxonomically important anatomical characters of R. nitida, com- 
ment on the taxonomic status oi Pavoraja, describe the new species, and then 
comment on the relationship of/?, polyommata to the other species. 

Materials and Methods 

Specimens of Raja nitida, R. polyommata, and the new species were obtained 
from the Australian Museum, Sydney (AMS); British Museum (Natural History) 
(BMNH); Tasmanian Museum, Hobart (TMH); and Western Australian Museum, 
Perth (WAM). Several individuals of R. nitida and one of the three specimens of 
the undescribed species were dissected to reveal the structure of the claspers, 
neurocrania, and scapulocoracoids. Several additional specimens of/?, nitida and 
all three specimens of the new species were radiographed to verify the anatomical 
observations based on dissections and to count vertebrae, pectoral radials, and 
pelvic radials. Methods for making measurements and counts are described by 
McEachran and Compagno (1979, 1982). 



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a b 

Fig. I. Lateral view of right clasper, partially expanded to show components: a, R. nitida, AMS 
IB. 5275; b, Pavoraja alleni, WAM P191 18 (Holotype). of — cleft, hy — hypopyle, pr — pseudorhipidion, 
rh — rhipidion, si — slit, sp — spike, sr — spur, st — sentinel. 



Results 

Clasper s. — Raja nitida has very slender, short claspers which are constricted 
rather than expanded at origin of glans (Fig. la); without dermal denticles or 
pseudosiphon; inner dorsal lobe with pseudorhipidion extending from level of 
hypopyle to about distal one-third of glans, continuing distally as a fold of integ- 
ument; slit located lateral to pseudorhipidion; spur well developed; cleft medial 
to spur; rhipidion well developed, running from level of hypopyle to distal one- 
third of glans, distal section rotated laterally and running over base of sentinel; 
sentinel rod-shaped and covered with integument, extending from level of slit to 
near tip of glans; spike ventral to sentinel, located within sentina, disc-shaped 
with a sharp, naked lateral margin; axial cartilage forming a slender tip (Fig. 2a, 
b, c); dorsal marginal little expanded distally, distal margin truncate, with an 
inner distal extension entering glans and forming pseudorhidion; ventral marginal 
with a evenly convex distal margin; dorsal terminal 1 and ventral terminal mem- 
branous, broadly joined on ventral aspect of glans, forming a sheath-like covering 
of glans; dorsal terminal 2 broad, fused to distal and distolateral surface of dorsal 
marginal; dorsal terminal 3 fused with dorsal terminal 2, small with a distally 
pointed and laterally curved extension forming spur; ventral terminal V-shaped, 
lacking a sharp lateral margin, lying on dorsal surface of accessory terminal 1; 
accessory terminal 1 Y-shaped, fused with distal surface of ventral marginal, 
S-shaped distal extension forming sentinel; accessory terminal 2 short, attached 
to accessory terminal 1, with a dorsoventrally flattened, disc-shaped extension 
forming spike. 



VOLUME 95, NUMBER 1 




dmg 





1cm 



vmg 




Fig. 2a, b, c. Right clasper cartilages of R. nitida, AMS 1B5275: a. Lateral view, partially ex- 
panded with dorsal terminal and ventral terminal cartilages shown separately; b. Dorsal view; c, 
Ventral view; d, e, f, right clasper cartilages oi P. alleni, WAM P19118 (Holotype), d. Lateral view, 
partially expanded with dorsal terminal and ventral terminal cartilages shown separately; e. Dorsal 
view; f. Ventral view. Atrj — accessory terminal 1, atra — accessory terminal 2, ax — axial, dmg — dorsal 
marginal, dtrj — dorsal terminal 1, dtrg — dorsal terminal 2, dtrg — dorsal terminal 3, vmg — ventral mar- 
ginal, vtr — ventral terminal. 



Neurocranium. — The neurocranium of R. nitida has a short, slender rostral 
shaft, fused to flattened rostral node and appendices at tip of snout (Fig. 3a), 
widely separated from rostral base and neurocranium; rostral appendices free of 
rostral shaft posteriorly and possessing two foramina; propterygia of pectoral 
girdle reaching rostral appendices; nasal capsules of moderate size, set at about 
a 40° angle to transverse axis of neurocranium; foramen for profundus nerve on 
leading edge of nasal capsule; anterior foramen for preorbital canal on anterior 
margin of kidney-shaped basal fenestra of nasal capsule; precerebral space nar- 
row, inner walls of nasal capsules moderately bulging into precerebral space; 
interorbital region narrow (Table 1); preorbital processes poorly developed, sep- 
arated from supraorbital crests by shallow notch; postorbital processes well de- 
veloped; anterior fontanelle tear-shaped; posterior fontanelle trapezoid-shaped; 



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1cm 





postp 





Fig. 3. Neurocranium of R. nitida, AMS IB. 5275: a, Dorsal view; b, Lateral view; c, Posterior 
view; d, Ventral view, ac — anterior cerebral vein foramen, af — anterior fontanelle, ante — antorbital 
condyle, bf — basal fenestra, end — endolymphatic foramen, es — efferent spiracular artery foramen, 
hf — hyomandibular facet, ic — internal carotid artery foramen, into — intercerebral vein foramen, ja — 
jugal arch, ibX — lateralis branch of vagus nerve foramen, nc — nasal capsule, obf — otic branch of 
facial nerve foramen, of — orbital fissure, one — orbitonasal canal, os — optic stalk, peri — perilymphatic 
foramen, pf — posterior fontanelle, poc — preorbital canal foramen, postp — postorbital process, prep — 
preorbital process, prof — profundus nerve foramen, ra — rostral appendix, rb — rostral base, rn — ros- 
tral node, rs — rostral shaft, II — optic nerve foramen. III — oculomotor nerve foramen, IV — trochlear 
nerve foramen, VII — hyomandibular branch of facial nerve foramen, IX — glossopharyngeal nerve 
foramen, X — vagus nerve foramen. 



foramen for anterior cerebral vein posterior to line connecting foramina for preor- 
bital and orbitonasal canals (Fig. 3b); trochlear nerve foramen posterior and dor- 
sal to optic nerve foramen; oculomotor nerve foramen situated above optic stock; 
orbital fissure located on posterior aspect of orbit, anterior to foramen for hyo- 
mandibular branch of facial nerve and posterior to foramen for interorbital vein; 
efferent spiracular artery foramen on ventral rim of orbit; jugal arches moderately 
slender; vagus nerve foramen immediately ventral to foramen for lateralis branch 
of vagus nerve and medial to foramen for glossopharyngeal nerve (Fig. 3c); basal 
plate moderately narrow (Fig. 3d). 

Scapulocoracoid. — The scapulocoracoids of/?, nitida are moderately short and 
anteroposteriorly elongated (Table 2), without an anterior bridge (Fig. 5a); an- 
terior fenestra little expanded; postdorsal fenestra moderately expanded; meso- 
condyle not expanded; three postventral foramina, first greatly enlarged; neo- 
pterygial ridge between mesocondyle and metacondyle incomplete. 

Comments. — Raja nitida differs from all other species of Raja in clasper and 
rostral structure and differs from all other genera of skates in its combination of 
derived character states, suggesting that the genus Pavoraja should be resur- 
rected for this species. ''Raja'' nitida differs from Raja in possessing claspers 
with membranous dorsal terminal 1 and ventral terminal cartilages which are 
broadly joined along the ventral aspect of the glans and disc-like accessory ter- 
minal 2 cartilages; a reduced and incomplete rostral shaft which is widely sepa- 



VOLUME 95, NUMBER 1 




Fig. 4. Neurocranium of P. alleni, WAM P19117 (Paratype): a, Dorsal view; b, Lateral view; c, 
Posterior view; d, Ventral view. Abbreviations as in Fig. 3. 



rated from the rostral base, and propterygia of pectoral fins which reach rostral 
appendices. Reduction of the rostral base precludes this species from being clas- 
sified in Raja (Ishiyama 1958, Stehmann 1970, Hulley 1972, McEachran and Com- 
pagno 1982). 



Table 1 . — Neurocranial measurements of P. alleni and P. nitida expressed as percentage of na- 
sobasal length. 



p. alleni 
278 mm TL 9 



P. nitida 
335 mm TL S 



P. nitida 
357 mm TL S 



Nasobasal length (mm) 

Cranial length 

Rostral cartilage length 

Prefontanelle length 

Cranial width 

Interorbital width 

Rostral base 

Anterior fontanelle length 

Anterior fontanelle width 

Posterior fontanelle length 

Posterior fontanelle width 

Rostral appendix length 

Rostral appendix width 

Rostral cleft length 

Cranial height 

Width across otic capsules 

Least width of basal plate 

Greatest width of nasal capsule 

Internasal width 



26.7 


36.2 


40.1 


185 


168 


151 


23 


20 


24 


84 


70 


70 


92 


80 


80 


26 


27 


25 


13 


10 


11 


33 


29 


28 


14 


9 


8 


42 


47 


34 


12 


17 


11 


16 


14 


11 


15 


10 


11 


6 


6 


5 


24 


23 


22 


58 


50 


51 


24 


25 


24 


37 


37 


37 


13 


10 


11 



6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Table 2. — Scapulocoracoid measurements of P. alleni and P. nitida expressed as percentage of 
scapulocoracoid length. 





p. alleni 


p. nitida 


p. nitida 




278 mm TL 9 


335 mm TL S 


357 mm TL c? 


Scapulocoracoid length (mm) 


16.3 


21.5 


25.1 


Scapulocoracoid height 


71 


74 


75 


Premesocondyle 


39 


40 


41 


Postmesocondyle 


61 


60 


58 


Postdorsal fenestra length 


32 


32 


33 


Postdorsal fenestra height 


24 


25 


19 


Anterior fenestra length 


17 


15 


15 


Anterior fenestra height 


25 


24 


22 


Rear corner 


53 


59 


58 



Pavoraja Whitley, 1939 

Type species. — Raja nitida Giinther, 1880. 

Diagnosis. — Snout with small laterally compressed rostral process; caudal fin 
poorly developed, with an epichordal lobe and with or without a hypochordal 
lobe; claspers constricted at origin of glans; dorsal marginal with a distomedial 
extension forming pseudorhipidion; dorsal terminal 1 and ventral terminal mem- 
branous and broadly joined along ventral aspect of glans; dorsal terminal 3 with 
free distal tip forming spur; accessory terminal 1 with an S-shaped distal extension 
forming sentinel; accessory terminal 2 with a disc-shaped extension forming 
spike; rostral shaft slender and reduced, separated by a wide distance from nar- 
row rostral base; rostral appendices short, propterygia of pectoral girdle extending 
to rostral appendices; nasal capsules with basal fenestrae; anterior fontanelle and 
internarial plate moderately narrow; interorbital region narrow; preorbital pro- 
cesses poorly developed; foramen for anterior cerebral vein posterior to line 



scp 



pdfe 




pre 



mtc 



msc 



Fig. 5. Lateral view of scapulocoracoid: a, R. nitida, AMS IB. 8274; b, Pavoraja alleni, WAM 
19117 (Paratype). af — anterior fontanelle, msc — mesocondyle, mtc — metacondyle, pdfe — postdorsal 
fenestra, pre — procondyle, pvf — ^postventral foramina, scp — scapular process. 



VOLUME 95, NUMBER 1 7 

connecting foramina for precerebral and orbitonasal canals; scapulocoracoid sub- 
rectangular to almost rectangular, moderately short and anteroposteriorly ex- 
panded; without anterior bridge; three postventral foramina; precaudal mono- 
spondylous vertebrae ranging from 26 to 29 and predorsal caudal diplospondylous 
vertebrae ranging from 66 to 79. 

Remarks. — Arhynchobatis, Bathyraja, Breviraja, Gurgesiella, Psammobatis, 
Pseudoraja, ''Raja'' waitei, and Sympterygia also possess reduced rostra, a de- 
rived state (McEachran and Compagno 1979, 1982). Rostra of Arhynchobatis , 
Bathyraja (in part), Psammobatis, ''Raja'' waitei, and Sympterygia are basally 
segmented and could presumably have evolved into the P. nitida state by distal 
retraction of the rostral shaft. The rostral shaft in Psammobatis is partially re- 
tracted while that of Pseudoraja is nearly absent (McEachran and Compagno 
1979). However, all of these taxa differ from P. nitida in structure of the claspers, 
neurocrania and scapulocoracoids (Compagno and McEachran, in prep.) and thus 
it seems likely that rostral reduction has occurred separately several times within 
the skates. Pavoraja nitida shows a closer relationship to the genera with reduced 
but basally unsegmented rostra {Breviraja and Gurgesiella) in structure of the 
neurocranium and scapulocoracoid, and to a lesser degree, in clasper structure, 
and possibly could have been derived from a common ancestor of either of these 
taxa. Breviraja possesses a distally segmented rostral shaft which could have 
evolved into the P. nitida state by basal retraction of the proximal, unsegmented 
part of the shaft. However, uniqueness of the clasper and rostral structure of P. 
nitida precludes its classification with either Breviraja or Gurgesiella and sup- 
ports the resurrection oi Pavoraja. 

Based on the examination of several small immature specimens of R. polym- 
mata, this species is classified in Raja rather than Pavoraja. It possesses a stout 
rostral shaft and in all aspects agrees with the anatomical character states within 
Raja. Further comments on its relationships must await the procurement of large, 
mature specimens to make more extensive anatomical observations and to ex- 
amine the claspers. 

Pavoraja nitida (Giinther, 1880) 
(Fig. 6, Table 3) 

Diagnosis. — Disc width greater than 54% of total length; orbital length 1.7 to 
2.1 times as long as spiracle; anterior margin of pelvic fins less than 75% of 
distance from origin of anterior lobe to extreme posterior margin of fin; second 
dorsal fin and epichordal lobe of caudal fin confluent; epichordal lobe shorter than 
base of second dorsal fin; hypochordal lobe of caudal fin absent; dorsal surface 
dark brown with small light spots, some of which are arranged into occelli; mature 
males range from 312 to 363 mm TL. 

Comments. — It is very unlikely that the type locality of P. nitida is South 
America rather than Australia, as suggested by Whitley (1940). The holotype of 
P. nitida does not differ significantly from the specimens from Australia. No 
specimens resembling P. nitida have been reported from South America, and 
specimens resembling this species have not been discovered in the large samples 
of South American skates studied by McEachran. 

Material examined.— BMNH 1879.5.14.417 (Holotype), Two Fold Bay, New 



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Fig. 6. Pavoraja nitida, BMNH 1879.5.14.417 (Holotype). 



South Wales, Australia, HMS Challenger, 6 Aprils June 1874. AMS 1A.2493 
(1), AMS lA. 3904.05 (1), AMS IB.4324.25 (2), AMS 1B.5275 (2), AMS 1B.8274 
(3), AMS E.2165 (1), AMS E.5453 (1), AMS 1.10823.24 (2), AMS 1.16564.001 (1), 
AMS IA.3904.05 (1), TMH D816 (1). 



Pavoraja alleni, new species 

Holotype. — WAM P19118, 297 mm TL, mature male, collected off northwest- 
ern Australia, near Rowley Shoals, 17°17.0'S, 119°57.0'E, 350 m, 20 December 
1969. 

Paratypes. — WAM P19117, 278 mm TL, female, collected with holotype; 
WAM P8226, 159 mm TL, female, collected December 1963 or January 1964 in 
the eastern Indian Ocean, 17°05'S, 119°48'E, aboard the Umitaka Mara. 

Diagnosis. — Disc width less than 52% of total length; orbital length 2.2 to 3.2 
times as long as spiracle; anterior margin of pelvic fins 81 to 104% of distance 
from origin of anterior lobe of fin to extreme posterior margin of fin; second 
dorsal fin and epichordal lobe of caudal fin not confluent, distance from posterior 
margin of base of second dorsal fin to tip of tail considerably greater than length 
of base of second dorsal fin; hypochordal lobe of caudal fin small but present; 
dorsal surface light tan with minute dark spots loosely concentrated into ill de- 
fined blotches but not forming ocelli; males mature at 297 mm TL. 

Description. — Disc 1.1 times as broad as long; maximum angle in front of 
spiracles 107° in holotype (110° to 112° in paratypes); margin of disc convex 
except concave opposite orbits and spiracles; outer corners of disc broadly round- 



VOLUME 95, NUMBER 1 9 

Table 3. — Proportional measurements and meristic values for P. alleni and P. nitida. Pro- 
portions are expressed as percentage of total length. 





p. alleni 


p. alleni 


p. alleni 


p. alleni 


p. nitida 


p. n. 


itida 


p. nitida 




(holotype) 


(paratype) 


(paratype) 


X 


(holotype) 


n = 


15 


X 


Sex 


s 


? 


$ 




s 








Total length (mm) 


297 


278 


159 




209 


242- 


-358 




Disc width 


50.8 


49.3 


47.2 


49 


59 


54-62 


59 


Disc length 


AAA 


43.5 


42.8 


44 


50 


45- 


-52 


51 


Snout length (preocular) 


10.4 


9.6 


11.7 


10.4 


11.0 


8.3- 


-11.4 


9.5 


Snout length (preoral) 


9.8 


10.5 


12.9 


10.7 


11.8 


9.3- 


-12.6 


10.6 


Snout to maximum width 


28.6 


26.3 


27.0 


27 


29 


25- 


-31 


28.7 


Prenasal length 


8.1 


8.0 


9.7 


8.4 


8.2 


6.3- 


-9.1 


1.1 


Orbit diameter 


4.7 


3.9 


4.7 


4.4 


5.7 


5.2- 


-6.1 


5.6 


Distance between orbits 


2.7 


3.1 


3.3 


3.0 


3.2 


2.9- 


-3.7 


3.3 


Orbit and spiracle length 


5.3 


4.6 


5.2 


5.0 


6.3 


6.2- 


-8.9 


6.6 


Spiracle length 


1.5 


1.8 


1.4 


1.6 


3.2 


2.7- 


-6.3 


3.2 


Distance between spiracles 


6.6 


6.0 


6.4 


6.3 


6.5 


4.2- 


-6.7 


6.3 


Mouth width 


6.9 


6.1 


5.8 


6.3 


6.3 


5.9-7.9 


6.9 


Nare to mouth 


3.5 


3.5 


4.5 


3.7 


3.5 


3.2-4.7 


4.2 


Distance between nostrils 


4.8 


4.5 


5.2 


4.8 


4.1 


3.2^.7 


3.9 


Width of first gill opening 


0.9 


1.0 


0.8 


0.9 


1.7 


1.6-2.1 


1.7 


Width of third gill opening 


1.0 


1.1 


0.8 


1.0 


1.9 


1.5- 


-2.0 


1.7 


Width of fifth gill opening 


0.7 


0.9 


0.8 


0.8 


1.6 


1.3- 


-1.6 


1.4 


Distance between first gill openings 


12 


12 


12 


12 


13 


15- 


-12 


12 


Distance between fifth gill openings 


8 


8 


7 


8 


7 


6-8 


6 


Length of anterior pelvic lobe 


12 


11 


13 


12 


12 


12- 


-14 


13 


Length of posterior pelvic lobe 


13 


14 


13 


14 


16 


17- 


-18 


17 


Distance — snout to cloaca 


41 


41 


40 


41 


44 


41-45 


43 


Distance — cloaca to 1st dorsal fin 


47 


47 


48 


47 


46 


43^7 


45 


Distance — cloaca to caudal origin 


55 


57 


55 


55 


54 


51- 


-56 


53 


Distance — cloaca to caudal tip 


59 


59 


61 


60 


57 


54-59 


57 


Number of tooth rows (upper jaw) 


38 


35 


45 


39 


34 


31- 


-36 


33.9 


Sample size of radiographs 








3 




4 




Number of trunk vertebrae 


26 


28 


26 


26.7 


27 


26-29 


27.5 


Number of predorsal caudal vertebrae 


79 


71 


74 


74.6 


66 


66-72 


70.2 


Number of pectoral radials 


63 


67 




65.5 


72 


62- 


-73 


69.8 


Number of pelvic radials 


16 


15 




15.5 


20 


19- 


-20 


19.7 



ed. Tip of snout with small, laterally flattened process. Axis of greatest width 
76% (67 to 68%) of distance from tip of snout to axil to pectoral fins. Pelvic fins 
deeply incised, anterior lobe narrow and acutely pointed; anterior margin 85% 
(81 to 104%) as long as distance from origin of anterior lobe to posterior extreme 
of fin. Tail slender, little depressed, its width at midlength about two-thirds di- 
ameter of eye. Tail with narrow lateral fold along ventrolateral surface running 
from near tip of pelvic fins to origin of hypochordal lobe of caudal fin, widening 
near tip of tail fold to equal height of epichordal lobe of caudal fin. Length of tail 
from center of cloaca to distal tip 1.4 times (1.5 times) distance from tip of snout 
of center of cloaca. 

Preocular length 2.5 times (2.2 to 2.5 times) as long as orbit; preoral length 2.0 
times (2.4 to 2.5 times) internarial distance. Interorbital distance 0.6 (0.7 to 0.8) 
times length of orbit, orbit length 3.1 (2.2 to 3.3) times as long as spiracles. 



10 



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Fig. 7. Pavoraja alleni, WAM P19118 (Holotype): a, Dorsal view; b, Ventral view; c, Ventral 
view of head. 




Fig. 8. Pavoraja alleni, WAM P19117 (Paratype). 



VOLUME 95, NUMBER 1 11 

Anterior and posterior nasal flaps without fringes; without nasal pits. Upper and 
lower jaws moderately arched (little arched). Teeth with pointed cusps near sym- 
physes of jaws, with rounded cusps near margin of jaws (with rounded cusps 
throughout jaws), teeth arranged in more or less transverse series (in quincunx). 

Distance between first gill slits 2.5 (2.4 to 2.8) times as great as between nares; 
distance between fifth gill slits 1.6 (1.3 to 1.8) times as great as between nares; 
length of first gill slits 1.4 (1.1 to 1.2) times length of fifth gill slits and 0.1 (0.1 to 
0.2) times mouth width. First dorsal fin slightly higher and longer than second; 
interspace between dorsal fins equal to or slightly shorter than base of first dorsal 
fin; second dorsal fin separated from epichordal caudal-fin lobe by distance equal 
to one-half base of second dorsal fin; epichordal lobe low, length of base about 
equal to that of second dorsal fin; distance from end of base of second dorsal to 
tip of tail considerably greater than length of base of second dorsal fin; hypo- 
chordal caudal lobe small. 

Upper surface of disc, pelvics and tail densely covered with denticles. Ventral 
surface naked. Orbit with 3 thorns on anteromedial margin and 3 thorns on pos- 
teromedial margin; 3 pre nuchal and 1 nuchal thorns; 3 irregular rows of thorns 
on dorsal surface of tail, no interdorsal thorns. Holotype with 5 alar spines on 
left and 4 on right lateral aspect of disc and a patch of malar thorns on antero- 
lateral aspect of disc. 

The claspers of P. alleni are similar to those of P. nitida with the following 
exceptions: claspers very slender; rhipidion poorly developed; sentinel curved 
laterally and extending to tip of glans (Fig. lb); dorsal terminal 1 only loosely 
connected to ventral terminal along ventral aspect of glans; dorsal terminal 3 with 
a longer distal extension forming spur (Fig. 2d, e, f); ventral marginal with a 
truncated distal margin; accessory terminal 1 with a relatively longer and less 
curved distal extension forming sentinel; accessory terminal 2 with a more elon- 
gated shaft supporting disc-like extension forming spike. 

The neurocranium of P. alleni is similar to that of P. nitida with the following 
exceptions: nasal capsules set at about a 30° angle to transverse axis of neuro- 
cranium (Fig. 4); rostral base better developed, extending nearly to leading edge 
of nasal capsules; precerebral space broader (Table 1), inner walls of nasal cap- 
sules not appreciably bulging into precerebral space; postorbital processes poorly 
developed; jugal arches relatively slender. 

The scapulocoracoids of P. alleni are similar to those of P. nitida (Table 2) 
with the following exceptions: the anterior vertical margin is more perpendicular 
to horizontal axis (Fig. 5b); posterior corner more posteriorly located; postdorsal 
fenestra is oval-shaped rather than elliptical. 

Color. — Dorsal surface uniformly light tan with minute dark spots loosely con- 
centrated into ill defined, symmetrically arranged blotches; tail darker with four 
obscure brown bands; dorsal fins tan with brown blotches. Ventral surface light 
tan. 

Etymology . — Named after Gerald R. Allen (WAM) who furnished us with the 
specimens of the new species. 

Acknowledgments 

We wish to thank Alwyne Wheeler and Peter J. P. Whitehead for providing 
work space at the British Museum (Natural History); Gerald R. Allen (WAM), 



12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

A. P. Andrews (TMH), and Doug Hoese (AMS) for providing specimens; and 
Matthias Stehmann for providing a radiograph and photograph of the holoype of 
Pavoraja nitida. Figures la, 2a, b, c and 3 were prepared by Debbie Allen, other 
figures by Janice D. Fechhelm. Helen Feney labelled and mounted the figures. 

Literature Cited 

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the western North Atlantic, Part II. Sawfishes, 

guitarfishes, skates and rays, and chimaeroids. — Memoir Sears Foundation for Marine Re- 
search 1(2): 588. 
Fowler, H. W. 1941. The fishes of the groups Elasmobranchii, Holocephah, Isospondyli, and Ostar- 

ophysi obtained by the United States Bureau of Fisheries Steamer "Albatross" in 1907 to 

1910, chiefly in the Philippine Islands and adjacent seas. — United States National Museum 

Bulletin 100(13): 1-879. 
Hulley, P. A. 1972. The origin, interrelationships and distribution of southern African Rajidae (Chon- 

drichthyes, Batoidei). — Annals of the South African Museum 60:1-103, 
Ishiyama, R. 1958. Studies on the rajid fishes (Rajidae) found in the waters around Japan. — Journal 

Shimonoseki College of Fisheries 7:1-394. 
McEachran, J. D., and L. J. V. Compagno. 1979. A further description of Gurgesiella furvescens 

with comments on the interrelationships of Gurgesiellidae and Pseudorajidae (Pisces, Rajoid- 

ei).— BuUetin of Marine Science 29(4): 530-553. 
, and . 1982. Interrelationships of and within Breviraja based on anatomical structures 

(Pisces: Rajoidei). — Bulletin of Marine Science, in press. 
Stehmann, M. 1970. Vergleichend morphologische und anatomische Untersuchungen zur Neuord- 

nung der Systematik der nordostatlantischen Rajidae. — Archiv fiir Fischeriewissenschaft 

21:73-164. 
Whitley, G. P. 1939. Taxonomic notes on sharks and rays. — Australian Zoologist 9:227-262, 2 plates. 
. 1940. The fishes of Australia. Part I. The sharks, rays, devil fishes, and other primitive fishes 

of Australia and New Zealand. Royal Zoological Society, New South Wales, Sydney. 279 pp. 

Department of Wildlife and Fisheries Sciences, Texas A&M University, Col- 
lege Station, Texas 77843 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 13-26 

A NEW SPECIES OF GYMNURE, PODOGYMNURA, 

(MAMMALIA: ERINACEIDAE) FROM 

DIN AG AT ISLAND, PHILIPPINES 

Lawrence R. Heaney and Gary S. Morgan 

Abstract. — A new species, Podogymnura aureospinula, of Philippine gymnure 
is described from Dinagat, a small island off the northeast coast of Mindanao in 
the southern Philippines. This species is distinguished from other members of the 
subfamily by the possession of spiny pelage, inflation of the frontal region, and 
presence of a distinct cusp at the base of the talonid notch. This new species is 
second in size among the living Echinosoricinae only to Echinosorex gymnurus. 
The relationship of Podogymnura to the other extant genera of echinosoricines 
is discussed. Based on these comparisons, Podogymnura and Echinosorex are 
shown to share a number of cranial and dental characters and are considered to 
be more closely related to one another than either is to Hylomys or Neotetracus. 



In 1972 and 1975, Dioscoro S. Rabor and a field party from Mindanao State 
University collected mammals and birds on Dinagat and Siargao islands, which 
are located off the northeast coast of Mindanao in the southern Philippines. The 
itinerary, habitat descriptions, and a report on the birds from the 1972 trip may 
be found in duPont and Rabor (1973), and a report on the mammals in Heaney 
and Rabor (1982). Specimens from this collection are housed in the Delaware 
Museum of Natural History, University of the Philippines at Los Banos, and the 
U.S. National Museum of Natural History. Among the mammals collected on 
Dinagat are four specimens of a unique member of the family Erinaceidae. In this 
paper, we describe these specimens as a new species and discuss the relationships 
of the Philippine gymnures, Podogymnura, within the erinaceid subfamily Echi- 
nosoricinae. 

Methods 

External measurements were taken from collector's labels. Cranial measure- 
ments (Table 1) were taken by Heaney with dial calipers graduated to 0.1 mm. 
Dental measurements (Table 2) were taken by Morgan using an Anderson cra- 
niometer attached to a Bausch and Lomb binocular microscope (Anderson 1968). 
All cranial measurements are as defined in DeBlase and Martin (1974) except the 
following: rostral length, from midline at anterior tip of nasals to orbital margin 
of infraorbital canal; rostral breadth, taken at labial edge of premaxillae just 
posterior to P; post-palatal depth, depth of cranium measured at the point just 
posterior of palate to point above at 90° to occlusal plane of molars; P to M^, 
maximum labial length from anterior edge of P alveolus to posterior edge of M^ 
at alveolus; M^ to M^, greatest width of palate taken at labial margin of alveoli; 
palatal width at M^, alveolar distance between lingual margins; height of coronoid, 
maximum vertical height from ventral edge of mandible to tip of coronoid process; 
depth of mandible, vertical height from ventral edge of mandible to alveolar 



14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

surface between Mi and Mg; thickness of mandible, distance from lingual to labial 
edge of mandible taken between Mi and Mg. All dental measurements in Table 
1 are dimensions of the teeth, not their alveoli, and represent the maximum length 
and/or width of a specified tooth or of a series of teeth. Dental nomenclature 
follows Szalay (1969:202). Specimens from the following museums were used in 
this study (standard acronym follows in parentheses): Delaware Museum of Nat- 
ural History (DMNH), Field Museum of Natural History (FMNH), University of 
the Philippines at Los Banos, Museum of Natural History (UPLB), and the U.S. 
National Museum of Natural History (USNM). 

Podogymnura aureospinula, new species 

Holotype.—DMNn 4386, adult female, skin and skull. Obtained 23 April 1972 
by Dioscoro S. Rabor (original number 259) at Plaridel, Albor Municipality, 
Dinagat Island, Surigao del Norte Province, Republic of the Philippines. Skin well 
prepared and in good condition. Skull complete except for zygomatic arches, 
both of which are broken (Figs. 1,2). 

Referred specimens. — The holotype and one adult male from Balitbiton, Loreto 
Municipality, Dinagat (UPLB 3753) were examined and measured. Two additional 
specimens (one male, one female) from Kambinlio, Loreto Municipality, Dinagat, 
are in the collection at UPLB, but were not examined. 

Diagnosis. — Size large; dorsal pelage short and spinous, golden brown color 
overall; temporal, sagittal, and nuchal crests prominent; frontal region conspic- 
uously inflated; interorbital region strongly constricted; external pterygoid pro- 
cesses large and separated at base from internal pterygoid processes by deep 
groove; mandibular rami robust; P^ large; P^ broad lingually; metaconule prom- 
inent on M^ and M^; metacone present on M^; P4 with small, but distinct, meta- 
conid; distinct cusp at base of talonid notch between metaconid and entoconid 
on M1-M3. 

Description. — Size (Tables 1 and 2) large for extant members of subfamily. 
Dorsal pelage composed of three types of hairs: slate-gray underfur ca. 5 mm in 
length; stiff, bristly, or spiny hairs, black at base and remainder golden yellow, 
many with black tips, ca. 15 mm in length; black, spiny hairs, ca. 12 mm in 
length. Golden spines twice as abundant as black hairs in middle of back. Black 
spines densest at mid-dorsum, decreasing in abundance laterally, disappearing on 
sides. Black-tipped golden spines especially common at mid-dorsum, also dis- 
appearing on sides. Only golden spines present on sides. Overall color of dorsum 
golden-brown, with black spines and black-tipped golden spines adding a black 
speckling. On holotype, golden color distinctly metallic when viewed at proper 
angle. Other specimens somewhat faded, not as metallic. Ventral pelage lacks 
spines, grades evenly from dorsal color to brownish-gray over most of venter; 
throat darker on some specimens. Ventral hairs of two types: soft, gray underfur 
ca. 5 mm in length, and slightly coarser guard hairs ca. 9 mm in length, gray at 
base and tipped with light brown. Pelage of rostrum and around eyes short, dense, 
and spiny. Upper and lower lips clothed in very short, moderately dense, white 
or light brown fur. Vibrissae dark at base, very light for most of length, up to 55 
mm. Rhinarium long, naked, and distinctly bilobed, with nostrils opening later- 
ally. Ears relatively large, appearing naked, but with sparse covering of extremely 



VOLUME 95, NUMBER 1 



15 




Fig. 1. From top to bottom — dorsal, ventral, and left lateral views of cranium and lateral view of 
mandible of Podogymnura truei truei (FMNH 61453) from Mt. Apo, Davao Prov., Mindanao (1^) 
and Podogymnura aureospinula (DMNH 4386), holotype, from Plaridel, Dinagat, Surigao del Norte 
Prov. (5-8). Actual size. 



short, white hairs. Fore and hind feet with moderate covering of short, white or 
Hght brown hairs dorsally, nearly naked ventrally. Hind legs appear almost naked 
distal to knee joint. Dorsal base of tail with area sparsely furred, nearly naked, 
15 mm in diameter. Tail with sparse covering of short hairs. Two pairs of mam- 
mae, one pair pectoral, one pair inguinal. 

Skull large and robust. Temporal crests converge at or slightly anterior to 



16 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





^m 



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Fig. 2. Stereophotographs of upper (1, 1') and lower (2, 2') dentitions of Podogymnura aureo- 
spinula (DMNH 4836), holotype. x3. 



interorbital constriction to form sagittal crest. Sagittal crest prominent, 0.5 to 1.0 
mm in height anteriorly to 2.5 mm in height posteriorly where it meets nuchal 
crest. Nuchal crest well developed. Frontal region conspicuously inflated. Fron- 
tals expanded laterally into orbital fossa. Rostrum relatively broad posteriorly, 
resulting from inflation of posterodorsal portion of maxilla. Inflation causing con- 
vexity in dorsal profile of skull, beginning at anterior edge of frontals, approxi- 
mately dorsal to P^, and extending posteriorly to interorbital constriction. Highest 
point on skuU dorsal to orbits. Interorbital region constricted. Braincase relatively 
long and not noticeably inflated. Paroccipital and mastoid processes prominent. 
Mastoid exposure on ventrolateral corner of skuU gently concave and composed 
of thick bone. Post-tympanic process of squamosal large, broad posteriorly, and 
with well developed epitympanic sinus. Periotic not inflated. Periotic component 
of bulla reduced, having a distinct, rounded emargination in ventromedial edge. 
Basisphenoid component of bulla more vertically oriented, projecting ventral to 



VOLUME 95, NUMBER 1 17 

level of occipital condyles. Tympanic cavity broadly open ventrally. External 
pterygoid processes large, triangular- shaped, and separated at base from internal 
pterygoid processes by deep groove. Mandibular rami relatively thick. 

All incisors single-rooted, P enlarged. P and P much smaller, nearly identical 
in size. Canine double-rooted, long, and flared laterally. P^ single-rooted, small 
compared to other premolars. P three-rooted, some specimens with slight lingual 
expansion and small, but distinct, hypocone. P^ broad lingually, with hypocone 
and protocone parallel to palatal midline. Metaconule prominent on M^ and M^, 
protoconule absent. M^ relatively large, with small, but distinct metacone. Ii and 
I2 approximately same size, spatulate, and procumbent. I3 considerably smaller. 
Canine large and vertical, highest tooth in lower tooth row. Pg and P3 compara- 
tively robust, P2 one-half the size of P3. P4 large, talonid basin moderately to well 
developed, and distinct metaconid present. Lower molars with small cusp at base 
of talonid notch between entoconid and metaconid, sometimes absent on M3. 
Postcristid on Mi and M2 slopes lingually at hypoconulid to meet moderately to 
strongly developed postcingulid. 

Etymology. — L. aureus, golden; L. spinula, diminutive of thorn. The specific 
name refers to the golden spines which characterize the dorsal pelage of this 
species. We suggest "golden-spined gymnure" as an English name. 

Comparisons. — Podogymnura aureospinula is more closely related to Podo- 
gymnura truei than to any other species in the Echinosoricinae as judged by the 
combination of the following characters: long rostrum, absence of postorbital 
processes, constricted interorbital region, extreme anterior placement of upper 
molariform teeth relative to orbit and infraorbital foramen, relatively small P, P 
and P equal in size, large laterally flaring upper canines, loss of P}, comparatively 
large P, and M^ and M^ square in outline. On the other hand, P. aureospinula 
possesses at least three derived characters that are unique among the Echino- 
soricinae: spiny dorsal pelage, conspicuously inflated frontal region, and presence 
of a cusp at base of talonid notch on lower molars. These characters might justify 
generic distinction for P. aureospinula', however, rather than erect a monotypic 
genus, we choose to place this new species in the genus Podogymnura to indicate 
the presumed monophyletic nature of the two endemic Philippine erinaceids. 

Podogymnura truei is the only echinosoricine which requires detailed compar- 
ison with P. aureospinula. The other extant echinosoricine genera are compared 
to Podogymnura in more general terms in the Discussion section. Podogymnura 
aureospinula and P. truei are quite different in external appearance. Besides its 
smaller size, P. truei has a longer, softer pelage, with no indication of spines, or 
even coarse hairs. Its underfur is particularly long, about twice as long as that of 
P. aureospinula. The dorsal surfaces of the fore and hind feet of P. truei have 
longer, darker hairs. The overall color of the pelage is darker in P. truei; dorsally 
it varies from reddish to chestnut brown compared to the golden brown color of 
the large species, and ventrally it is a medium brown, whereas the venter of P. 
aureospinula is gray, with a brownish tinge. 

The most obvious difference between the skulls oi Podogymnura aureospinula 
and P. truei is greater size of the former, its skull being 20% longer than the 
largest skull of P. truei measured (Table 1). Perhaps as a result of allometric 
changes correlated with increasing skull size, the temporal, sagittal, and nuchal 
crests and paroccipital and mastoid processes are more prominent in P. aureo- 



18 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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20 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Selected dental and mandibular measurements (mm) of Podogymnura species. 



Podogymnura 
aureospinula 
Measurement holotype, UPLB 3753 


Podogymnura 
truei truei 
holotype 


Podogymnura 
truei truei 

(N = 5) 


Podogymnura 

truei minima 

(N = 4-5) 


Length of maxillary tooth row 


20.6, 20.4 


15.2 


15.5 ± 0.44 
(15.1-16.2) 


15.0 ± 0.65 
(14.4_15.9) 


Length from M^ to M^ 


9.8, 9.6 


6.7 


7.1 ± 0.21 
(6.9-7.4) 


7.0 ± 0.33 
(6.6-7.5) 


Length of F 


2.7, 2.5 


1.8 


1.9 ± 0.11 
(1.8-2.1) 


1.8 ± 0.08 

(1.7-1.9) 


Width of P 


2.4, 1.7 


1.1 


1.3 ± 0.17 
(1.2-1.6) 


1.4 ± 0.07 
(1.3-1.5) 


Length of P 


3.2,3.0 


2.2 


2.4 ± 0.05 
(2.3-2.4) 


2.4 ± 0.11 
(2.3-2.6) 


Width of P 


3.2, 3.0 


2.1 


2.4 ± 0.10 

(2.3-2.5) 


2.4 ± 1.54 

(2.2-2.5) 


Length of M^ 


3.8, 3.8 


2.6 


2.9 ± 0.04 

(2.8-2.9) 


2.8 ± 0.11 
(2.6-2.9) 


Width of W 


3.7, 3.4 


2.3 


2.7 ± 0.12 
(2.6-2.9) 


2.6 ± 0.08 

(2.5-2.7) 


Length of M^ 


3.5, 3.5 


2.4 


2.5 ± 0.11 

(2.4-2.7) 


2.5 ± 0.16 

(2.3-2.7) 


Width of M2 


3.7,3.3 


2.4 


2.6 ± 0.05 

(2.6-2.7) 


2.6 ± 0.11 

(2.5-2.7) 


Length of M^ 


2.7, 2.6 


1.7 


1.9 ± 0.15 

(1.7-2.1) 


1.8 ± 0.09 
(1.7-1.9) 


Width of M3 


2.8, 2.8 


1.8 


2.1 ± 0.08 
(2.0-2.2) 


2.0 ±0.11 
(1.9-2.1) 


Total length of mandible 


41.2,40.8 




30.1 ± 0.77 
(29.6-31.5) 


28.8 ± 0.58 
(28.2-29.6) 


Height of coronoid 


13.9, 13.4 


9.2 


9.7 ± 0.50 
(9.0-10.4) 


9.3 ± 0.59 
(8.6-10.0) 


Depth of ramus between Mj and Mg 


4.7,4.8 


3.1 


3.3 ± 0.22 
(3.0-3.6) 


3.2 ± 0.25 
(2.8-3.5) 


Breadth of ramus between Mj and 


2.6, 2.8 


1.6 


1.8 ± 0.05 


1.7 ± 0.07 


M2 






(1.7-1.8) 


(1.6-1.8) 


Length of mandibular toothrow 


21.4, 20.7 


15.8 


16.2 ± 0.54 
(15.7-17.1) 


15.7 ± 0.64 
(15.1-16.6) 


Length from Mj to M3 


11.2, 10.5 


7.8 


8.0 ± 0.26 

(7.6-8.2) 


7.8 ± 0.31 
(7.5-8.3) 


Length of P4 


3.0, 2.6 


1.9 


2.0 ± 0.10 
(1.9-2.1) 


2.0 ± 0.19 
(1.8-2.3) 


Width of P4 


1.8, 1.7 


1.1 


1.3 ± 0.04 
(1.2-1.3) 


1.2 ± 0.08 
(1.1-1.3) 


Length of Mi 


4.3, 4.1 


3.0 


3.1 ± 0.11 
(2.9-3.2) 


2.9 ± 0.09 

(2.9-3.2) 


Width of M, 


2.5, 2.5 


1.7 


1.9 ± 0.13 
(1.7-2.0) 


1.8 ± 0.04 
(1.8-1.9) 



VOLUME 95, NUMBER 1 21 

Table 2. — Continued. 

Measurement 

Length of Mg 
Width of M2 
Length of M3 
Width of M3 



Podogymnura 

aureospinula 

holotype, UPLB 3753 


Podogymnura 
truei truei 
holotype 


Podogymnura 
truei truei 

(N = 5) 


Podogymnura 
truei minima 

(N = 4-5) 


4.0, 3.7 


2.7 


2.8 ± 0.13 


2.7 ± 0.08 






(2.6-2.9) 


(2.6-2.8) 


2.4, 2.4 


1.6 


1.8 ± 0.11 


1.8 ± 0.09 






(1.7-1.9) 


(1.7-1.9) 


3.3, 3.1 


2.3 


2.3 ± 0.11 


2.3 ± 0.15 






(2.2-2.4) 


(2.1-2.5) 


2.1, 2.0 


1.4 


1.5 ± 0.05 


1.5 ± 0.16 






(1.5-1.6) 


(1.3-1.7) 



spinula, approaching the condition seen in the much larger Echinosorex. In most 
skulls of P. truei, the weak temporal crests meet near the anterior edge of the 
interparietal, forming a short, weak sagittal crest, while in larger skulls of the 
same species, the temporal crests meet somewhat farther forward. The temporal 
crests of P. aureospinula converge at or slightly anterior to the interorbital con- 
striction to form the strong sagittal crest, which is particularly high posteriorly 
where it bisects the interparietal. The nuchal crest of P. aureospinula is more 
strongly developed than in P. truei. 

Podogymnura truei shows no evidence of the frontal inflation characteristic of 
P. aureospinula. The dorsal margin of the skull of P. truei rises in a nearly 
straight line to reach a maximum height above the glenoid region, rather than 
dorsal to the orbits as in P. aureospinula. The inflation of the frontals and pos- 
terodorsal portion of the maxilla of P. aureospinula is not characteristic of any 
other modern echinosoricine, although some erinaceines (e.g. Paraechinus) have 
moderately inflated frontals. The interorbital region appears to be more constrict- 
ed in P. aureospinula, although this feature is certainly enhanced by the inflation 
anterior to the constriction. The more prominent development of the sagittal and 
nuchal crests gives the braincase of P. aureospinula the appearance of being 
longer, whereas the braincase of the smaller species appears shorter, broader, 
and more bulbous, particularly in smaller specimens. 

The mastoid exposure on the posterolateral corner of the skull is inflated in 
Podogymnura truei and the bone is thin and nearly transparent. The mastoids of 
P. aureospinula are gently concave and composed of thicker bone. The periotic 
of P. truei is also inflated. The periotic is not inflated in P. aureospinula and the 
periotic component of the bulla has a small, rounded emargination in its ventro- 
medial edge which is lacking in the smaUer species. The basisphenoid portion of 
the bulla of P. aureospinula is more vertically oriented, projecting ventral to the 
occipital condyles, whereas the basisphenoid bulla of P. truei is more horizontal, 
tending to enclose the tympanic cavity. Coupling the more vertical orientation of 
the basisphenoid bulla with the reduced periotic portion of the bulla, the tympanic 
cavity is more open ventrally in P. aureospinula. The post-tympanic process of 
the squamosal is relatively larger in P. aureospinula than in any other modern 
echinosoricine. Its external pterygoid processes are also relatively large, but these 
are smaller in P. truei. Podogymnura aureospinula has a deep groove at the base 



22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

of the external pterygoid processes which separates them from the internal pter- 
ygoid processes. There is no evidence of this groove in any specimen of P. truei 
examined. As with the skull, the mandible of P. aureospinula is larger and more 
robust than that of its smaller counterpart. The mandibular rami are particularly 
thickened in P. aureospinula. 

The upper incisors and canines of Podogymnura aureospinula and P. truei are 
similar in morphology. The P^ is somewhat longer and broader in P. aureospinula ; 
it is more conical in P. truei. In most specimens of P. truei, the P is two-rooted, 
thin, and blade-like, with no indication of a protocone or hypocone. Butler (1948) 
stated that the P of Podogymnura had only two roots; however, the larger series 
of specimens now available shows this character to be variable. Most specimens 
of P. truei have a two-rooted P^, but the few that have a slight lingual expansion 
of P^ have three roots. The P^ of P. aureospinula is three-rooted, and bears a 
small hypocone. The P* of P. aureospinula is broader lingually, the hypocone is 
larger, and the protocone is higher and more lingually placed. In comparison, the 
P* in the smaller species is rather narrow lingually, the hypocone is small, and 
the protocone is located closer to the paracone. The M^ and M^ are similar in the 
two species, except for the more prominent metaconule and cingula in P. au- 
reospinula. The M^ of P. aureospinula is larger and bears a small metacone, the 
latter being absent on the M^ of P. truei. 

As in the upper dentition, the Pg and P3 are relatively larger and more robust 
in P. aureospinula than in P. truei. The talonid basin is reduced on P4 in P. truei 
and a metaconid is absent or tiny. The P4 of P. aureospinula has a moderately 
to well developed talonid basin and the metaconid is distinct. The morphology 
of the lower molars is similar in the two species. P. aureospinula has a small, 
distinct cusp located at the base of the talonid notch between the entoconid and 
metaconid on M^, M2, and sometimes M3. The entoconulid is normally located 
slightly anterior to, but close to the entoconid, and therefore this cusp does not 
appear to be homologous with an entoconulid. This cusp is absent in P. truei and 
seems to be unique among modern echinosoricines. 

Ecology. — Virtually nothing is known of the natural history of Podogymnura 
aureospinula. The vicinity where the holotype of P. aureospinula was taken was 
described by duPont and Rabor (1973:4) as ". . .a logged area in rolling country 
and low hills where there were still many patches of remnant dipterocarp forests 
in the surrounding localities." These forests are dominated by Dipterocarpus, 
Shorea, Hopea, Anisoptera, and Pentacme among the Dipterocarpaceae, and 
include members of at least eleven other plant families. Undergrowth consists 
mainly of rattan and ferns. Other terrestrial mammals taken at the type locality 
of P. aureospinula include Urogale everetti, Tarsius syrichta, Cynocephalus vo- 
lans, Sundasciurus mindanensis , Exilisciurus surrutilus, Batomys sp., Rattus ev- 
eretti, and Rattus rattus (Heaney and Rabor 1981). 

Additional specimens examined. — Podogymnura truei truei (2 SS, 4 9$). 
PHILIPPINES, Mindanao, Davao Province: E slope of Mt. McKinley, 5800 ft. 
elev., FMNH 56129, 56172, 56181; N slope of Mt. Apo, Lake Linau, 7800 ft. 
elev., FMNH 61435; E slope of Mt. Apo, Baclayan, 5400 ft. elev., FMNH 61453; 
Mt. Apo, 6000 ft. elev., USNM 125286 (holotype). 

Podogymnura truei minima (2 ^ (5 , 3 $ ?). PHILIPPINES, Mindanao, Bukid- 
non Province: Mt. Katanglad, near Malay Balay, DMNH 5949-5953. 



VOLUME 95, NUMBER 1 23 

Echinosorex gymnurus albus (1 S, A 9$). INDONESIA, Borneo, Sempang 
River: USNM 145581-582, 145584-586. 

Hylomys suillus dorsalis (10 SS, 1 9). MALAYSIA, Borneo, Sabah, Mt. 
Kinabalu, Bundu Tuhan: USNM 292337-339, 292341-342, 292350-354, 292356. 

Neotetracus sinensis (1 c?, 2 9?, 4?). CHINA: Yunnan, Ho mu shu Pass, 
USNM 241402; Szechuan, Kwan Shien, 3000 ft. elev., USNM 258124-129. 

Discussion 

The genus Podogymnura and its type species, P. truei, were described by 
Mearns (1905) from a single specimen collected on Mount Apo, south-central 
Mindanao, in the Philippines. Until the late 1940's, this specimen, an adult female 
consisting of a complete body in alcohol and a skull lacking the zygomatic arches 
and braincase, was the only known specimen of the genus. Mearns provided a 
detailed description of the external characters of P. truei but he did not illustrate 
the cranium or mandible and his description of them was brief. He described the 
dentition of P. truei, comparing it with that of Hylomys, but his descriptions and 
comparisons are so general that they are of little use. Lyon (1909) noted that 
although Podogymnura and Hylomys appeared to be closely related with respect 
to size and external characters, they were distinct dentally. Cabrera (1925) pro- 
vided a key to the living echinosoricine genera in which he distinguished Podo- 
gymnura by the combination of loss of P}, P^ larger than P^, and larger upper 
canines. Butler (1948) regarded Podogymnura as intermediate between Echino- 
sorex and the smaller Hylomys and Neotetracus, although he pointed out that in 
the enlargement of the canines, relatively large P^, length of the rostrum, and 
position of P^ and M^ relative to the orbit and infraorbital foramen, Podogymnura 
is similar to Echinosorex. 

Sanborn (1952) reported on 64 specimens of Podogymnura truei collected on 
Mount Apo and Mount McKinley on Mindanao. He compared Podogymnura to 
the other genera of modern echinosoricines, reaching the same general conclusion 
as did Butler. Sanborn (1953) described a new subspecies of Podogymnura truei, 
P. t. minima, from four specimens collected on Mount Katanglad, Bukidnon 
Province, north-central Mindanao. His diagnosis was based exclusively on the 
smaller size of P. t. minima. Detailed cranial and dental measurements of P. t. 
truei (including the type) and topotypic specimens of P. t. minima (Tables 1 and 
2) indicate that, on the whole, the available specimens from Mount Katanglad are 
slightly smaller than specimens from Mount Apo and Mount McKinley, but there 
is broad overlap in size between them, especially in dental measurements. In 
fact, the teeth of the type specimen of P. t. truei from Mount Apo are actually 
smaller in many dental measurements than the teeth of the topotypic specimens 
of P. t. minima. Sanborn specifically noted that there was no difference in color 
between his series of P. t. minima and the nominate form. The DMNH specimens 
of P. t. minima, however, are lighter in color than topotypic specimens of P. /. 
truei, being slightly more reddish brown and having the tail uniformly lighter. 
Besides the references cited above, HolHster (1913), Taylor (1934), and Alcasid 
(1970) mentioned Podogymnura, but provided no new information. 

Comparison 6>/ Podogymnura with other living echinosoricines. — Butler (1948) 
made exhaustive comparisons of three of the five extant genera of echinosori- 



24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

cines: Hylomys, Neotetracus, and Echinosorex. His comparisons of Podogym- 
nura were general, as they were based solely on published descriptions and fig- 
ures of the type of P. truei, the only specimen of the genus known at that time. 
The fifth Hving genus of echinosoricine, Neohylomys, was not described until 
1959. Since 1948 many specimens of Podogymnura have been collected, but the 
genus has not been adequately compared with the other members of its subfamily. 
Our description of a new species of Podogymnura, which in some characters 
appears to bridge the morphological gap between Podogymnura and Echinosorex, 
calls for fuller comparisons and a reassessment of the phylogenetic relationships 
of the Philippine gymnures. 

As Butler (1948) and others have noted, Hylomys and Neotetracus are closely 
related forms, although in our opinion they do represent distinct genera, contrary 
to Van Valen (1967). The status of Neohylomys is uncertain; the only specimens 
are in China, and unavailable for study. Many of the characters unique to Hy- 
lomys and Neotetracus, such as the reduced antemolar dentition, the palatal 
perforations, the presence of posterior processes of the maxillae and anterior 
processes of the parietals which extend across the frontals and meet or nearly 
meet dorsal to the orbits, the prominent flange on the anterior edge of the orbit, 
and the strongly concave anterior portion of the zygomatic arch for attachment 
of the lateral snout muscles, are derived characters which demonstrate a close 
phylogenetic relationship between the two genera. Conversely, many of the char- 
acters that are shared by Podogymnura and Echinosorex, but are not present in 
the other two genera, appear to be primitive for the Echinosoricinae. Of these 
characters, the longer rostrum, better developed maxillary dentition anterior to 
P*, widely separated maxillae and parietals dorsal to the orbits, and the nonper- 
forated palate are certainly primitive. In the remaining characters shared by Po- 
dogymnura and Echinosorex, it is difficult to determine the primitive character 
state for the Echinosoricinae because no appropriate outgroup exists. Butler 
(1948) felt that the two distinct groups of living echinosoricines represented two 
separate specialized offshoots from the primitive condition. Suffice it to say that 
Podogymnura and Echinosorex share so many cranial and dental features that 
they undoubtedly represent closely related genera. 

Although they share many cranial characters, Podogymnura and Echinosorex 
are extremely different externally. Echinosorex is a larger animal with a tail equal 
to about 60% of the length of head and body. Podogymnura is medium-sized {P. 
aureospinula) or small {P. truei) and has a tail equal to about 35% of the length 
of head and body. Podogymnura is a chestnut brown {P. truei) to golden brown 
{P. aureospinula) color and has relatively short pelage. Echinosorex is either pure 
white {E. gymnurus albus) or predominantly black with white markings on the 
neck and face (all other forms of the species) and has long, coarse guard hairs on 
the dorsum. Based on its external appearance, especially its small size and short 
tail, Podogymnura truei more closely resembles Hylomys and Neotetracus than 
Echinosorex. 

Additional cranial characters shared by Podogymnura and Echinosorex not 
listed above include: well developed canines and P^, small supraorbital crests, 
presence of two longitudinal grooves in maxillary component of hard palate which 
extend from incisive foramina to small foramina located medial to P^ or P, pres- 



VOLUME 95, NUMBER 1 25 

ence of a ventral process on maxillary portion of zygomatic arch, and more 
anterior placement of upper cheek teeth relative to orbit and infraorbital foramen. 
The tympanic bone is slender and not firmly attached to the bulla in Podogymnura 
and Echinosorex, whereas in Hylomys and Podogymnura the tympanic is broader 
and is firmly attached to the edge of the bulla, contributing to its formation and 
restricting the size of the opening of the tympanic cavity. Although Butler (1948) 
stated that Echinosorex lacked the anterior process of the tympanic, the large 
series of echinosoricines at our disposal reveals that both Echinosorex and Po- 
dogymnura definitely have an anterior process, but it is smaller than that of 
Hylomys and Neotetracus. 

In addition to the differences in external morphology cited above, there are a 
number of cranial and dental characters that are diagnostic of Podogymnura and 
distinguish it readily from Echinosorex. These include: less prominent temporal, 
sagittal, and nuchal crests and mastoid and paroccipital processes; more concave 
maxillary portion of zygomatic arch; broader, more inflated braincase; more an- 
terior placement of upper molars relative to orbit and infraorbital foramen (M^ is 
anterior to front edge of orbit and M^ is below infraorbital foramen); smaller P; 
P and P equal in size; canines relatively larger and flared laterally; loss of ?}; 
M^ square in outline; absence of protoconule on M^; smaller metaconule on M^ 
and M^; reduced posterolingual apex of M^. 

Podogymnura aureospinula is intermediate between Echinosorex and P. truei 
in many characters, although some of the characters on which this observation 
is based are probably related to its intermediate size. The more prominent tem- 
poral, sagittal, and nuchal crests and mastoid and paroccipital processes of P. 
aureospinula relative to P. truei are almost certainly correlated with its greater 
size. These features of the bony crests and processes are best developed in Echi- 
nosorex, the largest genus among living echinosoricines, and are likely to be 
aflometric changes associated with increasing skull size. Other intermediate char- 
acters of P. aureospinula which are more difficult to ascribe to allometry are the 
larger P, more prominent metaconule on M^ and M^, the presence of a metacone 
on M^, and the deep groove separating the bases of the external and internal 
pterygoid processes. Whether these are derived characters indicating a closer 
relationship between Echinosorex and P. aureospinula or whether these represent 
primitive characters shared by Echinosorex and P. aureospinula and lost by P. 
truei cannot be determined from available data. Regardless of the similarities 
between P. aureospinula and Echinosorex, the two species of Podogymnura are 
certainly more closely related to one another than either is to Echinosorex and 
have probably been isolated in the Philippines for a considerable period of time. 

Acknowledgments 

We wish to thank M. D. Carleton, P. Myers, and R. W. Thorington for valuable 
comments on the manuscript, and G. Lake and S. F. Campbell for assistance in 
preparation of the manuscript and tables. Photographic assistance was provided 
by D. Bay and V. Krantz. We thank P. Freeman, D. Niles, D. S. Rabor, R. W. 
Thorington, and R. M. Timm for permission to examine specimens under their 
care. 



26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Literature Cited 

Alcasid, G. L. 1970. Checklist of Philippine mammals. — National Museum, Department of Education 

(Manila). Museum Publication No. 5:1-51. 
Anderson, S. 1968. A new craniometer and suggestions for craniometry. — Journal of Mammalogy 

49:221-228. 
Butler, P. M. 1948. On the evolution of the skull and teeth in the Erinaceidae, with special reference 

to fossil material in the British Museum. — Proceedings of the Zoological Society of London 

118:446-500. 
Cabrera, A. 1925. Genera Mammalium — Insectivora, Galeopithecia. — Museo Nacional Ciencia 

Natural, Madrid, 232 pp. 
DeBlase, A. F., and R. E. Martin. 1974. A manual of mammalogy. Wm. C. Brown Co., Dubuque, 

Iowa, 329 pp. 
duPont, J. E., and D. S. Rabor. 1973. Birds of Dinagat and Siargao, Philippines, an expedition 

report. — Nemouria (Occasional Papers of the Delaware Museum of Natural History) 10: 1-1 1 1 . 
Heaney, L. R., and D. S. Rabor. 1982. An annotated checklist of the mammals of Dinagat and 

Siargao islands, Philippines. — (Occasional Papers of the Museum of Zoology, University of 

Michigan) (In press.) 
HoUister, N. 1913. A review of the Philippine land mammals in the United States National Museum. — 

Proceedings of the United States National Museum 46:299-341. 
Lyon, M. W., Jr. 1909. Remarks on the insectivores of the genus Gymnura. — Proceedings of the 

United States National Museum 36:449^56. 
Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the Philippine Is- 
lands. — Proceedings of the United States National Museum 28:425-460. 
Sanborn, C. C. 1952. Philippine zoological expedition 1946-47: Mammals. — Fieldiana: Zoology 

33(2):89-158. 
. 1953. Mammals from Mindanao, Philippine Islands collected by the Danish Philippine Ex- 
pedition 1951-1952. — Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 

115:283-288. 
Szalay, F. S. 1969. Mixodectidae, Microsyopidae, and the insectivore-primate transition. — Bulletin 

of the American Museum of Natural History 140:193-330. 
Taylor, E. H. 1934. Philippine land mammals. — Monographs of the Bureau of Science, Manila 

30:1-548. 
Van Valen, L. 1967. New Paleocene insectivores and insectivore classification. — Bulletin of the 

American Museum of Natural History 135:217-284. 

(LRH) Museum of Zoology and Division of Biology, University of Michigan, 
Ann Arbor, Michigan 48109; (GSM) Division of Mammals, U.S. National Mu- 
seum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 

Present address of GSM: Florida State Museum, University of Florida, Gaines- 
ville, Florida 32611. 



PROC. BIOL. SOC. WASH. 

95(1), 1982, pp. 27-47 

CHECKLIST OF THE FISHES OF THE CENTRAL AND 
NORTHERN APPALACHIAN MOUNTAINS 

Jay R. Stauffer, Jr., Brooks M. Burr, Charles H. Hocutt, and 

Robert E. Jenkins 



Abstract. — A table lists 398 forms and 5 intergrade populations in 28 families 
in an area on the Atlantic slope from the Susquehanna River south to the Peedee 
River, including Ohio River basin drainages from the Monongahela River in Penn- 
sylvania to the Tennessee River in Alabama and Tennessee. 



The central Appalachians harbor a diverse fish fauna that includes numerous 
endemics. Jenkins, Lachner, and Schwartz (1972), as part of a zoogeographic 
analysis of this ichthyofauna, provided a table that lists the fishes of the central 
Appalachians by river drainage and general habitat. This table has been extremely 
valuable to ichthyologists, fisheries scientists, and environmental consultants 
throughout the past decade. 

Numerous studies have substantially increased our knowledge of fish distri- 
bution throughout the central Appalachians (Hambrick et al. 1973, Hocutt and 
Hambrick 1973, Hocutt et al. 1973, Stauffer et al. 1975, Stauffer et al. 1976, 
Hocutt et al. 1978, Stauffer et al. 1978, Hendricks et al. 1979, Hocutt et al. 1979, 
Lee et al. 1980) and indicated that the original table should be updated. Moreover, 
it was thought that the addition of the Susquehanna, Licking, Green, and Ken- 
tucky rivers would enhance the usefulness of the faunal list. 

The list (Table 1) includes 398 forms and 5 intergrade populations in 28 families. 
It covers an area on the Atlantic Slope from the Susquehanna River (New York 
and Pennsylvania) south to the Peedee River (North Carolina and South Caroli- 
na). Ohio River basin drainages that are included extend from the Monongahela 
River in Pennsylvania south to the Tennessee River in Alabama and Tennessee. 

It should be noted that the list is conservative. If a question exists as to the 
current or historic presence of a species, it is not included. No attempt is made 
to distinguish species that were historically present in the drainage from those 
that currently occur. Trinomials are used only when the distribution of subspecies 
could be accurately determined. 

The authors appreciate the encouragement of Dr. E. Lachner, who recognized 
the need for a revision of the original faunal list. 



28 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Fishes of the central and northern Appalachian drainages and their general habitat. X = 
category most frequently inhabited. Drainage occurrence: E = endemic, N = native, NP = probably 
present-native, NI = regarded as native but possibly introduced, I = introduced, IP = probably or 
possibly introduced, Ma = euryhaline or diadromous. Native extralimital distribution: So = south on 
Atlantic slope. No = north on Atlantic slope, O = predominantly Ohio basin form, M = lower and/or 
central Mississippi basin, G = Gulf of Mexico slope. 



Drainage occurrence 



Atlantic Slope 



Habitat 















13 




(U 


<u 






C 

3 


c 
D 


c 

B 
c 
o 


> 

s 


e 

55 


u 




X 




X 


X 




X 


X 




X 


X 






X 






X 


X 


X 


X 






X 


X 




X 






X 


X 


X 


X 




X 


X 




X 


X 






X 


X 


X 


X 






X 


X 



u 



PETROMYZONTIDAE 

Ichthyomyzon bdellium 
Ichthyomyzon castaneus 
Ichthyomyzon fossor 
Ichthyomyzon gagei 
Ichthyomyzon greeleyi 
Ichthyomyzon unicuspis 
Lampetra aepyptera 
Lampetra appendix 
Petromyzon marinus 

ACIPENSERIDAE 

Acipenser brevirostrum 
Acipenser fulvescens 
Acipenser oxyrhynchus 
Scaphirhynchus platorynchus 

POLYODONTIDAE 

Polyodon spathula 

LEPISOSTEIDAE 

Lepisosteus spatula 
Lepisosteus oculatus 
Lepisosteus osseus 
Lepisosteus platostomus 

AMIIDAE 
Amia calva 

ANGUILLIDAE 
Anguilla rostrata 

CLUPEIDAE 

Alosa aestivalis 
Alosa alabamae 
Alosa chrysochloris 
Alosa mediocris 
Alosa pseudoharengus 
Alosa sapidissima 
Dorosoma cepedianum 
Dorosoma petenense 

HIODONTIDAE 

Hiodon alosoides 
Hiodon tergisus 

SALMONIDAE 

Coregonus artedii 
Coregonus clupeaformis 



X X 



X 
X 
X 
X 



X 
X 

X 
X 



X 



X 



X 
X 
X 
X 



X 
X 
X 
X 



X 



X 
X 




X 
X 




X 


X 


X 




X 


X 


X 


X 


X 
X 


X 


X 
X 


X 


X 




X 




X 




X 





NNNPNNNNN 
N N N N N 
Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 



Ma Ma 
Ma Ma Ma Ma 



Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 



NNNNNNNNPNN 



Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 



Ma Ma Ma Ma I Ma Ma Ma Ma Ma 



Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 
Ma Ma Ma Ma I Ma Ma Ma Ma Ma 
Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 
Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 
I I I I I 



IP 
IP 



VOLUME 95, NUMBER 1 



29 



Table 1. — Continued. 



JS ^ 



Drainage occurrence 



Ohio Basin 



^ — i> 



Native 
extralimital 
distribution 



PETROMYZONTIDAE 

Ichthyomyzon bdellium 
Ichthyomyzon castaneus 
Ichthyomyzon fossor 
Ichthyomyzon gagei 
Ichthyomyzon greeleyi 
Ichthyomyzon unicuspis 
Lampetra aepyptera 
Lampetra appendix 
Petromyzon marinus 

ACIPENSERIDAE 

Acipenser brevirostrum 
Acipenser fulvescens 
Acipenser oxyrhynchus 
Scaphirhynchus platorynchus 

POLYODONTIDAE 

Polyodon spathula 

LEPISOSTEIDAE 

Lepisosteus spatula 
Lepisosteus oculatus 
Lepisosteus osseus 
Lepisosteus platostomus 

AMIIDAE 
Amia calva 

ANGUILLIDAE 
Anguilla rostrata 

CLUPEIDAE 

Alosa aestivalis 
Alosa alabamae 
Alosa chrysochloris 
Alosa mediocris 
Alosa pseudoharengus 
Alosa sapidissima 
Dorosoma cepedianum 
Dorosoma petenense 

HIODONTIDAE 

Hiodon alosoides 
Hiodon tergisus 

SALMONIDAE 

Coregonus artedii 
Coregonus clupeaformis 



NP N N NP NP N N N N 

N 
N NP N 

N N N N 

NP NP N N N 



N 
N 

N 
N 
N 



NNNNINNNNNNNN 



N 



N NP N N N 



N 



N NP NP N 



N No 
NoSo 

N 
N 



O 
GM 

GM 
O 
M 

GM 
M 



GM 
GM 



N 


N 


N 




N 




N 


N 


N 
Ma 


N 

Ma 
Ma 


N 

Ma 
Ma 




GM 

GM 
GM 


Ma 


Ma 


Ma 




Ma 


Ma 


Ma 


Ma 


Ma 
Ma 


Ma 
Ma 


Ma 
Ma 


NoSo 


GM 
GM 








I 






N 


N 


N 


N 


N 


NoSo 


GM 


Ma 


Ma 


Ma 


Ma 




Ma 


Ma 


Ma 


Ma 


Ma 
N 


Ma 

N 


NoSo 
NoSo 


GM 
GM 






Ma 


I 




Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


NoSo 

No 
NoSo 


GM 


Ma 




Ma 




Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


NoSo 


GM 






I 


I 








I 


Ma 


Ma 


Ma 




GM 


N 










N 


NP 


N 


N 


N 


N 




M 


N 












N 


N 


N 


N 


N 




GM 



No 



30 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table I. — Continued. 



Habitat 



Drainage occurrence 



Atlantic Slope 



^ -2 



^ oi 



Oncorhynchus kisutch 
Oncorhynchus nerka 
Salmo gairdnen 
Salmo trutta 
Salvelinus fontinalis 
Salvelinus namaycush 

OSMERIDAE 

Osmerus mordax 

UMBRIDAE 

Umbra limi 
Umbra pygmaea 

ESOCIDAE 

Esox a. americanus 

Esox a. vermiculatus 

Esox lucius 

Esox masquinongy 

Esox niger 

Esox reicherti 

CYPRINIDAE 

Campostoma a. anomalum 
Campostoma oligolepis 
Carassius auratus 
Clinostomus elongatus 
Clinostomus f. funduloides 
Clinostomus f. estor 
Clinostomus f. subsp. 
Couesius plumbeus 
Ctenopharyngodon idella 
Cyprinus carpio 
Ericymba buccata 
Exoglossum laurae 
Exoglossum maxillingua 
Hemitremia flammea 
Hybognathus hayi 
Hybognathus n. nuchalis 
Hybognathus n. regius 
Hybopsis aestivalis hyostoma 
Hybopsis amblops 
Hybopsis cahni 
Hybopsis d. dissimilis 
Hybopsis hypsinotus 
Hybopsis i. insignis 
Hybopsis i. eristigma 
Hybopsis labrosa 
Hybopsis monacha 
Hybopsis storeriana 
Hybopsis x-punctata 



X 
X 



X 
X 



X X 
X X 











I 








I 


I 


I 


I 


I 


I 


I 


I 


I 


I 


I 


I 


N 


N 


NI 


N 


N 

I 



I 
I 

X X X IP 



NI NI 

X X 

XXNNNNNNNNNN 

XXNNNNNNNNNN 
X X 

I I 
XX II II 

XXNNNNNNNNNN 

I 





X 


X 




X 


X 




X 


X 




X 


X 




X 








X 




X 








X 




X 


X 






X 
X 




X 


X 




X 


X 


X 






X 








X 






X 


X 




X 


X 




X 






X 






X 






X 








X 


X 






X 


X 




X 






X 


X 




X 


X 




X 


X 






X 




X 


X 






X 






X 


X 




X 




X 


X 






X 






X 






X 






X 






X 






X 






X 


X 




X 






X 






X 






X 






X 




X 


X 




X 








X 






X 





N N 
I 



I I I I II 



N 



N 



N N 
I 



I I II 

NI 
XNNNNNNNNNN 



IP 

I I I 

N N N 



N N N N N N 



NNNNNNNNNN 



VOLUME 95, NUMBER 1 



31 



Table 1. — Continued. 



Drainage occurrence 



Ohio Basin 



Native 
extralimital 
distribution 



iH 2: 



a S 



^ _2 0) 



Oncorhynchus kisutch 
Oncorhynchus nerka 
Salmo gairdneri 
Salmo trutta 
Salvelinus fontinalis 
Salvelinus namaycush 



I 




















I 




I 


I 


I 


I 


I 


I 


I 


I 


I 


I 


I 




I 


I 


I 


I 


I 




I 


I 


I 




I 




N 


I 


IP 


N 


N 








IP 


I 


N 


NoSo 

No 



OSMERIDAE 


























Osmerus mordax 


I 
























UMBRIDAE 


























Umbra limi 




















N 




M 


Umbra pygmaea 






















NoSo 


G 


ESOCIDAE 


























Esox a. americanus 


I 




















NoSo 


G 


Esox a. vermiculatus 










N 


N 


N 


N 


N 


N 




GM 


Esox lucius 


I 








I 




I 






I 




M 


Esox masquinongy 


N 


N 


N 


IP 


I 


N 


N 


N 


N 


N 






Esox niger 


I 




I 


I 










N 


N 


NoSo 


GM 


Esox reicherti 


























CYPRINIDAE 


























Campostoma a. anomalum 


N 


N 


N 


N N 


N 


N 


N 






N 


NoSo 





Campostoma oligolepis 
















N 


N 


N N 




GM 


Carassius auratus 


I 




I 






I 


I 


I 


I 


I I 






Clinostomus elongatus 


N 










N 










No 


M 


Clinostomus f. funduloides 


N 






N 


N 






N 




NI 


So 





Clinostomus f. estor 


















N 


N 






Clinostomus f. subsp. 




















N 


So 




Couesius plumbeus 






















No 




Ctenopharyngodon idella 


















I 








Cyprinus carpio 


I 


I 


I 


I I 


I 


I 


I 


I 


I 


I I 






Ericymba buccata 


N 


N 


N 


N N 


N 


N 


N 


N 


N 


N N 


No 


GM 


Exoglossum laurae 


NI 






N 



















Exoglossum maxillingua 








NI 














No 




Hemitremia flammea 


















N 


N 




G 


Hybognathus hayi 
















N 








GM 


Hybognathus n. nuchalis 










N 






N 


N 


N 




GM 


Hybognathus n. regius 






















NoSo 




Hybopsis aestivalis hyostoma 






N 


N 


N 


N 


N 


N 


N 


N 




GM 


Hybopsis amblops 


N 


N 


N 




N 


N 


N 


N 


N 


N 




M 


Hybopsis cahni 




















E 






Hybopsis d. dissimilis 


N 


N 


N 


N 


N 




N 


N 


N 


N 







Hybopsis hypsinotus 






















So 




Hybopsis i. insignis 


















N 


N 






Hybopsis i. eristigma 




















E 






Hybopsis labrosa 






















So 




Hybopsis monacha 




















E 






Hybopsis storeriana 


N 


N 


N 




N 


N 


N 


N 


N 


N 




GM 


Hybopsis x-punctata 
















N 








M 



32 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Continued. 



Drainage occurrence 









Habitat 












Atlantic 


; Slop 


e 












EX 

a 
U 


0) 

Z 




o 

c 

o 
oi 


E 


o 
>- 


o 

o 

c 
c 
ca 
j: 

o. 
o. 

oi 


o 

a 

B 
o 

1 


§ 




-a 

c 

1 

o 

-J 


■§ 

D 


0) 

c 

B 
c 
o 


> 
S 


s 


u 


c 

JS 
<u 

3 
O" 

c« 
3 

on 


Hybopsis sp. cf. zanema 


X 








X 




N 


N 


















Leuciscus idus 






























I 




Nocomis biguttatus 




X 






X 


X 






















Nocomis effusus 




X 






X 


X 






















Nocomis I. leptocephalus 




X 


X 




X 


X 


N 


N 


N 


N 


N 


N 


N 


IP 


IP 




Nocomis I. bellicus 




X 






X 


X 






















Nocomis I. interocularis 




X 


X 




X 


X 






















Nocomis micropogon 




X 


X 


X 


X 














N 


N 


N 


N 


N 


Nocomis platyrhynchus 




X 


X 


X 


X 
























Nocomis raneyi 




X 




X 


X 








N 


N 


N 


NI 










Notemigonus crysoleucas 


X 


X 






X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


Notropis albeolus 




X 






X 


X 




N 


N 


N 


N 












Notropis alborus 




X 






X 


X 


N 


N 






N 












Notropis altipinnis 


X 


X 








X 


N 


N 


N 


N 


N 












Notropis amnis 


X 






X 


X 
























Notropis amoenus 


X 


X 




X 


X 




NI 


N 


N 


N 


N 


N 


N 


N 


N 


N 


Notropis analostanus 




X 






X 




N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


Notropis ardens 




X 






X 


X 




IP 


N 


N 


N 


N 






IP 




Notropis ariommus 




X 




X 


X 
























Notropis atherinoides 


X 


X 




X 


X 
























Notropis baileyi 




X 








X 






















Notropis bellus 




X 






X 


X 






















Notropis bifrenatus 


X 


X 








X 






N 


NP 


N 


N 


N 


N 


N 


N 


Notropis blennius 


X 






X 


























Notropis boops 




X 






X 
























Notropis buchanani 


X 


X 




X 


X 
























Notropis camurus 


X 


X 




X 


X 






— - ._ 


















Notropis cerasinus 




X 






X 


X 




IP 






N 


IP 










Notropis chalybaeus 


X 










X 


N 


N 


N 


N 


N 








N 




Notropis chiliticus 




X 






X 


X 


N 








N 












Notropis chrysocephalus 




X 


X 




X 


X 






















Notropis coccogenis 




X 


X 




X 


X 






















Notropis cornutus 




X 


X 




X 


X 












N 


N 


N 


N 


N 


Notropis c. cummingsae 


X 








X 




N 


N 


N 


N 














Notropis d. dorsalis 




X 








X 






















Notropis emiliae 


X 


X 




X 


X 
























Notropis fumeus 


X 








X 


X 






















Notropis galacturus 




X 


X 




X 


X 






















Notropis heterodon 




X 






X 


X 




















N 


Notropis heterolepis 




X 






X 


X 




















N 


Notropis hudsonius 


X 


X 




X 


X 




N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


Notropis leuciodus 




X 


X 




X 


X 






















Notropis lirus 




X 






X 
























Notropis lutipinnis 




X 






X 


X 






















Notropis lutrensis 


X 


X 




X 


X 


X 










I 












Notropis maculatus 


X 








X 


X 


N 


N 


















Notropis mekistocholas 




X 






X 






E 


















Notropis niveus 




X 






X 




N 


N 


N 


N 














Notropis petersoni 


X 








X 




N 


N 



















VOLUME 95, NUMBER 1 



33 



Table 1. — Continued. 



i2 ^ 



Drainage occurrence 



Ohio Basin 



_tu _ra jj 



Native 
extralimital 
distribution 



a 



Hybopsis sp. cf. zanema 
Leuciscus idus 
Nocomis biguttatus 
Nocomis effusus 
Nocomis I. leptocephalus 
Nocomis I. bellicus 
Nocomis I. interocularis 
Nocomis micropogon 
Nocomis platyrhynchus 
Nocomis raneyi 
Notemigonus crysoleucas 
Notropis albeolus 
Notropis alborus 
Notropis altipinnis 
Notropis amnis 
Notropis amoenus 
Notropis analostanus 
Notropis ardens 
Notropis ariommus 
Notropis atherinoides 
Notropis baileyi 
Notropis bellus 
Notropis bifrenatus 
Notropis blennius 
Notropis boops 
Notropis buchanani 
Notropis camurus 
Notropis cerasinus 
Notropis chalybaeus 
Notropis chiliticus 
Notropis chrysocephalus 
Notropis coccogenis 
Notropis cornutus 
Notropis c. cummingsae 
Notropis d. dorsalis 
Notropis emiliae 
Notropis fume us 
Notropis galacturus 
Notropis heterodon 
Notropis heterolepis 
Notropis hudsonius 
Notropis leuciodus 
Notropis lirus 
Notropis lutipinnis 
Notropis lutrensis 
Notropis maculatus 
Notropis mekistocholas 
Notropis niveus 
Notropis petersoni 



NI 



M 



N N 



N 



N 



So 























N 




G 






















IP 


So 




N 


N N 


E 


N 


N 


N 


N 




N 


N 


N 


No 





N 


N 


IP 

N 








N 


N 
N 


N 
N 


I 


N 
N 


NoSo 

So 
So 

No 
No 


GM 
GM 






N 




N 


N 


N 


N 


N 


N 


N 







N 


N 






N 




N 


N 


N 




N 







N 


N 




N 


N 


N 


N 


N 


N 


N 


N 
N 
N 


No 
No 


GM 
G 
G 


N 


N 




N 


N 


N 
N 


N 
N 


N 
N 


N 
N 




N 
N 




M 
M 


N 


N 


N 
IP 




NP 


N 


N 


N 


N 




N 
N 


NoSo 


GM 
GM 


N 


N N 


IP 
IP 


N 


N 


N 


N 


N 


N 


N 


N 
N 


So 


GM 


N 


N 




















No 
So 


M 
G 


N 




NI 




N 






N 
N 


N 
N 
N 

N 


IP 


N 
N 
N 


So 

No 
No 


GM 

GM 

M 

M 


N 




IP 
IP 










N 


N 




N 
N 
IP 


NoSo 
So 

So 

So 

So 

So 


M 


G 

GM 

G 



34 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Continued. 



Habitat 






Drainage occurrence 



Atlantic Slope 



Notropis photogenis 




X 




X 


X 




Notropis procne procne 


X 


X 






X 


X 


Notropis p. longiceps 


X 


X 






X 


X 


Notropis pyrrhomelas 




X 






X 




Notropis rubellus 




X 


X 




X 


X 


Notropis rubricroceus 






X 




X 


X 


Notropis scabriceps 




X 




X 


X 




Notropis scepticus 




X 






X 




Notropis semperasper 




X 






X 




Notropis shumardi 


X 






X 






Notropis spectrunculus 














spectrunculus 






X 




X 


X 


Notropis spectrunculus subsp. 






X 




X 


X 


Notropis spilopterus 














spilopterus 




X 




X 


X 




Notropis stilbius 




X 






X 




Notropis stramineus stramineus 




X 








X 


Notropis telescopus 




X 


X 




X 


X 


Notropis umbratilus 














cyanocephalus 


X 


X 






X 


X 


Notropis volucellus 




X 




X 


X 




Notropis whipplei 




X 




X 


X 




Notropis sp. (paleband shiner) 




X 






X 




Notropis sp. (sawfin shiner) 




X 




X 


X 




Notropis sp. {longirostris 














group) 


X 








X 


X 


Phenacobius crassilabrum 






X 




X 




Phenacobius mirabilis 


X 


X 




X 


X 




Phenacobius teretulus 




X 






X 




Phenacobius uranops 




X 




X 


X 




Phoxinus cumberlandensis 




X 








X 


Phoxinus eos 












X 


Phoxinus erythrogaster 




X 








X 


Phoxinus o. ore as 




X 


X 






X 


Phoxinus o. subsp. 




X 


X 






X 


Pimephales notatus 




X 




X 


X 


X 


Pimephales promelas 


X 


X 






X 


X 


Pimephales vigilax 


X 


X 




X 


X 




Rhinichthys atratulus 




X 


X 




X 


X 


Rhinichthys cataractae 




X 


X 




X 


X 


Semotilus atromaculatus 




X 


X 




X 


X 


Semotilus corporalis 




X 




X 


X 




Semotilus lumbee 














Semotilus margarita 




X 


X 






X 


Tinea tinea 















X X N N N N N 
N 



N N 



IP 



N N N N 



N N 



IP 



N N N N N 



N N N N N 



N N N NI 



N N 



N N N N N IP IP 



IP 



N 



IP 


IP 




IP 


N 


N 


I 




I 




I 


I 


N 


N 


N 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 




N 


N 


N 


N 

N 
I 


N 
N 



CATOSTOMIDAE 

Carpiodes carpio 
Carpiodes cyprinus 



X 

X X 



N 



N N 



N N 



VOLUME 95, NUMBER 1 



35 



Table 1. — Continued. 



4? ^ 



Drainage occurrence 



Ohio Basin 



O 



TO ro aj 



Native 
extralimital 
distribution 



Notropis photogenis 
Notropis procne procne 
Notropis p. longiceps 
Notropis pyrrhomelas 
Notropis rubellus 
Notropis rubricroceus 
Notropis scabriceps 
Notropis scepticus 
Notropis semperasper 
Notropis shumardi 
Notropis spectrunculus 

spectrunculus 
Notropis spectrunculus subsp. 
Notropis spilopterus 

spilopterus 
Notropis stilbius 
Notropis stramineus stramineus 
Notropis telescopus 
Notropis umbratilus 

cyanocephalus 
Notropis volucellus 
Notropis whipplei 
Notropis sp. (paleband shiner) 
Notropis sp. (sawfin shiner) 
Notropis sp. {longirostris 

group) 
Phenacobius crassilabrum 
Phenacobius mirabilis 
Phenacobius teretulus 
Phenacobius uranops 
Phoxinus cumberlandensis 
Phoxinus eos 
Phoxinus erythrogaster 
Phoxinus o. oreas 
Phoxinus o. subsp. 
Pimephales notatus 
Pimephales promelas 
Pimephales vigilax 
Rhinichthys atratulus 
Rhinichthys cataractae 
Semotilus atromaculatus 
Semotilus corporalis 
Semotilus lumbee 
Semotilus margarita 
Tinea tinea 



NNNNNNNNNN 



N 



N N N 



IP 

N 
IP 
E 



N N N N N 



N N N 
N 



N 



N 



NP 

























E 
























E 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 
N 


N 


N 


N 
IP 


N 
IP 


N 


N 


N 


N 




N 
N 




N 
N 






N 




N 


N 


N 


N 


N 


N 




N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 




N 


N 




N 


N 


N 


N 


N 


N 
N 
N 


N 


N 
N 
N 

N 
E 






N 




N 


N 


N 


N 


N 


N 




N 



N 



No 
So 

So 

No 
So 

So 



No 



O 



M 



GM 



M 

GM 

M 

M 

M 
GM 
GM 



M 
GM 



























No 


M 






N 


N 


N 


N 


N 


N 


N 


N 


N 


N 
IP 
E 




M 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


No 


GM 


I 


I 


IP 


I 




N 


N 


N 


N 


N 


NI 


N 


No 


GM 


N 




N 






N 


N 


N 


N 


N 


N 


N 




GM 


N 
N 




N 
N 


N 
N 


N 


N 


N 


N 


N 


N 


N 


N 
N 


NoSo 
NoSo 


M 



N 
N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


NoSo 
No 

No 


GM 

M 



CATOSTOMIDAE 

Carpiodes carpio 
Carpiodes cyprinus 



N N N 



N 



N N N N N 
N N N N N 



N GM 

N NoSo GM 



36 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Continued. 



Habitat 



Drainage occurrence 



Atlantic Slope 



^ 4 



S ^ 



o 


a 


o 




55 




CL, 


n! 
U 




c3 


o 




o 




o 


3 

on 


X 






X 








N 






















X 




X 


X 
























X 


X 




X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 




X 


























X 


X 






X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 






X 


X 






















X 


X 


X 




X 
X 


X 


N 


N 


N 


N 


N 














X 


X 




X 


X 


N 




N 


N 


N 


N 


N 


N 


N 


N 




X 






X 


X 










E 












X 






X 






IP 




















X 






X 


























X 


X 




X 


X 


X 






















X 


X 




X 


X 




N 


N 


















X 


X 
X 
X 




X 


X 
X 
X 


X 


N 


N 


N 


N 


N 
E 












X 


X 
X 
X 
X 
X 


X 


X 

X 
X 


X 
X 
X 
X 
X 


X 
X 






N 


N 


N 

N 
E 


NI 
N 






IP 




X 


X 




X 


X 




N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 




X 


X 
























X 


X 




X 


X 




N 


N 


N 


N 


N 














X 


X 




X 


X 










N 


N 




IP 


N 






X 






X 




N 


N 




















X 


X 




X 


X 


IP 





















Carpiodes velifer 
Catostomus catostomus 
Catostomus commersoni 
Cycleptus elongatus 
Erimyzon o. oblongus 
Erimyzon o. claviformis 
Erimyzon sucetta 
Hypentelium etowanum 
Hypentelium nigricans 
Hypentelium roanokense 
Ictiobus bubalus 
Ictiobus cyprinellus 
Ictiobus niger 
Lagochila lac era 
Minytrema melanops 
Moxostoma anisurum 
Moxostoma ariommum 
Moxostoma atripinne 
Moxostoma carinatum 
Moxostoma cervinum 
Moxostoma duquesnei 
Moxostoma erythrurum 
Moxostoma hamiltoni 
Moxostoma m. macrolepidotum 
Moxostoma m. breviceps 
Moxostoma pappillosum 
Moxostoma rhothoecum 
Moxostoma robustum 
Moxostoma rupiscartes 

ICTALURIDAE 

Ictalurus brunneus 
Ictalurus catus 
Ictalurus furcatus 
Ictalurus melas 
Ictalurus natalis 
Ictalurus nebulosus 
Ictalurus platycephalus 
Ictalurus punctatus 
Noturus baileyi 
Noturus elegans 
Noturus eleutherus 
Noturus exilis 
Noturus flavipinnis 
Noturus flavus 
Noturus furiosus 
Noturus gilberti 
Noturus gyrinus 
Noturus insignis 
Noturus leptacanthus 



X X 



X X 



N N 



IP 



X 


X 


X 






N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 




X 






I 


I 


I 






I 




I 






X 


X 


X 


X 


X 


I 








I 








I 




X 


X 




X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 


X 


X 




N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 


X 


X 




N 


N 


NI 




N 












X 


X 
X 
X 
X 
X 
X 


X 


X 
X 
X 
X 
X 
X 




IP 


IP 


IP 


IP 


IP 


IP 


IP 


IP 


IP 


IP 




X X 


X 


X 






















X 


X 
X 




X 
X 








N 


N 


N 


NI 










X 






X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 






X 


X 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


X 


X 




X 

























VOLUME 95, NUMBER 1 



37 



Table 1. — Continued. 



•2 > 



c ■- 



Drainage occurrence 



Ohio Basin 



i^ a 



12 s 



Native 
extralimital 
distribution 



Carpiodes velifer 
Catostomus catostomus 
Catostomus commersoni 
Cycleptus elongatus 
Erimyzon o. oblongus 
Erimyzon o. claviformis 
Erimyzon sucetta 
Hypentelium etowanum 
Hypentelium nigricans 
Hypentelium roanokense 
Ictiobus bubalus 
Ictiobus cyprinellus 
Ictiobus niger 
Lagochila lacera 
Minytrema melanops 
Moxostoma anisurum 
Moxostoma ariommum 
Moxostoma atripinne 
Moxostoma carinatum 
Moxostoma cervinum 
Moxostoma duquesnei 
Moxostoma erythrurum 
Moxostoma hamiltoni 
Moxostoma m. macrolepidotum 
Moxostoma m. breviceps 
Moxostoma pappillosum 
Moxostoma rhothoecum 
Moxostoma robustum 
Moxostoma rupiscartes 

ICTALURIDAE 

Ictalurus brunneus 
Ictalurus catus 
Ictalurus fur cat us 
Ictalurus melas 
Ictalurus natalis 
Ictalurus nebulosus 
Ictalurus platycephalus 
Ictalurus punctatus 
Noturus baileyi 
Noturus elegans 
Noturus eleutherus 
Noturus exilis 
Noturus flavipinnis 
Noturus flavus 
Noturus furiosus 
Noturus gilberti 
Noturus gyrinus 
Noturus insignis 
Noturus leptacanthus 



N 




N 




N 


N 


N 


N 


N 


N 




N 


So 


GM 


N 
























No 


GM 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


NoSo 


M 


NP 


NP 


NP 






NP 


N 


NP 


N 

N 
N 


N 
N 




N 
N 
NI 


NoSo 
So 


GM 

GM 

GM 

G 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


N 


NoSo 


GM 


N 




N 




N 


N 


N 


N 


N 


N 




N 




GM 












N 


N 


N 


N 


N 




N 




GM 






N 








NP 


NP 
N 


N 
NP 


N 
N 




N 
N 




GM 
M 




N 


N 




N 


N 


N 


N 


N 


N 


N 


N 


So 


GM 


N 


NP 


N 




N 


N 


N 


N 


N 


N 


N 


N 


So 


M 



N N N 

N N N 

N N N IP 



N N N 



N N N N N N 

N N N N N N 
N N N N N N 



N N N N N N 



N 



N N N N N N 

NN IP NNNNN 
NNNNINNNNNN 
N N NI N N N N 

NNNNINNNNNN 

N 
NNNNN 
N N N 

NN NINNNNNN 



N 



GM 



N N N 



IP 



N 



N N GM 

N N GM 

NoSo M 

N O 
So 

So 

So G 



So G 

NoSo G 

N GM 

N N GM 

N N NoSo GM 

IP N NoSo GM 

So 

N N So GM 
E 

N O 

N M 

N M 
E 

N M 



N NoSo GM 
IP NoSo 
NI So G 



38 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Continued. 



Habitat 



Drainage occurrence 



Atlantic Slope 



^ c 3 







^ 

a 




o 


60 

s 


0) 

Sri 


u 

u 

u 


Oh 




Noturus miurus 






X 






X 








Noturus nocturnus 






X 




X 


X 








Noturus stigmosus 




X 


X 






X 








Noturus stanauli 






X 




X 


X 








Noturus sp. cf. leptacc 


inthus 


X 








X 




N 


N 


Pylodictis olivaris 




X 


X 




X 


X 




I 


I 



AMBLYOPSIDAE 

Amblyopsis spelaea 
Chologaster agassizi 
Chologaster cornuta 
Typhlichthys subterraneus 

APHREDODERIDAE 

Aphredoderus sayanus 
PERCOPSIDAE 

Percopsis omiscomaycus 

GADIDAE 

Lota lota 

CYPRINODONTIDAE 

Fundulus albolineatus 
Fundulus catenatus 
Fundulus d. diaphanus 
Fundulus heteroclitus 
Fundulus lineolatus 
Fundulus notatus 
Fundulus olivaceus 
Fundulus rathbuni 
Fundulus stellifer 
Fundulus waccamensis 
Fundulus sp. 

POECILIIDAE 

Gambusia a. affinis 
Gambusia a. holbrooki 
Heterandria formosa 

ATHERINIDAE 

Labidesthes sicculus 
Menidia extensa 

GASTEROSTEIDAE 

Apeltes quadracus 
Culaea inconstans 

COTTIDAE 

Cottus baileyi 
Cot t us b. bairdi 



X X 
X 

X 



X X 



X 



X X N N N N N 



XXNNNNNNNNN 



X 



N N 



NI 





X 




X 
























X 


X 


X 






















X 


X 


X 




Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


X 








Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


X 
X 


X 
X 


X 
X 
X 


X 


N 


N 


N 


N 


N 














X 


X 


X 


N 


N 


N 




N 














X 


X 


X 






















X 


X 




X 


N 








NP 












X 


X 


X 


X 






















X 


X 


X 


X 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 


Ma 




X 






X 


NP 


N 



















X X 
X X 



X X 
X X 



N N 



Ma Ma Ma IP 
IP 



N N N 



VOLUME 95, NUMBER 1 



39 



Table 1. — Continued. 



-2 ^ 



Drainage occurrence 



Ohio Basin 



O 



Native 
extralimital 
distribution 



Noturus miurus 
Noturus nocturnus 
Noturus stigmosus 
Noturus stanauli 
Noturus sp. cf. leptacanthus 
Pylodictis olivaris 

AMBLYOPSIDAE 

Amblyopsis spelaea 
Chologaster agassizi 
Chologaster cornuta 
Typhlichthys subterraneus 

APHREDODERIDAE 

Aphredoderus say anus 

PERCOPSIDAE 

Percopsis omiscomaycus 

GADIDAE 

Lota lota 

CYPRINODONTIDAE 

Fundulus albolineatus 
Fundulus catenatus 
Fundulus d. diaphanus 
Fundulus heteroclitus 
Fundulus lineolatus 
Fundulus notatus 
Fundulus olivaceus 
Fundulus rathbuni 
Fundulus stellifer 
Fundulus waccamensis 
Fundulus sp. 

POECILIIDAE 

Gambusia a. affinis 
Gambusia a. holbrooki 
Heterandria for mas a 

ATHERINIDAE 

Labidesthes sicculus 
Menidia extensa 

GASTEROSTEIDAE 

Apeltes quadracus 
Culaea inconstans 

COTTIDAE 

Cottus baileyi 
Cottus b. bairdi 



N N 



N 


N 


N 


N 


N 


N 


N 


N 


N 


GM 






N 




N 


N 


N 




N 


GM 


N 




N 


N 


N 


N 








M 



NNNNNNNNNNNN 



N 



GM 



N N N 



N N N 



O 

N N N M 

So 
N N N GM 

N N 



N 



NI NI 



N N 



N N N N 
N 



N 



N 


NoSo 


GM 




No 


M 




No 


M 


E 






N 


No 


M 




So 


G 


N 




GM 


N 
IP 


So 


GM 
G 



N 



IP IP Ma Ma IP Ma GM 

NoSo G 
So G 

NNN I NNNNNNNN SoGM 



NNNNNNNN 



N 



N No 



M 



40 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Continued. 



^ ^ 



Habitat 



5 2 



Drainage occurrence 



Atlantic Slope 



U 



Cottus bairdi subsp. 
Coitus c. carolinae 
Cottus carolinae subsp. 
Cottus cognatus 
Cottus girardi 
Cottus sp. (smoky sculpin) 
Cottus sp. 

PERCICHTHYIDAE 

Morone americana 
M or one chrysops 
Morone mississippiensis 
Morone saxatilis 

CENTRARCHIDAE 

Acantharcus pomotis 
Ambloplites cavifrons 
Ambloplites r. rupestris 
Centrarchus macropterus 
Elassoma evergladei 
Elassoma zonatum 
Elassoma sp. 
Elassoma sp. 
Enneacanthus chaetodon 
Enneacanthus gloriosus 
Enneacanthus obesus 
Lepomis auritus 
Lepomis cyanellus 
Lepomis gibbosus 
Lepomis gulosus 
Lepomis humilis 
Lepomis macrochirus 
Lepomis marginatus 
Lepomis megalotis 
Lepomis microlophus 
Lepomis punctatus 
Micropterus coosae 
Micropterus d. dolomieui 
Micropterus p. punctulatus 
Micropterus s. salmoides 
Pomoxis annularis 
Pomoxis nigromaculatus 

PERCIDAE 

Ammocrypta asprella 
Ammocrypta clara 
Ammocrypta pellucida 
Ammocrypta vivax 
Etheostoma acuticeps 
Etheostoma asprigene 



X 
X 
X 
X 
X 
X 
X 
X 



X 
X 
X 

X 
X 



X 




X 


X 


X 




X 


X 


X 




X 


X 


X 




X 


X 


X 


X 


X 


X 




X 


X 


X 



X 



X 
X 
X 
X 
X 
X 

X 
X 



X 
X X 
X X 
X X 
X X 



X 
X 

X 
X X 



X X 



X 



X 



X 



X X 

X X 
X 

X X 



X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 



X X 

X X 

X X 

X X 



X 



X 



X 



IP 

N 
N 
N 

E 

N 
N 
N 
N 
I 

N 
N 



X X I 



N 



N 
N N 



Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 
I I I 

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma 



N N 



N 
N 
N 
N 
I 

N 
N 



N 
N 



N N 



N 
N 
I 

N 



N N N 



N 
N 
N 
N 
I 

N 
N 



N 
N 
N 
N 
I 

N 
N 



N 
N 
N 
N 
I 

N 
N 



NI NI IP 

N N N 



I 



IP 



N 



N N N 
I I I 

N N N N 



N N N N 



IP 

N 



N 
N 
N 
I 

N 
N 



I I 



N 



I I 

NI NI 



N N N 

N NP N 

N N N 

I I 

N N 

N IP 



N 
N 



NI NI IP IP IP IP IP 

N N N N 



I I 



N 

N 
I 

N 
N 



IP IP IP 

I 

I I 



I 



N N IP IP IP IP 
II III 

N IP IP IP IP IP 



VOLUME 95, NUMBER 1 



41 



Table 1. — Continued. 



■2 ^ 



c -5 



Drainage occurrence 



Ohio Basin 



O 



C3 03 flj 



Native 
extralimital 
distribution 



Cottus bairdi subsp. 
Coitus c. carolinae 
Cottus carolinae subsp. 
Cottus cognatus 
Cottus girardi 
Cottus sp. (smoky sculpin) 
Cottus sp. 

PERCICHTHYIDAE 

Morone americana 
Morone chrysops 
Morone mississippiensis 
Morone saxatilis 

CENTRARCHIDAE 

Acantharcus pomotis 
Ambloplites cavifrons 
Ambloplites r. rupestris 
Centrarchus macropterus 
Elassoma evergladei 
Elassoma zonatum 
Elassoma sp. 

Enneacanthus chaetodon 
Enneacanthus gloriosus 
Enneacanthus obesus 
Lepomis auritus 
Lepomis cyanellus 
Lepomis gibbosus 
Lepomis gulosus 
Lepomis humilis 
Lepomis macrochirus 
Lepomis marginatus 
Lepomis megalotis 
Lepomis microlophus 
Lepomis punctatus 
Micropterus coosae 
Micropterus d. dolomieui 
Micropterus p. punctulatus 
Micropterus s. salmoides 
Pomoxis annularis 
Pomoxis nigromaculatus 

PERCIDAE 

Ammocrypta asprella 
Ammocrypta clara 
Ammocrypta pellucida 
Ammocrypta vivax 
Etheostoma acuticeps 
Etheostoma asprigene 



N 
N 



N N N 



M 



No 



N So 



























NoSo 




I 


NI 


NI 


IP 




NI 


NP 


N 


N 


N 
N 




N 
N 




GM 
GM 








I 




I 




I 


I 


I 




I 


NoSo 
NoSo 


G 
G 


N 


N 


N 


IP 


N 


N 


N 


N 


N 
N 

N 


N 
N 


N 


N 
N 

N 
E 


No 
So 
So 
So 

NuSo 
NoSo 
NoSo 


M 
M 

GM 
G 








IP 








I 




I 


I 


I 


NoSo 


G 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


N 


N 




GM 


NX 




N 


IP 






IP 






IP 




IP 


NoSo 


M 






N 


IP 






N 


N 


N 


N 


IP 


N 


So 


GM 




IP 










N 


N 


N 


N 


N 


N 




GM 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


N 


N 
N 


So 
So 


GM 
GM 


N 


N 


N 


I 


N 


N 


N 


N 


N 


N 


N 


N 




GM 


IP 










IP 


IP 


IP 


N 
N 


N 


I 


N 
N 


So 
So 


GM 

GM 

G 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


N 


N 




M 


N 


N 


N 


NI 


N 


N 


N 


N 


N 


N 


N 


N 




M 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


NI 


N 


So 


GM 


N 


N 


N 


IP 


N 


N 


N 


N 


N 


N 


NI 


N 




GM 


N 


N 


N 


IP 




N 


N 


N 


N 

N 
N 


N 

N 
N 




N 
N 


So 


GM 

GM 
GM 


N 


N 


N 




N 


N 


N 


N 


N 


N 




N 




M 
GM 



N N 



E 

N 



GM 



42 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table I. — Continued. 



Drainage occurrence 



Atlantic Slope 



Habitat 















T3 




« 


u 






C 


■a 




> 


^ 




o 


1 


o 


2 

60 

PQ 


55 


(L> 
u 

o 




X 






X 






X 






X 


X 




X 






X 


X 




X 




X 


X 


X 




X 


X 




X 


X 






X 




X 


X 




X 


X 




X 


X 



Etheostoma aquali 
Etheostoma atripinne 
Etheostoma barbouri 
Etheostoma bellum 
Etheostoma blennioides 

blennioides 
Etheostoma b. gutselli 
Etheostoma b. newmanii 
Etheostoma b.: newmanii x 

blennioides 
Etheostoma b.: n. x gutselli 
Etheostoma blennius 
Etheostoma boschungi 
Etheostoma caeruleum 
Etheostoma camurum 
Etheostoma chlorobranchium 
Etheostoma chlorosomum 
Etheostoma cinereum 
Etheostoma collis collis 
Etheostoma c. lepidinion 
Etheostoma duryi 
Etheostoma etnieri 
Etheostoma flabellare 
Etheostoma fusiforme fusiforme 
Etheostoma f. barratti 
Etheostoma gracile 
Etheostoma histrio 
Etheostoma jessiae 
Etheostoma kanawhae 
Etheostoma kennicotti 
Etheostoma longimanum 
Etheostoma luteovinctum 
Etheostoma m. maculatum 
Etheostoma m. sanguifluum 
Etheostoma m. vulneratum 
Etheostoma mariae 
Etheostoma meadiae 
Etheostoma microlepidum 
Etheostoma neopterum 
Etheostoma n. nigrum 
Etheostoma n. susanae 
Etheostoma n.: nigrum x 

susanae 
Etheostoma obeyense 
Etheostoma olivaceum 
Etheostoma o. olmstedi 
Etheostoma o.: o. x 

atromaculatum 
Etheostoma o. atromaculatum 



X 



X X 





X 




X 


X 




X 






X 




X 




X 


X 




X 


X X 








X 


X 




X 


X 




X 
X 




X 


X 






X 


X 


X 






X 




X 




X 


X 




X 




X 


X 




X 


X 


X 


X 


X 








X 


X 








X 


X 






X 


X 


X 


X 
X 
X 




X 
X 
X 






X 




X 


X 




X 


X 


X 


X 




X 




X 






X 




X 






X 




X 






X 


X 


X 




X 


X 




X 


X 




X 




X 


X 




X 




X 




X 


X 






X 




X 




X 


X 




X 






X 




X 




X 


X 




X 




X 


X 




X 






X 


X 


X 




X 


X 



N N 



NI 



N 



N N 



N 



N N N N N N 
N N N N N 
N 



N N 
N 



N N N N 



N N N N N 



N 



X 



X 



N N N N N N 



VOLUME 95, NUMBER 1 



43 



Table 1. — Continued. 



-a » 



c ■= 



Drainage occurrence 



Ohio Basin 



a 






Native 
extralimital 
distribution 



Etheostoma aquali 
Etheostoma atripinne 
Etheostoma barbouri 
Etheostoma bellum 
Etheostoma blennioides 

blennioides 
Etheostoma b. gutselli 
Etheostoma b. newmanii 
Etheostoma b.: newmanii x 

blennioides 
Etheostoma b.: n. x gutselli 
Etheostoma blennius 
Etheostoma boschungi 
Etheostoma caeruleum 
Etheostoma camurum 
Etheostoma chlorobranchium 
Etheostoma chlorosomum 
Etheostoma cinereum 
Etheostoma collis collis 
Etheostoma c. lepidinion 
Etheostoma duryi 
Etheostoma etnieri 
Etheostoma flabellare 
Etheostoma fusiforme fusiforme 
Etheostoma f. barratti 
Etheostoma gracile 
Etheostoma histrio 
Etheostoma jessiae 
Etheostoma kanawhae 
Etheostoma kennicotti 
Etheostoma longimanum 
Etheostoma luteovinctum 
Etheostoma m. maculatum 
Etheostoma m. sanguifluum 
Etheostoma m. vulneratum 
Etheostoma mariae 
Etheostoma meadiae 
Etheostoma microlepidum 
Etheostoma neopterum 
Etheostoma n. nigrum 
Etheostoma n. susanae 
Etheostoma n.: nigrum x 

susanae 
Etheostoma obeyense 
Etheostoma olivaceum 
Etheostoma o. olmstedi 
Etheostoma o.: o. x 

atromaculatum 
Etheostoma o. atromaculatum 



NNNNNNNN 



N N N N 



N 



N 



N 



N 



N 



N 



N 



NNNNNNNNNN 



NNNNNNNNNN 



E 

N 







E 


N 


N 


N 

E 

E 
E 


N 


N 


N 


N 




N 
E 


N 




N 


N 




N 



o 

M 



M 
O 

GM 



So 



N 


N 


N 


NoSo 

No 


M 






IP 


So 


GM 


N 


N 


N 




GM 


N 


N 


N 
N 




GM 


N 


N N N 




O 




N 


N 






N 


E 
E 


E 

N 
E 




O 


N 


N 


N 


No 


GM 



No 



No 



44 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table I. — Continued. 



Drainage occurrence 









Habitat 












Atlantic Slope 










u 




g 


Tar 

Roanoke 
James 
York 


1 
1 

06 


e 

o 


cd 




T3 

1 


o. 


c 
1 


> 

.SP 

PQ 


So 




1 

en 
3 
CO 


Etheostoma o.: o. x vexillare 




X 






X 


X 








N 


N 






Etheostoma o. vexillare 




X 






X 


X 










N 






Etheostoma o. maculaticeps 


X 


X 






X 


X 


N 


N 












Etheostoma osburni 




X 


X 




X 


















Etheostoma parvipinne 


X 


X 






X 


X 
















Etheostoma perlongum 


X 












E 














Etheostoma podostemone 




X 






X 


X 








E 








Etheostoma proeliare 


X 








X 


















Etheostoma rufilineatum 




X 


X 




X 


X 
















Etheostoma s. sagitta 




X 






X 


X 
















Etheostoma sagitta spilotum 




X 






X 


X 
















Etheostoma sellare 




X 






X 


X 














E 


Etheostoma serriferum 


X 








X 




N 


N 


N 


N N 








Etheostoma simoterum 




X 






X 


X 
















Etheostoma smithi 




X 








X 
















Etheostoma s. spectabile 




X 






X 


X 
















Etheostoma squamiceps 




X 






X 


X 
















Etheostoma striatulum 




X 








X 
















Etheostoma stigmaeum 




X 






X 


















Etheostoma swaini 


X 


X 






X 


















Etheostoma swannanoa 






X 




X 


X 
















Etheostoma tippecanoe 




X 




X 


X 


















Etheostoma tuscumbia 




X 








X 
















Etheostoma variatum 




X 


X 




X 


X 
















Etheostoma virgatum 




X 






X 


X 
















Etheostoma vitreum 




X 






X 








N 


N N N N 


N 


N 




Etheostoma z- zonale 




X 


X 




X 


X 














I 


Etheostoma sp. (duskytail darter) 






X 






X 
















Etheostoma sp. (Elk darter) 




X 






X 


















Etheostoma {Ulocentra) sp. 




























A — emerald darter 




X 






X 


X 
















Etheostoma {Ulocentra) sp. B 




X 






X 


X 
















Etheostoma {Ulocentra) sp. C 




























Etheostoma {Ulocentra) sp. 




























D — golden snubnose darter 




X 






X 


X 
















Etheostoma {Ulocentra) sp. 




























E — (Green River) 




X 






X 


X 
















Etheostoma {Ulocentra) sp. 




























F — (Barren River) splendid 




























darter 




X 






X 


X 
















Perca flavescens 


X 


X 




X 


X 




NI 


NI 


NI 


NI NI NI NI 


NI 


NI 


NI 


Percina aurantiaca 




X 


X 




X 


















Percina burtoni 




X 






X 


















Percina c. cap rode s 




X 


X 


X 


X 


















Percina c. semifasciata 




X 




X 


X 














N 


N 


Percina copelandi 




X 




X 


X 


















Percina crassa 




X 






X 




N 


N 












Percina e. evides 




X 


X 


X 


X 


















Percina evides subsp. 






X 




X 



















VOLUME 95, NUMBER 1 



45 



Table 1. — Continued. 



Drainage occurrence 



Ohio Basin 



-2 ^ 



TO ro fli 



Native 
extralimital 
distribution 



i^ a 



Etheostoma o.: o. x vexillare 
Etheostoma o. vexillare 
Etheostoma o. maculaticeps 
Etheostoma osburni 
Etheostoma parvipinne 
Etheostoma perlongum 
Etheostoma podostemone 
Etheostoma proeliare 
Etheostoma rufilineatum 
Etheostoma s. sagitta 
Etheostoma sagitta spilotum 
Etheostoma sellare 
Etheostoma serriferum 
Etheostoma simoterum 
Etheostoma smithi 
Etheostoma s. spectabile 
Etheostoma squamiceps 
Etheostoma striatulum 
Etheostoma stigmaeum 
Etheostoma swaini 
Etheostoma swannanoa 
Etheostoma tippecanoe 
Etheostoma tuscumbia 
Etheostoma variatum 
Etheostoma virgatum 
Etheostoma vitreum 
Etheostoma z- zonale 
Etheostoma sp. (duskytail darter) 
Etheostoma sp. (Elk darter) 
Etheostoma (Ulocentra) sp. 

A — emerald darter 
Etheostoma {Ulocentra) sp. B 
Etheostoma (Ulocentra) sp. C 
Etheostoma (Ulocentra) sp. 

D — golden snubnose darter 
Etheostoma (Ulocentra) sp. 

E — (Green River) 
Etheostoma (Ulocentra) sp. 

F — (Barren River) splendid 

darter 
Perca flavescens 
Percina aurantiaca 
Percina burtoni 
Percina c. cap rod es 
Percina c. semifasciata 
Percina copelandi 
Percina crassa 
Percina e. evides 
Percina evides subsp. 



N 



N 




N 


N 




N 


E 


E 





N 



N 



E 
E 



NI 



IP IP 



IP 



N 



NNNNINNNNNNN 
N N N N N N N 

N N N N N N 



So 



GM 



So 





> 


n 


NI 










N 




















N 


N 














N 


N 


N 
N 

N 


N 
N 

N 


N 
N 
E 
N 
N 
E 




M 


GM 
GM 


N 


N 






N 


N 


N 


N 


N 
E 







N 


N 


N 


N 


N 


N 




E 




No 





N 


N 


N 


N 


N 


N 


N 


N 
N 


N 
N 
E 




M 












N 




N N 






















N 




G 


















N 




M 



IP 


No 


M 


E 






N 






N 




M 




No 


M 


N 


So 


M 


N 




M 


E 







46 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Continued. 









Habitat 










Drainage 


occurrence 


















Atlantic Slope 
























o 
o 




SJ 




T3 

C 

'% 
o 


"5. 


c 

C 

o 


> 

2 

S 


1 


Creek 
Peedee 


ex 
cd 
U 


3 

Z 




c 
O 


>-> 


u 
O 


1 

OS 


o 


1 
■§ 

3 
C/3 


Percina gymnocephala 




X 


X 




X 






















Percina macrocephala 




X 




X 


X 






















Percina maculata 




X 




X 


X 






















Percina n. notogramma 




X 






X 












N 


N 


N 


N 




Percina n. montuosa 




X 






X 












E 










Percina ouachitae 


X 


X 




X 


X 






















Percina oxyrhyncha 




X 


X 


X 


X 






















Percina p. pel tat a 




X 






X 












N 


N 


N 


N 


N 


Percina p. nevisense 




X 






X 






N 


N 


N 












Percina peltata subsp. 




X 






X 










E 












Percina phoxocephala 




X 






X 






















Percina rex 




X 






X 










E 












Percina roanoka 




X 






X 






N 


N 


N 


NI 










Percina s. sciera 


X 


X 




X 


X 






















Percina shumardi 


X 


X 




X 
























Percina squamata 




X 


X 




X 






















Percina tanasi 




X 




X 
























Percina {Odontopholis) sp. 




X 






X 






















Stizostedion canadense 


X 


X 




X 
























Stizostedion v. vitreum 




X 




X 


X 


IP 




NI 




NI 


NI 






I 


N 


SCIAENIDAE 
































Aplodinotus grunniens 


X 


X 




X 

























Literature Cited 



Hambrick, P. S., C. H. Hocutt, M. T. Masnik, and J. H. Wilson. 1973. Additions to the West Virginia 

ichthyofauna with comments on the distribution of other species. — Proceedings of the West 

Virginia Academy of Sciences 45:58-60. 
Hendricks, M. L., J. R. Stauffer, Jr., C. H. Hocutt, and C. R. Gilbert. 1979. A preliminary checklist 

of the fishes of the Youghiogheny River. — Natural History Miscellanea 203:1-15. 
Hocutt, C. H., R. F. Denoncourt, and J. R. Stauffer, Jr. 1978. Fishes of the Greenbrier River, West 

Virginia, with drainage history of the southern Appalachians .^Journal of Biogeography 

5:59-80. 

, , and . 1979. Fishes of the Gauley River, West Virginia. — Brimleyana 1:47-80. 

, and P. S. Hambrick. 1973. Hybridization between the darters Percina crassa roanoka and 

Percina oxyrhyncha (Percidae, Etheostomatini), with comments on the distribution of Percina 

crassa roanoka in New River. — American Midland Naturalist 90:397-405. 
, , and M. T. Masnik. 1973. Rotenone methods in a large river system. — Archiv fur 



Hydrobiologia 72:245-252. 
Jenkins, R. E., E. A. Lachner, and F. J. Schwartz. 1972. Fishes of the central Appalachian drainages: 
their distribution and dispersal, pp. 43-117. In: P. C. Holt (ed.). The distributional history of 
the biota of the southern appalachians. Part III: Vertebrates. — Research Division Monographs 
4, Virginia Polytechnic Institute and State University. Blacksburg, Virginia. 



VOLUME 95, NUMBER 1 



47 



Table 1. — Continued. 



Drainage occurrence 



Ohio Basin 



Native 
extralimital 
distribution 



TO TO q> 



o 



o 



Percina gymnocephala 








E 




















Percina macrocephala 


N 




N 






N 


NP 


N 


N 


N 


N 







Percina maculata 


N 


N 


N 




N 


N 


N 


N 


N 


N N N 




GM 


Percina n. notogramma 
























No 




Percina n. montuosa 




























Percina ouachitae 


















N 




N 




GM 


Percina oxyrhyncha 


N 


N 


N 


N 


N 


N 


N 


N 












Percina p. peltata 
























No 




Percina p. nevisense 




























Percina peltata subsp. 




























Percina phoxocephala 
















N 


N 


N 


N 




M 


Percina rex 




























Percina roanoka 








IP 




















Percina s. sciera 




N 


N 




N 


N 


NP 


N 


N 


N 


N 




GM 


Percina shumardi 














N 


N 


N 


N 


N 




GM 


Percina squamata 




















N 


N 






Percina tanasi 






















E 






Percina {Odontopholis) sp. 
















N 


N 










Stizostedion canadense 


N 




N 




N 


N 


NP 


N 


N 


N 


N 




GM 


Stizostedion v. vitreum 


N 




N 


NI 


N 


N 


N 


NP 


N 


N 


N 




GM 


SCIAENIDAE 




























Aplodinotus grunniens 


N 


N 


N 




N 


N 


N 


N 


N 


N 


N 




GM 



Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. 
Atlas of North American freshwater fishes. — North Carolina State Museum of Natural History, 
Raleigh, N. C. 854 pp. 

Stauffer, J. R., Jr., K. L. Dickson, J. Cairns, Jr., and D. C. Cherry. 1976. The potential and realized 
influences of temperature on the distribution of fishes in the New River, Glen Lyn, Virginia. — 
Wildlife Monographs 50: 1-40. 

, C. H. Hocutt, and D. S. Lee. 1978. The zoogeography of the freshwater fishes of the Potomac 

River basin, pp. 44-54. In: K. C. Flynn and W. T. Mason (eds.). The freshwater Potomac: 
aquatic communities and environmental stresses. — Interstate Commission Potomac River Ba- 
sin. Rockville, Maryland. 

, , M. T. Masnik, and J. E. Reed, Jr. 1975. The longitudinal distribution of the fishes 

of the East River, West Virginia-Virginia. — Virginia Journal of Science 26:121-125. 



(JRS) Appalachian Environmental Laboratory, Center for Environmental and 
Estuarine Studies, University of Maryland, Frostburg State College Campus, 
Frostburg, Maryland 21532; (BMB) Department of Zoology, Southern Illinois 
University, Carbondale, Illinois 62901; (RE J) Department of Biology, Roanoke 
College, Salem, Virginia 24153; (CHH) Horn Point Environmental Laboratories, 
Center for Environmental and Estuarine Studies, University of Maryland, Cam- 
bridge, Maryland 21613. 



PROC. BIOL. SOC. WASH. 

95(1), 1982, pp. 48-57 

TWO NEW GENERA OF DEEP-SEA POLYCHAETE WORMS 

OF THE FAMILY AMPHARETIDAE AND THE ROLE OF 

ONE SPECIES IN DEEP-SEA ECOSYSTEMS 

Robert Zottoli 

Abstract. — Two new ampharetid genera, Decemunciger and Endecamera, each 
with one new species, D. apalea and E. palea, are described from wood panels 
placed on the deep-sea floor by Turner (1973). The role of Decemunciger in deep- 
sea ecosystems is discussed. 



Recently, I examined a collection of ampharetid polychaetes removed from 
pieces of wood collected by R. D. Turner, using the submersible DSRV Alvin, 
from four experimental bottom stations in the North Atlantic, at depths ranging 
from 1830 to 3995 meters. The wood had been placed at these sites by Turner to 
study moUuscan wood borers, and to "test the hypothesis that wood is an im- 
portant source of nutrients and contributes to diversity in the deep sea" (Turner 
1977:18). 

Bivalve molluscs of the subfamily Xylophagainae (Family Pholadidae, Genera 
Xyloredo and Xylophaga) mechanically excavate burrows in wood (Turner 1973, 
1977). The bivalves ingest wood particles, making wood by-products available 
within fecal pellets as food for detritus consumers. Adult and juvenile bivalves 
may be consumed by a variety of predators such as galatheid crabs. Turner (1977) 
suggests that young galatheid crabs may feed on recently settled Xylophaga lar- 
vae and, later, on larger invertebrates including adult borers. Stomach contents 
of older crabs contained sponge spicules, small pieces of wood, a nematode and 
polychaete setae (Turner 1977). Other groups in this deep-sea food chain include 
several families of polychaete worms, brittlestars, small sea urchins and predatory 
gastropods (Turner 1977). 

Two new genera from the family Ampharetidae, each with one new species, 
have been found among the ampharetids associated with Turner's wood panels. 
The external anatomy of these ampharetids, Decemunciger apalea n. sp. and 
Endecamera palea n. sp., is described, followed by a discussion of the role of 
the former species in deep-sea ecosystems. 

Materials and Methods 

Three experimental islands, each with 12 separate one foot cubes of spruce 
wood were placed by Turner (1977), for a period of five years, at the following 
locations: 

1. Deep Ocean Station 1 (DOS-1), 39°46'N, 70°41'W, 110 miles south of Woods 
Hole, Mass., in 1830 m. 

2. Deep Ocean Station 2 (DOS-2), 38°18.4'N, 69°35.6'W, 190 miles southeast of 
Woods Hole, Mass., in 3506 m. 



VOLUME 95, NUMBER 1 49 

3. Tongue of the Ocean, Bahama Islands (TOTO Tower 3), 24°53.2'N, 
77°40.2'W, in 2066 m. 

Each experimental island is encircled by wood panels, 24" x 5" x 1", which 
are removed and replaced each time the islands are visited. At the time of re- 
trieval, panels are enclosed in mesh bags to prevent loss of crumbled wood and 
specimens. The mesh bags and their contents are then placed in 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 previously placed in the retrieval boxes, thus releasing gluteraldehyde. Al- 
ternatively, the contents in certain cases may be preserved immediately upon 
reaching the surface. Specimens were well preserved. The use of submersibles 
in studies of benthic communities is described by Grassle (1980a). 

Systematics 

Ampharetid polychaetes generally are wide anteriorly, tapering gradually to- 
wards the posterior end (Fig. lA). The prostomium is generally trilobed. Seg- 
ments 1 and 2, which lie immediately behind the prostomium, are fused in most 
species, and ventrally form the lower lip. Segment 3, in some species, bears one 
lateral bundle of paleal setae on each side (Fig. 2A). The thorax begins at segment 

4. The above segmental numbering system is that of Malmgren (1865-1866) and 
Fauvel (1927), who recognized two segments in front of the paleal. 

Decemunciger, new genus 

Type-species. — Decemunciger apalea n. sp. Gender, masculine. 

Diagnosis. — Body short, of 13 thoracic setigerous segments, last 10 unciniger- 
ous, and of 14 abdominal uncinigerous segments. Segments 1 and 2 fused ventrally 
forming lower lip. Segment 3 lacking paleae. Prostomium lacking glandular ridges. 
Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae on 
dorsal surface of segments 3-5. Branchial groups separated mid-dorsally by nar- 
row space. Abdominal notopodia and notopodial and neuropodial cirri absent. 

Remarks. — In comparison with other ampharetid genera with a similar distri- 
bution of uncinigerous thoracic segments, Decemunciger differs from Melinnata 
(Hartman, 1965), Melinnopsides (Day, 1964) and Muggoides (Hartman, 1965), 
among other characters, by having 4 rather than 3 pairs of branchiae; from Mel- 
innopsis (Mcintosh, 1885) by lacking a fleshy ridge across dorsal surface of seg- 
ment 6; and from Mexamage (Fauchald, 1972) by having 4 pairs of branchiae 
inserted on 3 successive segments rather than 4. 

Etymology. — Generic name derived from the Latin, refers to number of tho- 
racic uncinigerous segments. 

Decemunciger apalea, new species 
Fig. lA-C 

Material examined. — (asb = asbestos-backed panel; D = DSRV Alvin). 

Description. — Maximum size 6.3 mm long, 0.9 mm wide. Sexually mature in- 
dividuals as small as 3.6 mm long, 0.54 mm wide. Holotype complete, 4.7 mm 



50 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table of Material examined: 







Submerged 


Removed 


Time 
submerged 


No. 


of specimens 


Panel 


Adult 


Juveniles 


DOS-1 














N28 

N34 (asb) 
N35 
N47 
N65 
N67 
N72 
N91 (asb) 


8-30-75(D597) 
8-30-75(D597) 
8-30-75(D597) 
6-15-76(D658) 
8-17-76(D685) 
8-17-76(D685) 
7-29-77(D773) 
9-28-77(D794) 


9-28-77(D794) 
7-29-77(D773) 
9-18-78(D834) 
7-30-77(D774) 
7-29-77(D773) 
8- 1-77(D776) 
9-18-78(D834) 
9-18-78(D834) 


2 yrs 

2 yrs 

3 yrs 
lyr 
lyr 
lyr 
lyr 
lyr 




2 
2 





1 


1 

32 
4 
3 

13 
7 
3 
1 


DOS -2 














N31 




9- 5-75(D601) 8- 3-77(D777) 2 yrs 16 
Holotype (USNM #71545) and 4 paratypes (USNM #71546) 


10 


TOTO TOWER 3 












T56 (asb) 


5-12-77(D755) 


11-11-78(D851) 


lyr 





1 



long, 0.9 mm wide. Color in alcohol white to pale orange. Prostomium indistinctly 
trilobed, lacking glandular ridges. About 14 smooth oral tentacles, each with deep 
ventral groove. Segments 1 and 2 fused, ventral part forming lower lip. Segment 
3 without paleae and not visually obvious. Four pairs of smooth branchiae, about 
Vs body length; 2 on segment 3, 1 on segment 4, and 1 on segment 5. Branchial 
groups separated mid-dorsally by narrow space. Notopodial lobes each bearing 
7-11 winged capillary setae from segments 4-16. Each seta about 0.4 mm long, 
7.5 jULm wide basally, and 10 /xm wide across the blade. Notopodia lobes (unci- 
nigerous pinnules) bearing toothed uncini from segment 7 to end of abdomen. 
Ten thoracic and 14 uncinigerous abdominal segments. Thoracic uncini in single 
transverse rows, 22-35 per row. Each with about 10 teeth, more or less in 3 
transverse rows, above a rounded basal prow (Fig. IB). Abdominal uncini in 
single tranverse rows, about 10-18 per row. Each with about 15 teeth in several 
transverse rows, above a rounded basal prow (Fig. IC). Abdominal notopodia 
and notopodial, neuropodial, and anal cirri lacking. Pygidium rounded. 

Remarks. — Mucus-lined tubes covered with particulate matter, about 3 times 
worm length. Female about 4 mm long and 0.5 mm wide with approximately 260 
elliptical eggs in body cavity ranging from 25 to 150 /xm across widest diameter. 
No gonoducts visible. 

Etymology. — Specific name, derived from the Latin, refers to lack of paleae. 



Endecamera, new genus 

Type-species. — Endecamera palea n. sp. Gender, feminine. 

Diagnosis. — Body short, of 14 thoracic setigerous segments, last 11 unciniger- 
ous and of 14 abdominal uncinigerous segments. Segments 1 and 2 fused, ventrally 
forming lower lip. Paleae present on segment 3. Prostomium lacking glandular 
ridges. Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae 



VOLUME 95, NUMBER 1 



51 




Fig. 1. Decemunciger apalea: A, Lateral view of entire worm, 5.2 mm long; B, Mid-thoracic 
uncini, lateral and frontal views, length ^ 16 />tm; C, Mid-abdominal uncini, lateral and frontal views, 
length = 14 /i,m. 



52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

on dorsal surface of segments 3-4. No mid-dorsal space between branchial groups. 
Abdominal notopodia and notopodial and neuropodial cirri absent. 

Remarks. — In comparison with other ampharetid genera with a similar distri- 
bution of thoracic uncinigerous segments, Endecamera differs from Ampharana 
Hartman, 1967; Amythasides Eliason, 1955, and Glyphanostomum Levinsen, 
1884, by having 4 rather than 3 pairs of branchiae; from Amage Malmgren, 
1865-1866, Grubianella Mcintosh, 1885, and Phyllampharete Hartman and Fau- 
chald, 1971, by lack of abdominal notopodia; from Pterampharete Augener, 1918, 
and Sabellides Milne Edwards in Malmgren, 1865-1866, by having smooth rather 
than papillose oral tentacles, and from Par amage Caullery, 1944, by notosetae 
beginning on segment 4 rather than segment 6. 

Etymology. — Generic name, derived from the Greek, and transcribed to the 
Latin with a feminine- singular ending, refers to number of thoracic uncinigerous 
segments. 



Endecamera palea, new species 
Fig. 2A-C 

Material examined.— Si. Croix Station, 17°57.63'N, 64°48.6'W, in 3995 m 
DSRV Alvin dive 876, 20 Dec 1978, "wild" wood about 6 feet long. Eighteen 
specimens. Holotype (USNM 71547); 3 paratypes (USNM 71548). 

Description. — Maximum size 5 mm long, 0.75 mm wide. Holotype complete, 
3.3 mm x 0.5 mm. Color in alcohol white to pale orange. Trilobed prostomium 
with middle lobe about same width as lateral lobes; lacking glandular ridges. 
Smooth, ventrally grooved, oral tentacles. Segments 1 and 2 fused ventrally form- 
ing lower lip. Two lateral groups of paleae, about 11 in each group on segment 
3. Each palea approximately 0.38 mm long, 5 /xm wide basally, tapering gradually 
to a fine point. Four pairs smooth branchiae, about 14 body length; 3 of each 
group in a straight line across dorsal surface of segments 3-4 with 4th inserted 
just anterior to most medial branchiae. No mid-dorsal gap between branchial 
groups. Notopodial lobes, each bearing 7-12 winged capillary setae from segment 

4 to end of thorax. Setae about 0.43 mm long, 8 ixm wide basally, and 10 /am 
wide across the blade. Notopodia of segment 4 minute while those of segments 

5 and 6 larger than those of segment 4, but smaller than those of subsequent 
segments. Fourteen thoracic setigerous segments. Neuropodial lobes bearing 
toothed uncini from segment 7 to end of abdomen; 1 1 thoracic and 14 uncinigerous 
abdominal segments. Thoracic uncini in single transverse rows, about 23-27 per 
row. Each with 10 teeth, more or less in 2 transverse rows, above a rounded 
basal prow (Fig. 2B). Abdominal uncini in single transverse rows, about 10-18 
per row. Each with about 13 teeth in several rows above a rounded basal prow 
(Fig. 2C). Abdominal notopodia and notopodial, neuropodial and anal cirri lack- 
ing. Pygidium rounded. 

Remarks. — Mucus-lined tubes covered with particulate matter, about 3 times 
body length. Sexually mature females were broken or twisted making it impos- 
sible to count the number of eggs in the body cavity. No gonoducts visible. 

Etymology . — Specific name, derived from the Latin, refers to presence of pa- 
leae on segment 3. 



VOLUME 95, NUMBER 1 



53 




Fig. 2. Endecamera palea: A, Lateral view of entire worm, 4 mm long; B, Mid-thoracic uncini, 
lateral and frontal views, length = 14 /Am; C, Mid-abdominal uncini, lateral and frontal views, 
length == 12 /am. 



54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Feeding of the Two Species 

Ampharetid polychaetes live in mucus-lined tubes covered by particulate mat- 
ter. The tube often extends some distance above the sediment surface. Jumars 
found that starved Hobsonia increased the length of their tubes more, over a 
period of several months, than those that were fed (Fauchald and Jumars 1979). 
They suggested that tube building may be a form of locomotion, allowing animals 
to enter new feeding areas. Fauchald and Jumars (1979) also suggested that hor- 
izontal or vertical tube orientation may depend on food availability; horizontal 
tube orientation would allow worms to cover a larger feeding area. 

During feeding, worms emerge from the tube opening and evert their oral ten- 
tacles equal to about their body length over the sediment surface (Fauchauld and 
Jumars 1979; personal observations on Asabellides oculata [Webster], Hobsonia 
[Amphicteis] floridus [Hartmen] and Melinna cristata [Sars]). Food adheres to 
mucus produced on the ventral side of the tentacles and is carried by cilia to the 
mouth. The tentacular apparatus of Decemunciger apalea and Endecamera palea 
appears similar to that of shallow water ampharetids and it is assumed that the 
method of feeding is also similar. Pieces of wood, detritus and bivalve larvae 
(Xylophaga sp.) were found in the gut cavity of Decemunciger apalea, while only 
detritus was noted in the digestive tract of Endecamera palea. 

The digestive tract of both species like most ampharetids consists of buccal 
cavity, oesophagus, stomach, intestine, and rectum. The buccal cavity houses 
about 14 ventrally grooved and ciliated, smooth oral tentacles. 

Food available to deep-sea organisms includes plankton, remnants of marine 
macrophytes such as Sargassum, remnants of land plants, particulate residues 
of deteriorating nekton and chemo-autotrophic bacteria found in hydrothermal 
vent areas (George and Higgins 1979, Rowe and Staresinic 1979). Rowe and 
Staresinic (1979), using sediment traps, found that about 4 g C/m^/yr, mainly in 
the form of fecal pellets, reached the deep-sea bottom. Marine macrophyte re- 
mains were next in importance. Pieces of the brown alga Sargassum accounted 
for an average of 0.4 g C/m^/yr while the contribution from other sources was 
relatively small, 

Decemunciger apalea most likely feeds on fecal pellets produced by other 
animals on the "wood islands," on bivalve larvae {Xylophaga sp.) that settle 
near them, and on any type of detrital particle that reaches them from the sur- 
rounding water column. Fungi and bacteria on small pieces of ingested wood may 
represent an added source of nutrients. 

Life History /Recruitment/Growth Rate of Decemuniger 

The rate of colonization in deep-sea sediments is about two orders of magnitude 
lower than in shallow water (Grassle 1977). In addition, larval recruitment and 
settlement, growth rates, and probably mortality rates, are generally lower in the 
deep sea (Grassle and Sanders 1973, Grassle 1977, Sanders 1979). Grassle (1980b) 
found fewer individuals and species in boxes of azoic sediment placed on the 
deep-sea floor, compared to samples taken from surrounding sediments. Because 
of these factors and others, deep-sea populations are commonly dominated by 
mature adults (Grassle 1977). Opportunistic wood boring, deep-sea bivalves 
(Subfamily Xylophagainae; Family Pholadidae) characterized by rapid growth. 



VOLUME 95, NUMBER 1 55 

Table 1. — Numbers of Decemunciger apalea (in parentheses) of various body lengths (in mm) for 
listed wood panels. 

N31 DOS 2: (1) 2.0; (4) 2.7; (3) 3.0; (2) 3.2; (5) 3.6; (1) 4.0; (7) 4.5; (1) 5.4; (1) 6.0; (1) 6.3. 

N32 DOS 1: (2) 0.4; (1) 0.6; (1) 0.65; (1) 0.85; (5) 1.0; (2) 1.2; (3) 1.4; (2) 1.75; (1) 1.8; (1) 1.9; (2) 
2.0; (3) 2.3; (3) 2.5; (1) 2.8; (1) 3.0; (1) 3.5; (1) 5.0. 

N34 and 

N65 DOS 1: (1) 0.75; (1) 0.8; (2) 1.0; (2) 1.25; (3) 1.5; (1) 1.75; (1) 1.75; (1) 2.5; (1) 3.0. 

N67 DOS 1: (1) 1.2; (1) 1.3; (2) 1.5; (1) 1.75; (1) 2.5; (1) 2.65. 

N72 DOS 1: (1) 1.5; (1) 3.0; (1) 4.0; (1) 4.5; (1) 5.25. 

N91 DOS 1: (1) 1.0; (1)4.5. 



early sexual maturity, and relatively large numbers of eggs, appear to be an 
exception to the generality above (Turner 1973, 1977). The larger number of eggs 
and the resultant large number of motile larvae allow at least a few individuals 
to reach geographically scattered pieces of wood (Turner 1973, Sanders 1979). 

Decemunciger appears to have a life cycle similar to these bivalves. The large 
number of small eggs in the body cavity implies a high reproductive rate. The 
apparent onset of sexual maturity within a year's time, documented below, sug- 
gests early maturity and a rapid growth rate. Thus, also considering its ability to 
colonize a transient habitat, Decemunciger may be characterized as an oppor- 
tunistic species. Evidence for this is as follows, the only unknown aspect being 
the method of larval dispersal. 

Decemunciger with well developed egg or sperm in the body cavity ranged 
from about 3.6 to 6.3 mm in total body length. Worms less than 3.6 mm long were 
therefore assumed to be juveniles. Eggs probably pass singly through certain 
nephridia and nephridiopores into the anterior part of the tube where they are 
fertilized by sperm, released by males in the same fashion. Hobsonia {Amphic- 
teis) floridus (Zottoli, 1974), Hypaniola kowalewski (Marinescu, 1964) and Mel- 
innexis artica (Annekova, 1931), which have eggs roughly the same diameter as 
Decemunciger, retain developing larvae in the maternal tube until they are able 
to crawl on the bottom. It is hypothesized that these brooding and colonization 
patterns are the same for Decemuniciger. 

Sexually mature worms as well as juveniles were found on wood panels N72 
and N91 which were submerged for one year. Thus, if one accepts the premise 
that wood panels are colonized solely by larvae, then sexual maturity is attained 
within a year (Table 1). The absence of sexually mature Decemunciger on wood 
panels N34, N65, and N67, which were also submerged for one year, suggests 
that larval worms settled on the wood late in the year and did not have sufficient 
time to reach sexual maturity. In addition, the presence of certain size classes of 
adults and juveniles on wood panels N31, N34, N72, and N91 suggests that this 
species reproduces only at certain times of the year. Rokop (1974) felt that deep- 
sea organisms generally reproduce throughout the year. Lightfoot, Tyler, and 
Gage (1979) suggest that cyclic seasonal breeding is more common in the deep- 
sea than previously supposed. Timing of reproduction most likely reflects sea- 
sonal abundance of food reaching the deep-sea floor (Lightfoot, Tyler, and Gage 
1979). 



56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Acknowledgments 

Thanks are due to Dr. J. Fred Grassle, Charlene D. Long, Dr. Meredith L. 
Jones, and Dr. Ruth D. Turner for making specimens available and for reviewing 
the manuscript. The study, conducted by Turner (1977), is supported by the 
Office of Naval Research (ONR Contract No. 14-76-C-1281, NR 104-687 to Har- 
vard University). This paper is dedicated to my recently departed friend and 
colleague. Dr. Fred (Eric) Davis. 

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Augener, H. 1918. Polychaeta. In: W. Michaelsen (ed.), Beitrage zur Kenntnis der Meersfauna 

Westafrikas 2(2):67-625. 
Caullery, M. 1944. Polychetes sedentaire de L'Expedition du Siboga: Ariciidae, Spionidae, Chae- 

topteridae, Chloraemidae, Opheliidae, Oweniidae, Sabellariidae, Sternaspidae, Amphictendiae, 

Ampharetidae, Terebellidae. — Siboga-Expeditie Monographie 24(2): 1-204. 
Day, J. H. 1964. A review of the family Ampharetidae (Polychaeta). — Annals of the South African 

Museum 48:97-121. 
Eliason, A. 1955. Neue oder wenig bekannte schwedische Ampharetiden (Polychaeta). — Goteborgs 

Kungelige Vetenskaps Handlingar 6B(17):1-17. 
Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adjacent 

areas in the eastern Pacific Ocean. — Allan Hancock Monographs Marine Biology 7:1-575. 
, and P. A. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds. — Ocean- 
ography Marine Biology Annual Review 17:193-284. 
Fauvel, P. 1927. Polychetes sedentaires. — Faune de France 16:1-494. 
George, R. Y., and R. P. Higgins. 1979. Eutrophic hadal benthic community in the Puerto Rico 

trench. In: The Deep-Sea Ecology and Exploitation. — Ambio Special Report, No. 6:51-58. 
Grassle, J. F. 1977. Slow recolonization of deep-sea sediment. — Nature 265:618-619. 
. 1980a. In Situ Studies of Deep-Sea Communities. In: Advanced Concepts in Ocean Mea- 
surements For Marine Biology. F. P. Diemer, F. J. Vernberg, and D. Z. Mirkes (eds.). — The 

Belle Baruch Library in Marine Science, No. 10:321-332. 
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, and H. L. Sanders. 1973. Life histories and the role of disturbance. — Deep-Sea Research 

20:643-659. 
Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and 

other North Atlantic areas. — Allan Hancock Foundation Occasional Paper 28:1-378. 
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cruises, chiefly from Antarctic seas. — Allan Hancock Monographs in Marine Biology 2:1-387. 
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Bermuda and other North Atlantic areas. Part 2. — Allan Hancock Monographs in Marine 

Biology 6:1-327. 
Levinsen, G. M. R. 1884. Systematik-geografisk. oversight over de nordiske Annulata, Gephyrea, 

Chaetognathi og Balanoglossi. — Videnskabelige Meddeleser Dansk Naturhistorisk Forening 

1883:92-348. 
Lightfoot, R. F., P. A. Tyler, and J. D. Gage. 1979. Seasonal reproduction in deep-sea bivalves and 

brittlestars.—Deep-Sea Research 26(8A): 767-773. 
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Forhandlingar 22:181-192; 355-410. 
Mcintosh, W. C. 1885. Report on the Annelida Polychaeta collected by the HMS Challenger during 

the years 1873-76.— Challenger Reports 12:1-554. 
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Bidrag fran Uppsala 29:79-91. 
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VOLUME 95, NUMBER 1 57 

Rokop, F. J. 1974. Reproductive patterns in the deep-sea benthos. — Science 186:743-745. 

Rowe, G. T., and N. Staresinic. 1979. Sources of organic matter to deep-sea benthos. In: The Deep 

Sea Ecology and Exploitation. — Ambio Special Report No. 6:19-24. 
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Fitchburg State College, Fitchburg, Massachusetts 01420. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 58-66 

THE KARYOTYPE OF THE EURASIAN FLYING SQUIRREL, 

PTEROMYS VOLANS (L.), WITH A CONSIDERATION OF 

KARYOTYPIC AND OTHER DISTINCTIONS IN 

GLAUCOMYS SPP. (RODENTIA: SCIURIDAE) 

V. R. Rausch and R. L. Rausch 

Abstract. — The karyotype of Pteromys volans (L.) (2n = 38; FN = 73) is de- 
scribed, based on material from Hokkaido and northeastern Siberia, and com- 
pared with those of Glaucomys volans (L.) (2n = 48; FN = 80) and Glaucomys 
sabrinus (Shaw) (2n = 48; FN = 78) from North America. Although the similar- 
ities of the fundamental numbers are suggestive of karyotypic evolution of the 
Robertsonian type, the fossil record of these flying squirrels is at present frag- 
mentary, and no conclusion is drawn as to their affinities. Definite differences 
were observed between the karyotypes of G. volans and G. sabrinus, which had 
been described as identical. In combination with karyotypic differences, other 
characteristics of the two North American species indicate both a divergence 
earlier and an age greater than have been considered on the basis of paleontologic 
evidence. Karyograms of the three species of flying squirrels are included. 



In the course of investigations concerning northern mammals, we studied chro- 
mosomal preparations from two specimens of the Eurasian flying squirrel, Ptero- 
mys volans (L.), from Hokkaido and northeastern Siberia. Chromosomes of the 
two species of flying squirrels of the Nearctic genus Glaucomys were compared. 
The purpose of the present report is to define the karyotype of P. volans and to 
describe some previously unrecognized differences in the karyotypes of the two 
Nearctic species. Other distinguishing characteristics oi Glaucomys spp. also are 
briefly discussed. -— ^ 

Materials and Methods 

Two male specimens of P. volans were utilized. The first, P. volans orii Ku- 
roda, was provided in 1967 by Dr. H. Abe, who captured the animal at Koshimizu, 
Abashiri, Hokkaido Island (ca. 43°58'N, 144°30'E). CeUs from bone marrow and 
from one testis were fixed and stained in acetic orcein, following standard meth- 
ods for mammalian karyology. The second, P. volans cf. incanus Miller, was 
trapped by us in August 1979 along the upper Kolyma River, Magadansk Oblast', 
near the settlement of Sibik-Tiellakh (ca. 62°N, 149°30'E). Cells from marrow 
(femur) were prepared in the field and processed with Giemsa blood stain (Sea- 
bright 1972). In both cases, cells were treated with colchicine. 

Preparations were made by the same methods for Nearctic flying squirrels, as 
follows: Glaucomys sabrinus yukonensis (Osgood), two males collected at An- 
chorage and near Fairbanks, Alaska, in October 1966 and April 1969, respective- 
ly; one male G. sabrinus bangsi (Rhoads), captured on Powwatka Ridge, WaUowa 
Mountains, Oregon (ca. 45°40'N, 117°27'W), in December 1980; and one male G. 
volans querceti (Bangs), collected in the vicinity of Tampa, Florida, in early 1980. 



VOLUME 95, NUMBER 1 59 

After chromosomes (in metaphase) were counted, representative cells were 
photographed on high-contrast film, and the component chromosomes in five cells 
from each animal were measured (excepting the squirrels from the upper Kolyma 
River and from Anchorage, in which only contracted chromosomes were ob- 
served in the preparations). Chromosomes exhibiting Giemsa-banding were seg- 
regated according to size and banding-pattern with use of photographs and light 
microscopy. The identification of the X-chromosome was based on measurements 
and pattern of banding. The Y-chromosome in Glaucomys spp. was selected 
subjectively, since the small submetacentric chromosomes in these animals, of 
which the Y is one, appeared to be similar in both size and banding. The termi- 
nology concerning the location of the centromere on individual chromosomes 
follows Levan et al. (1964). The number of major chromosomal arms (funda- 
mental number, or FN) was determined according to the procedure of Matthey 
(1945). The chromosomal components of 30 to 61 cells from each of the animals 
were counted, excepting those from the upper Kolyma River and Anchorage for 
which only 18 and 15 cells, respectively, were counted. 

Results 

/. Pteromys volans {2n = 38, n = 19). — The diploid complement was made up 
of 12 metacentric autosomes (arm-ratios ranged from 1.03 to 1.7), 22 submeta- 
centric-subtelocentric and 2 acrocentric autosomes (arm-ratios from 2.4 to 6.3), 
the submetacentric X-chromosome (arm-ratio from 1.6 to 1.8), and the acrocentric 
Y-chromosome (arm-ratio from 4.2 to 7.3). In 61 cells (53 mitotic and 8 meiotic) 
examined, 49 contained 38 chromosomes and 7 contained 19 bivalent elements. 
A karyogram from the male P. volans from Hokkaido is shown in Fig. 1. 

The set of chromosomes (in all of the 18 cells examined) from the animal 
collected in northeastern Siberia was morphologically like that of P. volans from 
Hokkaido, consisting of 6 pairs of metacentric and 12 pairs of submetacentric- 
acrocentric autosomes, a submetacentric X, and an acrocentric Y-chromosome, 
as shown in Fig. 2. 

The number of major autosomal arms was 70 in both animals. Two autosomes 
with thin and lightly stained short arms were considered to be acrocentric. The 
FN (autosomes -\- X and Y) was 73 for P. volans. 

II. Glaucomys spp. (2n = 48). — Karyograms from G. volans (Florida) and from 
G. sabrinus (Oregon and Alaska) are shown in Figs. 3-5. Thirty bi-armed (meta- 
centric-subtelocentric) chromosomes (with arm-ratios ranging from 1.03 to 8.8) 
and 16 acrocentric chromosomes (with centromeres in the terminal region; arm- 
ratios ranged from 4.5 to 13.8) constituted the autosomal set in G. volans. In G. 
sabrinus, from both localities, the autosomes included 28 metacentric-subtelo- 
centric (arm-ratios ranging from 1.07 to 8.6) and 18 acrocentric (arm-ratios from 
4.3 to 14.5) elements. The sex-chromosomes of the two species were similar 
morphologically and in pattern of banding: X submetacentric, of medium size 
(ranges of arm-ratios were 1.5 to 2.03 for G. volans, and 1.9 to 2.9 for G. sa- 
brinus), and Y submetacentric of small size (ranges of arm-ratios were 2.1 to 3.2 
for G. volans, and 2.0 to 3.3 for G. sabrinus). The patterns of bands indicated 
that these two species have some equivalent, or homologous, chromosomes. 
However, significant morphologic differences also were evident. 



60 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




(\7i uln ni) Am 

H i)i liii) ftifj Pn m 



H * 

X Y 



I1( a¥ IJL h« is .. X« 

X V 



2 



* * • 



i . 

X T 



II t'i r.ijt m » .| ( 

i'lt i'i iji «A »!i ii k{k A I .;. 

l|« III oh 6f<^ CA «'« 4.^ w* .1. 



fi ^ ;< 3 



A I V 



I A t. |A 



I' I Ii a iVi^ ni lift hd In 4r. 



1 6 I A /< •• on M^ « # rk 



X Y 



r. t 



f: L 



w 



y ! t 



w • 



X 



U g 8 .. i II II U \i nn 



11 



M n •» :iU 



A * a .; 



Figs. 1-5. Karyograms of Pteromys volans (2n = 38) and Glaucomys spp. (2n = 48). 1,P. volans, 
male, from Hokkaido Island; Orcein stain. 2. /*. volans, male, from the Magadansk Oblast; Giemsa 
blood stain. 3, G. volans with Giemsa-bands, male, from Florida. Arrow indicates metacentric ele- 
ments not present in G. sabrinus. 4, G. sabrinus with Giemsa-bands, male, from Oregon. Arrow 
indicates acrocentric elements not present in G. volans. 5, G. sabrinus, male, from Alaska; Orcein 
stain. Scale-lines have value of 5 jxra. 



VOLUME 95, NUMBER 1 61 

Discussion 

The gliding squirrels allocated to the genera Pteromys G. Cuvier, 1800, and 
Glaucomys Thomas, 1908, have had rather complex taxonomic histories and, as 
indicated by recent reviews (cf. Mein 1970, Black 1972), their origins and affinities 
are obscure. The grounds for acceptance of the genus Pteromys instead of Sci- 
uropterus F. Cuvier, 1825, for the flying squirrel in northern Eurasia have been 
discussed by Miller (1914) and by Ellerman and Morrison-Scott (1951), among 
others, but Simpson (1945) considered the latter to be probably the valid generic 
designation. In addition to the widely occurring P. volans, the genus includes a 
second species, P. momonga Temminck, 1846, which occurs on the Japanese 
Islands of Honshu, Kyushu, and Hondo. According to Ellerman and Morrison- 
Scott (1951), its cranial differences are so marked that it cannot be considered a 
race oi P. volans. 

In his revision of the genera and subgenera of the Sciuropterus-group of flying 
squirrels, Thomas (1908) recognized four subgenera in Sciuropterus, mainly on 
the basis of dental characters. These included Sciuropterus, with S. {Sciuropter- 
us) russicus Tiedemann, 1808 (=Sciurus volans Linnaeus, 1758) as type, and 
Glaucomys, which was erected for the Nearctic flying squirrels, with S. (Glau- 
comys) volans {—Mus volans Linnaeus, 1758) as type. Glaucomys was elevated 
to generic rank by Howell (1915), on grounds not stated, but he later (1918) 
defined cranial and dental characters that distinguish Glaucomys from Pteromys. 
These distinctions were confirmed by Ognev (1940), who also pointed out the 
marked differences in the structure of the os penis in P. volans and G. volans. 
Recently, from the study of both fossil and living flying squirrels, Mein (1970) 
placed Pteromys and Glaucomys in different generic groupings on the basis of 
dental characters. 

The fossil record provides no clear indication of the origin of Pteromys volans. 
However, the comparison of serum proteins by means of double immunodiffusion 
by Zholnerovskaia et al. (1980) suggested that Pteromys has its closest affinities 
with Sciurus and Tamias, rather than with terrestrial sciurids. In North America, 
Glaucomys is known from the middle to late Irvingtonian. The stratigraphic range 
of Glaucomys volans extends from the late Irvingtonian, whereas remains of the 
northern G. sabrinus are known only from deposits of late Wiirm age (Kurten 
and Anderson 1980). Thenius (1972) considered that Glaucomys appears to have 
arisen from terrestrial sciurids, which had their center of radiation in North 
America. 

The diploid number of 38 chromosomes, as reported by Tsuchiya (1979) and 
determined by us, alone distinguishes the karyotype of Pteromys volans from 
those of Glaucomys spp. (2n = 48) (Nadler and Sutton 1967, Schindler et al. 
1973). In the former, 35 chromosomes were metacentric-subtelocentric, and only 
3 (two autosomes and the Y-chromosome) were acrocentric. The relatively large, 
acrocentric Y of P. volans is markedly different from the male sex-chromosomes 
of Glaucomys spp. That the fundamental numbers in these squirrels are rather 
similar (73 in P. volans; 78 in G. sabrinus; and 80 in G. volans) suggests that 
karyotypic evolution involving chromosomal rearrangements of Robertsonian 
type might have occurred. However, no significance is attributed to the apparent 
similarities when nothing is known concerning the further relationships of the 
Eurasian and American species. 



62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

The karyotype of a female specimen of P. momonga has been pubhshed by 
Tsuchiya (1979, fig. 7). It appears to be quite similar to that of P. volans, but the 
available details do not permit an extensive comparison. The presence of boreal 
forest far to the south in the Japanese Islands during the glacial maximum of 
Wiirm time would seem to have favored the southward dispersal of Pteromys 
volans (cf. Kotani 1969). 

With reference to the karyotypes of Glaucomys spp. , the published information 
is discrepant. That G. volans and G. sabrinus are karyotypically identical has 
been generally accepted. However, for both, Schindler et al. (1973) considered 
the number of bi-armed (metacentric-subtelocentric) autosomes to be two more 
than did Nadler and Sutton (1967). Hsu and Benirschke (1973) concurred with 
Schindler et al. The karyotype defined by us for G. sabrinus yukonensis (Alaska) 
agreed more closely with that shown and discussed by Nadler and Sutton (1967), 
who studied preparations from G. sabrinus lascivus (Bangs) (California), than 
with the results of Schindler et al. (1973) for G. sabrinus from New Hampshire 
(an area where this species is marginally sympatric with G. volans). Accordingly, 
we examined new material from G. sabrinus as well as G. volans, from geographic 
regions far removed from areas of parapatry or sympatry (Oregon and Florida). 
The identities of the animals studied were confirmed from the distinctive char- 
acteristics of the OS penis, in addition to other criteria. 

Although the respective karyotypes of these flying squirrels appeared superfi- 
cially to be much alike, they exhibited distinct differences, principally in the 
presence in G. volans of two small metacentric autosomes that did not occur in 
G. sabrinus, and in G. sabrinus of two small acrocentric autosomes not observed 
in G. volans. As well, the total lengths of individual chromosomes differed be- 
tween the two species, including those in some presumed to be homologous; the 
relative length of the female haploid complement (autosomes + X) was recorded 
as 65 )Ltm in G. volans and 86 /xm in G. sabrinus. Morphologically, the sex- 
chromosomes were similar. 

Of the chromosomes of G. volans, the two small metacentric autosomes (see 
Fig. 3, arrow) were the most nearly metacentric, sensu stricto, with an arm-ratio 
ranging from 1.03 to 1.05. However, these and the two small acrocentric elements 
in G. sabrinus (see Fig. 4, arrow) were similar in total length; their value relative 
to the lengths of the respective haploid complements was the same (3.5%) in both 
species. The appearance of Giemsa-bands in these chromosomes suggests that 
they may be homologues, and that the small metacentric pair in G. volans arose 
as a result of a pericentric inversion involving the small acrocentric pair in G. 
sabrinus (or perhaps vice versa). The disparate total lengths of the respective 
complements further suggest quantitative differences in nuclear DNA, which 
were not defined by our methods. 

Our findings indicate that the two species of Glaucomys are karyotypically 
distinct, and they are in contrast with the conclusions of others concerning these 
flying squirrels. The karyogram from G. volans from Florida (Fig. 3) appears to 
be identical with those shown by Schindler et al. (1973) as representative of both 
G. volans and G. sabrinus in New Hampshire, where the two species are sym- 
patric. 

The two species of Glaucomys are distinguished additionally by major differ- 
ences in the form of the os penis, as is also Pteromys volans. Since detailed 



VOLUME 95, NUMBER 1 63 

Table 1. — Helminths oi Glaucomys volans and G. sabrinus. 





G. volans 

Midwest 

(n = 22) 

no. infected 


G. 


sabrinus 


Species of helminth 


Midwest 

(n = 5) 

no. infected 


Oregon 

(n = 26) 

no. infected 


Nematoda: 








Capillaria americana Read, 1949 


7 


— 


— 


Syphabulea thompsoni (Price, 1928) 


1 


5 


— 


Syphabulea sp.* 


— 


— 


17 


Lemuricola sciuri (Cameron, 1932) 


13 


— 


— 


Citellinema bifurcatum Hall, 1916 


2 


2 


7 


Cestoda: 








Andrya sciuri Rausch, 1947 


— 


2 


4 


Monoecocestus thomasi Rausch and Maser, 1977 


— 


— 


18 


Catenotaenia sp. 


— 


3 


— 


Hymenandrya sp. 


— 


— 


3 



Description in preparation by J. -P. Hugot, Museum National d'Histoire Naturelle, Paris. 



descriptions of these structures have been pubHshed, it suffices here to point out 
their diversity in the three species. The os penis of P. volans is small (ca. 4 mm 
long in our material) and consists of a slender, pointed bone with a well developed 
barb at the proximal end (cf. Ognev 1940, fig. 153). That of G. volans, first 
described by Pocock (1923), is long (ca. 14 mm in specimens from Florida) and 
very slender, whereas that of G. sabrinus is shorter (ca. 6.5 mm in our material), 
broader, and flattened (see Burt 1960, plate IV). Each differs markedly from the 
others, and we were unable to discern any fundamental similarities unless, per- 
haps, the barb near the distal end of the os penis of G. sabrinus is homologous 
with the basal barb in that of P. volans. 

Certain differences in the helminth faunas of the two species of Glaucomys 
provide a further indication of dissimilarity. Of the nine species of helminths 
recorded by us from flying squirrels in North America (Table 1), six occur also 
in sciurids of other species (Rausch and Tiner 1948, Davidson 1976, McGee 1980). 
Host-specific cestodes of three species, Andrya sciuri, Monoecocestus thomasi, 
and Hymenandrya sp., are known only from Glaucomys sabrinus. The assem- 
blages involving these cestodes would seem to have arisen independently through 
coevolution of helminth and host (Rausch 1981), indicating comparatively early 
divergence of the two species oi Glaucomys. These cestodes are Nearctic species, 
with no congeners known from Pteromys volans or other sciurids in Eurasia. The 
only helminth known from P. volans is Citellina petrovi Shul'ts, 1930, which is 
host-specific and has been recorded from European Russia, Japan, and Chukotka 
(Hugot 1980). Some of the characteristic ectoparasites of Pteromys volans and 
Glaucomys spp. are congeneric (Acarina and MaUophaga), but no species is 
shared. Consequently, these arthropods do not appear to provide any useful 
indications of the relationships of their hosts. 

Various hypotheses have been proposed to account for the essential allopatry 
of the Nearctic flying squirrels, but the geographic ranges of these mammals seem 
simply to reflect differences in specific ecologic requirements. The two species 
occupy disjunct ranges over the greater part of North America where they occur. 



64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

relative to the distribution of coniferous and deciduous forest. In the east, where 
their ranges are parapatric to sympatric, they appear as well to be segregated 
ecologically. As noted by Guilday (1962), G. sabrinus has a relict distribution in 
the Appalachian Mountains, where it occurs at elevations above ca. 1000 m as 
a consequence of climatic change (warming) during post-glacial time. 

That the characteristic habitats of the two species of Glaucomys have a fun- 
damental relationship to their different dietary requirements is evident. Glauco- 
mys volans utilizes buds, fruits, and insects during spring and summer, but later 
feeds mainly on acorns and other nuts and seeds, some of which are stored in 
large quantities and consumed during winter (cf. Muul 1968). Glaucomys sabrinus 
depends mainly on hypogeous fungi during the warmer months, and utilizes epi- 
phytic lichens (Usnea and Alectoria) during winter (McKeever 1960, Maser et 
al. 1978, Maser pers. comm.). The diet of Pteromys volans differs from these in 
that it feeds on sprouts or buds of Be tula and Acer and on male flowers of conifers 
in summer, and on catkins or buds of Betula and Alnus in winter (Ognev 1940, 
Tavrovskii et al. 1971). 

Suggestions have been made to account for the pattern of distribution of Glau- 
comys sabrinus on grounds other than ecologic requirements. Muul (1968) con- 
sidered that the southward spread of G. sabrinus is prevented by competitive 
interaction with G. volans, inasmuch as the latter breeds earlier seasonally and 
therefore would already occupy the limited number of tree-cavities available. 
However, as reported by Cowan (1936), and observed by us in northern Wiscon- 
sin, G. sabrinus commonly utilizes outside nests constructed mainly of twigs. 

Weigl (1975) observed in captive animals that infection by a nematode of the 
genus Strongyloides was tolerated by G. volans, but was associated with high 
mortality in G. sabrinus. He considered that differential pathogenicity of this 
nematode might account in part for the segregation of the two species of Glau- 
comys where they are sympatric. Barbehenn (1978) suggested that this might 
constitute a mechanism by which the larger G. sabrinus would be excluded from 
habitat otherwise suitable for G. volans. However, Strongyloides robustus Chan- 
dler, 1942 exhibits little host-specificity, occurring in sciurids of various species 
(Rausch and Tiner 1948, McGee 1980). Davidson (1976) considered it to be the 
most pathogenic of the more common helminths in squirrels, and reported that 
infections involving more than 150 individuals often caused severe enteritis in 
gray squirrels, Sciurus carolinensis Gmelin. It might be expected that transmis- 
sion of this nematode would be enhanced by conditions of captivity, and that 
differences in feeding habits or other behavior could account for more massive 
infections in G. sabrinus. That Weigl did not find northern fiying squirrels natu- 
rally infected by Strongyloides appears to be compatible with the results of other 
surveys of helminths in this sciurid. 

Altogether, the differences between the two flying squirrels of the genus 
Glaucomys seem to indicate earlier divergence and greater age of these species 
than has been considered on the basis of paleontologic evidence. The origins and 
affinities of Glaucomys spp. and of Pteromys volans may be established if the 
earlier fossil record of these sciurids can be traced. 

Acknowledgments 

Specimens of flying squirrels were kindly provided by H. Abe, Faculty of 
Agriculture, Hokkaido University, Sapporo; Ms. E. L. Bull, Pacific Northwest 



VOLUME 95, NUMBER 1 65 

Range and Habitat Laboratory, U.S. Department of Agriculture, La Grande, 
Oregon; R. Anderson, Wallowa Valley Ranger District, U.S. Department of Ag- 
riculture, Joseph, Oregon; and L L. Duncan, Center for Disease Control, Atlanta. 
Field work in the Soviet Union was carried out under the U.S.A./U.S.S.R. En- 
vironmental Protection Treaty, Area V, Subproject A-3. In the U.S.S.R., V. L. 
Kontrimavichus, D. L Berman, and Ms. A. N. Leirikh, Institute of Biological 
Problems of the North, Academy of Sciences of the U.S.S.R., Magadan, kindly 
provided support and assistance. We express sincere thanks for these contribu- 
tions. 

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Black, C. C. 1972. Holarctic evolution and dispersal of squirrels (Rodentia: Sciuridae). — Evolutionary 
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Burt, W. H. 1960. Bacula of North American mammals. Museum of Zoology, University of Michigan, 
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Davidson, W. R. 1976. Endoparasites of selected populations of gray squirrels (Sciurus carolinensis) 
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Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic and Indian mammals. 
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Hsu, T. C, and K. Benirschke. 1973. Glaucomys volans (southern flying squirrel). Folio 312. Atlas 
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Kurten, B., and E. Anderson. 1980. Pleistocene mammals of North America. Columbia University 
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Levan, A., K. Fredga, and A. A. Sandberg. 1964. Nomenclature for centromeric position on chro- 
mosomes. — Hereditas 52:200-220. 

Maser, C, J. M. Trappe, and R. A. Nussbaum. 1978. Fungal — small mammal interrelationships with 
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McGee, S. G. 1980. Helminth parasites of squirrels (Sciuridae) in Saskatchewan. — Canadian Journal 
of Zoology 58:2040-2050. 

McKeever, S. 1960. Food of the northern flying squirrel in northeastern California. — Journal of 
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66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Nadler, C. F., and D. A. Sutton. 1967. Chromosomes of some squirrels (Mammalia: Sciuridae) from 

the genera Sciurus and Glaucomys. — Experientia 23:249-251, 
Ognev, S. I. 1940. Zveri SSSR i prilezhashchikh stran. IV. Gryzuny . Akademiia Nauk SSSR. Moskva- 
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London (1923), pp. 209-246. 
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Symposium, La specificite parasitaire des parasites de vertebres. Museum National d'Histoire 

Naturelle, Paris, 13-17 April 1981. (in press) 
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Helminths of Sciuridae. — American Midland Naturalist 39:728-747. 
Schindler, A.-M., R. J. Low, and K. Benirschke. 1973. The chromosomes of the New World flying 

squirrels {Glaucomys volans and Glaucomys sabrinus) with special reference to autosomal 

heterochromatin. — Cytologia 38: 137-147. 
Seabright, M. 1972. The use of proteolytic enzymes for the mapping of structural rearrangements in 

the chromosomes of man. — Chromosoma 36:204-210. 
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the American Museum of Natural History 85:1-350. 
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PROC. BIOL. SOC. WASH. 

95(1), 1982, pp. 67-80 

A NEW SPECIES OF THE GENUS ECHININUS (MOLLUSCA: 

LITTORINIDAE: ECHINININAE) WITH A 

REVIEW OF THE SUBFAMILY 

Joseph Rosewater 

Abstract. — A new ovoviviparous species, Echininus viviparus, inhabiting high 
intertidal and supratidal areas in the Mariana Islands is described in the Httorinid 
subfamily Echinininae which previously consisted of the polytypic Indo-Pacific 
species Echininus cumingi (Philippi, 1846) and the west Atlantic species Tectin- 
inus nodulosus (Pfeiffer, 1839). Characteristics of the subfamily are reviewed. 



I received for identification from L. G. Eldredge of the University of Guam 
specimens of an Echininus which upon close examination proved to be not only 
a new species, but one which exhibits ovoviviparity. The species described here 
I originally thought to be Echininus cumingi spinulosus (Philippi, 1847), and the 
realization that it is not prompted me to take a fresh look at the subfamily Echin- 
ininae resulting in the following review. 

Family Littorinidae Gray, 1840 
Subfamily Echinininae Rosewater, 1972 

Diagnosis. — The subfamily Echinininae is characterized by spinose to bluntly 
spinose shells; umbilicate or imperforate; opercula multispiral; radulae with mod- 
erately to greatly narrowed rachidians. Distribution is tropical west Pacific and 
west Atlantic. 

The subfamily Echinininae was established by Rosewater (1972) to contain the 
genus Echininus which differs in matters of sculpture, radula, animal morphology, 
and ecology from either Littorininae or Tectariinae. This subfamily concept is 
strengthened by findings enumerated in the present paper. Members of the 
subfamily have a unique habitat among Littorinidae, for they inhabit highest shore 
levels, at times seeming almost to be terrestrial. The radula of Echininus, while 
said to be "not unusual" (Rosewater 1972:507), is distinctive when examined 
with the SEM (Fig. lA-D). The radula of Tectininus is unique (Fig. lE-F). The 
operculum is multispiral, a departure from all other known littorines. Character- 
istics of both radulae and opercula are here considered adaptations to a high 
shore habitat, as is the occurrence of ovoviviparity in the new species of Echin- 
inus described herein. All are indicative of a basic evolutionary trend in Echini- 
ninae toward a terrestrial habit. Shells of this group are also distinctive, especially 
that of E. cumingi cumingi, which is characterized by rather long, open spines 
and a deep umbilicus. These latter features, also, may be of significant adaptive 
value to an animal living on tropical shores enabling easier dissipation of heat. 

Tectininus Clench and Abbott, 1942 

Tectininus Clench and Abbott, 1942:4; type-species by original designation Echin- 
inus nodulosus (Pfeiffer, 1839).— Abbott, 1954:458^62. 



68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Diagnosis. — Shells bluntly spinose, succeeding spiral rows of spines non-syn- 
chronous; imperforate; opercula multispiral; radula with very reduced rachidian, 
modified lateral, and marginals with few denticles. Distribution is subtropical to 
tropical west Atlantic. 

As originally proposed by Clench and Abbott (1942) and elaborated by Abbott 
(1954) the monotypic subgenus, Tectininus, was distinguished from Echininus 
s.s. because its type, E. nodulosus, possesses a unique radula, although still 
exhibiting the formula 2-1-1-1-2 and a modified littorinid embayment (Fig. lE-F). 
It is unlike any other radula in the family, having massive lateral teeth and a 
diminutive rachidian which appears almost vestigial. The operculum is multispir- 
al. It possesses a penis bearing a subterminal penial gland surrounded with fleshy 
papillae, which differs from the condition in the nominate subgenus and combines 
characteristics noted in the genera Littorina, Nodilittorina, and Tectarius. Al- 
though Rosewater (1972) had not seen the paper and reported that nothing was 
known of reproduction in Echinininae, it had already been shown that T. nodu- 
losus is oviparous and produces a pelagic egg capsule (Borkowski 1971). 

Until further evidence for its true relationships comes to light, I am satisfied 
to permit Tectininus to remain in its present position. As stated by Abbott, after 
careful study of the type-species of Tectininus, it appears to be ''. . . a special- 
ization of the ancestral stock ... in the family Littorinidae" (Abbott 1954), whose 
apparent closest relations are in the Echinininae. Because of its unique radula 
Tectininus should be accorded full generic status. 

Echininus Clench and Abbott, 1942 

Echininus Clench and Abbott, 1942:3; type-species by original designation Tro- 
chus cumingii Philippi, 1846. — Rosewater, 1972:525. 

Diagnosis. — Shells spinose to bluntly spinose; often with partly open spines; 
umbilicate to imperforate; opercula multispiral; radulae with moderately nar- 
rowed rachidians, laterals and marginals with moderately numerous denticles. 
Distribution is west Pacific from southern Japan through western Pacific arc to 
Cook Islands. 

The genus Echininus was delineated as consisting of 2 subgenera containing a 
total of 2 living species and 1 subspecies (Rosewater 1972). The nominate sub- 
genus, Echininus, inhabits the west Pacific and contains E. cumingi cumingi 
(Philippi, 1846) known from the southern Philippines southward through Mela- 
nesia (including New Guinea, Solomon Islands, New Hebrides, New Caledonia) 
eastward to the Cook Islands, and, E. cumingi spinulosus (Philippi, 1847) which 
occurs in southern Japan, the Ryukyu Islands, Taiwan and northern Philippines. 
Echininus adelaidensis (Cotton, 1947) is a Pliocene fossil from South Australia, 
which resembles certain trochaceans (Rosewater 1972). New evidence from pre- 
served specimens indicates that the Mariana Islands population, previously be- 
lieved to be part of E. cumingi spinulosus, is different from either of the previ- 
ously known Pacific subspecies and is a new species. 

Echininus cumingi cumingi, type-species, is comparatively large, reaching 20.9 
mm (Table 1), with 3 spiral rows of conspicuous projecting, open spines on the 
body whorl; it has a narrow, deep umbilicus; a multispiral operculum is present; 
the radula has a moderately narrow rachidian, lateral and inner marginal teeth 



VOLUME 95, NUMBER 1 69 

each having one large and several smaller cusps, and slender outer marginals 
having 4-5 cusps. Echininus cumingi spinulosus is very close in appearance to 
E. c. cumingi, although it never reaches as large a size (18.3 mm), is less spinose, 
and has a narrower umbilicus which occasionally is closed. The specimens from 
the Mariana Islands which were included with the former by Rosewater (1972) 
are consistently different from either E. c. cumingi or E. c. spinulosus. Available 
records indicate they are limited in distribution to the southern Mariana Islands, 
and, so far have been reported only on the islands of Saipan, Tinian, Rota and 
Guam. Specimens are smaller in size, the largest measuring 12.3 mm in length; 
have only 2 major rows of blunt spines on the body whorl. In addition, the shell 
shape is different; obesity of the Mariana specimens averages 0.76 compared with 
0.87 in cumingi and 0.78 in spinulosus. There are a number of quite absolute 
anatomical and biological differences also. Females of the Mariana population 
reproduce ovoviviparously, and the expanded oviduct may contain large numbers 
of young. The embryos are visible through the transparent dorsal mantle tissue 
in various stages of development including young snails with shells of 1.25-1.5 
whorls. This condition is very similar to that described in Littorina saxatilis by 
Thorson (1946), and Fretter and Graham (1962). The reproductive modes of £". 
c. cumingi and E. c. spinulosus are not known, but there is no evidence from 
examination of preserved specimens to indicate ovoviviparity in them. There are 
also morphological differences between radulae and penes of the Mariana species 
and the more western and northern Echininus. 

Echininus viviparus, new species 
Figs. 1-6, Table 1 

Description. — Shell: reaching 12.3 mm (about V2 inch) in length, turbinate in 
shape, with blunt spines; average obesity (width/length) about 0.76 (76 specimens 
range from 0.65-0.84) relatively thick in structure, usually non-umbilicate or with 
only a thin slit between columella and parietal callus; suture usually evident 
although often partially obscured by spiral sculpture; whorls 3-6, moderately 
shouldered below suture, relatively flat-sided; usually with 2 major rows of blunt 
spines on body whorl and one on spire whorls; occasionally with one or more 
additional major rows of spines. External shell color varying from grayish to 
tannish orange, the darker color characteristic of shells retaining periostracum; 
periostracum frequently worn away on spines which then appear grayish white. 
Apertural coloration light to dark orange with occasional light or darker orange 
lines revolving within. Base moderately flattened, sculptured with weakly nod- 
ulose cords. Length of spire greater than half length of shell. Spire convex; spire 
angle quite variable: from 60-72°. Aperture subquadrate; outer lip thickened, 
smooth within, only slightly wavy at edge reflecting external sculpture; inner lip 
with small but distinct toothlike bulge. Suture evident when not partly obscured 
by sculpture. Primary sculptural feature: 2 spiral rows of irregular blunt spines 
on body whorl, and one row on spire whorls. Rows of spines not well aligned 
axially; body whorl spine count 1 1-16 in posterior row and 12-16 in anterior row; 
13-14 on penultimate whorl of spire. Secondary spiral sculpture consists of 3^ 
fat, non-spinose cords between each spinose row, which occasionally become 
nearly as spinose as spinose cords they separate. In addition, an overall micro- 



70 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




u 


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1 


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Fig. 1. Radulae: A, B, Echininus viviparus, 9 from northeast coast of Tinian, Mariana Islands 
(USNM 796244): A, Showing 2 complete transverse rows, Bar = 10 />t,m, 875x; B, Rachidian tooth, 
Bar = 5 /xm 1800x. C, D, Echininus cumingi cumingi, 9 from Davao, Mindanao, Philippines (USNM 
1A1165): C, Showing 2 complete transverse rows, Bar = 10 jxm, 700x; D, Rachidian tooth, Bar = 5 
ixm, 1400X. E, F, Tectininus nodulosus, $ from San Salvador, Bahamas (USNM 596683): E, Showing 
1 complete transverse row; note small, narrow rachidian tooth and massive laterals. Bar = 10 /xm, 
510x; F, Oblique view of rachidian tooth, Bar = 2 /xm, 3500x. 



VOLUME 95, NUMBER 1 



71 



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72 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Shells of Echininus viviparus: A, B, Holotype (USNM 792356), 9.7 x 7.4 mm; C, D, 
Paratype (USNM 803296) 10.7 x 7.6 mm. 



scopical sculpture of fine, closely-spaced threads. Axial sculpture consists of 
irregular lines of growth. Operculum small in size, multispiral (polygyrous spiral 
type of Fretter and Graham 1962) having 5-6 volutions, chitinous, light brown in 
color. Nuclear whorls usually present, smooth, light brown to reddish brown; 
reaching about 1.25-1.50 volutions before developing spiral sculpture. 

Animal: Radula littorinoid, 2-1-1-1-2; rachidian narrow, but with bulbous base 
having 3 basal denticles; lateral tooth with large central cusp and smaller lateral 
cusps and with distinct littorinoid embayment in which inner marginal tooth ar- 



VOLUME 95, NUMBER 1 



73 



GO 

b 




ro 
en 

3 
3 




Fig. 3. Echininus viviparus: A, $ with well developed young in expanded oviduct seen through 
dorsal mantle; B, Detail from A: shell of young snail of about 1.5 whorls; C, 6 showing arrangement 
of penial glands. All from northeast coast Tinian, Mariana Islands (USNM 796244). 



ticulates; inner marginal with 4 cusps of which third from medial is largest; outer 
marginal with 4-5 cusps increasing in size laterally. Animal littorinoid; in the 
male penis is unbranched; wide and deep sperm groove runs along its posterior 
surface, the edges of which are brown pigmented; penis with 5-8 penial glands 
arranged in straight line along outer edge; well developed ctenidium present in 



74 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



160° 



I4Q° 




100° 120° 140° ISO 

Fig. 4. Known distribution of Echininus species. 



dorsal mantle cavity; leaflets well formed; efferent branchial vessel and osphra- 
dium present (similar to that shown in Fretter and Graham 1962, Fig. 7). Fecal 
pellets oval in shape, white with dark flecks. Female ovoviviparous with oviduct 
modified as brood pouch; containing as many as 50 or more embryos in various 
stages of development up to the crawling stage in which sheU has 1.25-1.50 
whorls. 

Etymology. — Viviparus, an adjective, from the Latin, meaning "bearing active, 
living young." 

Measurements. — See Table 1. 

Types. — Holotype, USNM 792356; length 9.7 mm, width 7.4 mm; 15 paratypes, 
USNM 803296, ranging in length from 8.1-12.3 mm, and in width from 6.2-8.2 
mm. 

Type-locality. — Machong Point [=As Matmos or Dudu, personal communi- 
cation, L. G. Eldredge] (about 14°irN, 145°18'E), Rota, Mariana Islands; in 
spray zone 30 feet landward from top of 50 foot cliff; 4 February 1979; L. G. 
Eldredge, collector. 

Material examined. — Mariana Islands, West Pacific Ocean. 

Guam: 

[1*] Agfayan Point (13°16'N, 144°44'E) high intertidal on limestone; July 1980, 
R. H. RandaU, USNM 803297. 



* Figure "1" refers to column 1 in Measurements Table 1; specimens from Guam are summarized 
together. 



VOLUME 95, NUMBER 1 75 

Camel Rock (13°29'N, 144°42'E) at 20 foot elevation on limestone; 13 

June 1980, L. G. Eldredge, USNM 803298. 
Asan Point (13°29'N, 144°42'E); 1951, D. B. Langford, USNM 613687. 
Apra Bay (13°27'N, 144°38'E); November 1907, P. Bartsch, USNM 

233323. 
Guam, 29 August 1949, V. L. Haack, USNM 620383. 
Guam, 1946, Capt. Draeger, USNM 707190. 

Rota: 

[2] Machong Point [= As Matmos or Dudu] (about 14°1 1 'N, 145°18'E) in spray 
zone 30 feet landward from top of 50 foot cliff; 4 February 1979, L. G. 
Eldredge, type-lot, USNM 792356, 803296. 

[3] West side Poniya Point (14°06'N, 145°10'E) above terraced bench in spray 
zone, 2 February 1979, R. K. Kropp, USNM 803299. 

Tinian: 

[4] Northeast coast (about 15°06'N, 145'39'E) in spray and splash zone of large 
blowhole, 100 feet landward from 20 foot cliff, 8 February 1979, L. G. 
Eldredge, USNM 796244. 

Saipan: 

[5] Puntan Agingan (15°07'N, 145°42'E) in depressions in limestone 6-10 feet 
landward from 25-30 foot cliff, 19 November 1980, L. G. Eldredge, 
USNM 803300. 

[6] Puntan Magpi (15°16'N, 145°48'E) on limestone in spray of blowhole, 5-8 
foot cliff, 21 November 1980, L. G. Eldredge, USNM 803301. 

Ecology. — Lives in the high intertidal and supratidal areas on rough, pitted 
limestone apparently restricted in distribution by the extent of ocean spray; often 
in the vicinity of blowholes (Tinian) and sometimes found at considerable dis- 
tances landward from the tops of sea cliffs 20-50 feet high (Rota and Tinian) (L. 
G. Eldredge, personal communication) (Fig. 6). 

Echininus viviparus lives highest above the sea of any littorinid that I have 
observed. It is described as living where it is only wet by spray from surf or the 
blowholes characteristic of coral shores, where it probably feeds on vegetation 
which is kept moist by the same spray. The ovoviviparous reproduction of this 
species appears to be in close accord with its high position on the shore and 
permits it to survive without a pelagic stage in its development. Careful studies 
need to be carried out before it can be stated unequivocally, however, that E. 
viviparus has no need to return to the sea for any part of its life history. 

Geographical distribution. — Southern Mariana Islands: Guam, Rota, Tinian, 
and Saipan. 

Comparative remarks. — Morphometries of Echininus viviparus are compared 
with those oiE. c. cumingi and E. c. spinulosus in Table 1. Note that E. viviparus 
is smaller than the other two species, reaching only 12.3 mm in maximum length 
versus 20.9 and 18.3 mm respectively. On the average it is less obese, 0.76 versus 
0.87 and 0.78, although difficulties of measuring spinose shells may artifically 
enhance the obesity of E. c. cumingi. Numbers of postnuclear whorls and rows 



76 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






1 K'lifll^Eifl^^^l 



Fig. 5. A, Apex of Echininus viviparus, showing smooth protoconch of about L25 whorls (compare 
with Fig. 3B), specimen from northeast coast Tinian (USNM 796244), Bar = 100 />tm, lOOx; B, Apex 
of Littorina saxatilis, another ovoviviparous littorine, showing smooth protoconch of about 1.5 
whorls, specimen from Odiorne Point State Park, Rye, Rockingham County, New Hampshire (USNM 
803282), Bar = 100 ^tm, 80x ; C, Apertural view of young E. viviparus showing 2 major rows of blunt 
spines on body whorl separated by moderately strong intermediate row, same data as A, Bar = 250 
^im, 40x ; D, Apex oi E. cumingi cumingi, showing decollate tip, specimen from Philippines (USNM 
89449), Bar = 100 /^m, 185x. 



of tubercles are less in E. viviparus. The aperture length/shell length index is 
smaller in E. viviparus. These data show the new species is smaller in size and 
more slender than previously known species. It is also non-umbilicate where the 
other two usually have an umbilicus. There is a rather weak but persistent tooth- 
like bulge on the inner lip. 

The animal of E. viviparus is typically littorinoid, having a blunt snout and 
tentacles with eyes at the outer bases. It is significant that females oiE. viviparus 
produce young ovoviviparously (Fig. 3A) because this has not been noted in other 
Echininus species. The penis oiE. viviparus differs markedly from other species: 
in E. viviparus there are 5-8 penial glands in a straight line along the outer edge 
of the unbranched penis (Fig. 3C), while in E. c. cumingi, the glands occur only 



VOLUME 95, NUMBER 1 77 

at the junction of the bulbous base and the penial extremity, laterally and pos- 
teriorly (Rosewater 1972: plate 406, fig. B). The arrangement in E. c. spinulosus is 
similar to the latter. Radulae of the three species of the genus Echininus are 
similar, with differences in some details. In all species the teeth are small (see 
magnifications. Fig. 1) and reminiscent of the "pick type" (Rosewater 1980). 
Rachidian teeth of all three species of Echininus are narrow distally, but in E. 
viviparus the base is bulbous with 3 denticles (Fig. IB). The tip of the protoconch 
is frequently present in E. viviparus, but is smaller and often decollated in E. 
cumingi and E. spinulosus (Fig. 5). 

Discussion. — When I initially mapped the distribution oi Echininus (Rosewater 
1972, plate 407), information from museum specimens indicated that the ranges 
of the two subspecies, E. c. cumingi and E. c. spinulosus, were divided into 
northern and southern components by an east- west line drawn just south of Puerto 
Princessa, Palawan, Philippines (9°N). Some additional records have come to 
light for E. c. cumingi: Guimaras Island, Philippines [verifying the type-locality 
of E. cumingi] (10°35'N) (Kevin Marx, personal communication 1979); Lifou, 
Loyalty Islands (C. Lamb 1979, USNM) not very far from the Tana, New Heb- 
rides record reported by Rosewater (1972). The discovery of the new species, 
Echininus viviparus, changes the pattern of distribution of the genus Echininus 
in the Indo-Pacific (Fig. 4). Although obviously related generically, the Mariana 
species is quite distinct from the other Echininus taxa in its distribution, in certain 
aspects of its morphology and, so far as is known, in reproducing ovoviviparous- 
ly. In my previous evaluation of a few small, worn shells from Guam (Rosewater 
ibid.) I referred them to E. c. spinulosus because no anatomical evidence was 
available to the contrary. The additional freshly collected specimens provided by 
L. G. Eldredge have shown without doubt that E. viviparus is a separate species 
distinct from the subspecies group of E. cumingi. It is rather difficult to speculate 
upon the origin of E. viviparus. Little fossil evidence exists to trace the evolu- 
tionary history of the genus Echininus or, in fact, of any members of the family 
Littorinidae. Some littorinid fossils may have been assigned to such families as 
Turbinidae and Trochidae which are almost indistinguishable as fossils. The most 
similar in appearance is the fossil, Tectarius rehderi Ladd, 1966, from Lower 
Miocene, Marshall Islands (Ladd 1966, Rosewater 1972, pi. 404, figs. 5-7). Ac- 
cording to Ladd (1960) much of the mollusk fauna now inhabiting west Pacific 
islands and the East Indies may have migrated in successive waves from east to 
west during Cretaceous and Tertiary times when an archipelago of islands existed 
through which species could move with relative ease. Such species as Tectarius 
rehderi may be the precursors of Recent Echininus and Tectarius species now 
inhabiting the western Pacific arc. It seems unlikely that the fossil species Echin- 
inus adelaidensis (Cotton, 1947) from the Pliocene of South Australia belongs in 
the Littorinidae and probably should be referred to the Trochacea. In the absence 
of data indicating a more widespread distribution, it appears that Echininus vi- 
viparus has evolved as an endemic species in the Mariana Islands where its 
ancestors became established during one of these earlier migrations. Other lit- 
torinid inhabitants of the Mariana Islands fail to show any indication of such 
endemicity (Roth 1976, Rosewater 1970, 1972). 

Echininus viviparus joins the small but increasing group of mollusks whose life 
histories are known to include ovoviviparity, or brooding of young by the parent 



78 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. A, Machong Point, Rota Island, Mariana Islands, habitat of Echininus viviparus 30 feet 
landward from top of 50 foot cliff; B, Same, showing spray reaching habitat; C, Same, showing pitted 
Hmestone; white dots are Echininus; Bar = 20 cm; D, Northeast coast Tinian, Mariana Islands, where 
Echininus lives 100 feet landward from top of 20 foot cliff. (Photos courtesy of L. G. Eldredge.) 



through early stages of development. The occurrence of this form of reproduction 
in mollusks was reviewed by Van der Schalie (1936) who attributed its develop- 
ment to being of survival value for animals subjected to unfavorable environment. 
Ovoviviparity now is known to occur in several additional groups to those cited 
by Van der Schalie (1936). In Littorinidae it is known in at least three species, 
L. saxatilis {=L. littoralis of Van der Schalie), and E. viviparus. Littorina scabra 
scabra and L. scabra angulifera release young in various stages of development, 
as summarized by Mileikovsky (1975). Other instances of ovoviviparity in gas- 
tropods occur in Siliquariidae, Struthiolariidae, Nassariidae, and Turridae (Mor- 
ton 1958); Coralliophilidae (Wells and Lalli 1972); Janthinidae and Hydrobiidae 
(Fretter and Graham 1962); Acmaeidae, Hipponicidae, Planaxidae, Vermetidae, 
Thiaridae, a few opisthobranchs, many Stylommatophora, a number of chitons, 
and some Aplacorphora (Hyman 1967). Among the Bivalvia there are also scat- 
tered cases: Nuculidae (Drew 1901); freshwater bivalves QxcQpt Dreissensia [sic] 
(Cooke 1895); Arcidae, Ostreidae, Erycinacea (Morton 1958); Carditacea (Jones 
1963, Yonge 1969); Veneridae (personal observation in the genus Gemma); Ter- 
edinidae (Turner 1966). 

Ovoviviparity occurs within groups of mollusks often in what appears to be a 
rather haphazard fashion, although occasionally it will occur in an entire family 
such as Unionidae or Viviparidae. The development of this special type of pa- 
rental care, as Van der Schalie (1936) has suggested, appears associated with 
animals living under special stress for which ovoviviparity contributes survival 



VOLUME 95, NUMBER 1 79 

value, i.e. larvae of freshwater mussels are distributed on fish; crawl-away young 
of Echininus living on cliffs far above the sea will survive where a pelagic de- 
veloper could not. There is no way to predict when ovoviviparity will occur in 
a group. But the frequency with which it does occur lends credence to its value 
under the proper conditions. 

Summary. — As presently defined the littorinid subfamily Echinininae consists 
of the following units distributed as noted: 

Family Littorinidae Gray, 1840 

Subfamily Echinininae Rosewater, 1972 
Genus Echininus Clench and Abbott, 1942, type-species by original desig- 
nation Trochus cumingii Philippi, 1946 
Echininus cumingi cumingi (Philippi, 1846) — southern Philippines and 

along the western Pacific arc to New Hebrides and the Cook Islands 
Echininus cumingi spinulosus (Philippi, 1847) — southern Japan through the 

Ryukyu Islands to northern and western Philippines 
Echinininus viviparus, new species — southern Mariana Islands 
IfEchininus adelaidensis (Cotton, 1947) — Pliocene, South Australia 
Genus Tectininus Clench and Abbott, 1942, type-species by original desig- 
nation, Litorina nodulosa Pfeiffer, 1839 

Tectininus nodulosus (Pfeiffer, 1839) — southern Florida, Bermuda, the 
Bahamas, and the Greater Antilles 

Acknowledgments 

L. G. Eldredge, University of Guam, provided specimens oi Echininus vivip- 
arus which he sent to me via G. J. Vermeij and P. Signor. He also sent habitat 
photos and information on ecology. The female with brood pouch and details 
of male anatomy were drawn by I. Jewett. SEM preparations were made by 
P. Greenhall and scanning electron microscopy performed at the Smithsonian's 
SEM laboratory, under the direction of W. Brown, by S. Braden and M. Mann. 
R. S. Houbrick offered helpful suggestions on the manuscript. 

Literature Cited 

Abbott, R. T. 1954. Review of the Atlantic periwinkles, Nodilittorina, Echininus and Tectarius. — 

Proceedings of the U.S. National Museum 103 (3 3 28): 449-464. 
Borkowski, T. V. 1971. Reproduction and reproductive periodicities of south Florida Littorinidae 

(Gastropoda: Prosobranchia). — Bulletin of Marine Science 21(4):826-840. 
Clench, W. J., and R. Tucker Abbott. 1942. The genera Tectarius and Echininus in the western 

Atlantic. — Johnsonia 1(4): 1-4. 
Cooke, A. H., A. E. Shipley, and F. R. C. Reed. 1895. Molluscs, Brachiopods (Recent), Brachiopods 

(Fossil). — The Cambridge Natural History 3, Macmillan and Co., New York and London, 

xi + 535 pp. 
Cotton, B. C. 1947. Some Tertiary fossil mollusks from the Adelaidean Stage (Pliocene) of South 

Australia. — Records of the South Australian Museum 8(4):653-670. 
Drew, G. A. 1901. The life history of Nucula delphinodonta (Mighels). — Quarterly Journal of Mi- 
croscopical Science 44(3) N.S.:3 13-391. 
Fretter, V., and A. Graham. 1962. British prosobranch molluscs. — The Ray Society, London, 

xvi + 755 pp. 
Hyman, L. H. 1967. The invertebrates, 6, Mollusca 1. McGraw-Hill, New York, vii + 792 pp. 



80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Jones, G. F. 1963. Brood protection in three Southern California species of the pelecypod Cardita. — 
The Wasmann Journal of Biology 21(2):141-148. 

Ladd, H. S. 1960. Origin of the Pacific island molluscan fauna. — American Journal of Science, 
Bradley Volume 258-A: 137-150. 

. 1966. Chitons and gastropods (Haliotidae through Adeorbidae) from the Western Pacific 

Islands. — Geological Survey Professional Paper 531:1-98, 16 plates. 

Mileikovsky, S. A. 1975. Types of larval development in Littorinidae (Gastropoda: Prosobranchia) 
of the world oceans and ecological patterns of their distribution. — Marine Biology 30:129-135. 

Morton, J. E. 1958. Molluscs. Hutchinson and Co., Ltd., London, 232 pp. 

Pfeiffer, Louis. 1839. Bericht iiber die Ergebnisse meiner Reise nach Cuba in Winter 1838-1839. — 
Archiv fiir Naturgeschichte, BerUn, Funfter Jahrgang 1:346-358. 

Philippi, R. A. 1846. Description of a new species of Trochus, and of eighteen new species of 
Littorina, in the collection of H. Cuming, Esq. — Proceedings of the Zoological Society of 
London for 1845, part XIII: 138-143. 

. 1847-1851. Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien 

3:1-138. 

Rosewater, J. 1970. The family Littorinidae in the Indo-Pacific, Part I. The subfamily Littornininae. — 
Indo-Pacific Mollusca 2(ll):417-506, plates 325-386. 

. 1972. The family Littorinidae in the Indo-Pacific, Part II. The subfamilies Tectariinae and 

Echinininae.— Indo-Pacific Mollusca 2(1 2): 507-533, plates 388-408. 

. 1980. A close look at Littorina radulae. — Bulletin of the American Malacological Union for 

1979:5-8, 8 figures. 

Roth, A. 1976. Preliminary checkhst of the gastropods of Guam. — University of Guam Marine Lab- 
oratory Technical Report No. 27, Sea Grant Publication UGSG-76-03:vi + 99. 

Thorson, G. 1946. Reproduction and larval development of Danish Marine bottom invertebrates, 
with special reference to the planktonic larvae in the Sound (0resund). — Meddeleser Era Kom- 
missionen For Danmarks Fiskeri — og Havenders0gelser, serie: Plankton 4(1): 1-523. 

Turner, R. D. 1966. A survey and illustrated catalogue of the Teredinidae. The Museum of Com- 
parative Zoology, Harvard University, vii + 265 pp. 

Van der Schalie, H. 1936. Ovoviviparity among mollusks. — The Nautilus 50(1): 16-19. 

Wells, F. E., and C. M. Lalli. 1977. Reproduction and brood protection in Caribbean gastropods 
Coralliophila abbreviata and C. caribbaea. — Journal of Molluscan Studies 43:79-87. 

Yonge, CM. 1969. Functional morphology and evolution within the Carditacea (Bivalvia). — Pro- 
ceedings of the Malacological Society of London 38:493-527. 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

95(1), 1982, pp. 81-88 

A NEW DWARF SPHAERODACTYLUS FROM HAITI 
(LACERTILIA: GEKKONIDAE) 

Richard Thomas 

Abstract. — Sphaerodactylus omoglaux is described from the southeastern Cul- 
de-Sac Plain and adjacent montane foothills of Haiti. It is a ground dwelling 
species of xeric to semi-xeric situations and belongs to a group of Hispaniolan 
Sphaerodactylus including S. altavelensis Noble and Hassler, S. armstrongi No- 
ble and Hassler, S. cryphius Thomas and Schwartz, S. darlingtoni Shreve, S. 
nycteropus Thomas and Schwartz, and S. streptophorus Thomas and Schwartz. 



Recent fieldwork in Haiti resulted in the discovery of an undescribed small 
species of Sphaerodactylus in the southeastern part of the Cul-de-Sac Plain and 
adjacent foothills of the Massif de la Selle. This new sphaerodactyl is related to 
the recently described Sphaerodactylus cryphius and S. streptophorus (Thomas 
and Schwartz, 1977). 

Abbreviations. — ASFS refers to the Albert Schwartz Field Series, CM to the 
Carnegie Museum of Natural History, RT to Richard Thomas's personal collec- 
tion, UMMZ to the University of Michigan Museum of Zoology, and USNM to 
the National Museum of Natural History. 

Sphaerodactylus omoglaux, new species 
Figs. 1-3 

Holotype. — USNM 221840, adult female, taken on the eastern edge of the town 
of Fond Parisien near the shore of Etang Saumatre, Dept. de TOuest, Haiti, on 
1 August 1979 by Richard Thomas. 

Paratypes.—ASVS V50461-62, 0.7 km E Fond Parisien, 16 July 1978, R. Thom- 
as; ASFS V50463, RT 5681, same locality as preceding, 23 July 1978, R. Thomas; 
RT 7173, ca. 1 km E Fond Parisien, 2 July 1979, R. Thomas; RT 7682, UMMZ 
172100-01, CM 83302, USNM 221841, same data as holotype; RT 5591, 8 km 
airline NW Fond Verrettes, ca. 0.3 km by road west of the ford across the Riviere 
Soliette, 424 m, 19 July 1978, R. Thomas; all paratypes are from the Dept. de 
rOuest, Haiti. 

Diagnosis. — A small species oi Sphaerodactylus (Fig. 1), maximum snout-vent 
length 20 mm; dorsal scales flattened, keeled, imbricate (30-34 between axilla 
and groin), no area of middorsal granules or granular scales; smooth throat and 
ventral scales (27-30 between axilla and groin); 51-57 scales around midbody; 
dorsal body scales with 5 to 9 hair-bearing scale organs, each with one hair, along 
distal edge of scale; rostral with a large flat dorsal area bordered by a rim and 
sloping towards the tip; 2 postnasals; large cobble-like snout scales (10-13 across 
base of snout). Head (Fig. 2) with a bilobed light anterior figure having postero- 
lateral extensions; paired, transverse mid-nuchal marks; paired but separate scap- 
ular ocelli; traces of dorsolateral striping; dorsal body pattern of scattered dark 



82 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ST 
I'll 



ST 



CR 



CR 



JJ 



AL 



y 



^ L 



OM 



y 

AL 

I L 



id 



40 



50 




OM 



15 ^^ 25 

mm 



Fig. 1 . Line histograms of midbody scale counts (left side of figure) for Sphaerodactylus omoglaux 
(OM), S. cryphius (CR), S. streptophorus (ST, sample from the Pedernales area, Dominican Re- 
public), and 5'. altavelensis (AL). The altavelensis sample above the dashed line is from the area of 
sympatry with S. cryphius, that from below the line is from the area of sympatry with S. omoglaux 
(Fond Parisien localities). The right side of the figure shows histograms of snout-vent length in mm 
size classes; in all graphs the smallest vertical unit is a single individual. 



brown flecks; lineate sacral figure; lineate-oceUate caudal pattern; boldly lined 
throat in males; no ventral lines. 

Description of holotype . — An adult female, 19 mm snout- vent length, tail (part- 
ly regenerated), 13 mm; dorsal body scales flattened, strongly keeled, acute and 
imbricate, 31 between axilla and groin; throat scales smooth, not becoming ex- 
tremely minute on central throat area; ventral scales smooth, flattened, rounded, 
imbricate, 30 between axilla and groin; 52 scales around midbody; lamellae of 
fourth toe of left pes 9. Snout moderate, rostral scale with large dorsal flat area 
bordered by a rim and sloping toward tip; snout scales broad, subhexagonal, 
keeled, cobble-like and subimbricate; 1 internasal; 2/2 postnasals; 3/3 supralabials 
to mid-eye; temporal scales and dorsal head scales keeled, juxtaposed, subim- 



VOLUME 95, NUMBER 1 



83 






B 




Fig. 2. Dorsal patterns of S. omoglaux: A, RT 7686; B, RT 5591; C, USNM 221840 (holotype). 



bricate; first infralabial broader anteriorly than posteriorly, subrectangular; dorsal 
scales of tail acute, keeled, flattened and flat-lying; ventral scales of tail smooth 
and enlarged in median line. 

Dorsal ground color in life brown, markings darker brown to nearly black; head 
with dark preocular lines and a pale transverse snout bar; pale anterior cephalic 
figure bilobed, extending behind eyes onto parietal region and having a short pale 
extension from the posterolateral corner of each lobe; a small pale parietal spot 
in the space between the posterior parts of the lobes; a pair of pale, dark-edged, 
transverse mid-nuchal marks; paired pale (cream) dark-edged scapular ocelli dor- 
solaterally positioned and not joined by a scapular patch of dark pigment; broad, 
dark-edged dorsolateral stripes present in scapular region, the uppermost stripe 
edge on each side interrupted by a scapular ocellus; stripes fading out just beyond 
level of axilla; dorsum brown with irregular mottling and spotting of darker 
brown; sacral markings of indistinct dark lines; caudal pattern lineate (dorsolat- 
eral, lateral, and ventrolateral dark lines) with light spots or ocelli spaced along 
the dorsalmost pair; venter unpatterned, pale (off-white) in life. 

Variation. — The maximum snout-vent length is 20 mm (female); two egg-bear- 
ing females are 19 mm, and the smallest escutcheoned male is 16 mm (largest 
male 18 mm). The scale morphology of the paratypes is similar to that of the 
holotype. Eight specimens have one internasal; 4 have 2; postnasals are 2/2 in all 
but one having 1/2; scales across the snout between the posterior ends of the first 



84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

supralabials are 10 (3 specimens), 11 (4), 12 (5), 13 (1); upper labials to mid-eye 
are 3/3 in all; dorsal scales between axilla and groin are 30-33 (x = 31.6); ventral 
scales axilla to groin, 27- 30 (x = 28.8); scales around midbody, 51-56 (Jc = 53.5); 
subdigital lamellae 8 (1), 9 (9), or 10 (2); keeling of throat and ventral scales is 
absent in all. The escutcheon of males varies from 5 to 7 scales long and from 15 
to 19 scales wide. 

In color and pattern the paratypes are generally similar to the holotype. All 
have a bilobed anterior head figure, each lobe with a median postpalpebral line 
of dark pigment. All but one show some evidence of the posterolateral extensions 
of the lobes (in three the extensions are indistinct or irregular). All have the 
preocular lines; all but three have the transverse snout bar, either pale as in the 
holotype or with the dark posterior edge being the most salient feature and thus 
appearing as a dark snout bar; median snout pigmentation varies from pale, nearly 
unpigmented, to solidly pigmented with dark brown or pigmented but with a light 
center. Nine of the 12 paratypes have plainly evident paired transverse mid- 
nuchal marks; in two the marks are obscured by irregular mottling of pigment, 
and in one they are absent. All specimens show some indication of the dorsolat- 
eral stripes in the scapular area, even if only unilaterally. Paired scapular ocelli 
are found in all and are distinctly separate, not interconnected by a dark patch, 
although each ocellus is surrounded by a zone of dark pigment (Fig. 2). Dorsal 
body coloration varies from almost uniform brown through the presence of iso- 
lated small brown flecks on a paler brown ground color to being fairly heavily 
mottled with irregular dark scale clusters. The sacral pattern is not prominent 
and consists of, when present, the dark dorsal edges of the dorsolateral lines 
which reappear in the sacral region. Unregenerated tails show the lineate and 
ocellate condition described for the holotype, the most proximal pair of ocelli 
often being the most prominent. Throats are heavily streaked with dark brown 
lines in the three largest males (the fourth male is very small, and the throat lines 
are present but not bold). Females lack throat streaking or show it only very 
weakly. Ventral coloration is pale (largely unpigmented) in most, but some have 
a peppering of melanophores over much of the surface, concentrated around the 
scale edges; the venters are never lined. 

Distribution. — Known only from the southeastern part of the Cul-de-Sac Plain 
of Haiti from the region of Fond Parisien southeastward into the foothills of the 
La Selle near Soliette (Fig. 3). 

Comparisons. — Sphaerodactylus omoglaux belongs to a closely related group 
of generally smaU to medium-sized species that includes darling toni Shreve, al- 
tavelensis Noble and Hassler, armstrongi Noble and Hassler, streptophorus 
Thomas and Schwartz, cryphius Thomas and Schwartz, and nycteropus Thomas 
and Schwartz. The last four and S. omoglaux appear to form a more closely 
related cluster within the group (Thomas and Schwartz in press). With the ex- 
ception of 5. armstrongi, these are small species. These four share polythetically 
a suite of characters, and the distributions are largely allopatric or parapatric 
(Fig. 3). Although sympatry is likely for at least two of the group, conspecific 
relationship between others cannot be ruled out. Because of the commixture of 
characters, even in sympatric members of the (larger) group, it is unwise to 
assume subspecific relationship without clear evidence of intergradation. 

The most pertinent comparisons are with Sphaerodactylus cryphius, known 



VOLUME 95, NUMBER 1 



85 




Fig. 3. Map of a portion of southern Hispaniola. Hexagons indicate S. omoglaux localities; 
rhombs, S. cryphius; circles, S. streptophorus, and triangles, S. armstrongi. 



from the south side of the Valle de Neiba, some 50 km to the southeast, in the 
Dominican RepubHc, and with S. streptophorus of the western, mostly peripheral, 
parts of the La Selle-Baoruco massif (Fig. 3). Sphaerodactylus omoglaux seems 
to be the ecological equivalent and the geographical vicariant of S. cryphius. In 
comparison with omoglaux, cryphius has a reduced color pattern with less elab- 
orate and less contrasting markings; the most striking absolute difference between 
the two lies in the nature of the scapular ocelli: prominent and separate with no 
conjoining patch in omoglaux, absent or small and close-set and with a small, 
non-enclosing but conjoining patch in cryphius (Figs. 2, 4). Furthermore the two 
taxa are strongly different in midbody scale counts (Fig. 1; Student's r-test in- 
dicates P < 0.01 that the two samples are from the same population). Sphaero- 
dactylus cryphius also lacks sexual dichromatism in the throat pattern or any 
other feature. 

Sphaerodactylus streptophorus is a distinctly larger lizard. Its scale counts 
strongly overlap those of omoglaux, but its head pattern differs strongly from 
that of omoglaux (Fig. 4A), although it may have a cognate of the anterior ce- 
phalic figure (but not bilobed: Fig. 4B). The scapular ocelli, when present, are 
separate, as in omoglaux, but not as boldly dark-edged and are sometimes farther 
back on the body. The cognate of the mid-nuchal marks of omoglaux in strep- 
tophorus is usually a thin transverse light line (whence the specific epithet mean- 
ing ''wearing a necklace"), although it may be broken into two segments. Another 
frequent feature of streptophorus is a long diagonal postauricular mark that often 
meets part of the cephalic figure to form a mesially directed temporal wedge. 

Sphaerodactylus armstrongi is a medium to large-sized member of this group 



86 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




o o 





A 





d^ 



Fig. 4. Semi-diagrammatic anterior dorsal patterns of 5. sircprophorus (A. ASFS V2580; B, ASFS 
V2590) and 5. ciypluus (C. ASFS V2051 1). 



that occurs in the uplands and easternmost lowlands of the La Selle-Baoruco 
ranges. It lacks the bilobed cephalic figure, but it may show a light dorsal head 
pattern bounded by postocular stripes and an occipitonuchal M-shaped figure 
(subspecifically variable). The postauricular marks (legs of the M) are probably 
homologues of the shorter, posteroventral light marks that extend from the cor- 
ners of the cephalic figure in omoglau.x. Easternmost 5. armstrongi often has 
prominent, separate scapular ocelli somewhat resembling the condition in om- 
oglaiix. Sphaerodactyhis armstrongi further differs in having one postnasal and 
a rounded rostral that does not have a shaiply delimited flat area and does not 
slope towards the tip. Sphaerodactylus armstrongi also differs from omoglaux 
(and Streptophorus, ctyphius. and nycteropus) in its proportionately larger 
ear openings. 

Sphaerodactylus nycteropus lacks mid-nuchal marks or scapular ocelli: it does 
have, in some specimens, a cephalic figure but no posterolateral marks. Evidence 
of dorsolateral lines is present. The venter is strongly lined. In scale characters 
and size nycteropus does not differ markedly from omoglaux. Sphaerodactylus 
nycteropus is now known from a total of five specimens. 

It is pertinent to compare 5. omoglaux and 5. altavelensis, which occur syn- 
topically. Both are small sphaerodactyls of generally similar appearance, although 
altavelensis is somewhat larger (Fig. 1). Both have similar scale moiphology, 
although omoglaux has higher midbody scale counts and lacks the keeling on the 
throat that is present in altavelensis. Reflecting the size difference, the modal 
number of subdigital lamellae is 10 in altavelensis, 9 in omoglaux. Both species 
have bilobed anterior cephalic figures, and the posterior cephalic figure of alta- 
velensis may be the homologue of the mid-nuchal marks of omoglaux. Sphae- 
rodactylus altavelensis has small, close-set scapular ocelli that are situated on 



VOLUME 95, NUMBER 1 87 

the outer edges of an irregular scapular patch. The patch may have a short light 
anterior margin giving it a straight border anteriorly and may also have indications 
of a larger, enclosing but not delimiting, light border laterally and posteriorly (this 
feature and patch size varies in different subspecies of altavelensis; see Thomas 
and Schwartz (in press) for a discussion of variation in altavelensis). The bold 
throat pattern is lacking in either sex of altavelensis, although faint striping is 
present in some individuals, but the venter is lined. 

Remarks. — At the two localities near Fond Parisien all of the specimens of 
Sphaerodactylus omoglaux were collected in leaf litter and beneath rocks within 
about 200 m of the south shore of Etang Saumatre. The habitat is a mixture 
of xerophytic scrub {Acacia, Prosopis, Opuntia, Cephalocereus) and more mesic 
woods, which include Sabal palms, Swietenia, Mangifera, Tamarindus , Catalpa, 
etc. There is a moderate gradient going toward the shore of the lake, where 
the habitat becomes more mesic and densely vegetated, although open and closed 
situations are patchy due to cutting by man. The only other sphaerodactyl that 
we found was S. altavelensis, which occupies the same ground habitat as om- 
oglaux. We obtained altavelensis in greater numbers than omoglaux (about 3 to 
1). All of the omoglaux were obtained by one person, whereas native collectors 
contributed to our sample of altavelensis. Sphaerodactylus altavelensis is com- 
mon in all of the leaf-litter habitats in the more or less wooded situations, whereas 
omoglaux seems to be confined to shadier, lower canopy (or thicker canopy) 
situations. We were unsuccessful in attempting to collect any sphaerodactyl in 
the pure desert scrub a few kilometers to the northwest of Fond Parisien, although 
some species almost certainly occur there. 

Sphaerodactylus omoglaux exists at least macrosympatrically with S. copei, 
S. elegans, and S. cinereus, in addition to S. altavelensis, just discussed. The 
first three are large species and are inhabitants principally of trees, rock crevices, 
or other three-dimensional niches (e.g., wood piles, thatch roofs). The single 
specimen of S. omoglaux from 8 km NW Fond Verrettes was taken from a dead 
Agave rosette in xerophytic scrub, where the road runs along a hillside above 
the Riviere Soliette at an elevation of 427 m. This locality is about 15 km southeast 
of the type-locality and is the most xeric situation in which the species was found. 
A specimen that has been referred to S. streptophorus was taken at Soliette, 
probably less than a kilometer from the locahty for the specimen of omoglaux just 
mentioned. The road descends along the xeric hillside, fords the Riviere Soliette, 
and then proceeds along a shadier, more mesic riverside area that is Soliette 
proper. The surrounding hillsides are open — scrubby or cultivated. The two 
species, omoglaux and streptophorus, may coexist in this area, each restricted 
to patches of appropriate habitat. 

Etymology. — Omoglaux is from the Greek, omos, shoulder, and glaux, owl, 
which is derived from glaukos, gleaming, in reference to the eyes. The allusion 
is to the prominent, separate scapular ocelli. 

Acknowledgments 

I wish to thank Luis Rivera Cruz for his capable assistance in the field. The 
fieldwork during which the specimens of Sphaerodactylus omoglaux were col- 
lected was supported by National Science Foundation Grant SER 77-04629. 



88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Literature Cited 

Thomas, R., and A. Schwartz. 1977. Three new species of Sphaerodactylus (Sauna: Gekkonidae) 
from Hispaniola. — Annals of the Carnegie Museum of Natural History 46(4):33^3. 

, and . in press. Variation in two species of Hispaniolan Sphaerodactylus (Reptilia: 

Sauria: Gekkonidae). — Bulletin of the Museum of Comparative Zoology. 

Biology Department, University of Puerto Rico, Rio Piedras, Puerto Rico 
00931. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 89-92 

AMERIGONISCUS MALHEURENSIS, NEW SPECIES, 

FROM A CAVE IN WESTERN OREGON 

(CRUSTACEA: ISOPODA: TRICHONISCIDAE) 

George A. Schultz 

Abstract. — A new species of Amerigoniscus Vandel, a genus with 8 other 
species in the United States, is described and illustrated from Malheur Cave in 
western Oregon. 



The new species is the third trichoniscid and the second in the genus Ameri- 
goniscus Vandel (1950) to be described from Oregon. The first species is Ore- 
goniscus nearcticus (Arcangeli, 1932) from MacLeary Park, Portland, and the 
second is Amerigoniscus rothi (Vandel, 1953a) from 8 mi (12.9 km) east of Gold 
Beach, near the California border. Neither was from a cave although both are 
blind and have little or no body pigment. The new species described here is from 
Malheur Cave in Harney County in western Oregon. It also has no eyes or body 
pigment. 

The two male specimens were collected by E. H. Gruber and sent to me by 
Jerry Lewis, University of Louisville, Kentucky, and I thank him for sending 
them to me. 

Amerigoniscus Vandel 

The genus was recently reviewed by Vandel (1977) who included information 
on the four species and added four new ones. This then is the ninth species placed 
in the genus. All are from the United States, from Georgia through Texas to 
Oregon. Only A. rothi Vandel is not from a cave. Vandel separated the eight 
species by using the configurations of the tip of the exopod of male pleopod 1 of 
each species. He illustrated that structure for all eight species. Unfortunately for 
the four new species which he described that is all he illustrated, but the four 
species are sufficiently different morphologically and geographically so as not to 
be confused with the new species described here. 

The relation oi Amerigoniscus Vandel to Oregoniscus Hatch (1947) as dis- 
cussed by Vandel (1953a: 177) is still unsettled since O. nearcticus (Arcangeli) is 
based on a female. Vandel (1953b) also discussed some of the species of the 
genus, and species of related genera of Trichoniscidae. He placed the first de- 
scribed species of Amerigoniscus in Caucausonethes Verhoeff and the genus was 
included among the primitive members of the family Trichoniscidae in Tribe 1 . 
However, when later (Vandel 1977) he changed the generic name of the species 
from the United States back to Amerigoniscus he did not mention the placement 
of the genus in any subgroup within the Trichoniscidae. 

Amerigoniscus malheurensis, new species 
Figs. lA-N, 2A-C 

There are no illustrations of the whole animal of any species of the genus. 
Unfortunately the two specimens collected here were too distorted to be drawn, 
but the appendages are definitive and are illustrated. 



90 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Amerigoniscus malheurensis, holotype male 3.5 mm long: A, Detail tip exopod male 
pleopod 1; B, Reverse of A; C-G, Pleopods 1-5 respectively; H, Peraeopod I (twisted); I, Mandibles; 
J, Maxilla 1; K, Maxilliped; L, Fungal spores (small ones same scale as mandibles); M, Antenna 1; 
N, Antenna 2. 



Description. — Eyeless, pigmentless. Cephalon and body covered with tuber- 
clelike scale groups. Few scale groups on top and one row on posterior margin 
of cephalon; 2 rows on peraeonal segments, especially apparent on segments I-V; 
few on edges of pleonal segments. Antenna 1 with 8 aesthetascs. Antenna 2 short, 



VOLUME 95, NUMBER 1 



91 




Fig. 2. Amerigoniscus malheurensis, holotype male: A, Peraeopod VII; B, Uropod; C, Pleotelson. 



with 3 flagellar articles. Mandibles each with well developed molar process and 
2 setae in setal row. Right mandible with 3 teeth on incisor process; lacinia mobilis 
with apical crown of small teeth. Left mandible with 4 teeth on incisor with 2 on 
each of 2 sections. Endopod of maxilla 1 with 2 penicillate setae on apex and 1 
longer subapical compound seta. Exopod with 8 teeth and 1 broad seta on apex. 
Maxilliped with pointed endite and palp of 2 segments with apical segment large, 
pointed and fringed with small setae. 

Peraeopod I of male with 2 setae on proximal inner margin of propodus (prob- 
ably sexually dimorphic character, but females not obtained). Peraeopod VII of 
male with 3 large setae on inner margin of propodus; carpus with simple arrange- 
ment of setae. Featherlike dactylar organ present on all peraeopods. Pleopod 1 
with large exopod with configuration of tip unique (Fig. lA, B). Pleopod 2 with 
exopod with medial posterior margin produced; 1 seta medially placed on pos- 
terior margin. Endopod with proximal segment shorter than length of exopod; 
distal segment elongate and tapered to truncate end. Pleopods 1-3 as illustrated. 
Uropodal rami each with 1 large seta at tip. Pleotelson with posterior margin 
produced, rounded, and with 4 small setae. 

Length. — Both males 3.5 mm long. 

Type-locality. — Malheur Cave, about 18 mi (29 km) southeast of New Princeton 
(on Oregon route 78), Harney County, Oregon. 

Distribution. — Known only from type-locality. 

Etymology. — The name malheurensis means from Malheur, the name of the 
cave which is the type-locality. 

Disposition of types. — The type-specimens have been placed in the National 
Museum of Natural History (Smithsonian Institution), holotype male USNM 
184664; paratype male USNM 184665. 

Discussion. — The species differs from A. rothi in that the configurations of the 
tips of the exopods of male pleopods 1 are quite different (cf. Vandel, 1977:308, 
Fig. 1, and Fig. lA here). Although the various configurations illustrated by 
Vandel (1977) are quite different, the best match is between the tip of the exopod 
of the new species and that of A. rothi. There are, however, many differences in 
other structures if the illustrations of Vandel (1953a) are compared to those in- 
cluded here. Vandel illustrated 11 aesthetascs on antenna 1 and six flagellar ar- 
ticles on antenna 2 for A. rothi. There are eight aesthetascs and three flagellar 



92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

articles on the new species. The general shape of the exopods of pleopods 1 
(excluding details of tip) have different shapes in the two species. Male pleopods 
2 are also differently shaped — in the new species the exopod has the medial 
posterior edge or tip produced; it is a right angle in A. rothi. The proximal segment 
of the endopod is proportionately shorter and the tip of the distal segment elongate 
and narrowed to a truncate, not pointed, end in the new species. There are other 
minor differences which can be seen by comparing the drawings of the two 
species. 

Ecology. — The type-specimen was found under a rotted stick in the flood zone 
at the 1325 foot (397.5 m) level. The paratype was found under a plank in the 
flood zone at the 1400 foot (420 m) level. The guts from the mouth to the anus 
of both specimens were stuffed with fungal spores on which they were apparently 
feeding. The spores are two-chambered (Fig. IL) and most probably from a fun- 
gus of the Deuteromycetes, Didymosporae group. No mycelia were present. 

Literature Cited 

Arcangeli, A. 1932. Isopodi terrestri raccolti dal Prof. Silvestri nel Nord-America. — Bolletino La- 

boratoire di Zoologicae Generale e Agraria, Portici 26:121-141. 
Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. — University 

of Washington Publications in Biology 10(5): 155-274. 
Vandel, A. 1950. Campagne speologique de C. Bolivar et R. Jeannel dans TAmerique du Nord (1928). 

Isopodes terrestres recueillis par C. Bolivar et R. Jeannel (1928) et le Dr. Henrot (1946). — 

Archives Zoologie Experimentale et Generale 87(3): 183-210. 
. 1953a. A new terrestrial isopod from Oregon, Caucasonethes rothi n. sp. — Pacific Science 

7(2): 175-178. 
. 1953b. Remarques systematiques, morphologiques et biogeographiques sur un groupe de 

Trichoniscidae Nord-Atlantiques (Crustaces; Isopodes terrestres). — Bulletin du Museum Na- 
tional d'Histoire Naturelle, Serie 2, 25(4):368-375. 
. 1977. Les especes appartenant au gtnvQ Amerigoniscus Vandel, 1950 (Crustaces, Isopodes, 

Oniscoides). — Bulletin de la Societe d'Histoire Naturelle de Toulouse 1 13:303-310. 

15 Smith St., Hampton, New Jersey 08827. 



PROC. BIOL. SOC. WASH. 

95(1), 1982, pp. 93-98 

A NEW SPECIES OF DEEP-SEA ISOPOD, 

STORTHYNGURA MYRIAMAE, FROM THE 

WALVIS RIDGE OFF SOUTH AFRICA 

Robert Y. George and Noel A. Hinton 

Abstract. — A new species of abyssal isopod, Storthyngura myriamae, is de- 
scribed. This species was collected at 5220 meters during the French JEAN 
CHARCOT expedition to the Walvis Ridge off South Africa. The study also 
includes a detailed comparison of the new species with the most closely related 
species Storthyngura caribbea from 1260 meters off the Windward Island in the 
West Indies. 



Introduction 

The deep-sea fauna on the western and eastern sides of the Walvis Ridge off 
South Africa between latitudes 25° and 35° south was recently investigated. This 
study was conducted aboard the R/V Jean Charcot by the French deep-sea group 
at the Brest Oceanographic Laboratory under the leadership of Dr. Myriam Si- 
buet. The deep-sea samples contained many isopods belonging to more than 
fifteen genera including the abyssal genus Storthyngura. 

The isopod fauna on both sides of the Walvis Ridge is thus far poorly known. 
However, the Walvis Ridge abyssal region is of zoogeographic interest in view 
of the theory that the genus Storthyngura originated in the Antarctic slope 
(George and Menzies 1968b) and the distribution of Storthyngura species is some- 
what related to the pattern of the Antarctic Bottom Water flow (George, unpub- 
lished data). The Walvis Ridge material contained a hitherto undescribed species 
of Storthyngura and the new species is named in honor of Myriam Sibuet for 
graciously providing the isopod material for study. 



Order: Isopoda, Suborder Asellota 

Family: Eurycopidae 

Storthyngura myriamae, new species 

Fig. lA-H 

Diagnosis. — Storthyngura with cephalon devoid of any spines. First pereonal 
somite lacking dorsal spines; somites 2 to 4 having single, prominent, anterior 
dorsal spine; in addition, somites 2 to 4 with posterior transverse ridge exhibiting 
median tubercle. Somites 5 to 7 displaying a pair of well-developed median dorsal 
spines. First pleonal somite lacking spination. Pleotelson with anterior spine and 
pair of posterior dorsal tubercles; 2 well-defined lateral spines and triangular apex. 
Basis of uropod approximately same length as endopod; exopod about two- thirds 
length of endopod. 

Material and station data. — Holotype female: length = 21 mm, width 8 mm 
(U.S.N.M. Cat. No. 184668). 

Type-locality: Angola Basin west of Walvis Ridge, JEAN CHARCOT Station 



94 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. I. Storthyngura mynamae: Holotype female, length 21 mm. A, Dorsal view; B, First antenna; 
C, Left mandible; D, Maxillule; E, Maxilla; F, Maxilliped; G, Sixth peraeopod; H, Uropod. 



DS09, one female specimen, 5220 m, 26°59.9'S-27°00.0'S, r06.7'E-r06.2'E. 
Date: 6-Jan-79. 

Other localities: JEAN CHARCOT Station CPIO, one female specimen, 5210 
m, 26°15.9'-27°00.35'S, r07.r-r06.4'E. Date: 7-Jan-79 (returned to Dr. Sibuet 
for deposition in the Brest Reference Museum); JEAN CHARCOT Station CP12, 



VOLUME 95, NUMBER 1 




20* E 



Fig. 2. Area map showing the Walvis Ridge separating the Angola and Cape basins. Stations with 
S. myriamae shown in soHd circles and stations lacking S. myriamae shown in open circles. 



two female specimens, 4660 m, 27°37.6'-27°38.5'S, 0°50.8'-0°51.3'E. Date: 9-Jan- 
79. Fragments from Station DS07 (See Fig. 2). 



General Description 

Body shape ovate, approximately 3 times longer than wide. Cephalon including 
a pair of mounds on dorsal surface. Lateral margin concave; frontal margin trun- 
cate. 

First pereonal somite without spination. Antero-lateral margin rounded, coxal 
plates small and acicular; second pereonal somite having prominent median spine, 
antero-lateral margin rounded; coxal plates protuberant and bilobed, anterior lobe 
prominent. Posterior ridge on somites 2 to 4 exhibiting median tubercle; somites 
3 and 4 each with produced median spine, antero-lateral margins acutely pro- 
duced, prominent and bilobed coxal plates with distinct anterior lobes; somites 
5, 6, and 7 with paired prominent median spines, concave posterior margins, 
lateral margins produced into spine-like process, coxal plates absent. 

Pleon with prominent anterior pleonal somite, smooth and lacking any spines; 
pleotelson shield-like, with well-developed acute antero-lateral angles and pro- 
jecting postero-lateral spines; apex triangular and converging to a point. Pleotel- 
son with anterior spine and pair of conical tubercles located anterior to postero- 
lateral spines. 

First antennae with capacious basal article, lateral margin entire. Peduncle with 



96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

one broad basal article, second article shorter than elongated third article: fourth 
article narrow, about one-half length of third article. Flagellum with more than 
60 annulate articles. Mandible with prominent palp consisting of 3 articles; second 
article slightly longer than first article; third article inflexed into semi-circular 
pattern; inner basal margin displaying a row of thick setae, distal margin with 7 
to 8 blunt, short spines and terminal setae. Well-developed cutting phase with 
pronounced molar and very dense middle row of approximately 20 spines. Incisor 
uniform with quadradentate lobe. Maxillule with outer lobe two times as broad 
as inner lobe; both lobes with dense setae at distal ends; outer lobe with approx- 
imately 15 stout, spine-like setae. Maxilla trilobed with outer lobes slender, each 
furnished with 2 to 4 long setae; inner lobe stout with approximately 20 terminal 
setae. Maxilliped with 11 coupling hooks on endite; palp with elongate basal 
article, somewhat narrower at base; second and third articles expanded, second 
article devoid of marginal setae, third article containing approximately 28 mar- 
ginal setae on inner margin; fourth article distally produced into lobe one-third 
length of article, with about 10 terminal setae; fifth article narrow, with tuft of 
apical setae. 

Posterior 3 legs morphologically similar; basis elongated, about as long as car- 
pus and ischium combined. Merus and propodus expanded, propodus with dense 
row of plumose setae on inner and outer margins; dactyl relatively long and 
narrow. Uropod with basis as long as endopod; exopod three-fourths length of 
endopod. 

Morphological Affinities with S. caribbea 

In terms of pleotelson configuration, S. myriamae exhibits a superficial resem- 
blance to S. unicornalis and in fact, to all member species of the Bg Storthyngura 
group according to the scheme proposed by George and Menzies (1968b). How- 
ever, this species is so unique in possessing a transverse ridge in somites two to 
four which is posterior to the prominent median dorsal spines in all three somites. 
This transverse ridge also exhibits a small median tubercle. The only other species 
in the genus Storthyngura that has a similar transverse ridge with tubercles on 
the second through fourth somites is S. caribbea which was originally described 
by Benedict as a species belonging to the genus Eurycope; subsequently, 
Richardson (1905) redescribed this species. Wolff (1962) properly affiliated this 
species to the genus Storthyngura but treated it as a subspecies of S. pulchra. 
George and Menzies (1968a, b) offered reasons for distinguishing S. caribbea as 
a distinct species. 

We believe that the new species S. myriamae, found in the Angola Basin at 
5220 m, is somehow very closely related to S. caribbea which occurs off the 
Windward Island in the West Indies at a depth of 1260 m. We examined the type 
specimen from the U.S. National Museum (courtesy of Dr. Tom Bowman), com- 
pared the two species, and arrived at the following major morphological differ- 
ences between the two species. 

Close observation of ^. myriamae and S. caribbea revealed a striking similarity 
in the shape of the head and general body form. One major difference is found 
in the second somite; the antero-lateral angle is produced in S. caribbea but 
rounded in S. myriamae. Also, comparison of the mouthparts showed conspic- 
uous anatomical differences in the mandible and maxilliped. The palp of the 



VOLUME 95, NUMBER 1 97 

maxilliped in S. caribbea has a short basal article widened at its base; S. myria- 
mae has a palp with an elongated basal article that is narrow at its base. 

Both S. caribbea and S. myriamae have single non-lobed coxal plates on the 
first somite, but S. caribbea has coxal plates that are far more produced than 
those found in S. myriamae. Storthyngura caribbea has elongated dorsal spines 
while S. myriamae has spines of a moderate length. There are also differences 
between the two species in the shape of the first pleonal somite and the pleotelson. 
In S. myriamae the median dorsal length of the first flat pleonal somite is two 
times that of its lateral extremity, but in S. caribbea it is of even length through- 
out. 

We believe that S. myriamae shows a close affinity with S. caribbea, but based 
on the morphological differences, S. myriamae is a distinct new species. 

Distribution: Angola Basin vs. Cape Basin 

The primary purpose of the French deep-sea expedition was to look for any 
faunal differences at the abyssal depths between the Angola Basin (on the western 
side of the Walvis Ridge) and the Cape Basin (on the eastern side of the Walvis 
Ridge, see Fig. 2). Both basins contained oligotrophic clay-Hke ooze (Sibuet 1980) 
but the Angola Basin is rich in CaCOg (70%) and the Cape Basin is poor in CaCOs 
(not more than 5%). In terms of bottom currents, the Angola Basin is subject to 
possible inflow of the Antarctic Bottom Water whereas the Cape Basin is blocked 
by the Walvis Ridge which possibly acts as a topographic and therefore biogeo- 
graphic barrier for the expansion of deep-sea fauna from the Weddell Sea-Scotia 
Sea Antarctic region. This geographic zone was considered as a center of origin 
for the genus Storthyngura (George, 1980). It is also known that the Antarctic 
Bottom Water extends north even beyond the equator into the North Atlantic 
and abyssal regions off the West Indies. 

In this present study we have carefully examined the isopod material from the 
deep-sea stations on both sides of the Walvis Ridge, four stations from the Angola 
Basin and six stations from Cape Basin (Fig. 2). We encountered Storthyngura 
myriamae only in the Angola Basin occurring at all four stations. Storthyungura 
myriamae was not found in any one of the six stations at comparable depths in 
the Cape Basin. It is of interest to point out that S. myriamae belongs to Stor- 
thyngura Bg group which includes six species of similar pleotelson configuration 
{S.fragilis from the North Pacific, S. caribbea from the North Atlantic, S. uni- 
cornalis from the South Pacific, S. gordonae from the Indian Ocean, and S. 
challengeri from the Antarctic Ocean). Although the Bg group is not represented 
in the Cape Basin, C-group (5. symmetrica) and D-group {S. triplispinosa) are 
represented in the Cape Basin (George and Menzies 1968b). Such a pattern of 
difference in biogeographic distribution between B-group and C & D-groups sug- 
gests apparently different evolutionary pathways for the species within the genus 
Storthyngura. 

This new species from the Angola Basin shows close morphological similarity 
with S. caribbea from the West Indies region which is separated from the Angola 
Basin by a series of abyssal basins such as the Guinea Basin, Sierra Leone Basin, 
Cape Verde Basin, and Guiana Basin. These basins have a deep sill-depth and 
therefore, faunal affiliation between these basins at the abyssal depths should be 



98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

carefully investigated. However, our knowledge of the abyssal zoogeography 
and faunistic link between these basins is very limited due to paucity of deep-sea 
samples. 

It is of interest to point out that S. myriamae is also related to S. challenged 
from the Antarctic Ocean. We can not, however, offer any explanation of the 
evolutionary directions and speciation pattern of these closely related B-group 
Storthyngura species. On the basis of plesiomorphic features, the Antarctic slope 
species from the Weddell Sea region appear to be somewhat primitive (George 
1980), although it is difficult to point out which one is the most primitive or 
parental species. Nevertheless, the global distribution pattern of Storthyngura 
species suggests that their speciation and radiation into the abyssal regions of the 
World Oceans since Miocene period is possibly promoted by the flow of the 
Antarctic Bottom Water which originates in the Weddell Sea region. This hy- 
pothesis obviously calls for careful reevaluation and further investigation. 

Literature Cited 

George, R. Y. 1980. Antarctica as a center of origin for the deepsea isopod genus Storthyngura. 
Symposium on Biology and Evolution of Crustacea, Sydney, Australia (May, 1980). 

, and R. J. Menzies. 1968a. Species of Storthyngura (Isopoda) from the Antarctic with de- 
scriptions of six new species. — Crustaceana 14(3):275-301. 

, and . 1968b. Distribution and probable origin of the species in the deep-sea isopod 

genus Storthyngura. — Crustaceana 15(2): 171-187. 

Richardson, H. E. 1905. A monograph on the isopods of North America. — Bulletin of the United 
States National Museum 54: 1-727. 

Sibuet, M. 1980. A report on Campagne Walvis I, JEAN CHARCOT expedition. December 12, 1978 
to January 15, 1979. (Unpublished Report.) 

Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda, Asellota. — Galathea 
Report 6:7-320. 

Institute for Marine Biomedical Research, University of North Carolina at 
Wilmington, Wilmington, North Carolina 28403. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 99-113 

PHYTOPLANKTON DISTRIBUTION ALONG THE EASTERN 

COAST OF THE USA IV. SHELF WATERS BETWEEN 

CAPE LOOKOUT, NORTH CAROLINA, AND 

CAPE CANAVERAL, FLORIDA 

Harold G. Marshall 

Abstract. — The phytoplankton composition is discussed for southeastern shelf 
waters of the United States with the average concentrations at near and far shore 
stations given for 328 species. A mixed ultraplankton group, composed of un- 
identified species, predominated in numbers, and was part of the shelf assem- 
blage of dominant forms with diatoms, dinophyceans, and haptophyceans. Larg- 
est cell concentrations were noted near shore with the diatoms and the 
ultraplankton component most abundant. Phytoplankton assemblages are given 
for near and far shore stations over the shelf. 



Phytoplankton composition in the continental shelf waters off the southeastern 
coast of the United States has been discussed by Hulburt (1967), Marshall (1969, 
1971), and Hulburt and MacKenzie (1971). These studies have indicated diatoms 
abundant in the shelf populations with the coccolithophorids and other phytofla- 
gellates becoming proportionally more significant beyond the shelf break and in 
the Gulf Stream. In addition to diatoms, dinophyceans, and coccolithophores, 
representatives from a few other phytoplankton groups have been found common 
in these waters. Hulburt and MacKenzie (1971) noted large concentrations of the 
cryptophycean Rhodomonas amphioxeia south and north of Cape Hatteras. Dun- 
stan and Hosford (1977) reported an abundance of the cyanophyceans Oscillatoria 
thiebautii and Oscillatoria erythraea in the south Atlantic bight, with Marshall 
(1981) noting 16 cyanophycean species in these shelf waters. Bishop et al. (1980) 
indicated little seasonal change in nutrient or phytoplankton concentrations in 
this area. They emphasized more variability occurred on the outer shelf which 
may be related to intrusions of the Gulf Stream and upwelling along the shelf 
break. They also suggested a short-term response time of days to weeks by the 
phytoplankton to these events and that responses may differ throughout the year, 
resulting in changes in the concentrations among the various populations. 

These past studies suggest this section of the continental shelf contains a di- 
verse phytoplankton flora with regional variations and changes in population con- 
centrations common. The purpose of this study was to evaluate the phytoplankton 
composition in this area of approximately 600 km along the shelf between Cape 
Lookout, N.C. and Cape Canaveral, Florida. Emphasis was placed on charac- 
terizing the populations from near and far shore stations over this section of the 
continental shelf. 

Methods 

Surface water samples were taken from 24 October to 16 November 1973 and 
6-20 September 1978 during MARMAP cruises of the South Carolina Marine 



100 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Resources Program. These collections were generally taken along transects di- 
rected from the coastal area seaward to the vicinity of the shelf break, between 
Cape Lookout, N.C. and Cape Canaveral, Florida (Fig. 1). There were 43 stations 
in the 1973 collections and 48 stations in 1978. Station depths ranged from 9 
m to 318 m, with 45 of the 91 stations located at water depths of less 
than 44 m. There were four stations in waters between 200 and 318 m deep. 
Reference to near shore stations are those located within 35 km from the 
nearest coastline, with those beyond this distance referred to as far shore stations. 
In this study 35 of the 91 stations were classified as near shore stations. 

At each station a 500 ml surface sample was obtained with a Van Dorn collec- 
tion bottle using standard hydrocast procedures. The samples were preserved 
immediately with buffered formalin and returned to the laboratory for subsequent 
settling. A modified Utermdhl method was used with the samples siphoned to a 
20 ml concentrate, transferred to a settling chamber and examined with a Zeiss 
inverted plankton microscope. The classification format of Hendey (1974), Parke 
and Dixon (1976), Drouet and Daily (1956), and Drouet (1968) is mainly followed 
in this study. Salinity values and other station data were provided by personnel 
from the South Carolina Marine Resources Program. 

Results 

There were differences in the range and average values for temperature and 
salinity for the two cruises (Table 1). The 1973 collections came from water that 
was cooler and slightly less saline, with a broader temperature range represented 
than what was present in the 1978 samples. The combined average temperature 
for both of these cruises was 26.42°C, which was higher than the average fall 
temparature (22.4°C) during the previous collections in this area by Marshall 
(1971). 

A total of 328 phytoplankters was identified in this study from the two cruises 
(Table 2). These consisted of Bacillariophyceae (194), Dinophyceae (83), Hap- 
tophyceae (13), Cyanophyceae (16), Euglenophyceae (7), Prasinophyceae (4), 
Chlorophyceae (3), Chrysophyceae (5), and Cryptophyceae (3). In addition, there 
were high concentrations of an unidentified ultraplankton component that was 
divided into three size categories, <3 fjum, 3-5 /xm, 5-10 /itm. These consisted of 
round, oval, and irregularly shaped cells, with apparent light green color. Most 
numerous was the <3 ptm size class which appeared similar to coccoid cyano- 
phyceans, whereas the 3-5 ^tm category resembled chlorophyceans with the 5-10 



Table I. — Temperature and salinity values for surface water at stations during the fall 1973 and 
1978 cruises. 



Temperature °C 



Salinity %c 





1973 


1978 


1973 


1978 


Range 


14.9-27.8 


27.2-28.6 


34.22-37.07 


34.87-38.81 


Mean 


24.1 


27.8 


35.91 


36.04 


Near Shore Mean 


23.2 


27.8 


35.69 


35.84 


Far Shore Mean 


26.1 


27.8 


36.03 


36.10 



VOLUME 95, NUMBER 1 



101 



80" 



3£ 



33" 



32° 



31: 



20! 



29° 



28° 



27° 



26° 





CAPE CANAVERAL 



ATLANTIC 
OCEAN 





Fig. I. Station locations for water samples along the southeastern continental shelf taken Octo- 
ber-November 1973 (®) and September 1978 (•). 



102 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Phytoplankton composition and average cell concentrations (no's/1) for combined stations 
in the 1973 and 1978 cruises. Counts for the filamentous cyanophyceans are in filaments per liter. 



Stations 



Near shore 



Far shore 



BACILLARIOPHYCEAE 

Achnanthes longipes Agardh 

Actinoptychus senarius Ehrenberg 

Actinoptychus vulgaris Schumann 

Amphiprora sp. 

Amphiprora alata (Ehrenberg) Kutzing 

Amphora sp. 

Amphora binodis Gregory 

Amphora cuneata Cleve 

Amphora egregia var. interrupta Peragallo & Peragallo 

Amphora grevilleana contracta Cleve 

Amphora marina (W. Smith) Van Heurck 

Amphora obtusa Gregory 

Amphora ostrearia Brebisson 

Amphora ovalis Kutzing 

Amphora peragalli Cleve 

Amphora t err oris Ehrenberg 

Asterionella glacialis Castracane 

Asterionella kariana Grunow 

Asteromphalus flabellatus (Brebisson) Greville 

Auliscus sculpt us (W. Smith) Ralfs 

Bacteriastrum comosum Pavillard 

Bacteriastrum delicatulum Cleve 

Bacteriastrum hyalinum Lauder 

Bacteriastrum hyalinum var. princeps (Castracane) Ikari 

Bacteriastrum varians Lauder 

Biddulphia alternans (Bailey) Van Heurck 

Biddulphia aurita (Lyngbye) Brebisson 

Biddulphia longicruris Greville 

Biddulphia mobiliensis (Bailey) Grunow 

Biddulphia sinensis Greville 

Biddulphia tridens (Ehrenberg) Ehrenberg 

Campylodiscus limbatus Brebisson 
Campylodiscus rutilis Skvortzow 
Chaetoceros affine Lauder 
Chaetoceros atlanticum Cleve 
Chaetoceros breve Schutt 
Chaetoceros coarctatum Lauder 
Chaetoceros compressum Lauder 
Chaetoceros convolutum Castracane 
Chaetoceros curvisetum Cleve 
Chaetoceros decipiens Cleve 
Chaetoceros densum Cleve 
Chaetoceros didymum Ehrenberg 
Chaetoceros diver sum Cleve 
Chaetoceros gracile Schutt 
Chaetoceros laciniosum Schutt 
Chaetoceros lorenzianum Grunow 
Chaetoceros messanense Castracane 
Chaetoceros pelagicum Cleve 
Chaetoceros pendulum Karsten 



0.29 
0.57 
0.19 

0.38 
7.90 
8.38 



0.09 
0.09 
0.09 

0.09 

0.87 



— 


0.04 


— 


0.17 


0.38 


1.04 


0.19 


0.09 


4.00 


0.52 


0.95 


0.70 


— 


0.17 


4.57 


— 


3.05 


2.09 


— 


4.00 


2.10 


0.43 


— 


0.09 


— 


1.13 


4.38 


9.74 


3.62 


12.91 


— 


0.17 


— 


0.96 


97.05 


0.57 


4.67 


0.17 


0.57 


0.52 


2.95 


0.52 


0.10 


0.04 


0.76 


— 


— 


0.09 


— 


0.04 


17.32 


1.0 


6.48 


6.26 


29.52 


28.48 


10.86 


3.45 


— 


0.70 


— 


0.52 


— 


0.35 


63.24 


25.87 


0.38 


1.13 


— 


0.09 


22.29 


3.48 


— 


0.35 


— 


0.26 


22.86 


12.17 


0.57 


— 


11.43 


— 



1.04 



VOLUME 95, NUMBER 1 



103 



Table 2. — Continued. 



Stations 



Near shore 


Far shore 


6.10 


6.39 


2.86 


— 


— 


0.43 


— 


1.65 


— 


0.35 


35.29 


23.91 


5.71 


— 


1.14 


— 


9.14 


1.22 


0.38 


0.96 


0.38 


— 


0.76 


0.09 


5.81 


0.61 


0.38 


— 


9.14 


1.13 


4.95 


2.74 


29.05 


3.61 


28.95 


0.61 


64.00 


9.00 





0.09 



Chaetoceros peruvianum Brightwell 

Chaetoceros pseudocurvisetum Mangin 

Chaetoceros radians Schutt 

Chaetoceros sociale Lauder 

Chaetoceros wighami Brightwell 

Climacodium frauenfeldianum Grunow 

Cocconeis sp. 

Cocconeis molesta var. crucifera Grunow 

Cocconeis pinnata Gregory 

Corethron criophilum Castracane 

Coscinodiscus argus Ehrenberg 

Coscinodiscus asteromphalus Ehrenberg 

Coscinodiscus centralis Ehrenberg 

Coscinodiscus gigas Ehrenberg 

Coscinodiscus grani Gough 

Coscinodiscus granulosus Grunow 

Coscinodiscus lineatus Ehrenberg 

Coscinodiscus marginatus Ehrenberg 

Coscinodiscus nitidus Gregory 

Coscinodiscus nobilis Grunow 

Coscinodiscus perforatus Ehrenberg 

Coscinodiscus radiatus Ehrenberg 

Coscinodiscus stellaris var. symbolophora (Grunow) Jorgensen 

Coscinodiscus wailesii Gran and Angst 

Cyclotella sp. 

Cyclotella meneghiniana Kutzing 

Cylindrotheca closterium (Ehrenberg) Reiman and Lewin 

Cymatosira belgica Grunow 

Cymatosira lorenziana Grunow 

Dactyliosolen antarcticus Castracane 
Dactyliosolen mediterraneus Peragallo 
Diploneis crabro Ehrenberg 
Diploneis crabro var. pandura (Brebisson) Cleve 
Ditylum brightwellii (West) Grunow 

Eucampia zoodiacus Ehrenberg 

Eunotia sp. 

Eunotia bidentula W. Smith 

Fragilaria sp. 

Fragilaria crotonensis Kitton 

Fragilariopsis cylindrus (Grunow) Helmcke and Krieger 

Grammatophora sp. 

Grammatophora angulosa Ehrenberg 

Grammatophora marina (Lyngbye) Kutzing 

Guinardia flaccida (Castracane) Peragallo 

Gyrosigma sp. 

Gyrosigma balticum (Ehrenberg) Cleve 

Hantzschia marina (Donkin) Grunow 
Hemiaulus hauckii Grunow 
Hemiaulus membranaceus Cleve 
Hemiaulus sinensis Greville 



0.38 



0.09 



0.38 


— 


0.38 


— 


— 


6.35 


— 


0.13 


63.05 


21.36 


151.05 


33.83 


18.29 


1.13 


— 


0.87 


— 


3.39 


29.52 


7.48 


11.24 


0.35 


14.76 


0.65 


— 


0.09 


0.10 


— 


— 


0.09 


— 


0.52 


— 


0.09 


334.29 


13.30 


37.52 


18.04 


0.76 


5.30 


— 


0.43 


50.48 


41.91 


— 


0.09 


0.38 


— 


0.19 


— 


23.24 


34.48 


49.52 


9.83 


47.81 


26.80 



104 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Continued. 



Stations 



Leptocylindrus danicus Cleve 

Licmophora sp. 

Licmophora flabellata (Carmichael) Agardh 

Mastogloia smithii Thwaites 

Melosira distans (Ehrenberg) Kutzing 

Melosira granulata (Ehrenberg) Ralfs 

Melosira granulata var. angustissima Muller 

Melosira islandica Muller 

Melosira moniliformis (MuUer) Agardh 

Melosira nummuloides (Dillwyn) Agardh 

Navicula sp. 

Navicula abrupta (Gregory) Cleve 

Navicula annulata Grunow 

Navicula cancellata Donkin 

Navicula clavata Gregory 

Navicula for cipata Greville 

Navicula lyra Ehrenberg 

Navicula opima (Grunow) Cleve 

Navicula praetexta Ehrenberg 

Navicula pusilla W. Smith 

Nitzschia sp. 

Nitzschia angularis W. Smith 

Nitzschia distans Gregory 

Nitzschia insignis Gregory 

Nitzschia longissima (Brebisson) Ralfs 

Nitzschia lorenziana var. densistriata (Peragallo and Peragallo) Hustedt 

Nitzschia lorenziana var. incerta Grunow 

Nitzschia lorenziana var. subtilis Grunow 

Nitzschia panduriformis Gregory 

Nitzschia pungens Grunow 

Nitzschia seriata Cleve ^- 

Nitzschia sigma (Kutzing) W. Smith 

Nitzschia sigma var. intercedens Grunow 

Nitzschia socialis Ralfs 

Nitzschia spathulata Brebisson 

Paralia sulcata (Ehrenberg) Cleve 

Pinnularia sp. 

Plagiogramma staurophorum (Gregory) Heilberg 

Plagiogramma vanheurckii Grunow 

Planktoniella sol (Wallich) Schutt 

Pleurosigma sp. 

Pleurosigma angulatum (Quekett) W. Smith 

Pleurosigma elongatum W. Smith 

Pleurosigma hamuliferum Brun 

Pleurosigma nicobaricum (Grunow) Grunow 

Pleurosigma normanii Ralfs 

Pleurosigma obscurum W. Smith 

Podosira stelliger (Bailey) Mann 

Rhaphoneis amphiceros Ehrenberg 
Rhaphoneis surirella (Ehrenberg) Grunow 
Rhizosolenia acuminata (Peragallo) Gran 



Near shore 


Far shore 


19.24 


29.17 


0.19 


0.39 


3.81 


0.09 


10.29 


0.48 


49.81 


29.57 


17.33 


12.09 


22.14 


85.74 


— 


4.75 


— 


0.04 


— 


1.57 


0.38 


0.13 


— 


0.26 


55.05 


6.83 


16.57 


1.74 


16.19 


1.65 


25.33 


0.52 


15.71 


0.70 


0.38 


6.35 


0.09 


— 


— 


0.17 


— 


0.35 


4.19 


0.43 


9.52 


0.61 


1.52 


— 


16.19 


8.00 


It 1.14 


1.39 


0.76 


0.78 


0.76 


— 


0.76 


0.43 


56.19 


2.96 


— 


6.78 


0.38 


— 


— 


0.87 


— 


0.17 


4.29 


— 


531.05 


87.09 


— 


0.26 


448.00 


25.57 


— 


0.70 


— 


0.43 


0.19 


0.52 


5.24 


2.09 


3.05 


1.09 


30.29 


4.51 


1.71 


— 


3.43 


— 


1.52 


— 


0.19 


— 


3.43 


1.22 


9.14 


3.13 


2.29 


0.26 



VOLUME 95, NUMBER 1 



105 



Table 2. — Continued. 



Stations 



Near shore 


Far shore 


713.52 


336.46 


99.62 


19.65 


1029.62 


404.04 


1.33 


0.52 


37.62 


31.65 


0.57 


2.52 


0.76 


7.26 


0.57 


3.30 


0.38 


0.09 


— 


2.17 


53.33 


23.28 


3.71 


5.30 


84.94 


14.61 


37.81 


167.78 


25.24 


11.96 


0.10 


1.83 


— 


0.78 


36.10 


0.96 


3.43 


0.43 


32.67 


8.04 


— 


0.09 


0.19 


— 


1.90 


1.04 


70.10 


27.48 


0.95 


— 


0.38 


— 


— 


0.09 


0.19 


0.09 


133.52 


1.00 


97.52 


25.22 


808.67 


144.70 


— 


0.43 


13.62 


2.09 


4.38 


1.74 


25.71 


1.61 


— 


0.13 


— 


1.30 


99.43 


19.61 


12.00 


6.63 


0.10 


0.13 


0.19 


— 


0.19 


0.04 


0.57 


0.35 


201.14 


54.43 


0.38 


8.78 


— 


0.04 


— 


0.91 


— 


0.52 





0.39 



Rhizosolenia alata Brightwell 
Rhizosolenia alata f. gracillima (Cleve) Grunow 
Rhizosolenia alata f. indie a (Peragallo) Gran 
Rhizosolenia bergonii Peragallo 
Rhizosolenia calcar-avis Schultze 
Rhizosolenia castracanei Peragallo 
Rhizosolenia delicatula Cleve 
Rhizosolenia fragilissima Bergon 
Rhizosolenia hebetata f. hiemalis Gran 
Rhizosolenia hebetata f. semispina (Hensen) Gran 
Rhizosolenia imbricata Brightwell 
Rhizosolenia robusta Norman 
Rhizosolenia setigera Brightwell 
Rhizosolenia stolterfothii Peragallo 
Rhizosolenia styliformis Brightwell 
Rhizosolenia temperei Peragallo 

Schroederella delicatula (Peragallo) Pavillard 

Skeletonema costatum (Greville) Cleve 

Stephanopyxis palmeriana (Greville) Grunow 

Stephanopyxis turns (Greville) Ralfs 

Surirella crumena Brebisson 

Surirella pandura var. contracta Peragallo and Peragallo 

Synedra crystallina (Agardh) Kutzing 

Synedra fulgens (Greville) W. Smith 

Synedra gaillonii (Bory) Ehrenberg 

Synedra robusta Ralfs 

Synedra tabulata (Agardh) Kutzing 

Synedra toxoneides Castracane 

Synedra undulata Bailey 

Tabellaria fenestrata var. asterionelloides Grunow 
Thalassionema nitzschioides Hustedt 
Thalassiosira baltica (Grunow) Ostenfeld 
Thalassiosira eccentrica (Ehrenberg) Cleve 
Thalassiosira gravida Cleve 
Thalassiosira nordenskioeldii Cleve 
Thalassiosira subtilis (Ostenfeld) Gran 
Thalassiothrix delicatula Cupp 
Thalassiothrix frauenfeldii Grunow 
Thalassiothrix mediterranea Pavillard 
- Triceratium favus Ehrenberg 
Triceratium formosum var. pentagonalis (Schmidt) Hustedt 
Tropidoneis lepidoptera (Gregory) Cleve 
Tropidoneis seriata Cleve 

Unidentified diatoms 
DINOPHYCEAE 

Amphidinium acutissimum Schiller 
Amphidinium acutum Lachmann 
Amphidinium bipes Herdman 
Amphidinium globosum Schroder 
Amphidinium klebsii Kofoid and Swezy 



106 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Continued. 



Stations 



Near shore 


Far shore 





0.04 


— 


3.04 


0.19 


0.04 


0.19 


0.39 


— 


0.04 


0.19 


0.48 


11.90 


3.83 


7.71 


0.93 


0.38 


— 


0.76 


— 


— 


0.09 


4.76 


3.30 


0.19 


— 


0.38 


— 


3.33 


0.83 


— 


0.09 


0.38 


1.83 


— 


0.09 


— 


0.17 


— 


0.22 


2.48 


3.43 


0.48 


0.39 


0.57 


0.78 


11.52 


2.22 


— 


0.26 


— 


0.43 


— 


0.09 


0.76 


0.09 


29.14 


— 


— 


0.70 


— 


0.35 


— 


0.09 


0.76 


7.87 


1.52 


0.57 


1.71 


0.09 


— 


0.17 


— 


1.22 


36.57 


0.70 


— 


1.17 


— 


0.70 


— 


0.09 


15.7 


12.85 


— 


0.61 


0.19 


— 


— 


0.22 


— 


0.09 


— 


0.61 


0.19 


— 


— 


0.09 


— 


0.09 


17.90 


0.52 



Amphidinium lanceolatum Schroder 
Amphidinium schroederi Schiller 

Ceratium contortum (Gourret) Cleve 

Ceratium contortum var. karsteni (Pavillard) Soumia 

Ceratium digitatum Schutt 

Ceratium extensum (Gourret) Cleve 

Ceratium f urea (Ehrenberg) Claparede and Lachmann 

Ceratium fusus (Ehrenberg) DuJardin 

Ceratium geniculatum (Lemmermann) Cleve 

Ceratium horridum (Cleve) Gran 

Ceratium kofoidi Jorgensen 

Ceratium lineatum (Ehrenberg) Cleve 

Ceratium longirostrum Gourret 

Ceratium macroceros (Ehrenberg) VanHoffen 

Ceratium massiliense (Gourret) Jorgensen 

Ceratium minutum Jorgensen 

Ceratium pentagonum Gourret 

Ceratium ranipes Cleve 

Ceratium setaceum Jorgensen 

Ceratium teres Kofoid 

Ceratium trichoceros (Ehrenberg) Kofoid 

Ceratium tripos (MuUer) Nitzsch 

Ceratium tripos var. atlanticum (Ostenfeld) Paulsen 

Dinophysis caudata Kent 
Glenodinium sp. 
Gonyaulax diegensis Kofoid 
Gonyaulax fragilis (Schutt) Kofoid 
Gonyaulax minuta Kofoid and Michener 
Gonyaulax monilata Howell 
Gymnodinium sp. #1 
Gymnodinium sp. #2 
Gymnodinium coeruleum Dogiel 
Gymnodinium danicans Campbell 
Gymnodinium variabile Herdman 
Gyrodinium sp. 

Gyrodinium dominans Hulburt 
Gyrodinium estuariale Hulburt 
Gyrodinium fusiforme Kofoid and Swezy 
Heterocapsa triquetra (Ehrenberg) Stein 

Katodinium sp. 

Katodinium asymetricum (Massart) Fott 

Katodinium rotundatum (Lohmann) Loeblich 

Oxytoxum gracile Gran 

Oxytoxum sceptrum (Stein) Schroder 

Oxytoxum variabile Schiller 

Peridiniopsis assymetrica Mangin 
Podolampas bipes Stein 
Podolampas elegans Schutt 
Podolampas palmipes Stein 
Podolampus curvatus Schiller 
Prorocentrum aporum (Schiller) Dodge 



VOLUME 95, NUMBER 1 



107 



Table 2. — Continued. 



Stations 



Near shore 



Far shore 



Prorocentrum balticum (Lohmann) Loeblich 
Prorocentrum cassubicum (Woloszynska) Dodge 
Prorocentrum compressum (Bailey) Abe 
Prorocentrum gracile Schutt 
Prorocentrum lima (Ehrenberg) Dodge 
Prorocentrum maximum (Gourret) Schiller 
Prorocentrum micans Ehrenberg 
Prorocentrum nanum Schiller 
Protoperidinium sp. 

Protoperidinium biconicum (Dangeard) Balech 
Protoperidinium breve (Paulsen) Balech 
Protoperidinium brochii (Kofoid and Swezy) Balech 
Protoperidinium cerasus (Paulsen) Balech 
Protoperidinium claudicans (Paulsen) Balech 
Protoperidinium conicum (Gran) Balech 
Protoperidinium crassipes Kofoid 
Protoperidinium depressum (Bailey) Balech 
Protoperidinium divergens (Ehrenberg) Balech 
Protoperidinium grande (Kofoid) Balech 
Protoperidinium leonis (Pavillard) Balech 
Protoperidinium oceanicum (VanHoffen) Balech 
Protoperidinium pendunculatum (Schutt) Balech 
Protoperidinium pentagonum (Gran) Balech 
Protoperidinium quarnerense (Schroder) Balech 
Protoperidinium solidicorne (Mangin) Balech 
Protoperidinium sphaericum (Okamura) Balech 
Protoperidinium steinii (Jorgensen) Balech 

Unidentified dinoflagellate cysts 
Unidentified dinoflagellates 

HAPTOPHYCEAE 

Acanthoica aculeata Kamptner 

Chrysochromulina sp. 

Cyclococcolithus leptoporus (Murray and Blackman) Kamptner 

Emiliania huxleyi (Lohmann) Hay and Mohler 

Gephyrocapsa oceanica Kamptner 

Hymenomonas carterae (Braarud and Fagerland) Braarud 

Rhabdosphaera claviger Murray and Blackman 
Rhabdosphaera hispida Lohmann 
Rhabdosphaera sty lifer Lohmann 
Syracosphaera sp. 
Syracosphaera molischii Schiller 
Syracosphaera pirus Halldal and Markali 
Syracosphaera pulchra Lohmann 

Unidentified coccolithophorids 
CHRYSOPHYCEAE 

Calycomonas ovalis Wulff 

Dictyocha fibula Ehrenberg 

Distephanus speculum (Ehrenberg) Haekel 

Ochromonas sp. 

Ochromonas caroliniana Campbell 



0.76 



9.43 



0.22 
0.09 
1.04 
0.17 



0.38 


— 


0.76 


— 


168.48 


9.09 


6.38 


3.83 


— 


0.09 


— 


0.04 


0.19 


0.13 


0.10 


0.09 


1.14 


0.09 


0.19 


0.09 


2.29 


0.26 


— 


0.35 


5.14 


0.78 


0.76 


— 


— 


0.04 


0.38 


— 


— 


0.35 


0.57 


0.04 


0.57 


0.09 


— 


0.17 


— 


0.04 


0.38 


0.43 


— 


0.96 


0.19 


0.70 


62.10 


49.87 


0.76 


0.43 


— 


0.09 


14.29 


28.57 


50.86 


60.48 


— 


12.96 


0.38 


0.09 


3.05 


1.04 


3.43 


— 


15.62 


— 


0.19 


— 


— 


0.09 


0.76 


0.26 


1.90 


2.70 


469.90 


127.48 


0.09 





50.14 


4.00 


0.38 


0.35 


158.29 


3.22 


0.19 






108 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Continued. 



Stations 



Near shore 



Far shore 



CYANOPHYCEAE 

Agmenellum quadruplicatum (Meneghini) Brebisson 
Agmenellum thermale (Kutzing) Drouet and Daily 
Anacystis aeruginosa Drouet and Daily 
Anacystis dimidiata (Kutzing) Drouet and Daily 
Anacystis marina (Hansg) Drouet and Daily 

Entophysalis deusta (Meneghini) Drouet and Daily 

Gomphosphaeria aponina Kutzing 

Johannesbaptistia pellucida (Dickie) Taylor and Drouet 

Nostoc commune Vaucher 

Oscillatoria sp. 

Oscillatoria erythraea (Ehrenberg) Kutzing 

Oscillatoria submembranacea Ardissone and Strafforello 

Richelia intracellularis Schmidt 

Schizothrix calcicola (Agardh) Gomont 
Schizothrix tenerrima (Domont) Drouet 
Spirulina subsalsa Oersted 

EUGLEN OPH YCE AE 

Euglena sp. 

Euglena ehrenbergii Klebs 

Euglena fusca (Klebs) Lemmermann 

Eutreptia lanowii Steuer 

Eutreptia viridis Perty 

Trachelomonas sp. 

Trachelomonas hispida (Perty) Stein 

CHLOROPHYCEAE 

Chlorella sp. 

Crucigenia tetrapedia (Kirchner) West and West 

Staurastrum quadricuspidatum Turner 

PRASINOPHYCEAE 

Pyramimonas sp. 

Pyramimonas torta Conrad and Kuff 

Tetraselmis sp. 

Tetraselmis gracilis (Kylin) Butcher 

CRYPTOPHYCEAE 

Chroomonas sp. 
Cryptomonas sp. 
Cryptomonas sp. #2 

OTHERS 

Unidentified green cells (<3.0 microns) 
Unidentified green cells (3-5 microns) 
Unidentified green cells (5-10 microns) 



12.38 


— 


4.19 


— 


38.48 


— 


87.62 


12.43 


543.05 


29.83 


2.29 


— 


45.81 


21.43 


395.85 


35.83 


500.95 


126.67 


0.76 


— 


119.90 


116.87 


17.52 


1.74 


— 


3.13 


1.52 


— 


2.29 


— 


0.57 


— 


0.19 




0.19 


— 


0.19 


1.22 


1.71 


0.35 


0.95 


0.17 


0.10 


0.09 


0.38 


0.17 


8.57 


1.04 


1.90 


0.87 


0.76 


— 


0.19 




— 


0.09 


0.67 


— 


— 


0.09 


0.09 




0.38 


3.39 


0.09 


— 


32,154.76 


12,844.57 


105.95 


290.22 


3.05 


2.61 



VOLUME 95, NUMBER 1 109 

Table 3. — Average concentrations of cells (no's/1) for the various phytoplankton groups at near and 
far shore stations in 1973 and 1978. 





Near shore 


Far shore 






1973 1978 


1973 1978 


Combined 


Bacillariophyceae 


7850 4753 


1656 2073 


3161 


Dinophyceae 


621 83 


160 53 


185 


Haptophyceae 


949 113 


78 126 


227 


Chrysophyceae 


89 <1 


10 <1 


17 


Cyanophyceae 


2513 1037 


396 299 


778 


Euglenophyceae 


8 1 


<1 <1 


2 


Chlorophyceae 


<1 17 


<1 2 


3 


Cryptophyceae 


<1 <1 


3 3 


2 


Prasinophyceae 


1 <1 


<1 <1 


<1 


Ultraplankton* 


4952 56,642 


4158 20,435 


17,322 



* Combined size groups of unidentified cells less than 10 microns in size. 

/xm group more indiscriminate to a specific taxonomic group. In addition to these 
non-flagellated forms, there were also some phytoflagellates included in these size 
categories. An ultraplankton component to estuarine and marine habitats has 
been recognized in recent years (Malone 1971; McCarthy et al. 1974; among 
others). These cells may include the cyanobacteria (blue-green algae) found 
widely distributed in the western north Atlantic by Waterbury et al. (1979), and 
Johnson and Sieburth (1979). They are also similar to those found in the Chesa- 
peake Bay plume by Marshall (in press) and off the northeastern U.S. coast by 
Marshall and Cohn (1981). 

The study area represents a broad, crescent-shaped segment of the southeast- 
ern continental shelf, that reaches its greatest width in the area between Jack- 
sonville and Savannah. In this region the shelf break is approximately 120 km 
from the coast. The phytoplankton populations varied over the shelf with distinct 
groups more characteristic of either the near or far shore stations. Average total 
concentration of cells was generally greater near shore as was the presence of 
the taxonomic groups represented in the samples, with the exception of the cryp- 
tophyceans, haptophyceans, and an unidentified ultraplankton component (Table 
3). During both cruises, the far shore populations of the cryptophyceans, although 



Table 4. — Numbers and percentages of species within each group that were noted limited to near 
and far shore stations, or found at both stations. 







Only at 


Only at 






Total 


near shore 


far shore 


In both areas 


Bacillariophyceae 


194 


*32 17% 


53 27% 


109 56% 


Dinophyceae 


83 


12 14% 


39 46% 


32 40% 


Haptophyceae 


13 


3 23% 


4 31% 


6 46% 


Chrysophyceae 


5 


2 40% 





3 60% 


Cyanophyceae 


16 


8 50% 


1 6% 


7 44% 


Euglenophyceae 


7 


2 29% 





5 71% 


Chlorophyceae 


3 


1 33% 





2 67% 


Cryptophyceae 


3 


2 67% 





1 33% 


Prasinophyceae 


4 


2 50% 


2 50% 






110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

not high, were greater than what was found at near shore stations. However, the 
total phytoplankton composition was dominated by diatoms, dinoflagellates, hap- 
tophyceans, cyanophyceans, and the ultraplankton component during both cruis- 
es over the shelf. The total near shore populations included more of the smaller 
sized diatoms, whereas over the central and far shelf, larger sized diatoms were 
abundant. Of the 194 diatoms, 56% of the species were found at both near and 
far shore stations, with another 17% limited to the near shore and 27% noted only 
at far shore stations (Table 4). The prominent diatoms over the shelf were Cy- 
matosira belgica, Paralia sulcata, Plagiogramma staurophorum, Rhizosolenia 
alata, R. alata indica, R. stolterfothii, and Thalassionema nitzschioides (Table 5). 
Rhizosolenia alata averaged 713 and 404 cells/1 at near and far shore stations, 
with R. alata indica averaging 1029 and 404 cells/1 at near and far shore sites. In 
1978, Rhizosolenia alata indica reached 18,472 cells/1 directly off Savannah, with 
other "pockets" of high concentration scattered over the shelf. These two 
species, with Rhizosolenia alata gracillima, R. calcar avis, R. hebetata semis- 
pina, R. setigera, R. stolterfothii, and R. styliformis represented a common dia- 
tom and generic assemblage throughout the shelf. 

The phytoflagellates were not found in high concentrations in these collections 
but were generally widely distributed. Only 14% of the dinophyceans (Pyrrho- 
phyceans) were limited to the near shore stations, with 40% of the species found 
at both near and far shore sites and 46% limited to the far shore stations. The 
haptophyceans, (Prymnesiophyceans) consisted mostly of coccolithophores, with 
46% of this class common across the shelf and 31% of the species limited to the 
far shore sites. The euglenophyceans, chlorophyceans, and chrysophyceans were 
found in low concentrations, but widely distributed over the shelf, with cyano- 
phyceans, and prasinophyceans more common near shore. The prominent near 
shore dinophyceans were Prorocentrum micans, P. aporum, Gyrodinium fusi- 
forme, Gonyaulax monilta, with Ceratium furca, Dinophysis caudata, and Ka- 
tyodinium rotundatum common over the entire shelf. Other characteristic near 
shore species include Emiliania huxleyi, Ochromonas sp., Dictyocha fibula, An- 
acystis marina, and Johannesbaptistia pellucida. A far shore dominant was Emi- 
liania huxleyi, with Cyclococcolithus leptoporus and Oscillatoria erythraea found 
over the shelf in significant numbers. There was no increase in cell concentrations 
at stations near the shelf break during either of these cruises. Total phytoplankton 
populations were generally low, but gave evidence of patchiness in cell concen- 
trations over the entire area during both cruises. 

Discussion 

The shelf phytoplankton possessed a diverse assemblage of 328 species from 
ten taxonomic categories. Diatoms, dinophyceans, haptophyceans, cyanophy- 
ceans, and the ultraplankton component represented the most abundant forms 
and were distributed over the entire shelf. Each of these categories had charac- 
teristic species in the near and far shore stations, with several species common 
to both (e.g. Rhizosolenia alata, R. alata indica, Emiliania huxleyi, etc.). The 
cyanophyceans were more concentrated in the near shore areas, with many 
species widely distributed over the shelf. It is also suggested that many of the 
ultraplankters (size <3 fjum) are cyanophyceans which may indicate added sig- 



VOLUME 95, NUMBER 1 



111 



Table 5. — Prominent phytoplankton associated with near and far shore stations during the 1973 and 
1978 collections. 



Near shore assemblage 



Far shore assemblage 



Diatoms 
Biddulphia alternans 
Cymatosira belgica 
Chaetoceros decipiens 
Fragilariopsis cylindrus 
Cylindrotheca closterium 
Guinardia flaccida 
Hemiaulus sinensis 
Melosira distans 
Paralia sulcata 
Plagiogramma staurophorum 
Rhizosolenia alata 
Rhizosolenia alata gracillima 
Rhizosolenia alata indica 
Rhizosolenia setigera 
Synedra fulgens 
Synedra undulata 

Tabellaria fenestrata asterionelloides 
Thalassionema nitzschioides 
Thalassiosira frauenfeldii 

Dinophyceans 

Gonyaulax monilta 
Gyrodinium fusiforme 
Prorocentrum micans 
Prorocentrum aporum 
Ceratium furca 
Dinophysis caudata 
Katodinium rotundatum 

Others 

Emiliania huxleyi 
Ochromonas sp. 
Dictyocha fibula 
Anacystis marina 
Johannesbaptistia pellucida 
Ultraplankton component 



Diatoms 

Chaetoceros decipiens 
Climacodium frauenfeldium 
Guinardia flaccida 
Rhizosolenia alata 
Rhizosolenia alata indica 
Rhizosolenia stolterfothii 
Thalassionema nitzschioides 
Melosira granulata angustissima 

Dinophyceans 

Ceratium furca 
Dinophysis caudata 
Katyodinium rotundatum 

Others 

Emiliania huxleyi 
Cyclococcolithus leptoporus 
Oscillatoria erythraea 
Cryptomonas sp. 
Ultraplankton component 



nificance in the presence of this group to the region. These cells also appear 
widely distributed over the eastern shelf of the United States (Marshall and Cohn, 
1981). Other representatives within this size category and the 3-5 fxm group 
appear to include chlorophyceans, among other types. Many of these phyto- 
plankters may represent the so called "lesser" systematic categories (rather than 
the dinophyceans, diatoms, and haptophyceans). Of note are the cryptophyceans, 
whose concentrations were not high in these collections in comparison to the 
more prominant groups, but maintained a broad distribution pattern over the 
shelf. However, the total significance of this and other groups is not completely 
represented in this study. Although the present collections include an extensive 
shelf coverage from two different cruises, there was a temporal limitation of the 



112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

data (both cruises were in the fall and each covered a limited time span for the 
collections). Some smaller scale variations in composition and concentrations 
also occurred along the transects between adjacent stations. This patchiness in- 
volved high concentrations of one or more species at scattered locations during 
both cruises and was more typically found among species of diatoms and dino- 
flagellates. There was also a pattern of high cell counts for the ultraplankton 
component at stations nearest the estuaries with decreasing concentrations sea- 
ward. However, the contributions of this unidentified ultraplankton component 
to the biomass of the standing crop appeared to be consistently small in com- 
parison to the amount attributed by diatoms and dinoflagellates. Unknown is the 
long term impact of these cells to the productivity and annual biomass totals for 
this area. 

Pollen and moth scales were also common in the samples and both were noted 
at stations up to 100 km off the coast. Rhaphoneis surinella was found only 
attached to grains of sediment that were collected in the water column. Richelia 
intracellularis was not found in a free state but as an endosymbiont in Rhizoso- 
lenia styliformis and R. hebetata semispina. Epibiotic fungi (chytridiaceans) were 
also noted on Rhizosolenia alata indica. The reported dinoflagellate cysts con- 
sisted mainly of Protoperidinium pentagonum. 

In summary, the results from collections at 91 shelf stations indicated higher 
concentrations for each taxonomic group, except the cryptophyceae, and hap- 
tophyceans, near shore during both collection periods. This pattern was also 
noted for the unidentified ultraplankton component. Exceptions to this pattern 
were common, and occurred as patchiness of species dominance and major shifts 
in individual concentration values along transects. The shelf populations were 
dominated by numerous large sized centric diatoms (e.g. Rhizosolenia alata and 
R. alata indica), with clusters and single filaments of the cyanophycean Oscil- 
latoria erythraea also common. However, numerous small sized cyanophyceans, 
diatoms and phytoflagellates were also characteristic of the shelf region. Another 
prominent group was the cyanophyceans, these were widespread over the shelf, 
often illustrating patchiness and high concentrations of cells near estuaries. Mar- 
shall (1981) associates many of the coastal marshes bordering the inner shelf area 
as a possible origin for many of these cyanophyte species. 

Not observed in this study was the pattern of increased cell concentrations 
from the middle to outer shelf areas, as noted by Bishop et al. (1980). However, 
a similar increase in cell concentrations near the shelf break was noted by Mar- 
shall (in press) off the Virginia coast. In contrast, the results at shelf stations 
from these cruises were generally uniform in their composition and concentra- 
tions, with occasional patchiness found broadly distributed, but more centered 
near and slightly beyond coastal estuaries. However, it is noted that these cruises 
were both limited to fall collections and covered a brief collection period. 

Acknowledgments 

Appreciation is given to the National Marine Fisheries Service (MARMAP 
Program) in cooperation with the South Carolina Marine Resources Institute of 
Charleston, South Carolina, which provided the samples and supplementary station 
data, with special thanks to Drs. Victor Burrell and Charles Baran. Further ap- 



VOLUME 95, NUMBER 1 113 

preciation is given to graduate assistants Charles K. Rutledge, Stephen Cibik, 
Brad Fawley, and Laurie Kalenak for their contributions in this study, and to Dr. 
Paul Kirk for identification of the chytridiacean fungi. 

Literature Cited 

Bishop, S., J. Yoder, and G. Paffenhofer. 1980. Phytoplankton and nutrient variability along a cross- 
shelf transect of Savannah, Georgia, U.S.A. — Estuarine and Coastal Marine Science 

2:359-368. 
Drouet, F. 1968. Revision of the classification of the Oscillatoriacea. Monogr. 15. — The Academy of 

Natural Sciences of Philadelphia, Fulton Press, Lancaster, Pa., 370 pp. 
, and W. A. Daily. 1956. Revision of the coccoid Myxophyceae. — Butler University Botanical 

Studies. Vol. XIL Hafner Press, New York, N.Y., 222 pp. 
Dunstan, W., and J. Hosford. 1977. The distribution of planktonic blue green algae related to the 

hydrography of the Georgia Bight. — Bulletin of Marine Science 27:824-829. 
Hendey, N. I. 1974. A revised check-list of British marine diatoms. — Journal of the Marine Biological 

Association United Kingdom 54:277-300. 
Hulburt, E. M. 1967. Some notes on the phytoplankton off the southeastern coast of the United 

States. — Bulletin of Marine Science 17:330-337. 
, and R. S. MacKenzie. 1971. Distribution of phytoplankton species at the western margin of 

the North Atlantic Ocean. — Bulletin of Marine Science 21:603-612. 
Johnson, P. W., and J. McN. Sieburth. 1979. Chroococcoid cyanobacteria in the sea: A ubiquitous 

and diverse phototrophic biomass. — Limnology and Oceanography 24:928-935. 
Malone, T. C. 1971. The relative importance of nannoplankton and netplankton as primary producers 

in tropical oceanic and neritic phytoplankton communities. — Limnology and Oceanography 

16:633-639. 
Marshall, H. G. 1969. Phytoplankton distribution off the North Carolina coast. — American Midland 

Naturalist 81:241-257. 
. 1971. Composition of phytoplankton off the southeastern coast of the United States. — Bul- 
letin of Marine Science 21(4):806-825. 
. 1981. Occurrence of bluegreen algae (Cyanophyta) in the phytoplankton off the southeastern 

coast of the United States. — Journal of Plankton Research 3:163-166. 
. In Press. Phytoplankton assemblages within the Chesapeake Bay plume and adjacent waters 

of the continental shelf. — Estuarine, Coastal and Shelf Science. 
, and M. S. Cohn. 1981. Phytoplankton community structure in northeastern coastal waters 

of the United States. I. October 1978.— NO AA Technical Memorandum NMFS-F/NEC-8, 

57 p. 
McCarthy, J. J., W. R. Taylor, and J. Loftus. 1974. Significance of nanoplankton in the Chesapeake 

Bay estuary and problems associated with the measurement of nanoplankton productivity. — 

Marine Biology 24:7-16. 
Parke, M., and P. S. Dixon. 1976. Checklist of British marine algae. Third Revision. — Journal of the 

Marine Biological Association United Kingdom 56:527-594. 
Waterbury, J., S. Watson, R. Guillard, and L. Brand. 1979. Widespread occurrence of a unicellular, 

marine, planktonic, cyanobacterium. — Nature 277:293-294. 



Department of Biological Sciences, Old Dominion University, Norfolk, Virgin- 
ia 23508. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 114-115 

NOMENCLATURAL STATUS OF THE FORAMINIFERAL 
GENUS CUBANELLA SAIDOVA, 1981 

Richard W. Huddleston and Drew Haman 



The Carpenteriinae, as defined by Saidova (1981) contains Carpenteria Gray, 
1858, Haerella Belford, 1960, Carpenterella Bermudez, 1949, and Cubanella Sai- 
dova, 1981. Carpenterella Bermudez is a junior homonym of Carpenterella Col- 
lenette in Collenette and Hale-Carpenter 1933, and was renamed Bermudezella 
by Thalmann (1951). Loeblich and Tappan (1964) considered Bermudezella and 
Haerella junior synonyms of Carpenteria. 

Saidova (1981) established the genus Cubanella and selected Carpenteria cu- 
bana (Cushman and Bermudez, 1936) as the type-species. Originally this species 
was described as the type and only member of Neocarpenteria by Cushman and 
Bermudez (1936), and they differentiated Neocarpenteria from Carpenteria based 
on its large, semicircular aperture with distinct lip. Loeblich and Tappan (1964) 
noted that the aperture of the type-specimen of Neocarpenteria cubana was an 
irregular artifact of the broken final chamber. Loeblich and Tappan (1964:C708) 
further added, "As the type species of Carpenteria does not grow upward into 
a cylindrical form and the apertural distinction is nonexistent, the name Neocar- 
penteria is a junior synonym [of Carpenteria}.'' Cubanella Saidova is a junior 
objective synonym of Neocarpenteria (International Code of Zoological Nomen- 
clature, ICZN, Article 61b) with Neocarpenteria a junior subjective synonym of 
Carpenteria (Loeblich and Tappan, 1964). Cubanella Saidova also is a junior 
homonym of Cubanella Pelegrin Franganillo (1926), an arachnid. 

Cubanella Saidova 1981, cannot be given a new name (ICZN, Article 60), but 
is to be replaced by Carpenteria, the oldest available synonym (ICZN, Article 
60a); or if the synonymy of Neocarpenteria and Carpenteria is rejected, by Neo- 
carpenteria. 

Acknowledgments 

We thank K. L. Finger, Chevron Oil Field Research Company for reading this 
manuscript and helpful comments, and Chevron Oil Field Research Company for 
permission to publish. 

Literature Cited 

Belford, D. J. 1960. Upper Cretaceous Foraminifera from the Toolonga calcitutite and Gingin Chalk, 
western Australia. — Australian Bureau of Mineral Resources, Geology and Geophysics, Bul- 
letin 57, 198 pp. 

Bermudez, P. J. 1949. Tertiary smaller Foraminifera of the Dominican Republic. — Cushman Labo- 
ratory of Foraminiferal Research, Contributions, Special Publication 25, 322 pp. 

Collenette, C. L., and G. D. Hale-Carpenter. 1933. New species and description of larva of Lyman- 
triidae from Uganda. — Entomology Monthly Magazine 69:258-270. 

Cushman, J. A., and P. J. Bermudez. 1936. New genera and species of Foraminifera from the Eocene 
of Cuba. — Cushman Laboratory of Foraminiferal Research, Contributions 12(2):27-63. 

Gray , J . E . 1858. On Carpenteria and Dujardina , two genera of a new form of Protozoa with attached 



VOLUME 95, NUMBER 1 115 

multilocular shells filled with sponge, apparently intermediate between Rhizopoda and Porif- 

era. — Zoological Society of London, Proceedings 26:266-271. 
Loeblich, A. R. Jr., and H. Tappan. 1964. Treatise on Invertebrate Paleontology, Part C, Protista 2, 

Chiefly Thecamoebians and Foraminiferida (ed. R. C. Moore). — University of Kansas Press, 

Lawrence, 900 pp. 
Pelegrin Franganillo, R. P. 1926. Nuevos o poco conocidos de la isla de Cuba. — Boletin Sociedad 

Entomologica Espana 9(3^):42-68. 
Saidova, Kh. M. 1981. O sovremennom sosteyanii sistemy Nadvidovykh Taksonov Kajnozojskihh 

bentosnykh foraminifer. — Akademiia Nauk SSSR Isstytut Okeanologiia, 73 pp. 
Thalmann, H. E. 1951. Mitteilungen uber Foraminiferen. — Eclogae Geologicae Helvetiae, pt. 

9(43):22 1-225. 

Chevron Oil Field Research Company, P.O. Box 446, La Habra, California 
90631. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 116-160 

SOME NEW AND OLD SPECIES OF THE PRIMNOID GENUS 

CALLOGORGIA GRAY, WITH A REVALIDATION OF 

THE RELATED GENUS FANELLIA GRAY 

(COELENTERATA: ANTHOZOA) 

Frederick M. Bayer 

Abstract. — One new and three previously known species of Callogorgia Gray 
are described and illustrated by scanning electron microscopy. The diagnostic 
characters of the genus are discussed and the species divided between two genera, 
for the second of which Gray's generic name Fanellia is restored to validity. 
Keys to the species of both (exclusive of the Atlantic and Mediterranean) are 
provided. 



Introduction 

In the course of reviewing the genera of Gorgonacea, it was found that the 
species traditionally assigned to the genus Callogorgia (Family Primnoidae) fall 
into two separate groups on the basis of external skeletal ornamentation, loosely 
correlated with colonial growth form. Typical Callogorgia, based upon Gorgonia 
verticillata Pallas, has pinnate growth form and sclerites varying from externally 
smooth to granulated or wrinkled, in a few species with the abaxial scales sculp- 
tured by prominent radial crests derived from the marginal ridges of the inner 
rim. A second group of species has dichotomous branching (or pinnate branching 
so lax that it may appear dichotomous, which is here called 'quasi-dichotomous') 
and sclerites with crowded, radially oriented tubercles extensively coalesced as 
ridges. This group of species is now restored to generic status under the name 
Fanellia, proposed for Primnoa compressa Verrill by J. E. Gray (1870:46). This 
genus as now constituted occurs from the Bering Sea and Gulf of Alaska south 
and west to Hawaii and New Caledonia, across the Pacific to Japan and Indonesia. 
It is not known from the eastern Pacific, from either side of the Atlantic or from 
the Mediterranean, and has not been reported from the Indian Ocean. Species of 
Callogorgia occur in the Mediterranean Sea, on both sides of the Atlantic Ocean, 
from the west coast of the Americas westward to Indonesia, and throughout most 
of the Indian Ocean. The few Callogorgia species reported from the Southern 
Ocean (C. antarctica, C. kuekenthali, C. nodosa and possibly C. ventilabrum), 
all with dichotomous growth form, are referable to still another genus, Ascolepis 
Thomson and Rennet, having sculpture completely different from that oi Fanellia. 

Methods 

The Scanning Electron Micrographs (SEM) of intact polyps were made with 
a Cambridge Stereoscan model S4-10, and the sclerites were examined and pho- 
tographed with a Coates and Welter model 106 field-emission instrument. 

For examination by SEM, sclerites were dissociated and cleaned of surrounding 
tissues by commercial 5.25% sodium hypochlorite bleach, followed by weak (3%) 



VOLUME 95, NUMBER 1 117 

hydrogen peroxide and thorough washing in water. After a rinse in 100% ethanol, 
the bulk sample was dried and subjected to an oxygen plasma for about 4 hours, 
before mounting with PVA emulsion adhesive on 10-mm coverglasses that can 
be affixed to standard Cambridge SEM stubs, or held in a custom-made spring- 
clip holder compatible with the Cambridge stage. Specimens were precoated with 
carbon and sputter-coated with gold/palladium. 

Samples for examination as whole mounts were selected for undamaged con- 
dition using a conventional stereomicroscope. After removal of superficial tissue 
by a very brief immersion in sodium hypochlorite solution followed by hydrogen 
peroxide and thorough rinsing in 70% ethanol, air-dried samples of suitable length 
(no more than 2 cm) were mounted vertically in the center of standard Cambridge 
stubs by means of PVA emulsion adhesive. After appropriate coating with carbon 
and gold/palladium, these preparations were placed in the Cambridge series 200 
microanalytical stage using an adapter specially made to accommodate the stan- 
dard Cambridge stub. The smaller Cambridge series 100 stage will not accept 
samples of this size. The stage was tilted to a full 90° for examination of such 
wholemounts, and stereo pairs prepared by rotation (8°) rather than by tilting, in 
order to preserve the original orientation more closely. 



Taxonomic Characters 

The principal characters available for the classification and identification of 
primnoids are: 

1 Colonial form and manner of branching. 

2 Size and arrangement of polyps. 

3 Number, arrangement, form and ornamentation of sclerites on the polyps and, 
to a lesser extent, on the coenenchyme. 

Colonial form. — Although "pinnate" and ''dichotomous" branching have been 
employed as key characters, the two forms intergrade, and where they do so they 
are ambiguous. In true pinnate growth form, only branches of the first order are 
produced; the first-order branchlets are terminal "pinnae" that rarely subdivide, 
except to produce a lateral plume that is structurally identical to the main one. 
In the dichotomous growth form, branching proceeds to several orders, each 
branch repeatedly forking into two. If the first-order branchlets in a pinnate col- 
ony are widely spaced and are not consistently "terminal" but produce second- 
and third-order branchlets, the branching pattern simulates dichotomy. This con- 
dition is illustrated in Figs. 15 and 18, and is here called "quasi-dichotomous." 

Size and arrangement of polyps. — The distribution of polyps in the colony 
appears to be a consistent character at generic and specific levels. The verticillate 
arrangement is common in the family and, in genera such as Primnoella, Calyp- 
trophora, and Narella, is never departed from. In all known species of Callo- 
gorgia save one, the polyps are in whorls, and the one exception differs in so 
many other regards that it probably does not belong in Callogorgia at all. In 
Plumarella, the polyps may be arranged either in tight spirals all around the 
branchlets, or biserially (but not in opposite pairs, which really are "whorls" of 
two). In Thouarella, various species are described with polyps paired, in whorls. 



118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

and in loose, irregular spirals even within one subgenus of this large and per- 
plexing genus. It is probable that when all the species have been reviewed in 
modern terms, a generic realignment in which the arrangement and spiculation 
of the polyps is more consistent will emerge. 

The direction in which the polyps face is quite consistent. Except in the genus 
Primnoa, polyps face upward in all genera in which they are not in whorls; the 
polyps of Primnoa face downward, apart from occasional strays. In Primnoella, 
Callogorgia, and Fanellia they always face upward, again except for strays 
(which may be induced by the presence of epizoites, such as echinoderms). In 
Calyptrophora they always face upward, in Paracalyptrophora, Arthrogorgia, 
and Narella always downward, and in Candidella, Parastenella, Pterostenella, 
and the verticillate species of Thouarella, directly outward or slightly upward. 

Sclerites. — Although irregular tuberculate rods and spindles occur in the Prim- 
noidae and may be the original sclerite form in this family, they are of only 
sporadic occurrence and the predominant forms are flat scales or plates, some- 
times very thick. On the body of the polyps, these scales are basically arranged 
in 8 longitudinal rows as well as in transverse circlets, of which the uppermost 
constitutes in most genera an operculum of 8 triangular scales distinctly differ- 
entiated from the rest of the body scales below them. The sizes and shapes of 
the sclerites are influenced by the extent of inward curvature achieved by the 
polyps during contraction. A special terminology of sclerites in various positions 
on the polyps has developed over the years, differing even from genus to genus, 
depending upon the degree of modification that the skeletal armature has under- 
gone. Because these terms are generally used in keys and taxonomic descriptions, 
those applicable to forms with strongly in- turned polyps, such as Callogorgia, 
are briefly explained. 

Because of the inward curvature of the polyps, those rows on the side of the 
body turned toward the axis are called adaxial, those on the side facing outward 
abaxial, and the intermediate ones outer lateral and inner lateral. Owing to the 
bilateral symmetry of the octocoral polyp, the scale rows are paired, so the 
members of the pairs lie on opposite sides of the body, with the "sagittal" plane 
passing between the members of the abaxial and adaxial scale rows. The scales 
of the transverse circlet immediately below and surrounding the operculars are 
termed marginal, but no special terminology applies to those below. To save 
space in print, the scale rows are designated by abbreviations (Abax = abaxial; 
OL = outer lateral; IL = inner lateral; Adax = adaxial) and the individual scales 
are numbered from the operculum downward, as shown in Fig. 1. 

In Callogorgia the outer surface of the sclerites may be smooth, wrinkled, or 
ornamented by sharp granules often aligned in radial or reticulating rows (see 
Figs. 9, 10). The inner surface of the body scales always has a more or less 
smooth distal margin marked by radial ridges that may be low, or high and crest- 
like, throwing the edge into dentations (see Figs. 7,8), and sometimes extending 
outward as tall crests on the distal outer surface of the scales (see Figs. 2-5). 

In Fanellia the outer surface is covered by crowded, fluted or serrated tubercles 
commonly coalescing to form radial ridges (see Figs. 13, 14; 16, 17; 19-22; 25, 
26; 28, 29). In both genera, the central inner surface of all scales is covered by 
crowded, complex tubercles that serve to anchor the sclerites in the mesogloea. 



VOLUME 95, NUMBER 1 



119 




Abax-O 



Abax-1 



Abax-2 



Abax-3 



Abax-4 



Abax-5 



Abax-6 




Abax-7 
Abax- 



Abax-6 



Abax-7 



Fig. 1. Arrangement and terminology of body sclerites, diagrammatic: A, Fanellia with both 

outer-lateral and inner-lateral rows reduced in number; B, CaUogorgia with outer-lateral rows well 

developed, inner laterals reduced. Abax = abaxial; OL = outer lateral; IL = inner lateral; Adax - 

adaxial (not visible in side view). O = opercular scale; 1 = marginal; 2 — = remaining body scales 

numbered proximad. 



CaUogorgia Gray, 1858 

Gorgonia. — Pallas, 1766:160 (part). — Linnaeus, 1767:1289 (part). — Ellis and So- 

lander, 1786:67 (part). 
Muricea. — Dana, 1846:675 (part). 
Prymnoa. — Ehrenberg, 1834:357 (part). 
Primnoa. — Milne Edwards and Haime, 1857:139 (part). — von Koch, 1878:457; 

1887:85. 
CaUogorgia Gray, 1857 [1858]: 286 (type-species, Gorgonia verticiUata Pallas, 

1766, by monotypy). — Bayer, 1961 [1962]:296 (part). — Carpine and Grasshoff, 

1975:102. 
CaUigorgia Gray, 1870:35 (unjustified emendation). — Studer, 1878:51. 
XiphoceUa Gray, 1870:56 (type-species, Gorgonia verticiUata: sensu Esper, 

1797:156, by monotypy). 
? CalUceUa Gray, 1870:37 (type-species, CaUiceUa elegans Gray, 1870, by mono- 
typy). 
Caligorgia Wright and Studer, 1889:75 (unjustified emendation). — Versluys, 

1906:55.— Kukenthal and Gorzawsky, 1908:19.— Kinoshita, 1908:34.— Nutting, 



120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

1908:574.— Kiikenthal, 1912:320(7); 1915:146; 1919:362; 1924:267.— Deich- 
mann, 1936:158. 

Description.— Colonics of moderate (30 cm) or large (over 1 m) size, branched 
in one plane, pinnate, with branchlets usually alternating, rarely opposite (1 
species), new branchlets originating only apically, not between pre-existing 
branchlets, sometimes openly pinnate or quasi-dichotomous, rarely dichotomous 
(1 species?). Polyps in whorls of 3-6 or more, directed toward apex of branches, 
in some species beginning distally as pairs and increasing in numbers proximad, 
bent inward toward the axis, the adaxial side more or less closely appressed to 
the axial coenenchyme. Sclerites of the polyps in the form of scales curved to 
conform with the contours of the body, placed in longitudinal rows of which only 
the two abaxials are complete, the lateral and adaxial rows with reduced numbers 
of scales, in some species absent adaxially except for the operculars. Opercular 
scales triangular, shape abruptly different from those of marginals, which do not 
fold inward over the operculars. Tentacles without sclerites in most species, a 
few with minute rods. Coenenchymal sclerites in the form of plates, from broadly 
polygonal or ovate to strongly elongate. Axis heavily calcified, irregularly grooved 
longitudinally, yellowish, brownish or greyish, often with metallic or iridescent 
sheen. 

Discussion. — This genus, established by J. E. Gray (1858) for the eastern At- 
lantic and Mediterranean Gorgonia verticillata Pallas, 1766, is widespread and, 
apart from some trans-Atlantic confusion of the type-species (see Carpine 
1963:30; Carpine and Grasshoff 1975:102), has been comparatively free of prob- 
lems. However, a number of species have been ascribed to it erroneously, and 
examination of a wide array of species reveals that they fall into rather well- 
defined groups on the basis of spicular ornamentation. 

Kiikenthal (1924:267) characterized the genus as follows (translation mine): 
Colonies abundantly branched and mostly in one plane, sometimes pinnate with 
alternate or opposite terminal branchlets, sometimes more dichotomous. The 
polyps occur in whorls, only on the larger stems scattered as well. Their adaxial 
wall is more or less naked. The scales of the polyps are set in longitudinal rows, 
of which the adaxials are always reduced; they are mostly strong and conspicu- 
ous, beset on the inner surface with numerous closely placed small warts, on the 
outer surface mostly with thornlike projecting ridges or other sculpture. The 
opercular scales are well developed abaxially and distinctly pointed, becoming 
smaller adaxially. The marginal scales do not overreach the operculars and are 
not movable inward. The coenenchymal scales are unlike those of the polyps, not 
imbricated but set side by side and mostly quite thick, commonly elongated. 

It is clear from published descriptions that Caligorgia antarctica Kiikenthal 
(1912:321), Caligorgia nodosa Molander (1929:60), and Caligorgia ventilabrum 
Studer sensu Gravier (1914:85) actually belong to Ascolepis Thomson and Rennet 
(1931:20), which I regard as a valid genus (Bayer 1981:936). The growth form of 
all of these species is quite unlike the pinnate colonies of most species of Cal- 
logorgia . 

Versluys (1906:83), in his exemplary monograph on the Primnoidae of the Si- 
boga Expedition, recognized that the species of Callogorgia reviewed by him fall 
into groups. Unfortunately, he grouped the species first on the basis of branching. 



VOLUME 95, NUMBER 1 121 

which, as can be seen in other genera of Primnoidae, is an unreliable character, 
partly because it is difficult to distinguish pinnate branching, if widely spaced, 
from dichotomous, and partly because both "pinnate" and "dichotomous" 
branching can occur in a single genus. 

However, among the dichotomously branched species, Versluys (1906:84) clear- 
ly recognized the distinctively different kind of sculpturing characteristic of the 
sclerites of C. compressa and C. tuberculata. The subsequent description of two 
other species (C granulosa, C. aspera) from Japan by Kinoshita (1907, 1908a) and 
one from the Marshall Islands by me (Bayer 1949:207), together with the discov- 
ery among the Albatross Philippine octocorals of yet another species, all with the 
same kind of external ornamentation, makes it clear that this character, not the 
colonial growth form, provides a reliable basis for grouping of species. Further, 
neither in the literature nor among the specimens examined have I found any 
examples of sculpture intermediate between the tuberculata-compressa type and 
the verticillata-flabellum type. Consequently, these two groups are discontinuous 
and merit full generic rank. A new generic name is not required for the tuber- 
culata-compressa complex as Gray (1870:46) already has proposed for C. com- 
pressa Verrill a nominal genus Fanellia, which is here restored to good standing. 

Although the remaining species of Callogorgia have generally similar sculpture, 
two sub-types can be recognized among them: (1) those with the scales more or 
less smooth externally, sculptured at most by smooth granules, usually scattered 
but sometimes merging to form wrinkles that may anastomose into reticular or- 
namentation (sertosa Wright and Studer, verticillata Pallas, grimaldii Studer, 
pennacea Versluys, joubini Versluys, similis Versluys, minuta Versluys, affinis 
Versluys, kinoshitae Kiikenthal, gilberti Nutting, ventilabrum Studer, gracilis 
Milne Edwards and Haime); and (2) those with the body scales sculptured ex- 
ternally by prominent radial crests that are outward extensions of the radial ridges 
developed around the inner distal margin of the body scales in most species, if 
not all {flabellum Ehrenberg, weltneri Versluys, robusta Versluys, cristata Au- 
rivillius, verticillata sensu Deichmann). Even though these two kinds of sculpture 
are usually quite distinct, there is some overlap. In some (but not all) specimens 
of C. gracilis (Milne Edwards and Haime), the inner marginal ridges of the abaxial 
marginal scales are reflected outward onto the outer surface as low ridges con- 
verging toward the nucleus of the scale. These ridges have a quite different aspect 
from those of C. flabellum and related forms {robusta, weltneri, cristata), but 
they seem to be identical structurally. Therefore, there seems to be no infallible 
basis for subdividing those species into two genera, or even subgenera. 

The disposition of Ascolepis Thomson and Rennet, 1931, is equivocal. Both of 
the original species, A. splendens and A. spinosa, have sclerites of unusual 
form — not so distinctly cuplike as originally described and illustrated from ob- 
servation with the light microscope, but certainly with a concave outer part more 
or less clearly set off from a tuberculated base. The same is true of Caligorgia 
nodosa Molander and the specimens reported as C ventilabrum by Gravier, 
appears to be true also of C. antarctica Kiikenthal, and could be true of C. 
kiikenthali Molander. If C. antarctica represents an intermediate between Cal- 
logorgia and Ascolepis, thus making the latter a junior synonym, then the genus 
Callogorgia will include species that do not remotely resemble the majority of 
species of Callogorgia. Because of the difficulty in defining Callogorgia that 



122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

would result from merging Ascolepis with it, the preferable course is to maintain 
both as distinct. 

Other problems with the definition of Callogorgia remain to be solved, notably 
the case of those Antarctic species of C. antarctica growth form in which the 
regular alignment of abaxial and lateral body scales is completely disrupted. 
Whether these merit separate recognition, or must be accommodated in a more 
broadly defined genus Callogorgia is a question that must await further study. 
For the time being, I propose to recognize Callogorgia, Fanellia, and Ascolepis 
as separate genera. 

Key to Indo-Pacific Species of Callogorgia^ 

1(20). Branching typically pinnate. 

2(3). Branches strictly opposite C. formosa Kiikenthal, 1907 

3(2). Branches alternate. 

4(7). Outer-lateral scale-rows well developed. 

5(6). Polyps 1.3-1.5 mm tall, whorls separated by conspicuous intervals, 

5 in 1 cm C. sertosa Wright and Studer, 1889 

6(5). Polyps larger, up to 2 mm tall, whorls closely placed, 5 in 1 cm . . . 

C. kinoshitae Kiikenthal, 1913 

7(4). Scales in outer lateral rows sharply reduced in number. 
8(19). Operculum tall and prominent. 
9(16). Apex of opercular scales with a single point. 
10(13). 10 scales in each abaxial row. 

11(12). Apex of opercular scales prolonged into a rodlike point 

C ramosa Kiikenthal & Gorzawsky, 1908 

12(11). Apex of opercular scales not prolonged into a rodlike point 

C. flabellum (Ehrenberg, 1834) 

13(10). 6-7 scales in each abaxial row. 

14(15). Polyps 2 mm tall C. robusta Versluys, 1906 

15(14). Polyps 1 mm tall C. joubinii Versluys, 1906 

16(9). Abaxial opercular scales with 2-A apical points. 
17(18). 5-6 whorls of polyps in 1 cm; 8-10 scales in abaxial rows; no inner 

lateral scales C. cristata Aurivillius, 1931 

18(17). 4-5 whorls of polyps in 1 cm; 7 scales in abaxial rows; 1 inner-lateral 

scale C weltneri Versluys, 1906 

19(8). Operculum low C. pennacea Versluys, 1906 

20(1). Branching dichotomous or quasi-dichotomous. 
21(26). Outer-lateral scale rows well developed. 

22(23). 8 scales in each abaxial row C. ventilabrum Studer, 1878 

23(24). 9 scales in each abaxial row C. laevis Thomson and Mackinnon, 191 P 

24(25). 10 scales in each abaxial row C. versluysi Thomson, 1905 

25(22). 12-13 scales in each abaxial row C. elegans (Gray, 1870) 

26(21). Scales in outer lateral rows reduced in number. 

27(28). 3 scales in each outer-lateral row C. indie a Versluys, 1906 



1 Modified from Kukenthal (1919). 

^ Examination of the type-specimens in the Australian Museum since this manuscript went to 
press has enabled me to determine that this species does not belong to Callogorgia but to Primnoella. 



VOLUME 95, NUMBER 1 123 

28(27). 1 or 2 scales in each outer-lateral row. 

29(32). 5 scales in each abaxial row. 

30(31). 2-3 polyps per whorl, rarely 4; 8-9 whorls in 1 cm; body scales with 

low marginal ridges C. minuta Versluys, 1906 

31(30). 3^ polyps per whorl, sometimes 5; 9-1 1 whorls in 1 cm; body scales 

with distal margins strongly reflexed, exposing high crestlike ridges 

C. chariessa, n. sp. 

32(29). 7 scales in each abaxial row. 

33(34). Body scales with strong marginal ridges; 4-5 polyps in each whorl, 

9-10 whorls in 1 cm C. affinis Versluys, 1906 

34(33). Body scales with weak marginal ridges; 2, usually 3 polyps in each 

whorl, 7-8 whorls in 1 cm C. similis Versluys, 1906 



Callogorgia flabellum (Ehrenberg) 
Figs. 2, 3 

Prymnoa flabellum Ehrenberg, 1834:358 (locality not given). 

Caligorgia flabellum.— Versluys, 1906:69, figs. 75-78, pi. 5, fig. 13, pi. 6, fig. 14 

(Kei Islands, 5°28.4'S, 132°0.2'E, 204 m). 
not Caligorgia flabellum. — Nutting, 1912:60 (=C. aspera Kinoshita, 1908a = C. 

tuberculata Versluys, 1906). 

Material examined. — Moluccas, off Makian, west of Halmahera, 0°15'00"N, 
127°24'35"E, 545 m, sand and coral, Albatross sta. D5621, 28 November 1909: 
several large branches without base, and smaller detached branches, USNM 
57545 (Figs. 2, 3). 

Description. — See Versluys, 1906:69. 

Discussion. — As Versluys (1906:69) has given a full description of the Siboga 
material, description of the present material would be superfluous. However, the 
validity of Versluys's identification of the Siboga material rests solely on his own 
comparison with a badly preserved fragment of Ehrenberg' s type in the Berlin 
Museum, as Wright and Studer did not compare the Challenger specimen with 
Ehrenberg's type as Versluys thought (1906:71), but with Gray's type of Callicella 
elegans (Wright and Studer, 1889:79). 

The specimen taken by the Albatross off Makian, west of Halmahera in the 
Moluccas, agrees satisfactorily with Versluys's account of the Siboga specimens 
from the Kei Islands. The characters that are diagnostically important are: 8-10 
abaxial scales in each row, with strong radial ridges extending from the inner 
marginal ridges; 3 outer laterals, 2 inner laterals, 1 adaxial; operculars tall, tapered 
to a single blunt point; abaxial scales sculptured with strong radial crests that 
extend distally as marginal serrations, most prominent on the distal sclerites, 
decreasing in strength proximad until on the basalmost one or two they are re- 
duced to low ridges or merely conspicuous wrinkles. The cortical plates are 
irregularly elongate, commonly with a central outer projection and crowded, 
pointed tubercles that are more or less extensively joined to form irregularly 
meandering low ridges or wrinkles. 

The present material differs from Versluys's description to a small degree, 
which records 2 outer laterals and 1 inner lateral but, in fact, his drawing (Fig. 
85) suggests 3 outer laterals and 2 inner laterals, as is usually the case in the 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Callogorgia flabellum (Ehrenberg), USNM 57545: left and right lateral views of polyp. 
SEM stereo pairs, x40. 



specimen before me. As can be seen from the accompanying SEM photos (Fig. 
2), the sculpture of the body scales obscures the distinction between individual 
scales and renders counting and drawing by camera lucida uncertain. The shapes 
and positions of scales shown in Versluys's drawing make it difficult to interpret 
the scales labeled 'G' as outer laterals. Their location requires them to be inner 
laterals, hence 2 in number, so the adaxial part of the 3 distalmost abaxial scales 
can only be 3 outer laterals. 

Callogorgia flabellum is very similar to C. cristata in aspect, difficuli to dis- 
tinguish with the dissecting microscope. The most reliable recognition features 
are: (1) the presence of 3 OL, 2 IL, and 1 Adax; (2) the single points on the 
abaxial operculars; (3) the extension of the inner marginal ridges of the abaxial 



VOLUME 95, NUMBER 1 



125 




Fig. 3. Callogorgia flabellum (Ehrenberg), USNM 57545: opercular, adaxial marginal, abaxial, 
outer-lateral sclerites of polyp. Scale = 0. 1 mm. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Callogorgia cristata Aurivillius. Top, USNM 57544; bottom, USNM 44179, part of type 
in Zoological Museum, University of Uppsala: intact whorls of polyps. SEM stereo pairs, x27. 



scales onto the outer surface as strong radial crests; and (4) the persistence of 
external sculpture, though reduced, on the proximal abaxial scales. 



Callogorgia cristata Aurivillius 
Figs. 4, 5 

ICaligorgia weltneri Versluys, 1906:73, figs. 80-82, pi. 4, fig. 10 (southwest of 
Waigeu, between Jef Fam and Gagi, 0°29.2'S, 130°05.3'E, 469 m).— Kukenthal, 
1924:274. 

Caligorgia cristata Aurivillius, 1931:262, fig. 52, pi. 6, fig. 1 (Japan, Goto Is., 
west of Kyushu, 160 m). 



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127 




Fig. 5. Callogorgia cristata Aurivillius, USNM 57544: opercular, abaxial, outer-lateral, and mod- 
ified opercular scales of polyp. Scale = 0.1 mm. 



128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Material examined. — Japan, Goto Islands, off Pallas Rock, 32°15'N, 128°12'E, 
160 m, coral bank, 15 May 1914, coll. S. Bock: small branch of type, USNM 
44179 (Fig. 4, bottom). 

Moluccas: off Ternate, 0°49'30"N, 127°25'30"E, 240 m, coral; Albatross sta. 
D5617, 27 November 1909: large branch and detached smaller branches, USNM 
57544 (Fig. 4, top).— off Makian, 0°15'00°N, 127°24'35"E, 545 m, sand and coral; 
Albatross sta. D5621, 28 November 1909: one pinnate branch without base, 
USNM 60285. 

Description. — See Aurivillius, 1931:262. Compare Versluys, 1906:73. 

Discussion. — The specimens collected by the Albatross from Ternate and Mak- 
ian, both off the west coast of Halmahera in the Moluccas, agree well with Au- 
rivillius 's account of C. cristata, and with Versluys' s description of C. weltneri. 
The principal difference between C. cristata and C. weltneri cited by Aurivillius 
(1931:266) is the "somewhat greater number of polypal spicules, amounting to 
8-9(-10) as against only 7" in weltneri. The present specimens tend to have 
somewhat more scales in the abaxial rows than was usual in C. cristata, but this 
character is subject to variation in all species of Callogorgia. Direct comparison 
by SEM of USNM 57544 (Fig. 4a) with type-material of C. cristata (Fig. 4b) 
reveals no other significant difference, so they can safely be considered conspe- 
cific. Even though Versluys (1906:74) reports no variation from 7 abaxials in C. 
weltneri, deviations almost certainly occur. A difference of 1 in the prevalent 
number of abaxial sclerites is a weak justification for specific separation so it is 
very likely that cristata and weltneri represent a single species. However, as 
type-material of C. weltneri has not been examined in this context, C. cristata 
is maintained. The inner basal angle of the inner-lateral opercular scales (IL-0) 
is in many cases expanded as a rounded lobe that extends into the position that 
would be occupied by the inner-lateral marginal scale (IL-1) if such a sclerite 
were developed (Fig. 5). This condition was not noticed by Aurivillius, but its 
inconsistent development may have led to its oversight. 

Callogorgia chariessa, new species 
Figs. 6-8 

Material. — One pinnately divided branch; Tawitawi Group, Sulu Archipelago, 
5°18'10"N, 120°02'55"E (Tocanhi Point S.27°E, 2.1 miles), 49 fath. (90 m), 
coral sand and shell; Albatross sta. D5153, 19 February 1908. Holotype, USNM 
58398. 

Description. — Callogorgia with alternate pinnate branching in one plane, the 
nodes 5-10 mm apart and the undivided terminal branchlets about 2 cm long, 
producing the open type of pinnate branching that easily can be mistaken for 
dichotomous. Polyps in whorls of 3-5, commonly 4; in the terminal branchlets, 
9 whorls commonly occur in 1 cm, uncommonly as many as 10 or 11. Diameter 
of undivided terminal branchlets 0.5 mm exclusive of polyps; axis alone about 
0.3 mm in diameter, tapering to a hair-fine apex. Axis heavily calcified, longitu- 
dinally striated, creamy yellow with golden iridescence. 

Polyps with only the two abaxial rows of scales well developed, 4 or 5 scales 
in each; one large outer lateral on each side, although this is occasionally absent, 
its place being occupied by the distalmost abaxial which then is sufficiently broad- 



VOLUME 95, NUMBER 1 



129 




Fig. 6. Callogorgia chadessa, new species; holotype colony, USNM 58398. Scale in cm. 



ened to cover the space; distal margins of the body scales strongly reflexed out- 
ward exposing the marginal radial ridges, which are high and crestlike. Opercular 
scales roughly triangular, with serrated edges, rounded basal angles and 1^ ser- 
rated ridges on both outer and inner surfaces of the blunt apex; adaxial and inner 
lateral operculars conspicuously smaller than the abaxials and outer laterals. Ex- 
posed surface of scales with sparse granular sculpture, partly aligned in irregular 
radial rows, inner surface with compound tubercles. Cortical scales large, polyg- 
onal, closely interlocking by their marginal serrations. 

Comparisons. — This species is similar to C. minuta and C. similis (Versluys, 
1906:76, 78); the latter, in turn, is "im Habitus der C. ventilabrum ausserst ahn- 
lich" (Versluys 1906:74, 76). C. ventilabrum, which originally was collected in 
162 m north of New Zealand by the Gazelle Expedition, has 7 scales in the abaxial 
rows (as determined by Versluys 1906:75, who examined a part of the type col- 
ony) compared with 5 or fewer in C. chariessa. The inner distal edges of the 
body scales of C. ventilabrum have sharp radial ridges but they are comparatively 
low and not conspicuously exposed by the outward reflexion of scale margins as 
in C chariessa. 

Callogorgia chariessa resembles C. minuta even more closely, but the whorls 
tend to be more closely placed (as many as 10 or 11 per cm, though commonly 
9) than in that species (8 or 9 per cm), and to consist of a larger number of polyps 
(commonly 4, sometimes 3 or 5, compared with 2 or occasionally 3 but rarely 4 
in minuta). The margins of the body scales are strongly reflexed in C. chariessa, 
conspicuously exposing the marginal ridges to view. In this regard, C. affinis 
approaches C. chariessa more closely, but the polyps have 7 scales in the abaxial 
rows as opposed to 5 (or even 4) in C. chariessa, and the margins of the body 
scales are not so conspicuously reflexed. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. Callogorgia chariessa, new species; holotype, USNM 58398: abaxial and right lateral 
views of polyp. SEM stereo pairs, x67. 



When more material becomes available for an assessment of variation, it may 
prove that all of these basically similar forms belong to a single variable species, 
but for the present time it is preferable to call attention to the differences between 
them. 



Callogorgia formosa Kiikenthal 
Figs. 9-10 

Caligorgia formosa Kiikenthal, 1907:208 (Siidwestlich von Gross-Nikobar, 362 

m); 1919:366, figs. 155-159, pi. 30, fig. 1, pi. 40, fig. 47; 1924:269, fig. 153. 
Primnoella indica Kiikenthal, 1907:210. 

Material examined. — Hawaiian Islands: north Bank of Necker Island, 250 fath. 



VOLUME 95, NUMBER 1 



131 




Fig. 8. Callogorgia chariessa, new species; holotype, USNM 58398: opercular, adaxial marginal, 
abaxial and outer-lateral sclerites of polyp. Scale = 0.05 mm. 



(458 m), Townsend Cromwell 76-06-73, sta. 4, 15 October 1976; large colony 
broken into 4 pieces: USNM 60692.— Off Necker Island, 23°39'N, 164°3rW, 
80-162 fath. (146-296 m), Townsend Cromwell sta. 6, 16 October 1976; two badly 
damaged pinnate fragments: USNM 60291. 

Description. — A large, plumose colony, broken in 4 pieces measuring 29.5, 
18.5, 12.2 and 8.3 cm, corresponds in all essentials with the description and 
figures of C.formosa given by Kiikenthal (1919). Evidently broken off well above 
the holdfast, the intact colony must have approached 1 m in height. It is a rather 
stiff and brittle pinnate plume with strictly opposite, undivided branches up to 
14.5 cm long, the distal ones rather quickly decreasing in length toward the apex. 



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Fig. 9. Callogorgia formosa Kiikenthal, USNM 60692: left lateral and abaxial views of polyp. 
SEM stereo pairs, x38. 

The polyps occur on the branches in rather well- separated whorls of 3 or 4 dis- 
tally, increasing proximad to 5, usually 4 in 1 cm; they are in whorls also on the 
main axis near its apex, but this arrangement is soon lost and only widely scat- 
tered individuals persist on the lower parts of the trunk. The size, shape, and 



Fig. 10. Callogorgia formosa Kiikenthal, USNM 60692: sclerites of coenenchyme, three outer, 
one inner view; opercular, adaxial marginal, abaxial and outer-lateral sclerites of polyp. Scale = 
0.1 mm. 



VOLUME 95, NUMBER 1 



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134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

armature of the polyps is essentially as described by Kiikenthal (1919). He makes 
the point (1919:367) that only the distalmost abaxial scales are wider than high, 
the rest being rounded or trapezoidal in outline. However, judging by his drawing 
(op. cit.: fig. 155), it appears that the proximal abaxial scales may have suffered 
some breakage, as is the case in the present specimens. When intact, the proximal 
abaxial scales are broad and tapering, extending well around the sides of the 
polyp. The outer-lateral rows are unusually well developed for the genus, com- 
posed of 6 or more scales. Except for the distal scales, it is difficult to confirm 
Kiikenthal 's counts for the inner-lateral and adaxial rows, as the proximal scales 
imperceptibly merge with the coenenchymal sclerites, which in full development 
are long, irregular spindles sometimes with blunt lobes. The body scales have 
only fine granulation externally, but no trace of radial ridges or crests; internally, 
they are covered with crowded complex tubercles of usual form, and the distal 
margins are sharply and conspicuously toothed. The tentacles are devoid of scler- 
ites. 

Relationships . — Kiikenthal (1919:369) considered this species so unusual that 
it might merit a new genus distinct from Callogorgia. Although he considered 
that in form of the polyps C. formosa conformed with other species of Callo- 
gorgia, he had actually mistaken detached branches for a new species of Prim- 
noella (Kukenthal, 1907:210, 1919:368). This demonstrates the difficulty of even 
generic identification because of the ambiguity of many taxonomic characters in 
the Octocorallia. 

Distribution. — In view of its type-locality in the Indian Ocean off Great Nicobar 
Island, the discovery of C. formosa in Hawaiian waters is of no little interest. Its 
distribution therefore parallels that of Fane Ilia tuberculata, first found off the 
Sulu Islands and now reported from Hawaii, but it has not been reported even 
as a synonym from intermediate localities as has F. tuberculata. It is probable 
that this large distributional gap results only from inadequate sampling. The other 
species of Callogorgia known from Hawaii, C. gilberti Nutting, is more closely 
related to C. kinoshitae Kukenthal from California and C. verticillata (Pallas) sensu 
Deichmann from the Caribbean than to any species from the Indo-west Pacific. 



Fanellia J. E. Gray, 1870 

Fanellia J. E. Gray, 1870:46 (type-species, Primnoa compress a Verrill, 1865; by 

monotypy). 
Caligorgia. — Auct. (in part). 

Diagnosis. — Primnoid colonies branched in dichotomous or lax pinnate manner 
simulating dichotomous, rarely close pinnate, usually but not always in one plane. 
Polyps upturned, verticillate, in whorls of 2-12; sclerites of polyps arranged in 
longitudinal rows in which the distalmost circle of 8 triangular or lanceolate scales 
(operculars) is always present; the two abaxial rows of body scales always fully 
developed but the outer-lateral, inner-lateral and adaxial rows are more or less 
reduced in number of scales owing to inward curvature of the polyps in contrac- 
tion (i.e., the adaxial body wall is more or less naked); tentacles devoid of scler- 
ites. Coenenchymal sclerites in the form of thick, closely fitting, rounded or 



VOLUME 95, NUMBER 1 135 

polygonal plates presenting a cobblestone-like aspect. Sclerites sculptured exter- 
nally by closely set, angular tubercles that may fuse to form serrated ridges 
irregularly meandering or more or less distinctly radiating outward from the cen- 
ter, internally by very complex, crowded tubercles not regularly aligned. 

Comparisons . — This genus closely resembles Callogorgia but is distinguished 
by the external sculpturing of the sclerites, which does not intergrade with the 
type of ornamentation developed in Callogorgia. The growth form is predomi- 
nantly dichotomous, or so openly pinnate that it appears to be dichotomous (here 
called 'quasi-dichotomous'), but only rarely is it closely pinnate (F. granulosa). 
In most Callogorgia species it is regularly pinnate, in some becoming more lax, 
but in a few species described (e.g., C. antarctica) it is dichotomous as in Fanellia 
compressa. However, several of these dichotomous species seem to be referable 
to the genus Ascolepis Thomson and Rennet on the basis of sclerite form. If not, 
the branching oi Fanellia and Callogorgia will be equivalent, in that both pinnate 
and dichotomous conditions occur, together with the quasi-dichotomous inter- 
mediate. In Fanellia, branching is predominantly dichotomous, in Callogorgia 
predominantly pinnate. In Ascolepis, branching varies even more, ranging from 
unbranched {nodosa) to typical dichotomous (splendens, antarctica), lax pinnate 
or quasi-dichotomous (spinosa) and bottle-brush form (abies). 

Geographical distribution. — Fanellia appears to be confined to the Pacific 
Ocean, occurring from the Gulf of Alaska and Aleutian Islands to Japan, Indo- 
nesia, Marshall Islands, Hawaii, and as far south as New Caledonia. No species 
recognizable as belonging to Fanellia has been reported from the Indian Ocean 
or from the Southern Ocean. In contrast, Callogorgia occurs in all seas (unless 
all of those from the Southern Ocean prove to be species of Ascolepis), and 
Ascolepis occurs only in Antarctic and Subantarctic waters. 



Key to Species of Fanellia 

1(4). Terminal branches long (15-30 cm), branching dichotomous or quasi- 
dichotomous; polyps in whorls of 8-12. 

2(3). Polyps with 7-8 scales in each abaxial row, 2 in outer-lateral rows . . . 

F. fraseri (Hickson) 

3(2). Polyps with 10-11 scales in each abaxial row, 5-7 in outer-lateral rows 

F. compressa (Verrill) 

4(1). Terminal branches shorter (7 cm or less), branching clearly pinnate or 
quasi-dichotomous; polyps in whorls of 2-5. 

5(6). Branching distinctly pinnate F. granulosa (Kinoshita) 

6(5). Branching quasi-dichotomous. 

7(8). Colonies flabellate, compressed or in one plane; polyps with 5-8 scales 

in abaxial rows, 1 or 2 outer laterals, 1 inner lateral 

F. tuberculata (Versluys) 

8(7). Colonies corymbose, not in one plane; polyps with 5 (occasionally 6) 
scales in abaxial rows, one wide marginal scale occupying the combined 
inner and outer lateral positions on both sides; no separate inner 
lateral F. corymbosa, n. sp. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. IL Fanellia compressa (Verrill), USNM 60281: complete colony. Divisions of scale are cm. 



Fanellia compressa (Verrill, 1865) 
Figs. 11-14 



Prymnoa verticillaris. — Ehrenberg, 1834:357. 
non Gorgonia verticillata Pallas, 1766:177. 



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137 




Fig. 12. Fanellia compressa (Verrill), USNM 60281: detail of colony. Divisions of scale are mm. 



non Gorgonia verticillaris Linnaeus, 1767:1289. — Ellis and Solander, 1786:83. — 

Statius Muller, 1775:753.— Houttuyn, 1772:309, pi. 132, fig. 1. 
non Callogorgia verticillata. — Gray, 1858:286. 
Primnoa compressa Verrill, 1865:189 (Aleutian Islands). 



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Fig. 13. Fanellia compressa (Verrill), USNM 57543: top, whorl of polyps, x27; bottom, oblique 
view of polyp, x44. SEM stereo pairs. 



Fanellia compressa. — Gray, 1870:46 (new genus for Primnoa compressa Verrill). 
Caligorgia compressa. — Versluys, 1906:81. — Kiikenthal, 1924:276. 

Material examined.— Amchi\kdi, Aleutian Islands, 51°32'N, 179°15'W, 278-289 
m, K. K. Chew, coll. 2 September 1968: 4 colonies, complete or nearly so, USNM 
60281 (Figs. 11, 12, 14). 

Bering Sea, 52°05'N, 177°40'E, 100 m. Albatross sta. 3599, 9 June 1894: 1 
colony, USNM 57542. 

Bering Sea, 52°14'N, 174°13'E, 882 m. Albatross sta. 4781, 7 June 1906: 1 
colony, somewhat broken, USNM 57543 (Fig. 13). 

Description. — Colonies flabellate, branched in a very loose pinnate manner that 
simulates dichotomous (i.e., quasi-dichotomous), lateral branches arising alter- 



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139 




Fig. 14. Fanellia compressa (Verrill), USNM 60281: opercular, abaxial, outer-lateral sclerites of 
polyp. Scale = 0.1 mm. 



140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

nately, mostly 30^0 mm apart, the undivided terminal branches up to 30 cm 
long, nearly straight. The polyps are situated around the branches in closely 
placed whorls of 8-12 distally, increasing in numbers proximad until in the major 
branches there may be 14-16; on the larger branches some of the whorls may be 
oblique, some incomplete, resulting in a spiral appearance; 4-6 whorls occur in 
1 cm of length. The polyps are about 1.5 mm tall in the contracted state; the two 
abaxial rows of sclerites consist of 10-11 scales each, the outer laterals 5-7, the 
inner laterals 2-3 , and the adaxials 1 . Because of the greater number of inner and 
outer lateral scales in F. compressa as compared with F. fraseri, the winglike 
lateral expansion of the abaxial scales has room for development only on those 
in the more proximal positions, so it occurs only from Abax-7 to 10; the outer 
laterals, beginning between OL-4 and 6, depending upon the number of inner 
laterals present, also develop lateral expansions, as may one or more of the inner 
laterals. All of the body scales are sculptured externally by closely placed, mi- 
nutely aculeate ridges irregularly radiating outward from the nucleus. Where ex- 
posed, the coenenchymal sclerites are large, closely fitting polygonal plates sim- 
ilarly sculptured, but where covered by the contracted polyps they are smaller 
tuberculate spheroids. Small, warty spheroids also occur sparsely in the meso- 
gloea between the longitudinal stem canals and immediately surrounding the axis 
(i.e., the axial sheath). 

Comparisons. — This species resembles F. fraseri but seems to grow larger. 
The polyps are larger, the lateral scale rows better developed, and the sculpturing 
more complex. 

Remarks. — Although Verrill's original specimens lacked coenenchyme, the lo- 
cality from which they came makes it probable that they were the species here 
described. Prymnoa verticillaris Ehrenberg could have been either this species 
or F. fraseri, as Studer (1878:647) pointed out that Ehrenberg' s material was 
obtained in the North Pacific by the Rurik Expedition. Ehrenberg 's statement 
that the polyps had 8 scales in the abaxial rows ("scutellorum serie dorsuali 
longitudinali duplici, trasversis 8") makes it slightly more likely that he hsid fraseri 
in hand, although the difficulty of counting scales accurately leaves room for 
doubt. The matter is of no nomenclatural significance because Ehrenberg used 
the variant spelling employed by Linnaeus for Gorgonia verticillata Pallas. 

Fanellia fraseri (Hickson, 1915) 
Figs. 15-17 

Caligorgia fraseri Hickson, 1915:553, fig. 4, pi. 1, fig. 2 (Albatross and Portlock 
banks. Gulf of Alaska, 50-100 fath. on halibut lines); 1917:23. — Kiikenthal, 
1924:279. 

Material examined. — Gulf of Alaska: Albatross and Portlock banks, 50-100 
fathoms (92-183 m); two loosely pinnate branches, paratypes, British Museum 
(Nat. Hist.) 1962.7.20.821 (figured by Hickson) and 1962.7.20.822 (Fig. 15B).— 
Albatross Bank, 100-125 fath. (183-229 m); M/S Dorothy, Alaska King Crab 
Investigations, sta. C-45, 28 March 1941; three branches, probably from a single 
colony, USNM 51284 (Fig. 15A). 

Description. — Like Hickson' s original specimens, the present topotypic ma- 
terial consists of branches possibly from a single colony. Two of the branches 



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Fig. 15. Fanellia fraseri (Hickson): A, USNM 51284; B, paratypes, BM(NH) 1962.7.20.821 (fig- 
ured by Hickson) and 1962.7.20.822. Scale in cm. 



(one illustrated in Fig. 15A, right) are very loosely pinnate just as Hickson 
(1915:553) described the types, but the third (Fig. 15A, left) is more distinctly 
pinnate and probably came from a position lower in the colony. The longest 
undivided terminal branchlet is 15 cm long. Distally the polyps are set in whorls 
of 5-8, increasing to 12 in the lowest parts of the colony preserved; 6-7 whorls 
occur in 1 cm of length. The polyps are about 1 mm tall in the contracted state; 
the two abaxial rows of sclerites consist of 7-8 scales each (although the distinc- 
tion between the lowest abaxial scale and the adjoining coenenchymal sclerites 
is often so unclear that an accurate count is difficult to make), and the 3rd, 4th 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 16. Fanellia fraseri (Hickson), USNM 51284: top. whorl of polyps, x25; bottom, oblique 
view of polyp, x42. SEM stereo pairs. 



and 5th on each side have inconspicuous to moderate "wingHke expansions" (as 
observed by Versluys, 1906:81, Fig. 95, for C. tuberculata); the outer-lateral rows 
usually have 2 scales each, the inner laterals 1, and the adaxials 1 small scale 
(sometimes missing) below each opercular; the larger coenenchymal sclerites are 
closely fitting polygonal plates externally covered by irregular, close-set ridges; 
where covered by the contracted polyps, they are smaller, tuberculated grains of 
irregular shape. 

Comparisons. — This species is generally similar to F. compressa, with the 
following differences: branching tends to be somewhat closer and therefore more 
distinctly pinnate; polyps smaller, fewer scales in the abaxial and outer-lateral 



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Fig. 17. Fanellia fraseri (Hickson), USNM 51284: opercular, adaxial marginal, abaxial, outer- 
lateral and inner-lateral sclerites of polyp. Scale = 0. 1 mm. 



144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

rows; winglike lateral expansions of the body scales weakly or moderately de- 
veloped on Abax-3, 4 and 5, and not at all on outer laterals or inner laterals. 

Remarks. — The type-lot was obtained by Prof. Arthur Willey from halibut lines 
on Albatross and Portlock banks in the Gulf of Alaska, where it was reported as 
common. The color in life was described as pink. 

Fanellia granulosa (Kinoshita, 1907) 

Caligorgia granulosa Kinoshita, 1907:231 ("Sagami- und Kagoshima-See"); 

1908a:37, pi. 2, figs. 13-14, pi. 6, fig. 46 ("Westkiiste von Satsuma"); 1908b: 

pi. 18, fig. 2; 1909:2. 
Caligorgia tuberculata. — Kiikenthal, 1924:278 (part). 

Material examined. — None. 

Discussion. — Kinoshita (1908a: 40) stated that the sculpture of the sclerites of C. 
aspera and C. granulosa is the same, but that the species differ in branching and 
arrangement of polyps. Although his very clear photographs (PI. 2, figs. 13, 14) 
of C. granulosa show a distinctly pinnate colony, they also show that the distal 
parts of branches, if isolated, could easily be interpreted as dichotomous. More- 
over, the differences between the polyps of the two are negligible; even the 
presence of 1-3 adaxial body scales below the operculum in aspera compared 
with "vereinzelten Schuppen" in granulosa is of no significance. In view of the 
intraspecific variation in colonial form that can occur under different ecological 
conditions, it seems likely that C. granulosa, C. aspera and C. tuberculata are 
indeed but a single species as interpreted by Kiikenthal. Although there is little 
doubt that Caligorgia aspera Kinoshita is a junior synonym of Fanellia tuber- 
culata (Versluys), I prefer for the moment to retain Kinoshita' s C. granulosa as 
a distinct species, now referred to the genus Fanellia, because no authentic ma- 
terial has been available for examination. 

Diagnosis. — Colonies pinnately branched. 

Fanellia tuberculata (Versluys) 
Figs. 18-26 

Caligorgia tuberculata Versluys, 1906:80, figs. 95, 96, pi. 6, fig. 15 (Sulu-Inslen, 

5°43.5'N, 119°40'E, 522 m).— Kukenthal, 1924:278 (part). 
Caligorgia aspera Kinoshita, 1908a:39, pi. 2, figs. 15, 16; pi. 6, fig. 47 (Westkiiste 

von Satsuma). — Nutting, 1912:61. 
Caligorgia flabellum. — Nutting, 1912:60. 
Caligorgia pseudo flabellum Bayer, 1949:207, fig. 2a-c, pi. 4, fig. 2 (Bikini Atoll, 

off Enyu Pass, 11°29'28"N, 165°31'40"E, 116-120 fath.). 

Material examined.— YLsiwausin Islands: Middle Bank, 22°47.5'N, 161°02.2'W, 
to 22°47.9'N, 161°02.3'W, 382 m, Townsend Cromwell TC78, 11 January 1978; 
broken branches, possibly from one large colony, USNM 56791 (Fig. 19). — Kaena 
Point, Oahu, 2r35.85'N, 158°24.55'W, 275^45 m, "Sango XII" haul 1, 27 July 
1971; broken branches and part of main trunk of a large colony, USNM 60348. — 
Makapuu, Oahu, 2ri8.0'N, 157°32.7'W, 362 m, "Sango XIV" haul 1, 18 January 
1972; 1 small colony lacking holdfast, USNM 60290 (Figs. 18, 20). 



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Fig. 18. Fanellia tuberculata (Versluys) from Hawaii, USNM 60290: typical quasi-dichotomous 
colony with sterile twig tips. Scale divisions are mm. 



Marshall Islands: Bikini Atoll, off Enyu Pass, ir29'28"N, 165°3r40"E, 212-220 
m, dredge sta. 30, 22 August 1947; one branch, broken in two pieces, from a 
larger colony, holotype of Caligorgia pseudoflabellum Bayer, USNM 44089 (Figs. 
21, 22). 

Japan: Honshu, Ose Zaki S.36°, W 0.8 mi, 65-125 fath., volcanic sand, shells, 
rock, Albatross sta. 3716, 11 May 1900; large dichotomously divided branch 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 19. Fanellia tuberculata (Versluys) from Hawaii, USNM 56791: right and left lateral views of 
polyp. SEM stereo pairs, x56. 



lacking holdfast, USNM 49582 (Fig. 23); smaller branch, part of preceding?, 
USNM 57523 .—Kyushu, off Kagoshima Wan, 30°54'40"N, 130°37'30"E, 103fath., 
stones. Albatross sta. 4936, 16 August 1906; flabellate colony with holdfast, with 
broken branches, Caligorgia flabellum det. Nutting, USNM 30029 (Figs. 24, 26); 
branch without holdfast, part of preceding?, Caligorgia aspera det. Nutting, 
USNM 30075. — Kyushu, Westkiiste von Satsuma; small branch of colony det. 



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Fig. 20. Fanellia tuberculata (Versluys) from Hawaii, USNM 60290: opercular, adaxial marginal, 
abaxial, and outer-lateral sclerites of polyp. Scale = 0.1 mm. 



K. Kinoshita in Zoological Institute, University of Tokyo, USNM 50120. — Kyu- 
shu, Kagoshima, 70 fath., 15 July 1899; small branch of colony det. K. Kinoshita 
in Zoological Institute, University of Tokyo, USNM 60291 (Fig. 25). 

Description. — Branching quasi-dichotomous or loosely pinnate, alternate, com- 
pressed or in one plane. Polyps in whorls of 2 or 3 near tips of terminal twigs, 
increasing proximally to 5 in some colonies, and as many as 7 or 8 on larger 
branches, often becoming irregularly scattered; usually 7 or 8 whorls in 1 cm of 
axial length in the terminal twigs, sometimes as few as 6 or as many as 9. Polyps 
typically with 6 scales in the abaxial rows, but occasionally as few as 4 and as 
many as 8; outer lateral rows with 1 or 2 scales; inner lateral rows with only 1 
and this may be absent on one or both sides, its place occupied by wide lateral 
extension of the outer-lateral scale (OL-1); OL-2 if present may extend laterally 
as a curved lobe beneath the inner-lateral marginal, but a separate second inner 
lateral (IL-2) has not been observed; 1-3 small scales, commonly 1 or 2, in the 
adaxial rows beneath the adaxial operculars. Tentacles without sclerites. Coe- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2L Fanellia tuber culata (Versluys) from Marshall Islands, USNM 44089, holotype of Cal- 
logorgia pseudoflabellum Bayer: top, whorl of polyps, x44; bottom, lateral view of polyp, x56. SEM 
stereo pairs. 



nenchyme covered with closely fitted, rounded-polygonal plates. All sclerites 
sculptured externally by closely placed, serrated, fluted or ridged tubercles 
aligned in rows radiating outward from the depositional center ("nucleus"), in- 
ternally by crowded, complex tubercles. 

Comparisons. — Kinoshita (1908a:40) remarked upon the similarity of his Cali- 
gorgia aspera to C. tuberculata Versluys, but justified its status as a distinct 
species on the basis of differences in the form of the external warts of the scales, 
and on the absence of the sterile twig tips reported in C. tuberculata by Versluys 
(1906:80). These sterile tips are of inconsistent occurrence in Hawaiian material, 



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Fig. 22. Fanellia tuberculata (Versluys) from Marshall Islands, USNM 44089, holotype of Cal- 
logorgia pseudoflabellum Bayer: opercular, adaxial marginal, abaxial and outer-lateral sclerites of 
polyp. Scale = 0.1 mm. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 23. Fanellia tuberculata (Versluys), quasi-dichotomous colony from Japan, USNM 49582. 
Scale in cm. 



but it is not possible to determine whether they are present in some colonies but 
not in others, because the material was so badly broken during collection. Some 
of the specimens with sterile branch tips are infested with colonies of a thecate 
hydroid which seem to grow only on these sterile tips, but I cannot determine 



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Fig. 24. Fanellia tuberculata (Versluys), loose pinnate colony from Japan, USNM 30029. Scale 
divisions are mm. 



whether the hydroids stimulate their production. Such branch tips are not present 
in any of the Japanese specimens examined. 

It should be noted that Nutting (1912:60, 61) reported two specimens from one 
and the same Albatross station, probably parts of one colony, as two different 
species, Caligorgia flabellum (wrongly attributed to KoUiker) and C. aspera Ki- 
noshita, the former described as "regularly alternate," the latter as "dividing 
dichotomously." 

Although the holotype colony of Caligorgia pseudoflabellum is pinnately 
branched (Bayer 1949:207), the branchlets are widely spaced as well as further 
subdivided, and the main axis bends slightly away from the branch at each node, 
resulting in a loose pinnate pattern that can easily pass for dichotomous. This is 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



rim^f^ 



V-* 

f 




Fig. 25. Fanellia tuberculata (Versluys) from Japan; original material of C. aspera determined by 
Kinoshita, USNM 60291. Right and left lateral views of polyps. SEM stereo pairs, x56. 



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153 




Fig. 26. Fanellia tuberculata (Versluys) from Japan, USNM 30029: opercular, adaxial marginal, 
abaxial, and outer-lateral sclerites of polyp. Scale = 0.1 mm. 



154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

clearly seen in the photograph (Bayer 1949: pi. 4, fig. 2), in which the distalmost 
part of the right-hand fragment would justifiably be described as dichotomous if 
it were broken off from the lower part. As this growth form does not differ 
significantly from that illustrated for C. aspera by Kinoshita (1908a: pi. 2, figs. 15, 
16), and as Kinoshita observed that "die junge Kolonie sowohl als der Stam- 
mabschnitt der erwachsenen sind beinahe federartig verzweigt" it is evident that 
this character is unreliable. The distribution of the polyps and the arrangement 
of their body sclerites are in such close agreement in C. tuberculata, C. aspera, 
C. granulosa, and C. pseudoflabellum that it is probable all four nominal taxa 
belong to a single species. Although no authentic material of C. granulosa has 
been available to me for examination, the specimen from Albatross sta. 4936 off 
Kyushu (Fig. 24) is loosely pinnate and approaches (but does not duplicate) the 
close-pinnate form originally described for that species. However, on the strength 
of existing data, this specimen must be referred to as C. aspera, leaving C. 
granulosa as a distinct nominal taxon pending detailed study. 
Distribution. — Sulu Archipelago, Japan, Marshall Islands, Hawaii. 

Fanellia corymbosa, new species 
Figs. 27-29 

Material examined. — One colony, broken in several pieces: Moluccas, off Do- 
worra Island, 0°50'00"S, 128°12'E, 205 fath. (375 m), coral sand; Albatross 
sta. D5629, 2 December 1909. Holotype colony, USNM 50146. 

Description. — Branching irregularly dichotomous, not in one plane, producing 
colonies of corymbose form (Fig. 27). Polyps in pairs, the individuals opposite 
or slightly offset, infrequently in whorls of 3 even on the larger branches; polyps 
irregularly scattered on the largest branches, absent from the side opposite the 
direction in which the branchlets grow; 6 or 7 pairs or whorls in 1 cm of branchlet. 
Branches stiff and rather brittle, the terminal branchlets commonly 10 mm long 
or less, and 0.75 mm in diameter (exclusive of polyps), with the axis 0.3-0.4 mm 
in diameter proximally, tapering to a rather coarse point. Diameter of the largest 
main stem about 3.5 mm exclusive of cortex; axis heavily calcified, brittle, surface 
inconspicuously grooved longitudinally, appearing nearly smooth, distally creamy 
yellow with golden iridescence, proximally brown with bronzy gloss. 

Polyps (Fig. 28) with only the 2 abaxial rows of sclerites well developed, 5 or 
6 thick plates in each, the boundary between proximal abaxial and cortical plates 
difficult to determine; one wide lateral scale in marginal position on both sides, 
bearing the outer-lateral and inner-lateral opercular scales; occasionally the lateral 
marginal plate is divided into two, thus representing outer-lateral and inner-lateral 
plates; one or two (usually one) small adaxial scale beneath each adaxial oper- 
cular, but the remaining adaxial surface of the polyps naked. Body plates (Fig. 
29) thick, the external tubercles near the distal margin forming blunt dentations 
that interlock with the proximal tubercles of the following sclerite, inner surface 
covered with fine, complex tubercles; opercular scales triangular, the laterals 
only slightly asymmetrical, the adaxials somewhat smaller than the others but 
not markedly different in shape; on the inner surface of the operculars a low, 
strong longitudinal ridge extends from center to apex, most prominent on the 
dominant abaxial, progressively smaller on the outer and inner laterals and adax- 



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Fig. 27. Fanellia corymbosa, new species; holotype colony, USNM 50146. Width 35 cm. 



ials; outer surface convex, a prominent rounded boss at the nucleus, covered 
with closely set hemispherical tubercles that become lower and less prominent 
toward the edges and elongated in shape toward the apex of the scale. Tentacles 
without sclerites. 

Coenenchymal sclerites rounded or spheroidal pebbles with hemispherical tu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 28. Fanellia corymbosa, new species; holotype, 
of polyp. SEM stereo pairs, x45. 



USNM 50146: abaxial and right lateral views 



bercles on outer surface, finer complex tubercles on inner surface; largest sclerites 
become thick, irregular bodies resembling coarse cobblestone pavement. 

Comparisons. — In many ways, this species resembles Kinoshita's Callogorgia 
aspera and C. granulosa, the first definitely and the second possibly a synonym 
of Fanellia tuberculata (Versluys), but differs from both in growth form. F. 
tuberculata, from the Sulu Archipelago, usually has 2 plates in the outer-lateral 
rows and 1 in the inner-lateral rows, whereas F. corymbosa has only a single 
outer-lateral plate on each side, extending into the inner-lateral position where 
no separate plate is developed. The lower abaxial plates of corymbosa are much 
wider (to 0.6 mm) than those of tuberculata (0.3-0.4 mm). No sterile terminal 
branch tips were observed in corymbosa. 



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Fig. 29. Fanellia corymbosa, new species; holotype, USNM 50146: opercular, abaxial marginal 
and abaxial proximal body sclerites. Scale = 0. 1 mm. 



158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Acknowledgments 

I am grateful to Dr. P. F. S. Cornelius and Mr. Gordon Paterson of the Coel- 
enterate Section, British Museum (Nat. Hist.), who made available for study 
paratypes of Hickson's Caligorgia fraseri. Dr. K. K. Chew supplied, through the 
Smithsonian Oceanographic Sorting Center, some of the material of Fanellia 
compressa here described. The specimens of Callogorgia formosa and Fanellia 
tuberculata from Hawaiian waters were provided by Dr. Richard Grigg. The speci- 
mens of C aspera identified by K. Kinoshita were made available to me in Tokyo 
by Dr. K. Takewaki of Tokyo University. All scanning electron micrographs were 
made by Mr. Walter R. Brown, chief of the SEM Laboratory, National Museum 
of Natural History, Smithsonian Institution. Photographs of colonies were made 
by Mr. Michael R. Carpenter, who also prepared the photographic prints re- 
produced in this paper. Dr. Stephen D. Cairns kindly reviewed the manuscript. 
To all of these I express my thanks. 

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dique de Roret. 
Molander, A. R. 1929. Die Octactiniarien. — Further Zoological Results of the Swedish Antarctic 

Expedition 1901-1903, 2(2):i-iv + 1-86, pis. 1-5. 
Nutting, C. C. 1908. Descriptions of the Alcyonaria collected by the U.S. Bureau of Fisheries steamer 

Albatross in the vicinity of the Hawaiian Islands in 1902. — Proceedings of the United States 

National Museum 34:543-601, pis. 41-51. 
. 1912. Descriptions of the Alcyonaria collected by the U.S. Fisheries steamer "Albatross," 

mainly in Japanese waters, during 1906. — Proceedings of the United States National Museum 

43:1-104, pis. 1-21. 
Pallas, P. S. 1766. Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum 

cognitarum succinctas descriptiones cum selectis auctorum synonymis. Hagae Comitum. 

i-xvi + 17-28 + 1^51 pp. 
Statins Miiller, P. L. 1775. Von den Corallen. In Des Ritters Carl von Linne . . . vollstandiges 

Natursystem . . . 2(6):i-xvi + 641-960 [1068], pis. 20-37. Niirnberg, Gabriel Nicolaus Raspe. 
Studer, T. 1878. Ubersicht der Anthozoa Alcyonaria, welche wahrend der Reise S.M.S. Gazelle um 

die Erde Gesammelt wurden. — Monatsbericht der Koniglich Preussischen Akademie der Wis- 
senschaften zu Berlin 1878:632-688, pis. 1-5. 

. 1887. Versuch eines Systemes der Alcyonaria. Archiv fiir Naturgeschichte 53(1): 1-74, pi. 1. 

Thomson, J. A., and N. I. Rennet. 1931. Alcyonaria, Madreporaria, and Antipatharia. — Australasian 

Antarctic Expedition 191 1-14. Scientific Reports (C-Zoology and Botany) 9(3): 1-46, pis. 8-14. 
Verrill, A. E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedition, 



160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

under Commodore C. Ringgold and Captain John Rodgers. U.S.N. . from 1853 to 1856. Col- 
lected by Dr. Wm. Stimpson. naturalist to the Expedition. With descriptions of some additional 
species from the west coast of North America. — Proceedings of the Essex Institute, Salem, 
4:181-196. pis. 5-6. 

Versluys. J. 1906. Die Gorgoniden der Sihoga Expedition II. Die Primnoidae. — Siboga-Expeditie 
Monographie 13a. Pp. 1-187. figs. 1-178, pis. 1-10. chart. 

Wright. E. P.. and T. Studer. 1889. Report on the Alcyonaria collected by H.M.S. Challenger during 
the years 1873-1876. — Voyage of the Challenger, Zoology, 31:i-lxxii + 1-314, 43 pis. 

Department of Invertebrate Zoology. Smithsonian Institution, Washington, 
D.C. 20560. 



PROC. BJOL. SOC. WASH. 
95nj, 1982, pp. 161-166 

FRESHWATER TRICLADS TTURBELLARIA) OF NORTH 

AMERICA. XII J. FHAGOCATA HAMPTONAE, 

NEW SPECIES, FROM NEVADA 

Roman Kenk 

Abstract. — A new species of the planarian genus Pha^ocata, P. hamptonae, 
is described. It differs from all other North American species of the genus by 
having 4 to ] 1 eyes in the usual position. In the copulatory apparatus, the struc- 
ture of the penis deviates from the general plan by having a wide ejaculatory duct 
filled with an eosinophilic secretion. The species forms spermatophores that are 
inserted into the bursa of the copulating partner. 



In an extensive field study of the freshwater triclads of the Lake Tahoe basin 
and the surrounding area, Dr. Anne M. Hampton discovered a white planarian 
with more than one pair of eyes, that she recognized as being a new taxon. She 
kindly sent me a number of live specimens for a more detailed study. Some of 
the worms were sexually mature. 

Phagocata hamptonae, new species 

Type-material. — All type-specimens are deposited in the National Museum of 
Natural History, Smithsonian Institution. Washington. D.C. Holotype. set of 
sagittal sections on 3 slides f L'SNM 68009j: paratypes. 5 sets of sagittal sections 
on 3 slides each (USNM 680KJ-68014) and one set of transverse sections of 
posterior part of body on 3 slides (USNM 68015). 

External features (Fig. \). — The species is unpigmented. white, and at first 
glance resembles the eastern Phagocata morgani (Stevens and Boring). Mature 
specimens reach a length of 1 1 mm and a width of 1.3 mm. The anterior end is 
truncate, with a straight or some v. hat wavy frontal margin (slightly convex in the 
center), changing somewhat during gliding locomotion, and with rounded lateral 
corners. Behind the anterior end. a slight narrowing or "'neck" may be seen in 
the gliding animal. The body soon reaches its maximum width, the lateral margins 
running parallel in the greater part of the body length, converging in the last fifth 
of the body, and meeting at the bluntly pointed posterior end. 

The most characteristic feature of the species is the plurality of the eyes. While 
most species of Phagocata have only one pair of eyes or are blind. P. hamptonae 
has a small but variable number of eyes (Fig. Ic). Dr. Hampton, who analyzed 
37 specimens, found their number to vary between 4 and 11, the mean number 
being 6. arranged generally in two longitudinal rows separated by about one-fifth 
the transverse diameter of the head. The eyes are rather small, of about equal 
size, but fully functional, each equipped with a well-developed pigment cup. 

The pharynx measures about 1 5 the body length, its root being inserted anterior 
to the middle of the body in mature specimens. The intestine is amply ramified, 
which makes it difficult to count the branches of the intestinal trunks. By feeding 
some specimens a mixture of beef-liver tissue and carbon powder and later ex- 



162 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




B 








C 



Fig. 1. Phagocata hamptonae: A, Photograph of living specimen after ingestion of a mixture 
of beef-liver tissue and carbon powder to show the intestine, x7.5; B, Outline drawing with indica- 
tion of eyes, pharynx, and copulatory apparatus; C, Arrangement of eyes, redrawn from sketches 
prepared by A. M. Hampton. 



amining them under slight compression, the approximate number of branches 
could be established: 4-6 heavily ramified branches on either side of the anterior 
trunk; the two posterior trunks each bear about 14-19 less ramified lateral branch- 
es and a considerable number of short medial diverticula. Postpharyngeal anas- 
tomoses between the two trunks may occur. 

Anatomy. — The anterior end shows no special structures (glands or muscles) 
that could be interpreted as corresponding to an adhesive organ. The zone of 
eosinophilic adhesive glands, that runs along the ventral side of the body margins, 
is interrupted in the central area of the frontal margin, where there is a field of 
infranucleate ciliated epithelium, apparently a sensory organ. The pharynx has 
the normal structure characteristic of the Planariidae, its internal muscular zone 
consisting of a thick layer of circular fibers surrounding the pharyngeal canal, 
followed by a layer of longitudinal muscles. 

The two ovaries or germaria are situated on the medial side of each ventral 
nerve cord at the level of the first lateral branches of the intestine. Each ovary 
is accompanied by a group of cells about the size of the ovary, situated anterior 
to it. Such cell accumulations adjoining the ovaria are widely distributed in tri- 



VOLUME 95, NUMBER 1 



163 



0.1mm, 




Fig. 2. Phagocata hamptonae , semidiagrammatic view of copulatory apparatus in sagittal section: 
am, male atrium; b, copulatory bursa; bd, bursal duct; bp, penis bulb; de, ejaculatory duct; gp, 
gonopore; m, mouth; ode, common oviduct;/?/?, penis papilla; vd, vas deferens; vs, seminal vesicle. 



clads and are usually termed "parovaria." In P. hamptonae these cells corre- 
spond entirely in size, stainability, and cytoplasmic inclusions (yolk granules) to 
the cells of the vitellaria, a fact that confirms the often expressed assumption that 
the parovaria have a genetic, if not always functional, relation to the yolk glands. 

The numerous testicles are of moderate size, usually measuring less than half 
the dorsoventral diameter of the body. They are predominantly ventral and are 
arranged in longitudinal rows beginning a short distance posterior to the ovaries 
and extending to almost the hind end of the body. The two sperm ducts or vasa 
deferentia run along the ventral nerve cords medial to the oviducts. In the region 
of the pharynx they expand to form the spermiductal vesicles or "false seminal 
vesicles" which, filled with sperm, proceed posteriorly to their entrance into the 
penis bulb. 

The copulatory apparatus (Fig. 2) closely adjoins the pharyngeal pouch. The 
genital atrium is confined to the male atrium {am), since the bursal duct {bd) 
proceeds directly to the gonopore {gp) without entering a separate atrial com- 
partment. The penis consists of a relatively large bulb {bp) and a conical papilla 
ipp). The bulb has its musculature developed principally near its periphery, while 
its central part is remarkably devoid of muscle fibers. The lumen of the penis is 
clearly divided into two sections. An anterior cavity, the seminal vesicle {vs), 
located in the bulb, is lined with an epithelium of cuboidal, apparently apocrine, 
cells, the secretions of which must have dissolved during the procedures used in 
preparing the sections, so that the cells look empty (see Fig. 3). The posterior 
part {de) of the lumen is a rather wide, elongated cavity, separated from the 



164 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 









. ^ T'^^"^ 



Fig. 3. Phagocata hamptonae , photomicrograph of a sagittal section through the copulatory com- 
plex of a specimen that has two spermatophores in the copulatory bursa, xl20. 



seminal vesicle by a slight constriction and extending from the penis bulb to the 
tip of the papilla, tapering gradually in its posterior portion. It is almost com- 
pletely filled with strands of a granular, slightly eosinophilic secretion, the origin 
of which is not quite clear. It may be in part produced by the epithelial cells 
lining the cavity, but traces of apparently the same secretion are seen also in the 
parenchymal part of the penis papilla, which would hint at gland ducts entering 
from the mesenchyme surrounding the penis bulb. This posterior part of the penis 
lumen corresponds by its location to the ejaculatory duct of related species, 
although it may have a different function, possibly related to the production of 
spermatophores. The two sperm ducts or vasa deferentia {vd) enter the penis 
bulb ventrolaterally, turn upward, and open into the seminal vesicle separately. 

The two oviducts or ovovitelloducts run along the dorsal side of the ventral 
nerve cords up to the level of the male atrium. There they turn dorsally, approach 
the midline, and unite in the space between the atrium and the bursal duct. The 
common oviduct {ode) thus formed proceeds posteroventrally in a short curve 
along the atrial wall and opens into the posterior part of the atrium. The paired 
oviducts, after they have left the nerve cords, and the upper part of the common 
oviduct receive the highly eosinophilic shell glands. 

The copulatory bursa {h) is of variable size and shape. In some of the specimens 
it contained remnants of spermatophores (see Fig. 3). Spermatophores were also 
visible in some living specimens as glittering spherules shining through the trans- 
parent body wall. 

The bursal stalk or duct {bd) runs from the bursa posteriorly above the penis 
and atrium, then curves ventrally, slightly increasing in diameter, and opens at 
the gonopore {gp) to the left of the outlet of the male atrium. No differentiated 



VOLUME 95, NUMBER 1 165 

end part or vagina is developed and the duct is histologically uniform throughout. 
Its musculature consists of a subepithelial layer of circular fibers, surrounded by 
a layer of longitudinal muscles. 

All epithelia of the copulatory complex are nucleate, including the common 
oviduct, while the paired oviducts show an infranucleate epithelium. 

Distribution and ecology. — Phagocata hamptonae was collected by Dr. Anne 
M. Hampton in an irrigation ditch 400 m east of the ranch house on the Gene 
Scossa Ranch, Gardnerville, Douglas County, Nevada, on 5 April 1981 (water 
temperature, 15°C, pH 7.5) and 5 July 1981. Dr. Hampton sent me samples from 
both collections, containing some sexually mature and some immature specimens. 
The worms were placed in cultures kept at 14°C. They readily accepted beef liver 
as food. 

Taxonomic position. — The genus Phagocata is admittedly a very heteroge- 
neous genus, distributed widely in the Northern Hemisphere in both Eurasia and 
North America (see also discussion by Ball and Gourbault 1975: 11-13). The mem- 
bers of the genus conform in the basic structure of the copulatory apparatus, 
though they may differ in many other characters that are of taxonomic signifi- 
cance. In some species the testicular zone extends to the tail end, in others only 
to the level of the mouth; some are pigmented, others white; they generally have 
one pair of eyes, but some are blind, and the present species has multiple eyes 
(as does P. uenoi Okugawa from Manchukuo, see Okugawa 1939:157). It must 
be stressed that the eyes are completely formed eyes with no intergrades between 
functional eyes and apparently nonfunctional pigment clumps, such as are seen 
in P. morgani polycelis Kenk (1935:103), Dendrocoelopsis americana (Hyman) 
(Kenk 1973:14) and some species of Dendrocoelum. The structure of the penis 
may vary in the genus within wide limits even in species that are closely related 
both morphologically and zoogeographically. Within certain geographic regions 
we may find groups of species that are undoubtedly monophyletic and could be 
gathered into separate subgenera or even genera. Phagocata hamptonae has the 
zone of testes approaching the posterior end of the body, a character it shares 
with nine of the North American species of the genus. On the other hand, P. 
morgani (Stevens and Boring), P. bursaperforata Darlington, P. bulbosa Kenk, 
P. angusta Kenk, and P. holleri Kenk have no testes behind the copulatory 
apparatus. Phagocata hamptonae is monopharyngeal as are all the other North 
American species except P. gracilis (Haldeman), P. woodworthi Hyman, and P. 
nordeni Kenk. The new species may well deserve to be placed in a separate 
subgenus, principally on account of the aberrant morphology of its penis. This 
should be done, however, within the framework of a reexamination of the entire 
genus. 

The species is named in honor of its collector. Dr. Anne Marie Hampton of 
Tahoe Paradise, California. 

Acknowledgments 

Thanks are due to Dr. Hampton for the transmittal of the material and for her 
generous collaboration in the study of the new species. Dr. John C. Harshbarger 
of the Smithsonian Institution was helpful in the preparation of the photomicro- 
graph and Dr. Marian H. Pettibone kindly reviewed the manuscript. 



166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Literature Cited 

Ball, I. R., and N. Gourbault. 1975. The morphology, karyology, and taxonomy of a new freshwater 
planarian of the genus Phagocata from California (Platyhelminthes: Turbellaria). — Life Sci- 
ences Contributions, Royal Ontario Museum 105:1-19. 

Kenk, R. 1935. Studies on Virginian triclads. — Journal of the Elisha Mitchell Scientific Society 
51:79-133. 

. 1973. Freshwater triclads (Turbellaria) of North America. VI. The genus Dendrocoelopsis . — 

Smithsonian Contributions to Zoology 138:1-16. 

Okugawa, K. 1939. Probursalia (Triclada-Paludicola) of Manchoukuo. — Annotationes Zoologicae Ja- 
ponenses 18:155-165. 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 167-187 

CLASSIFICATORY REVISIONS IN GAMMARIDEAN 
AMPHIPODA (CRUSTACEA), PART 2 

J. Laurens Barnard and Gordan S. Karaman 

Abstract. — The new family Paracalliopidae is established and the family Pseud- 
amphilochidae is recognized from provisional proposal. 

New genera are described as follows: Relictomoera for Paramoera relicta and 
Sternomoera for P. yezoensis and P. Japonica; Nasageneia for Pontogeneia 
nasa; Aurohornellia for Tulearogammarus sinuatus; Lupimaera for Maera lu- 
pana; Tegano for Melita seticornis; Maleriopa for Eriopisella dentifera; Texi- 
weckeliopsis for Texiweckelia insolita ; Holsingerius for T. samacos ; Indocalliope 
for Paracalliope indica ; Feriharpinia for Harpinia ferenteria ; Torridoharpinia 
for Proharpinia hurley i. 

New synonymies are provided for Afrochiltonia {=Austrochiltonia), Cerado- 
copsis {=Maeracunha) , Hornellia {=Metaceradocus). Echiuropus is raised to full 
generic level. New lists of species are provided for Pseudomoera, Paracalliope 
and Proharpinia. 



More "armchair" revisions in Amphipoda must be presented before we can 
issue our forthcoming "The Families and Genera of Gammaridean Amphipoda," 
a sequel to that done by J. L. Barnard (1969b). This time, freshwater taxa will 
be included with the marine in the same treatise. 

Our first part is cited as Karaman and Barnard (1979). 

Ceinidae 
Afrochiltonia K. H. Barnard, new synonymy 

Afrochiltonia K. H. Barnard, 1955:93 (Chiltonia capensis K. H. Barnard, 1916, 
original designation). 

Austrochiltonia Hurley, 1958:767 (Hyalella australis Sayce, 1901, original des- 
ignation). 

Griffiths (1976) made a case for synonymizing these two genera but mistakenly 
assumed Austrochiltonia took priority. It does not and therefore the situation 
should appear as shown above. 

Eusiridae 
Pseudomoera Schellenberg, revised 

Pseudomoera Schellenberg, 1929:281 {Atyloides gabrieli Sayce, 1901, monotypy). 

Until now this genus has been monotypic but we transfer into this genus Aty- 
loides fontana Sayce (1902) which for years has been placed in Paramoera. The 
type-species is characterized by the loss of medial setae on the inner plate of 
maxilla 1 whereas P. fontana has full setal armament on that plate like Para- 
moera. However, Pseudomoera can now be distinguished from Paramoera on 
the presence of geniculate lobes on the wrists of the gnathopods and this action 



168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

removes from Paramoera a clearly alien species and places it precisely contig- 
uous to its apomorph. This creates an unusual genus in which maxillary setae are 
variable but this is perhaps preferable to adding yet another monotypic genus to 
the pool. 

Species . Montana (Sayce, 1902a); gabrieli (Sayce, 1901); Australia, Victoria, 
streams between 457 and 915 m of altitude. 

Relictomoera, new genus 

Type-species. — Paramoera relicta Ueno, 1971a. 

Body slender, compressed, smooth, urosomites free. Rostrum obsolescent; 
lateral cephalic lobes sinusoid; with notch in middle, anteroventral margin of 
head weakly produced. Eyes round, tiny, or vestigial, or absent. 

Antennae elongate, 1 longer than 2, peduncular articles of antenna 1 progres- 
sively shorter (ratio 21:19:12), article 1 as long as or longer than head, articles 
2-3 scarcely shorter than article 1; article 3 not produced; article 1 of primary 
flagellum ordinary, accessory flagellum 1 -articulate, barrel shaped. 

Labrum [?entire, ?subrounded, broader than long]; epistome unproduced. Mo- 
lar triturative, columnar, article 2 of palp poorly lobed, article 3 shorter than 2 
(ratio = 4:16:13). Labium: inner lobes [?absent]. 

Maxilla 1: inner plate with 5 apical setae, outer plate with 9 spines, palp long, 
article 1 short. Maxilla 2: inner plate not broader nor longer than outer plate, 
narrow, inner plate with facial row of 4 setae but no other medial setae. Maxil- 
liped: inner plate relatively long, outer plate slightly shorter than inner, not spi- 
nose medially; palp of 4 articles, 4 slightly shorter than 3, 3 unlobed, 4 not spinose 
along inferior margin, unguiform, with nail. 

Coxae short but contiguous, almost glabrous, coxa 1 not produced anteriorly 
nor expanded ventrally, coxa 4 with weak posterior lobe, excavate. 

Gnathopods feeble, slender, diverse, 2 larger than 1, subchelate, not eusirid, 
medium, wrist of both as long as hand, without posterior lobe, with few short 
posterior setae, propodi rectangular or weakly expanded, gnathopod 2 especially. 

Pereopods 3-7 ordinary, simple, dactyls simple, article 2 not anteriorly lobate, 
on pereopods 5-7 weakly expanded, weakly or moderately lobate. 

Epimeron 3 smooth. Pleopods with peduncle dominating rami. 

Outer rami of uropods 1-2 slightly shortened or not; rami with lateral and 
dorsal spines. Uropod 3 ordinary, scarcely extended beyond uropod 1, peduncle 
without large process, rami lanceolate, aequiramous, 1 -articulate. 

Telson ordinary, cleft two thirds, apices with short or thin apical armaments. 

Coxal gills present on somites 2-6, ovate, some pediculate. Oostegites broad. 
Sternal gills absent. 

Variables. — Telson elongate {tsushimana). 

Relationship. — Like Paramoera but head with unusually sinusoid anterolateral 
margin. See Sternomoera. Differing from Awacaris Ueno by the normal palp of 
maxilla 1, the normal head and the stronger hand and palm of gnathopod 1. 
Differing from Apherusa in the stronger sinusoid cephalic lobes, the deeply cleft 
telson, and the presence of an accessory flagellum. 

Species. — relicta (Ueno, 1971a); tsushimana (Ueno, 1971b); Goto and Tsu- 
shimana Islands south and west of Kyushu, Japan, hypogean. 



VOLUME 95, NUMBER 1 169 

Etymology. — Name composed of root from species name plus "moera" a com- 
mon root of genera in Eusiridae. Feminine. 



Sternomoera, new genus 

Type-species. — Paramoera yezoensis Ueno, 1933. 

Body ordinary, compressed, smooth, urosomites free. Rostrum very small, 
lateral cephalic lobes rounded or mammilliform, anteroventral margin of head 
scarcely produced, with normal sinus. Eyes reniform. 

Antennae elongate, 1 longer than 2, peduncular articles progressively shorter 
(ratio = 30:18:12), article 1 shorter than head, article 3 not produced; article 1 of 
primary flagellum short, basal articles proliferate, accessory flagellum 1-articu- 
late, scale-like. 

Lab rum entire, subrounded, as long as broad; epistome unproduced. Molar 
triturative, columnar, article 2 of palp unlobed, article 3 as long as 2 (ratio = 
4:9:10). Labium: inner lobes absent. 

Maxilla 1: inner lobe with many medial setae, outer plate with 7 spines, palp 
long, article 1 short. Maxilla 2: inner plate not broader nor longer than outer, 
inner plate with scarcely submarginal facial row of many setae and several other 
medial setae. Maxilliped: inner plate not relatively long, outer plate slightly short- 
er than inner, medially spinose, palp of 4 articles, 4 shorter than 3, 3 unlobed, 4 
not spinose along inferior margin, weakly unguiform, [?with nail]. 

Coxae ordinary, poorly setose, coxa 1 not produced anteriorly or expanded 
ventrally, coxa 4 with posterior lobe, excavate. 

Gnathopods alike in female, almost feeble, subchelate, not eusirid, small wrist 
of both scarcely shorter than hand, without posterior lobe, with numerous pos- 
terior setae, hands rectangular; male gnathopods larger, 2 larger than 1, wrists 
broadly lobate, hands ovate, posterior margins swollen, palms oblique, with few 
large clavate or peg spines. 

Pereopods 3-7 ordinary, simple, dactyls simple; article 2 of pereopods 3^ not 
anteriorly lobate, of pereopods 5-7 poorly expanded, lobate, poorly armed; or of 
pereopod 6 slightly narrowed. Pleopods [?ordinary]. Epimeron 3 serrate sparsely 
and minutely. 

Outer rami of uropods 1-2 not or scarcely shortened; rami with lateral and 
dorsal spines. Uropod 3 ordinary, not extended beyond uropod 1, peduncle with- 
out large process, rami lanceolate. 

Telson ordinary, cleft, apices with small apical armaments. Coxal gills [2-7], 
ovate. Oostegites broad. Thorax with sternal gills. 

Variables. — Apices of telson poorly {yezoensis) to densely setose (japonica) 
apically. 

Relationship. — Like Paramoera but sternobranchiate. Not of modern crango- 
nyctid affinity because of precisely aequiramous uropod 3, short peduncle of 
antenna 1 , basally proliferate flagellum of antenna 1 . Not bogidiellid because of 
the general gnathopodal facies and presence of sternal gills. Differing from Eo- 
niphargus in the magniramous, aequiramous uropod 3 and vestigial accessory 
flagellum. Differing from Relictomoera in the presence of sternal gills and the 
normal head. 



170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Species. — hayamenensis (Stephensen, 1944); japonica (Tattersall, 1922); ye- 
zoensis (Ueno, 1933); Japan, streams. 

Etymology. — Name composed of "sterno" referring to sternal gills and 
"moera" a root used commonly for genera in Eusiridae. Feminine. 

Nasageneia, new genus 

Type-species. — Pontogeneia nasa J. L. Barnard, 1969c. 

Body slender, compressed, smooth. Rostrum large, lateral cephalic lobes or- 
dinary, anteroventral margin of head scarcely produced. Eyes reniform. 

Antennae subequal, peduncular articles of antenna 1 progressively shorter, 
article 1 shorter than head, article 3 weakly produced; article 1 of primary fla- 
gellum ordinary to short, accessory flagellum absent. 

Labrum entire, sub rounded, broader than long, epistome unproduced. Molar 
triturative, columnar, article 2 of palp unlobed, article 3 shorter than 2. Labium: 
inner lobes absent. 

Maxilla 1: inner lobe with medial and 2 apical setae, palp long, article 1 short. 
Maxilla 2: inner plate not broader but slightly longer than outer, inner plate 
without facial row of setae but with other medial setae, few, large, at least one 
slightly submarginal. Maxilliped: inner plate not relatively long, outer plate slight- 
ly shorter than inner; palp of 4 articles, 4 slightly shorter than 3, 3 unlobed, 4 not 
spinose along inferior margin. 

Coxae ordinary to short, coxa 1 not produced anteriorly or expanded ventrally, 
coxa 4 without posterior lobe, excavate. 

Gnathopods diverse, of similar size, subchelate, not eusirid, medium, wrist of 
both shorter than hand, only gnathopod 2 with strong posterior lobe extending 
distad, wrist without numerous long posterior setae, hands rectangular in female, 
inflated in male, in latter with posterior spines outside limits of oblique palm. 

Pereopods 3-7 ordinary, simple, dactyls simple, article 2 not anteriorly lobate. 

Epimeron 3 serrate. 

Outer rami of uropods 1-2 shortened; rami with lateral and dorsal spines. 
Uropod 3 ordinary, not extended beyond uropod 1, peduncle with small process, 
rami lanceolate. 

Telson ordinary, weakly cleft, apices without long apical armaments. 

Relationship. — Like Tethygeneia but epimeron 3 serrate and hands of male 
gnathopods with posterior spines well outside palmar limits as in Gondogeneia. 
Calceoli tending to be much more strongly anthurial than in Tethygeneia, with 
one lobe quite linguiform. 

Species. — nasa (J. L. Barnard, 1969c, 1979); quinsana J. L. Barnard, 1964, 
1969a, 1979); marine, warm temperate California and Mexico, 0-1 m. 

Etymology. — Name composed of "nasa" from species name plus "geneia" a 
name fragment commonly encountered in Eusiridae. Feminine. 

Paramoera stephenseni Barnard and Karaman, new name 

Paramoera brachyura Stephensen, 1949:18, figure 6 [homonym to Paramoera 
brachyura Schellenberg, 1931:201-202, figure 102]. 

Etymology. — Named for K. Stephensen. 



VOLUME 95, NUMBER 1 171 

Gammaridae 
Aurohornellia, new genus 

Type-species. — Tulearogammarus sinuatus Ledoyer, 1967a. 

Body [?ordinary], pleon [?and urosome] dorsally crenulate transversely on pos- 
terior segmental margins, each of pleosomite 2 to urosomite 2 with 2 dorsal 
spines, [?urosomites free]. Rostrum medium, lateral cephalic lobe sharply con- 
ical, sinus present. 

Antennae elongate, antenna 1 shorter than 2, ratio of peduncular articles = 
23:25:10, primary flagellum longer than peduncle, accessory flagellum 2-articu- 
late. Antenna 2 slender. 

Lab rum broader than long, entire, rounded. Mandibular incisor toothed, molar 
triturative, ratio of palp articles = 4:17:15, article 3 Hnear, stubby, setae = ADE. 
Inner lobes of labium present. Maxillae moderately setose medially, inner plate 
of maxilla 1 ovate, very short, moderately setose apicomedially, outer plate with 
[?7] spines, palps 2-articulate, [?symmetrical]. Inner plate of maxilla 2 [?with 
oblique facial row of setae]. Outer plate of maxilliped [?spinose medially, article 
3 of palp unlobed, dactyl shorter than 3, unguiform, with nail]. 

Coxae long, with long setae, coxa 1 slightly expanded apically, coxa 3 much 
smaller than 1 or 2, coxa 4 long but unlobed. Gnathopods feeble, slender, almost 
simple, wrists elongate, hands elongate, thin, palms scarcely evident, gnathopod 
2 thinner than gnathopod 1 (male not clarified). 

Pereopods 3^ ordinary. Article 2 of pereopods 5-7 expanded, moderately to 
strongly lobate, serratosetulose posteriorly; pereopod 7 elongate, dactyl 
[?elongate and setose]. 

Pleopods [?ordinary]. Rami of uropods 1-2 marginally spinose, outer rami 
slightly shortened, peduncle of uropod 1 [?with basofacial spine]. Uropod 3 slight- 
ly extended, magni- or variramous, rami thin, outer ramus with long article 2, 
inner ramus reaching apex of article 1 on outer ramus. Telson of ordinary length, 
fully cleft, lobes tapering, poorly armed but with dorsal spinules. 

Coxal gills [?2-7, ovate]. Oostegites [?narrow]. 

Relationship. — Standing between Hornellia and Megaluropus; like Hornellia 
but coxa 3 reduced and like Megaluropus but rami of uropod 3 thin. 

Species. — sinuata (Ledoyer, 1967a); Madagascar, shallow water. 

Etymology. — From "aurora," dawn, and '"Hornellia,'''' a related genus. Fem- 
inine. 

Ceradocopsis Schellenberg, new synonymy 

Ceradocopsis Schellenberg, 1926:364 (Ceradocopsis kergueleni Schellenberg, 

1926, monotypy). 
Maeracunha Stephensen, 1949:22 (Maeracunha tristanensis Stephensen, 1949, 

monotypy) [new synonym]. 

Lateral cephalic lobes broadly rounded, sinus present but mandible inserted 
there, not antenna 2. 

Antennae elongate, antenna 1 longer than 2, ratio of peduncular articles = 
14:12:6, flagellar ratio = 20:7, accessory flagellum with 4 articles. Antenna 2 
slender. 



172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Mandibular incisor almost smooth, molar small and poorly triturative, palp 
small, ratio of articles = 6:15:12, article 3 linear, setae = E and sparse. Inner 
lobes of labium absent. Maxillae medially setose, inner plate of maxilla 1 pyriform 
to triangular, with medial setae on apical half only, outer plate with 7-11 spines, 
palps [?symmetrical]. Inner plate of maxilla 2 with oblique row of facial setae. 
Outer plate with naked medial margin, palp article 3 weakly lobate, dactyl shorter 
than 3, unguiform, lacking or bearing nail. 

Coxae of medium extension, poorly setose, coxa 1 expanded below, coxa 2 
with anteroventral acuity, coxae 3-4 weakly shorter than 1-2, coxa 4 unexcavate 
posteriorly, coxa 5 as long as 4. Gnathopods diverse, gnathopod 1 small, of 
Melitid form, wrist scarcely elongate, scarcely lobate, hand longer, rectangular, 
palm weakly oblique, short, gnathopod 2 enlarged, wrist short to medium, lobate 
or weakly, hand elongate, large, rectangular, palm weakly oblique, weakly sculp- 
tured or strongly spinose. 

Article 2 of pereopods 5-7 expanded, alike, lobate, poorly setose, serrate and 
straight posteriorly, appendages otherwise stout. 

Rami of uropods 1-2 subequally extended, marginally spinose, peduncle of 
uropod 1 with basofacial spine. Uropod 3 not extended, short, magniramous, 
dispariramous, outer ramus with small article 2. Telson of ordinary length, fully 
cleft, lobes tapering, moderately to strongly armed apically. 

Coxal gills [?2-6]. Oostegites [?slender]. 

Variants. — Outer plate of maxilla 1 with 11 spines ipeke) but other species 
poorly known; though uropod 3 stated as magniramous, actually appearing to be 
parviramous kind with reduced outer ramus now so small as to match inner. 

Relationship. — Differing from Maera in the strongly setose maxilla 2 (facial) 
and the strong article 2 on the outer ramus of uropod 3. Differing from Ceradocus 
by uropod 3 in the same way stated for Maera. Differing from Paraceradocus in 
the short dispariramous uropod 3 and short, slender antenna 2. Differing from 
Ceradocoides in the short, dispariramous uropod 3 and fully cleft telson. Differing 
from Ceradocus in the short dispariramous uropod 3. Differing from various 
genera near Melita in the absence of inner lobes on the labium and the miniatur- 
ized uropod 3. 

Species. — kergueleni Schellenberg, 1926 (Bellan-Santini and Ledoyer 1974); 
peke J. L. Barnard, 1972b; tristanensis (Stephensen, 1949); Antarctica and an- 
tiboreal, especially insular. 

Echiuropus Sowinsky, new synonymy 
Echiuropus Sowinsky, 1915:55 {Echiuropus macronychus Sowinsky, 1915, mono- 

typy). 

[Asprogammarus Bazikalova, 1975:38 (no type-species, therefore unavailable)]. 
[Smaragdogammarus Bazikalova, 1975:64 (no type-species, therefore unavail- 
able)]. 

Bazikalova' s new taxa belong with Echiuropus; she reduced Echiuropus to 
subgeneric level under Asprogammarus which is illegal under ICZN rules. Until 
type-species are selected by Bazikalova the two new taxa are unavailable. We 
trust biologists will allow Bazikalova to rectify this problem rather than capturing 
the taxa for themselves. 



VOLUME 95, NUMBER 1 173 

Hornellia Walker, new synonymy 

Hornellia Walker, 1904:268 {Hornellia incerta Walker, 1904, monotypy). — Le- 

doyer, 1973:29. 
Metaceradocus Chevreux, 1925:304 {Metaceradocus perdentatus Chevreux, 

1925, monotypy) [new synonym]. 
Tulearogammarus Ledoyer, 1967:129 (Tulearogammarus peresi Ledoyer, 1967, 

original designation, = Hornellia incerta Walker). 

Body ordinary, pleon and urosome dorsally crenulate transversely on posterior 
segmental margins, urosomites free, often with dorsal articulate spines. Rostrum 
small, lateral cephalic lobes rounded-quadrate, sinus present. Eyes present. 

Antennae elongate, extending subequally or antenna 2 longer, ratio of pedun- 
cular articles on antenna 1 = 20:13:7, primary flagellum longer than peduncle, 
accessory flagellum 2-5 articulate. Antenna 2 ordinary. 

Labrum broader than long, notched (type) or entire. Mandibular incisor 
toothed, molar triturative, ratio of palp articles = 4:11:10 or 4:18:15, article 3 
linear (type) or falcate, setae = (AB)DE. Inner lobes of labium present. Inner 
plate of maxilla 1 ovate, apically and part to all medially setose, outer plate with 
11 spines, palps [?symmetrical]. Inner plate of maxilla 2 with oblique facial row 
of setae. Outer plate of maxilliped medially spinose, palp article 3 unlobed, dactyl 
[?shorter than 3, with nail]. 

Coxae ordinary, poorly setose, coxa 1 apically expanded, coxa 4 lobate. 
Gnathopods feeble, slender, scarcely subchelate, wrists elongate, lobed (type) or 
unlobed, hands ovate to rectangular, hand of gnathopod 2 elongate, palms 
oblique, weak, poorly defined. 

Pereopods 3^ ordinary. Article 2 of pereopods 5-7 slightly expanded, poorly 
setose, scarcely lobate. 

Pleopods ordinary, rami of uropods 1-2 marginally spinose, only outer of uro- 
pod 2 shortened, uropod 1 with basofacial spine [in several species, type 
unknown]. Uropod 3 extended, magniramous, aequiramous, peduncle slightly 
elongate, rami elongate, lanceolate, setose, outer 1-2 articulate. Telson elongate, 
deeply cleft, lobes tapering, notched (type), weakly spinose apically and medially 
and dorsally (type) and laterally. 

Coxal gills 2-7, ovate. Oostegites slightly broadened to slender. 

Variants. — Uropod 3 without article 2 on outer ramus {perdentata, occiden- 
talis, micramphopus , vesentiniae), telson fully cleft {occidentalis, micrampho- 
pus): gnathopod 2 very linear {micramphopus)\ Metaceradocus definable as sub- 
genus with micramphopus, occidentalis, perdentata and vesentiniae: telson fully 
cleft, article 2 of outer ramus on uropod 3 absent. 

Relationship. — Differing from Elasmopoides and Maeropsis and Ceradocus in 
the feeble gnathopod 2 of both sexes; somewhat ancestral to Maerella and Jer- 
barnia but anterior coxae normal, not reduced or diversified and maxillipedal 
dactyl well developed; differing from the Eriopisella group in the magniramous 
uropod 3 and well setose maxillae. 

Species. — incerta Walker, 1904 (= peresi Ledoyer, 1967) (Ruffo 1969); mi- 
cramphopus (Stebbing, 1910); occidentalis (J. L. Barnard, 1959); perdentata 
(Chevreux, 1925); vesentinae (Ruffo, 1969); whakatane (J. L. Barnard, 1972b); 
marine, warm-temperate and tropics, shallow water. 



174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Lupimaera, new genus 

Type-species. — Maera lupana J. L. Barnard, 1969a. 

Body slender, urosomites free, naked. Rostrum obsolescent, lateral cephalic 
lobes mammilliform. Antennae medium to short, antenna 1 scarcely longer than 
2, ratio of peduncular articles = 16:8:3, primary flagellum not longer than pedun- 
cular article 1, with 5 articles, accessory flagellum 3-articulate, more than half as 
long as primary flagellum. Antenna 2 also short, flagellum 4-articulate, not longer 
than article 5 of peduncle. Ratio of mandibular palp articles = 2:7:5, article 3 
linear, setae = DE. Inner lobes of labium present. Maxillae not setose medially, 
inner plate of maxilla 1 rectilinear, with 3 apical setae, outer plate with 9 spines, 
palps symmetrical. Inner plate of maxilla 2 with 2 medial and 1 facial setae. 
Outer plate of maxilliped mediafly spinose, palp article 3 unlobed, dactyl shorter 
than 3, very short, unguiform, with nail. 

Coxae of ordinary dimensions, poorly setose, coxa 1 weakly expanded and 
lobed anteroventrally, coxa 4 not lobate, coxa 5 as long as 4. Gnathopods diverse, 
not sexually dimorphic, gnathopod 1 small, wrist elongate, unlobed, hand sub- 
rectangular, palm short, slightly oblique; gnathopod 2 enlarged, wrist short, 
lobed, hand large, subrectangular, palm oblique, short, sculptured. 

Pereopods 3^ ordinary. Article 2 of pereopods 5-7 weakly expanded, weakly 
lobate, posterior margins poorly setose, weakly convex; article 2 of pereopod 5 
not longer than coxa 5 (distinction from Maera); pereopods short. 

Outer rami of uropods 1-2 slightly shortened, all rami marginally densely spi- 
nose, spines small, peduncle of uropod 1 with 1-2 basofacial spines. Uropod 3 
not extended, very short, magniramous, almost aequiramous, rami short, scarcely 
longer than peduncle, spinose, spines short (distinction from Maera). Telson 
short, almost fully cleft, lobes tapering, apices weakly spinose, notched. 

Coxal gills 2-6, ovate. Oostegites [?narrow]. 

Relationship. — Differing from Maera, Paraweckelia, and Meximaera in the 
small article 2 of pereopods 5-7, article 2 of pereopod 5 not being longer than 
coxa 5; rami of uropods 1-3 with spines ah shortened; article 2 of antenna 1 only 
half as long as article 1 . More than 55 species of Maera do not conform to this 
group of distinguishing characters noted for Lupimaera. Distinguished from Ce- 
radocus, which it closely resembles, in the poorly setose maxillae, short uropod 
3 and short article 2 of antenna 1 ; from Ceradomoera in the symmetrical gnath- 
opod 2 and smooth pleon; from Maeropsis in the poorly setose maxilla 2; from 
Ifalukia in the normally long article 2 of mandibular palp; from Paraceradocus 
in the poorly setose maxilla 2 and linear mandibular palp; from Anelasmopus by 
the poorly setose maxillae; and from Ceradocoides in the deeply cleft telson and 
poorly setose maxillae. 

Species. — lupana (J. L. Barnard, 1969a); California, intertidal. 

Etymology. — Name composed of "lupana" and ''Maera,'' a related genus. 
Feminine. 

Lupimaera lupana (J. L. Barnard), new combination 

Fig. 1 

Maera lupana J. L. Barnard, 1969a: 122, figure 20. 

Added description. — Right and left incisors each with 3 teeth; right lacinia 
mobilis bifid and complex (see illustration), left 5-dentate; right rakers 6, left 7; 



VOLUME 95, NUMBER 1 



175 





Fig. 1. Lupimaera lupana (J. L. Barnard), male "h" 3.95 mm: A, Lower lip; B, Maxilla 2; C, 
Maxilla 1; D, Left incisor and lacinia mobilis; E, Maxilliped; F, Right mandible. 



molars of medium size, weakly triturative, each with long seta; palps alike on 
either side, article 1 short, article 2 with 3 inner subbasal setae, article 3 shorter 
than 2, weakly curved but not considered falciform, of linear form but short, 
setae = D2E3. Lower lip with fleshy inner lobes bearing sublobes; outer lobes 
normal. Inner plate of maxilla 1 bluntly subconical, apex with 2-3 setae; outer 
plate with 9 normal spines; palps symmetrical, 2-articulate; article 1 slightly elon- 
gate, article 2 truncate, with 6 apical setae and 2 medial setules. Plates of maxilla 
2 extending equally, outer broader than inner, latter with 2 thin medial setae, one 
facial seta near apex assumed to represent facial row. Plates of maxillipeds normal 
(illustrated), though article 2 elongate, palp smaller than usual in family, dactyl 
short, scarcely unguiform, with large nail. 

Coxal gills on segments 2-6 ovate. Oostegites unknown. 

Material. — J. L. Barnard Station 41, California, Goleta, 6 July 1961, 3 m, rhi- 
zomes of Macrocystis pyrifera, male ''h" 3.95 mm (mouthparts illustrated). 

Remarks. — Now that this species is segregated in its own genus the mouthparts 
are herein illustrated; they are not significantly distinct from those of Maera as 



176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

originally implied. The generic distinctions from Maera lie in the posterior per- 
eopods (see above). 
Distribution. — California, Goleta, in Macrocystis rhizomes, 3 m. 

Maleriopa, new genus 

Type-species. — Eriopisella dentifera Ledoyer, 1978. 

Body slender, urosomites [?free, naked]. Rostrum small, lateral cephalic lobes 
weak, rounded, sinus absent. Eyes present. 

Antennae moderately extended, ratio of peduncular articles = 26: 19:8, primary 
flagellum slightly longer than peduncle, accessory flagellum 2-articulate. Antenna 

2 ordinary. 

Labrum [?entire, rounded]. Mandibular incisor toothed, right mandible with 3 
rakers, molar weakly triturative, ratio of palp articles = 3:10:9, article 3 linear, 
setae = E, sparse. Inner lobes of labium present, fleshy, mandibular lobes ex- 
tended. Maxillae not setose medially, inner plate of maxilla 1 subrectangular, 
with 2 distal setae, outer plate with about 7 spines, palps 2-articulate, 
[?symmetrical]. Inner plate of maxilla 2 without facial and medial setae. Outer 
plate of maxilliped [?medially spinose, article 3 of palp unlobed, dactyl shorter 
than 3, unguiform, with nail]. 

Coxae of ordinary length, poorly setose, coxa 1 tapering, coxa 4 [?unlobed]. 
Gnathopods small to moderate, gnathopod 1 with elongate wrist, hand subrect- 
angular, palm weakly oblique; gnathopod 2 moderately enlarged, wrist short, 
strongly lobed, lobe bending distad, hand subrectangular, weakly expanded api- 
cally, palm oblique, well defined, strongly spinose. 

Pereopods 3^ ordinary. Article 2 of pereopods 5-6 weakly expanded, mod- 
erately lobate, posterior margin straight, of pereopod 7 expanded, lobate, pos- 
terior margin convex, all posterior margins weakly setose; dactyls ordinary. 

Pleopods [?ordinary]. Rami of uropods 1-2 extending equally, marginally spi- 
nose except for outer ramus of uropod 1, latter [?with basofacial spine]. Uropod 

3 slightly extended, parviramous, article 2 on outer ramus short. Telson short, 
deeply cleft, lobes broad but weakly tapering, each apex with subapical spine. 

Coxal gills [?2-6, ovate]. Oostegites [?narrow]. 

Relationship. — Differing from Eriopisa, Victoriopisa and Psammogammarus 
in the loss of medial setae on the maxillae; from Tegano in the presence of article 
3 on the mandibular palp, the short article 2 of antenna 1 and the loss of medial 
setae on maxilla 2; from Eriopisella in the slightly enlarged gnathopod 2 with 
strong, spinose palm. 

The type-species is assumed to have a direct origin distinct from Paraniphargus 
as reflected in the tapering coxa 1, presence of eyes, large lobe on wrist of 
gnathopod 2, more strongly lobate and larger article 2 of pereopods 5-7, presence 
of article 2 on the outer ramus of uropod 3 and the loss of marginal spines on the 
outer ramus of uropod 3 . 

Species. — dentifera (Ledoyer, 1978); Mauritius, sublittoral, 1 species. 

Etymology . — Name contrived from parts of "Mauritius" and ''Eriopisa'' with 
an "ell" thrown in between for euphony. Feminine. 

Tegano, new genus 
Type-species. — Melita seticornis Bousfield, 1970. 



VOLUME 95, NUMBER 1 177 

Body smooth, urosomite 2 with small middorsal posterior mucronation. Ros- 
trum obsolescent, lateral cephalic lobes strongly mammilliform. 

Antennae elongate, slightly thickened, antenna 1 scarcely longer than 2, ratio 
of peduncular articles = 12:15:5, ratio of flagella = 30:3, accessory flagellum 2- 
articulate. Antenna 2 gland cone large. 

Ratio of mandibular palp articles = 4:8:0 or 3:10:0 (article 3 absent), apical 
setae of article 2 = E (one long only). Inner lobes of labium well developed, 
fleshy. Maxillae moderately setose medially, inner plate of maxilla 1 ovate, with 
6 apical and medial setae, outer plate with "9-10" (but see figure) spines, palps 
[?asymmetric]. Inner plate of maxilla 2 with several marginal medial setae, non- 
facial setae. Outer plate of maxilliped minutely spinose medially, dactyl with tiny 
nail. 

Coxae of ordinary length, poorly setose, coxa 1 undilate, coxa 4 unlobed. 
Gnathopods 1-2 diverse, gnathopod 1 the smaller, of Melitid form, wrist weakly 
elongate, hand subrectangular or trapezoidally expanded apically, palm trans- 
verse, convex in female, acquiring rugose process in terminal males, gnathopod 
2 slightly enlarged, wrist of medium length, scarcely lobed (broadly), hand elon- 
gate, subrectangular in female, with oblique short palm, in male larger, more 
ovate, palm longer, softly excavate, dactyl longer and more curved. 

Article 2 of pereopods 5-7 scarcely expanded, scarcely lobate or not postero- 
ventrally, poorly setose (only setulose). 

Rami of uropods 1-2 extending equally, marginally spinose, peduncle of uropod 
1 with basofacial spine. Uropod 3 extended, parviramous, outer ramus elongate, 
with medium article 2. Telson short, cleft to base, lobes leaf-like, tapering api- 
cally, sparsely setose apically. 

Coxal gills 2-6 ovate, that on pereonite 2 pediculate. Oostegites narrow. 

Description. — Antennae of male with whorls of setae on peduncles and flagella. 

Relationship. — Differing from Melita in the reduced mandibular palp. 

Species. — seticornis (Bousfield, 1970); Solomon Islands (Rennell) and Bis- 
marck Archipelago (Mussau), anchialine. 

Etymology. — Named for Lake Tegano, Rennell Island, type-locality of the 
type-species. Masculine. 

Hadziids and Weckeliids 

A Weckeliid lacks eyes and has aequiramous uropod 3. A Hadziid lacks om- 
matidial eyes though occasionally has ocular pigment but uropod 3 is disparira- 
mous. Hadziids further differ from neighboring blind taxa in the elongate uropod 
3, cleft telson and loss of inner lobes on the lower lip. 

The two groups at times have been considered congruent but there remains the 
possibility that two or more ancestries occur, the Weckeliids from a Paraweck- 
elia-Ceradocus ancestry and the Hadziids from a Melitid ancestry. As shown in 
the Key to the Hadziids and Weckeliids, the groups are so contrived that elevation 
to family level is impossible. 

Key to Hadziids and Weckeliids 

1 . Outer ramus of uropod 3 with 2 articles (Hadziids) 2 

- Outer ramus of uropod 3 with 1 article (Weckeliids) 10 



178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

2. Inner plate of maxilla 1 lacking medial setae, inner plate of maxilla 2 
lacking oblique facial row of setae, medial margin setose or not 3 

- Inner plates of maxillae 1-2 strongly setose medially, maxilla 2 with 
oblique facial row of setae 4 

3. Uropod 3 magniramous Paraweckelia (twice) 

- Uropod 3 parviramous Psammoniphargus 

4. Coxa 4 with large posterodistal lobe Saliweckelia 

- Coxa 4 unlobed 5 

5. Wrists of gnathopods 1-2 unlobed, their setae marginal 6 

- Wrists of gnathopods 1-2 lobed, some of their lateral setae facial 9 

6. Telson shortened (uropod 3 variramous, gnathopods and telsonic spi- 
nation like Metahadzia couplet below) Metaniphargus 

- Telson of ordinary length or elongate 7 

7. Gnathopod 1 lacking medial setal brush on article 5, palm of male gnath- 
opod 2 densely spinose, palm of female gnathopod 2 distinct, weakly 
spinose, telson with medial spines, uropod 3 magniramous . . . Metahadzia 

- Gnathopod 1 with medial setal brush, palm of male gnathopod 2 poorly 
spinose, palm of female gnathopod 2 indistinct, telson lacking medial 
spines, uropod 3 parviramous 8 

8. Palm of male gnathopod 2 densely setose, telson naked laterally 

Dulzura 

- Palm of male gnathopod 2 not densely setose, telson with lateral spines 

Protohadzia 

9. Telson with lateral spination, article 5 of gnathopod 1 elongate 

Liagoceradocus 

- Telson naked laterally, article 5 of gnathopod 1 as long as 6 Hadzia 

10. [In contradistinction to couplet 1, uropod 3 actually with vestigial article 
2 on outer ramus], otherwise gnathopod 2 of both sexes of typical Ce- 
radocid or Melitid form, hand well inflated, palm long, minutely serrate, 
lacking Hadziid setae, wrist very short, strongly lobed, maxilla 2 lacking 
oblique facial row of setae, inner plate of maxilla 1 with setae mostly 
apical Paraweckelia (twice) 

- [Like couplet 1, uropod 3 lacking article 2 on outer ramus], otherwise 
gnathopod 2 not typical of Melitids or Ceradocids, hand either poorly 
inflated or palm poorly defined in either sex or short, or not serrate, or 
bearing Hadziid setae, wrist in either sex relatively elongate, maxillae 
weU setose medially, inner plate of maxilla 2 with oblique row of facial 
setae 11 

1 1 . Coxa 4 with posterodistal lobe 12 

- Coxa 4 without posterodistal lobe 13 

12. Inner lobes of lower lip present, palp of mandible 1 -articulate . . . Weckelia 

- Inner lobes of lower lip absent, palp of mandible 3-articulate 

Alloweckelia 

13. Coxa 1 not enlarged, right lacinia mobilis present 14 

- Coxa 1 much larger than coxa 2, right lacinia mobilis absent 16 

14. Wrist of gnathopods lobate Mexiweckelia 

- Wrist of gnathopods not lobate 15 



VOLUME 95, NUMBER 1 179 

15. Gnathopod 1 not merochelate Mexiweckelia particeps 

- Gnathopod 1 merochelate Mayaweckelia 

16. Mouthparts projecting anteriorly as far as article 2 of antenna 1 17 

- Mouthparts (except maxilliped) projecting less than halfway along article 

1 of antenna 1 18 

17. Dactyl of maxilliped short, article 3 stout, inner plates of maxillae 1-2 
elongate, outer plate of maxilla 1 with 7 spines, inner plate of maxilliped 
ordinary Holsingerius 

- Dactyl of maxilliped elongate, article 3 thin, inner plates of maxillae 1-2 
ordinary, outer plate of maxilla 1 with 14 spines, inner plate of maxilliped 
broadly expanded Texiweckeliopsis 

18. Hand of gnathopod 1 like that of gnathopod 2, longer than wrist, gnatho- 
pods of sexes alike, article 2 of pereopods 3^ strongly expanded 

Allot exiweckelia 

- Hand of gnathopod 1 different from gnathopod 2, shorter than wrist, 
gnathopods of sexes strongly distinct, article 2 of pereopods 3^ thin . . 

Texiweckelia 

Texiweckeliopsis, new genus 

Type-species. — Texiweckelia insolita Holsinger, 1980. 

Body slender, urosomites 1-3 each with 2, 4, and 2 dorsal spines respectively. 
Lateral cephalic lobes truncate, without sinus. Eyes absent. 

Antennae elongate, antenna 1 longer than 2, ratio of peduncular articles = 
23:9:6, ventral margin of article 1 lacking spines. Accessory flagellum vestigial 
or absent (smaller than in other Hadziids). 

Mandibles and maxillae projecting forward as far as article 2 of antenna 1. 
Mandibular right lacinia mobilis absent, palp absent. Labium without inner lobes, 
not gaping. Maxillae medially setose, inner plate of maxilla 1 triangular, fully 
setose medially, outer plate with 14-15 serrate spines, palps [?asymmetrical]. 
Plates of maxilla 2 slightly broadened, short, inner with oblique facial row of 
setae. Inner plate of maxilliped extremely broad, outer plate medially setose; 
article 3 of palp thin, with very small lobe forming weak apical chela, dactyl very 
long, thin, with nail (maxilliped palp thus weakly prehensile). 

Coxae of medium size to short, but coxa 1 larger than coxa 2, coxa 4 unlobed, 
coxa 5 [?shorter than coxa 4]. Gnathopods subchelate, scarcely dimorphic sex- 
ually, feeble; gnathopods 1-2 almost alike, mittenform, wrists elongate, broadly 
lobate, setation facial, hands slightly shorter than wrists, narrow but not perfectly 
linear or rectangular, slightly expanded in middle and then tapering towards palm, 
palms weakly oblique, short, spines tiny or sparse, weakly bifid, Hadziid setae 
few, elongate only on gnathopod 2 and sparse on palm, male gnathopods slightly 
stouter than in female, especially on hands. 

Pereopod 5 slightly shorter than 6-7, article 2 of pereopods 5-7 slightly ex- 
panded, weakly lobate, dactyls with several setules on inferior margin, article 6 
weakly setose or spinose. 

Rami of uropods 1-2 subequally extended, uropod 1 with 2 basofacial spines, 
comb of uropod 2 absent, outer ramus of uropod 2 dorsally naked. Uropod 3 



180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

elongate, magniramous, almost aequiramous, outer ramus 1-articulate, apices of 
rami weakly truncate, with several spines of which one elongate. Telson of or- 
dinary length, cleft about two thirds, each lobe with 2 apical spines, main setules 
M. 90. 

Coxal gills 2-6, ovate, with long stalks, 2-articulate. Oostegites narrow. 

Relationship . — Differing from Texiweckelia in the strong forward projection of 
the mandibles and maxillae, and mittenform gnathopods. See Holsingerius. 

Species. — insolita (Holsinger, 1980), Texas, hypogean, 1 species. 

Etymology. — Name composed of Texiweckelia and "opsis" for "likeness." 
Feminine. 

Holsingerius, new genus 

Type-species. — Texiweckelia samacos Holsinger, 1980. 

Body slender, urosomites 1-3 each with 2, 2 and 4 dorsal spines respectively. 
Rostrum obsolescent, lateral cephalic lobes sharply mammilliform. Eyes absent. 

Antennae elongate, antenna 1 longer than 2, ratio of peduncular articles = 
30:16:9, ventral margin of article 1 lacking spines. Accessory flagellum vestigial 
or absent (smaller than in other Hadziids). 

Mandibles and maxillae projecting forward as far as article 2 of antenna 1. 
Mandibular right lacinia mobilis absent, palp absent. Labium without inner lobes, 
gaping. Maxillae medially setose, inner plate of maxilla 1 enlarged, triangular, 
fully setose medially, outer plate with 7 serrate spines, palps [?asymmetrical]. 
Inner and outer plates of maxilla 2 greatly elongate, inner with oblique facial row 
of setae. Outer plate of maxilliped medially setose; article 3 of palp stout, apically 
expanded and weakly chelate, dactyl of ordinary length, thin, with nail (palp 
scarcely prehensile). 

Coxae of medium size to short, but coxa 1 larger than coxa 2, coxa 4 unlobed, 
coxa 5 [?shorter than coxa 4]. Gnathopods subchelate, sexually dimorphic, those 
of female feeble; wrist of both pairs in both sexes of medium length, strongly 
lobate, with facial setation; hand of female gnathopod 1 shorter than wrist, rect- 
angular, palm short, weakly oblique, minutely spinose, male hand enlarged, palm 
long, strongly oblique, well spinose; hand of female gnathopod 2 as long as wrist 
(thus elongate), thin, rectangular, palm short, weakly oblique, minutely spinose 
and with several Hadziid setae; anterior margin of hand lined with setae; hand 
of male gnathopod 2 enlarged, palm long, rounded-oblique, densely spinose 
(spines weakly bifid), with 2 Hadziid setae, dactyl very long. Article 2 of per- 
eopods 3^ scarcely expanded, poorly spinose. Pereopod 5 shorter than 6-7; 
article 2 of pereopods 5-7 expanded, lobate, dactyls short, with several setules 
on inferior margin, article 6 weakly setose or spinose. 

Rami of uropods 2-3 subequally extended, uropod 1 with several basofacial 
spines, comb of uropod 2 absent. Uropod 3 elongate, magniramous, almost ae- 
quiramous, outer ramus 1-articulate, apices of rami weakly truncate, with several 
spines, of which one elongate. Telson scarcely elongate, cleft almost three- 
fourths, each lobe with 3 apical spines and occasional lateral spine, main setules 
M. 65. 

Coxal gills 2-6, at least some ovate, with long stalks, 2-articulate. Oostegites 
narrow [but full adult unknown]. 



VOLUME 95, NUMBER 1 181 

Relationship. — Differing from Texiweckeliopsis in the ordinary dactyl of the 
maxilHped, ordinary outer plates of maxillae and maxillipeds but the elongate 
inner plates of the maxillae. 

Species. — samacos (Holsinger, 1980), Texas, San Marcos Well, hypogean, 1 
species. 

Etymology. — Named for the eminent authority on freshwater amphipods, Dr. 
John R. Holsinger. 

Paracalliopiidae, new family 

Head, eyes, mouthparts, coxae 1-3, pereopods 3-6, uropods 1-2 ordinary. 
Accessory flagellum vestigial. Coxa 4 poorly excavate posteriorly. Female 
gnathopods feeble, mittenform, male gnathopods larger, gnathopod 2 enlarged, 
wrist small, hand large (usually rotated inward on death), palm oblique. Pereopod 
7 elongate, dactyl elongate and setose. Uropods 1-3 extending equally, peduncle 
of uropod 3 slightly elongate, rami short, equal, lanceolate, outer 1-articulate. 
Telson laminar, entire. Urosomites 2-3 coalesced. 

See Eusiridae, Pontogeneiidae, Calliopiidae, Oedicerotidae. 

Male antennae with calceoli; antenna 2 longer than antenna 1. Setosity of inner 
plates on maxillae variable. Wrists of female gnathopods elongate, weakly to 
strongly lobate posteriorly, lobes rounded and pointing terminad, hands slender, 
palms oblique, short; wrists of male gnathopods not elongate, lobes pointing 
almost perpendicular to axis of appendage, hands of ontogenetic appearance, 
palmar defining corners softly rounded, palms of both gnathopods with long setae, 
palm of gnathopod 2 also with stout spines; gnathopods of both sexes with patches 
of pubescence. Pleon well developed, epimera large. Body not carinate. 

Distinguished from Eusiridae, Pontogeneiidae, Calliopiidae, and Oedicerotidae 
by the fused urosomites. Standing as an evolutionary grade between Calliopiidae 
and Oedicerotidae by mixing the head, body form, and unspecialized mouthparts 
of Calliopiidae with the elongate pereopod 7, elongate dactyl of pereopod 7, 
setose dactyl of pereopod 7, slightly elongate peduncle of uropod 3, and rudiments 
of fossorial adaptations in the slightly increased setosity of pereopods character- 
istic of Oedicerotidae. The fused urosomites, however, show that the group is 
divergent from the main evolutionary track. 

Key to the Genera of Paracalliopiidae 

1. Inner plates of maxillae densely setose medially Paracalliope 

- Inner plates of maxillae not setose medially Indocalliope 

Paracalliope Stebbing 
Paracalliope Stebbing, 1899:210; 1906:279.— J. L. Barnard, 1972b:70. 

Type-species. — Calliope fluviatilis Thomson, 1879 (original designation). 

Inner plates of maxillae 1-2 densely setose medially. Uropods 1-2 normally 
spinose. 

Species. — australis (Haswell, IS^O); fluviatilis Thomson (J. L. Barnard, 1972b); 
karitane J. L. Barnard, 1972b; novaecaledoniae Ruffo and Vesentini-Paiotta, 1972; 



182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

novizealandiae (Dana) (?J. L. Barnard, 1972b); Dubious: fluviatilis of Chilton, 
1921; New Zealand, Australia, middle IndoPacific to India; fresh and brackish 
waters, marine littoral. 

Nomenclatural changes. — Paracalliope fernandoi Wignarajah (1958) belongs 
in the family Talitridae. 

Indocalliope, new genus 

Type-species. — Paracalliope indica K. H. Barnard, 1935. 

Inner plates of maxillae not setose medially. Spines of uropods 1-2 sparse or 
absent. 

Species. — indica K. H. Barnard, 1935 (Nayar, 1959); India, brackish waters, 
1 species. 

Etymology. — Name composed from "India" and ''Calliope.'' Feminine. 

Phoxocephalidae 
Feriharpinia, new genus 

Type-species. — Harpinia ferenteria Gurjanova, 1977. 

Rostrum unconstricted. Eyes absent. Article 2 of antenna 1 short, ventral setae 
widely spread. Article 1 of antenna 2 [?ensiform], article 3 with 3 facial setules, 
facial spines on article 4 in [?1 row, ?all spines thin], article 5 ordinary. Right 
mandibular incisor with [?3 teeth, right lacinia mobilis ?bifid, molar not triturative, 
with ?3+ basally fused splayed spines; ?special spines; ?palpar hump small], apex 
of palp article 3 oblique. Inner plate of maxilla 1 with 1 seta, palp biarticulate. 
Maxillipeds ordinary, apex of palp article 3 not strongly protuberant, dactyl elon- 
gate, apical nail [?distinct]. 

Gnathopods ordinary, small, [?dissimilar, gnathopod 2 ?weakly enlarged], ar- 
ticle 5 of gnathopods 1-2 very short, free, palms oblique, hands ordinary, ova- 
torectangular, poorly setose anteriorly. 

Article 5 of pereopods 3^ [? without posteroproximal setae, article 6 with ?thin 
armaments]. Article 2 of pereopod 5 of [?narrow] form, articles 4-5 of pereopods 
5-6 narrow; pereopod 7 ordinary, article 3 slightly enlarged, dactyl ordinary. 

Epimera 1-2 [?without long facial brushes or posterior setae], epimeron 3 of 
ordinary classification, bearing 4 or more long setae. Urosomite 3 [? without dorsal 
hook]. 

Peduncle of uropod 1 [? without interramal spike, ? without major displaced 
spine, rami of uropods 1-2 ?not continuously spinose to apex, inner ramus of 
uropod 1 with ?1 row of marginal spines]. Inner ramus of uropod 2 [?ordinary]. 
Uropod 3 [?ordinary], one of rami [?longer than peduncle], bearing [?article 2 on 
outer ramus with ?2 apical setae]. Telson ordinary, but slightly elongate. 

Relationship. — Like Harpinia but male armaments distinctive: instead of 
brushes being present on article 1 of antenna 1 and article 1 of flagellum of antenna 
1 and on articles 3-4 of antenna 2, and instead of article 1 of primary flagellum 
on antenna 1 being enlarged and dominant, the male of Feriharpinia has a brush 
of aesthetascs on article 3 of the peduncle of antenna 1 , article 1 of the primary 
flagellum is not grossly enlarged, and article 5 of antenna 2 has a row of large 
dorsal calceoli, not found in Harpinia. 



VOLUME 95, NUMBER 1 183 

Species .^erenteria Gurjanova, 1977; Okhotsk Sea, west Kamchatka, 196-230 
m, 1 species. 

Etymology. — Name composed from "ferenteria" and related genus "Harpinia.'' 
Feminine. 

Proharpinia Schellenberg, (new composition) 

Proharpinia Schellenberg, 1931:80 {Proharpinia antipoda Schellenberg, 1931, 
monotypy). — J. L. Barnard, 1960:311. — Barnard and Drummond, 1978:532. 

Rostrum unconstricted. Eyes present. 

Article 2 of antenna 1 short, ventral setae almost confined apically (by fiat). 
Article 1 of antenna 2 not ensiform, article 3 with 4-5 facial setules, facial spines 
on article 4 in 1 main row, all spines thin, article 5 short. Right mandibular incisor 
with [?3 teeth, right lacinia mobilis ?bifid, ?weakly flabellate. ?molar not tritu- 
rative, with ?3 splayed spines]; palpar hump small, apex of palp article 3 oblique. 
Inner plate of maxilla 1 naked, palp biarticulate. Maxillipeds ordinary, apex of 
palp article 3 not strongly protuberant, dactyl body not elongate, but apical nail 
distinct and elongate. 

Gnathopods small, scarcely dissimilar, gnathopod 2 weakly to moderately en- 
larged, article 5 of gnathopods 1-2 very short, free on gnathopod 1. cryptic on 
gnathopod 2, palms oblique, hands of gnathopods 1-2 ovatorectangular, elongate, 
poorly setose anteriorly. 

Article 5 of pereopods 3-4 with posteroproximal setae, article 6 with thin 
armaments. Article 2 of pereopod 5 of narrow form, articles 4-5 of pereopods 
5-6 medium to narrow; pereopod 7 ordinary, but article 3 enlarged, dactyl ordi- 
nary. 

Epimera 1-2 without long facial brushes or posterior setae, epimeron 3 of 
ordinary classification, bearing 3 or more long setae. Urosomite 3 without dorsal 
hook. 

Peduncle of uropod 1 without interramal spike, without major displaced spine, 
rami of uropods 1-2 continuously spinose to apex, with subapical spines or nails, 
inner ramus of uropod 1 with 1 row of marginal spines. Inner ramus of uropod 
2 ordinary. Uropod 3 ordinary, 1 of rami longer than peduncle, bearing article 2 
on outer ramus, with vestigial or no apical setae. Telson ordinary. 

Variants. — Proharpinia stephenseni, the second species of the genus, is re- 
tained here only provisionally; it differs from the type in the presence of setae on 
the inner plate of the maxilliped and the absence of apical spination on the rami 
of uropods 1-2. However it matches the type in the telson. and loss of strong 
setation on article 2 on the outer ramus of uropod 3. 

Relationship . — Differing from Heterophoxus in the absence of an ensiform pro- 
cess on antenna 2; more plesiomorphic than Torridoharpinia in spination of uro- 
pods 1-2 and normal condition of outer ramus of uropod 1 (not shortened), and 
normal armament of telson but more apomorphic in loss of setae on inner plate 
of maxilla 1 . Proharpinia hurleyi and P. tropicana are removed to Torridohar- 
pinia below. 

Species. — See J. L. Barnard, 1960; antipoda Schellenberg, 1931; stephenseni 
Schellenberg, 1931; Magellan area, Falkland Islands. 2-274 m. 



184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Torridoharpinia, new genus 

Type species. — Proharpinia hurleyi J. L. Barnard, 1958, 1960. 

Rostrum unconstricted. Eyes present. 

Article 2 of antenna 1 short, ventral setae ventrally spread or almost confined 
apically. Article 1 of antenna 2 not ensiform, article 3 with 4 facial setules, facial 
spines on article 4 in 1 main row, all spines thin, article 5 short. Right mandibular 
incisor with 3 teeth, right lacinia mobilis bifid, flabellate, molar not triturative, 
with 2 splayed spines; palpar hump small, apex of palp article 3 oblique. Inner 
plate of maxilla 1 with 2-4 (type) setae, palp biarticulate. Maxillipeds ordinary, 
apex of palp article 3 not strongly protuberant, dactyl elongate, apical nail dis- 
tinct. 

Gnathopods small, dissimilar, gnathopod 2 weakly to moderately enlarged, 
article 5 of gnathopods 1-2 short, free on gnathopod 1, cryptic on gnathopod 2, 
palms oblique, hands ovatorectangular, broadened on gnathopod 2, poorly setose 
anteriorly. 

Article 5 of pereopods 3^ with posteroproximal setae, article 6 with thin ar- 
maments. Article 2 of pereopod 5 of narrow form, articles 4-5 of pereopods 5-6 
medium to narrow; pereopod 7 ordinary, article 3 enlarged, dactyl ordinary. 

Epimera 1-2 without long facial brushes or posterior setae, epimeron 3 of 
ordinary classification, bearing 3 or more long setae. Urosomite 3 without dorsal 
hook. 

Peduncle of uropod 1 without interramal spike, without major displaced spine, 
rami of uropods 1-2 not continuously spinose to apex, inner ramus of uropod 1 
only in male with 2 rows of marginal spines. Inner ramus of uropod 2 ordinary. 
Uropod 3 ordinary, one of rami longer than peduncle, bearing article 2 on outer 
ramus, with 1 or 2 (type) apical setae. Telson ordinary, but 1 apical element stout 
(contrast Proharpinia). 

Relationship . — Differing from Proharpinia in the lack of ramal spines on female 
uropod 2, the shortened outer ramus of uropod 1, the presence of 1-2 long apical 
setae on article 2 of the outer ramus on uropod 3, the presence of a stout spine 
on each lobe of the telson and the presence of setae on the inner plate of maxilla 
1. Proharpinia has only thin setae or setules on the telson. 

Species. — hurleyi J. L. Barnard, 1958, 1960 (=stephenseni of Hurley, 1954, not 
Schellenberg, 1931); tropicana J. L. Barnard, 1960; New Zealand, Auckland Is- 
lands, Campbell Island, Galapagos Islands, 0-46 m, 2 species. 

Etymology . — Name composed from "torrid" and ''Harpinia.'' Feminine. 

Pseudamphilochidae Schellenberg, 1931 

Only coxa 4 slightly broadened, coxae 2^ with continuous margins overlap- 
ping, not rabbeted, coxae 1-2 not hidden, coxa 1 not reduced, no anterior coxa 
hidden. Telson cleft. 

Upper lip weakly excavate but unnotched. Mandibular molar vestigial or ab- 
sent. Lower lip with well developed inner lobes. 

Palp of maxilliped lacking process on article 3. 

Outer ramus of uropod 1 shortened. Peduncle of uropod 3 not greatly elongate, 
rami almost twice as long as peduncle. 



VOLUME 95, NUMBER 1 185 

Remarks. — This family has not been properly heralded in the literature before 
and is brought to light from a provisional proposal (Schellenberg, 1931: 92). 

Pseudamphilochus Schellenberg 

Pseudamphilochus Schellenberg, 1931:92. 

Type-species. — Pseudamphilochus shoemakeri Schellenberg, 1931 (monotypy). 
With familial characters. 

Gnathopods of medium size, wrists short, weakly lobate, not carpochelate, 
hands broad, scarcely 1.4 times as long as broad, palms subtrans verse. 
Species. — shoemakeri Schellenberg, 1931; South Georgia, littoral. 

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PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 188-193 

HEMIODOPSIS OCELLATA, A NEW HEMIODONTID 

CHARACOID FISH (PISCES: CHARACOIDEA) 

FROM WESTERN SURINAM 

Richard P. Vari 

Abstract. — Hemiodopsis ocellata, an open water species occuring in the black 
water drainages of western Surinam, is described from several localities within 
the Corantijn River system basin. The pattern of zig-zag lines on the dorsal por- 
tion of the body appears to be unique in the genus. 



Introduction 

The new species described herein, Hemiodopsis ocellata, is widespread in the 
black waters of the Corantijn River system of western Surinam. Habitats of in- 
dividuals captured or observed in the wild show that during low water levels this 
species is primarily an inhabitant of the main river channels and larger open pools 
and streams rather than the shaded creeks and smaller streams of the rain forest. 
During high water levels, in contrast, H. ocellata apparently travels up smaller 
tributaries into the flooded rain forest to feed and reproduce. The species has 
been taken most often in the more readily sampled still or slowly flowing waters. 
Nonetheless, the capture of a single specimen in the rapidly-flowing waters of 
Dalbana Creek and observations over several seasons indicate that H. ocellata 
also occurs in waters with high flow velocities. This species is usually found in 
association with the much more abundant Hemiodopsis goeldii, which it closely 
resembles in overall body form, maximum adult size, and pigmentation pattern 
(see Bohlke, 1955, fig. 7). 

Hemiodopsis ocellata, new species 
Fig. 1, Table 1 

Holotype. — National Museum of Natural History (USNM) 221175, 170.0 mm 
standard length (SL), collected by R. P. Vari, 7 December 1979 in the main stream 
of Dalbana Creek, approximately 150 m upstream of its junction with the Ka- 
balebo River, Nickerie District, Surinam (approx. 4°47'N, 57°29'W). 

Paratypes.—l specimen, USNM 225592, 141.5 mm SL, collected by R. P. Vari 
and L. R. Parenti, 8 September 1980, in a slow-flowing side channel of the Cor- 
antijn River about 180 km from its mouth, Nickerie District, Surinam (approx. 
5°08'N, 57°18'W); 2 specimens, USNM 225593, 104.7-113.0 mm SL [1 cleared 
and counterstained for cartilage and bone] coUected by R. P. Vari and L. R. 
Parenti, 17 September 1980, in a still pool near "Camp Hydro" on an island in 
the middle of the Corantijn River, Nickerie District, Surinam (approx. 4°22'N, 
57°58'W); 4 specimens, USNM 225594, 129.0-142.0 mm SL, coUected by H. M. 
Madarie, 15 May 1980, in a small creek and the surrounding flooded rain forest 
of the Corantijn River, Nickerie District, Surinam (approx. 5°32'N, 57°10'W). 

Diagnosis. — Within the family Hemiodontidae the new species is assignable to 



VOLUME 95, NUMBER 1 



189 




Fig. 1. Hemiodopsis ocellata, new species, holotype, USNM 221175, 170.0 mm SL. 



Hemiodopsis based on the very slightly protractile jaws (in contrast to the highly 
protractile jaws of Bivibranchia and Argonectes), multicuspidate dentition limited 
to the upper jaw (in contrast to unicuspidate teeth in both jaws in Michromis- 
chodus), 3. dorsal fin of moderate height (in contrast to an elongate, anteriorly 
filamentous dorsal fin in Pterhemiodus), and a moderate vertical gradation in 
scale size (in contrast to a pronounced vertical gradation in scale size in He- 
miodus). Within Hemiodopsis (see Gery 1963 for a discussion of the redefinition 
of the genus) the 88-96 lateral line scales occurring in H. ocellata readily distin- 
guish it from the majority of the species in the genus {H. gracilis, H. goeldii, H. 
fowleri, H. thayeri, H. semitaeniata, H. ternetzi, H. immaculata, H. rodolphoi, 
and H. parnaguae) which as a unit demonstrate a range from 42 to 83 lateral line 
scales, and from H. microlepis and H. argentea which have a higher number of 



Table 1. — Morphometries oi Hemiodopsis ocellata, new species. Standard length is expressed in 
mm; measurements 1 to 1 1 are percentages of standard length; 12 to 15 percentages of head length. 





Holotype 


Paratypes (7) 






Range 


Average 


Standard length 


170.0 


113.0-141.5 


129.6 


1 . Greatest body depth 


29.1 


27.4-29.8 


29.1 


2. Snout to dorsal-fin origin 


49.2 


42.8-54.0 


49.3 


3. Snout to anal-fin origin 


80.9 


80.7-82.9 


81.6 


4. Snout to pelvic-fin origin 


52.8 


52.9-54.0 


53.4 


5. Snout to anus 


78.0 


77.0-77.9 


78.1 


6. Origin of rayed dorsal to hypural joint 


56.0 


53.9-56.3 


55.0 


7. Least depth of caudal peduncle 


10.1 


9.1-11.2 


10.0 


8. Pectoral-fin length 


20.0 


19.0-21.2 


19.7 


9. Pelvic-fin length 


21.8 


21.3-24.1 


22.6 


10. Dorsal-fin height 


26.4 


25.1-30.2 


27.7 


1 1 . Head length 


26.1 


25.0-26.4 


25.6 


12. Orbital diameter 


30.0 


29.0-32.4 


30.1 


13. Snout length 


31.0 


29.2-34.2 


31.6 


14. Postorbital length 


38.4 


37.3-40.0 


38.8 


15. Interorbital width 


36.8 


34.9-38.1 


36.1 



190 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Map of the lower Corantijn River basin region, Surinam and Guyana, showing collecting 
localities for the following specimens of Hemiodopsis ocellata (dotted line depicts the road to Camp 
Matapi and Camp Amotopo): 1, collection locality of holotype, USNM 221175; 2, collection locality 
of 2 paratypes, USNM 225593; 3, collection locality of 1 paratype, USNM 225592; 4, collection 
locality of 4 paratypes, USNM 225594. 



VOLUME 95, NUMBER 1 191 

pored lateral line scales (110 to 112 and 120 to 125 respectively). The only He- 
miodopsis species having a lateral line scale count approximating that of H. 
ocellata is H. parnaguae of the upper Parnaiba River system (80-83 lateral line 
scales; Eigenmann and Henn 1916:87). However, in addition to the differences 
in the number of pored lateral line scales, these species are distinguishable by 
the hyaline anal fin of H. ocellata which contrasts with the presence of a distal 
black band on the fin in H. parnaguae (see Eigenmann and Henn 1916, PI. 17). 
Furthermore, H. parnaguae lacks the distinctive dorsal zig-zag patterning char- 
acteristic of H. ocellata. 

Description. — Table 1 gives morphometries of the holotype and paratypes. 
Body relatively slender, slightly compressed laterally. Greatest body depth in 
region of origin of rayed dorsal fin. Dorsal profile of body gently curved from tip 
of snout to interorbital region, slightly convex or straight from that area to origin 
of rayed dorsal fin. Body profile at base of rayed dorsal fin straight, posteroven- 
trally sloped. Dorsal profile of body nearly straight from rear of insertion of rayed 
dorsal fin to upper margin of caudal peduncle. Ventral profile of head smoothly 
convex from tip of lower lip to below base of pectoral fin; nearly straight from 
that point to origin of anal fin. 

Head relatively small, snout obtuse with mouth subinferior. Anterior margin 
of lower jaw cresent-shaped. Upper jaw moderate, very slightly protractile, pos- 
terior margin of maxilla extends slightly posterior of a vertical through posterior 
border of rear nostril. Nostrils approximate, anterior opening round, posterior 
cresent-shaped. Eye relatively large. An extensive, horizontally ovoid "adipose 
eyelid" (a thick transparent connective tissue layer) extends from posterior mar- 
gin of rear nostril to middle of opercle (less developed posteriorly in smaller 
specimens). Adipose eyelid with an ovoid, vertically-elongate opening overlying 
pupil. Fronto-parietal fontanel extensive, extending into rear of ethmoid. Parietals 
completely separated, frontals in contact only at epiphyseal bar. 4 branchiostegal 
rays on each side; 1 on posterior ceratohyal, 3 on anterior ceratohyal. Osteolog- 
ical characters are overall very similar to those of Hemiodus (=Hemiodopsis) 
semitaeniata as illustrated by Roberts (1974). Vertebrae 39 or 40 including those 
of Weberian apparatus and counting fused PUj + Ui as a single element. 

Lower jaw edentulous, anterior edge sectorial. Functional teeth in a single 
series in upper jaw. All teeth multicuspidate; number of cusps on each tooth 
decreasing from 1 1 on medial teeth on 7 on lateral teeth. Sixteen to 18 teeth along 
each side of upper jaw, 10 maxillary and 8 premaxillary teeth on each side of jaw 
in cleared and stained specimen. Teeth extending along entire anterior edge of 
maxilla (see Roberts, 1974, fig. 9 for comparable situation in Hemiodopsis sem- 
itaeniata). One or two rows of partially formed replacement teeth internal to 
functional tooth row. Premaxillary replacement tooth rows embedded in flesh on 
inner surface of premaxilla; maxillary replacement teeth arranged in a shallow 
maxillary replacement-tooth trench. Dermopalatine and ectopterygoid edentu- 
lous. Ceratobranchial 5 bearing a relatively narrow band of posteriorly-directed 
teeth along medial and posteromedial borders. Upper pharyngeal tooth-plates 4 
and 5 with bands of small, conic teeth. Ceratobranchials 2, 3, and 4 and hypo- 
branchial 3 bearing ventrally-directed processes extending lateral to ventral aorta. 
Gill-rakers expanded distally, fan-shaped, with a series of pointed, digitiform 
processes along distal edges. Number and size of distal gill-raker processes in- 



192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

creasing towards ceratobranchial-epibranchial joint. Gill-rakers on first gill arch 
38 on epibranchial and 50 on ceratobranchial in cleared and counterstained spec- 
imen. 

Scales cycloid, thin. Pored lateral line scales between supracleithrum and hy- 
pural joint 93 in holotype (88 in 1 paratype, 92 in 1 paratype, 93 in 2 paratypes, 
95 in 2 paratypes, 96 in 1 paratype). Five to 7 pored lateral line scales extending 
beyond hypural joint onto caudal fin. Scales above lateral line in a transverse 
series to origin of rayed dorsal fin 22 in holotype (22 in 3 paratypes, questionably 
21 in 4 partially descaled paratypes). Scales below lateral line in a transverse 
series to origin of anal fin 14 in holotype (12 in 1 paratype, 13 in 1 paratype, 14 
in 2 paratypes, 15 in 1 paratype, 2 paratypes descaled in region). Body squamation 
extending onto base of caudal rays. Low sheath of scales along base of rayed 
dorsal fin and anterior portion of anal fin. Axillary process of pelvic fin composed 
of a series of scales. Size of body scales only slightly graduated in a vertical 
series. 

Rayed dorsal fin pointed but not filamentous anteriorly; second unbranched 
and first branched rays longest, subequal. Dorsal-fin rays ii,9 in all specimens. 
Adipose dorsal fin moderate, unsealed; length about two-thirds diameter of orbit. 
Anal fin falcate, anterior branched rays more than twice as long as posteriormost 
branched rays. Anal-fin rays iii-9 in holotype (ii,9-i in 1 paratype, iii-9 in 2 para- 
types, ii-10 in 4 paratypes), first 2 unbranched anal rays very short. Pectoral fin 
pointed, reaching two-thirds distance to a vertical through insertion of pelvic fin. 
Pectoral-fin rays i-16-ii in holotype (i-16-ii in 3 paratypes, i-16-iii in 2 paratypes, 
i-17-ii in 2 paratypes). Pelvic fin pointed, reaching three-quarters distance to anus. 
Pelvic-fin rays i-10 in holotype (i-10 in 6 paratypes, i-11 in 1 paratype). 

Coloration in alcohol. — Overall coloration silvery. Head dark dorsally, a much 
more intensely pigmented spot about size of pupil located on anterodorsal corner 
of opercle and surrounding region. Fleshy lip of premaxilla and maxilla dark. 
Body darker dorsal to lateral line, silvery-white ventrally. A diffuse dark lateral 
band slopes slightly posteroventrally from dorsal portion of opercle to base of 
caudal fin. Body band interrupted by an intensely pigmented, longitudinally elon- 
gate, ocellated spot. Spot straddling but located largely dorsal to lateral line, 
extending from slightly behind a vertical through rear of dorsal-fin insertion nearly 
to a vertical through posterior limit of pelvic fin; spot extending about 12 scales 
longitudinally and 7 vertically. Unpigmented region surrounding spot approxi- 
mately 5 or 6 scales wide. Dorsal portion of body above lateral line marked by 
31 to 35 dark, strongly-angled bars having the form of a "V" on its side with 
angle of flexure directed posteriorly. Each bar about one and one-half or two 
scales wide. Pigmentation pattern less apparent anteriorly and posteriorly. A 
series of randomly arranged spots on sides of body below lateral line, each spot 
about size of exposed portion of scales in that region. Rayed dorsal fin dusky, 
with no apparent pattern. Adipose dorsal fin dusky. Anal fin clear. Pectoral fin 
with narrow bands of light brown chromatophores outlining unbranched lateral 
fin ray and first branched ray. Anal fin with series of pale brown chromatophores 
outlining distal portions of rays on second through fourth branched anal rays. 
Lower lobe of caudal fin with a dark stripe continuous with lateral band on body. 
Procurrent rays and first and second principal rays of lower caudal lobes unpig- 
mented. Procurrent rays of upper caudal lobe outlined by dark chromatophores. 



VOLUME 95, NUMBER 1 193 

Coloration in life. — Overall coloration silvery golden. Iris intense gold. Silvery 
coloration somewhat masking lateral body band and dorsal bars. Fins pinkish or 
light red. 

Relationships. — Our present understanding of relationships within Hemiodop- 
sis are quite poor. Indeed, no characters supporting an hypothesis of monophyly 
of the genus have been proposed. Within Hemiodopsis as presently delimited, H. 
ocellata is most similar to although necessarily most closely related to H. par- 
naguae. 

Etymology . — Ocellata, from the Latin for little eyes, in reference to the lateral 
eye-like body spot. 

Acknowledgments 

The specimens that served as the basis for this paper were collected during 
surveys carried out as a part of preimpoundment studies associated with the 
Kabalebo Hydroelectric Project. The assistance of Drs. M. P. Panday-Verheuvel 
and S. Niekoop is gratefully acknowledged as is field assistance by Dr. L. R. 
Parenti, Mr. H. M. Madarie, Ms. S. Engel, and Mr. S. Silos. This manuscript 
benefited from critiques by L. R. Parenti and S. H. Weitzman. Figure 1 is by 
Susan L. Jewett. 

Literature Cited 

Bohlke, J. 1955. Studies on fishes of the family Characidae. — No. 10. Notes on the coloration of the 
species of Hemiodus, Pterhemiodus, and Anisitsia, with the description of a new Hemiodus 
from the Rio Negro at the Brazil-Colombia border. — Notulae Naturae 278:1-15. 

Eigenmann, C. H., and A. W. Henn. 1916. Description of three new species of characid fishes. — 
Annals of the Carnegie Museum 10:87-90. 

Gery, J. 1963. Sur la nomenclature et la systematique du genre Hemiodus Miiller (Pisces, Chara- 
coidei). — Bulletin Museum National d'Histoire Naturelle, Series 2, 35:589-605. 

Roberts, T. R. 1974. Osteology and classification of the Neotropical characoid fishes of the families 
Hemiodontidae (including Anodontinae) and Parodontidae. — Bulletin of the Museum of Com- 
parative Zoology 146:41 1-472. 

Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 194-197 

EVIDENCE OF ONTOGENETIC SETAL CHANGES 
IN HETEROMASTUS FILIFORMIS 
(POLYCHAETA: CAPITELLIDAE) 

Thomas J. Fredette 

Abstract. — Heteromastus filiformis undergoes ontogenetic setal replacement in 
the fourth and fifth setiger. Setigers 4 and 5 initially bear hooded hooks which 
are gradually lost and replaced by capillary setae. 



The possible occurrence of ontogenetic setal changes in the polychaete Het- 
eromastus filiformis was first observed in samples taken during a colonization 
experiment in the lower York River, Virginia (Fredette 1980). The first sample 
set (0.25 mm sieve), taken shortly after deployment of azoic substrates, contained 
newly set juveniles of several different species of benthos, including a capitellid 
polychaete. The general body morphology of these newly set capitellids was 
similar to that of the 2 locally common species, Heteromastus filiformis and 
Mediomastus ambiseta. However, the specific characters fit neither. The juve- 
niles had capillary setae on the first 3 setigers while the remaining setigers bore 
hooded hooks. This is quite unlike H. filiformis which has the first 5 anterior 
setigers with capillary setae and M. ambiseta which has capillary setae in the 
first 4 anterior setigers and the last several notopodia. Specific taxonomic place- 
ment of these juvenile capitellids did not seem possible. 



60-1 



3 CAPILLARY SETIGERS 



4 CAPILLARY SETIGERS 




5 CAPILLARY SETIGERS 



10 15 



TIME IN DAYS 
Fig. 1. Progression of setal changes observed in a population of H. filiformis colonizing azoic 
substrates in the York River, Virginia. Volume of area indicates percent of population exhibiting the 
indicated character. 



VOLUME 95, NUMBER 1 



195 




Fig. 2. SEM micrographs of juvenile capitellids: A, Anterior end of a juvenile capitellid with both 
capillary and hooded hook setae on fifth setiger; B, Mixed setal bundle of a juvenile capitellid. Setal 
bundle in background bears all capillary setae. 



Successive samplings taken during the colonization process revealed serial 
changes in setal pattern in the population of newly set capitellids (Fig. 1). The 
individuals from these later samples constituted a continuum from the 3-capillary 
setiger stage to a 5-setiger stage. The continuum was complete with intermediate 
stages (individuals with setal bundles containing both capillary setae and hooded 
hooks) (Fig. 2) and a gradation of sizes. The graph shown in Figure 1 is based on 
the assumption that all the juveniles are H. filiformis. However, individuals that 
were obviously M. ambiseta also occurred in the samples from day 73 and day 
144. The large time gaps in the sampling series made a final conclusion on the 
specific placement of these juveniles difficult. 

Additional documentation of this phenomenon was obtained by examining sam- 
ples from an ongoing benthic study in the lower York River in which weekly 
samples are being taken (Diaz, unpublished data). It was felt that these samples 
would clarify the progression of setal changes and help to better isolate the ap- 



196 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



100- 



% 50- 



lOOn 



7o 50- 



lOOi 
% 50 

lOO-i 
% 50- 



lOO-i 
% 50- 



I I 



21 APRIL 
n = 2 



28 APRIL 
n--0 



I ' •- 







5 MAY 
n = 7 












.1. ■ 


1 c^n 



13 MAY 
n= 25 



. rn . rr i 



19 MAY 
n = 28 







28 MAY 


■ 






n = 60 


'— 1 — ' 




1 . ' . ' . 1 





I i I 



I ' I ■ f 



I " * 



2 JUNE 
n = 6l 



9 JUNE 
n = 94 



16 JUNE 
n= 132 



23 JUNE 
n = 73 



ABCDE F ABCDE F 

Fig. 3. Weekly distribution of setal pattern in a field population of capitellids. A. 3 anterior 
capillary setigers; B, 4th setiger with both capillary and hooded hook setae; C, 4 anterior capillary 
setigers (no posterior capillaries); D, 5th setiger with both capillary and hooded hook setae; E, 5 
anterior capillary setigers (H. filiformis pattern); F, 4 anterior capillary setigers, posterior capillaries 
(M. ambiseta pattern). Areas indicated are percents of total population. 



pearance of M. ambiseta and H. filiformis in the benthic community. Individuals 
in these samples were placed into 6 classes: (A) 3 anterior capillary setigers, (B) 
fourth setiger with mixed capillaries and hooded hooks (no posterior capillaries), 
(C) 4 anterior capillary setigers (no posterior capillaries), (D) fifth setiger with 
mixed capillaries and hooded hooks, (E) H. filiformis setal pattern, (F) M. am- 
biseta setal pattern. 

The occurrence of M. ambiseta in these samples was sporadic, in low densities 
and seemed to bear no relationship to the development of the juvenile capitellids 
(Fig. 3). Also, a few small individuals of M. ambiseta were found that were 
similar in size to the other juvenile capitellids, indicating that M. ambiseta may 
have the same setal pattern in juveniles and adults. 

The other five classes appear in a pattern similar to the age-class distributions 
one expects from a population with synchronous breeding and recruitment (Fig. 
3). Recruitment occurs in early May and the population quickly passes through 
the various setal changes until 7 weeks later when all the individuals exhibit the 
typical H. filiformis pattern. 



VOLUME 95, NUMBER 1 197 

Table 1. — Width measurement (fjLm) of fifth setiger, means or ranges. 

Class Width 

3 capillary setigers 146.2, n = 4 
4th mixed 185.6, n = 9 

4 capillary setigers 208.7, n = 7 
5th mixed 317.2, n = 16 
H. filiformis (adults) 300-600 
juvenile M. ambiseta 140-200 



To illustrate the progression of sizes that occur, several individuals from each 
class were measured by recording the width of the fifth setiger. The juvenile M. 
ambiseta were also measured for comparison (Table 1). As indicated earlier these 
individuals are similar in size to the 3-capillary setiger individuals. The juvenile 
capitellids showed a steady progression of sizes and change to a 5-capillary setal 
pattern. 

That H. filiformis undergoes an ontogenetic development from a 3 anterior 
capillary stage to a 5 anterior capillary stage is strongly indicated by the evidence 
presented. Setigers 4 and 5 initially bear hooded hooks which are gradually lost 
and replaced by capillary setae. Mediomastus ambiseta appears not to undergo 
such a morphological change. Further substantiation of the observed setal 
changes could be facilitated by examination of cultured populations of these two 
capitellid species. 

The separation of M. ambiseta and H. filiformis in benthic samples (0.5 mm 
sieve) by anterior capillary setiger counts alone (see keys of Ewing and Dauer 
1981, Fauchald 1977) is not sufficient. Examination for posterior notopodial cap- 
illaries is necessary. 

Acknowledgments 

I would like to express my thanks to Dr. Robert J. Diaz for the loan of several 
samples and critical review of this manuscript, Dr. Frank O. Perkins for advice 
on SEM techniques and preparations, and Michael J. Kravitz and Stephanie A. 
Vay for valuable suggestions and discussion. 

Contribution No. 1039 from the Virginia Institute of Marine Science, The Col- 
lege of William and Mary. 

Literature Cited 

Ewing, R. M., and D. M. Dauer. 1981. A new species of Amastigos (Polychaeta: Capitellidae) from 
the Chesapeake Bay and Atlantic coast of the United States with notes on the Capitellidae of 
the Chesapeake Bay. — Proceedings of the Biological Society of Washington 94:163-168. 

Fauchald, K. 1977. The polychaete worms, definitions and keys to the orders, families and genera. — 
Natural History Museum Los Angeles County, Science Series 28:1-190. 

Fredette, T. J. 1980. Macrobenthic colonization of muddy and sandy substrates in the York River, 
Virginia. M.S. thesis. The College of William and Mary, Williamsburg, Virginia, 62 pp. 

Department of Invertebrate Ecology, Virginia Institute of Marine Science and 
School of Marine Science, The College of William and Mary, Gloucester Point, 
Virginia 23062. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 198-202 

A NEW SPECIES OF OPHIOGOMPHUS (INSECTA: 

ODONATA: GOMPHIDAE) FROM THE WESTERN 

HIGHLAND RIM IN TENNESSEE 

Jerry A. Louton 

Abstract. — Ophiogomphus bouchardi, a new species of Gomphidae, is de- 
scribed from the Western Highland Rim of central Tennessee. The description 
includes both sexes and the exuviae of the holotype male. Affinities of the new 
species apparently lie with O. carolinus and O. mainensis. 



While collecting crayfish on the Western Highland Rim in Dickson Co., Ten- 
nessee, in 1971, Dr. Raymond W. Bouchard collected a single female nymph of 
Ophiogomphus which he subsequently succeeded in rearing. The reared female 
and exuviae were placed in the University of Tennessee collection of aquatic 
insects (as O. mainensis) and there remained until examined by me in 1976. The 
female and nymphal skin were at once seen to be distinct from other species of 
the genus by several outstanding characters. I now have a series of seventeen 
reared adults to support the original supposition of distinctness. Dr. Bouchard 
has kindly consented to the inclusion of his 1971 collection as part of the type 
series. It is with great pleasure that I name this new species in recognition of Dr. 
Bouchard's many contributions to aquatic biology. 

Ophiogomphus bouchardi, new species 
Figs. 1-13 

Material— RoXoiy^Q male, TENNESSEE, Dickson Co., Will Hall Creek at 
U.S. Hwy. 70 (entrance to Montgomery Bell State Park), nymph collected 21 
Mar 1981, adult emerged 13 Apr 1981; allotype female, same location and date, 
adult emerged 14 Apr 1981; paratypes, same location and date, 5 males and 3 
females (all reared); same location, 1 female, nymph collected 26 Mar 1971, date 
adult emerged unknown. Additional material not in type series, same location 
and date, 3 reared males and 2 reared females; same location, 25 Sep 1980, 5 final 
instar nymphs; Lewis Co., Little Swan Creek at Natchez Trace Parkway, 20 Mar 
1979, 1 final instar nymph. All types will be placed in the Florida State Collection 
of Arthropods, Gainesville, Florida. 

Diagnosis. — Dorsal stripes of synthorax absent or vestigial; face not cross- 
striped with black at sutures; yellow of tibiae restricted to one-fifth length of 
tibiae; lateral process of male epiproct not strongly projected; anterior occipital 
horns of females erect, separated by a distance equal to their height; posterior 
occipital horns of females vestigial; prementum of nymphs narrowed anteriorly, 
ligula narrow and strongly projected. 

Description of holotype male. — Total length 47.4 mm, abdomen 34.1 mm, hind 
wing 28.2 mm. Form and coloration generally typical for eastern North American 
species. 



VOLUME 95, NUMBER 1 



199 




Figs. 1-13. Ophiogomphus bouchardi, n. sp.: 1, Dorsum of synthorax of holotype male; 2, Lateral 
view of synthorax of holotype male; 3-5, Extremes in form of occipital horns of females; 6, Anterior 
hamule of holotype male; 7, Posterior hamule of holotype male; 8, Penis and vesicle of holotype 
male; 9, Lateral view of terminal abdominal appendages of holotype male; 10, Dorsal view of terminal 
abdominal appendages of holotype male; 11, Right antenna of exuviae of holotype male; 12, Ventral 
view of subgenital plate of allotype female; 13, Prementum of exuviae of holotype male with inset of 
detail of ligula. 



200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Anterior of frons yellow-green without cross stripes at sutures, dorsum nar- 
rowly infuscated with brown between antennal bases, six black granules antero- 
laterally, entire frons covered with black hairs. Vertex blackish across anterior 
half, yellowish at hairy prominences, brownish on posterior half. Occiput yellow 
with dense line of black hairs at crest, scattered black hairs on posterior surface. 

Prothorax with two broad longitudinal black bands on dorsum. Synthorax light- 
ly striped with brown; brown stripes adjacent to dorsal crest vestigial, only a 
faint brown streak adjacent to middle prominence of crest (Fig. 1); second (an- 
tehumeral) stripe of thorax present, continuous with brown of katepisternum 
below, upper end not connected to brown of antealar ridge; stripe of second 
plural suture (humeral) complete, wider in upper half and constricted somewhat 
just as it joins antealar ridge; middle suture with faint stripe ending at spiracle; 
stripe of third pleural suture absent except for faint streak on upper one-third 
(Fig. 2). Femora greenish; streaked with black that is apically coalesced; tibia 
and tarsi all black except for small narrow proximal spot of green on outer surface 
of tibia. 

Abdominal segments with dorsal yellow spots, wide on segments two and three, 
narrowing to segment eight, wider on segments nine and ten; in lateral view 
abdominal segments dark in upper one-half, gray-green below with black en- 
croaching down into light areas along supplementary transverse carinae and as 
dark postero-lateral spots; light areas of segments seven to ten tinged with yellow; 
auricles yellow-green with brownish crescent behind. Male genitalia black-tipped, 
vesicle shining black; anterior hamule bifid, tip claw-like (Fig. 6); posterior ham- 
ule tapering, bent anteriorly forming smooth curve (Fig. 7); penis with moderate 
flagella (Fig. 8). Terminal abdominal appendages yellow; superior appendages 
taper to sharp tip, in dorsal view bowed in middle and proximate at bases and 
tips (Fig. 10), in lateral view appendages arched in middle (Fig. 9), black denticles 
scattered on ventral surface near tips; inferior appendage bifid to one-half its 
length, lateral prominences are low tubercles placed near midlength of appendage. 

Description of allotype female. — Total length 46.7 mm, abdomen 31.9 mm, 
hind wing 30.2 mm. Similar in general form and coloration to holotype male 
except: occiput with pair of large erect sharp-tipped horns separated by distance 
equal to their length (Fig. 5) and surrounded by black hairs; no vestige of brown 
stripes adjacent to dorsal crest of synthorax; posterolateral black spots on ab- 
dominal segments continue anteriorward as streaks on middle abdominal seg- 
ments; vulvar lamina as long as ninth abdominal sternite (Fig. 12). 

Exuviae of holotype male. — Total length 26.3 mm. Exuviae light brown with 
coarse pigmented cuticular granules. 

Antennae broad and flat, two and one-fourth times longer than wide (Fig. 11). 
Prementum narrowed anteriorly; palpal lobes short, two-fifths length of premen- 
tum and bearing only ten teeth; ligula narrow, less than one-third width of pre- 
mentum at widest point and bearing 18 teeth and a dense brush of piliform setae 
on anterior margin (Fig. 13). Lateral spines present on abdominal segments seven 
to nine, dorsal hooks well developed on segments two to nine, ante-apical tu- 
bercles of epiproct located at three-fifths of distance from base to tip. 

Variations. — Adult males: total lengths 45.4-50.2 (avg. 48.8) mm, abdominal 
lengths 31.4-35.3 (avg. 34.1) mm, hind wing lengths 27.0-28.6 (avg. 28.0) mm. 



VOLUME 95, NUMBER 1 201 

Adultfemales: total lengths 47.5-51.0 (avg. 48.7) mm, abdominal lengths 33.9-35.8 
(avg. 34.7) mm, hind wing lengths 25.8-31.2 (avg. 29.3) mm. Exuviae: total 
length (both sexes) 24.0-28.0 (avg. 26.4) mm. Above measurements from pre- 
served material. Live final instar nymphs 23-24 mm. 

The form and coloration of all adult specimens is generally uniform except that 
the brown wash adjacent to the dorsal crest of the synthorax is often absent. The 
occiput of females displays some variation in shape of the posterior outline viewed 
dorsally often showing a slight convexity or concavity that may be artif actual. 
The occipital horns display great variability in thickness and position (see Figs. 
3-5). In addition, there are often adventitious spinules isolated some distance 
from the main horns (Fig. 3), as a row between the horns and/or as lateral out- 
growths of the large paired horns (Fig. 4). 

Relationships. — Relationships among species of Ophiogomphus have not been 
discussed. Species in eastern North America are morphologically distinctive and 
have highly sympatric distribution areas. Closely related species pairs are un- 
common in Ophiogomphus unlike many other genera and subgenera of Gomphi- 
dae (Louton 1981). Three species of the genus share structural similarities that 
seem to indicate a close relationship. Ophiogomphus bouchardi, O. carolinus, 
and O. mainensis share: 1. similarly structured anterior occipital horns of the 
female (though those of O. mainensis are adjacent), 2. rudimentary posterior 
occipital horns and 3. terminal abdominal appendages of similar structure (though 
exaggerated in O. mainensis). The close relationship of these three species is 
further suggested by the allopatric nature of their distributional areas. 

The simple matrix below facilitates the separation of the three above mentioned 
similar species. 

O. bouchardi O. carolinus O. mainensis 

Yellow tibial stripe no^ yes no 

Middorsal thoracic stripe no yes yes 

^ Proximal yellow spot about one-fifth length of tibia. 

Nymphal habitat. — Nymphs of Ophiogomphus bouchardi were found in cherty 
limestone gravel pockets in a small second order tributary (Will Hall Creek) of 
the Harpeth River (Cumberland River System), The stream is predominantly 
bedrock with gravel and sand deposits limited to fissures and pockets in pool 
areas. Nymphs were also located in sand trapped by the roots of sedges growing 
where bedrock fissures intersected the streambanks. The most common odonate 
associates in this habitat were Stylogomphus albistylus, Boyeria vinosa and, less 
commonly, Gomphus lividus. All three associates are widespread lotic general- 
ists. 

Acknowledgments 

I would like to thank Christine Eason Louton and John L. Harris for assistance 
in collecting nymphal material, Leah R. Baker for typing the manuscript, and Dr. 
David A. Etnier for critically reviewing the manuscript. 



202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Literature Cited 

Louton, J. A. 198 L Lotic dragonfly (Anisoptera: Odonata) larvae of the southeastern United States: 
identification, distribution, and a consideration of the historical biogeography of the groups. 
Unpublished dissertation, University of Tennessee, Knoxville. 

Department of Zoology, University of Tennessee, Knoxville, Tennessee 37916. 



Note added to galley proof: After this paper had gone to press, Dr. Minter 
Westfall (University of Florida, Gainesville) informed me that the forthcoming 
(December, 1981) issue of Odonatologica contained a description of a new spe- 
cies of Ophiogomphus that would be conspecific with the one that I was de- 
scribing (Carle, F. L. 1981. A new species of Ophiogomphus from eastern North 
America, with a key to the regional species. Odonatologica 10(4): 27 1-278). 
Mr. Carle submitted his paper October 14, 1981, with the knowledge that I had 
submitted a description of this same species a month earlier. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 203-209 

TWO NEW SPECIES OF ONUPHIS (ONUPHIDAE: 
POLYCHAETA) FROM URUGUAY 

Kristian Fauchald 

Abstract. — Two new species of Onuphis {Onuphis) from Uruguay, O. (O.) 
difficilis and O. (O.) orensanzi are described, based on material identified as 
Onuphis setosa Kinberg by Orensanz (1974). 



Kinberg (1865) described several species of Onuphis from the southern Atlantic 
Ocean collected during the EUGENIE expedition. The material of most of these 
species is in Riksmuseet, Stockholm, Sweden, with the exception of the material 
of Onuphis setosa which has been lost (Roy Oleroed, in litt.). 

The specimens of O. setosa must have been lost before 1930 since Augener 
(1931:297) remarked that they were absent when he reviewed Kinberg's collection 
of onuphids from the La Plata region. Orensanz (1974:88) applied the name to a 
species found in the type-area. Some discrepancies between Kinberg's (1865, 
1910) description and illustrations and those made by Orensanz suggested that it 
was necessary to review Orensanz' material. Three samples were made available 
by the Director, M.N.H.N., Montevideo, Uruguay. As suspected, these speci- 
mens cannot belong to Onuphis setosa Kinberg and are described below as two 
new species. 

The generic definitions and terminology are as in Fauchald (1980). 

Onuphis (Onuphis) difficilis, new species 
Fig. 1 , Table 1 

Onuphis setosa. — Orensanz, 1974:89, in part (not Kinberg, 1865:560). 

Material.— Ai\2iX\i\c Ocean off Uruguay. 34°5rS, 54°04.5'W, 39 m depth, 14 
April 1965, coll. A. Knipovich, st. 263 (holotype, M.N.H.N., Montevideo, I 1382, 
1 paratype, M.N.H.N., 1 paratype, USNM 69917). 

Description. — The holotype is an incomplete specimen with 50 setigers that is 
16.5 mm long and 1.7 mm wide with parapodia. It is pale buff-colored and lacks 
distinct color patterns. Eyes are absent. 

The prostomium (Fig. la) is a rounded lobe; the frontal palps are short and 
triangular and the short peristomial cirri barely reach the middle of the prosto- 
mium. The outer lateral occipital antennae reach setiger 1, the inner lateral an- 
tennae reach setiger 7, and the median antenna reaches setiger 4. The occipital 
ceratophores have up to 7 distinct rings on the lower half. Each ring is very 
narrow and the rings are crowded near the base of the antennae making them 
difficult to observe. 

Branchiae are first present from setiger 6; all branchiae are branched; the first 
branchia has 3 filaments; where best developed, posterior to setiger 21, each 
branchia has 6 filaments. In one of the paratypes (USNM 69917), which consists 
of 200 setigers, the last 50 setigers have reduced branchiae with only 3 or 4 fila- 
ments. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




O.'jmm c,f 



0.002 mm b,d,e 



1 mm 



Fig. L Onuphis (Onuphis) difficilis holotype, N.M.H.N., Montevideo, I 1382: a, Anterior end, 
dorsal view; b, Pectinate seta, median parapodium; c. Third parapodium, anterior view; d. Large 
hook, third parapodium; e, Pseudocompound hook, third parapodium; f. Eighth parapodium, anterior 
view; g, Anterior end, lateral view. 



Ventral cirri (Fig. Ig) are cirriform in the first 8 setigers and postsetal lobes are 
distinctly digitiform in the first 17 setigers in the holotype and in the first 11 or 
23 setigers in the 2 paratypes. The first parapodia are no longer than those in the 
second or third setiger (Fig. Ic). In each of these parapodia the acicular lobe is 
distally rounded, the presetal lobe follows the outline of the acicular lobe closely 
except on the lower side where it is cut away from the base of the setae. A 
distinct contraction fold is present across the middle of the frontal face. The 
ventral cirrus and postsetal lobes are both triangular and of the same size. The 
dorsal cirrus is more distinctly digitiform and twice as long as the ventral cirrus. 
The ventral cirri and postsetal lobes become increasingly tapered posteriorly (Fig. 



VOLUME 95, NUMBER 1 205 

If). The dorsal cirrus remains of about the same length in all setigers but becomes 
increasingly slender in posterior setigers. 

Limbate and pectinate setae, pseudocompound, large and subacicular hooks 
are present. Limbate setae are present in all setigers but are especially numerous 
in the anterior third of the body. In the first 4 or 5 setigers they are short, slender, 
and limited to the superior part of the fascicles. Each of the median and posterior 
parapodia has 2 pectinate setae; each (Fig. lb) is distinctly oblique distally and 
has about 14 or 15 coarse teeth. The distal edges of the pectinate setae are curved. 
Tridentate pseudocompound hooks (Fig. le) are present in the first 5 setigers; the 
distal tooth is extremely long and slender and projects well beyond the edge of 
the hood and the median and proximal teeth are short and slender. Large, tri- 
dentate hooks (Fig. Id) are present in setigers 3-8; each has a long, relatively 
slender distal tooth, the median tooth is blunt and conical, and the proximal tooth 
is slender and narrow. Bidentate subacicular hooks are first present in setiger 18 
in the holotype and in setigers 15 and 18 in the 2 paratypes. 

The maxillary formula (observed in the paratype in USNM 69917) is 1 + 1, 9+9, 
11+0, 9+8, and 1 + 1. 

Fragments of a ragged tube were present in the vial with the 3 specimens. The 
inner lining of the tube appears to be soft, but with considerable tensile strength; 
it is covered externally with loose debris in no recognizable pattern. 

Kinberg's illustrations and descriptions of Onuphis setosa show a species with 
cirriform ventral cirri on less than 5 setigers, branchiae starting between setigers 
10 and 20. The pseudocompound hooks are distally shown as bidentate and with 
short, blunt hoods. Orensanz' material from Uruguay have ventral cirri in at least 
6 setigers, branchiae are present from setiger 6 and the pseudocompound hooks 
are tridentate. The Uruguayan specimens resemble O. setosa basically in having 
the median occipital antenna clearly shorter than the inner lateral ones, but this 
is certainly not a unique character for this species. 

Onuphis difficilis and O. orensanzi (see below) both resemble the same complex 
of species, including O.fragilis Kinberg (1865:561; see also Fauchald 1980:808), 
O. pulchra Fauchald (1980:814), O. simoni Santos, Day and Rice (1981:663), O. 
vermillionensis Fauchald (1968:41) and O. virgata Fauchald (1980:819) in that all 
these species have exclusively tridentate pseudocompound hooks, branched 
branchiae starting at setigers 5-10 and ventral cirri present in at least 5 setigers. 
A comparison of these species is given in Table 1 . 

Etymology. — The problems in differentiating species in this group prompted 
the name. 

Distribution. — Onuphis difficilis is known from one locality in shelf depths off 
Uruguay in the southwestern Atlantic Ocean. 



Onuphis (Onuphis) orensanzi, new species 
Fig. 2, Tables 1 and 2 

Onuphis setosa. — Orensanz, 1974:94, in part (not Kinberg, 1865:560). 

Material.— AWdiniic Ocean off Uruguay, 34°5rS, 52°35'W, 83 m, 12 April 1965, 
coll. A. Knipovich, st. 250 (holotype, M.N.H.N., Montevideo, I 1380, 129 para- 
types, M.N.H.N., USNM 69918). Atlantic Ocean off Uruguay, 35°20.8'S, 



206 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table I. — Comparison of selected species of Onuphis (Onuphis). The columns are: 1, Number of 
rings in the occipital ceratophores; 2, Number of segments with cirriform ventral cirri; 3, Number of 
segments with pseudocompound hooks; 4, Last segment with large hooks; 5, First branchia present; 
6, Maximal number of branchial filaments; 7, Number of teeth in the pectinate setae; 8, First segment 
with subacicular hooks. 



Name 


\ 


2 


3 


4 


5 


6 


7 


8 


difficilis 


7-8 


7-9 


5 


8 


6 


6 


15 


18 


fragilis 





7 


7 


12 


6 


2 


? 


25 


orensanzi 


3-4 


5 


4 


6 


6 


3 


10 


13 


pulchra 


4-5 


9 


6 


15-19 


6 


4 


50 


16-20 


simoni 


3 


6-8 


5 


16 


6-9 


7-8 


16-19 


17 


vermillionensis 


4-5 


9 


8 


10 


7 


4 


14 


12 


virgata 


4-5 


11-13 


7 


21-22 


5-7 


5 


8 


22-23 



52°50.4'W, 130 m, 14 April 1965, coll. A. Knipovich, st. 260 (15 paratypes, 
M.N. H.N. , Montevideo, I 1383). 

Description. — The holotype is a complete specimen with 153 setigers that is 
33.15 mm long and 2.50 mm wide with parapodia. The body is cylindrical and 
pale pink in color; distinct color patterns are absent. The prostomium is a short 
rounded lobe with the occipital antennae attached medially. The frontal palps are 
short and slender. At the posterior margin of the prostomium is a small, distinct 
tubercle. The outer lateral occipital antennae reach setiger 2, the inner lateral 
antennae reach setiger 5 and the median antenna reaches setiger 3. The inner 
lateral ceratophores have up to 4 basal rings. The peristomial antennae are short 
and slender and do not reach beyond the base of the occipital antennae. 

Branchiae are present from setiger 6 to setiger 92 in the holotype. The first 15 
and the last 30 pairs of branchiae are single filaments; all other branchiae have 
either 2 or 3 filaments where best developed. 

The first parapodia are all similar in size; the acicular lobes are rounded and 
the presetal lobe follows the outline of the acicular lobe closely. The postsetal 
lobes and the ventral cirri are of the same length in the first parapodium (distorted 
in Fig. 2a), in the second and third parapodia the ventral cirri are distinctly longer 
than the postsetal lobe. The ventral cirri are digitiform; the postsetal lobes are 
triangular. The dorsal cirri are longer than the ventral cirri in all setigers; each 
is digitiform but becomes slender and thread-like in posterior setigers. Ventral 
cirri are cirriform in the first 5 setigers in all specimens. Postsetal lobes are 
digitiform in the first 10 setigers. 

Limbate and pectinate setae, pseudocompound, large and subacicular hooks 
are present. Limbate setae are present in all setigers, but are especially common 
in the first third of the body. Distally oblique pectinate setae with about 10 teeth 
each are present in median and posterior setigers. Usually 2 pectinate setae are 
present in a parapodium. Tridentate pseudocompound hooks with short blunt 
hoods are present in the first 3-5 setigers; the distal tooth is only slightly longer 
than the median and proximal teeth and does not project beyond the hood. Large 
hooks are present from setiger 3 to about 6; each large hook is tridentate with 2 
large, strongly curved teeth basally and a short, slender tooth distally. The distal 



VOLUME 95, NUMBER 1 



207 




. a,g 



Fig. 2. Onuphis (Onuphis) orensanzi: a, c-e and g, holotype M.N.H.N., Montevideo, I 1380, b and 
f, paratype USNM 69918. a, Anterior end, lateral view; b, Seventh parapodium, anterior view; c. 
Pectinate seta, median parapodium; d. Large hook, third parapodium; e, Pseudocompound hook, 
third parapodium; f, Third parapodium, anterior view; g. Anterior end, dorsal view. 



tooth is often broken, but the base of the tooth can always be recognized. Bi- 
dentate subacicular hooks are present from setiger 12 or 13. 
The maxillary formula (investigated in 3 paratypes from USNM 69918) is 1 + 1, 

5 + 5, 7+0, 4+6-7, and 1 + 1. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2. — Survey statistics for Onuphis (Onuphis) orensanzi. 



Range 



Mean 



s.d. 



Occipital antennae 

Outer lateral reach number 
Inner lateral reach number 
Median reaches number 
Number of rings 

Branchiae 

From setiger number 

To setiger number 

Number of filaments 
Cirriform ventral cirri to setiger num- 
ber 
Digitiform postsetal lobes to setiger 

number 
Pseudocompound hooks to setiger 

number 
Number teeth in pseudocompound hooks 
Subacicular hooks first present from 

number 
Volume through setiger 10 

Mean length per setiger (x 100): 18.10 

Mean length per setiger (x 100), st. 250: 16.12 
Mean length per setiger (x 100), st. 260: 22.74 



1-2 


1.03 


0.16 


0.03 


39 


3-9 


5.23 


1.03 


1.06 


35 


2-4 


3.19 


0.66 


0.44 


37 


3-4 


3.78 


0.42 


0.18 


40 


6 




invariant 






76-92 


87.00 


5.60 


31.33 


7 


1-3 


2.28 


0.60 


0.36 


40 


5 




invariant 






8-10 


9.83 


0.50 


0.25 


40 


3-5 


3.90 


0.44 


0.19 


40 


3 




invariant 






10-14 


12.53 


0.82 


0.67 


40 


0.03-2.68 


1.05 


0.54 


0.29 


40 



Tubes are slender, with a thin inner Hning and thin cover of sediment particles. 

All specimens from station 260 were investigated in detail, as were 25 of the 
130 specimens from station 250. The remaining 105 specimens were only cursorily 
examined, but do not appear to differ in the distribution of ventral cirri, branchiae, 
postsetal lobes, or in the number of rings on the ceratophores. 

The differentiation of O. orensanzi and related species is indicated in Table 1. 

Etymology. — The species is named in honor of the Argentinian polychaetologist 
Jose Maria Orensanz whose papers on the southwest Atlantic fauna have added 
considerably to our knowledge of the polychaetes. 

Distribution. — O. orensanzi is known from two localities off the coast of Uru- 
guay and may be widespread in the southwest Atlantic Ocean. 



Acknowledgments 

I would like to thank Dr. Jose Maria Orensanz for his efforts in arranging the 
loan of the specimens housed at the Museo Nacional de Historia Natural, Mon- 
tevideo, and the Director of that Institution, Dr. Miguel A. Klappenbach for 
allowing me the loan of the material. My colleague, Dr. Meredith L. Jones, kindly 
read and commented on the manuscript. 

Literature Cited 

Augener, H. 193 1 . Die bodensassigen Polychaeten nebst einer Hirudinee der Meteor-Fahrt. — Mittei- 
lungen aus dem Hamburgischen Zoologische Staatsinstitut und Museum 44:279-313. 



VOLUME 95, NUMBER 1 209 

Fauchald, K. 1968. Onuphidae (Polychaeta) from western Mexico. — Allan Hancock Monographs in 
Marine Biology 3:1-82. 

. 1980. Onuphidae (Polychaeta) from Belize, Central America, with notes on related taxa. — 

Proceedings of the Biological Society of Washington 93(3):797-829. 

Kinberg, J. G. H. 1865. Annulata nova. — Oefversigt af Kungliga Vetenskaps-Akademiens For- 
handlingar, Stockholm, 1864 (vol. 21):559-574. 

. 1910. Annulater. In: Kungliga Svenska Fregatten Eugenie's Resa omkring jorden under befal 

af C. A. Virgin aren 1851-1853. — Vetenskapliga lagttagelser pa Konung Oscar den Forstes 
Befallning utgifna delen. Zoologi 3:33-78 (issued posthumously by Hajalmar Theel). 

Orensanz, J. M. 1974. Los anelidos pohquetos de la provincia biogeografica Argentina V. Onuph- 
idae. — Physis, Buenos Aires, Seccion A 33(86):75-122. 

Santos, S. L., R. Day, and S. A. Rice. 1981. Onuphis simoni, a new species of polychaete (Poly- 
chaeta: Onuphidae) from south Florida. — Proceedings of the Biological Society of Washington 
94(3):663-668. 

Department of Invertebrate Zoology, Smithsonian Institution, Washington, 
D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(1), 1982, pp. 210-221 

REMARKS ON THE STYLASTERINE FAUNA OF THE 

WEST INDIES, WITH THE DESCRIPTION OF 

STYLASTER ANTILLARUM, A NEW SPECIES 

FROM THE LESSER ANTILLES 

(CNIDARIA: HYDROZOA: 

STYLASTERINA) 

Helmut Zibrowius and Stephen D. Cairns 

Abstract. — Stylaster antillarum, a new species of stylasterine coral from the 
Lesser Antilles, is described and illustrated. The 23 species of Stylasterina now 
known from the West Indies are listed and briefly discussed. Inaccurate distri- 
butional patterns of some of these species, usually resulting from incorrect iden- 
tifications, are noted. 



Introduction 

Most of the information on the Stylasterina of the West Indies (between Flor- 
ida, Central America and South America), comprising descriptions of species, 
nomenclatural changes, additional records and inventories, can be found in the 
following publications: Pallas (1766), Duchassaing and Michelotti (1861, 1864), 
Pourtales (1867, 1868, 1871, 1874, 1878), Saville Kent (1870, 1871), Lindstrom 
(1877), Moseley (1879, 1881), Broch (1936, 1942), Boschma (1951, 1955, 1957, 
1962, 1964a, 1964b, 1964c, 1965), Squires (1962), Roos (1971), Vervoort & Zi- 
browius (1981) and Zibrowius (in press). 

Until now, stylasterine corals have been reported from the West Indies under 
28 specific and one subspecific names which are listed below. Six of these species 
and the one subspecies are considered to be junior synonyms or dubious records, 
resulting in 23 valid species records (including the new species) for the area. The 
invalid and dubious species records are marked by an asterisk. If not otherwise 
indicated, the original descriptions of these species are based on material from the 
West Indies. 

Stylaster complanatus Pourtales, 1867 

Stylaster duchassaingi Pourtales, 1867 

(new name for Stylaster elegans Duchassaing & Michelotti, 1864) 

Stylaster echinatus Broch, 1936 

Stylaster antillarum, new species 

^Stylaster elegans Duchassaing & Michelotti, 1864 

Gunior homonym of Stylaster elegans Verrill, 1864, a species from the Marshall Islands, Pa- 
cific; renamed S. duchassaingi by Pourtales, 1867) 

Stylaster erubescens Pourtales, 1871 

*Stylaster eximius Saville Kent, 1871 

(new name for Stylaster elegans Duchassaing & Michelotti, 1864; junior synonym of S. du- 
schassaingi) 

*Sty taster eximius forma atlanticus Broch, 1936 
Gunior synonym of S. duchassaingi) 

Stylaster filogranus Pourtales, 1871 

Stylaster punctatus Pourtales, 1871 



VOLUME 95, NUMBER 1 211 

Stylaster roseus (Pallas, 1766) 

(originally described as Madrepora) 
*Stylaster sanguineus Milne Edwards & Haime, 1849 

(type-locality Australia; dubious record for western Atlantic: see text) 
Allopora miniata Pourtales, 1868 
Stenohelia challengeri Boschma, 1951 

(new name for Stenohelia profunda Moseley, 1881) 
"^Stenohelia maderensis (Johnson, 1862) 

(originally described as Allopora, type-locality Madeira, East Atlantic; dubious record for 

western Atlantic: see text) 
Stenohelia virginis (Lindstrom, 1877) 

(originally described as Crypthelia) 
Crypthelia peircei Pourtales, 1867 
^Crypthelia pudica Milne Edwards & Haime, 1849 

(type-locality Philippine Islands; dubious record for western Atlantic) 
Distichopora barbadensis Pourtales, 1874 
Distichopora cervina Pourtales, 1871 
Distichopora contorta Pourtales, 1878 
Distichopora foliace a Pourtales, 1868 
Distichopora sulcata Pourtales, 1867 
*Errina (Errina) aspera (Linne, 1767) 

(originally described as Millepora, type-locality Mediterranean; not present in western Atlantic: 

misidentification of Broch, 1942) 
Errina (Lepidopora) carinata Pourtales, 1867 

(originally described as Heliopora) 
Errina (Lepidopora) cochleata Pourtales, 1867 
Errina (Lepidopora) decipiens Boschma, 1964 
Errina (Lepidopora) glabra Pourtales, 1867 
Pliobothrus symmetricus Pourtales, 1868 
Pliobothrus tubulatus (Pourtales, 1867) 

(originally described as Heliopora) 

General remarks. — A review of the literature and a preliminary study of nu- 
merous specimens in various museum collections (National Museum of Natural 
History, Washington, D.C.; Yale Peabody Museum; Museum of Comparative 
Zoology; Museum National d'Histoire Naturelle, Paris; Rijksmuseum van Na- 
tuurlijke Historic, Leiden; Zoologisch Museum, Amsterdam; British Museum 
(Natural History), London; Royal Scottish Museum, Edinburgh; Zoologisk Mu- 
seum, Copenhagen; Naturhistoriska Riksmuseet, Stockholm) provide the bases 
for the following remarks on the synonymy and the distribution of the West Indian 
Stylasterina; however, a monographic study including a revision of all previous 
records is greatly needed. 

Stylaster elegans (name preoccupied), Stylaster eximius, S. eximius forma at- 
lantica and Stylaster duchassaingi are all names given to the same species for 
which the latter name must be retained. 

Stylaster complanatus has characters somewhat intermediate between typical 
Stylaster and Stenohelia. Here it is listed with the former genus although some 
authors referred it to Stenohelia (Saville Kent 1870; Broch 1936; Boschma 1957, 
1964b,c). Broch (1936) incorrectly synonymized it with Stenohelia virginis. 

The shallow-water specimens of Stylaster punctatus (which Pourtales seemed 
to consider as the typical form) are identical with Stylaster roseus. 

The new name Stenohelia challengeri had been introduced by Boschma (1951) 
for Stenohelia profunda Moseley, 1879, because of homonymy with Allopora 



212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

profunda Moseley, 1879, in case the closely related genera Stylaster, Allopora 
and Stenohelia were synonymized, as has been suggested previously by some 
authors. However, Boschma's later papers considered these as separate genera 
(a point of view now generally accepted), and therefore there was no real need 
for that new name. 

Errina (Lepidopora) cochleata has been incorrectly described as a bryozoan, 
Hornera galeata Smitt, 1872 (see Zibrowius, in press). 

Most of the West Indian species appear to be endemic to that area, or at least 
to a larger area in the West Atlantic, whereas only a few species are amphi- 
Atlantic. The occurrence in the eastern Atlantic of Stylaster erubescens (off west- 
ern Brittany, between Hebrides and Faeroe, between Orkney and Faeroe, off 
southeast Iceland) and of Pliobothrus symmetricus (Hyeres Banc, Azores, off 
western Brittany, off western Ireland, Faeroe, Norway, off southeast Iceland) is 
confirmed. In addition, a still unnamed species of Crypthelia known from the 
eastern Atlantic (Hyeres Banc, Azores, Madeira) has been recognized among 
Pourtales material from the Lesser Antilles (Museum of Comparative Zoology). 

Not all indications in the literature on amphi-Atlantic distributions are correct. 
Errina (Errina) aspera has been reported by Broch (1942) from the West Indies 
(detailed origin unknown) based on a small fragment of pink Errina (Naturhis- 
toriska Riksmuseet, Stockholm, No. 45). However, this is not E. aspera, oth- 
erwise known from the southwest Mediterranean and the eastern Atlantic, but 
probably an Antarctic or Subantarctic species (mislabelled), as already suggested 
by Boschma (1965:2). Nothing similar of authentic West Indian origin has yet 
been found in the various museum collections. 

The case of Stenohelia maderensis, reported by Boschma (1964b, 1964c) from 
the West Indies (St. Vincent), needs further investigation. 

Indications in the literature of species of even wider distribution, occurring in 
both the West Indies and in the Indo-Pacific, are also doubtful and are possibly 
all caused by mislabelling (origin confused), inversion of labels in the collections, 
or misidentification. This is sufficiently confirmed in some cases. For instance, 
Stylaster erubescens (?) sensu Moseley, 1881, from the Kermadec Islands, is a 
Conopora (British Museum (Nat. Hist.) 1880.11.25.178). 

Typical specimens of Stylaster sanguineus, a pink shallow-water species well 
known from the central and southwest Pacific, are labelled as being from Florida 
in the Museum of Comparative Zoology and the Yale Peabody Museum. They 
may well be those mentioned by Pourtales (1871:83) as undistinguishable from 
authentic Pacific specimens. A label-inversion error with specimens of the pink 
shallow- water species Stylaster roseus from the West Indies, might best explain 
this unusual, still unverified distribution. Furthermore, S. roseus is mentioned on 
some of the old labels, either as the original identification or as a synonym of S. 
sanguineus. Boschma (1957) reported the reverse case of Stylaster roseus oc- 
casionally being mentioned from the Pacific (Hawaii, Samoa). 

Boschma (1957) further points out that material from the Loyalty Islands used 
in an anatomical study (England 1926) had erroneously been cited as Stylaster 
filogranus, the latter being a West Indian species. 

A detailed study will probably confirm that the Stylaster from Mauritius, In- 
donesia, the Philippine Islands and the Sea of Okhotsk referred to as 5. eximius 
(see Boschma 1957) are not only "forma" or "facies" but species distinct from 



VOLUME 95, NUMBER 1 213 

the West Indian one, for which the name Stylaster duchassaingi has priority over 
S. eximius. A similar result can reasonably be expected from the comparison of 
Stenohelia virginis (Lindstrom, 1877) with Stylaster virginis sensu Hickson and 
England, 1905, from Indonesia. 

Boschma (1964c) affirms that Moseley's (1881) two records of Stenohelia pro- 
funda (renamed Stenohelia challengeri) from the West Indies (type-locality) and 
the Kermadec Islands are indeed the same species. If he is correct, it would be 
a very unusual distribution, but it should be remembered that specimens of var- 
ious zoological groups in the "Challenger" collections have confused origins. On 
the other hand, specimens from the Galapagos Islands originally reported as 
Stenohelia profunda by Marenzeller (1904) have been described by Boschma 
(1964b) as a distinct species, Stenohelia robusta. 

World-wide, several species have probably been identified as Crypthelia pu- 
dica. Moseley's (1881) record of C. pudica from the West Indies (Sombrero) is 
considered with suspicion; until more detailed comparisons are made, we do not 
assume that it occurs both in the western Pacific (type-locality: Phillipine Islands) 
and in the western Atlantic. 

A new species of Stylaster, distinct from all those previously reported from the 
West Indies, is described below. The unstudied material from the cruise of the 
United States Coast Survey steamer "Blake" in 1878/79 to the Caribbean Islands 
(eight stations) was discovered in the collection of the Museum of Comparative 
Zoology during a visit in summer, 1980. The original labels printed for that cruise 
read: "U.S. Coast Survey, C. P. Patterson, Supi. -Caribbean Islands exploration. 
U.S.C.S., S. Blake, Alex. Agassiz, 1878-79." They are marked in pencil with the 
station number, depth (in fathoms), and the name of the nearest island. Comple- 
mentary information on the stations was obtained from Smith's compilation 
(1889). No preliminary identification, generic or specific, was found with the 
specimens. Pourtales' (1880) report on this cruise of 1878/79 deals only with the 
Scleractinia and Antipatharia. He probably intended to publish a separate report 
on the Stylasterina, whereas his previous papers include both Scleractinia and 
Stylasterina. 

Stylaster antillarum, new species 
Figs. 1-4 

Types. — The nearly complete colony from "Blake" Sta. 241 is designated as 
holotype. The other colonies, branches, and fragments from "Blake" stations 
213, 215, 216, 218, 219, 231, and 238 are designated as paratypes. Depository: 
holotype and most paratypes at MCZ; two paratypes at USNM ("Blake" sta. 
216, USNM 60350; "Blake" sta. 219, USNM 60349). 

Description. — The "Blake" material is in good condition; all had been collected 
alive and then dried. For the present study it has been cleaned of the dry tissue 
and centenary dust with sodium hypochlorite solution. 

The holotype is a regularly flabellate and moderately branched colony rising 
from a strong, encrusting base that is 7.5 mm wide. About 40 mm high and 39 
mm wide, the holotype is the largest of the specimens from the "Blake" stations. 
Most of the other specimens are just branches or fragments and considerably 
smaller. Colonies comprising the entire base or at least the basal part of the trunk 
are rare. The coenosteum bears distinct, discontinuous carinae, which are most 



214 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Stylaster antillamm type-material from "Blake" cruise 1878/79. 



Station 



Date 



Area 



Coordinates 



Depth 



Quantity 



213 12.2.1879 Martinique 14°32'38"N, 61°06'40"W 357 fm-653 m 1 branch 

215 15.2.1879 St. Lucia 13°51'30"N, 61°03'45"W 226fm^l4m 2 branches 

216 15.2.1879 St. Lucia 13°51'45"N, 61°03'30"W *153 fm-280 m 15 colonies, branches, 

fragments 

218 15.2.1879 St. Lucia 13°49'12"N, 61°04'40"W 164 fm-300 m 1 colony, 1 branch 

219 15.2.1879 St. Lucia 13°49'50"N, 6r03'50"W 151 fm-276 m 9 colonies, branches, 

fragments 
231 20.2.1879 St. Vincent 13°12'10"N, 6ri7'18"W 95 fm-174 m 1 branch 

238 23.2.1879 Grenadines: 12°46'10"N, 6r23'35"W *126 fm-231 m 1 branch 

Cannuan 
241 24.2.1879 Grenadines: 12°28'22"N, 6r32'18"W 163 fm-298 M 1 colony 

Carriacou 

* The original labels indicate slightly different depths, respectively 154 fathoms and 127 fathoms. 



well developed near a cyclosystem, one carina corresponding to each dactylo- 
pore. The carinae are about 32-37 /xm wide, about 64 />tm tall, and variable in 
length, ranging from 0.1 mm to several millimeters. The distance between two 
adjacent carinae varies between 0.20-0.30 mm. The coenosteum lying between 
the carinae is usually slightly concave and is penetrated by coenosteal slits which 
are also aligned parallel to the carinae. The slits are variable in length, ranging 
from 20-80 /xm, but are consistently 8-9 /xm wide. Distinct, rounded coenosteal 
granules are not present; however, the coenosteal surface is covered by irregular 
deposits of approximately 3-6 x 2-3 /xm in size, particularly noticeable at the 
edges of coenosteal slits (Fig. 2D). 

Branching is strictly in one plane as the result of simultaneous budding to the 
left and to the right from below the top part of older cyclosystems. Otherwise, 
branches lengthen by alternate budding to the left and right. Angle of budding 
about 60-70°. This sympodial pattern persists even on the main branches. Cyclo- 
systems occur only on the lateral branch edges; the anterior and posterior sides 
of the colony are entirely devoid of cyclosystems. 

Well-developed young distal and peripheral cyclosystems are long, cylindrical 
structures, circular in cross section, except distally, where they are compressed 
in the flabellar plane. They are about 3.2-3.6 mm high and 0.9 mm in diameter 
at the base. The thickness of the wall of a cyclosystem is about 0.2 mm, and the 
lesser axis of the distal part of a cyclosystem is about 0.9-1.0 mm, the greater 
axis, 1.2-1.3 mm. Older cyclosystems are not much taller but do become pro- 
nouncedly thicker (thickening of the wall), becoming proportionally more com- 
pressed at the top (lesser axis 1.0 mm, greater axis up to 1.6 mm). Initial stages 
of new cyclosystems start as low annular ridges, 0.7 mm wide, below the tops of 
older ones. 

Seen from the anterior and posterior sides of the colony, the top of the cyclo- 
system is distinctly flabellate with a rounded outline, the pseudosepta and dac- 
tylopore openings near the greater (longitudinal) axis being much lower than those 
near the lesser (transverse) axis. Adjacent to the greater axis the dactylopores 



VOLUME 95, NUMBER 1 



215 




Fig. 1. Types of S. antillarum: A, Holotype colony from "Blake" sta. 241, MCZ, x2.0; B, 
Branchlet of holotype with prominent ampullae and circular depressions left by ruptured ampullae, 
x6.3; C, Detail of paratype colony from "Blake" sta. 216, MCZ, with ampullae occurring singly or 
in pairs below the flared top of the cyclosystems, x6.3; D, paratype colony from "Blake" sta. 219, 
MCZ, with large acrothoracic cirriped crypt in the fractured base, x4.3. 



216 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Paratypes of S. antillarum from "Blake" sta. 216, USNM 60350: A-B, Lateral and oblique 
views of cyclosystems and coenosteal ridges, one ridge corresponding to each dactylopore, x44, 
x34, respectively; C, Coenosteal texture, x 106; D, Coenosteal slit, x640; E, Oral view of cyclosystem 
with 15 dactylopores, x50, SEM stereomicrographs. 



VOLUME 95, NUMBER 1 



217 




Fig. 3. Paratypes of 5'. antillamm: A, "Blake" sta. 216, longitudinal section of gastropore re- 
vealing gastrostyle, x24; B, Gastrostyle of figure 3A, xllO; C, "Blake" sta. 219, USNM 60349 
gastrostyle and inner ring of granules, xllO, SEM stereomicrographs; D, "Blake" sta. 216, dacty- 
lostyle, x680; E, Gastrostyle tip of fig. 2C, x340. 



218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

are closely spaced, separated by very distinct lamellar pseudosepta; adjacent to 
the lesser axis they are widely spaced, separated by thick pseudosepta which are 
less prominent and more flattened toward the gastropore tube. Upper rim of 
cyclosystem crenulate; upper part of outside surface carinate, with carinae cor- 
responding to dactylopores and depressions to pseudosepta. 

Dactylopores are deep, narrow pits parallel to the central gastropore tube, 
separated from the latter by a thin wall except at the upper part where the narrow 
slit-like dactylotomes permit open communication. The gastropore tube is cylin- 
drical, up to 3 mm long, and about 0.40-0.43 mm in diameter. The basal 0.5 mm 
of the tube, the gastrostyle chamber, is narrower, usually only 0.3 mm in diam- 
eter. At the boundary of the gastrostyle chamber with the larger, upper part there 
is a distinct ring of granules, the ring being about 0.2 mm thick. The granules are 
thick and blunt, about 50 /i,m tall and 35-40 fjum thick, and are closely spaced, 
often fusing. This ring of granules further constricts the gastropore tube to a 
diameter of about 0.20-0.25 mm. 

Gastrostyles are conical to bullet-shaped, up to 0.55 mm tall and 0.20 mm 
wide. Height : width ratios of those styles measured ranged from 2.67-2.90. The 
style occupies the lower 15-19% of the gastropore tube, the deUcate tip usually 
extending just above the ring of blunt granules. Styles are highly sculptured with 
spines which are particularly well developed at the tip where they measure up to 
36 /xm long and 8.5 /jun wide (Fig. 3E). No ridges are present on the styles. 

Dactylostyles are very poorly developed, composed of small, irregular, angular 
deposits measuring about 15x5 /xm. These deposits are usually fused in a line 
forming the dactylostyle. 

The number of dactylopores varies from 10 to 18 in the 200 cyclosystems 
examined in detail; 14 dactylopores per cyclosystem is both the median and the 
mode, 14.35 is the average. 

Dactylopores per cyclosystem 10 11 12 13 14 15 16 17 18 

Number of cyclosystems 7 9 16 28 44 40 31 17 8 

The ampullae are prominent outgrowths on both the anterior and posterior 
sides of the colony, located just below the flared top of the cyclosystem, at the 
same level from which the younger cyclosystems bud. They measure up to 0.57 
mm tall and 0.61 mm broad. Ampullae occur singly or in pairs and the same 
cyclosystem may have ampullae on both its anterior and posterior sides. Their 
roughly hemispherical shapes are obscured by irregular crests and small knobs. 
Some ampullae have a tiny pore near the base, probably an efferent duct. Rather 
frequently ampullae are ruptured, leaving a circular depression. 

Remarks. — Cavities caused by acrothoracic cirripeds are present in the base 
of two colonies, one from "Blake" sta. 216, the other from "Blake" sta. 219. 
No other symbionts are known from Stylaster antillarum. 

Discussion. — The combination of characters described above distinguishes S. 
antillarum from all previously described species oi Stylaster (see Boschma 1957; 
Vervoort & Zibrowius 1981). As far as could be deduced from the literature, only 
Stylaster multiplex Hickson & England, 1905, and Stylaster crassior Broch, 1936, 
show some resemblance to S. antillarum, mainly by the somewhat elongate tops 
of their cyclosystems. However, comparison with S. multiplex (type-material 
from Indonesia) and 5. crassior (from the Comoros, "Le Suroit" cruise, Ben- 



VOLUME 95, NUMBER 1 



219 




Fig. 4. Paratypes of S. antillarum from "Blake" sta. 216, USNM 60350: A, Cross section of 
gastropore tube just above ring of granules; gastrostyle tip in center; rudimentary dactylopore tubes 
encircle gastropore, x50; B, Two fused granules from the ring of granules of fig. 4A, x680; C, 
cyclosystem flanked by irregularly shaped ampullae, x30. 



thedi, 1977; the type-material from Mauritius was not available) indicates that 
these forms, similar to each other, differ considerably from the West Indian 
species. The tops of their cyclosystems are not as prominent or flabellate as in 
S. antillarum, and their dactylopores and pseudosepta are more uniformly 
spaced. Other differences concern the depth of the gastropore tube, the surface 
structure of the branches, and the ampullae. 



Acknowledgments 

We would like to thank Dr. Herbert W. Levi (Museum of Comparative Zool- 
ogy, Harvard University) for making available to us the specimens reported in 



220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

this paper and for the cooperation extended toward the senior author during his 
visit to the MCZ in 1980. 

Literature Cited 

Boschma, H. 195L Notes on Stylasterina (Hydrocorallia). — Proceedings Koninklijke Nederlandse 
Akademie van Wetenschappen (C)54(5):45 1-458. 

. 1955. The specific characters of the coral Stylaster roseus. — Deep-Sea Research 3, Supple- 
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. 1957. List of the described species of the order Stylasterina.— Zoologische Verhandelingen, 

Leiden 33:1-72. 

. 1962. Notes on the Stylasterine coral Allopora miniata. — Proceedings Koninklijke Neder- 
landse Akademie van Wetenschappen (C)65(3): 195-204, pi. 1-2. 

. 1964a. Errina (Lepidopora) decipiens, 2l new Stylasterine coral from the West Indies. — 

Proceedings Koninklijke Nederlandse Akademie van Wetenschappen (C)67(2): 55-63, pi. 1. 

. 1964b. On the Stylasterina of the genus Stenohelia. — Proceedings Koninklijke Nederlandse 

Akademie van Wetenschappen (C): 67(2): 64-73, pi. 1-2. 

. 1964c. Further notes on the Stylasterine corals Stenohelia challenged and Stenohelia mader- 

ensis. — Proceedings Koninklijke Nederlandse Akademie Wetenschappen (C)67(2): 78-84, pi. 
1-2. 

. 1965. Further notes on Stylaster roseus (Pallas). I & II. — Proceedings Koninklijke Neder- 
landse Akademie Wetenschappen (C)68(2):229-250, pi. 1^. 

Broch, H. 1936. Untersuchungen an Stylasteriden (Hydrokorallen). Teil 1. — Skrifter utgitt av Det 
Norske Videnskaps-Academi i Oslo, I Mat.-Naturv. Klasse, 8:1-103, pi. 1-13. 

. 1942. Investigations on Stylasteridae (Hydrocorals). — Skrifter utgitt av Det Norske Viden- 
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Duchassaing, P., and J. Michelotti. 1861. Memoire sur les coralliaires des Antilles. — Memoires de 
r Academic des Sciences de Turin (2)19:279-365, pi. 1-10. 

. 1864. Supplement au memoire sur les coralliaires des Antilles. — Memoires de 1' Academic 

des Sciences de Turin (2)23:97-206, pi. 1-11. 

England, H. M., 1926. Development of the gonophores of the Stylasteridae. — Proceedings of the 
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Hickson, S. J., and H. M. England. 1905. The Stylasterina of the Siboga expedition. — Siboga-Ex- 
peditie 8:1-26, pi. 1-3. 

Lindstrom, G. 1877. Contributions to the actinology of the Atlantic Ocean. — Kongliga Svenska Ve- 
tenskaps-Akademiens Handlingar, Stockholm 14(2): 1-26, pi. 1-3. 

Marenzeller, E. von. 1904. Stein- und Hydro-korallen. — Bulletin of the Museum of Comparative 
Zoology 43(2):75-87, pi. 1-3. 

Moseley, H. N. 1879. On the structure of the Stylasteridae, a family of the Hydroid stony corals. — 
Philosophical Transactions of the Royal Society of London 169, 1878:425-503, pi. 34-44. 

. 1881. Report on certain Hydroid, Alcyonarian and Madreporarian corals procured during the 

voyage of H. M.S. Challenger in the years 1873-1876. — Report on the Scientific Results of the 
Voyage of H. M.S. Challenger during the years 1873-1876. Zoology 2:1-248, 32 pi. 

Pallas, P. S. 1766. Elenchus zoophytorum . . . Hagae Comitum xvi-28-451 pp. 

Pourtales, L. F. de. 1867. Contributions to the fauna of the Gulf Stream at great depths. — Bulletin 
of the Museum of Comparative Zoology 1(6): 103-120. 

. 1868. Contributions to the fauna of the Gulf Stream at great depths (2^ series). — Bulletin of 

the Museum of Comparative Zoology 1(7): 121-142. 

. 1871 . Deep-sea corals — Illustrated Catalogue of the Museum of Comparative Zoology 4: 1-93, 

pi. 1-8. 

1874. The zoological results of the Hassler Expedition. Deep-sea corals. — Illustrated Cata- 
logue of the Museum of Comparative Zoology 8:33^9, pi. 6-9. 

. 1878. Report on the results of dredging, under the supervision of Alexander Agassiz, in the 

Gulf of Mexico, by the U.S. coast survey steamer "Blake." Corals. — Bulletin of the Museum 
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Roos, P. J. 1971. The shallow-water stony corals of the Netherlands Antilles. — Studies on the Fauna 
of Curasao and other Caribbean Islands 37(130): 1-108, pi. 1-53. 



VOLUME 95, NUMBER 1 221 

Saville Kent, W. 1870. On a new genus of Madreporaria or stony corals (Stenohelia). — Annals and 

Magazine of Natural History, London (4)5:120-123. 
. 1871. On some new and little-known species of Madrepores, or stony corals, in the British 

Museum collection. — Proceedings of the Zoological Society of London (1871): 275-286, pi. 

23-25. 
Smith, S. 1889. Lists of the dredging stations of the U.S. Fish Commission, the U.S. Coast Survey, 

and the British steamer Challenger, in North American waters, from 1867 to 1887, together 

with those of the principal European government expeditions in the Atlantic and Arctic 

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Vetenskaps-akademiens Handlingar, Stockholm 10(11): 1-20, pi. 1-5. 
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stitution, Washington, D.C. 20560. 



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CONTENTS 

A new species of skate from Western Australia with comments on the status of Pavoraja 

Whitley, 1939 (Chondrichthyes: Rajiformes) John D. McEachran and Janice D. Fechhelm 1 
A new species of Gymnure, Podogymnura, (Mammalia: Erinaceidae) from Dinagat Island, 

Philippines Lawrence R. Heaney and Gary S. Morgan 13 

Checklist of the fishes of the central and northern Appalachian Mountains 

Jay R. Stauffer, Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jenkins 27 
Two new genera of deep-sea Polychaete worms of the family Ampharetidae and the role of one 

species in deep-sea ecosystems Robert Zottoli 48 

The karyotype of the Eurasian flying squirrel, Pteromys volans (L.), with a consideration of 

karyotypic and other distinctions in Glaucomys spp. (Rodentia: Sciuridae) 

V. R. Rausch and R. L. Rausch 58 
A new species of the genus Echininus (MoUusca: Littorinidae: Echinininae) with a review 

of the subfamily Joseph Rosewater 67 

A new dwarf Sphaerodactylus from Haiti (Lacertilia: Gekkonidae) Richard Thomas 81 

Amerigoniscus malheurensis, new species, from a cave in western Oregon (Crustacea: Isopoda: 

Trichoniscidae) George A. Schultz 89 

A new species of deep-sea isopod, Storthyngura myriamae, from the Walvis Ridge off South 

Africa Robert Y. George and Noel A. Hinton 93 

Phytoplankton distribution along the eastern coast of the USA IV. Shelf waters between Cape 

Lookout, North Carolina, and Cape Canaveral, Florida Harold G. Marshall 99 

Nomenclatural status of the foraminiferal genus Cubanella Saidova, 1981 

Richard W. Huddleston and Drew Haman 1 14 
Some new and old species of the primnoid genus Callogorgia Gray, with a revalidation of the 

related genus Fanellia Gray (Coelenterata: Anthozoa) Frederick M. Bayer 116 

Freshwater triclads (Turbellaria) of North America. XIII. Phagocata hamptonae , new species, 

from Nevada Roman Kenk 161 

Classificatory revisions in gammaridean Amphipoda (Crustacea), Part 2 

J. Laurens Barnard and Gordon S. Karaman 167 
Hemiodopsis ocellata, a new hemiodontid characoid fish (Pisces: Characoidea) from western 

Surinam Richard P. Vari 188 

Evidence of ontogenetic setal changes in Heteromastus filiformis (Polychaeta: Capitellidae) 

Thomas J. Fredette 194 
A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from the western Highland 

Rim in Tennessee Jerry A. Louton 198 

Two new species of Onuphis (Onuphidae: Polychaeta) from Uruguay Kristian Fauchald 203 

Remarks on the Stylasterine fauna of the West Indies, with a description oi Stylaster antillarum, 

a new species from the Lesser Antilles (Cnidaria: Hydrozoa: Stylasterina) 

Helmut Zibrowius and Stephen D. Cairns 210 



(ISSN 0006-324X) 



Proceedings 

of the 

BIOLOQICAL SOCIETY 

of 

WASHINGTON 



Volume 95 11 August 1982 Number 2 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

1981-1982 
Officers 



President: Raymond B. Manning 
Vice President: Paul J. Spangler 



Frederick M. Bayer 
Kristian Fauchald 



Elected Council 



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Secretary: Michael A. Bogan 
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Isabel C. Canet 
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Editor: Brian Kensley 



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PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 223-231 

ATROPHECAECUM LOBACETABULARE, N. SP. 
(DIGENEA: CRYPTOGONIMIDAE: ACANTHOSTOMINAE) 

WITH DISCUSSION OF THE GENERIC STATUS OF 
PARACANTHOSTOMUM FISCHTHAL AND KUNTZ, 1965, 

AND ATEUCHOCEPHALA COIL AND KUNTZ, 1960 

Daniel R. Brooks and Janine N. Caira 

Abstract. — A new species of Atrophecaecum is described from various sea 
snakes collected in northern Australia and Malaysia. The new species most 
closely resembles A. burminis, A. simhai, and Paracanthostomum cerberi in 
having one atrophied cecum, preovarian vitellaria, vitelline follicles not extending 
anteriorly to the posterior margin of the seminal vesicle, eggs attaining lengths 
greater than 30 /itm, and a short prepharynx. It is unique among acanthostomes 
in possessing a flap of tissue on the posterior margin of the acetabulum, giving 
that organ a bipartite appearance. The new species further resembles A. burminis 
and-A. simhai by possessing oral spines, and resembles P. cerberi in having a 
subterminal mouth. Inclusion of Paracanthostomum cerberi and Ateuchocephala 
marina in Atrophecaecum provides the most parsimonious hypothesis of phylo- 
genetic relationships among the taxa involved, and produces a more efficient and 
stable classification. 



Brooks (1980) recently revised the digenean subfamily Acanthostominae Poche 
based on a phylogenetic analysis. In that monograph a new species of Atrophe- 
caecum Bhalerao was discussed in terms of its effect on the classification of 
acanthostomes and two other cryptogonimid genera, Paracanthostomum Fisch- 
thal and Kuntz, and Ateuchocephala Coil and Kuntz. Specifically, the possibility 
was raised that Ateuchocephala and Paracanthostomum could not be recognized 
as natural (monophyletic) taxa unless Atrophecaecum was somehow subdivided. 
This report describes the new species of Atrophecaecum and, based on phylo- 
genetic analysis, shows that the most useful classification for the taxa involved 
is one including Ateuchocephala and Paracanthostomum in the genus Atrophe- 
caecum. 

Specimens upon which the description of the new species is based were col- 
lected from sea-snakes quick-frozen shortly after capture and kept frozen until 
returned to the laboratory, where they were thawed and examined. Most were 
partially thawed, eviscerated, and the viscera refrozen and shipped to the Uni- 
versity of Alberta. Collected helminths were fixed with cold AFA, stained with 
Harris' hematoxylin and mounted in Permount (Fisher). Measurements are in 
micra unless otherwise stated, with mean values (i) and number of observations 
(n) for some traits; figures were drawn with the aid of a drawing tube. 

Atrophecaecum lobacetabulare, new species 
Figs. 1^ 

Description (based on 34 specimens). — Body elongate, 4.32-12.30 mm long 
(n = 27) by 0.24-0.75 mm wide (n = 27); widest point variable in hindbody; ratio 



224 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 1-4. Atrophecaecum lobacetabulare: 1, Ventral view of holotype; 2, Close-up of posterior 
end of paratype, showing position of seminal receptacle, which is not apparent in holotype; 3, Close- 
up of ootype region; 4, Dorsal view of acetabular region of holotype, showing terminal genitalia. 
Abbreviations: HD = hermaphroditic duct; LC = Laurer's canal; MG = Mehlis' gland; O = ovary; 
SR = seminal receptacle; SV = seminal vesicle; U = uterus; VR = vitelline reservoir; VD - vitel- 
line duct. 



of body width to length 1:9.1-33.8 (jc = 1:18.1, n = 27). Tegument spinose; extent 
of spination not determined due to fixation methods. Eyespot pigment lacking. 
Oral sucker cup-shaped, terminal with subterminal mouth, 154-297 (n = 24) long 
by 143-231 (n = 24) wide, surrounded by single uninterrupted row of 24-26 
spines; spines 22-38 (n = 20) long by 12-29 (n = 20) wide. Acetabulum 105-231 
long (n = 23) by 95-209 wide (n = 23), with posterior lobe giving bipartite ap- 
pearance; lobe not apparent in many specimens. Forebody 6.4-14.1% of total 
body length. Ratio of oral sucker width to acetabular width 1:0.64-1.07 {x = 
1:0.84, n = 20). Prepharynx 60-242 long (n = 21), thin- walled. Pharynx barrel- 
shaped, 108-148 long (n = 22) by 99-214 wide (n = 22). Ratio of oral sucker 
width to pharyngeal width 1:0.50-0.89 (i = 1:0.71, n = 16). Ratio of acetabular 
width to pharyngeal width 1:0.59-1.15 (x = 1:0.87, n = 16). Esophagus usually 
extremely short, not measured. Cecal bifurcation less than 5% of total body length 
pre-acetabular; ceca lined with epithelium, opening separately and laterally near 
posterior end of body; one cecum atrophied. 

Testes spherical to subspherical, smooth, tandem, not contiguous; anterior 
testis 209-345 long by 132-208 wide, posterior testis 220-396 long by 132-352 
wide; post-testicular space 3-11% of total body length (x = 7%, n = 26). Seminal 
vesicle consisting of posterior saccate part and anterior sinus part, sinuous, me- 
dian, intercecal, extending 4.6-11.7 times acetabular length postacetabular (x = 
9.1, n = 18). Prostatic duct surrounded by few prostatic cells free in parenchyma. 
Preacetabular pit without spines, with transverse aperture 80-83 wide, surround- 



VOLUME 95, NUMBER 2 225 

ed by gland cells; gonotyl lacking. Genital pore immediately preacetabular, open- 
ing immediately posterior to preacetabular pit. Postacetabular pit lacking. 

Ovary approximately one ovarian diameter pretesticular, not contiguous with 
anterior testis, spherical to subspherical, 120-264 long by 99-242 wide. Seminal 
receptacle posterodorsal to ovary, 132-506 long by 77-198 wide. Ootype region 
as in Fig. 3. Uterus wound in ascending loops intercecal between ovary and 
acetabulum; loops occupying 63-75% of total body length (x = 70%, n = 23); 
joining hermaphroditic duct posterior to acetabulum. Vitellaria follicular; follicles 
in 2 longitudinal rows dorsal and lateral to ceca, extending from immediately 
preovarian to 8-50% of total body length postacetabular (x = 27%, n = 24), ter- 
minating at uneven levels. Left vitellarium follicles extending anteriorly to 8^6% 
of total body length postacetabular, right vitellarium follicles extending anteriorly 
to 9-50% of total body length postacetabular. Follicles not reaching anteriorly to 
posterior margin of seminal vesicle, 14-33 long by 12-29 wide. Eggs 25-35 (jc = 
28, n = 34) long by 13-20 (i = 16.5, n = 34) wide. Excretory vesicle Y-shaped; 
bifurcation posterodorsal to acetabulum; pore terminal with muscular sphincter 
surrounded by gland cells. 

Hosts. — Disteira major (Shaw), Hydrophis caerulescens (Shaw), H. spiralis 
(Shaw), Enhydrina schistosa (Daudin) (type), Lapemis hardwicki Gray (Ophidia: 
Hydrophiidae: Hydrophiinae: Hydrophiini). 

Site of infection. — Small intestine. 

Localities. — Penang, Malaysia (type) {E. schistosa, H. spiralis, L. hardwicki); 
Western Gulf of Carpenteria, Australia {D. major, H. caerulescens). 

Holotype.—USNU Helm. Coll. No. 77161. Paratypes.—USNM Helm. Coll. 
No. 77162, 77163, 77164. 

Etymology. — The species is named for its lobate acetabulum, a feature unique 
among acanthostomes. 

Atrophecaecum lobacetabulare most closely resembles the species A. bur- 
minis (Bhalerao, 1926) Khalil, 1963, and A. simhai Khalil, 1963, and Para- 
canthostomum cerberi Fischthal and Kuntz, 1965, by having one atrophied cecum, 
preovarian vitellaria, vitelline follicles not extending anteriorly to the pos- 
terior margin of the seminal vesicle, eggs attaining lengths greater than 30 /xm, 
and a very short prepharynx. The new species is unique among known acan- 
thostomes by virtue of its possessing a lobate acetabulum. It resembles A. bur- 
minis and A. simhai by having oral spines, although they are relatively much 
smaller than those of most other acanthostomes. It resembles P. cerberi by pos- 
sessing a subterminal mouth with terminal oral sucker. Thus, A. lobacetabulare 
occupies a systematic position intermediate between the taxa listed above. 

Discussion 

Brooks (1980) alluded to possible taxonomic problems arising from the discov- 
ery of A. lobacetabulare. The nature of the problem, and of its solution, may be 
seen best by examining the relative positions of A. lobacetabulare, P. cerberi, 
and Ateuchocephala marina in a cladogram depicting the phylogenetic relation- 
ships of Atrophecaecum spp. (Fig. 5). Synapomorphies (shared special traits) 
linking the new species and the other two taxa above to Atrophecaecum are listed 
on the cladogram (simplified from that of Brooks 1980). 

Figure 5 clearly shows that A. lobacetabulare, P. cerberi, and Ateuchocephala 



226 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




no prepharynx 
short seminal vesicle 
secondary vitelline clusters 
around testes 
no oral spines (2° lossj 



nnouth subternninal 
uterine loops occupying more than 50% TBL 
eggs longer than 30gm 



one cecum atrophied 



short prepharynx 



vitellaria terminating preovarially 



Fig. 5. Cladogram, partly modified from that of Brooks (1980), depicting phylogenetic relationships 
of Atrophecaecum spp. plus Paracanthostomum cerberi and Ateuchocephala marina. Modifications 
comprise deletion of some characters from the cladogram for the sake of clarity in viewing, and 
addition of A. lobacetabulare and P. cerberi and A. marina. 



marina form a monophyletic group most closely related to Atrophecaecum bur- 
minis and A. simhai, hence our choice of comparisons above. This arrangement 
would require that Paracanthostomum and Ateuchocephala be considered junior 
synonyms of Atrophecaecum if A. lobacetabulare and all other members of the 
genus are to be retained in a single genus. Despite the perceived morphological 
distinctness of oral structures possessed by P. cerberi and Ateuchocephala ma- 
rina, they represent only secondary loss of traits (oral spines) already reduced in 
size in their closest relative and a tilting of the mouth orientation. Paracantho- 
stomum and Ateuchocephala are therefore characterized in part by the absence 
of traits. Conversely, Atrophecaecum would not exist as a natural taxon if P. 
cerberi and Ateuchocephala marina are excluded from it. 

Three types of objections commonly raised concerning cladistic classification 
include: (1) they involve loss of information about "gaps" or degrees of difference 
between taxa (anagenetic information), (2) they produce confusing and unusable 
diagnoses and keys, and (3) any attempts to preserve cladistic information pro- 
duce long unwieldy classifications. All of these objections have been treated 
empirically in papers published by various authors primarily in Systematic Zo- 
ology during the past ten years. This study provides an opportunity to demon- 
strate the efficacy of phylogenetic classification using a set of real taxa. 

Loss of information. — Consider the three genera Atrophecaecum, Paracan- 
thostomum, and Ateuchocephala in the classification presented by Yamaguti 
(1971). Paracanthostomum and Atrophecaecum (as a subgenus of Acanthosto- 
mum) belong in the Acanthostominae and Ateuchocephala comprises the sole 



VOLUME 95, NUMBER 2 227 

member of the Ateuchocephalinae. Such a classification does not represent any 
information concerning any members of Atrophecaecum which exhibit traits in- 
termediate between the other taxa, nor does it recognize any of the similarities 
between Ateuchocephala and Paracanthostomum. Yamaguti's classification pro- 
posed a set of relationships opposite to the phylogenetic relationships supported 
by known data; Ateuchocephala was considered the sister-Hneage of Paracan- 
thostomum plus all other acanthostomes, and Paracanthostomum was considered 
the sister-lineage of all armed acanthostomes, including Atrophecaecum. Even 
a Paracanthostomum were considered a member of the Ateuchocephalinae, the 
classification would suggest that Atrophecaecum excluding Paracanthostomum 
and Ateuchocephala comprises a monophyletic group. Such a notion is not sup- 
ported by any known data. In such cases, because Atrophecaecum lobacetabu- 
lare possesses traits intermediate between Atrophecaecum and Paracanthosto- 
mum plus Ateuchocephala, the "gap" between the taxa no longer exists, and 
existed originally only as an artifact of sampling error. Insofar as gaps represent 
either large numbers of character differences, which may be an artifact of data 
type or of sampling, or large-scale character differences, which may also be an 
artifact of data type, information concerning such occurrences should not be a 
basis for classification. Such occurrences are represented in every cladogram but 
do not alter the branching pattern or classification produced. A cladistic classi- 
fication does not indicate directly such gaps, but the diagnoses for which the 
taxon names stand do. 

Confusion in diagnoses and keys. — It has been suggested (Mayr 1969; Sneath 
and Sokal 1973) that cladistic classifications are not very stable. One aspect of 
that instability would be wholesale changes in diagnoses and keys every time a 
new taxon is included in the classification. It is true that the inclusion of the taxa 
in question within Atrophecaecum requires changes in some diagnoses and keys. 
However, we think the changes are few and minor, especially when compared 
with the number of unnecessary redundancies in diagnoses required if Paracan- 
thostomum and Ateuchocephalus were maintained separately from the other 
acanthostomes. 

Inclusion of Paracanthostomum and Ateuchocephalus in the Acanthostominae 
requires the following changes (in italics) in the subfamilial diagnosis presented 
by Brooks (1980): 

Acanthostominae Poche, 1926 

Diagnosis (emended from that of Brooks 1980). — Body elongate, eyespotted 
or not. Oral sucker terminal, with terminal or subterminal mouth, surrounded by 
uninterrupted single row of spines or lacking spines. Prepharynx and esophagus 
present and variable in length or lacking. Ceca extending to near posterior end 
of body. Acetabulum median, embedded in parenchyma or enclosed in body fold. 
Preacetabular pit present. Postacetabular pit present or lacking. Testes 2, interce- 
cal, tandem or oblique, in hindbody. Seminal vesicle present. Cirrus sac lacking. 
Gonotyl present or lacking. Genital pore immediately preacetabular, not in pre- 
acetabular pit. Ovary spherical or subspherical, pretesticular in hindbody. Sem- 
inal receptacle and Laurer's canal present. Vitellaria follicular, in lateral fields in 
hindbody. Uterus postacetabular, usually not extending postovarially. Eggs em- 



228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

bryonated, not filamented. Excretory vesicle Y- or V-shaped, with post- or pre- 
acetabular bifurcation. Parasites of estuarine and freshwater fishes and reptilians. 
Pantropical. Type-genus: Acanthostomum Looss, 1899. 

Inclusion of Par acanthostomum and Ateuchocephala in Atrophecaecum re- 
quires the following emendations to the generic diagnosis presented by Brooks 
(1980): 

Atrophecaecum Bhalerao, 1940 

Ateuchocephala Coil and Kuntz, 1960:145-150. 
Paracanthstomum Fischthal and Kuntz, 1965:124-136. 

Diagnosis. — Acanthostominae. Tegumental spines uniform in size. Oral sucker 
armed with single uninterrupted row of spines or lacking spines. Mouth terminal 
or subterminal. Prepharynx and esophagus variable in length or one lacking. 
Ceca opening separately and laterally; ceca not atrophied, one cecum atrophied 
or one cecum lacking. Gonotyl lacking. Seminal vesicle coiled. Vitelline follicles 
terminating preovarially; secondary cluster of follicles surrounding testes may be 
present. Seminal receptacle posterodorsal to ovary. Uterine loops preovarian. 
Excretory vesicle Y-shaped, with pre- or po^^acetabular bifurcation. Parasites of 
crocodilians and ophidians. India, Pakistan, Burma, Malaysia, Australia. Type- 
species: Atrophecaecum burminis (Bhalerao, 1926), Bhalerao, 1940. 

The above additions to the diagnoses of Acanthostominae and of Atrophecae- 
cum eliminate the need for separate diagnoses of Paracanthostomum, Ateucho- 
cephala and the Ateuchocephalinae. Additionally, the phylogenetic relationships 
of the taxa involved are represented in the classification and the new classification 
differs little from the previous classification of the acanthostomes with the ex- 
ception of the addition of three new species. Thus, the cladistic system is de- 
monstrably more stable and less ambiguous than any other. The above additions 
do not require any changes in the generic key presented by Brooks (1980) but do 
require a new key to the species oi Atrophecaecum, as follows: 

la. Prepharynx less than 3 times longer than pharynx, one cecum partially 

or completely atrophied, parasites of ophidians 3 

lb. Prepharynx at least 3 times longer than pharynx, ceca not atrophied, 

parasites of crocodilians 2 

2a. Oral spines 18-19 in number slusarskii 

2b. Oral spines 22 in number indicum 

3a. One cecum lacking 4 

3b. Both ceca present 5 

4a. Oral spines 20-22 in number, vitellaria confluent preovarially 

proctophorum 

4b. Oral spines 24-28 in number, vitellaria not confluent preovarially 

asymmetricum 

5a. Oral spines 23 in number, vitelline follicles extending anteriorly to pos- 
terior margin of seminal vesicle pakistanense 

5b. Oral spines 24-29 in number or lacking, vitelline follicles not extending 
anteriorly to posterior margin of seminal vesicle, eggs reaching more 
than 30 ^xm in length 6 



VOLUME 95, NUMBER 2 229 

6a. Cecal bifurcation more than 10% of total body length preacetabular, 

uterine loops occupying less than 50% of total body length simhai 

6b. Cecal bifurcation less than 5% of total body length preacetabular, uterine 

loops occupying more than 50% of total body length 7 

7a. Mouth terminal burminis 

7b. Mouth subterminal 8 

8a. Oral spines present, acetabulum bipartite lobacetabulare 

8b. Oral spines lacking, acetabulum unipartite 9 

9a. Esophagus present, prepharynx lacking, secondary vitelline cluster sur- 
rounding testes, seminal vesicle short marinum 

9b. Esophagus lacking, prepharynx short, secondary vitelline cluster lack- 
ing, seminal vesicle elongate cerberi 

Coil and Kuntz (1960) reported lateral ani for Ateuchocephala marina, but 
Yamaguti (1971) reported no ani after examining a paratype. We examined the 
specimens in question and found lateral ani and slight cecal atrophy. Yamaguti 
further reported a fairly long prepharynx in Paracanthostomum cerberi, but we 
found the prepharynx of the holotype and of specimens reported by Brooks and 
Palmieri (1981) to be as short as or shorter than the pharynx. 

A number of species oi Ateuchocephala have been described in addition to A. 
marina, but their validity and identities remain in question pending acquisition 
of material for study. Therefore, we make only the following new combinations: 
Paracanthostomum cerberi Fischthal and Kuntz, 1965 becomes Atrophecaecum 
cerberi (Fischthal and Kuntz, 1965), n. comb., and Ateuchocephala marina Coil 
and Kuntz, 1960 becomes Atrophecaecum marinum (Coil and Kuntz, 1960), n. 
comb.; Ateuchocephalinae Yamaguti, 1971 becomes a junior subjective synonym 
of Acanthostominae Poche, 1926. 

Inclusion of all cladistic information in classifications produces long unwieldy 
classifications. — Do attempts to preserve all the information in a cladogram nec- 
essarily produce long unwieldy classifications? In order to examine this question, 
we formulated a complete classification of the acanthostomes according to the 
convention of phyletic sequencing (see Wiley 1979) in which every taxon is con- 
sidered the sister-group of all taxa of equivalent position below it. That classifi- 
cation: 

Subfamily Acanthostominae 
Genus Timoniella 
Subgenus 1 

species praeterita 
species imbutiformis 
Subgenus 2 

species incognita 
species scyphocephala 
species unami 
species loossi 
Genus Gymnatrema 
species gymnarchi 
species pambanense 
Genus Proctocaecum 
Subgenus 1 
Infrasubgenus 1 



230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



species coronarium 
species vicinum 
Infrasubgenus 2 
species productum 
species gonotyl 
Subgenus 2 
Infrasubgenus 1 
species nicolli 
Infrasubgenus 2 

species atae sedis mutabilis 
species elongatum sedis mutabilis 
species crocodili sedis mutabilis 
Genus Caimanicola 
species caballeroi 
species pavidus 
species marajoara 
species brauni 
Genus Acanthostomum 
Subgenus 1 

species absconditum 
species knobus 
species niloticum 
species spiniceps 
Subgenus 2 
Infrasubgenus 1 
species gnerii 
species minimum 
species astorquii 
Infrasubgenus 2 

species americanum 
species megacetabulum 
Genus Atrophecaecum 
Subgenus 1 

species indicum 
species slusarskii 
Subgenus 2 

species pakistanense 
Subgenus 3 

species proctophorum 
species asymmetricum 
Subgenus 4 
species simhai 
species burminis 
species lobacetabulare 
species cerberi 
species marinum 



This classification is only 16 lines longer than a classification listing only genera 
and their included species, and only 6 lines longer than a classification using 
subgeneric designations. Thus, a fully-resolved cladistic classification need not be 
substantially longer than a less informative syncretistic scheme. 

This report demonstrates the utility of a cladistic classification scheme as a 
general reference system for helminth systematics. Theoretical claims in support 
of the stability, consistency, predictivity, information content, and applicability 
of Hennigian systematics are upheld in this practical demonstration. We strongly 



VOLUME 95, NUMBER 2 231 

urge helminthologists to investigate the use of such an approach in classifying 
other taxa. 

Acknowledgments 

We express appreciation to Dr. John C. Holmes, University of Alberta, for pro- 
viding the specimens of Atrophecaecum lobacetabulare. Funds for this study were 
provided through a grant to DRB from the Natural, Applied and Health Sciences 
Grant Committee of the University of British Columbia. 

Literature Cited 

Brooks, D. R. 1980. Revision of the Acanthostominae Poche, 1926 (Digenea: Cryptogonimidae). — 

Zoological Journal of the Linnean Society 70:313-382. 
, and J. R. Palmieri. 1981. Six platyhelminths from Malaysian reptiles including Paradisto- 

moidella cerberi n. g., n. sp. (Digenea: Dicrocoeliidae). — Journal of Helminthology 55:39-43. 
Coil, W. H., and R. E. Kuntz. 1960. Three new genera of trematodes from Pacific sea serpents. — 

Proceedings of the Helminthological Society of Washington 27:145-150. 
Fischthal, J. H., and R. E. Kuntz. 1965. Digenetic trematodes of amphibians and reptiles from North 

Borneo (Malaysia). — Proceedings of the Helminthological Society of Washington 32:124-136. 
Mayr, E. 1969. Principles of systematic zoology. — McGraw-Hill, New York. 
Sneath, P. H. A., and R. R. Sokal. 1973. Numerical taxonomy. — W. H. Freeman and Co., San 

Francisco. 
Wiley, E. O. 1979. An annotated Linnean hierarchy, with comments on natural taxa and competing 

systems. — Systematic Zoology 28:308-337. 
Yamaguti, S. 1971. Synopsis of digenetic trematodes of vertebrates. — Keigaku Publ. Co., Tokyo. 



(DRB) Department of Zoology, University of British Columbia, 2075 Wesbrook 
Mall, Vancouver, B.C. V6T 2A9, Canada; (JNC) Harold W. Manter Laboratory, 
Division of Parasitology, University of Nebraska State Museum, W-529 Nebraska 
Hall, Lincoln, Nebraska 68588-0514. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 232-237 

A PARTIAL REVISION OF THE GENUS NOTOMASTUS 

(POLYCHAETA: CAPITELLIDAE) WITH A DESCRIPTION OF 

A NEW SPECIES FROM THE GULF OF MEXICO 



R. Michael Ewing 

Abstract. — An emended diagnosis for the genus Notomastus (Polychaeta: Cap- 
itelUdae) is proposed with two closely related genera, Rashgua Wesenberg-Lund 
and Paraleiocapitella Thomassin, considered as junior synonyms, A new species 
oi Notomastus from the Gulf of Mexico, N. daueri, is described. 



Recent benthic ecological studies in the northern Gulf of Mexico revealed that 
an undescribed capitellid polychaete was a prominent component of the infauna 
of relatively shallow silty sand habitats. Description of this species necessitated 
expansion of the generic diagnosis oi Notomastus Sars, 1851. Close examination 
of the literature indicated that two closely related genera, Rashgua Wesenberg- 
Lund, 1949 and Paraleiocapitella Thomassin, 1970 are sufficiently similar to be 
considered as synonyms oi Notomastus. 

The holotype and one set of paratypes of the new species are deposited in the 
National Museum of Natural History (USNM), Smithsonian Institution, Wash- 
ington, D.C. An additional paratype is deposited in Allan Hancock Foundation 
(AHF), University of Southern California, Los Angeles, California. 

Family Capitellidae Grube, 1862 
Genus Notomastus Sars, 1851, emended 

Rashgua Wesenberg-Lund, 1949:336. 
Paraleiocapitella Thomassin, 1970:86. 

Type-species. — Notomastus latericeus Sars, 1851:199. 

Diagnosis. — Thorax with an achaetous peristomium and 11 setigerous seg- 
ments; epithelium wholly or partly smooth, tessellated, or areolated, Prostomium 
conical or triangular, with or without palpode; eyespots present or lacking. First 
setiger with or without capillary setae in neuropodia; thereafter thoracic segments 
with capillary setae only in both rami or with hooks in neuropodia of last 1-3 
setigers. Nephridial apertures absent, limited to either thorax or abdomen, or 
present on a few segments in both regions. Abdominal neuropodia with hooded 
hooks only; notopodia with hooks only, present throughout the abdomen, re- 
stricted to anterior segments, or completely absent. Branchiae present or lacking; 
if present, as nonretractile, simple expansions or prolongations of noto- and/or 
neuropodia or as eversible branched tufts from notopodial ridges. Pygidium with 
or without appendages. 

Discussion 

Remarks on Rashgua Wesenberg-Lund, 1949: Wesenberg-Lund (1949:336) 
erected the genus Rashgua, characterized as having a thorax consisting of an 



VOLUME 95, NUMBER 2 233 

achaetous peristomium and 11 segments with capillary setae only in both noto- 
and neuropodia and an abdomen with hooded hooks only. Wesenberg-Lund pre- 
sumably distinguished Rashgua from the closely related genus Notomastus on 
the basis that the former genus had abdominal notopodia "perfectly devoid of 
hooks" (1949:337) while the latter had (by original definition) hooks in both rami 
of the abdomen. However, Hartman (1947:415) described a species of Notomas- 
tus, N. lobatus, which agrees with Rashgua in lacking abdominal notosetae. 
Although Hartman (1947:416) did not emend the generic diagnosis oi Notomastus 
to accommodate N. lobatus, her observation that "notopodial tori are believed 
to be absent" was apparently overlooked by Wesenberg-Lund (1949). 

The original description of the genotype oi Rashgua, R. rubrocincta, was based 
on anterior fragments only. This species agrees with N. lobatus in most characters 
but, based strictly on the literature account of R. rubrocincta, the 2 appear to 
differ slightly in the configuration of abdominal tori; these 2 species are the only 
known capitellids completely lacking abdominal notosetae. 

A new species of Notomastus described in this paper has notopodial hooks in 
anterior abdominal segments only. This structure is intermediate between the 
conditions of no abdominal notosetae {Rashgua and Notomastus lobatus) and 
notosetae throughout the abdomen (all other known Notomastus). The presence 
or absence (complete or partial) of notosetae in the abdomen is now recognized 
as a variable character in the genus Notomastus. Therefore, the genus Rashgua 
is herein designated a junior synonym oi Notomastus Sars. Notomastus rubro- 
cinctus (Wesenberg-Lund, 1949) is considered a new combination. 

Remarks on Paraleiocapitella Thomassin, 1970: Thomassin (1970:86) erected 
the genus Paraleiocapitella which is characterized as having a thorax with an 
achaetous peristomium and 11 setigerous segments; an incomplete first setiger 
(i.e. notopodia only) is followed by 9 setigers with capillary setae only in both 
rami and a last thoracic segment with capillary setae only in the notopodia and 
hooded hooks only in the neuropodia; abdominal segments are provided with 
hooks only in both noto- and neuropodia. 

Paraleiocapitella was presumably easily distinguished from the nearly identical 
genus Notomastus based on the presence of hooks in thoracic neuropodia in the 
former genus and by original description, capillary setae only in the thorax of 
Notomastus. However, prior to 1970, descriptions of at least 2 species of No- 
tomastus with thoracic neuropodial hooks had been published although the ge- 
neric description was not technically revised. Notomastus precocis Hartman, 
1960, has hooks only in the last 3 thoracic neuropodia. Hartman (1965) also 
described Notomastus teres which is characterized as having hooded hooks only 
in the last 2 neuropodia. Day (1973), in agreement with Hartman' s expanded 
diagnosis of Notomastus but seemingly unaware of the genus Paraleiocapitella, 
described Notomastus americanus which has a thoracic setal formula identical 
to that of the genotype oi Paraleiocapitella, P. mossambica. 

In the present study, numerous immature specimens representing 3 additional 
species oi Notomastus, N. hemipodus Hartman, 1947, N. lobatus Hartman, 1947, 
and N. daueri, n. sp., were found with hooded hooks only or mixed fascicles of 
capillary setae and hooks in as many as 5 posterior thoracic neuropodia and rarely 
in 1-2 notopodia. With very few exceptions, a distinct pattern of setal develop- 
ment toward the adult arrangement (i.e. thoracic setal formula) was recognized 



234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

in each of these species. Similar conditions were also noted in juveniles of several 
other capitellid genera. The subject of setal development will be discussed in 
greater detail in a forthcoming paper. Tentatively outlined, the replacement of 
hooded hooks by capillary setae in the thoracic neuropodia of Notomastus is as 
follows: 1) juveniles are provided with hooks only in neuropodia of several seg- 
ments in the posterior one-half of the thorax, 2) as the worm grows, hooks are 
apparently lost (shed, broken or resorbed?) from the ventral position in the setal 
fascicle and replaced by capillary setae emerging from the superior position; the 
replacement of setae progresses until all hooks in a "changing" fascicle are re- 
placed by capillary setae and the process continues in a relatively predictable 
manner toward the posterior thoracic segments until the adult condition is 
reached. 

In summary, the presence and/or location of hooded hooks in thoracic neuro- 
podia of both juveniles and adults is recognized as a highly variable character in 
the genus Notomastus and perhaps many other capitellid genera; thus the genus 
Paraleiocapitella Thomassin is herein designated a junior synonym of Notomas- 
tus Sars. Notomastus mossambicus (Thomassin, 1970) will be considered a new 
combination pending personal examination of type-material. 

Notomastus daueri, new species 
Fig. la-g 

Material examined. — LOUISIANA: Gulf of Mexico: Approx. 29.3 km SSW 
Grand Isle: 28°56'12"N, 90°04'07"W, 27.7 m, silty clay, holotype (LW&F, col., 
16 April 1980, USNM 71442), 2 paratypes (USNM 71443), 18 specimens; 8 spec- 
imens, 21 Aug. 1980, same location; 2 specimens, 8 Sept. 1980, same location. 
Approx. 33.9 km SSW Grand Isle: 28°53'06"N, 90°0r30"W, 33.5 m, clayey silt, 
1 paratype (LW&F, col., 8 Sept. 1980, AHF Poly 1361). Approx. 37.0 km SSW 
Grand Isle: 28°51'06"N, 90°04'21"W, 33.2 m, 1 specimen, 21 Aug. 1980. MISSIS- 
SIPPI: Gulf of Mexico: lEC 732 MO Sta. 014, approx. 5.9 km S Ship Island pass, 
30°10.32'N, 88°55.00'W, 10 m, muddy sand, 1 specimen (lEC, col., June 1980); 
Mississippi Sound: sta. 043, approx. 3.6 km N Petit Bois Island, 30°14.48'N, 
88°25.63'W, 5.6 m, muddy sand, 1 specimen (Vittor & Assoc, col., 22 Oct. 1980). 

Description. — Length of largest complete specimen (holotype) approximately 
65 mm, width 1.1 mm, 234 setigerous segments. Lengths of 7 additional complete 
specimens ranged from 40 to 52 mm, widths 0.3 to 0.9 mm, with up to 197 setigers. 

Color light tan to brown in alcohol. Thorax slightly inflated through setigers 4- 
5, surface epithelium faintly areolated; following thoracic segments nearly smooth 
except for ventral biannulation. Abdominal epithelium smooth throughout, 

Prostomium short, broadly triangular in dorsal view, with 2 inconspicuous nu- 
chal slits near posterior border; eyespots absent. Achaetous peristomium slightly 
wider than long, approximately XVi to 2 times as long as following segment; 
eversible pharynx bulbous, coarsely papillated on proximal two-thirds, smooth 
distally. Thoracic setigers about 3 times as wide as long. Anteriormost notopodia 
dorsolateral, well separated, but approach middorsally by setigers 6-7; neuropodia 
ventrolateral in position throughout thorax. Anterior 10 setigers with 15-20 cap- 
illary setae per fascicle in both rami; last thoracic segment (setiger 11) with cap- 
illary setae only in notopodia and fascicles of 12-20 hooded hooks only in neu- 
ropodia (Fig. la). Nephridial apertures (1 pair on each segment) located in 



VOLUME 95, NUMBER 2 



235 




mm 



I .0 mm 




0.5mm 





0. 1 mm 



Fig. 1. Notomastus daueri: a, Lateral view of anterior end showing thorax and first 2 abdominal 
segments; b, Dorsolateral view of midabdominal segments; c-d, Lateral and dorsal views of posterior 
abdominal segments; e, Dorsal view of pygidium; f-g, Lateral and frontal views of neuropodial hooded 
hook from midabdomen. 



segmental groove between each of last 4 thoracic segments. Lateral organs pres- 
ent on all thoracic setigers as a minute pore between noto- and neuropodium; 
appearing as a small ovoid structure just above neuropodial tori in abdominal 
segments, present at least through midabdomen but becoming increasingly less 
conspicuous moving posteriorly. Transition from thorax to abdomen marked by 
setael change in notopodia from capillary setae to hooks, increase in number of 
neurosetae per fascicle and slight broadening of segments. 

Anterior abdominal segments of approximately same length as those of the 
posterior thorax, gradually lengthening to the midabdominal region where they 
are Wi to 2 times as long as wide; thereafter segments becoming increasingly 
shorter. 

Branchial lamellae first emerging from anterior abdominal segments (about se- 
tiger 60 in holotype) as short posteriolateral swellings of the notopodia, increasing 
in length posteriorly (Fig. lb), developing into large triangular projections which 
overlap the following segment in the posterior region (Figs. Ic-d). 

Abdominal parapodia with multidentate hooded hooks only; hooks consisting 



236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

of a main fang surmounted by 3 teeth in a triangular arrangement and a second 
row of 4-5 smaller denticles (Figs, le-f), similar in structure to those in the last 
thoracic neuropodia. Anteriormost notopodia with 15-20 setae per fascicle on 
short, slightly elevated tori; notosetae decreasing rapidly in number posteriorly, 
disappearing completely 5-15 segments after the appearance of branchial lobes. 
Neurosetal fascicles on well-defined glandular ridges at posterior margin of each 
segment, extending from lateral to near midventral position; neurosetae increas- 
ing in number posteriorly to as many as 40 hooks per fascicle by midabdomen, 
then decreasing in number through posterior abdomen to 1-2 per fascicle in last 
8-10 segments; noticeable reduction in size (and perhaps number) of denticles 
above the main fang of hooks in far posterior region with some hooks appearing 
unidentate under oil immersion in last few segments. Neuropodial tori of latter 
segments enlarged at posterior margin as a shallow, somewhat cup-shaped pro- 
cess (dorsal view) which may also be branchial in function. 

Pygidium funnel-shaped with single digitate median cirrus (Fig. Ig). 

Remarks. — Variations from the thoracic setal arrangement of adults were ob- 
served in numerous small specimens (juveniles?) of Notomastus daueri. 

Notopodia of setigers 10 and 11 were rarely found with mixed fascicles of 
capillary setae and hooks. Neuropodia of setigers 7-11 may have capillary setae 
only, mixed setal fascicles, hooded hooks only, or combinations of these in a 
given segment. 

Notomastus daueri differs from most other species of the genus in having some 
thoracic neuropodia with hooded hooks instead of capillary setae only. The 4 
known species of Notomastus with hooks in one or more thoracic neuropodia, 
N. precocis Hartman, 1960, N. teres Hartman, 1965, N. mossambicus (Thomas- 
sin, 1970; new combination proposed in this paper), and N. americanus Day, 
1973, all have notopodia only on setiger 1 whereas N. daueri has a complete first 
setiger. 

Notomastus daueri is known to occur in the northern Gulf of Mexico off Lou- 
isiana and Mississippi in shallow subtidal muddy sands. 

Etymology. — This species is named in honor of Dr. Daniel M. Dauer, my good 
friend and former graduate advisor, in an attempt to express my sincere appre- 
ciation for his unselfish guidance over the past several years. 

Acknowledgments 

Some of the specimens examined in this study were collected at stations selected, 
sampled and processed under the supervision of John deMond of Louisiana Wild- 
life and Fisheries under contract to LOOP, Inc. Special thanks are extended to 
the Seafood Division (LW&F) and A. J. Heikamp (LOOP, Inc.) for permission 
to use this material. Additional specimens were obtained from Interstate Elec- 
tronics Corporation (lEC) under contract no. 68-01-4610 from the Environmental 
Protection Agency. 

Barry A. Vittor & Associates, Inc. provided the laboratory space and materials 
used in this study and I sincerely thank the following employees for their con- 
tributions: Caroline Coker, Michael Milligan, Donna Valentine and Jewel Walt- 
man assisted in assembling the collection. Jerry Gathof, Gary Goeke, and Paul 
Wolf commented on the manuscript. Figures were prepared for press by Ben 
Jordan. Several drafts of the paper were typed by Dottie Cotton. 



VOLUME 95, NUMBER 2 237 

I also thank Susan Williams of Allan Hancock Foundation for arranging loans 
of type-material. 

My studies of capitellid polychaetes have benefitted greatly from discussions 
with Dr. Kristian Fauchald of the Smithsonian Institution. I am especially ap- 
preciative of his many helpful suggestions and of his constructive criticism of 
this manuscript. 

Literature Cited 

Day, J. H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North 

Carolina. — National Marine Fisheries Service Circular 375:1-140. 
Grube, A. E. 1862. Noch ein Wort iiber die Capitellen und ihre Stellung im Systeme der Anneliden. — 

Archiv fiir Naturgeschichte (Berlin) 28:366-378. 
Hartman, O. 1947. Polychaetous annelids. Pt. 7. Capitellidae. — Allan Hancock Pacific Expeditions 

10:391^81. 
. 1960. Systematic account of some marine invertebrate animals from the deep basins off 

Southern California. — Allan Hancock Pacific Expeditions 22:69-216. 
. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other 

North Atlantic areas. — Allan Hancock Foundation Publications Occasional Paper 28:1-378. 
Sars, M. 1851. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Loften og Fimmarken. — 

Nyt Magazin for Naturvidenskabeme Christiania 6:121-211. 
Thomassin, B. 1970. Contribution a I'etude des polychetes de la region de Tulear (SW de Mada- 
gascar). Sur les Capitellidae des sables corralliens. — Recueil des Travaux de la Station Marine 

d'Endomme, Supplement 10:71-101. 
Wesenberg-Lund, E. 1949. Polychaetes of the Iranian Gulf. — Scientific Investigations in Iran 4:247- 

400. 

Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Road, Mobile, Alabama 
36609. Present address: Department of Biological Sciences, Old Dominion Uni- 
versity, Norfolk, Virginia 23508. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 238-250 

SOME SPECIES OF ONUPHIS (POLYCHAETA: ONUPHIDAE) 
FROM THE ATLANTIC OCEAN 

Kristian Fauchald 

Abstract. — Five new species of Onuphis (Polychaeta: Onuphidae) are de- 
scribed. Three, O. {Nothria) australatlantica, O. (N.) heterodentata, and O. 
(N.) lithobiformis come from southern Atlantic waters; the other two, O. {On- 
uphis) declivorum, and O. (O.) texana are from the east coast of the United 
States. 



Introduction 

A review of material of onuphids deposited in the collections of the National 
Museum of Natural History, Smithsonian Institution, has revealed the presence 
of five previously undescribed species. Part of the material was previously treated 
by Hartman (1967) in her study of Antarctic polychaetes; other specimens come 
from benthic surveys along the east coast of the United States. 

Generic definitions and terminology are those in Fauchald (1980). 

Family Onuphidae Kinberg, 1866 

Genus Onuphis Audouin and Milne Edwards, 1833 

Subgenus Nothria Malmgren, 1866 

Onuphis (Nothria) australatlantica, new species 

Fig. 1, Table 1 

Nothria cf. conchylega. — Hartman, 1967:91 (not M. Sars, 1835:61). 

Material examined. — Antarctic Ocean, off South Shetland Islands, 3768-3816 
m, Menzies- trawl, 2 August 1962, Eltanin Sta. 129 (17 paratypes, USNM 58296). 
Antarctic Ocean, off South Shetland Islands, 3138-3239 m, Blake-trawl, 8 Sep- 
tember 1963, Eltanin Sta. 722 (holotype, USNM 58195, one paratype, USNM 
67488). 

Description. — The holotype is an incomplete specimen with 29 setigers that is 
23 mm long and 2.75 mm wide, with parapodia; it is a mature female with large 
eggs in the body cavity. The body is dorsoventrally flattened with the parapodia 
attached at the margins. All types are pale pink and lack color patterns. Eyes are 
absent. The holotype is described in detail below; a summary of information for 
all types is given in Table 1. 

The prostomium (Fig. lb) is rounded frontally and is wider than long. The 
frontal palps are shorter than the length of the prostomium. The outer lateral 
occipital antennae reach setiger 1 , the inner lateral antennae reach setiger 4 and 
the median antenna reaches setiger 7. The occipital ceratophores are short and 
have up to 3 rings. Branchiae are absent. 

The first 2 setigers (Fig. la) are enlarged. The first parapodia are directed 
forward and laterally so that the morphologically anterior face of each parapodium 
is medial. In the first pair of parapodia (Fig. li-j) the flattened, auricular presetal 



VOLUME 95, NUMBER 2 



239 




Fig. 1. Onuphis (Nothria) australatlantica (holotype, USNM 58195): a, Anterior end, lateral view; 
b. Anterior end, dorsal view; c, Pseudocompound hook, second setiger; d, Intrafascicular hook, 
median setiger; e, Pectinate seta, median setiger; f, Pseudocompound hook, first setiger; g. Second 
parapodium, anterior view; h. Second parapodium, posterior view; i, First parapodium, posterior 
view; j, First parapodium, anterior view. All scales are 1 mm except where otherwise indicated. 



240 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Summary statistics for Onuphis {Nothria) australatlantica. 





Range 


Mean SD 


V 


N 


Occipital antennae 










Outer lateral reach # 


1 


invariant 




19 


Inner lateral reach # 


4-8 


5.89 1.08 


1.16 


18 


Median reaches # 


6-10 


8.14 1.23 


1.52 


14 


# of rings 


2-3 


2.24 0.44 


0.19 


19 


Ventral cirri cirriform to setiger # 


2 


invariant 




19 


Pseudocompound hooks to setiger # 


2-3 


2.05 0.23 


0.05 


19 


Intrafascicular hooks from setiger # 


9-11 


9.95 0.71 


0.50 


19 



lobe enfolds the dorsal side of the superiormost setae. The digitiform postsetal 
lobe is directed posteriorly. The tapering dorsal cirrus is about twice as long as 
the postsetal lobe. The ventral cirrus is similar to the postsetal lobe in size and 
shape (the postsetal lobe projects posteriorly and is thus foreshortened in Fig. 
li). 

The second parapodia (Fig. Ig-h) are shorter than the first and directed later- 
ally. The presetal lobe is less than half as wide as that of the first setiger, but is 
still distinctly flattened and auricular. The postsetal lobe is about as long as that 
of the first setiger, but is tapering from the base, rather than digitiform. The dorsal 
cirrus resembles the one in the first setiger. The ventral cirrus is very short and 
sharply tapering. 

All other parapodia resemble each other; the presetal lobes are completely 
reduced and the ventral cirri are replaced by glandular pads. The postsetal lobes 
become increasingly threadlike towards the posterior end, but are distinct in all 
setigers present; the dorsal cirri also become increasingly slender towards the 
posterior end. 

Pectinate setae, limbate setae, and anterior pseudocompound and intrafasci- 
cular hooks are present. Compound spinigers, large hooks, and subacicular hooks 
are absent. Pectinate setae (Fig. le) are present from the second setiger; each 
seta is distally scoop-shaped and has about 15 teeth. About 12 pectinate setae are 
present per parapodium in the anterior third of the body. The limbate setae have 
extremely long shafts with a short, slightly geniculate limbation distally. The 
pseudocompound hooks of the first setiger (Fig. If) are very thick, indistinctly 
bidentate, and have short, blunt hoods. The hooks of the second setiger are less 
than half as wide as those of the first setiger (Fig. Ic); the hinge-line is distinct 
and each hook is clearly bidentate with a short, blunt hood. Pseudocompound 
hooks are present in the first 2 setigers only. A pair of intrafascicular hooks is 
present in each setiger from setiger 10. The intrafascicular hooks are oriented 
parallel to the acicula and the limbate setae, and are about as long as the latter. 
Each intrafascicular hook is bidentate distally and the head of the hook is tilted 
so that both teeth are axial. 

The maxillary formula (examined in the paratype from USNM 67488) is 1 + 1 , 
8+10, 3+0, 4+9 and 1 + 1; the unpaired maxilla III has 2 poorly defined basal 
teeth and a large distal fang. 

The tubes are quadrangular in cross-section. The sides of the tubes that cor- 



VOLUME 95, NUMBER 2 241 

respond to the dorsal and ventral surface of the worm are somewhat more flat- 
tened than the 2 lateral sides. The inner lining is tough; the outer surface is 
covered with black, volcanic sand and other large particles that are not arranged 
in a recognizable pattern. 

Only one previously described species of Onuphis (Nothria) combines pro- 
longed first parapodia with a lack of branchiae. Onuphis (N.) text or (Hartman 
and Fauchald, 1971:78) was described from deep water in the Atlantic Ocean. It 
is a very small species; a mature female with 15 setigers is about 2.5 mm long, 
and has extremely long, slender first parapodia projecting well beyond the tip of 
the prostomium. Onuphis {N.) australatlantica is a much larger and more massive 
species; a mature female is 23 mm for 29 setigers, and the first parapodia, while 
enlarged, do not project beyond the tip of the prostomium. 

These specimens were originally identified as Nothria cf. conchylega by Hart- 
man (1967); they differ clearly from Onuphis {Nothria) conchylega Sars, 1835, in 
that they lack branchiae, present in the latter. 

Etymology. — The specific name, australatlantica, refers to the localities in the 
southern Atlantic Ocean from which material of the species was collected. 

Distribution. — Onuphis (N.) australatlantica is known from two localities in 
abyssal depths in the southern Atlantic Ocean. 

Onuphis (Nothria) heterodentata, new species 
Fig. 2, Table 2 

Nothria liridescens. — Hartman, 1967:91 (in part, not Johnson, 1901:408). 

Material examined. — South Atlantic Ocean, off Falkland Islands, 587-595 m, 
rock dredge, 3 December 1962. Eltanin Sta. 338 (holotype, USNM 58299, 13 
paratypes, USNM 67489). 

Description. — The holotype is an incomplete specimen with 46 setigers, is 18 
mm long and 1.5 mm wide, with parapodia. Other specimens are up to 24 mm 
long for 55 setigers; none are complete. The anterior end of the specimens is 
cylindrical; the dorsum is flattened and the ventrum is curved in median and 
posterior setigers. The holotype (Fig. 2c) has lost the styles on both the inner 
lateral and median occipital antennae. The outer lateral antennae reach setiger 2 
and the ceratophores are distinctly longer than the styles. The inner lateral cer- 
atophores, which are the longest, have up to 13 rings. In the paratypes, the inner 
lateral antennae reach setiger 7 and the median antenna reaches setigers 4-5 
(Table 2). The short frontal palps are ovate; eyes are absent. 

Branchiae are present from setiger 2 in the holotype and from setigers 1 or 2 in 
the paratypes. If branchiae are present from setiger 1, the first branchiae may be 
missing from one side of the animal. Branchiae are present to the end of all 
fragments; each is simple and strapUke, and flattened, especially in posterior 
setigers. 

The first 2 pairs of parapodia are directed anteriorly. Ventral cirri are cirriform 
in the first 5 setigers (Fig. 2f) and the postsetal lobes are digitiform in the first 11 
setigers. The first 2 parapodia (Fig. 2a) have distinct contraction folds at the bases 
of the dorsal cirri and rounded presetal and acicular lobes. The postsetal lobes 
have thickened, pad-shaped bases. Median and posterior parapodia are barely 
raised ridges with the setae emerging in a crescentic opening ventrally. The ven- 



242 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 2. Onuphis {Nothria) heterodentata (holotype, USNM 58299): a, Second parapodium, an- 
terior view; b, Pectinate seta, median setiger; c, Anterior end, dorsal view; d-e, Superior and inferior 
pseudocompound hook, second parapodium; f. Anterior end, lateral view; g-h, Superior and inferior 
pseudocompound hooks, third parapodium. All scales are 1 mm except where otherwise indicated. 



tral glandular pads that replace the ventral cirri are very large and extend beyond 
the tips of the subacicular hooks in most setigers. 

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres- 
ent. Compound spinigers and large hooks are absent. Limbate setae are most 
common in anterior parapodia except in the first 2 where only 1 or 2 are present. 
One or 2 pectinate setae (Fig. 2b) are present in each parapodium; each pectinate 
seta is slightly oblique distally, forming a 120° angle with the long axis of the seta, 
and has 13-14 teeth; the 2 marginal teeth are not prolonged. Tridentate pseudo- 
compound hooks (Fig. 2d-e, g-h) with short hoods are present in the first 3 or 
4 parapodia. Only a single hook is present in setiger 4. The middle tooth projects 
beyond the other teeth. The proximal tooth is broadly conical, the middle tooth 
is curved, nearly beaked, and the distal tooth is thick and abruptly tapering 



VOLUME 95, NUMBER 2 



243 



Table 2. — Summary of statistics of Onuphis (Nothria) heterodentata. 





Range 


Mean 


SD 


V 


N 


Occipital antennae 












Outer lateral reach # 


2 


invariant 


t 




12 


Inner lateral reach # 


5-10 


7.17 


2.14 


4.57 


6 


Median reaches # 


3-6 


4.67 


1.53 


2.33 


3 


# of rings 


12-13 


12.20 


0.42 


0.18 


10 


Branchiae first present from setiger # 


1-2 


1.60 


0.52 


0.27 


10 


Ventral cirri cirriform to setiger # 


5 


invariant 




14 


Pseudocompound hooks present to 












setiger # 


4 


invariant 




14 


Subacicular hooks present from 












setiger # 


12-14 


12.50 


0.65 


0.42 


12 



distally. Two bidentate subacicular hooks are present in each parapodium from 
setiger 12. 

The maxillary formula (investigated in 2 paratypes) is 1 + 1, 9+9, 8—9+0, 
7-8+9-10 and 1 + 1. 

Onuphis (N.) heterodentata resembles O. (N.) holobranchiata Marenzeller 
(1879:132), O. (N.) iridescens (Johnson, 1901:408), O. (TV.) opalina (Verrill, 
1873:102), and O. (N.) lithobiformis (see below). All these species have simple 
branchiae starting on setiger 1 and tridentate pseudocompound hooks exclusively. 
Onuphis heterodentata differs from the other 4 species in that the median tooth 
of the pseudocompound hooks is larger and projects further than the other 2 
teeth; in all the other species the middle tooth is either of the same size as the 
proximal tooth or intermediate in size between a long distal tooth and a short 
proximal tooth. The relationships between the other 4 species are discussed be- 
low. 

Etymology. — The specific name refers to the different shapes of the teeth in 
the pseudocompound hooks. 

Distribution. — Onuphis (N.) heterodentata is known from a single locality in 
slope-depths off the Falkland Islands. 



Onuphis {Nothria) lithobiformis, new species 

Fig. 3 

Nothria liridescens. — Hartman, 1967:91 (in part, not Johnson, 1901:408). 

Material examined. — South Atlantic Ocean, off Falkland Islands, from 53°08'S, 
59°23'Wto 53°07'S, 59°2rW, 578-567 m, Menzies-trawl, 3 December 1962, Eltanin 
Sta. 340 (holotype, USNM 58300, 4 paratypes, USNM 67490). South Atlantic 
Ocean, Straits of Magellan, from 53°48'S, 70°53'W to 53°45'S, 70°53'W, 485 m, 
Blaketrawl, 6 February 1964, Eltanin Sta. 963 (2 paratypes, USNM 58301). 

Description. — All specimens are incomplete; the holotype consists of 58 seti- 
gers, is 21 mm long and 1.20 mm wide, with parapodia. The anterior part of the 
body (Fig. 3e) is cylindrical and the median and posterior parts are dorsally 
flattened. Color patterns and eyes are absent. The prostomium (Fig. 3a) is a short, 
rounded lobe (not clearly visible in the illustration) with a pair of triangular frontal 



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Fig. 3. Onuphis (Nothria) lithobiformis (holotype, USNM 58300): a, Anterior end, dorsal view; 
b, Pseudocompound hook, third parapodium; c. Pectinate seta, median parapodium; d. Third para- 
podium, anterior view; e. Anterior end, lateral view. All scales are 1 mm except where otherwise 
indicated. 



palps. The ceratophores of the occipital antennae are all strongly ringed and reach 
well beyond the edge of the prostomium. The inner lateral ceratophores are the 
longest and have up to 13 rings. The outer lateral antennae reach setiger 2, the 
inner lateral antennae reach setigers 8-12 and the median antenna reaches setigers 
3-7. The outer lateral ceratostyle is clearly shorter than its ceratophore. 

Branchiae are present from the first setiger in most specimens. The occurrence 
of branchiae on the first 3 setigers may be irregular; in the holotype branchiae 
are absent on the second setiger on one side and first present from the third 
setiger on the other side. Branchiae are present to the end of all the specimens; 
each branchia is simple and straplike and reaches nearly halfway across the dor- 
sum in median and posterior setigers. 

The first parapodia (Fig. 3d) have rounded acicular lobes; the presetal lobes 
follow the outlines of the acicular lobes closely. Distinct contraction folds are 
present at the bases of the dorsal cirri. Digitiform postsetal lobes are present in 
the first 11-12 setigers. Cirriform ventral cirri are present in the first 4 or 5 
setigers; setigers 5-6 have shortened, blunt cirri; all following setigers have glan- 
dular pads replacing the ventral cirri. 



VOLUME 95, NUMBER 2 245 

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres- 
ent. Compound spinigers and large hooks are absent. Limbate setae are present 
in all setigers, but are most numerous in anterior setigers. Each of the pectinate 
setae (Fig. 3c) is distally oblique and has about 12 teeth; there are 1 or 2 pectinate 
setae in a parapodium in median and posterior setigers. Tridentate pseudocom- 
pound hooks (Fig. 3b) with short, blunt hoods are present in the first 4 setigers. 
The 2 proximal teeth are similar in size and orientation in all hooks; both teeth 
are narrowly conical with blunt tips. Bidentate subacicular hooks are present 
from setigers 9-15. 

The maxillary formula (investigated in one paratype from USNM 67490) is 1 + 1 , 
6+8, 4+0, 6+9 and 1 + 1; in the unpaired maxilla III the distal tooth is a large 
fang, separated from the other 3 teeth by nearly half the length of the maxilla. 

The holotype had a flimsy tube, consisting of a thin, fragile inner Uning sparsely 
covered with sand-grains. 

Onuphis {N.) lithobiformis resembles O. (N.) holobranchiata Marenzeller 
(1879:132), O. (N.) iridescens (Johnson, 1901:408), O. (N.) opalina (Verrill, 
1873:102), and O. (N.) heterodentata, new species (see above) in that all 5 species 
have simple branchiae present from the first setiger and have tridentate pseudo- 
compound hooks exclusively. The differentiation of O. heterodentata from the 
other 4 species was indicated above. Onuphis holobranchiata differs from the 
remaining 3 species in that it has the outer lateral ceratophores longer than the 
other ceratophores and in that it has digitiform postsetal lobes at least through 
setiger 33. The inner lateral ceratophores are the longest in the other 3 species 
and none of them have digitiform postsetal lobes in more than 16 setigers. The 
middle tooth of the pseudocompound hooks is by far the widest, and is bluntly 
conical in O. opalina and O. iridescens; the proximal and the middle tooth are 
of the same size and shape in O. lithobiformis , in which no teeth are bluntly 
conical. 

Etymology. — The general shape of the body resembles that of a centipede, 
Lithobius being the name of a well-known genus in that group. 

Distribution. — Onuphis lithobiformis is known from upper slope-depths in the 
southern Atlantic Ocean. 

Onuphis {Onuphis) declivorum, new species 

Fig. 4 

Material examined.— Ai\2intic Ocean, off New Jersey, 38°45.2'N, 73°01.0'W, 
350 m, R.V. Pierce Sta. J1-BLM-04B (holotype, USNM 57639). 

Description. — The holotype is an incomplete specimen with 39 setigers, is 17 
mm long and 1.6 mm wide, with parapodia. The specimen is a female with large 
oocytes in the body-cavity. The body is cyUndrical throughout and lacks a color 
pattern. Eyes are absent. 

The prostomium (Fig. 4a) is shorter than wide and has a pair of elongate frontal 
palps directed ventrally. The 5 occipital ceratophores are all longer than the length 
of the prostomium. The maximal number of rings is 5 and the rings are limited 
to the proximal half of each ceratophore. The inner lateral and median cerato- 
styles had loosened from the ceratophores, but were retained in a mucus coat 
surrounding the specimen. They are all of the same length and would have 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Onuphis {Onuphis) declivorum (holotype, USNM 57639): a, Anterior end, dorsal view; 
b-d, Inferior, median and superior pseudocompound hooks, second parapodium; e. Second parapo- 
dium, anterior view; f, Sixth parapodium, anterior view; g. Ninth parapodium, anterior view; h, 
Anterior end, lateral view. All scales are 1 mm except where otherwise indicated. 



reached about setiger 6, if attached. The outer lateral ceratostyles are about as 
long as their ceratophores; the outer lateral antennae reach setiger 2. 

The peristomium bears a pair of short, slender peristomial cirri. The first 5 
pairs of parapodia (Fig. 4h) are directed forward and slightly ventrally. The para- 
podial bases are well developed in the first 3 parapodia and become reduced 
posteriorly. In the first several parapodia (Fig. 4e-f) the acicular lobes are evenly 
rounded distally and have a contraction fold running across the frontal face at 
about the bases of the dorsal cirri. The postsetal lobes are at least as long as the 
base of the parapodium and are tapered. Ventral cirri are long and digitiform in 



VOLUME 95, NUMBER 2 247 

the first 6 parapodia and abruptly replaced by glandular pads from setiger 7. The 
dorsal cirri increase in length from the first through the fifth setiger where they 
are longer than half the width of the body. From about setiger 10 they remain of 
about the same length, but become increasingly slender posteriorly. The postsetal 
lobes decrease in size from the first setiger, but are distinct as short, conical 
tubercles, even in the last setigers present. 

Branchiae are present from setiger 7 to the end of the fragment. The first 
branchia is simple; all others are branched (Fig. 4g) with up to 4 long, slender 
branchial filaments. The filaments are as long as the dorsal cirri or longer in all 
except the first branchial setiger. 

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres- 
ent. Compound spinigers and large hooks are absent. Tridentate pseudocom- 
pound hooks (Fig. 4b-d) with short hoods are present in the first 5 setigers; all 
teeth are short and very strongly curved. Bidentate subacicular hooks are present 
from setiger 18. Each pectinate seta is distilly transverse and has about 15 teeth. 
Limbate setae are present in all setigers, but are especially prominent in superior 
fascicles in anterior parapodia. 

The structure of the maxillae was not observed in the single specimen available. 

Onuphis (O.) declivorum resembles O. {O.) acapulcensis Rioja (1944:139), O. 
(O.) aucklandensis Augener (1924:418), and O. (O.) vexillaria Moore (1911:266). 
All 4 species have branched branchiae starting from an anterior setiger other than 
the first and have tridentate pseudocompound hooks exclusively. Of the 4 species, 
O. vexillaria has large hooks in some anterior setigers; the other 3 species lack 
such hooks. The 3 remaining species can be differentiated on a set of characters. 
Onuphis acapulcensis has branchiae from setiger 6 with up to 12 branchial fila- 
ments, the occipital ceratophores are strongly ringed, and ventral cirri are cirri- 
form in 5 setigers. Onuphis aucklandensis has branchiae from setiger 2 with up 
to 5 branchial filaments, the occipital ceratophores are strongly ringed, and ven- 
tral cirri are cirriform in the first 7 setigers. Onuphis declivorum has branchiae 
from setiger 7 with up to 4 filaments, the occipital ceratophores are poorly ringed, 
and ventral cirri are cirriform in the first 6 setigers. These differences are well- 
supported by numerical studies on material of 2 of the other species with which 
O. declivorum was compared (O. aucklandensis and O. vexillaria). 

Etymology. — declivorum, living on the slope, refers to the depth at which the 
holotype was collected. 

Distribution. — Onuphis declivorum is known from one locality in the Atlantic 
Ocean off New Jersey in upper slope-depths. 

Onuphis {Onuphis) texana, new species 
Fig. 5 

Material examined. — Gulf of Mexico, off Brownsville, Texas, 26°16'29.6"N, 
97°10'16.3"W, 13 m, slightly silty sand (87% sand), coll. Thomas Calnan, sta. 243 
(holotype, USNM 67491, 2 paratypes, USNM 67492). 

Description. — All 3 specimens are incomplete. The holotype consists of 39 
setigers, is 10.8 mm long and 0.75 mm wide, with parapodia. The anterior part 
of the body is cylindrical, the posterior part is dorsally flattened. The anterolateral 
edges of each of the first 10 setigers has a dark brown, nearly black, pigment bar, 
visible from both the dorsal and the ventral side. The peristomium has a pair of 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 5. Onuphis (Onuphis) texana (holotype, USNM 67491): a, Anterior end, dorsal view; b, 
Pseudocompound hook, second parapodium; c. Second parapodium, anterior view; d. Anterior end, 
lateral view; e. Pectinate seta, median parapodium. AU scales are 1 mm except where otherwise 
indicated. 



dark brown lateral bars, and each of the ceratophores has 3 equally spaced brown 
rounded spots. 

The prostomium (Fig. 5a) is narrowed anteriorly and has a pair of ovate frontal 
palps. A pair of black eyespots is near the frontal bases of the outer lateral 
occipital antennae and another, larger and somewhat diffuse pair of eyespots 
(Fig. 5d) is near the outer base of the inner lateral occipital antennae. All cera- 
tophores reach well beyond the edge of the prostomium. The inner lateral cera- 
tophores are the longest and have up to 18 rings. The styles of the outer lateral 
antennae are much shorter than their ceratophores; the outer lateral antennae 
reach setiger 2. The inner lateral antennae reach setigers 10-12 and the median 
antenna reaches setiger 5 in all 3 specimens. 

Branchiae are present from the first setiger to the end of the fragment. The first 
18-20 pairs are simple, the next 10 pairs are bifid and, in the last several segments, 
either 2 or 3 branchial filaments are found in an irregular arrangement. 

The first 2 pairs of parapodia are directed forward. The acicular lobe (Fig. 5c) 
is conical and the presetal lobe follows the outline of the acicular lobe closely, 
except on the ventral side where it leaves the bases of the setae exposed. The 
postsetal lobes are digitiform in the first 10 setigers; thereafter they are reduced 



VOLUME 95, NUMBER 2 249 

to short, truncate knobs that remain distinct in the remainder of the fragments. 
Ventral cirri are cirriform in the first 6 setigers and are replaced by transversely 
elongated glandular pads in all setigers thereafter. 

Limbate and pectinate setae, pseudocompound hooks, and subacicular hooks 
are present. Compound spinigers and large hooks are absent. Limbate setae are 
present in all setigers, but are most common in anterior setigers, starting at setiger 
3. Median and posterior setigers each have a single, narrow pectinate seta (Fig. 
5d) which is distally transverse and has 8-10 teeth. Bidentate pseudocompound 
hooks (Fig. 5b) with short, blunt hoods are present in the first 2 setigers only. 
Bidentate subacicular hooks are first present from setiger 10. 

The maxillary formula (investigated, in one paratype) is 1 + 1, 8+9, 9+0, 5+9 
and 1 + 1. 

Tubes were absent. 

Onuphis (O.) texana resembles O. (O.) dibranchiata Willey (1905:272) in 
having poorly developed branched branchiae present from the first setiger; it 
differs in that it has exclusively bidentate pseudocompound hooks; O. (O.) di- 
branchiata has both bi- and tridentate kinds. Other species of Onuphis with 
branched branchiae present from the first setiger include O. (O.) chinensis Us- 
chakov and Wu (1962b: 118) and O. (O.) fukianensis Uschakov and Wu 
(1962a: 93); again both can be separated from O. (O.) texana by having both bl- 
and tridentate pseudocompound hooks. 

A unique feature of O. (O.) texana appears to be the presence of pseudocom- 
pound hooks in only 2 setigers; in all other species of Onuphis such hooks are 
present in at least 3 setigers. 

Etymology. — The species is known from off Brownsville, Texas, hence texana. 

Distribution. — The species is known from a single locality in shelf-depths in 
the Gulf of Mexico. 

Acknowledgments 

I would like to thank Mr. Thomas R. Calnan, Bureau of Economic Geology, 
University of Texas at Austin, for sending me the interesting material from the 
Texan coast and for securing permission to publish the description. The type- 
material of Onuphis (Onuphis) declivorum was collected during the offshore in- 
vestigations of the Bureau of Land Management, U.S. Department of Interior. 
I would like to thank my colleague. Dr. Meredith L. Jones, for advice and for a 
critical review of the manuscript. 

Literature Cited 

Augener, H. 1924. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. XVIII. Polychaeta 

II. Polychaeten von Neuseeland I. Errantia. — Videnskabelige Meddelelser fra Dansk Natur- 

historisk Forening, Copenhagen 75:241-441. 
Fauchald, K. 1980. Onuphidae (Polychaeta) from Belize, Central America, with notes on related 

taxa. — Proceedings of the Biological Society of Washington 93(3): 797-829. 
Hartman, O. 1967. Polychaetous annelids collected by the USNS Eltanin and Stat en Island cruises, 

chiefly from Antarctic Seas. — Allan Hancock Monographs in Marine Biology 2:1-387. 
, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to 

Bermuda and other North Atlantic areas (Part 2). — Allan Hancock Monographs in Marine 

Biology 6:1-327. 



250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Johnson, H. P. 190L The Polychaeta of the Puget Sound region. — Proceedings of the Boston Society 

for Natural History 29:381-437. 
Marenzeller, E. von. 1879. Suedjapanische Anneliden I. — Denkschrifte der Mathematisch-Naturwis- 

senschaftliche Classe der Kaiserlichen Akademie der Wissenschaften, Wien 41(2): 109-152. 
Moore, J. P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of 

southern California in 1904. III. Euphrosynidae to Goniadidae. — Proceedings of the Academy 

of Natural Sciences of Philadelphia 63:234-318. 
Rioja, E. 1944. Estudios anelidologicos XI. Notas sobre algunas especies de poliquetos de las costas 

mexicanas del Pacifico. — Anales Instituto Biologia, Mexico 15:139-145. 
Sars, M. 1835. Beskrivelser og lagttagelser over nogle maerkelige eller nye i Havet ved den Bergenske 

Kyst levende Dyr af Polypemes, Acalephemes, Radiatemes, Annelidenes og MoUuskemes 

Classer, med en kort Oversigt over de hidtil af Forfatteren sammesteds fundne Arter og deres 

Forekommen. Bergen, xii and 81 pp. 
Uschakov, P. V., and B. L. Wu. 1962a. [Preliminary report on the Polychaetes from Chekian and 

Fukian, China].— Studia Marine Sinica [Hai Yang K'o Hsueh Chi K'an] 1(1):89-108. 
, and . 1962b. [The Polychaetes from the Yellow Sea. VI. Additions to Errantia]. — 

Studia Marina Sinica [Hai Yang K'o Hsueh Chi K'an] 2(2): 110-134. 
Verrill, A. E. 1873. Results of recent dredging expeditions on the coast of New England. — American 

Journal of Science, New Haven (3) 5:98-106. 
Willey, A. 1905. Supplementary report XXX. Report on the Polychaeta collected by Professor Herd- 
man, at Ceylon in 1902. — Report to the Government of Ceylon on the Pearl Oyster Fisheries 

of the Gulf of Manaar by W. A. Herdman, D.Sc, F.R.S., P.L.S. with Supplementary Reports 

upon the Marine Biology of Ceylon by Other NaturaHsts 4:243-324. 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 251-255 

A NEW SPECIES OF THE GENUS CAMBARINCOLA 

(CLITELLATA: BRANCHIOBDELLIDA) FROM ILLINOIS 

WITH REMARKS ON THE BURSA OF CAMBARINCOLA 

VITREUS ELLIS, 1919, AND THE STATUS OF 

SATHODRILUS HOLT, 1968 



Perry C. Holt 

Abstract. — Cambarincola illinoisensis, new species, is described and illustrat- 
ed. Similarities between the everted bursa of this species, that of C. vitreus Ellis, 
1919, and those of species of Sathodrilus Holt, 1968, are illustrated and com- 
mented on. The involved genera are not synonymized, since it is expected future 
work will result in the erection of new genera to contain some species now 
assigned to each genus. 



Recently, in the process of redescribing Cambarincola elevatus Goodnight, 
1940, and reassigning the species to Sathodrilus Holt, 1968, specimens of Cam- 
barincola vitreus Ellis, 1919, and a previously unrecognized species of Camba- 
rincola were found with everted bursae. To describe the newly found species and 
comment on the bursae and penes of the 2 species and those of Sathodrilus are 
the objectives of this report. 

The methods I use have been fully stated (Holt 1960): here it might be empha- 
sized that the drawings, made with the aid of a camera lucida, are semi-diagram- 
matic, with little or no effort to indicate cellular detail, and those of bursae and 
penes are of optical sections done from animals mounted entire. Measurements 
are to be regarded as approximations and are given in millimeters with the ranges 
in parentheses. The initials "USNM" refer to the catalog numbers of specimens 
in the collections of the National Museum of Natural History; "PCH" to those 
of specimens in my collection. 

I wish to acknowledge the financial aid of the Virginia Polytechnic Institute 
and State University; the help given me by Horton H. Hobbs, Jr., and his iden- 
tifications of the host crayfishes; the assistance in collecting of my wife, Virgie 
F. Holt, and our daughter, Susan E. H. West. 

Cambarincola illinoisensis, new species 
Fig. lA-E 

Type-specimens. — Holotype, USNM 65225, 3 paratypes, USNM 65226, 5 para- 
types, PCH 840, taken on Orconectes virilis (Hagen 1870) from a prairie stream 
(? Sugar Creek) north of Stockland, Iroquois County, Illinois, by Perry C. and 
Virgie F. Holt, 25 July 1958. 

Diagnosis. — Medium-large, relatively robust worms (holotype 3.2 mm in 
length); upper lip with 4 obscure lobes, lower with 2; no oral papillae; low dorsal 
ridges; upper jaw twice that of lower in size, dental formula 5/4; bursa about V3 
body diameter in length, ovate; atrial fold present; protruded penis slender; tu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Cambarincola illinoisensis : A, B, Holotype; A, Outline of entire animal, lateral view; B, 
Lateral view of jaws; C-E, Paratype; C, Lateral view of reproductive systems (b, bursa; p, prostate; 
sg, spermiducal gland; spb, spermathecal bulb; spd, spermathecal duct); D, Optical longitudinal sec- 
tion through penis and bursa (a, atrium; ab, atrial part of bursa; af, atrial fold; ed, ejaculatory duct; 
p, penis; ps, penial sheath); E, Optical longitudinal section through everted bursa and penis. Cam- 
barincola gracilis from Lincoln County, Oregon: F, Optical longitudinal section through bursa and 
penis. Sathodrilus dorfus from Lincoln County, Oregon: G, Optical longitudinal section through bursa 
and penis. Sathodrilus lobatus from Lincoln County, Oregon: H, Optical longitudinal section through 
bursa and penis. 



VOLUME 95, NUMBER 2 253 

bular spermiducal gland about Vs body diameter in length, and about Vi its length 
in diameter; prostate slender, in diameter V4 to Vs that of spermiducal gland, 
composed of densely granular cells, with prominent ental bulb; spermatheca 
greater than body diameter in length, with long ectal duct, ovate bulb. 

Etymology. — For the state of Illinois. 

Description. — Five specimens of Cambarincola Illinois e nsis , including the ho- 
lotype, have the following dimensions: total length, 2.9 (2.5-3.3); greatest di- 
ameter, 0.5 (0.4-0.6); head length, 0.5 (0.4-0.7); head diameter, 0.3 (0.3-0.4); 
diameter, segment I, 0.3 (0.2-0.4); diameter, sucker, 0.3 (0.2-0.4). 

The lobes of the upper lip are very short, broad, and not easily seen in most 
specimens; there are no oral papillae. External sulci of the head are shallow and 
broad; the one such sulcus other than the peristomial one is a slight ventral 
depression. There is one internal pharyngeal sulcus. The dorsal ridges are low 
and indistinct, but weakly developed supernumerary segmental muscles are pres- 
ent. 

The jaws are distinctive: of an irregular triangular shape in lateral view, the 
upper bears a proportionately huge median tooth and 2 almost undetectable lateral 
teeth on each side; the lower jaw is from ^/s to Vi the upper in length and bears 
2 prominent paramedian teeth and 2 minute lateral ones. 

The spermiducal gland is not unusual in any respect. It is approximately % the 
body diameter in length and about Vi its length in diameter, tapering slightly to 
its junction with the ejaculatory duct, and usually lying obhquely in the coelom 
of its segment. The vasa deferentia enter the gland at widely separate points, 
creating the appearance, from some perspectives, of a prominent anterior deferent 
lobe. 

The prostate is unusual: the glandular cells which compose it are finely and 
densely granular, unlike those of the spermiducal gland which are of the common 
type, large and coarsely granular. In shape and size, the prostate is slender, in 
diameter about Vk that of the spermiducal gland and subequal to the latter in 
length. The ental bulb is prominent, but the prostate itself, lying partly obscured 
in lateral view alongside the spermiducal gland, is often difficult to see. 

The bursa is elongate ovate, approximately twice its diameter and about V3 the 
diameter of the body in length. There is a prominent median atrial fold. The penial 
sheath comprises about V3 of the total length of the bursa. 

One specimen of the paratype series has an everted bursa (Fig. IE). The penis 
is carried to the outside as a relatively slender tube, but it is impossible to tell if 
it itself is everted. 

The ejaculatory duct presents no features of note. 

The male efferent apparatus as a whole is proportionately small, lying entirely 
beneath the gut in some specimens and never extending to the dorsal border of 
the gut. The necessity of viewing the structures against a background of the gut 
contents makes it difficult to determine the extent and nature of the spermiducal 
gland and related structures in whole mounts, the only material available. For- 
tunately, and most unusually, in one paratype-specimen the ovarian, VII, segment 
is parasitized by a nematode and, presumably as a result, the spermatozoa that 
normally fill the coeloms of segments V and VI are absent and all of the male 
secondary sexual organs are easily seen. 

The spermatheca is somewhat greater than the body diameter in total length 



254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

and its ectal duct is about twice the length of its bulb which is slender ovate in 
shape. There is no ental process. 

Variations. — None, other than those associated with size and degree of con- 
raction, were noticed. 

Affinities. — Cambarincola illinoisensis seems to be closer to C. jamapaensis 
Holt, 1973, than to any other of its congeners. C. illinoisensis is composed of 
larger worms. The reproductive systems are of similar proportions, but differ in 
the proportionately greater size of the prostate of C. jamapaensis which was 
earlier considered as related to species from the Appalachians with differentiated 
prostates (Holt 1973:20). C. acudentatus Holt, 1973, also has a prostate composed 
of cells that are densely filled with small granules, but differs from C. jamapaensis 
and C. illinoisensis in the absence of dorsal ridges and in the shape and size of 
its jaws: delicate and light in color with a dental formula of 7/6 (Holt 1973:11- 
13). The "differentiated" prostate of these 3 species differ from those of such 
Appalachian species as C. philadelphicus (Leidy, 1851) and C.fallax Hoffman, 
1963, which have prostates that are composed of heavily vacuolated cells: those 
heretofore described as "differentiated." The Mexican species, C. jamapaensis, 
thus remains for now as the closest relative of C. illinoisensis, but the two are 
readily separated by the disparity in the size of the jaws in the latter. 

Host. — Orconectes virilis (Hagen, 1870). 

Distribution. — Known only from the type-locality; it is expected that C. illi- 
noisensis will be found widely throughout the Central Plains of the United States. 

Discussion. — As noted, a specimen among the paratypes has an everted bursa. 
In addition, 2 specimens of C. vitreus with everted bursae were found in the 
material studied for the paper referred to above (Holt 1978). The occurrence of 
specimens collected with everted bursae and protruded or everted penes is rare 
and in this case furnishes an opportunity to comment further on the structure of 
the bursa and penis in the genera Cambarincola and Sathodrilus. 

The penis of members of Cambarincola has been considered non-eversible 
(Moore 1895:498; Ellis 1912:481; Holt 1949:554; Hoffman 1963:289-290; inter 
alia), while that of species of Sathodrilus has been described as eversible (Holt 
1968:294). To my knowledge, no one has actually seen either the bursa or the 
penis evert in specimens of these genera and all these statements (certainly mine 
and Hoffman's) are inferences based on the observable structures of dead ani- 
mals. 

If Cambarincola vitreus and C. illinoisensis were not clearly in other respects 
congeneric with the type of the genus {Cambarincola macrodontus Ellis, 1912), 
the shape of their almost identical penes carried outward by everted bursae might 
align them with species of Sathodrilus. Indeed, a comparison of illustrations of 
optical sections, showing in each case the retracted penis (Fig. IF-H) of repre- 
sentative species of these genera does not reveal any significant differences: those 
of shape, size, proportionalities of parts and atrial folds are common among the 
members of both genera. Species of Sathodrilus differ from those of Cambarin- 
cola in lacking prostates, having only what I have called a "prostatic protuber- 
ance" (Holt 1968:298), or in having prostates that arise from the spermiducal 
gland entad to its junction with the ejaculatory duct. 

Taxonomists customarily recognize genera as groups of species separated from 
other such similar groups by one or more morphological "gaps." Often subse- 



VOLUME 95, NUMBER 2 255 

quent discoveries or additional studies lead to the obliteration of the "gaps" (Holt 
1968:5). Such seems to be the case here: the eversible penis attached by cyto- 
plasmic strands to the inner wall of the penial sheath, if indeed this is the correct 
description, of some species of Sathodrilus and the size and shape of the prostate 
vary through a series of species until the "gap" between Sathodrilus and Cam- 
barincola is virtually bridged. However, I do not choose to merge the currently 
recognized 44 species assigned to Cambarincola with the 15 that nominally com- 
pose the genus Sathodrilus. Rather, I suspect more than 2 genera should be 
erected for these 59 species and with a refinement of taxonomic concepts and 
procedures in the study of the branchiobdellids, I expect that this will be done. 

Literature Cited 

Ellis, Max M. 1912. A new discodrilid worm from Colorado. — Proceedings of the United States 
National Museum 42(1912):481-486. 

. 1919. The branchiobdellid worms in the collections of the United States National Museum, 

with descriptions of new genera and new species. — Proceedings of the United States National 
Museum 55(2267): 24 1-265, pis. 10-13. 

Goodnight, Clarence J. 1940. The Branchiobdellidae (Oligochaeta) of North American crayfishes. — 
Illinois Biological Monographs 17(3): 1-75, pis. 1-3. 

Hoffman, Richard L. 1963. A revision of the North American annelid worms of the genus Camba- 
rincola (Oligochaeta: Branchiobdellidae). — Proceedings of the United States National Museum 
114(3470):271-371. 

Holt, Perry C. 1960. The genus Ceratodrilus Hall (Branchiobdellidae, Oligochaeta) with the descrip- 
tion of a new species. — Virginia Journal of Science, new series, ll(2):53-73, pis. 1-4. 

. 1968. New genera and species of branchiobdellid worms (Annehda: Clitellata). — Proceedings 

of the Biological Society of Washington 81:291-318. 

. 1973. A summary of the branchiobdellid worms (Annehda: Clitellata) fauna of Mesoameri- 

ca. — Smithsonian Contributions to Zoology 142:i-iii, 1^0. 

. 1978. The reassignment of Cambarincola elevatus Goodnight, 1940 (Clitellata: Branchiob- 

dellida) to the genus Sathodrilus Holt, 1968. — Proceedings of the Biological Society of Wash- 
ington 9 1(2): 472^82. 

Leidy, Joseph. 1851. Contributions to helminthology. — Proceedings of the Academy of Natural Sci- 
ence of Philadelphia 5:205-209. 

Moore, J. Percy. 1895. The anatomy of Bdellodrilus illuminatus, an American discodrilid. — Journal 
of Morphology 10(2): 497-540, pis. 28-32. 

Department of Biology, Virginia Polytechnic Institute and State University, 
Blacksburg, Virginia 24061. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 256-264 

A NEW SPECIES OF OREOHELICID LAND SNAIL 
FROM THE SAN AGUSTIN PLAINS, NEW MEXICO 

Celinda R. Crews and Artie L. Metcalf 

Abstract. — A new species of pulmonale land snail, Oreohelix litoralis, is de- 
scribed from populations inhabiting wave cut cliffs in the southern part of the San 
Agustin Plains, west-central New Mexico. Taxonomic evaluation is based on 
qualitative and quantitative characters of shell and genital structures. A rede- 
scription of Oreohelix magdalenae Pilsbry is provided, since it is considered to 
be closely related to Oreohelix litoralis. Both species are members of the Oreo- 
helix metcalfei Cockerell complex within the Oreohelix yavapai Pilsbry group. 



Most of the basic taxonomic work on the genus Oreohelix in New Mexico was 
done by H. A. Pilsbry and J. H. Ferris s, culminating in Pilsbry' s (1939) mono- 
graphic treatment of known species. A number of New Mexican oreohelicids 
were assigned to the wide-ranging Oreohelix yavapai Pilsbry, 1905, group. Taxa 
occurring in the Black Range and peripheral mountains in south-central New 
Mexico were ascribed to the Oreohelix metcalfei Cockerell, 1905, complex. Met- 
calf (1974) discussed and referred several species to this complex, including Or- 
eohelix magdalenae, Pilsbry, 1939, as a northeastern derivative. The new species 
described below is judged to be a northern derivative of this complex and to be 
closely related to Oreohelix magdalenae . 

Oreohelix litoralis, new species 
Figs. lA, B, C; 2A; 3A 

Description of shell of holotype (from Locality 6). — Shell slightly depressed, 
convex dorsally and ventrally; spire forming angle of 118°; 16.4 mm in diameter 
and 9.7 mm in height; upper lip descending only slightly; angular peripherally 
with angularity at about mid-height of body whorl; first 2 whorls keeled; aperture 
slightly ovate horizontally, 7.7 mm wide and 6.2 mm high; aperture oriented at 
angle of 54° to vertical axis of shell; umbilicus open, not contracting rapidly 
within, 3.9 mm wide, contained 4.21 times in diameter; sutures moderately im- 
pressed; 5.10 whorls; dorsal surface generally smooth, with sculpture limited to 
growth lines, weak and irregularly spaced on early whorls, becoming stronger on 
body whorl; ventral surface smooth with only low growth lines (no spiral striae 
or Hrae); earliest 2.25 whorls of dorsal surface uniformly cinnamon brown, grading 
to grayish brown mottling on a whitish background by whorl 3 with mottled 
pattern extending to body whorl; thin brown band appears at 2.25 whorls, ex- 
tending to body whorl; this band situated approximately one-third whorl- width 
inward from suture on earlier whorls and approximately one-third whorl-width 
above angulation on body whorl; ventral surface with grayish-brown mottling 
mainly descending ("bleeding") from thin brown band present on body whorl 
immediately beneath angulation. 

Variation in shells. — Variation in shell measurements and proportions for 20 



VOLUME 95, NUMBER 2 



257 




Fig. 1. A, B, C, Shell of holotype of Oreohelix litoralis (diameter, 16.4 mm); D, E, F, Shell of 
Oreohelix magdalenae from Loc. 9 (diameter, 15.7 mm). 



Specimens each from Localities 6 and 7 is given in Table 1 . A few paratypes are 
less depressed than the holotype. Descent of the body whorl ranges from weak 
to marked. Shape of the aperture ranges from round to oval. Banding and other 
coloration varies little among paratypes from Localities 1-7. There is a variable 
degree of fading, due to wear, on the ventral surface of the last half of the body 
whorl. 

Description of genitalia (based on dissections of 5 paratypes from Locality 6; 
collected in Dec. 1979; see Figs. 2A and 3A). — Lower penis swollen, upper penis 
bearing small, rounded appendix laterally; internally, lower penis bears 2^ ir- 
regular longitudinal pilasters, upper penis with close-set pustules, varying in size; 



258 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Some shell and genital measurements (mm) and proportions for Oreohelix litoralis and 
Oreohelix magdalenae. Mean and standard deviation (in parentheses) are given above and range 
below. D = diameter. 





O. litoralis 
Loc. 6; n = 20 


O. litoralis 
Loc. 7; n = 20 


O. magdalenae 
Loc. 9; n = 20 




O. litoralis 
Loc. 6; n = 5 


O. magdalenae 
Loc. 9; n = 5 


Diameter 


18.9(1.01) 
17.2-21.5 


17.5 (0.86) 
16.2-19.9 


14.0 (0.88) 
12.8-15.7 


Penis 


8.3 (0.41) 
7.8-8.9 


8.9 (0.67) 
7.9-9.6 


Height 


11.7(0.89) 
10.0-13.6 


11.2(1.03) 
10.0-13.6 


8.5 (0.61) 

7.5-9.5 


Epiphallus 


3.4 (0.41) 
2.8-3.8 


3.4 (0.37) 
2.8-3.8 


Aperture 
width 


8.8 (0.64) 
7.5-10.3 


8.4 (0.75) 
7.1-10.4 


6.5 (0.38) 
5.9-7.1 


Vas deferens 


6.3 (0.47) 
5.8-7.0 


9.8 (0.44) 
9.4-10.5 


Aperture 
height 


7.1 (0.42) 
6.2-7.8 


7.8 (0.57) 
6.5-8.8 


6.2 (0.29) 

5.7-6.7 


Vagina 


3.2 (0.34) 
2.9-3.8 


3.2 (0.21) 
3.0-3.5 


Umbilicus 
width 


5.1 (0.47) 
4.5-6.2 


4.4 (0.39) 
3.9-5.1 


3.7 (0.30) 
3.2-4.3 


Free oviduct 


3.2 (0.41) 
2.7-3.6 


2.7 (0.29) 
2.3-3.0 


No. of whorls 


5.33(0.122) 
5.10-5.55 


5.16(0.141) 
4.90-5.40 


4.61 (0.206) Spermathecal 
4.25-5.00 duct plus sac 


12.6(1.03) 
11.5-13.6 


11.9(0.72) 
10.9-12.6 


D/height 


1.62(0.102) 
1.38-1.84 


1.58(0.096) 
1.35-1.72 


1.65 (0.068) D/penis 
1.55-1.83 


2.00(0.147) 
1.82-2.15 


1.56(0.106) 
1.44-1.69 


D/aperture 
width 


2.14(0.077) 
2.01-2.31 


2.10(0.097) 
1.91-2.29 


2.15(0.089) D/epiphallus 
2.00-2.31 


4.92 (0.553) 
4.42-5.79 


4.12(0.379) 
3.55-4.61 


D/aperture 
height 


2.67(0.135) 
2.43-2.99 


2.26(0.106) 
2.05-2.49 


2.25 (0.136) D/vas deferens 

2.02-2.53 


2.62 (0.210) 
2.39-2.88 


1.42(0.083) 
1.33-1.51 


D/umbilicus 
width 


3.75 (0.237) 
3.38-4.22 


4.01 (0.264) 
3.49-4.39 


3.76 (0.210) D/vagina 
3.34-4.11 


5.15(0.591) 
4.42-5.76 


4.41 (0.366) 
4.03-4.77 


D/no. of whorls 


3.55(0.179) 
3.23-4.02 


3.40(0.145) 
3.11-3.69 


3.03(0.112) D/free oviduct 

2.83-3.27 


5.26 (0.574) 
4.67-6.19 


5.16(0.606) 
4.70-6.22 










D/spermathe- 
cal duct 


1.32(0.042) 
1.26-1.36 


1.17(0.077) 
1.07-1.27 










plus sac 







Upper and lower penis approximately equal in length; penial verge short; penial 
retractor muscle strands attached to both upper penis and epiphallus at their 
juncture; epiphallus relatively long and slender, tapering at upper end; vas def- 
erens inserts centrally on upper terminus of epiphallus; vagina stout, widening at 
upper end; free oviduct stout, widening at base; albumen gland short and muzzle- 
shaped with slightly darker pigmentation; talon hook-like, black in color, slightly 
swollen and of moderate size relative to albumen gland; spermathecal sac oval 
to suboval. Lengths of some genital organs and ratios of shell diameter to organ 
lengths are given in Table 1 . 

Description of embryonic shells. — Embryos taken from a single dissected spec- 
imen from Locality 6 possess 2.55, 2.60, and 2.65 whorls and diameters measure 
4.2, 4.1, and 4.2 mm, respectively. Overall color is cinnamon brown as seen in 
live adult shells for approximately the earliest 2.5 whorls (first 2.25 whorls in the 
holotype). Banding is faint, but discernible dorsally after the second whorl. Em- 
bryonic shells are keeled, but possess no cuticular projections. 

Remarks. — On 11 October 1980, Oreohelix litoralis was found in crevices and 
in rock rubble of wave cut cliffs on the southern margin of the San Agustin Plains 
at an elevation of approximately 2105 m (6900 feet) (Loc. 6). Rocks were igneous. 



VOLUME 95, NUMBER 2 



259 





Fig. 2. A, Genitalia of Oreohelix litoralis; B, Genitalia of Oreohelix magdalenae. Ip = lower 
penis; up = upper penis; ap = appendix; pr = penial retractor muscle; e = epiphallus; vd = vas def- 
erens; pg = prostate gland; v = vagina; fo = free oviduct; u = uterus; ag = albumen gland; t = tal- 
on; hd ==: hermaphroditic duct; sd = spermathecal duct; s — spermathecal sac; a = atrium; go = 
common gonopore. 



mainly rhyolite tuff, welded tuff, and scoria. The area is grassland with sparse 
shrubs, in contrast to the forested habitats observed in association with most 
occurrences of Oreohelix. Common plants included Atriplex canescens (four- 
winged saltbush), Rhus aromatic a (lemon sumac), Brickellia sp. (brickellia), 
Lycium sp. (wolfberry), and Bouteloua gracilis (blue grama grass). 

Live, aestivating individuals were found attached to small rocks in rubble, on 
the underside of larger stones in talus, and beneath low ledges at the base of the 
cliffs. Oreohelix litoralis at this locality seemed colonial, with up to 11 individuals 
clustered under one rock. Both live and dead snails were abundant. The harsher 
environment occupied by these populations (relative to montane populations) 
may be related to the high degree of colonialism observed. 

Disposition of types. — The holotype (Fig. lA, B, C) is a fresh shell with des- 
iccated soft parts (National Museum of Natural History 784661). The paratypes 
are fresh shells with desiccated soft parts (National Museum of Natural History 
784662; University of Texas at El Paso 7725, 8467, 8593-8594; Academy of Nat- 
ural Sciences of Philadelphia 354156; University of Arizona 19016; Dallas Mu- 
seum of Natural History 5364). 

Etymology. — The epithet litoralis (L., of the shore) refers to occurrence of 
populations of the species along wave cut cliffs of the former shoreline of Pluvial 
Lake San Agustin. 



Oreohelix magdalenae Pilsbry 
Figs. ID, E, F; 2B; 3B 

Oreohelix socorroensis magdalenae Pilsbry, 1939: 515, Fig. 336b (Type: 

dalena Mountains, Socorro County, New Mexico; ANSP 158166). 
Oreohelix magdalenae. — Metcalf, 1974:99. 



Mag- 



260 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 3. A, Longitudinal dissection of the penis of Oreohelix litoralis; B, Longitudinal dissection 
of the penis of Oreohelix magdalenae . pr ^ penial retractor muscle; e = epiphallus; po = pore; pu = 
pustules; pi = longitudinal pilasters; v = vagina; a = atrium; go = common gonopore. 



Description of shells (based on 20 specimens from Locality 9). — Shell slightly 
depressed to, in a few specimens, pyramidal; diameter 12.8-14.7 mm (mean 14.0 
mm); height 7.5-9.5 mm (mean 8.5 mm); descent of body whorl slight to none; 
approximately first 2.25 whorls keeled, becoming angular peripherally with an- 
gularity at about mid-height of body whorl; aperture rounded to oval perpendic- 
ular to parietal wall, 5.9-7.1 mm wide (mean 6.5 mm) and 5.7-6.7 mm high (mean 
6.2 mm); umbilicus open and ample, 3.2-4.3 mm wide (mean 3.7 mm), contained 
3.34-4.11 times within diameter (mean 3.76 times); 4.25-5.00 whorls (mean 4.61 
whorls); dorsal surface generally smooth with faint lirae on earliest approximately 
2.5 whorls, not extending to later whorls; transverse growth lines on first 2.5 
whorls, becoming more irregular and increasing in strength on outer whorls; 
ventral surface sculpture limited to transverse growth lines; dorsally, first 2.5 
whorls dark gray, grading to a dull gray and brown mottled pattern intersected 
by irregular transverse white bands that increase in area on outer whorls; thin 
brown band evident dorsally after first whorl, becoming stronger on later whorls 
and extending to body whorl, band located approximately one-third whorl- width 
above suture on earlier whorls and one-third whorl-width above peripheral an- 
gulation on body whorl; ventrally, irregular gray, white, and brown transverse 
zones of coloration with last 0.5 body whorl worn to tan on some specimens; 
ventral band located immediately below peripheral angulation, distinct, and ap- 
proximately 3 times size of dorsal band. Additional data on variations in shell 
measurements and proportions are given in Table 1 . 

Description of genitalia (based on dissections of 5 specimens from Locality 9, 
collected on 20 April 1980; see Figs. 2B and 3B). — Lower penis greatly swollen, 



VOLUME 95, NUMBER 2 261 

approaching a bulbous condition; large, prominent lateral appendix on upper 
penis; internally, lower penis with 2 to 3 irregularly shaped longitudinal pilasters; 
upper penis with numerous suboval pustules, varying in size; upper and lower 
penis approximately equal in length; penial verge slight; penial retractor muscle 
with attachment to both upper penis and epiphallus; epiphallus long and slender; 
central insertion of the vas deferens onto the apex of the epiphallus; vagina stout, 
slightly swollen at emergence of spermathecal duct; free oviduct also stout, slight- 
ly larger at base than top; spermathecal sac oval; albumen gland more darkly 
pigmented, relatively small; talon hook-like and black, large relative to albumen 
gland. 

Of a second group of 5 dissections done on specimens of O. magdalenae col- 
lected at Locality 9 on 19 May 1981, 3 specimens of adult size (diameters 14.0, 
12.5, and 12.5 mm) had the same overall development of the genitalia as seen in 
the first group of dissections. However, 2 seemingly subadult specimens showed 
varying degrees of development of the genital organs. One specimen (diameter 
10.0 mm) had only slight evidence of the lower genitalia (penis-vagina area only), 
with no development of the uterus-prostate area. Another specimen (diameter 
10.6 mm) showed the beginning of development of the uterus-prostate area, with 
lower genitalia developed, but not of full size. 

Lengths of some genital structures and ratios of shell diameter to organ lengths 
for 5 adult specimens are given in Table 1. 

Remarks. — At Locality 9, O. magdalenae was found living under loose igneous 
stones at 2230 m (7320 feet) elevation. Vegetation included Pseudotsuga menziesii 
(Douglas fir), Pinus ponderosa (ponderosa pine), and Quercus gambelii (Gam- 
bel's oak) on the slope where collections were made, with Populus angustifolia 
(narrowleaf cotton wood) common along a brook below the slope. A few speci- 
mens were collected (Robert H. Weber) at another site in the range (Locality 8) 
near the summit of North Baldy Peak at approximately 3000 m (9850 feet). 

At Locality 9, on 20 April 1980 and 19 May 1981, aestivating individuals oc- 
curred under stones both attached and unattached with epiphragms intact. One 
individual was active under a stone on 20 April 1980. Two to 3 live individuals 
occurred together under some stones. 

Embryos were absent in uteri of specimens dissected, indicating that devel- 
opment to observable size (or perhaps fertilization) had not yet occurred or that 
deposition of young had occurred before these (spring) dates noted. It is logical 
to suppose that deposition of young should occur before or during the early phase 
of the summer rains to allow subsequent development under favorable conditions. 
No detailed account of the life history has been published for any species of 
'Oreohelix in New Mexico. 

Distribution 

Oreohelix litoralis, as now known, occurs in local populations along the south- 
ern margin and on an isolated hill arising from the floor of the San Agustin Plains. 
Oreohelix magdalenae occurs in local populations in the Magdalena Mountains, 
east of the Plains. These species seem to represent northern, peripheral deriva- 
tives of the Oreohelix metcalfei complex. This complex is centered in the Black 
Range, to the south, with several peripheral species in outlying ranges (Metcalf 



262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

1974:99). Ancestors of the 2 species possibly migrated northward during Pleis- 
tocene pluvial episodes. The occurrence at Locality 1 is described by Robert H. 
Weber (in litt.) as follows: "an isolated outcrop on the floor of the lake plain, 
separated from the cliffs ... by nearly two miles of open grassland. The hill was 
submerged at water levels about 6885 feet [2098 m], and was an island between 
that level and a little over 6800 feet [2073 m]." Weber (pers. comm.) indicates 
that the hill was submerged in Pluvial Lake San Agustin during the late Wisconsin 
rise of the lake and that it was, for a time, an island during the time of lake 
recession in the earlier Holocene. Thus, colonization of the hill by O. litoralis 
seems to have taken place during the past 10,000 years. Mountain ranges im- 
mediately south of the San Agustin Plains have not been investigated malacol- 
ogically. 

Differential Diagnoses and Discussion 

Although O. litoralis seems closely related to O. magdalenae, it is judged 
sufficiently distinctive to warrant specific rank. Shells of O. litoralis are larger, 
overall, than shells from populations of O. magdalenae observed (Table 1), and 
possess more strongly indented sutures and a slightly higher spire. In O. litoralis, 
no lirae are present either dorsally or ventrally, whereas the dorsal surface of O. 
magdalenae possesses faint lirae on the earliest ca. 2.5 whorls. Although a mot- 
tled color pattern is observed in both species, shells of O. magdalenae are dom- 
inated by a darker gray and brown and exhibit less diffuse mottling, with white 
areas being restricted and more distinctly demarcated. Both species possess a 
thin brown band dorsally and a second band below the peripheral angulation. 
However, in O. litoralis the 2 bands are approximately equal in size with the 
lower band diffusing ventrally ("bleeding"), whereas the lower band in O. mag- 
dalenae is 3 times the width of the upper and discrete. 

The overall shell size and number of whorls observed in O. litoralis and O. 
magdalenae generally are closer to those recorded for taxa in the O. metcalfei 
complex (Pilsbry 1939:507-514; Metcalf 1974:95-98) than to those observed for 
other members of the Oreohelix yavapai group occurring in New Mexico. Both 
O. litoralis and O. magdalenae have peripheral angulation at about mid-height 
of the body whorl in adult shells. Other geographically peripheral taxa (see above) 
of the O. metcalfei complex also are angular peripherally, in contrast to the 
carinate condition seen in centrally located (i.e.. Black Range) taxa. The periph- 
eral taxa of the complex, along with O. litoralis and O. magdalenae, also have 
shells that are less depressed, and with more rounded (biconvex) whorls than 
those seen in centrally located members of the O. metcalfei complex. Banding 
is relatively prominent and surface sculpture is not elaborate, generally lacking 
prominent striae and lirae, in O. litoralis, O. magdalenae, and other peripheral 
taxa of the O. metcalfei complex. 

We regard the characters exhibited by O. litoralis, O. magdalenae, and other 
peripheral taxa — angulation opposed to strong keeling, higher elevation of the 
spire, biconvex shells, relatively smooth surface sculpture, and relative promi- 
nence of banding — as conservative and opposed to more advanced characteristics 
seen in taxa in the center of distribution (i.e.. Black Range) of the Oreohelix 
metcalfei complex. These latter species possess varying degrees of carination and 



VOLUME 95, NUMBER 2 263 

depression of shells, elaborate spiral and/or radial surface sculpturing, and lack 
prominent banding. Metcalf (1974:99) noted that banding in O. confragosa Met- 
calf, 1974 (which occurs west of the Black Range and which he believed to be 
more closely related to the peripheral taxa) is weak in living specimens, but more 
prominent in fossil shells. If the two-banded pattern in Oreohelix is an ancestral 
feature of the genus in the process of disappearing (Pilsbry 1939:413), then the 
evolution towards the loss of banding observed by Metcalf would seem to support 
the view that the strongly-banded peripheral taxa of the Oreohelix metcalfei com- 
plex are more conservative. 

Pilsbry (1939:510) reported the presence of prominent "triangular processes" 
(cuticular projections) on the keel and dorsal and ventral surfaces of embryonic 
shells of taxa in the O. metcalfei complex in both peripheral and centrally located 
taxa. Unfortunately, embryonic shells of O. magdalenae were not available for 
comparison. However, embryonic shells of O. litoralis differ from those reported 
for other members of the O. metcalfei complex in possessing a relatively smooth 
surface and lacking these cuticular projections. 

The genitalia of O. litoralis are similar to those of O. magdalenae in possessing 
a swollen lower penis, a lateral appendix situated on the upper penis, and a 
relatively reduced albumen gland (Figs. 2A, B). However, the degree of swelling 
of the lower penis is much greater in O. magdalenae , approaching a bulbous 
condition, and reduction in size of the albumen gland is greater. The appendix 
also is larger and more pronounced in specimens of O. magdalenae. Several 
subspecies of O. metcalfei were reported by Pilsbry (1939:511) as having the 
lower penis "much swollen." However, the genitalia of most subspecies have 
not been described in detail or illustrated for comparison. No information is 
available regarding the presence of a lateral appendix on the upper penis in this 
complex, with the exception of O. confragosa (Metcalf 1974:96-98), which pos- 
sesses a small, lateral, cornuted appendix. Insertion of the vas deferens is central 
in O. litoralis and O. magdalenae, but has not been reported for other taxa of 
the O. metcalfei complex. 

List of Localities 

Localities are listed west to east. Localities 2-7 are in areas of wave-cut cliffs 
along the generally southern shore of Pluvial Lake San Agustin. 

1 New Mexico: Catron Co.: on isolated outcrop on floor of Pluvial Lake San Agustin (former 
island): T6S, R14W, SW14 SE1/4 NW1/4 Sec. 28:2040 m (6700 feet): 33°45'34"N; 108°17'29"W (Robert 
H. Weber, collector; Nov. 1981). 

2 New Mexico: Catron Co.: TVS, R14W, NW1/4 NE1/4 NW1/4 Sec. 2: 1950 m (6400 feet): 33°44'05"N; 
108°15'26"W (Robert H. Weber, collector; Sept. 1981). 

3 New Mexico: Catron Co.: T6S, R14W, NE1/4 NE1/4 SW^A Sec. 26: 1950 m (6400 feet): 33°45'26"N; 
108°15'22"W (Robert H. Weber, collector; Sept. 1981). 

4 New Mexico: Catron Co.: shore of Pluvial Lake San Agustin: T6S, R14W, NE14 SWA SW14 Sec. 
35: 2040 m (6700 feet): 33°44'24"N; 108°15'18"W (Robert H. Weber, collector; Sept. 1981). 

5 New Mexico: Catron Co.: T6S, R14W, SE1/4 SE1/4 SE1/4 Sec. 24: 2075 m (6800 feet): 33°46'00"N; 
108°13'44"W (Robert H. Weber, collector; Sept. 1981). 

6 Type locality of Oreohelix litoralis. New Mexico: Catron Co.: center, boundary T6S, Rl IW, Sec. 
6 and T6S, R12W, Sec. 1: 2105 m (6900 feet): 33°48'54"N; 108°01'16"W (collected by Robert H. Weber, 
Dec. 1979, and by A. L. Metcalf and C. R. Crews, 11 Oct. 1980). 



264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

7 New Mexico: Catron Co.: T5S, Rl IW, SW1/4 NW1/4 SW1/4 Sec. 27: 2105 m (6900 feet): 33°50'28"N; 
107°59'01"W (Robert H. Weber, collector; 28 Nov. 1979). 

8 New Mexico: Socorro Co.: Cibola Natl. Forest: Magdalena Mts.: summit of North Baldy Peak: 
TBS, R3W, SEi/t SWi/i NW1/4 Sec. 17: 3005 m (9858 feet): 34°03'02"N; 107°10'56"W (Robert H. Weber, 
collector; May 1976). 

9 New Mexico: Socorro Co.: Cibola Natl. Forest: Magdalena Mts.: T3S, R3W, SWV^ SEV4 Sec. 
21: North Fork Cn.: 2.6 km (1.6 mile) by road from Water Cn.: base of N-facing slope: 2230 m (7320 
feet): 34°01'47"N; 107°09'16"W (A. L. Metcalf and C. R. Crews, collectors; 20 April 1980; 19 May 
1981). 

Acknowledgments 

We are grateful to Dr. Robert H. Weber, New Mexico Bureau of Mines and 
Mineral Resources, who first apprised us of Oreohelix lit oralis and generously 
supplied specimens from several localities. 

Literature Cited 

Metcalf, A. L. 1974. Peripheral species of the Oreohelix metcalf ei CockereU complex (Pulmonata: 

Oreohelicidae).— The Nautilus 88:94-100. 
Pilsbry, H. A. 1939. Land Mollusca of North America (north of Mexico). — Academy of Natural 

Sciences, Philadelphia, Monograph 3:l(l):i-xvii + 1-573 + i-ix. 

Laboratory for Environmental Biology, University of Texas at El Paso, El 
Paso, Texas 79968. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 265-268 

A NEW TREE SNAIL, GENUS DRYMAEUS (BULIMULIDAE) 

FROM SOUTHEASTERN PERU 

Fred G. Thompson and Jane E. Deisler 

Abstract. — Drymaeus aurantiostomus , n. sp. is described from Madre de Dios 
Province, Peru. It belongs to the D. expansus species group and is the southern- 
most member of the group. 



In this paper we describe a new species of tree snail belonging to a group that 
is widely distributed in the Andean region of northwestern South America and 
adjacent Panama. The snail comes from the Department of Madre de Dios in 
southeastern Peru and is the southernmost member of the species group to which 
it belongs. The molluscan fauna of this department is very poorly known. No 
paper deals specifically with the region. We are grateful to Roy W. McDiarmid, 
Department of Amphibians and Reptiles, National Museum of Natural History 
for sending us the snail for study and to Alan Solem, Field Museum of Natural 
History (FMNH) for the loan of comparative specimens. 

Drymaeus aurantiostomus new species 
Figs. 1, 2, 5, 6 

Diagnosis. — A species of the subgenus Drymaeus Albers, and of the expansus 
species group because of its broadly expanded peristome, it has a relatively ru- 
gose axial sculpture and a compressed imperforate umbilicus. It differs from 
related species by its color. It is white with a subperipheral and basal spiral band, 
and a spiral series of rust colored flames above each band. The interior of the 
peristome is bright yellow, thus the Latin derivation of the specific name auran- 
tiostomus. 

Shell (Figs. 1, 2). — Medium sized, about 30 mm long; conical with a broadly 
reflected peristome. Thin, weakly translucent. Spire nearly straight-sided. Um- 
bilicus compressed-rimate. Whorls 5.8, nearly flat-sided with a distinct but weakly 
impressed subperipheral suture. Whorls regularly increasing in size and descent; 
insertion of peristome rising onto periphery of previous whorl; 2.1 embryonic 
whorls. Post embryonic whorls sculptured with distinct, regularly spaced, axial 
riblets that are crossed by much finer incised spiral striations; axial sculpture 
strongest on last whorl behind lip. Peristome broadly expanded throughout, con- 
spicuously dilated baso-laterally. Aperture oblique in lateral profile, lying at about 
22° to shell axis, nearly tangential to body whorl. Columellar lip broadly expanded 
in front of umbilical area. Columella straight, nearly vertical. Color: Embryonic 
whorls grayish yellow; postembryonic whorls white with a subperipheral and a 
basal purple band on last whorl; each band bordered above by regularly spaced 
rusty flames; those above subperipheral band continuing on to penultimate whorl 
as a series of small reddish brown spots (Fig. 2); interior of aperture white with 
outer bands and flames clearly distinguishable. Peristome with a broad yellow 
zone that continues onto columella; edge of peristome white. Parietal callus thin, 
clear, colorless. 



266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 1-4. Drymaeus spp.: 1, 2, D. aurantiostomus , holotype; 3, 4, D. latitesta, holotype. 



Measurements of holotype: shell height, 29.2 mm; shell width, 19.7 mm; ap- 
erture height 17.1 mm; aperture width, 16.0 mm. 

Radula. — Lost during specimen preparation. 

Reproductive system (Figs. 5, 6). — Ovotestis large, occupying nearly entire 
digestive gland and consisting of numerous closely packed alveolae. Hermaph- 
roditic duct highly convoluted; lower half greatly swollen. Albumen gland 8.9 mm 
long, lying completely beneath right side of stomach, crescent-shaped with a deep 
transverse intestinal groove across distal end. Talon 3.6 mm long, club-shaped, 
flexed in middle, completely exposed. Carrefour lying along columellar side of 
albumen gland. Uterus about 30 mm long, voluminous and highly convoluted. 
Spermatheca globose, lying along columellar side of uterus beneath pericardium 
and just below junction of uterus and albumen gland. Spermathecal duct 23 mm 
long, tapered. Vagina 5.3 mm long, moderately voluminous. Atrium short. Penis 
about 7 mm long (severed during dissection but apparently recovered entire). 
Penis simple, slender, with 5 longitudinal branching pilasters internally; demar- 
cated from epiphallus by slight internal constriction of lumen. Base of penis with 
a short thick penis sheath about 2 mm long. Epiphallus 7 mm long, slender, 
cylindrical, terminated by an epiphallic flagellum 1.6 mm long. Lumen of epi- 
phallus with low, narrow tubercles anastomosing in lower portion; upper portion 
with 5-6 longitudinal branching pilasters that continue diminished into flagellum. 
Penis retractor muscle short, 3 mm, inserting on inner wall of lung about V3 of 
distance from pericardium to mantle collar; originating on apex of flagellum. Vas 
deferens emerging at junction of epiphallus and flagellum, completely embedded 
in epiphallic and vaginal wall; partially exposed above penis sheath. 

Type-locality. — Peru, Dept. Madre de Dios, Tambopata Reserve, on Rio Tam- 
bopata at Rio La Torre, about 30 km SSW Puerto Maldonado (12°49' S, 69°17'W); 
280 m alt. Holotype: UF 26605; collected 19 November 1979 by Roy W. Mc- 
Diarmid on a shrub along the Tres Chimbadas Trail, about 20 minutes from the 
Explorers Inn. 

Discussion. — The Bulimulidae are anatomically conservative. Subfamilies are 
weakly differentiated. Genera and subgenera usually differ only by shell char- 
acters, and occasionally by slight anatomical features. Studies on Drymaeus 
anatomy have failed to find anatomical traits that are useful for group classifi- 
cation (Strebel 1882; Baker 1925; Pilsbry 1946; Solem 1955; van Mol 1971; Breure 
1976; Breure and Eskens 1977). Breure (1979) demonstrated that the digestive 
and reproductive systems in Drymaeus are highly conservative. Minor variations 



VOLUME 95, NUMBER 2 



267 



ntestinal groove 



ovotestis 




albumen gland 



albumen gland 

penis retractor 
flagellum 
spermatheca 

uterus ( -L — epiphallus 



penis 
vas deferens 
penis sheath 




uterus 



vagina, 



5mm 



Figs. 5-6. Drymaeus aurantiostomus, holotype: 5, Reproductive system; 6, Upper part of female 
system showing exposed talon. Scales in mm. 



in the spermatheca, penis complex and radula have evolved independently in 
different species groups and have little phylogenetic significance, although they 
may be useful for comparing closely related species. It is apparent that in Dry- 
maeus the shell is an adequate indicator of group relationships and specific dis- 
tinctions. Data from other morphological systems have not provided a better 
basis for classification. The reproductive system of D. aurantiostomus is de- 
scribed so that other species may be compared with it. However, other members 
of the expansus group have not been studied anatomically. 

D. aurantiostomus is related to a group of Andean species with a broadly 
expanded peristome and relatively rugose axial sculpture. The group includes D. 
expansus (Pfeiffer, 1848), and its subspecies vanattai Pilsbry, 1898, balboa Pils- 
bry, 1926, subprotractus Pilsbry, 1902, flavilabrum Weyrauch, 1967, ore e si Wey- 
rauch, 1958, altorum Weyrauch, 1958 Siud pereninus DaCosta, 1901, as well as 
D. inca Smith, 1943, D. latitesta Haas, 1952, D. protractus (Pfeiffer, 1885), D. 
bartletti (Adams, 1866), D. scitus (Adams, 1885), D. eusterius Pilsbry, 1944, and 
D. weeksi Pilsbry, 1926. The group is widely distributed in the northern Andean 
region in Panama, Columbia, Ecuador, and Peru. D. aurantiostomus and D. 
latitesta are the southernmost members of the group and come from adjacent 
areas in Peru. 

D. aurantiostomus is similar in shape and size to D. expansus. D. expansus 
differs by having more rugose sculpture, the peristome is not as protracted baso- 
laterally, the color pattern consist of purpHsh flammulate blotches on a grayish 
white background, a basal band is absent, the interior of the aperture is purple 
tinted, and a subperipheral band is usually absent, except in some specimens in 
which the flames may fuse to form a band. 



268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

D. aurantiostomus is similar to D. latitesta Haas, a species described from 
Dept. Cuzco, Peru. The holotype of D. latitesta (FMNH 38120) (Figs. 3, 4) is a 
partly weathered specimen with a chipped peristome and broken apex, but im- 
portant characteristics of shape and color pattern are clear. They are alike in 
having a subperipheral band that enters the aperture below the insertion of the 
outer lip (the band is faded on the front of the holotype of D. latitesta) and in 
having a basal band that enters the aperture at the insertion of the columellar lip. 
Both species have flammulate markings above the bands, though they are better 
defined and more periodic in D. aurantiostomus. D. latitesta differs from D. 
aurantiostomus in structure by having more inflated whorls with a more deeply 
impressed suture. It is larger (37.2 mm long), is more corpulent in shape, and the 
axial striations are weaker (Fig. 4). The columella is nearly vertical and the outer 
peristome is nearly uniformly arched. The plane of the aperture in lateral profile 
is vertical and basal in position, not tangential to the body whorl. The color 
pattern differs from that of D. aurantiostomus by having an additional supraper- 
ipheral band with flames above it (Fig. 4). 

Literature Cited 

Baker, H. B. 1925. The Mollusca collected by the University of Michigan- Williamson Expedition in 

Venezuela. IV. — Occasional Papers of the Museum of Zoology University of Michigan 167:1- 

49, pis. 12-19. 
Breure, A. S. H. 1976. Notes on Bulimulidae (Gastropoda, Euthyneura): Some BuUmulidae from 

French Guyana and Surinam with notes on their anatomy. — Zoologische MededeUngen Rijks- 

museum Natuurlijke Historic Leiden 50:107-115. 
. 1979. Systematics, phylogeny and zoogeography of BuHmulidae (Mollusca). — Zoologische 

Mededelingen Rijksmuseum Natuurlijke Historic Leiden (168): 1-215, pis. 1-3. 
, and A. A. C. Eskens. 1977. Observations on the formation of spermatophores in a bulimulid 

land snail, Drymaeus canaliculatus (Pfeiffer, 1845) (Mollusca, Gastropoda, Pulmonata). — 

Netherlands Journal of Zoology 27: 271-276. 
van Mol, J. -J. 1971. Notes anatomiques sur les BuHmulidae (Mollusques, Gastropodes, Pulmones). — 

Annales de la Societe Royale Zoologique de Belgique 101:183-225. 
Pilsbry, H. A. 1946. Land Mollusca of North America (north of Mexico). — Monographs (3) Academy 

of Natural Sciences of Philadelphia 2(1): 1-520. 
Solem, A. 1955. Mexican molluscs collected for Dr. Bryant Walker in 1926, IX. Drymaeus. — Oc- 
casional Papers of the Museum of Zoology University of Michigan (566): 1-20, pis. 1-5. 
Strebel, H. 1882. Beitrag zur Kenntniss der Fauna mexikanischer Land — und Susswasser-Conchy- 

lien. Pt. 5. Orthalicidae. 1-144, pis. 1-19.— Hamburg. 

Florida State Museum, University of Florida, Gainesville, Florida 32611. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, p. 269 

YOCHELSONIELLA, NOM. NOV., A NEW NAME FOR 

ELLISELLA ROHR, 1980 (GASTROPODA) 

NON GRAY, 1858 (COELENTERATA) 

David M. Rohr and Richard W. Huddleston 



Rohr (1980) erected the new genus Ellisella (type-species E. greggi Rohr, 1980) 
for Middle-Upper Ordovician Archaeogastropoda from the Kangaroo Creek For- 
mation, California. Rohr (1980) tentatively assigned this form to the family Euom- 
phalidae. Gray (1858) previously used the name Ellisella for Recent anthozoan 
coelenterates. This renders Ellisella Rohr, 1980 a junior homonym of Ellisella 
Gray, 1858 and requires a new name (International Code of Zoological Nomen- 
clature Article 53, 60a). 

We propose the new name Yochelsoniella to replace Ellisella Rohr, 1980 non 
Gray, 1858. 

Yochelsoniella is characterized by possessing "Nearly flat-spired, rounded 
nodes at upper-outer angulation; small rounded tubes at mid- whorl; subovate 
whorl section; widely umbilicate" (Rohr 1980:166). 

Yochelsoniella is constructed in honor of Dr. ElHs Yochelson for his contri- 
butions in Paleontology. 

Acknowledgments 
We thank Chevron Oil Field Research Company for their assistance. 

Literature Cited 

Gray, J. E. 1858. Synopsis of the families and genera of axiferous zoophytes or barked corals. — 

Proceedings of the Zoological Society of London 25 (1857):278-294. 
Rohr, D. M. 1980. Ordovician-Devonian Gastropoda from the Klamath Mountains, California. — 

Palaeontographica Abteilung A 171:141-199. 

(DMR) Department of Geology, Sul Ross State University, Alpine, Texas 
79830; (RWH) Chevron Oil Field Research Company, La Habra, California 90632 
(current address: Scientific Research Systems, 11044 McGirk, El Monte, Cali- 
fornia 91731). 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 270-281 

PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS 
I. THE MARSHALL ISLANDS 

C. Allan Child 

Abstract. — This first report on Pycnogonida from the Marshall Islands lists 6 
shallow water species. These are: Ammothella stauromata, new species; Ano- 
plodactylus glandulifer Stock, previously known from the Red Sea to Singapore; 
Anoplodactylus marshallensis , new species; Endeis nodosa Hilton, known only 
from Hawaii in past reports, with a male figured for the first time; specimens very 
near Callipallene novae zealandiae (Thomson), a species not known from the 
western north Pacific; and Nymphon micronesicum, new species. The new 
species are apparently not closely related to Pacific mainland pycnogonids. 



There are no previous reports on pycnogonids from the Marshall Islands and, 
indeed, practically nothing is known of the pycnogonids inhabiting the vast ex- 
panse of Micronesia. Only 2 Micronesian species are reported in the literature, 
each from a single locality in the Caroline Islands (Stock 1968:10, 49). Tropical 
Pacific island pycnogonids have received some study in limited localities by Lo- 
man (1908) for Indonesia, Stock (1953) for Indonesia and the Philippines, Hilton 
(1942a) for Hawaii, and Stock (1968) for several Pacific localities, with a few short 
reports by others (Child 1970, Society Islands and Tuamotus; Child and Hedgpeth 
1971, Galapagos; Clark 1973, New Britain and Tonga; Child 1977, French Ocean- 
ia). The pycnogonid fauna of Australia, New Zealand, and continental Pacific 
borders is better known, probably because it is more accessible to collectors. 

The Micronesian fauna could be expected to contain species common to other 
parts of the tropical Pacific and at least 3 of the Marshall Island species reported 
here are found elsewhere in the Pacific. The remaining 3 species are new, reflect- 
ing the lack of benthic collections from Micronesia. The atolls and small volcanic 
islands of Micronesia are of particular interest to the taxonomist because of the 
restriction or absence of habitats such as muddy estuaries, algal encrusted rocky 
shores, and beds of sea grasses. These restrictions limit many marine faunules, 
including pycnogonids, to coral or coral sand and rubble, sponges, and a few 
other sessile sources of food. Coelenterates are a preferred food of many pyc- 
nogonids, many of which have been collected in association with coral reefs. I 
predict that there will be a much larger pycnogonid fauna found associated with 
the reefs of the Marshall Islands and other Pacific islands and atolls than is 
described in this short report. 

The majority of these specimens were collected by the author during a Smith- 
sonian Enewetak Atoll survey in 1969. Additional specimens were loaned by the 
B. P. Bishop Museum of Honolulu, Hawaii. 

I am grateful for field support and transportation supplied by the Atomic Energy 
Commission who operated the Enewetak Marine Biological Laboratory under the 
management of the University of Hawaii, for base support supplied by Kentron, 
Inc., and the U.S. Air Force, and for the field support of my fellow divers from 
the Smithsonian and the U.S. Geological Survey. I wish to thank Dr. Dennis M. 



VOLUME 95, NUMBER 2 271 

Devaney of the B. P. Bishop Museum for loaning the additional specimens. The 
specimens marked (BPBM) are deposited in the Bishop Museum. All other spec- 
imens are deposited in the National Museum of Natural History under the catalog 
numbers of the old U.S. National Museum (USNM). I wish to thank Dr. Thomas 
E. Bowman, Department of Invertebrate Zoology (Crustacea), National Museum 
of Natural History, for critically reviewing the manuscript. 

Family Ammotheidae 

Ammothella stauromata, new species 

Fig. 1 

Material examined. — N end of Enewetak Island, on pilings of large marine pier 
in 0.3-1.8 m, ll°2r48"N, 162°21'10"E, Child sta. 39-69, 12 Oct. 1969, 1 male 
holotype, USNM 189275, and 6 juvenile paratypes, USNM 189276. Pilings of 
small wooden pier at N end of Enewetak Island in 1.5 m, 11°21'52"N, 162°21'15"E, 
Child sta. 25-69, 29 Sept. 1969, 7 juvenile paratypes, USNM 189277. 

Description. — Moderately small, adult leg span just under 14 mm. Trunk As- 
corhynchus -likQ with slender median tubercles taller than their segment diameter. 
Trunk segmentation flaring with each segment anterior inserted into posterior 
large cowl of each preceding segment. Lateral processes separated by distances 
slightly greater than their diameters, twice as long as their maximum diameters, 
and armed with short slender dorsodistal tubercles and lateral setae shorter than 
segment diameter. Lateral setae arranged only on posterior 6 lateral processes: 
1 on posterior of second lateral processes; 1 on anterior and posterior of third 
and fourth processes. First lateral processes without setae. Ocular tubercle a 
cylinder as tall as median trunk tubercles, placed at anterior of cephalic segment, 
with darkly pigmented eyes at rounded tip. Abdomen cylindrical, slightly swollen 
distally, longer than median tubercles, carried almost erect, armed with several 
short distal setae. 

Proboscis moderately inflated with broad flat lips just distal to slight constric- 
tion. 

Chelifores 3-segmented, slender, with small ovoid vestigial chelae having only 
hint of fingers. First segment short, with slender dorso-distal tubercle longer than 
segment diameter, without setae. Second segment 2.5 times length of first, with- 
out tubercles but with fringe of distal setae as long or longer than segment di- 
ameter. 

Palps 9-segmented, originating from bulbous tubercles placed anterolaterally 
on cephalic segment ventral to ocular tubercle. Each bulbous tubercle with a 
slender, short, obliquely-pointing tubercle. Palp slender, second and fourth seg- 
ments with single long lateral seta and several short setae. Distal segments with 
many short ventral and lateral setae. 

Oviger 10-segmented, originating ventral to first lateral processes. First 5 seg- 
ments with few short setae, sixth with 8 or 9 short setae, seventh with 2 long 
lateral setae, and terminal 3 segments (strigilis) with 2 denticulate spines on each 
segment. Spines with many serrations per side. 

Third leg: first coxa with slender dorsodistal tubercle not as long as segment 
diameter, with shorter tubercle anteriolateral to this, segment armed with ante- 
rior, posterior and ventral setae. Coxa 2 with 1 dorsal seta and several short distal 
setae, with slender ventrodistal sexual pore tubercle having several distal setae. 



272 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Ammothella stauromata, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view; 
c, Third leg with enlargement of cement gland; d. Terminal segments of third leg, enlarged; e. Palp; 
f, Oviger; g. Terminal segments of oviger, enlarged; h. Juvenile chelafore. 



VOLUME 95, NUMBER 2 273 

Tubercle almost as long as segment diameter. Coxa 3 with several ventral setae. 
Femur armed with single long anterior and posterior midlateral setae, several 
ventral short setae, fringe of distal setae, and dorsodistal broad tubercle carrying 
cement gland tube along its dorsal length. Tibia 1 the longest segment, tibia 2 
shorter than femur, both armed with few dorsal and lateral setae longer than 
segment diameter and several shorter ventral setae. Tarsus very short, triangular, 
with 1 dorsal seta and 4 or 5 ventral short setae. Propodus robust, strongly curved, 
armed with 3 stout heel spines, 10 or 11 short sole spines, several dorsal setae, 
the longest slightly longer than segment diameter, and a fringe of distal setae. 
Claw moderately curved, about 0.4 length of propodus. Auxiliary claws almost 
as long as main claw. 

Measurements ofholotype (in mm). — Trunk length (tip of ocular tubercle to tip 
4th lateral processes), 1.74; trunk width (across 2nd lateral processes), 1.06; pro- 
boscis (lateral), 1.23; abdomen (lateral), 0.71; 3rd leg, coxa 1, 0.32; coxa 2, 0.69; 
coxa 3, 0.47; femur, 1.35; tibia 1, 1.42; tibia 2, 1.25; tarsus, 0.18; propodus, 0.68; 
claw, 0.27. 

Distribution. — Known only from the type-locahty, Enewetak Island, Enewetak 
Atoll, in the intertidal. 

Etymology. — The species name is Greek and means a palisade or stockade, 
this pertaining to the palisaded appearance of the tall ocular tubercle, median 
trunk tubercles and abdomen as seen in lateral view. 

Remarks. — There are few Ammothella species with tall median trunk tubercles. 
Those that appear to be closest to this new species are Ammothella setosa Hilton, 
1942b, A. menziesi Hedgpeth, 1951, A. thetidis Clark, 1963, and A. exornata 
Stock, 1975b. Of these, A. stauromata is least like A. exornata, a very small 
compact species with many coxae, chelifore and lateral process tubercles and a 
tuberculate ocular base. The median trunk tubercles of A. menziesi are shorter 
than the ocular tubercle and are blunt finger-like projections, unlike those of A. 
stauromata. Hedgpeth' s species is also notably larger than A. stauromata. The 
2 species most similar to this new species are A. setosa and A. thetidis. They are 
both slender graceful species with lateral processes well separated, but unlike A. 
stauromata, neither has a median trunk tubercle on the cephalic segment and 
their chelifores are much longer and more slender. 

Family Phoxichilidiidae 
Anoplodactylus glandulifer Stock 

Anoplodactylus spec. Caiman, 1923:289. 

Anoplodactylus glandulifer Stock, 1954:80-84, fig. 36; 1958:3; 1968:49; 1974:16- 
17._Arnaud, 1973:955, figs. 1, 2. 

Material examined. — N end of Enewetak Island, pier pilings of large marine 
pier in 0.3-1.8 m, 11°21'48"N, 162°21'10"E, Child sta. 39-69, 12 Oct. 1969, 1 6 
with eggs. 

Remarks. — This single male (a larval specimen accompanied the male, but it 
is too juvenile for determination and may not be this species) is slightly smaller 
than the measurements given by Stock (1954:84). It varies in other small details: 
the lateral processes are more closely spaced; there are fewer teeth on the chela 



274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

fingers; there are only very low palp buds with no discernable segmentation line; 
the propodal lamina is shorter, but is still longer than half the sole length; and 
the second oviger segment is longer than that shown for the type. There are also 
3 cement glands on each femur of all legs. Stock found either 2 or 3 glands on 
legs of the type. This egg-carrying specimen is otherwise the same as the type- 
specimen, also a male. 

The known distribution of this species, previously from the Red Sea and east 
coast of Africa to Burma and Singapore in depths from littoral to 5 meters, is now 
extended to the Marshall Islands littoral. 

Anoplodactylus marshallensis , new species 
Fig. 2a-f 

Material examined. — N end of Enewetak Island, on pier pilings of large marine 
pier in 0.3-1.8 m, ir21'48"N., 162°21'10"E., Child sta. 39-69, 12 Oct. 1969, 1 
c^ with eggs, holotype (USNM 189278), 2 S, I 9 ovigerous, 1 $, 4 juveniles, 
paratypes (USNM 189279). N end of Enewetak Island, pilings of small wooden 
pier in 1.5 m, ll°2r52"N, 162°21'15"E, Child sta. 25-69, 29 Sept. 1969, 1 c^ with 
eggs, 1 larva, paratypes (USNM 189280). 

Description. — Very small, leg span only slightly over 4 mm. Trunk robust, 
ovoid in dorsal aspect, unsegmented. Lateral processes glabrous, separated by 
distances equal to half their diameter, each only slightly longer than its diameter. 
Ocular tubercle a moderately tall cone inflated at midlength with unpigmented 
eyes, with lateral papillae distal to eyes. Neck fairly short, without setae. Ab- 
domen slightly shorter than ocular tubercle, curved dorsally, armed with 3 or 4 
distal setae. 

Proboscis a cylinder tapering distally to rounded lips, without constrictions. 

Palps entirely lacking. 

Chelifores robust, club-shaped, curved ventrally in lateral aspect, armed with 
1 or 2 dorsodistal setae. Chela tiny, palm ovoid, longer than fingers, armed with 
3 or 4 short distal setae. Immovable finger slightly curved, with 1 seta, movable 
finger with greater curve, crossing immovable finger at tip, armed with 3 setae. 
Fingers without teeth. 

Oviger moderately short, second segment curved, third straight and the longest 
segment, both armed with several short setae. Fourth segment only slightly longer 
than fifth, sixth round, no longer than wide. Fifth and sixth armed with several 
setae longer than segment diameters. 

Legs moderately short, armed with very few setae and a very long dorsodistal 
seta on each major segment. Femur the longest segment, with tibia 1 longer than 
tibia 2. Single cement gland a flask-shaped distally-pointing tube shorter than 
segment diameter. Body of tube creased with many circumferential constrictions 
when seen under high magnification. Tarsus very short, almost triangular, armed 
with 3 or 4 short ventral setae. Propodus robust, almost straight, with marked 
heel bearing 1 stout spine and 4 short setae. Sole armed with 5 or 6 short curved 
spines, a very short lamina and several lateral and distal setae. Claw moderately 
curved, robust, two-thirds propodal length. Auxiliary claws entirely lacking. 

Measurements of holotype (in mm). — Trunk length (chelifore insertion to tip 
4th lateral processes), 0.68; trunk width (across 1st lateral processes), 0.45; pro- 



VOLUME 95, NUMBER 2 275 

boscis length (lateral), 0.28; abdomen length (lateral), 0.14; 3rd leg, coxa 1, 0.14; 
coxa 2, 0.24; coxa 3, 0.14; femur, 0.38; tibia 1, 0.33; tibia 2, 0.28; tarsus, 0.06; 
propodus, 0.25; claw, 0.17. 

Distribution. — Known only from the type-locality, Enewetak Island, Enewetak 
Atoll, in littoral depths. 

Etymology. — Named for the Marshall Islands, its type-locality. 

Color notes. — When freshly collected, all 11 specimens displayed a distinctive 
red color pattern. The white trunk bore a purplish red longitudinal stripe mid- 
dorsally from the abdomen tip to the posterior surface of the ocular tubercle. 
This stripe was bordered in darker red and was a dark blood-red at the ocular 
tubercle. The tibiae each had the same dorsal coloration with tibia 2 slightly 
darker. The remainder of the legs and all other appendages were white. After 
several months in alcohol, these colors had entirely disappeared. The coloration 
appeared to be in the chitinous integument itself and not associated with the gut 
diverticula where coloration is more often found among the pycnogonids. 

Remarks. — This small species is without many distinctive characters except 
for the coloration described above. It appears to be morphologically related to 
Anoplodactylus quadratispinosus Hedgpeth, 1943, and to A. inermis Losina-Lo- 
sinsky, 1961. Both share with the new species the characters of a tall conical 
ocular tubercle, a distinctive large heel on the propodus, and a trunk with lateral 
processes separated by distances equal to or less than their diameter. Several 
other characters immediately separate this new species from the others. It has 
much shorter leg segments, shorter and more robust chelifores, a smaller pro- 
boscis, shorter lateral processes, and an unsegmented trunk. Losina-Losinsky's 
species is only known from females, but the characters above would serve equally 
to separate A. marshallensis from a male of A. inermis. 

The small robust shape of this new species is so far unique to the genus An- 
oplodactylus from the western Pacific. 

Family Endeidae 

Endeis nodosa Hilton 

Fig. 2g-i 

Endeis (Phoxichilus) nodosa Hilton, 1942a:47-48, fig. 4. 
Endeis nodosa. — Stock, 1968:59 [key]. 

Material examined. — Runit (site name Yvonne) Island, pier pilings on lagoon 
side in 0-0.7 m, ir32'40"N, 162°2r53"E, Child sta. 36-69, 8 Oct. 1969, 2 $ 
(USNM). Enewetak Island, on lagoon rocks in 4 meters, J. Lamberson, Jan. 
1976, 1 9 (BPBM). Hawaii, Honolulu Harbor on boat hawser, O. Degener, Apr. 
1947, 1 6 with eggs (USNM). 

Remarks. — This species has never been adequately figured and since the male 
has never been recorded in the literature, I include a set of figures here of the 
Honolulu male specimen. The species is rather distinctive in a genus with many 
species difficult to separate. The intestinal diverticula in each leg have many blind 
caeca or pockets and the femur has a large ventral triangular projection on each 
leg. Endeis flaccida Caiman, 1923, is a similar species, but is without the ventral 
femoral projection. 



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Fig. 2. Anoplodactylus marshallensis, holotype, male; a, Trunk, dorsal view; b, Trunk, lateral 
view; c. Chela; d, Third leg with enlargement of cement gland; e, Terminal segments of third leg, 
enlarged; f, Oviger. Endeis nodosa, male: g. Trunk, dorsal view; h, Third leg, showing gut diverticula; 
i. Terminal segments of oviger with eggs. 



VOLUME 95, NUMBER 2 277 

This Species is probably not rare but only rarely reported because its habitats 
are seldom sampled. It was known previously only from Hawaii in the littoral 
and its distribution is now extended westward to the Marshall Islands at Enewetak 
Atoll. 

Family Callipallenidae 
Callipallene sp. cf. C. novae zealandiae (Thomson) 

Pallene novae-zealandiae Thomson, 1884:246-247, pi. 14, figs. 1-4. — Hutton, 

1904:247. 
Callipallene brevirostris ssp. novae-zealandiae . — Stock, 1954:48-50, fig. 21a-h. 
Callipallene brevirostris cf. ssp. novaezealandiae . — Arnaud, 1972:162. 
Callipallene brevirostris novaezealandiae . — Child, 1975:10. 
Callipallene brevirostrum ssp. novaezealandiae . — Stock, 1975a: 132. 

Material examined. — Large coral knoll in lagoon near Chinieero (site name 
Alvin) Island in filamentous green algae at 3-5 m, 11°29'05"N, 162°23'25"E, Child 
sta. 05-69, 22 Sept. 1969, 1 9 (USNM). Enewetak Island, lagoon side, in rubble 
at 4 m, Brock, 27 Sept. 1975, 1 S (BPBM). 

Remarks. — These 2 specimens are quite close to Thomson's species and agree 
for the most part with his figures. The ovigers bear fewer denticulate spines than 
either Thomson's figure 3, or Stock's descriptions of his specimens. The speci- 
mens in hand have a denticulate spine count of 6:5:5:5, and 6:5:5:6, while Thom- 
son's female has a count of 8:8:7:8, and Stock's male has 8:8:8:7 and his female 
8:7:6:8 spines. This is the only significant difference I can find between these 
Enewetak specimens and the detailed figures and descriptions given by Thomson 
and Stock. It is possible that these specimens represent a new species, but with 
only 2 specimens in hand and the close agreement between these and most char- 
acters of Thomson's species, I regard them provisionally as near to, if not con- 
specific with his New Zealand species. 

This species has never been reported from tropical western Pacific localities. 
Its known distribution has been from New Zealand and Australia to the western 
Indian Ocean at Madagascar and Tanzania, in mostly shallow depths. If these 
specimens are in fact Thomson's species, then the distribution is extended to the 
northern hemisphere for the first time, in the Marshall Islands. 

Family Nymphonidae 

Nymphon micronesicum, new species 

Fig. 3 

Material examined. — Bikini Atoll, Namu Island, outer reef, 11°42'N, 165°17'E, 
M. W. Johnson, 4 Apr. 1946, 1 $, holotype (USNM 189274). 

Description. — Female (male unknown): of moderately small size, leg span al- 
most 15 mm. Trunk robust, segments slightly inflated, fully segmented. Lateral 
processes separated by distances slightly greater than or equal to their diameters, 
only as long as their diameters, armed with few tiny setae. Neck moderately long, 
with segmentation line (possible artifact) at narrowest diameter directly anterior 
to oviger implantation which is slightly anterior to but not touching first lateral 



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Fig. 3. Nymphon micronesicum, holotype, female: a, Trunk, dorsal view; b, Ocular tubercle, 
lateral view; c, Palp; d. Third leg with enlargement of ventral spine; e, Distal propodus and claws, 
enlarged; f. Chela; g. Fingers of chela, enlarged; h, Oviger; i, Terminal segment of oviger with a 
denticulate spine, enlarged. 



processes. Ocular tubercle slightly taller than its diameter, implanted over first 
lateral processes, rounded, with 2 lateral papillae and tiny apical papilla. Eyes 
large, slightly pigmented. Abdomen short, erect, no taller than ocular tubercle, 
armed with 2 laterodistal setae. 



VOLUME 95, NUMBER 2 279 

Proboscis a slightly swollen cylinder tapering distally to rounded lips. 

Chelifore scapes large, cylindrical, 4 times as long as their diameters, armed 
with several lateral setae. Chela shorter than scape, palm longer than fingers, 
armed with several dorsal and distal fringe of setae. Fingers broad, immovable 
finger straight, armed with 11 or 12 bicuspidate teeth. Movable finger moderately 
curved, armed with 12 or 13 bicuspidate teeth. Finger tips not overlapping. 

Palp short, second segment longest, terminal segment only slightly shorter. 
Third segment 0.6 length of second, armed with ventrodistal setae. First and 
fourth segments only slightly longer than wide. Fourth and fifth segments armed 
with many setae slightly longer than segment diameter. 

Oviger segments 4 and 5 subequal, armed with several distal setae. Sixth seg- 
ment cylindrical, 0.5 length of fourth, armed with lateral and dorsal setae. Strigilis 
4 segments armed with dorsal and lateral setae and denticulate spines in the 
formula 11:9:9:10. Spines with 2 proximal large serrations and several smaller 
distal serrations per side. Terminal claw as long as terminal segment, without 
endal teeth or denticulations. 

Leg segments slender, tibia 2 the longest segment, 0.3 longer than tibia 1 and 
femur which are subequal in length. Major segments armed with short dorsal and 
ventral setae increasing in numbers distally. Tibia 2 armed with 2 ventrodistal 
spines, each with several endal crenulations. Tarsus and propodus short; tarsus 
0.4 length of propodus, both armed with many slender setae and broader spines 
on sole. Propodus without heel or larger heel spines. Major claw short, very 
curved, armed with 2 rows of low conical rugosities endally. Auxiliary claws 
longer than main claw, each armed with 2 rows of tiny serrations or teeth endally. 

Measurements (in mm). — Trunk, first segment, 0.89; posterior 3 segments, 0.7; 
width (across 2nd lateral processes), 0.67; proboscis length, 0.67; abdomen 
length, 0.16; 3rd leg, coxa 1, 0.23; coxa 2, 0.64; coxa 3, 0.4; femur, 1.4; tibia 1, 
1.48; tibia 2, 2.08; tarsus, 0.2; propodus, 0.47; claw, 0.1; auxiliary claws, 0.13. 

Distribution. — Known only from the type-locality, Namu Island, Bikini Atoll, 
in a shallow but unrecorded depth. 

Etymology. — Named for Micronesia, a place of "small islands," a number of 
which form Bikini Atoll, the type-locality for this species. 

Remarks. — Nymphon micronesicum is morphologically similar to A^. giraffa 
Loman, 1908, N. aequidigitatum Haswell, 1884, N. floridanum Hedgpeth, 1948, 
and perhaps is closest to N. biformidens Stock, 1974. It shares with the latter 
species a short main propodal claw with longer auxiharies with all claws having 
endal rugosities or tiny denticles, legs and palps with very similar length propor- 
tions, and similar trunk and chelifore shapes. The new species differs from N. 
biformidens in having a taller ocular tubercle with prominent lateral papillae, a 
more cylindrical proboscis, a larger oviger claw without denticulations or rugos- 
ities, longer 4th and 5th oviger segments, and more teeth on both fingers of the 
chela with all chela teeth bicuspidate. 

The other 3 similar species share short main claws with longer auxiliaries (those 
of N. aequidigitatum lack endal rugosities), long palp terminal segments, biden- 
tate chela fingers (except for N. giraffa), long second tibiae, and long oviger 
terminal claws without denticulations. The new species differs from N. giraffa 
in having a more compact trunk and appendages, a chela palm which is more 
cylindrical than globular, a much shorter tarsus in relation to propodal length. 



280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

and very different oviger denticulate spines. Both of these species have a con- 
striction or segmentation line around the narrowest part of the neck. This is 
possibly an artifact due to repeated bending either during the life of the animal 
or during capture. 

Nymphon micronesicum differs from N. aequidigitatum in having more slender 
and shorter chela fingers with many more rounded teeth, a longer tarsus in relation 
to propodal length, a longer palp second segment, and a different trunk config- 
uration. The new species differs from A'^. floridanum in having a shorter neck 
without parallel sides, shorter chela fingers with fewer bicuspidate teeth, a more 
cylindrical proboscis, and a terminal palp segment longer than the third segment 
and almost equal to the second. 

No other known Pacific species of Nymphon is as similar to this new species 
as are the 4 above. 

Literature Cited 

Amaud, F. 1972. Pycnogonides des recifs coralliens de Madagascar. 3. Famille des Callipallenidae. — 

Tethys, Supplement 3:157-164, figures 1-6. 
. 1973. Pycnogonides des recifs coralliens de Madagascar. 4. Colossendeidae, Phoxichilidiidae 

et Endeidae.— Tethys 4(4):953-960, 8 figures. 
Caiman, W. T. 1923. Pycnogonida of the Indian Museum. — Records of the Indian Museum 25(3):265- 

299, 17 figures. 
Child, C. A. 1970. Pycnogonida of the Smithsonian-Bredin Pacific Expedition, 1957. — Proceedings of 

the Biological Society of Washington 83(27): 287-308, 5 figures. 
. 1975. Pycnogonida of Western Australia. — Smithsonian Contributions to Zoology 190:1-28, 

11 figures. 
. 1977. On some Pycnogonida of French Oceania. — Proceedings of the Biological Society of 

Washington 90(2):440^46, 1 figure. 
, and J. W. Hedgpeth. 1971. Pycnogonida of the Galapagos Islands. — Journal of Natural His- 
tory 5:609-634, 8 figures. 
Clark, W. C. 1963. Australian Pycnogonida. — Records of the Australian Museum 26(1): 1-81, 38 

figures. 
. 1973. New species of Pycnogonida from New Britain and Tonga. — Pacific Science 27(1):28- 

33, 3 figures. 
Haswell, W. A. 1884. On the Pycnogonida of the Australian coast, with descriptions of new species. — 

Proceedings of the Linnean Society of New South Wales 9:1021-1034, plates 54-57. 
Hedgpeth, J. W. 1943. Reports on the scientific results of the Atlantis expeditions to the West Indies, 

under the joint auspices of the University of Havana and Harvard University. — Proceedings 

of the New England Zoological Club 22:41-58, plates 8-10. 
. 1948. The Pycnogonida of the western north Atlantic and the Caribbean. — Proceedings of 

the United States National Museum 97 (3216): 157-342, figures 4-53, 3 charts. 
. 1951. Pycnogonids from Dillon Beach and vicinity, California, with descriptions of two new 

species. — Wasmann Journal of Biology 9(1): 105-1 17, 3 plates. 
Hilton, W. A. 1942a. Pycnogonids from Hawaii. — Occasional Papers from the B. P. Bishop Museum, 

Honolulu, Hawaii 17(3):43-55, 10 figures. 
. 1942b. Pycnogonids from the Pacific. Family Ammotheidae. — Pomona Journal of Entomology 

and Zoology 34(4): 93-97. 
Hutton, F. W. 1904. Pp. 246-247 in Index Faunae Novae Zealandiae. — London, Dulau. 
Loman, J. C. C. 1908. Die Pantopoda der Siboga-Expedition. — Siboga-Expeditie Monograph 40:1- 

88, 15 plates. 
Losina-Losinsky, L. 1961. [Pantopoda of the Far Eastern Seas of the U.S.S.R.] — Issledovanniya 

Dalnyevostochnykh Morei SSSR, Leningrad 7:47-117, 27 figures. [In Russian]. 
Stock, J. H. 1953. Biological results of the SneUius Expedition XVII. Contribution to the knowledge 

of the pycnogonid fauna of the East Indian Archipelago. — Temminckia 9:276-313, 18 figures. 



VOLUME 95, NUMBER 2 281 

. 1954. Pycnogonida from Indo-West-Pacific, Australian, and New Zealand waters. — Viden- 

skabelige Meddelser fira Dansk Naturhistorisk Forening (Copenhagen) 116:1-168, 81 figures. 

. 1958. The Pycnogonida of the Erythrean and of the Mediterranean coasts of Israel. Contri- 
butions to the knowledge of the Red Sea. 5. — Bulletin of the Sea Fisheries Research Station, 
Haifa 16:3-5. 

. 1968. I*ycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific 

Oceans. — Videnskabelige Meddelser fi-a Dansk Naturhistorisk Forening (Copenhagen) 131:7- 
65, 22 figures. 

. 1974. Medio- and infralittoral Pycnogonida collected during the I.I.O.E. near the landbase 



on Nossi-Be, Madagascar. — Bulletin of the Zoological Museum, University of Amsterdam 
4(3): 11-22, 4 figures. 

. 1975a. Infralittoral Pycnogonida from Tanzania. — Travais du Museum d'Histoire Naturelle 

"Gr. Antipa," Bucharest 16:127-134, 12 figures. 

. 1975b. Biological Results of the University of Miami Deep-Sea Expeditions, 108. Pycnogon- 
ida from the Continental Shelf, Slope, and Deep Sea of the Tropical Atlantic and east Pacific. — 
Bulletin of Marine Science 24(4): 957- 1092, 59 figures. 

Thomson, G. M. 1884. On the New Zealand Pycnogonida with descriptions of new species. — Trans- 
actions and Proceedings of the New Zealand Institute 16(1 883): 242-248, plates 14-16. 

Department of Invertebrate Zoology (Crustacea), NHB W 323-163, Smithson- 
ian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 282-291 

THREE NEW SPECIES OF PYCNOGONIDA 
FROM SAGAMI BAY, JAPAN^ 

Koichiro Nakamura and C. Allan Child 

Abstract. — Three new species of Pycnogonida are described from Sagami Bay, 
Japan; Ascorhynchus utinomii, Anoplodactylus shimodaensis and Anoplodacty- 
lus perforatus, and their relationships to known species in the two genera are 
discussed. A short resume of pycnogonid literature on Sagami Bay is given. 



Introduction 

Ohura or Oura Bay, locally called Nabeta Bay (34"39'30"N, 138°56'40"E), and 
the immediate vicinity, is the collecting locality for 3 new pycnogonid species. 
This locality is near Shimoda and is situated on the southwestern side of Sagami 
Bay, an area that has received substantial coverage in pycnogonid literature. 
Utinomi (1971), in his summary of known Japanese pycnogonids to that date, 
listed 32 species as residents of Sagami Bay and he stated (p. 340) that "Sagami 
Bay and its adjacent waters is . . . the richest of all regional faunae in Japanese 
waters, as represented by more than 17 genera and 46 species." Fourteen of the 
species listed are from off Sagami Bay proper, in deeper water. 

The early literature on pycnogonids of the area includes Bohm (1879) who 
described Ammothea hilgendorfi, Ascorhynchus ramipes and Propallene longi- 
ceps, all from Enoshima. Ortmann (1890) described Anoplodactylus gestiens, 
Ascorhynchus cryptopygium and A. bicornis (=A. auchenicum (Slater)), and list- 
ed Ascorhynchus ramipes and Pycnogonum littorale var. tenue (=P. tenue (Sla- 
ter)) from Sagami Bay. Ives (1891) described Ascorhynchus japonicum from Ja- 
pan, listing many collecting localities, one of which was Sagami Bay. Loman 
(1911) described 4 new species and listed 9 other species from Sagami Bay. 

Several other authors, among them Ohshima (1936), listed subsequent records 
of one or more species from this area. The next significant paper was by Hedgpeth 
(1949), in his report on Japanese pycnogonids collected by the Albatross. He 
listed 3 new and 3 known species from Sagami Bay (including adjacent Tokyo 
Bay). Stock (1954) and Utinomi (1959, 1962, 1971) hsted and described many 
additional species from Sagami Bay. 

Very little collecting has been done, or at least recorded in the literature, 
recently for shallow water pycnogonid species in Sagami Bay, but the senior 
author gathered a large collection of pycnogonids from there as a secondary result 
of collecting live specimens for purposes of embryological studies. Most of the 
specimens were taken from shallow water, averaging about 10 meters in depth, 
in the vicinity of the Shimoda Marine Research Center, the University of Tsukuba 
(formerly Shimoda Marine Biological Station, Tokyo Kyoiku University). Among 
the specimens collected were the three new species of pycnogonids described in 
this paper. 



Contribution No. 378 from the Shimoda Marine Research Center, the University of Tsukuba. 



VOLUME 95, NUMBER 2 283 

The types are deposited in the collections of the National Museum of Natural 
History, Smithsonian Institution, under the catalog numbering system of the 
United States National Museum, except for several paratypes retained by the 
senior author to be deposited in Japanese collections. 

Pycnogonida 

Family Ammotheidae Dohrn, 1881 

Ascorhynchus Sars, 1877 

Ascorhynchus utinomii, new species 

Fig. 1 

ry/7^5.— Holotype (USNM 184555), male with eggs, coll. Nabeta Bay, 7 to 15 
meters, 10 July 1970. Paratypes (USNM 184556), 2 males with eggs, 3 females, 
same collection as holotype. Paratypes (USNM 184557), 3 males with eggs, 1 
male, 3 females, coll. Nabeta Bay, 7 to 15 meters, 13 June 1970. Paratypes 
(USNM 184558), 3 males with eggs, 3 females, 1 juvenile, coll. Nabeta Bay, 7 to 
15 meters, 3 October 1969. Paratypes (KN collection), 1 male with eggs, 2 fe- 
males, coll. Nabeta Bay, 7 to 15 meters, 8 August 1973. 

Description. — Trunk elongate, lateral processes well separated by about their 
own diameter. Body smooth, without trunk tubercles, but with low rounded tu- 
bercle on distal end of all lateral processes. Anterior of cephalic segment without 
tubercles over chelifore insertion. Ocular tubercle placed over anterior of oviger 
bases, low, rounded at tip, with 4 large eyes lightly pigmented in alcohol. 

Proboscis almost half length of trunk, long and oval with one constriction 
towards base giving bilobed shape, rounded in cross section. 

Abdomen long, reaching almost to tips of second coxae of last pair of legs, 
swollen distally. 

Chelifore scape 1- segmented, longer than third palp segment, glabrous. Chela 
small with atrophied tiny curved fingers. 

Palp 10-segmented, basal 2 segments very small, not longer than wide, third 
segment longest. Fifth segment with one large seta endally and proximally, sev- 
eral shorter and 2 longer setae distally. Sixth to tenth segments densely setose 
ventrally, setae subequal in length. Terminal 4 segments long, slender. 

Oviger 10-segmented, with tiny terminal claw. Fourth and fifth segments long- 
est, armed with row of lateral setae. Terminal 4 segments with compound spines 
arranged in 2 or 3 rows, the longest row in the formula 12:9:8:10 for the holotype 
and 9:8:8:10 for a juvenile. 

Legs slender. First coxa with 2 dorsodistal tubercles. Coxa 2 with one low 
tubercle middorsally. Femur almost as long as combined length of coxae, armed 
with single dorsodistal seta. Cement glands not found. Tibia 1 the longest seg- 
ment, armed with few tiny setae dorsally. Tibia 2 about equal in length to femur. 
Tarsus slightly shorter than propodus, both without ventral setae or spines. An- 
terior pair of legs with longer tarsus and propodus, armed with extremely tiny 
terminal claw. Posterior 6 legs with shorter tarsus and propodus having claw 
almost equal to tarsus length. Male genital pores on posterior 4 legs only. In 
female, tiny pores without projections ventrally on second coxae of all legs. 

Measurements. — (in mm). First trunk segment length, 1.58; total trunk length 
(to tip 4th lateral processes), 3.46; trunk width (across 2nd lateral processes), 
1.92; proboscis length, 1.55; abdomen length, 1.03; third leg, coxa 1, 0.53; coxa 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Ascorhynchus utinomii, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view; c, 
Palp; d, Third leg; e. Distal segments of first leg with enlargement of tip; f, Distal chelifore, enlarged; 
g, Oviger; h, Oviger terminal segments, enlarged; i, Oviger tip, further enlarged. 



VOLUME 95, NUMBER 2 285 

2, 0.59; coxa 3, 0.43; femur, 1.22; tibia 1, 1.78; tibia 2, 1.27; tarsus, 0.66; prop- 
odus, 0.76; claw, 0.67. 

Distribution. — Known only from the type-locality, Nabeta Bay in Sagami Bay, 
in depths of from 7 to 15 meters. 

Etymology. — This proposed new species is dedicated to the late Dr. Huzio 
Utinomi, for his many contributions to our understanding of the pycnogonids of 
Japan. The Japanese compound word ""utinomi,'' translated literally (pronounced 
oo-chi-noh-me), means "inside the sea." This compound might in itself prove to 
be a good name for a new species, but the authors prefer the patronym for this 
new species. 

Remarks. — This species was collected together with Propallene longiceps and 
Ascorhynchus ramipes, but in fewer numbers than the others. When living, the 
proboscis is light yellow in color. 

This species is characterized by the shape of the proboscis, the location of the 
ocular tubercle in relation to oviger implantation, lack of dorsal trunk tubercles, 
and length of the abdomen. Its body size is almost the same as Ascorhynchus 
minutum, but A. utinomii proves to be distinct from A. minutum in the following 
details: the proboscis is bilobed with one constriction in the new species, but is 
trilobed with 2 constrictions in A. minutum, and in cross section, the proboscis 
is rounded in the new species and not triangular; the ocular tubercle is placed 
over the anterior of the oviger implantation bulges in A. utinomii, but is entirely 
anterior to the oviger bases in A. minutum; median trunk tubercles are absent in 
A. utinomii, but A. minutum has small median tubercles; the abdomen is longer 
in the new species, reaching almost to the tip of the second coxae of the fourth 
pair of legs, but is shorter and does not reach the distal end of the first coxae in 
A. minutum. 

Ascorhynchus utinomii differs from the other small Japanese species such as 
A. ramipes and A. auchenicum in its bilobed proboscis and lack of median trunk 
tubercles. The 2 latter species have a trilobed proboscis and median trunk tu- 
bercles of various sizes. 

Although the cement glands of this species are not evident, they are probably 
a row of extremely tiny pores such as those found on some of the larger species 
of the genus. 

Family Phoxichilidiidae Sars, 1891 

Anoplodactylus Wilson, 1878 

Anoplodactylus shimodaensis, new species 

Fig. 2 

Types.— Holoiype (USNM 184559), male with eggs, coll. Nabeta Bay, 7 to 15 
meters, 13 May 1970. Paratypes (USNM 184560), 2 males with eggs, 3 females, 
same collection as holotype. Paratype (USNM 184561), 1 male with eggs, coll. 
Nabeta Bay, 7 to 15 meters, 3 October 1969. Paratypes (USNM 184562), 1 male 
with eggs, 5 females, coll. off Kisami, 30 meters, 22 May 1970. Paratypes (USNM 
184563), 3 males, 1 female, coll. Nabeta Bay, 7 to 15 meters, 19 January 1971. 
Paratypes (KN collection), 3 males with eggs, 2 females, coll. Nabeta Bay, 7 to 
15 meters, 5 October 1970. 

Description. — Trunk robust, first and second segmentation lines complete. No 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Anoplodactylus shimodaensis , holotype, male: a, Trunk, dorsal view; b. Trunk, lateral 
view; c. Third leg, with enlargement of cement gland and propodal heel; d, Chela. 



VOLUME 95, NUMBER 2 287 

segmentation between third and fourth trunk segments. Lateral processes long, 
over twice as long as their diameters, separated by their diameters or less, without 
tubercles, armed with 3 or 4 dorsodistal setae. Fourth pair of lateral processes 
shortest. Neck narrow, ocular tubercle about 1.5 times as tall as width at base, 
rounded at tip, with low apical cone and 4 large lightly pigmented eyes. Lateral 
sensory papillae large, prominent. 

Proboscis short, robust, blunt at tip with constriction near distal fourth of 
length. 

Abdomen short, erect, approximately as tall as ocular tubercle, tapering to 
rounded tip, armed with several short setae. 

Chelifore scape straight proximally, strongly curved ventrodistally, armed with 
few setae. Chela small, with curved fingers overlapping at tips. Movable finger 
with 2 tiny tooth-like bumps, immovable finger without teeth. 

Oviger of 6-segments, third with proximal construction. Terminal 2 segments 
armed with many setae shorter than segment diameter. 

Legs moderately long. First coxa with distal setae, second coxa longest. Femur 
longest of major segments, armed with few setae and a long dorsodistal seta. 
Cement gland a stout tube shorter than segment diameter, opening dorsally at 
distal fourth of segment. First and second tibiae almost equal in length, armed 
with single long dorsodistal seta each. Tarsus short, almost triangular, armed 
with several ventrodistal setae. Propodus slightly curved, sole straight, with 
marked heel armed with 2 short stout spines and 2 setae. Sole with propodal 
lamina over entire length flanked by few short setae. Claw 0.7 propodal length, 
slender. Auxiliaries tiny. 

Measurements. — (in mm). Trunk length (without chelifores), 1.45; trunk width 
(across 2nd lateral processes), 1.26; proboscis length, 0.59; abdomen length, 0.24; 
third leg, coxa 1, 0.35; coxa 2, 0.53; coxa 3, 0.44; femur, 0.91; tibia 1, 0.82; tibia 
2, 0.79; tarsus, 0.11; propodus, 0.5; claw, 0.36. 

Distribution. — Known from the type-locality, Nabeta Bay on Sagami Bay in 
depths of from 7 to 15 meters. Also found off Kisami in 30 meters. 

Etymology. — Named for the town of Shimoda, near where the species was 
collected. The town of Shimoda was made famous, if fleetingly, as the first place 
where Commodore Perry landed in 1853, to open diplomatic relations with Japan. 

Remarks. — This species is easily distinguishable among Japanese species of 
Anoplodactylus by its small robust size and shape, the short barrel-shaped pro- 
boscis with construction, and the wide distally-placed tube of the cement gland 
in the male. No other known Japanese species in this genus has such robust 
lateral processes separated from each other by their own diameter, in combination 
with a short stout proboscis. The single femoral cement glands of this species are 
placed in a relatively rare distal position. Where the cement gland configuration 
is known, there are few species in this crowded genus which have the cement 
gland placed in the distal fourth or even third of the femur. We are unable to find 
any other known species of the genus from Japan or vicinity with a broad tubular 
cement gland placed this far forward on the femur. 

There are several known species of Anoplodactylus with similarities shared by 
this species, but none of them have the above characters in combination with 
chela fingers without teeth, no genital spurs on the second coxae of the third and 
fourth legs, and a moderate length ocular tubercle with conical apex. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Anoplodactylus perforatus, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view; 
c. Terminal segments of oviger, enlarged; d. Third leg, 3 cement glands enlarged; e. Third leg terminal 
segments; f, Propodal lamina enlarged; g, Chela. 



VOLUME 95, NUMBER 2 289 

Anoplodactylus perforatus , new species 
Fig. 3 

ry/7^5.— Holotype (USNM 184564), male with eggs, coll. Nabeta Bay, 7 to 15 
meters, 3 June 1970. Paratypes (USNM 184565), 1 male with eggs, 2 males, 2 
females, 1 juvenile male, same collection as holotype. Paratype (KN collection), 
1 male with eggs, coll. Nabeta Bay, 7 to 15 meters, 3 October 1969. Paratype 
(USNM 184566), 1 male, R/V Tansei Mam, cruise KT 69-12, sta. 21, 35°00'54"N, 
139°08'36"E, 113 meters, 17 July 1969. 

Description. — Trunk elongate, fully segmented between first 3 segments, third 
and fourth unsegmented. Lateral processes separated by less than their diameters, 
armed with 2 or 3 dorsodistal setae, without tubercles. Neck short, slender. 
Ocular tubercle rounded with conical apex, 1.5 times taller than width at base, 
with 4 large darkly pigmented eyes. 

Proboscis slender, blunt at tip, slightly constricted distally. 

Abdomen erect, tapering to rounded tip, glabrous, about equal in length to 
ocular tubercle. 

Chelifore scape slender, slightly curved laterally, armed with several dorsal 
setae. Chela large, palm cylindrical, with 2 or 3 setae. Fingers placed almost at 
right angles to palm, curved, overlapping at tips, armed laterally and dorsally 
with several setae, without teeth. 

Oviger 6-segmented, long, third segment longest, with proximal constriction. 
Distal 2 segments armed with many curved setae. Fifth segment armed with single 
short recurved spine endally. 

Legs slender. Coxae armed with several dorsal and ventrodistal setae. Second 
coxae without genital spur. Femur almost equal in length to tibia 1, both with 
small dorsodistal tubercle bearing a seta. Femoral cement gland openings dorsal, 
arranged in single row of cribellate pores numbering 17 to 25 per leg. First tibia 
slightly longer than second, both armed with many dorsal, lateral and ventral 
setae shorter than segment diameter and single dorsodistal seta longer than seg- 
ment diameter. Tarsus short, semitriangular, with several ventral and single dor- 
sal setae. Propodus moderately curved, with marked heel bearing 2 spines. Sole 
without spines proximally, with about 10 spines and several lateral setae along 
distal three-quarters of length and very short propodal lamina distally, armed 
dorsally and laterally with several setae. Terminal claw long, moderately curved, 
about two-thirds propodal length. Auxiliary claws tiny. 

Measurements. — (in mm). Trunk length without chelifores, 1.97; trunk width 
(across 2nd lateral processes), 1.27; proboscis length, 1.34; abdomen length, 0.37; 
third leg, coxa 1, 0.47; coxa 2, 0.9; coxa 3, 0.65; femur, 1.68; tibia 1, 1.64; tibia 
2, 1.4; tarsus, 0.13; propodus, 0.78; claw, 0.54. 

Distribution. — Known from the type-locality, Nabeta Bay in from 7 to 15 me- 
ters, and from off Shimoda in 113 meters. 

Etymology. — Named for the perforated appearance of the 17 to 25 cement 
glands along the dorsal surface of the femur. 

Remarks. — This species closely resembles Anoplodactylus gestiens, except 
that it is not quite as slender or tenuous and has many cement glands instead of 
the single gland. A set of figures of A. gestiens is included here (Fig. 4) because 
this species has never, in our opinion, been adequately figured in the literature. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Anoplodactylus gestiens, specimen from TS-69, station 21, male: a, Trunk, dorsal view; 
b, Trunk, lateral view; c, Third leg; d. Terminal segments of third leg; e. Chela. 



There is no known species of Anoplodactylus having so many cement glands 
per femur. The nearest species in number of cement glands is A. cribellatus which 
is reported in the literature to have fifteen or so per leg. All other known species 
have 10 or less glands per femur. The 14 known species of Anoplodactylus with 
multiple glands are as follows: A. cribellatus Carpenter, with about 15 glands; A. 
longiformis Child, with 8 to 10; A. australis (Hodgson), about 7; A. xenus Stock, 
about 8; A. pycnosoma (Heifer), from 5 to 8; A. multiclavus Child, 3 to 5; A. 
glandulifer Stock, 2 to 4; A. angulatus (Dohrn), 4 or 5; A. dentimanus Stock, 3; 
A. oculatus Carpenter, 5; A. virescens (Hodge), 2 or 3; A. robustus (Dohrn), 3; 
A. longiceps Stock, 2; and the present new species with from 17 to 25 glands. 



VOLUME 95, NUMBER 2 291 

There are several species of the closely related genus Phoxichilidium which have 
multiple cement glands on the legs of males. 

The proposed new species has no other conspicuous characters to set it apart 
from many of the genus, but the cement gland number and shape have proved to 
be sufficiently stable in providing a primary source of differentiation among the 
many species of Anoplodactylus. 

Acknowledgments 

We acknowledge the assistance and generosity of the Shimoda Laboratory 
staff, particularly Hajime Ueda, without whose help this collection of new species 
could not have been made. We are grateful also to Dr. Koichi Sekiguchi of 
the University of Tsukuba, and chief scientist of the Tansei Mam Expedition, 
for his collaboration in collecting material. 

Literature Cited 

Bohm, R. 1879. Uber zwei neue von Herrn Dr. Hilgendorf in Japan gesammelte Pycnogoniden. — 

Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1879(4): 53-60, figs. 

a-c. 
Hedgpeth, J. W. 1949. Report on the Pycnogonida collected by the Albatross in Japanese waters in 

1900 and 1906.— Proceedings of the United States National Museum 98(3231):233-321, figs. 

18-51. 
Ives, J. E. 1891. Echinoderms and arthropods from Japan. — Proceedings of the Academy of Natural 

Sciences of Philadelphia, 1891-1892:210-221, plate XII. 
Loman, J. C. C. 1911. Japanische Podosomata; Beitrage zur Naturgeschichte Ostasiens, herausgeben 

von F. Doflein. — Abhandlungen der Kaiserlich Bayerischen Akademie der Wissenschaften 

(Mathematisch-Naturwissenschaftliche Klasse), Suppl. 2(4): 1-18, pis. I-II. 
Ohshima, H. 1936. [A list of Pycnogonida recorded from Japanese and adjacent waters.] — Dobut- 

sugaku Zasshi 48(8- 10): 86 1-869. [In Japanese] 
Ortmann, A. E. 1890. Bericht iiber die von Herrn Dr. Doderlein in Japan gesammelten Pycnogoni- 
den.— Zoologische Jahrbucher (Systematik) 5(1): 157-168, pi. XXIV. 
Stock, J. H. 1954. Pycnogonida from Indo-West Pacific, Australian and New Zealand waters. Papers 

from Dr. Th. Mortensen's Pacific Expedition 1914-1916. — Videnskabelige Meddelelser fra 

Dansk Naturhistorisk Forening i Kjobenhavn 116:1-168, figs. 1-81. 
Utinomi, H. 1959. Pycnogonida of Sagami Bay. Publications of the Seto Marine Biological Laboratory 

7(2): 197-222, figs. 1-9. 
. 1962. Pycnogonida of Sagami Bay — Supplement. — Publications of the Seto Marine Biological 

Laboratory 10(1):91-104, figs. 1-6. 
. 1971. Records of Pycnogonida from shallow waters of Japan. — Publications of the Seto 

Marine Biological Laboratory 18(5):3 17-347. 

(KN) 2-25-3, Nakamachi, IVlusashino-shi, Tokyo 180, Japan; (CAC) Depart- 
ment of Invertebrate Zoology (Crustacea), NHB W323, stop 163, Smithsonian 
Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 292-296 

A GYNANDROMORPH OF THE JAPANESE PYCNOGONID 
ANOPLODACTYLUS GESTIENS (ORTMANN) 

C. Allan Child and Koichiro Nakamura 

Abstract. — A single sexual mosaic or gynandromorph specimen was found 
among a very large series of the Japanese pycnogonid Anoplodactylus gestiens 
(Ortmann). The rarity and morphology of the specimen are examined and de- 
scribed, and its relationship with other arthropod and pycnogonid gynandro- 
morphs is discussed. 



Pycnogonids are normally dioecious and sexually dimorphic with testes or ova- 
ries situated in each leg. In the few instances where sexual habits have been 
observed, the male accepts eggs released from the female's sexual pores, fertilizes 
them externally, cements the eggs together by use of femoral cement glands, and 
carries the eggs cemented to his ovigers until after hatching. The sexual habits, 
as far as we have been able to discover in the literature, are unknown for many 
genera, including those few species having parasitic or commensal larvae and 
young not carried by the male. It is therefore significant to discover a single 
apparent gynandromorph specimen among a very large sample of the pycnogonid 
Anoplodactylus gestiens (Ortmann, 1891), collected in Sagami Bay, Japan. 

The occurrence of hermaphrodites and gynandromorphs among the Pycnogon- 
ida is extremely rare according to the literature. There are only three examples 
of these abnormalities on record. The first two references (to one hermaphroditic 
species and a gynandromorphic specimen of another species) are reviewed by 
Child (1978:134), and the third example, a gynandromorph series, is discussed in 
the same paper (Child 1978:135-141, figs. 1-4). Such abnormalities are apparently 
equally rare in other marine arthropods and there is only a very limited literature 
on the subject (Chace and Moore 1959:226-231, figs. 1—4; Froglia and Manning 
1978:700, fig. 5; Manning and Holthuis 1981:62, fig. 14). The majority of papers 
concern bilateral gynandromorphs where half the animal has male characters 
including obvious coloration, and the other half has female characters with female 
coloration. It is to be assumed that where coloration is not a factor and where 
micro- or semimicroorganisms are concerned, a small number of hermaphrodites 
or gynandromorphs would pass unobserved, particularly when examining a large 
series of these organisms. It was therefore rewarding when one of us (Nakamura) 
discovered a single abnormal specimen while determining the sex of a series of 
598 specimens of Anoplodactylus gestiens (Ortmann). 

This pycnogonid is a fairly commonly captured endemic in the shallow waters 
of Japan and is listed in most taxonomic reports concerning Japanese collections. 
The collection in which this single gynandromorph was found was taken in a 
depth of 113 meters during a dredging-trawling survey by the R/V Tansel Mam 
of the Ocean Research Institute of the University of Tokyo, while sampling for 
certain common pycnogonids to be used in seasonal fluctuation studies of both 
numbers and sex frequency over a 28 month period (Nakamura and Sekiguchi, 
in press). 



VOLUME 95, NUMBER 2 



293 




Fig. 1. Anoplodactylus gestiens, gynandromorph appendages: A, Normal female leg with ova; B, 
Leg L 4 of gynandromorph with enlargement of cement gland; C, Normal male leg; D, Proximal 
segments of gynandromorph ovigers, left at top, right at bottom. Abbreviations: ov, ovary with ova; 
id, intestinal diverticulum; eg, cement gland and tube; t, testes; sp, sexual pore. 



Description of the Specimen 

This specimen has characters of both male and female pycnogonids of the genus 
Anoplodactylus. Normally, gut diverticula extend out through most segments of 
each leg of a pycnogonid. The ovaries of the female lie dorsally to these diver- 
ticula and are usually confined to the first four segments of each leg (Fig. lA). In 



294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

the male, the testes lie ventrally to the diverticula and are usually confined to the 
three coxae or the second and third coxae only (Fig. IC). Testes are normally 
found in the posterior four legs or the posterior pair of legs only. Both sexes have 
sexual discharge pores placed ventrally on the second coxae; the female's on all 
eight legs and the male's corresponding to the occurrence of testes in the posterior 
legs. Normally only males of this genus have ovigers, the egg-carrying appendages 
to which the extruded eggs are cemented (except for a few enigmatic females of 
Anoplodactylus jonesi Child, 1974). 

This apparent gynandromorph has ovigers like the male but with the left one 
slightly deformed (Fig. ID), has female ovaries distended with ova in all eight 
legs, has sex pores on all eight legs, and has a femoral cement gland present but 
reduced or atrophied on leg L 4 (Fig. IB). 

Each set of coxae was carefully examined for the presence of testes, but aside 
from the darker opaque tubes of the gut diverticula and ovaries, none of the eight 
legs have any matter resembling a testes in the coxae. Unlike the hermaphrodite 
species (of a different genus) found by Marcus (1952), this specimen is apparently 
incapable of self-fertilization and thus is more properly termed a sexual mosaic 
or gynandromorph rather than an hermaphrodite. 

The right oviger is normal to all appearances, but the left is slightly shorter 
with the two proximal segments reduced in diameter. The second segment tapers 
proximally and is expanded distally, forming a club-like shape. The normal ce- 
ment gland in Anoplodactylus gestiens is an oblong opaque sac placed mid-dor- 
sally in the femur and extending outside the integument in the form of a slanted 
tube as long as one-third the diameter of the segment. The cement gland present 
on leg L 4, the single leg out of eight with the gland in this specimen, is a tiny 
tube expanded at its base, and measures in length less than one-fourth the femur 
diameter. The gland within the segment has the shape of a small dark bifurcated 
tube. The space which would normally be taken up by a full-sized gland is lighter 
colored than the surrounding tissues as though the space were empty. The smaller 
cement gland is placed more toward the distal end of the femur than the normal 
mid-dorsal placement in the male leg. The seven other legs have no suggestion 
of cement glands and are apparently normal in all respects for an ovigerous 
female. All eight legs have normal sexual discharge pores on the second coxae. 

There are other differences between this specimen and our series of Anoplo- 
dactylus gestiens specimens. These differences in the gynandromorph appear to 
fall within the range of non-sexual variation in the species. This is verified, except 
for a difference in specimen size, by the large series in hand. The ocular tubercle 
is shorter in the gynandromorph, but this tubercle varies in length through the 
series. The propodal lamina is slightly longer than usual in this specimen and 
corresponds to Stock's (1954:69) figure 31a of a propodus from a young female 
of the species. Some of the female lamina are shorter than this in our series of 
specimens. 

The single major difference in the gynandromorph specimen is its reduced size. 
It measures only about three-fourths the size of a normal male of the species, 
and in pycnogonids the male is almost always smaller than the female. This could 
be attributed to the mosaic characters of the specimen, but with other sexual 
mosaic pycnogonids (Child 1978), little or no difference in size was found between 
normal and abnormal specimens. 



VOLUME 95, NUMBER 2 295 

Discussion 

This single gynandromorph among 598 specimens of the same Anoplodactylus 
species affords no hint of the cause or origin of this abnormahty in the population 
as sampled. Records were not made at the time of capture concerning associated 
fauna or bottom conditions, but the depth of 113 meters at which the sample was 
taken would preclude the influence of wave action, ship's wake, or other surface 
phenomena as a disturbance in the embryonic determination of sexual characters. 
On the other hand, it is entirely possible that this specimen developed in a dif- 
ferent habitat and depth and only reached the sampling depth as an adult. 

The fact that this isolated abnormality is not unique among the pycnogonids is 
of some interest. Gynandromorphy has been discovered among species of the 
gQiitrsi Anoplodactylus Sind Ascorhynchus. Anoplodactylus gestiens is the second 
species of this genus in which gynandromorphs are known, the other being A. 
port us Caiman, 1927 (Child 1978). The littoral gynandromorphs of A. port us were 
possibly caused by agitation of the eggs during the time of early embryonic de- 
termination and development by fast ships' propellers. This could account for the 
random nature of visible abnormalities rather than a bilateral appearance of the 
resulting gynandromorphs. A random gynandromorph pycnogonid has one leg 
with male characters, the next with female characters, the next with both male 
and female characters, and the next with only female characters, and so on, with 
no apparent pattern. This gynandromorph of A. gestiens has only one random 
character, that of the reduced cement gland on the left fourth leg. All of the legs 
have developing eggs and seven of the eight legs are normal female legs in ap- 
pearance, while the specimen has fully developed male ovigers with the left oviger 
slightly deformed. There is no randomness in these last characters even though 
the presence of both ovigers and ovaries in a single specimen is contrary to the 
definition of this genus. The fact that both the reduced cement gland and the 
malformed oviger occur on the left side of the specimen may be due to coinci- 
dence or it may signify a form of bilateral mosaic, but there is no further evidence 
to support bilateralism. 

The two species with abnormalities in the genus Ascorhynchus are A. abyssi 
Sars, 1877, and A. corderoi du Bois-Reymond Marcus, 1952. Losina-Losinsky 
(1964) described but did not figure an apparent bilateral of A. abyssi having male 
legs on one side and female legs with eggs on the other. Marcus, in describing 
her new species A. corderoi, listed several specimens with both testes and ovaries 
in the same legs, making this the only true hermaphrodite species known. 

There is one other species oi Anoplodactylus , A. jonesi, in which a few spec- 
imens out of all of those known, while not apparently gynandromorphs, have 
female legs lacking cement glands while at the same time having male ovigers. 
Based on our knowledge of the genus, there is as little explanation for this sexual 
combination as there is for the gynandromorph in A. gestiens. Perhaps it will be 
necessary to redefine the genus based on this occasional presence of otherwise 
absent characters for a few of the known species. 

Acknowledgments 

We express our gratitude to Dr. F. A. Chace, Jr., Department of Invertebrate 
Zoology, Smithsonian Institution, for his critical review of the manuscript, to 



296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Professor Koichi Sekiguchi of the University of Tsukuba and chief scientist of 
the Tansei Mam Expedition, for his collaboration in collecting material (with 
KN), to Mr. H. Ueda of the M.R.C. for his excellent technical assistance in 
collecting the specimens (with KN), and to the Department of Invertebrate Zoology 
(Crustacea), Smithsonian Institution, for use of visitor facilities (by KN). 

Literature Cited 

Chace, F. A., Jr., and G. M. Moore. 1959. A bicolored gynandromorph of the lobster, Homarus 

americanus . — Biological Bulletin 11 6(2): 226-231, figs. 1-4. 
Child, C. A. 1978. Gynandromorphs of the ^ycnogomd Anoplodactylus partus. — Zoological Journal 

of the Linnean Society 63:133-144, 4 figures. 
FrogHa, C, and R. B. Manning. 1978. Brachynotus gemmellari (Rizza, 1839), the third Mediterranean 

species of the genus. — Proceedings of the Biological Society of Washington 91(3):691-705, figs. 

1-5. 
Losina-Losinsky, L. 1964. [Pantopoda from the collections of the F. Litke in 1955 and the Ob in 

1956, Scientific results of the oceanographic expeditions to the northern parts of the Greenland 

Sea and neighboring regions of the Arctic Basin in the years 1955-1958]. — Trudy arkticheski 

i antarkicheski Nauchnovo-issiedovatetskovo institut 259:330-339 [In Russian]. 
Manning, R. B., and L. B. Holthuis. 1981. West African Brachyuran Crabs (Crustacea: Decapoda). — 

Smithsonian Contributions to Zoology 306:i-xii, 1-379, figs. 1-88. 
Marcus, E. du B.-R. 1952. A hermaphrodite pantopod. — Anais da Academia Brasileira de Ciencias 

24(l):23-30. 
Nakamura, K., and K. Sekiguchi. (in press). Seasonal occurrence of four species of pycnogonids in 

Nabeta Bay, Shimoda, Japan. 
Ortmann, A. 1891. Bericht iiber die von Herm Dr. Doderlein in Japan gesammelten Pycnogoniden. — 

Zoologische Jahrbiicher (Systematik) 5(1): 157-168, plate 24. 
Stock, J. H. 1954. Pycnogonida from the Indo- West-Pacific, Australian and New Zealand waters. 

Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916. — Videnskabelige Meddelelser 

fra Dansk Naturhistorisk Forening i Kj0benhavn 116:1-168, 81 figures. 

(CAC) Department of Invertebrate Zoology, Smithsonian Institution, Wash- 
ington, D.C. 20560; (KN) 2-25-3 Nakamachi, Musashino-shi, Tokyo 180, Japan. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 297-318 

THE ENTOCYTHERID OSTRACOD FAUNA OF 
NORTHERN GEORGIA 

Horton H. Hobbs, Jr. and Daniel J. Peters 

Abstract. — The entocytherid ostracod fauna of the Tennessee and Coosa river 
basins (exclusive of the Tallapoosa sub-basin of the latter) in Georgia consists of 
13 species that infest one or more of 26 of the 27 epigean crayfishes frequenting 
these watersheds. None of the ostracods is restricted to the area, and there is no 
evidence that any of them require a specific host. Spot maps depicting the locality 
records for each ostracod, summaries of their ranges, and their associations with 
the crayfish hosts and other entocytherids occurring in the area are tabulated. A 
key for recognizing the ostracods is also included. 



Introduction 

During a survey of the crayfish fauna of Georgia (Hobbs 1981), many of the 
entocytherid ostracods infesting these decapods were preserved for future study. 
Those available from the Coosa (exclusive of the Tallapoosa) and Tennessee river 
basins, an area herein designated as "northern Georgia," constitute the material 
on which this is based. 

The entocytherid ostracods are obligate symbionts of aquatic amphipods, iso- 
pods, crayfishes, and crabs. Only members of the nominate subfamily occur in 
Georgia, and all are associated with crayfishes. The entocytherid fauna of the 
area treated here consists of 13 species that infest one or more of 26 of the 27 
epigean species of cambarine crayfishes frequenting the Coosa and Tennessee 
watersheds. 

Not one of the 13 entocytherids is restricted to northern Georgia, but we an- 
ticipate that the southeastern limits of the range of at least two of them (Asce- 
tocy there bouchardi and Dactylocythere prominula) will prove to lie within the 
area. Few of the members of the genus Dactylocythere are expected to be found 
to the south except in the headwaters of the Chattahoochee and Savannah rivers. 

The area under consideration. — The Coosa and Tennessee rivers (Fig. 1) drain 
some 12,000 and 7250 square kilometers, respectively, of the northern part of 
Georgia. Within this area, there are segments of four of the five physiographic 
regions represented in the state (Fig. 2). Furthermore, elevations range from 
approximately 152 to 1450 meters. 

The headwater streams and most of the creeks and rivers outside of the Ridge 
and Valley Province flow, for the most part, over granitic or sandstone beds and 
thus have low mineral content. Those coursing at lower elevations of the Ridge 
and Valley Province flow over eroded beds of Paleozoic limestone, dolomite, and 
shale and are enriched by calcium and magnesium salts. 

A broad array of habitats ranging from seepage areas and mountain rills to 
rather large, sluggish streams and crayfish burrows is available to the ento- 
cytherids in this section of the state, and whereas all appear to have been ex- 
ploited by the ostracods, we have been able to recognize few correlations of their 



298 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. I. River basins in northern Georgia. 



distribution with ecological conditions. Unlike entocytherid faunas in other areas 
in which unique species are symbionts of those crayfishes largely restricted to 
burrows, the ostracods infesting the burrowing crayfishes in northern Georgia are 
members of some of the same species that are found on the stream-dwelling 
crayfishes frequenting nearby epigean habitats. The ostracods occurring in the 
physiographic provinces and drainage basins are summarized in Table 1. 

Entocytherid associates and infestations. — As we stated in our summary of the 
entocytherids of North Carolina (Hobbs and Peters 1977:4), "In . . . [northern 
Georgia], as elsewhere, frequently more than one entocytherid infests the same 
crayfish, and in this study the ostracods were, for the most part, obtained from 
collections of crayfishes in which all of the specimens collected in one locality 
were preserved in the same container. Thus, if two or three species of crayfishes 
were obtained at one station, we were unable to determine which of the ostracods 
retrieved from the container infested which host species. The only instances in 
which we can be certain that an entocytherid was associated with a particular 
host species are those in which collections contain only one species of crayfish. 
In Table 2, the associations indicated are based in part upon records of ostracods 
occurring in the same locality, perhaps infesting more than one species of crayfish 
(open circles) rather than occurring on the same host species (solid circles)." Our 
collections were obtained from 154 localities (Fig. 3). 

Whereas an obligate association with one or more crayfishes exists for all of 
the entocytherids of the area, evidence exists that none requires a specific host. 



VOLUME 95, NUMBER 2 



299 



35- 



34"- 




Fig. 2. Physiographic provinces in northern Georgia. 

The only ostracod known to be associated with a single host species in the region 
is Dactylocythere prominula which was found in a single locality on a small 
mountain stream in which only one crayfish, Cambarus (C) bartonii, was col- 
lected. In several localities in Tennessee, however, it has been found on other 
crayfishes (Hobbs and Walton 1977:609), and we beheve it highly Hkely that it 
will be found on species other than C. (C) bartonii in northern Georgia. 

Entocytherid distribution in northern Georgia. — The numbers of entocytherids 
(Table 1) and crayfishes (Hobbs 1981: table 2) occurring in the four provinces in 
northern Georgia are as follows: 



Provinces 


Entocytherids 


Crayfishes 


Appalachian Plateau 
Ridge and Valley 
Blue Ridge 
Piedmont 


6 
10 

7 
7 


8 

19 
13 

8 



Obviously the Ridge and Valley Province supports the largest number of both 
entocytherids and crayfishes: 10 of the 13 known ostracods and 20 of the 27 
crayfishes. Despite the relatively small area of northern Georgia encompassed by 
the Appalachian Plateau, almost half of the species comprising the entocytherid 
fauna have been found within it. 

Entocythere elliptica and Uncinocythere simondsi are the most widespread 
entocytherids in northern Georgia, both occurring in all of the physiographic 



300 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Distribution of entocy therids in the physiographic provinces and drainage basins of north- 
em Georgia. 





ENTOCYTHERIDS 
















« 






CO 

• lO 

CO 

<^ 












•i^ 




H 




H 


-Cf 


« 









CO 

• fO 
















C3 








CO 




CO 






o 
O 


o 






-0 


C3 

S«i 


-Si 

so 

& 




CO 




• pO 


• 10 

CO 


l-O 


• rO 
SO 

.5- 


CO 

• fO 

se 

•10 

'-O 

•»o 


CO 

•lO 

CO 




K 


03 




so 


so 


so 


so 


So 


so 


C 










^ 


^ 





f^ 


Q 


Q 


Q 


Q 


Q 


•Q 


(^ 


(^ 


b 


PROVINCES 




























Appalachian Plateau 




• 




• 






• 






• 


• 




• 


Ridge and Valley 


• 




• 


• 


• 


• 


• 




• 




• 


• 


• 


Blue Ridge 






• 




• 


• 




• 




• 


• 




• 


Piedmont 


• 




• 




• 


• 








• 


• 




• 


DRAINAGE BASINS 




























Tennessee 


• 


• 




• 


• 




• 






• 


• 


• 


• 


Hiwassee 










• 


• 


• 






• 


• 




• 


Little Tennessee 










• 


• 








• 








Coosa 






• 




• 












• 




• 


Conasauga 






• 




• 


• 




• 


• 




• 




• 


Coosawattee 






• 




• 


• 










• 




• 


Etowah 


• 








• 


• 








• 


• 




• 


Chattooga 






• 




± 












• 




• 



provinces and in all major drainage basins except the Little Tennessee. Only 
slightly less broadly distributed is Dactylocythere falcata, which has not been 
found on the Appalachian Plateau. In contrast, Ascetocythere bouchardi, Dac- 
tylocythere brachystrix, Dt. prominula, Dt. suteri, and Entocythere illinoisensis 
have the most restricted ranges. The first of these has been found in four localities 
in the Tennessee Basin lying on the Appalachian Plateau. Dactylocythere bra- 
chystrix is a little more widespread, for while known only from the Tennessee 
Basin, it occurs in streams of both the Appalachian Plateau and Ridge and Valley 
provinces. Dactylocythere prominula was discovered in only a single locality, 
that lying in the Conasauga Basin of the Blue Ridge Province. Dactylocythere 
suteri was obtained in several localities in the same river basin, but in the Ridge 
and Valley instead of the Blue Ridge Province. Like Dt. prominula, E. illinois- 
ensis was found in a single locality in the area, but one lying in the Tennessee 
watershed of the Ridge and Valley Province. 



VOLUME 95, NUMBER 2 



301 



Table 2. — The entocytherids and their crayfish hosts in northern Georgia (solid circles = known 
host-commensal relationships; open circles = occurrence in same locality; see "Entocytherid asso- 
ciates and infestations" for more detailed explanation). 



HOSTS 


ENTOCYTHERIDS 


o 

so 

si 


^^ 
o 

o 

CO 




o 


so 


o 

s- 

so 


o 

CO 

o 
o 
<^ 

so 


l-O 

Si 

• lO 

o 

so 


■ fO 

k. 
So 

Si 

CO 

so 


CO 

* lO 

CO 

si 

<^ 

O 

CO 

Ri 
Ki 

O 

si 


• lO 

so 

■S< 

l-O 
l-O 


CO 

• rO 

CO 

<^ 

CO 

• lO 

o 

• lO 
l-O 
l-O 

• lO 


• lO 

CO 

-^ 

si 

o 

• fO 

CO 


C. (C.) hartonii 










o 


• 


• 


• 




• 


• 




• 


C. (D.) cymatilis 










o 












o 




o 


C. (D.) latimanus 


O 




o 




• 


• 


o 








• 




• 


C. (D.) striatus 


O 




• 




• 




o 




• 


O 


• 


o 


• 


C. (H.) coosawattae 










o 


o 










o 




o 


C. (H.) fasciatus 


o 




o 




o 


o 








O 


• 




o 


C. (H.) girardianus 










o 




• 






• 





o 


• 


C. (H.) longirostris 


o 








• 




o 








• 




o 


C. (H.) manningi 










• 












o 




o 


C. (H.) speciosus 










• 


o 










• 




• 


C. (J.) conasaugaensis 






o 




o 


• 








o 


o 




• 


C. (J.) distans 




















• 








C. (J.) nodosus 




















o 


o 






C. (J.) parvoculus 




• 




o 












o 






o 


C. (J.) unestami 




• 




• 






o 






• 


• 




o 


C. (L.) acanthura 






o 




o 








o 




o 




o 


C. (P.) coosae 






o 




• 


• 






• 




• 




• 


C. (P.) extraneus 


o 








o 




o 








o 


o 


o 


C. (P.) georgiae 




















• 








C. (P.) hiwasseensis 












o 


o 






o 


• 




o 


C. (P.) parrishi 












• 








• 


• 






C. (P.) scotti 






o 
















• 




o 


O. erichsonianus 


o 




o 








o 






o 


• 


o 


• 


0. forceps 




























0. spinosus 






o 




• 




o 




o 




o 




o 


P. (0.) lophotus 










o 




o 








o 


o 


o 


P. (Pe.) spiculifer 


o 




o 




o 


• 










• 




o 



302 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Localities from which entocytherids have been examined. 



Finding Ankylocythere telmocea in the Tennessee Basin, and especially in the 
Ridge and Valley Province, came as a real surprise to us for elsewhere in its 
range it has been reported only from the Piedmont and Coastal Plain provinces 
(see Hart and Hart 1974:32-33). This isolated locality, and perhaps others in the 
province, should be investigated further to assure the validity of the occurrence 
of the species in the Tennessee River basin. 

In respect to their hosts, Entocythere elliptic a is apparently the least selective 
of the entocytherids occurring in northern Georgia, having been definitely asso- 
ciated with 13, and possibly occurring on as many as 23, of the 27 crayfishes 
inhabiting the area. Only slightly less selective are Dactylocythere falcata and 
Uncinocythere simondsi. The former has been found on 12 species of crayfishes 
frequenting the area, and the possibility exists that 10 additional host species are 
also infested by Dt. falcata. Uncinocythere simondsi has found the exoskeleton 
of all of the crayfishes acceptable for colonization except perhaps those of Cam- 
barus (J.) nodosus, C. (J.) distans, C. (P.) georgiae, C. (P.) parrishi, and O. 
forceps. 

Although most studies have indicated that crayfishes of the genus Orconectes, 
in general, support smaller numbers of ostracods than do most members of other 
cambarine genera, we are not confident that Orconectes forceps lacks ento- 
cytherid infestations. The few collections of this crayfish from Georgia streams 
that are available to us were not maintained in the original containers in which 
they were collected, and the fluid in which they were killed had been discarded. 



VOLUME 95, NUMBER 2 



303 



Table 3. — Associations of entocytherids in northern Georgia (solid circles = known utilization of 
same host species in at least one locality; open circles = occurrence in same locality; see "Ento- 
cytherid associates and infestations" for more detailed explanation). 



ENTOCYTHERIDS 


ENTOCYTHERID 
ASSOCIATES 


o 


o 

CO 




H 

cc 
o 


' — ;> 


SO 


-Si 

Oh 

CO 

O 


o 

Oh 


CO 


CO 
CO 

O 

CO 

O 

si 




CO 
CO 

<^ 

CO 

o 

• lO 


CO 

o 

CO 


An. telmoecea H 






o 


o 


O 








O 




o 


As. houchardi 


■ 


o 












O 






O 


C. cyma 




■ 


o 


o 






o 




O 




O 


Dt. brachystrix 




o 


■ 










O 


• 






Dt. falcata 


O 




O^ 


■• 


O 




• 




• 




• 


Dt. leptophylax 


O 




o 




• ■o 


• 




• 


• 




• 


Dt. mecoscapha 


o 








o 


o 


■ 




• 


o 


O 


• 


Dt. prominula 












• 


■ 






• 






Dt. suteri 






o 




• 








■1 


o 




• 


Dn. donnaldsonensis 




o 




o 




• 


• 






■ 


• 




• 


E. elliptic a 


o 




o 


• 


• 


• 


o 


• 


o 


• 


■ 


O 


• 


E. illinoisensis 














o 








o 


■ o| 


U. simondsi 


o 


l2. 


o 




• 


• 


• 




• 


• 


• 


o 


■ 



Cambarus (J.) distans has been found in only two localities, both on an inter- 
mittent stream in the northern part of Dade County, and in one of them it is 
infested with Donnaldsoncythere donnaldsonensis, one of the most widely rang- 
ing entocytherids in the eastern half of the United States. Cambarus (P.) geor- 
giae, known from a single locality in the Georgia segment of the Little Tennessee 
River, harbors two of the three ostracods known to occur in the headwaters of 
that stream. 

The burrowing Cambarus (D.) cymatilis and C. (J.) nodosus and the riffle- 
inhabiting C. (H.) manningi are infested by three, two, and three entocytherid 
species, respectively, but the ostracods on them are among the commonest in the 
area, infesting a comparatively large variety of crayfish hosts (Table 1). 

In contrast, the least ecologically restricted crayfish in the area, Cambarus (D.) 
striatus, has been found to host five and possibly nine entocytherids. Cambarus 
(C.) bartonii and C. {P.) coosae harbor six and five species of entocytherids, 
respectively, and possibly an additional one occurs on each. 



304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

The host species in the area have been treated by Hobbs (1981), and synony- 
mies, ranges, and notes on their biology are recorded therein. The epigean cray- 
fish fauna of northern Georgia, as delimited here, consists of the following 
species: Cambarus {Cambarus) bartonii (Fabricius, 1798); C. {Depressicamba- 
rus) cymatilis Hobbs, 1970; C {D.) latimanus (LeConte, 1856); C. (D.) striatus 
Hay, 1902; C. {Hiaticambarus) coosawattae Hobbs, 1981; C. (H.) fasciatus 
Hobbs, 1981; C. (//.) girardianus Faxon, 1884; C. (H.) longirostris Faxon, 1885; 
C. (H.) manningi Hobbs, 1981; C. (H.) speciosus Hobbs, 1981; C. {Jugicam- 
barus) conasaugaensis Hobbs and Hobbs, 1962; C. (J.) distans Rhoades, 1944; 
C. (J.) nodosus Bouchard and Hobbs, 1976; C. {J .) parvoculus Hobbs and Shoup, 
1947; C. (/.) unestami Hobbs and Hall, 1969; C. (Lacunicambarus) acanthura 
Hobbs, 1981; C. {Puncticambarus) coosae Hobbs, 1981; C. {P.) extraneus Ha- 
gen, 1870; C. (P.) georgiae Hobbs, 1981; C. {P.) hiwasseensis Hobbs, 1981; C. 
(P.) p arris hi Hobbs, 1981; C. (P.) scotti Hobbs, 1981; Orconectes erichsonianus 
(Faxon, 1898); O. forceps (Faxon, 1884); Orconectes spinosus (Bundy, 1877); 
Procambarus (Ortmannicus) lophotus Hobbs and Walton, 1960; P. (Pennides) 
spiculifer (LeConte, 1856). 

Key to Entocytherid Fauna of Northern Georgia 
(Based on male copulatory complex) 

1. External border of horizontal ramus of clasping apparatus with sim- 
ple excrescence Ankylocythere telmoecea 

1 ' . External border of horizontal ramus of clasping apparatus entire . . 2 

2(1'). Penis with prostatic and spermatic elements distinctly diverging along 
part of their length; ventral part of peniferum deeply cleft posteriorly 

Ascetocythere bouchardi 

2'. Penis with prostatic and spermatic elements contiguous throughout 
their lengths; ventral part of peniferum never deeply cleft posteriorly 
3 

3(2'). Ventral part of peniferum bulbous Cymocythere cyma 

3'. Ventral part of peniferum never bulbous 4 

4(3'). Finger guard absent 5 

4'. Finger guard present Dactylocythere 8 

5(4'). Rami of clasping apparatus disposed at angle of no more than 70 

degrees Donnaldsoncythere donnaldsonensis 

5'. Rami of clasping apparatus disposed at angle of at least 90 degrees 6 

6(5'). Clasping apparatus with internal border of horizontal ramus not ser- 
rate, bearing maximum of 3 teeth Uncinocythere simondsi 

6'. Clasping apparatus with internal border of horizontal ramus serrate, 

bearing 5 teeth Entocythere 1 

7(6'). Clasping apparatus with external border of junction of horizontal and 

vertical rami produced in angle Entocythere illinoisensis 

1 .' Clasping apparatus with external border of junction of horizontal and 

vertical rami broadly rounded Entocythere elliptic a 

8(4'). Clasping apparatus "U" or "C" shaped, distal part expanded 

Dactylocythere leptophylax 



VOLUME 95, NUMBER 2 305 

8'. Clasping apparatus "L" shaped, distal part tapering 9 

9(8'). Accessory groove short, never reaching above dorsal margin of sper- 
matic loop 10 

9' . Accessory groove long, reaching distinctly dorsal to spermatic loop 

11 

10(9). Accessory groove never reaching dorsal margin of spermatic loop; 

vertical ramus of clasping apparatus bowed 

Dactylocythere brachystrix 

10'. Accessory groove reaching dorsal margin of spermatic loop; vertical 

ramus sometimes with shoulder but never distinctly bowed 

Dactylocythere suteri 

1 1(9'). Finger guard jutting anteroventrally Dactylocythere prominula 

11 ' . Finger guard directed ventrally 12 

12( 11 '). External border of junction of horizontal and vertical rami of clasping 
apparatus angular; more than one tooth on internal border of hori- 
zontal ramus Dactylocythere mecoscapha 

12' . External border of junction of horizontal and vertical rami of clasping 
apparatus rounded; never more than one tooth on internal border of 
horizontal ramus Dactylocythere falcata 

Annotated List of Species 

In the following annotated list of ostracods, complete synonymies are furnished 
for only those species {Ascetocythere bouchardi and Dactylocythere prominula) 
that have been described since the publication of a monograph of the family (Hart 
and Hart 1974) in which the remaining species are fully treated. Locality records 
are included only for those species that are known from as few as 10 localities. 
For information concerning the crayfish hosts, Hobbs (1981) should be consulted. 

Ankylocythere telmoecea (Crawford) 
Fig. 4 

Entocythere telmoecea Crawford, 1959:167-173, figs. 24-30. 

Ankylocythere telmoecea. — Hart, 1962:128. — Hart and Hart, 1974:32-33, pi. 4, 

figs. 1-5; pi. 46.— Hobbs, 1981:498-501. 
Uncinocythere lucifuga. — Hart and Hart, 1974:131 [in part; not Entocythere 

lucifuga Walton and Hobbs, 1959]. 

Range. — "From the FHnt River drainage in Georgia to the York Basin in Vir- 
ginia" (Hobbs and Peters 1977:18). 

Previous records in northern Georgia. — Hart and Hart (1974) reported the oc- 
currence of Uncinocythere lucifuga in northern Georgia (see locality 4 below). 
We have examined the specimen on which this record was based and it appears 
to us to be a member of A. telmoecea. 

Distribution in northern Georgia. — Except for a single locality in the Chicka- 
mauga Creek watershed (Tennessee Basin), all of them are situated in the Etowah 
drainage system. Our seven locality records are as follows: Cherokee County: 
(1) stream 2 mi W of Free Home on St Rte 20 (hosts: C. (D.) latimanus, P. (Pe.) 



306 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Distribution of Ankylocythere telmoecea, Cymocythere cyma, and Dactylocythere bra- 
chystrix in northern Georgia (encircled spot and open triangle denote previously reported localities). 



spiculifer; associates: Dt. leptophylax, U. simondsi); (2) stream 1 mi W of Free 
Home and 1.5 mi S of Rte 20 (hosts: C.(D.) latimanus, C. (D.) striatus, P. (Pe.) 
spiculifer; associate: E. elliptica); (3) Scott Mill Creek at St Rte 140, 2 mi SE of 
Canton (hosts: C. (D.) latimanus, P. (Pe.) spiculifer; associates: Dt. falcata, Dt. 
leptophylax, E. elliptica, U. simondsi); (4) stream 3.0 mi N of Canton (Hart and 
Hart 1974:131) (hosts: C. (D.) latimanus, P. (Pe.) spiculifer; no associate cited). 
Lumpkin County: (5) Etowah River at St Rte 52 (hosts: C. (D.) latimanus, C. 
{H.) fasciatus, P. {Pe.) spiculifer; associate: U. simondsi). Paulding County: 
(6) Possum Creek 1.3 mi NE of U.S. Hwy 278 on St Rte 92 (hosts: C. {D. 
latimanus, P. {Pe.) spiculifer; no associates). Walker County: (7) trib to Chicka- 
mauga Creek 0.5 mi N of St Rte 143 on Rte 341 (hosts: C {H.) longirostris, C. 
{P.) extraneus, O. erichsonianus; associate: Dt. mecoscapha). 



Ascetocythere bouchardi Hobbs and Walton 

Fig. 5 

Ascetocythere bouchardi Hobbs and Walton, 1975:7-10, fig. la-d. 

Range. — This ostracod ranges in the Tennessee River basin, on the Cumber- 
land Plateau, from Cumberland County, Tennessee southward to Dade County, 
Georgia. 



VOLUME 95, NUMBER 2 307 

Previous records in northern Georgia. — None. 

Distribution in northern Georgia. — In Georgia, Ascetocythere bouchardi has 
been found in only four localities, all situated on Sand Mountain, Dade County, 
in tributaries of the Tennessee River: (1) trib to Warren Creek on unnumbered 
road 0.3 mi E of Alabama line and 1.0 mi N of St Rte 301 (host: C. (J.) parvoculus; 
no associates); (2) trib to Warren Creek 0.4 mi N of St Rte 301 (hosts: C. (J.) 
parvoculus, C. (J.) unestami; associates: Dt. brachystrix, U. simondsi)', (3) creek 
at Alabama line about 1.5 mi S of St Rte 301 (hosts: C. (/.) parvoculus, C. (J.) 
unestami; associates: Dt. brachystrix, Dn. donnaldsonensis); (4) Higdon Creek 
about 2.0 mi W of Stephensville on St Rte 143 (hosts: C. (J.) parvoculus, C. (J.) 
unestami; associate: Dn. donnaldsonensis) . 

Cymocythere cyma (Hobbs and Walton) 
Fig. 4 

Entocythere cyma Hobbs and Walton, 1960:18-19, figs. 11-16. 
Cymocythere cyma. — Hart, 1962:129. — Hart and Hart, 1974:44, pi. 9, figs. 7-11; 
pi. 47.— Hobbs, 1981:215, 499, 500. 

Range. — From Anderson County, Tennessee, to Jefferson County, Alabama, 
and from Murray County, Georgia, to Hardin County, Tennessee. Encompassing 
the Altamaha, Coosa- Alabama, and Tennessee river basins. 

Previous records in northern Georgia. — Only one locality has been cited for 
this species in the northern part of the state, this in a small tributary to the 
Oostanaula River (Hart and Hart 1974; see below). 

Distribution in northern Georgia. — Cymocythere cyma is widely distributed but 
uncommon throughout most of the Coosa River basin (including the Chattooga) 
of Georgia, although it has not been found in the Etowah watershed. The type- 
locality (Four Mile Creek, 1.8 mi SW of Benton, Polk County, Tennessee) is 
situated in the Hiwassee drainage system some 20 air miles from the upper reach- 
es of the basin in Georgia, but we have no specimens from segments of the river 
system in the state. Localities from which our specimens were collected are the 
following. Floyd County: (1) stream 5.0 mi W of Rome on St Rte 20 (hosts: C 
(/).) striatus, C. (L.) acanthura, C. (P.) coosae, O. spinosus, P. (Pe.) spiculifer; 
associates: Dt.falcata, U. simondsi). Gilmer County: (2) Hell's Creek between 
St Rtes 5 and 156, 2.5 mi NW of Pickens Co line (hosts: C. (D.) latimanus, C. 
(H.) fasciatus, C. (J.) conasaugaensis, C. (L.) acanthura; associates: Dt. lep- 
tophylax, U. simondsi). Gordon County: 1.8 mi N of Calhoun on U.S. Hwy 
41 (Hart and Hart 1974) (hosts: C. (D.) striatus, C. (L.) acanthura, P. (Pe.) spic- 
ulifer; no associate cited). Murray County: (4) trib of Conasauga River 0.8 mi 
N of St Rte 2 on U.S. Hwy 411 (hosts: C. (D.) striatus, C. (L.) acanthura, C. 
(P.) coosae; associates: Dt. falcata, Dt. suteri, U. simondsi); (5) drainage ditch 
2.1 mi E of Conasauga River on St Rte 2 (hosts: C. (D.) striatus, C. (L.) acan- 
thura; associates: Dt. suteri, U. simondsi). Polk County: (6) Little Cedar Creek 
on Esom Hill Rd, upstream from Hematite Branch (hosts: C. (D.) latimanus, C. 
(D.) striatus, C. (P.) coosae; associate: E. elliptica). Walker County: (7) 6 mi 
S of Lafayette at jet of U.S. Hwy 27 and St Rte 143 (hosts: C. (D.) striatus, C. 
(P.) scotti, O. erichsonianus; associate: Dt. falcata). 



308 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 5. Distribution of Ascetocythere bouchardi, Dactylocythere falcata, and Dactylocythere 
prominula in northern Georgia (encircled spot denotes previously reported locality). 



Dactylocythere brachystrix Hobbs and Walton 

Fig. 4 

Dactylocythere brachystrix Hobbs and Walton, 1966:2-3, fig. la-d. — Hart and 
Hart, 1974:50-51, pi. 11, figs. 7-11; pi. 48. 

Range. — This ostracod is known only from the Tennessee River basin, in which 
it ranges on the Cumberland Plateau and Highland Rim from Cumberland County 
southwestward to Lawrence County, Tennessee, and on the Plateau and in the 
Ridge and Valley Province in Dade and Walker counties, Georgia. 

Previous records in northern Georgia. — None. 

Distribution in northern Georgia. — Dactylocythere brachystrix has been found 
in five localities in the Tennessee Basin on Sand and Lookout mountains. Dade 
County: (1) trib to Warren Creek, 0.4 mi N of St Rte 301 (hosts: C. (J.) par- 
voculus, C. (P.) unestami; associates: As. bouchardi, U. simondsi). (2) creek at 
Alabama line about 1.5 mi S of St Rte 301 (hosts: C. (J.) parvoculus, C. (P.) 
unestami; associates: As. bouchardi, Dn. donnaldsonensis) . (3) trib to Lookout 
Creek on St Rte 143, 2.5 mi W of Walker Co line (host: C. (J.) unestami; no 
associates); (4) Bear Creek at St Rte 157 (host: C. (J.) unestami; no associates). 
Walker County: (5) Rock Creek at old St Rte 193 on Lookout Mt (host: C 
(J.) unestami; associate: E. elliptica). 



VOLUME 95, NUMBER 2 309 

Dactylocythere falcata (Hobbs and Walton) 
Fig. 5 

Entocythere falcata Hobbs and Walton, 1961:379-381, figs. 2, 3, 7, 8. 
Dactylocythere falcata.— RdiVU 1962:130.— Hart and Hart, 1974:58-59, pi. 15, 
figs. 1-5; pi. 49.— Hobbs, 1981:231, 238, 257, 295, 499. 

Range. — "From the Coosa River drainage in Georgia and northeastern Ala- 
bama to the Tennessee Basin in Virginia, and east of the Appalachian Divide in 
the upper James and Roanoke drainage systems in Virginia. The records from 
Missouri and Canada cited by Hart and Hart (1974:48, 49) should be confirmed" 
(Hobbs and Peters 1977:30). 

Previous records in northern Georgia. — Hart and Hart (1974) reported the oc- 
currence of this crayfish in six localities in the Coosa Basin of Georgia: three in 
Chattooga (the Floyd County record, 23 mi NW of Rome, is actually in Chattooga 
County), and one each in Murray and Whitfield counties (see open circles in Fig. 
5) and an unknown one in Walker County. 

Distribution in northern Georgia. — This ostracod, one of the commonest in the 
area, has been found in 54 localities where it occurs in all of the physiographic 
provinces except the Appalachian Plateau and in all of the drainage basins. Hosts 
include all of the crayfishes reported from the area except C. (J.) distans, C. (J.) 
parvoculus, C. (J.) nodosus, C. (J.) unestami, and O. forceps. It has been found 
in association with the following entocytherids: C. cyma, Dt. leptophylax, Dt. 
mecoscapha, Dt. s uteri, E. elliptic a, and U. simondsi. 

Dactylocythere leptophylax (Crawford) 
Fig. 6 

Entocythere leptophylax Crawford, 1961:238-242, figs. 9-14. 
Dactylocythere leptophylax.— Hart, 1962:130.— Hart and Hart, 1974:61-62, pi. 
16, figs. 1-5; pi. 48.— Hobbs, 1981:70, 257, 450, 598, 500. 

Range. — From the Savannah River basin in Oconee County, South Carolina, 
westward through the upper Chattahoochee, Ocmulgee, and Coosa basins in 
Georgia, and in the French Broad, Little Tennessee, and Hiwassee drainage 
systems in Tennessee. A single specimen from the Duck River watershed in 
Lewis County, Tennessee (Little Swan Creek in Meriweather Lewis Monument 
Park), suggests the occurrence of an outlying population there. 

Previous records in northern Georgia. — The only report of the occurrence of 
this ostracod in the area is that of Hart and Hart (1974:60) who recorded it from 
Neel's Gap, Union County, on Cambarus (C.) bartonii. 

Distribution in northern Georgia. — Dactylocythere leptophylax has been found 
in 40 localities in the Ridge and Valley, Blue Ridge, and Piedmont provinces of 
northern Georgia where it occurs in the Conasauga, Coosawattee, Hiwassee, 
Little Tennessee, and upper Etowah basins. Its hosts in the area are C. (C.) 
bartonii, C. (D.) latimanus, C. (H.) coosawattae, C. (H.) fasciatus, C. (H.) 
speciosus, C. (J.) conasaugaensis, C. (P.) coosae, C. (P.) hiwasseensis, C. (P.) 
parrishi, and P. (Pe.) spiculifer. The entocytherid associates are An. telmoecea, 



310 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Distribution of Dactylocythere leptophylax and Dactylocythere mecoscapha in northern 
Georgia (encircled spot and open triangle denote previously reported localities). 



C. cyma, Dt. falcata, Dt. mecoscapha, Dt. prominula, Dn. donnaldsonensis, E. 
elliptica, and U. simondsi. 

Dactylocythere mecoscapha (Hobbs and Walton) 

Fig. 6 

Entocythere mecoscapha Hobbs and Walton, 1960:19-23, figs. 17-20. 
Dactylocythere mecoscapha Hart, 1962:130. — Hart and Hart, 1974:62-63, pi. 17, 
figs. 1-5; pi. 48.— Hobbs, 1981:380, 499, 500. 

Range. — This ostracod occurs in the tributaries of the westwardly flowing seg- 
ment of the Tennessee River from Polk County, Tennessee, and Dade, Walker, 
Catoosa, and Union counties, Georgia, to Colbert and Lauderdale counties, Al- 
abama. The record from Hampton County, South Carolina, cited by Hart and 
Hart (1974:63), is almost certainly erroneous, for this low-lying area of the Coastal 
Plain Province furnishes unlikely habitats for any member of the genus Dactylo- 
cythere except Dt. suteri. 

Previous records from northern Georgia. — This ostracod was reported by Hart 
and Hart (1974:63) from the Chickamauga National Military Park near Rossville 
(Chickamauga Creek basin). Its hosts were P. (O.) lophotus, O. erichsonianus, 
and C. (D.) striatus. 

Distribution in northern Georgia. — In Georgia, Dactylocythere mecoscapha 



VOLUME 95, NUMBER 2 311 

has been found in 17 localities, all except one of which occurs in the Appalachian 
Plateau and Ridge and Valley provinces. There are two records in the Blue Ridge 
Province (Wolf Creek and Nottely River) in Union County, so close together as 
to be recorded herein by a single spot on Map 6. All 17 localities are in tributaries 
of the Tennessee River. In the area under investigation, the hosts of this ostracod 
are: C. (C.) bartonii, C. (D.) latimanus, C. (D.) striatus, C. (H.) girardianus , C. 
(H.) longirostris, C. (J.) unestami, C. (P.) extraneus, C. (P.) hiwasseensis, O. 
erichsonianus , O. spinosus, and P. (O.) lophotus. Its entocytherid associates are 
An. telmoecea, Dt. falcata, Dt. leptophylax, Dn. donnaldsonensis , E. elliptica, 
E. illinoisensis, and U. simondsi. 

Dactylocythere prominula Hobbs and Walton 

Fig. 5 

Dactylocythere prominula Hobbs and Walton, 1977:606-609, fig. 3. 

Range. — Except for the single new locality cited herein, this ostracod is known 
to occur only in the Tennessee River basin from Hawkins County, Tennessee, 
southward to Walden Gorge and a single outlying locality in the Elk River wa- 
tershed in Lincoln County, Tennessee. In this survey, it was found in a tributary 
to the Conasauga River in extreme northeastern Murray County, Georgia. 

Previous records in northern Georgia. — None. 

Distribution in northern Georgia. — Dactylocythere prominula was found in- 
festing C. (C.) bartonii in a small unnamed tributary to the Conasauga River 
(Blue Ridge Province) about 4.0 mi E of U.S. Hwy 411 on St Rte 2. At this 
locality it shared its host with Dt. leptophylax and E. elliptica. 

Dactylocythere suteri (Crawford) 
Fig. 7 

Entocythere suteri Crawford, 1959:162-167, pi. 3. 

Dactylocythere suteri.— RdiVi, 1962:131.— Hart and Hart, 1974:72-73, pi. 21, figs. 

11-15; pi. 49.— Peters, 1975:28, figs. 4a, 9.— Hobbs and Peters, 1977:41-43, fig. 

20, map 10.— Hobbs, 1981:231, 295, 498-500. 

Range. — From the Susquehanna Basin in York County, Pennsylvania, south- 
ward on the Atlantic slope to the Oconee River drainage in Morgan County, 
Georgia. In the Gulf watershed, it occurs in the Conasauga Basin in Murray 
County, Georgia, and in the Tennessee Basin in McMinn County, Tennessee. 

Previous records in northern Georgia. — Hart and Hart (1974:73) reported the 
presence of this ostracod in Holly Creek, Murray County (see below). 

Distribution in northern Georgia. — In this area Dactylocythere suteri appears 
to be restricted to the Ridge and Valley Province where it has been found in the 
following localities in the Conasauga watershed. Murray County: (1) Holly 
Creek 1 mi E of Chatsworth on U.S. Hwy 76 (Hart and Hart 1974:73) (hosts: 
Cambarus (D.) striatus, C. (P.) coosae, and O. spinosus; associates: Dt. falcata 
and U. simondsi); (2) trib to Conasauga River 0.8 mi N of St Rte 2 on U.S. Hwy 
411 (hosts: C. (D.) striatus, C. (L.) acanthura, and C. (P.) coosae; associates: 
C. cyma, Dt. falcata, and U. simondsi); (3) trib to Conasauga River on St Rte 
225 between jet of St Rtes 2E and 2W (hosts: C. (D.) striatus and C. (P.) coosae; 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. Distribution of Dactylocythere suteri, Donnaldsoncythere donnaldsonensis , and Ento- 
cythere illinoisensis in northern Georgia (encircled spot and open triangle denote previously reported 
localities). 



associate: U. simondsi); (4) drainage ditch 2.1 mi E of Conasauga River on St 
Rte 2 (hosts: C. (D.) striatus and C. (L.) acanthura; associates: C. cyma and U. 
simondsi); (5) stream 0.2 mi W of St Rte 225 on Rte 286 (hosts: C (H.) coosae 
and O. spinosus; associates: Dt. falcata and £. elliptica). 



Donnaldsoncythere donnaldsonensis (KUe) 
Fig. 7 

Entocythere donnaldsonensis Klie, 1931:334-341, figs. 1-9. 

Donnaldsoncythere donnaldsonensis. — Hart, 1962:131-132. — Hart and Hart, 
1974:79-80, pi. 23, figs. 11-14; pi. 50.— Hobbs and Peters, 1977:43-45, fig. 21, 
map 4 [including a complete synonymy]. — Hobbs, 1981:70, 156, 231, 450, 498- 
500. 

Entocythere hiwasseensis. — Hobbs, 1981:70. 

Donnaldsoncythere hiwasseensis. — Hobbs, 1981:70, 156, 231, 450. 

Range. — "From northern Georgia to Indiana and Maine" (Hobbs and Peters 
1977:44). 

Previous records in northern Georgia. — Hart and Hart (1974:79) reported it as 
Donnaldsoncythere hiwasseensis from Shoal Creek in the Etowah Basin in Daw- 
son County where it infested "C. {Puncticambarus) sp." (probably C. (H.)fas- 
ciatus). 



VOLUME 95, NUMBER 2 



313 




Fig. 8. Distribution of Entocythere elliptica in northern Georgia (encircled spots denote previously 
reported localities). 



Distribution in northern Georgia. — Donnaldsoncy there donnaldsonensis has 
been found in 22 localities in the Appalachian Plateau, Blue Ridge, and Piedmont 
provinces in tributaries of the Tennessee, Hiwassee, and Little Tennessee rivers, 
and in headwaters of the Etowah River. Possible hosts in the area are C. (C.) 
bartonii, C. (D.) striatus, C. (H.) fasciatus, C. (//.) girardianus, C. (J.) cona- 
saugaensis, C. (J.) distans, C. (J.) nodosus, C. (J.) parvoculus, C. (J.) unestami, 
C. (P.) georgiae, C. {P.) hiwasseensis, C. (P.) parrishi, and O. erichsonianus . 
The entocytherid associates are As. bouchardi, Dt. brachystrix, Dt. leptophylax, 
Dt. mecoscapha, E. elliptica, and U. simondsi. 



Entocythere elliptica Hoff 
Fig. 8 

Entocythere elliptica Hoff, 1944:345-349, figs. 15-21.— Hart and Hart, 1974:87- 
88, pi. 26, figs. 1, 2; pi. 51.— Hobbs, 1981:71, 231, 295, 312, 399, 403, 431, 450, 
498-501. 

Range. — Hart and Hart (1974:227) depicted the range of this ostracod as ex- 
tending from extreme southeastern Texas to South Carolina and the northern part 
of the peninsula of Florida, and northward through most of eastern Mississippi, 
Alabama, and Georgia. 

Previous records in northern Georgia. — Hart and Hart (1974:88) reported this 



314 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

species from two localities in Murray and one each in Fannin and Whitfield 
counties. 

Distribution in northern Georgia. — As noted in Fig. 8, this ostracod has been 
found in 54 localities scattered through all of the physiographic provinces of the 
region as well as in all of the watersheds except the Little Tennessee River. Its 
23 hosts and 11 entocytherid associates in the area are summarized in Tables 2 
and 3. 

Remarks. — In treating the entocytherid fauna of North Carolina, Hobbs and 
Peters (1977) recognized five species of the genus Entocythere in the state. They 
were convinced that Entocythere harrisi Peters (1975:32), E. internotalus Craw- 
ford (1959:152), and E. reddelli Hobbs and Walton (1968:243) were all repre- 
sented, and although some difficulty was encountered in separating them, they 
believed that combinations of characteristics of the male and female genitalia and 
those of the pectinate prominence at the base of the apical claws of the antenna 
of the female provided a means of recognizing the three. 

In studying the Georgia representatives of the genus, we have found those 
combinations of characters to be unreliable. Whereas specimens assignable to 
each are present in the northern Georgia fauna, there are also individuals that 
appear to be typical of Hoff s Entocythere elliptica. Both of us were surprised 
when in one specimen of the genus we found one of the paired clasping apparatus 
of the male to be typical of that of E. internotalus and the other characteristic of 
E. elliptica. A similar asymmetry of the corresponding elements resembling those 
of E. internotalus and E. reddelli was encountered later in several specimens. 

Employing the key we had prepared to the North Carolina fauna, we found 
that among the specimens from one of the localities in Georgia all of the males 
were members of E. internotalus but the females appeared to belong to E. red- 
delli. Repeated examination of the material available to us from northern Georgia 
has almost convinced us that insofar as the development and position of the 
"internal talon" on the clasping apparatus is concerned, much of the difference 
noted results from the position of the apparatus when viewed, and whereas the 
position and development of the teeth (often appearing to be absent) of the talon 
may be variable, the direction from which they are viewed minimizes or enhances 
the marginal structures of the talon. 

The female genitalia are indeed highly variable regardless of the angle at which 
they are examined, even more so than those illustrated by Hobbs and Peters 
(1977: figs. 25f, 26e, and 27f). The presence of a vermiform tube in E. harrisi, a 
simple sinuous sclerotized thickening in E. reddelli, and the compound sclerotized 
element in E. internotalus does not characterize the triunguis females collected 
in northern Georgia. In at least three of them there are no discernible genitalic 
structures. Among other specimens the genitalia are intermediate in their com- 
plexity between those of E. internotalus and E. reddelli. Finally, a vermiform 
tube was observed in at least one of the females from the Coosa Basin to be 
incorporated in a genital complex that is otherwise typical of E. internotalus. 

The pectinate prominence at the base of the apical claws of the antenna of the 
triunguis female from northern Georgia surprisingly varies from two to at least 
10 pectins. Generally the fewer the number the more prominent the pectin. In 
seven specimens from a single locality in Cherokee County the full range of 
variability was represented, and even in localities in which the number and po- 



VOLUME 95, NUMBER 2 



315 



Table 4. — Variation in size of Entocythere elliptica in the Coosa Basin of northern Georgia. 





isin 


Sex 


Number 


Length in 


mm 


Height in 


mm 


Sub-b5 


Range 


Average 


Range 


Average 


Conasauga 




S 


22 


0.434-0.490 


0.473 


0.189-0.210 


0.197 






9 


32 


0.560-0.651 


0.590 


0.273-0.343 


0.280 


Coosa 




$ 


17 


0.413-0.511 


0.447 


0.175-0.231 


0.199 






? 


16 


0.546-0.630 


0.578 


0.266-0.294 


0.281 


Coosawattee 




S 


14 


0.420-0.483 


0.450 


0.182-0.224 


0.199 






9 


18 


0.532-0.616 


0.571 


0.245-0.294 


0.266 


Etowah 




6 


23 


0.406-0.476 


0.435 


0.175-0.210 


0.193 






9 


24 


0.504-0.630 


0.548 


0.245-0.315 


0.278 


Entire upper 


Coosa Basin 


6 


76 


0.406-0.511 


0.450 


0.175-0.231 


0.197 






9 


90 


0.504-0.651 


0.577 


0.245-0.343 


0.276 



sition of the pectins exhibit little variability, always one or two individuals 
were, in this respect, atypical of the remainder of the population. 

In view of the range of the complex, from Texas to Virginia (Hobbs and Peters 
1977:52, 55), and the broad areas within it from which no specimens are available, 
we are loath to synonymize any of the three with Hoff s E. elliptica. Because we 
are uncertain as to the identity of the highly variable populations in northern 
Georgia, however, we are tentatively assigning our specimens from the area to 
the first named of the four, E. elliptica. 

Measurements were made of the length and height of the males and females 
throughout the study area (Table 4). The average length of specimens from the 
Conasauga is greater than that of individuals from the other three sub-basins in 
the Coosa watershed and is least in those from the Etowah. In the latter, speci- 
mens from the upper part of the drainage system in Dawson and Lumpkin coun- 
ties have shorter average lengths (c?, 0.429 and 0.417; 9, 0.504 and 0.526 mm, 
respectively) than in the middle {6, 0.440; 9, 0.573 mm) and lower parts {S , 
0.462; 9, 0.602 mm) of the sub-basin. 



Entocythere illinoisensis Hoff 
Fig. 7 

Entocythere illinoisensis Hoff, 1942:67-69, figs. 1-8.— Hart and Hart, 1974:88- 
90, pi. 26, figs. 3-4; pi. 51. 

Range. — From Michigan southward to northwestern Georgia, northern Ala- 
bama and Mississippi, and west of the Mississippi River in Arkansas. 

Previous records in northern Georgia. — None. 

Distribution in northern Georgia. — In this area, Entocythere illinoisensis is 
known from a single locality, a tributary to Chickamauga Creek 9.8 mi E of the 
Dade County line on St Rte 143, Walker County. This stream lies in the Ridge 
and Valley Province, and the crayfish hosts were Cambarus (D.) striatus, C. (H.) 
girardianus, C. (P.) extraneus, Orconectes erichsonianus, and P. (O.) lophotus. 
It was associated with the following entocytherids: Dactylocythere mecoscapha, 
Entocythere elliptica, and Uncinocythere simondsi. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 9. Distribution of Uncinocythere simondsi in northern Georgia (encircled spots denote pre- 
viously reported localities). 



Uncinocythere simondsi (Hobbs and Walton) 

Fig. 9 

Entocythere simondsi Hobbs and Walton, 1960:17, 18, figs. 1-10. — Hobbs, 

1981:70,71. 
Uncinocythere simondsi. — Hart, 1962:138. — Hart and Hart, 1974:133-135, pi. 37, 

figs. 4-7, pi. 55.— Hobbs and Peters, 1977:63-64, fig. 33, map 5.— Hobbs, 

1981:70, 71, 97, 215, 231, 295, 380, 450, 598-501. 

Range. — Hart and Hart (1974:231) reported this crayfish to occur from Illinois 
and Kentucky southward to the panhandle of Florida and from Mississippi east- 
ward to Georgia and South Carolina. Hobbs and Peters (1977:70) added 25 lo- 
calities in the western half of the state of North Carolina. 

Previous records in northern Georgia. — This ostracod was reported by Hart 
and Hart (1974:134) from seven localities in the area under consideration (Fig. 9, 
open circles). 

Distribution in northern Georgia. — Uncinocythere simondsi has been collected 
in 68 localities in all of the physiographic provinces in the area. Except in the 
Nottely and the Hiwassee rivers in Union and Towns counties and in the Little 
Tennessee River in Rabun County, it is widespread in northern Georgia where 
it has been found on all of the crayfishes in the area except Cambarus (J.) distans, 
C. (J.) nodosus, C. (P.) georgiae, C. (P.) parrishi, and Orconectes forceps and 



VOLUME 95, NUMBER 2 317 

was associated with all of the entocytherids occurring in northern Georgia except 
Dactylocythere brachystrix and Dt. prominula. 

Acknowledgments 

Many persons aided in collecting the specimens on which this report is based. 
We are grateful to all of them, and especially to T. A. English, Jr., of Eastern 
Air Lines, Atlanta, Georgia, E. T. Hall, Jr., of the Georgia Department of Natural 
Resources, and J. E. Pugh, of Christopher Newport College, Virginia, who de- 
voted considerable time and effort to assisting us in acquiring the crayfish hosts 
from which the ostracods were taken. We extend our thanks to Margaret A. 
Daniel for rendering the maps and tables, and to her, T. E. Bowman, and C. W. 
Hart, Jr., all of the Smithsonian Institution, for their criticisms of the manuscript. 

Literature Cited 

Crawford, E. A., Jr. 1959. Five new ostracods of the genus Entocythere (Ostracoda, Cytheridae) 

from South Carolina. — University of South Carolina Publications, Biology, series III 2(4): 149- 

189, 5 plates. 
. 1961. Three new species of the genus Entocythere (Ostracoda, Cytheridae) from North and 

South Carolina. — American Midland Naturalist 65(1): 236-245, 21 figures. 
Hart, C. W., Jr. 1962. A revision of the ostracods of the family Entocytheridae. — Proceedings of the 

Academy of Natural Sciences of Philadelphia 114(3): 121-147, 18 figures. 
Hart, Dabney G., and C. W. Hart, Jr. 1974. The ostracod family Entocytheridae. — Academy of 

Natural Sciences of Philadelphia Monograph 18:ix + 239 pages, 49 figures, 52 plates. 
Hobbs, Horton H., Jr. 1981. The crayfishes of Georgia. — Smithsonian Contributions to Zoology 

318:viii + 549 pages, 262 figures. 
, and Daniel J. Peters. 1977. The entocytherid ostracods of North Carolina. — Smithsonian 

Contributions to Zoology 247:4 + 73 pages, 33 figures. 
, and Margaret Walton. 1960. Three new ostracods of the genus Entocythere from the 

Hiwassee drainage system in Georgia and Tennessee. — Journal of the Tennessee Academy of 

Science 35(1): 17-23, 20 figures. 
, and . 1961 . Additional new ostracods from the Hiwassee drainage system in Georgia, 

North Carolina, and Tennessee. — Transactions of the American Microscopical Society 80(4): 

379-384, 8 figures. 
, and . 1966. A new genus and six new species of entocytherid ostracods (Ostracoda, 

Entocytheridae). — Proceedings of the United States National Museum 1 19(3542): 1-12,2 figures. 
, and . 1968. New entocytherid ostracods from the southern United States. — Proceed- 
ings of the Academy of Natural Sciences of Philadelphia 120(6): 237-252, 3 figures. 
, and . 1975. New entocytherid ostracods from Tennessee with a key to the species of 

the genus Ascetocythere. — Proceedings of the Biological Society of Washington 88(2):5-20, 

2 figures. 

, and . 1977. New entocytherid ostracods of the genus Dactylocythere. — 'Proceedings 

of the Biological Society of Washington 90(3):600-614, 4 figures. 
Hoff, C. Clayton. 1942. The subfamily Entocytherinae, a new subfamily of fresh-water cytherid 

ostracods, with descriptions of two new species of the genus Entocythere. — American Midland 

Naturalist 27(l):63-67, 13 figures. 
. 1944. New American species of the ostracod genus Entocythere . — American Midland Nat- 
uralist 32(2):327-357, 33 figures. 
Klie, W. 1931. Campagne speologique de C. BoHvar et R. Jeannel dans I'Amerique du Nord (1928), 

3: Crustaces Ostracodes. — Biospeologica: Archives de Zoologie Experimentale et Generale 

71(3):333-344, 20 figures. 
Peters, Daniel J. 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:iii + 50 pages, 15 figures. 



318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Walton, Margaret, and Horton H. Hobbs, Jr. 1959. Two new eyeless ostracods of the genus Ento- 
cythere from Florida. — Quarterly Journal of Florida Academy of Sciences 22(2): 114-120, 20 
figures. 



(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Wash- 
ington, D.C. 20560; (DJP) York High School, P.O. Box 547, Yorktown, Virginia 
23690. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 319-324 

TWO NEW SPECIES OF LEPTOMYSINID MYSIDS 

(CRUSTACEA, MYSIDACEA) FROM 

SOUTHERN CALIFORNIA 

Linda G. Gleye 

Abstract. — Two new species of leptomysinid mysids, Cubanomysis myste- 
riosa, the second species of the genus described by Bacescu (1968), and Mysi- 
dopsis cathengelae are described from the San Onofre region of the Newport- La 
Jolla Shelf, California. 



Only one species of the genus Cubanomysis is known, C. jimenesi Bacescu, 
1968 from Cuba. The species described below has been collected in shallow water, 
out to a depth of 8 meters, off the coast of Southern California. 

Cubanomysis mysteriosa, new species 
Figs. 1-2 

Material examined. — Southern California, San Onofre State Beach, 8 meters 
depth, 15 January 1977: c^ holotype (USNM 184074), 2^,2$ paratypes (USNM 
184075). 

Etymology. — From the Greek ''mysterion' (mystery), referring to the unpre- 
dictable occurrence of this species. 

Description. — Small, delicate mysid, 3 mm length. Carapace rounded, slightly 
produced into rounded rostrum between eyes. Eyes large, globose. Antennal 
scale 7 times as long as broad, sparsely setose all around, with large apical seg- 
ment, apex bluntly pointed, extending beyond peduncle of antennule by Va {S) 
to Vs (?) its length. Distal joint of antennal peduncle slightly shorter than pre- 
ceding joint, whole peduncle extending ^/lo (c^) to Vio (9) length of antennal 
scale. 

Mandible with large masticatory surface, with small teeth on cutting edge. 
Distal segment of palp with 14 plumose setae and 1 terminal simple seta. Maxilla 
with small exopod bearing 5 setae. 

Endopod of first thoracic limb strong, distal article with dactyl claw and 6 
simple setae. Last article of endopod of second thoracic limb with 3 strong setae, 
plumose and recurved on the interior face and 7 simple and one plumose setae 
on the distal curvature. Endopods of third to eighth thoracic limbs delicate, car- 
popropodus divided into 2 subjoints; 8 large articles on exopods. 

Sixth abdominal somite twice as long as preceding somite. 

First pleopod of adult male without exopod. Pleopods II-V biramous. Pleopod 
IV with 2 branches of 6 articles each, exopod longer than endopod, terminating 
in a long curved seta bearing 2 rows of fine spines on distal third. Seta extends 
past distal tip of telson when pleopod IV is parallel to abdomen. 



320 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 









Fig. 1. Cubanomysis mysteriosa: a, Whole body, lateral, 9 ; b, Rostrum, eyes antennular pedun- 
cle, antennal scale, dorsal, c? ; c, Antennal scale and antennal peduncle, ventral c? ; d. Right mandibular 
palp, S; e. Maxilla, 6; f. Thoracic endopod 1, distal segments, S • 



Telson short, tip extending to distal end of statocyst, constricted distally. Lat- 
eral margins bare. Apex emarginate bearing 16-24 closely set spines. 

Endopods of uropods shorter than exopods; inner lower margin with 10 short 
spines. Statocysts large with hyaline area appearing swollen. 

Remarks. — The present new species is closely allied to C. jimenesi (Bacescu 
1968, Brattegard 1969) but differs from the latter in the following features: 1) 
more robust form of the endopods of the first and second thoracic appendages, 
2) setation of the last article of the endopod of the second thoracic limb, 3) 
spination rather than serration of the terminal seta of the exopod of the fourth 
pleopod of the adult male and 4) spination of the telson. 

There are four known species of the genus Mysidopsis in Southern California, 
M. calif ornica Tattersall, 1951, M. onofrensis and M. brattegartii Bacescu and 
Gleye, 1969, and M. intii Holmquist (personal observation). The species de- 
scribed below has occurred regularly out to a depth of 12 meters in our nearshore 
epibenthic samples since the beginning of our studies in 1976. 



VOLUME 95, NUMBER 2 



321 








Fig. 2. Cubanomysis mysteriosa: a, Thoracic endopod 2, distal segments, S', b, Pleopod 1, d; c, 
Pleopod 4, (J; d, Same, terminal seta exopod, distal tip; e, Telson, S; f, Uropod, S . 



Mysidopsis cathengelae, new species 
Figs. 3^ 

Material examined. — Southern California, San Onofre State Beach, 10 October 
1979: 6 holotype (USNM 184076), 2 d, 5 $, 1 Imm c^ paratype (USNM 18077), 
1 d, 2 9,2 Imm 6 paratype (USNM 184078). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 







Fig. 3. Mysidopsis cathengelae : a, Whole body, lateral, 9 ; b. Rostrum, eyes, antennular peduncle, 
antennal scale, dorsal, S; c, Antennular peduncle, antennal scale, antennal peduncle, ventral; S; d. 
Thoracic endopod 1, $ ; e. Thoracic endopod 2, 9 ; f , Thoracic limb 3,9. 



Etymology. — Named for the California zoologist, Dr. Catherine Engel. 

Description. — Medium sized, robust mysid, 10 mm. Carapace produced be- 
tween eyes into pointed, triangular rostral plate extending nearly to end of eye- 
stalks. Eyes of moderate size, cornea occupying half of whole eye in dorsal view. 
Antennal scale 9 times as long as broad, narrowly lanceolate, setose all around, 
without a distal joint, apex bluntly rounded, extending beyond peduncle of an- 
tennule by ^/s {S) to Vi (?) of its length. Distal joint of antennal peduncle % 
length of preceding joint, whole peduncle extending less than Vi length of antennal 
scale. 



VOLUME 95, NUMBER 2 



323 





a / V b 

Fig. 4. Mysidopsis cathengelae: a, Pleopod 4, d; b, Telson, 9; c, Uropod, ? 



Maxillulae with inner lobe small and armed with 2 setae. Maxilla characteristic 
of genus, exopod well developed, long and slender, setae present on outer margin, 
distal setae much longer than proximal setae. 

Endopod of first thoracic limb short and robust, of form characteristic of genus, 
dactylus with claw, outer margin armed with 7 heavy and 3 light simple setae. 
Endopod of second thoracic Hmb slightly longer than and less robust than en- 
dopod of first limb. Endopod of third thoracic limb longer and more slender than 
the second thoracic limb, sixth joint divided into 3 subjoints, second subjoint 
shortest, third subjoint longest. 

Sixth abdominal somite about 1% times as long as fifth. 

Pleopods in male typical for genus, fourth pair with exopod longer than en- 
dopod, exopod terminating in long, plumose seta, endopod terminating in two 
long, simple setae. 

Telson as long as sixth abdominal somite, extending % length of inner uropod, 
linguiform, apex entire and narrowly rounded. Lateral margins armed with about 
40 short spines extending throughout entire length, 5-6 spines distinctly set, re- 
maining spines closely set, gradually increasing in length distally, apex armed with 
a pair of long spines Vio length of telson. 

Inner margin of inner uropod armed with 12 spines, increasing in length distally, 
distal 8 lying more remotely from each other than proximal 4. Exopod of uropod 
more than V3 again as long as endopod. 

Remarks. — Because of the similarity in telsons, this species may be superfi- 
cially confused with M. calif ornica Tattersall. It can be distinguished from the 
latter by the following features: 1) larger size, 2) absence of distal suture on 
antennal scale, 3) fewer spines on inner margin of the endopod of uropod. 



324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Acknowledgments 

I wish to thank Dr. M. Bacescu, Museul de Istorie Naturala "Grigore Antipa," 
for his help with the genus Cubanomysis, and Dr. T. E. Bowman for his help 
with M. cathengelae and the preparation of this manuscript. 

Funding for this study was supplied by the Marine Review Committee, Inc. 

Literature Cited 

Bacescu, M. 1968. Etude des quelques Leptomysini (Crustacea Mysidacea) des eaux du Bresil et de 

Cuba; Description d'un genre et de cinq autres taxons nouveaux. — Annali Museo Civico di 

storia Naturale di Genova 77:232-249. 
, and L. G. Gleye. 1979. New Mysidacea from the California waters. — Travaux du Museum 

d'Historie Naturelle Grigore Antipa 20:131-141. 
Brattegard, T. 1969. Mysidacea from shallow waters in the Bahamas and Southern Florida. Part 1. — 

Sarsia 39:17-106. 
Tattersall, Walter M. 1951. A Review of Mysidacea of the United States National Museum. — United 

States National Museum Bulletin 201 :x + 292 pp. 

Marine Ecological Consultants, 5333 Stevens Ave., Solana Beach, California 
92075. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 325-333 

THE STATUS OF CIROLANA PARVA HANSEN, 1890 

(CRUSTACEA, ISOPODA, CIROLANIDAE) WITH 

NOTES ON ITS DISTRIBUTION 

Niel L. Bruce and Thomas E. Bowman 

Abstract. — The type-material of Cirolana parva Hansen is redescribed, and 
the distribution of the species reassessed. Cirolana parva is rehably known only 
from the Caribbean Sea and the Gulf of Mexico; most non-Caribbean records are 
probably erroneous. Cirolana diminuta Menzies, 1962 is accepted as a valid 
species. 



Hansen (1890) in his monograph of the Cirolanidae described and figured Cir- 
olana parva. All but one of his specimens originated from the West Indies. The 
species was later recorded from the Caribbean area (Richardson 1900, 1921; 
Moore 1901; Menzies and Glynn 1968) and also throughout the Indo-Pacific and 
the East Pacific. The only areas from which the species has never been recorded 
are the coasts of South America and Europe. 

In examining cirolanids from Australian coasts it became apparent that there 
were some inconsistencies amongst specimens being identified as C. parva (Bruce 
1980a, 1980b) and Cirolana cranchii australiense Hale 1925 (Holdich et al. 1981). 
Detailed study of specimens from selected localities eventually led to the conclu- 
sion that what has been considered Cirolana parva is in fact a complex of over 
a dozen sibling species, at least 8 of which occur in Australia. It was found to be 
necessary to examine the syntypes of Cirolana parva to determine convincingly 
that species' absence from Australia. 

Since these species are all so similar to one another we redescribe Cirolana 
parva to prevent further application of that name to the numerous closely related, 
but distinct species. 

Cirolana parva Hansen 
Figs. 1, 2 

Cirolana parva Hansen, 1890:340, pi. 2, figs. 6-6b, pi. 3, figs. 1-ld. — Richardson, 
1900:217; 1905:111, figs. 93-95; 1912:178.— Moore, 1901:167, pi. 8, figs. 6-8.— 
Menzies & Glynn, 1968:38, fig. 14C-D. 

Non Cirolana /7flrvfl.— Stebbing, 1905:12; 1910:217.— NobiH, 1907:421.— Bar- 
nard, 1914:353a; 1936:154; 1940:499.— Chilton, 1924:883, fig. 5; 1926:1980.— 
Boone, 1927:136.— Monod, 1930:fig. 5A, C; 1931:3; 1933:173, fig. 80; 1937:15.— 
Nierstrasz, 1931:151. — Edmondson, 1951:192, fig. 4a-g. — Menzies & Franken- 
berg, 1966:51, fig. 27 A-C— Miller, 1968:15, fig 4.— Schultz, 1969:185.— Jones, 
1976:216.— Bowman, 1977:653, figs. 1-3.— Hamsa & Nammalwar, 1978:517, 
figs. 1-11.— Kensley, 1978:69, fig. 289.— Bruce, 1980a: 110; 1980b: 158. 

Material. — Cirolana parva syntypes: 2 6 (5.6, 6.9 mm), 3 $ (5.1, 5.5, 7.6 
mm), St. Thomas, West Indies. S (5.9 mm), 9 (6.3 mm) 25°N, 34°E. 9 (7.9 mm, 



326 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Cirolana parva syntypes: A-D, F, H, 5.3 mm d; E, G, 6.3 mm S; I-M, 5.5 mm $. A, 
Lateral view; B, Cephalon, dorsal view; C, Pleon and pleotelson; D, Pleotelson, posterior border; E, 
Peneal processes; F, Clypeal region; G, Sternite 7; H, Pleon, lateral view; I, Pereopod 1; J, Dactyl 
of pereopod 1; K, Pereopod 2; L, Antennal peduncle; M, Antennule. Scale line represents 1.0 mm. 



ovigerous), West Indies. S (6.9 mm), no locality, West Indies. $ (5.0 mm) Samoa. 
2 specimens (4.4, 5.0 mm), in very poor condition, St. Croix, West Indies. Cir- 
olana parva non-types: Specimens from the localities shown in Fig. 4 from the 
collections of the National Museum of Natural History, Smithsonian Institution. 
Not Cirolana parva: Hundreds of Australian specimens of Cirolana spp., spec- 



VOLUME 95, NUMBER 2 



327 




Fig. 2. Cirolana parva syntypes: A-D, 5.5 mm 6; E-J, 6.9 mm S . A, Maxilliped; B, Uropod; C, 
Uropod exopod, apex; D, Uropod endopod, apex; E, Pereopod 7; F, Appendix masculina, apex; G, 
Pleopod 1; H, Pleopod 2; I, Pleopod 4; J, Pleopod 5. 



imens of both sexes of C. diminuta Menzies, 1962, from La JoUa, California, and 
of C. parva of Jones (1976). 

Type-locality. — Hansen (1890) gave no type-locality, but as one of the St. 
Thomas specimens has been dissected, it seems apparent that Hansen's descrip- 
tion is based on these specimens. St. Thomas is here recognized as the type- 
locality. The type-specimens are held by the Zoologisk Museum, Copenhagen, 
Denmark. 

Male. — Body about 2% times longer than greatest width, surface minutely 
punctate. Cephalon with rostral process; dorsal surface with anterior interocular 
carina, and entire faint furrow running between the dorsomedial margin of each 
eye, medial part of furrow posteriorly deflected; maxillipedal somite indicated by 
2 furrows at posterolateral margins. Pereonite 1 with 2 horizontal furrows; per- 
eonites 2-4 each with horizontal furrow, pereonites 5-7 each with oblique furrow; 
coxae of pereonites 2-7 each with entire furrow; coxae of pereonites 4-7 pro- 
jecting beyond posterior margins of segments. Pleon with segment 1 largely con- 



328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

cealed by pereonite 7; pleonite 2 with posterolateral margins acute; pleonite 3 
with posterolateral margins produced and encompassing pleonite 4, superior mar- 
gin convex; pleonite 4 with superior margin rounded, small projection where 
horizontal furrow meets margin. Pleotelson with smoothly curving lateral mar- 
gins; posterior border with 8 spines, each set within slight identation; short plu- 
mose setae lie on each side of spines. 

Antennule with peduncular articles 1 and 2 fused; peduncular article 3 with 
fused 4th article present; flagellum extending to pereonite 1, composed of 9 ar- 
ticles, first of which is longest. Antenna with peduncular articles 1-3 short, pe- 
duncular articles 4 and 5 long, subequal in length; flagellum composed of about 
26 articles, extending to pereonite 6. 

Frontal lamina pentagonal, lateral margins straight, slightly divergent, anterior 
margins concave; apex overlapped by downward projection of rostral process. 
Clypeus narrow. The fragility of the specimens did not allow a full dissection of 
the mouthparts, but they appear similar to those of other members of the genus 
(sensu strictu, Bruce 1981). Maxilliped with 2 coupling hooks and 6 plumose setae 
on endite. 

Pereopod 1 with setae at posterodistal angle of basis; ischium with single spine 
at posterodistal angle and 2 setae at superior anterodistal angle; merus with pos- 
terior margin slightly sinuate, provided with 5 robust and 2 acute spines; carpus 
with single seta and acute spine at posterodistal angle; propodus with 2 spines on 
palm, and third stout spine opposing the biunguiculate dactyl. Pereopods 2 and 

3 similar, less robust than pereopod 1 with merus, carpus and propodus propor- 
tionally longer, and with more numerous and larger spines. Pereopods 4-7 pro- 
gressively longer. Pereopod 7 with virtually no setae, groups of spines at anterior 
distal angle of ischium, and propodus with 2 groups of spines, merus and carpus 
with one; distal angles of propodus each with 2 spines. 

Penes present on sternite 7; short, triangular, projecting posteriorly, set close 
together. 

Pleopod 1 endopod with slightly concave lateral margin, broadest at Vs of its 
length from apex. Pleopod 2 with appendix masculina arising basally, extending 
beyond endopod by Vs its length, apex narrows rapidly to an acute slender point. 
Exopods of pleopods 3-5 with entire transverse suture. Endopod of pleopod 3 
with about 13 setae, pleopod 4 with about 9. Peduncles of pleopods 2^ each with 

4 coupling hooks. 

Uropods projecting slightly beyond apex of pleotelson. Exopod slightly shorter 
than endopod, lateral margin with 7 spines and short plumose marginal setae, 
medial margin with 3 spines; apex bifid with medial process more prominent than 
lateral. Endopod with 2 spines and sensory seta on lateral margin; medial margin 
with 4 spines amongst plumose marginal setae; apex bifid. 

Female. — Differs from the male only in the sexual characters. 

Color. — Pale tan in alcohol. Chromatophores bleached out. Related species 
show a variety of chromatophore patterns (pers. obs.). 

Size. — Syntypes range from 5.6-6.9 mm for the males, and 5.0-7.6 mm for 
females . 

Variation. — The dorsal cephalic furrow is not always easy to observe and the 
indentation of the posterolateral margin of pleonite 4 is only present in the type- 



VOLUME 95, NUMBER 2 



329 




Fig. 3. Cirolana diminuta: A-I, 6.3 mm S; La Jolla, California. A, Pleon, lateral view; B, Uro- 
pods; C, Uropod endopod, apex; D, Uropod exopod, apex; E, Pleotelson, dorsal view; F, Pleotelson, 
apex; G, Pleopod 1; H, Pleopod 2; I, Appendix masculina, apex. Cirolana parva, appendix masculina, 
apices: J, Montego Bay, Jamaica; K, Quintana Roo, Mexico; L, No Name Key, Florida. 



specimens. The apex of the appendix masculina varies sHghtly as shown in 
Fig. 3. 

The spination of the pleotelson and uropods, position of the penes, and mor- 
phology of pleopods 1 and 2 are constant throughout the specimens examined. 

Remarks. — The characters that most readily separate C. parva from other sim- 
ilar species are the cephalic furrowing, the spination and shape of the pleotelson 
and uropods, the shape of the endopod of pleopod 1, the morphology of the 
appendix masculina, the shape and position of the penes, the shape of the lateral 
margins of the pleonites and the form of the uropod apices. Cirolana diminuta 



330 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Known distribution of Cirolana parva, confirmed from syntypes and from other specimens 
in the National Museum of Natural History, Smithsonian Institution. 1, St. Thomas; 2, St. Croix; 3, 
San Juan, Puerto Rico; 4, La Parguera, Puerto Rico; 5, Northwestern Cuba; 6, Florida Keys, Florida; 
7, Dry Tortugas, Florida; 8, South of Apalachicola, Florida; 9, Montego Bay, Jamaica; 10, Espiritu 
Santo Bay, Quintana Roo, Mexico. 



can be separated by the differences in the characters figured here (Fig. 3A-L) 
and also by having the vasa deferentia opening flush with the surface of sternite 
7. 

Distribution. — Rehable records exist only for the Caribbean and the Gulf of 
Mexico. These are reviewed by Richardson (190.5). Menzies and Frankenberg 
(1966) record the species from Georgia but figure the uropod with 6 spines on the 
medial margin of the endopod, and as the appendix masculina is not quite the 
same, this record needs confirmation before it can be accepted. All other records 
are either without figures and cannot be confirmed without reference to the spec- 
imens, or can be refuted by reference to the figures given (Monod 1930; 1933; 
1976), or by reference to specimens (C. parva of Jones 1976, C. diminuta Men- 
zies, C. parva of Bruce 1980a, b). 

The specimen labelled as from "Samoa" appears identical to all the other 
syntypes, and it is likely that the locality is in error, a possibility considered by 
Hansen (1890). 



Discussion 

In his revision of the family Cirolanidae, Hansen (1890) described Cirolana 
parva, the description being accompanied by excellent figures. Several subse- 
quent records (Richardson 1900, 1912; Moore 1901) were from the Caribbean, the 



VOLUME 95, NUMBER 2 331 

source of the original specimens. Shortly after the turn of the century reports 
were published from further afield, from Ceylon (Stebbing 1905), the Red Sea 
(Stebbing 1910), Polynesia (Nobili 1907) and southeastern Africa (Barnard 1914). 
Although none of these early reports gave figures, they apparently led to general 
acceptance that Cirolana parva is cosmopolitan in distribution. This acceptance 
was reinforced by numerous subsequent records and by 1980 the species had 
been recorded from Kenya (Jones 1976), West Africa (Monod 1931), Thailand 
(Chilton 1924, 1926), Hawaii (Edmondson 1951; Miller 1968), Indonesia (Nier- 
strasz 1931), Cocos Island in the East Pacific (Bowman 1977) and Australia (Bruce 
1980a, 1980b). The synonymization of Cirolana diminuta extended the range to 
include the western coast of the United States. 

In examining ""Cirolana parva'' from Australia, there appeared to be a great 
deal of variation present in samples from one locality and from different localities. 
In some series the males had a dense fringe of setae along the inferior margins 
of the first pereopod articles. Specimens with this character could not satisfac- 
torily be identified as Cirolana parva, yet neither could they be identified as 
Cirolana crane hii australiense (see Bruce & EUis (1982) for discussion of this 
species). This prompted the detailed study of all Australian specimens available, 
and eventually 8 species were determined. These different species are primarily 
separated by differences in details of the pleon, pleopods 1 and 2, uropod and 
pleotelson morphology and spination, setation of the male first pereopod, position 
of the vasa deferentia, and articulation and proportional length of the antennule 
articles. In general the antenna, mouthparts, pereonal morphology and pereopod 
morphology were found ineffective in separating species. 

Comparison of the syntypes with specimens recorded by Bruce (1980a) which 
include two species, and by Bruce (1980b), and to other unrecorded material 
reveals that Cirolana parva does not occur in Australia. Similarly, Kenyan spec- 
imens proved not to be C. parva. The record of Hamsa & Nammalwar (1978) is 
of a species related to Natatolana woodjonesi (Hale, 1925), and Monod's (1933) 
record, one of the few with figures, is also not C parva. 

One further species needs discussion, Cirolana diminuta, from California. 
Menzies and Glynn (1968) concluded that the differences shown by C. diminuta 
were insufficient to keep it separate from C. parva. Specimens from La Jolla 
identified as C. parva show several differences from C. parva (see Fig. 3), and 
the shape of the pleotelson readily identifies them as adult C. diminuta. 

A further complication in dealing with the taxonomy of Cirolana parva resulted 
from confusion as to whether or not the species was distinct from C. cranchii 
Leach. This idea was initiated by Stebbing (1917), and supported by Nordenstam 
(1946). Monod (1976) contributed to the argument, and concluded that although 
the 2 species were not separable, they were not necessarily conspecific. Bruce 
and Ellis (1982) have shown that the 2 species are abundantly distinct, as in fact 
can be seen from Hansen's (1890) figures. 



Acknowledgments 

The authors thank Dr. J. Just, Universitet Zoologisk Museum, Copenhagen, 
for loan of type-material, and Dr. R. C. Brusca, Allan Hancock Foundation, for 
donation of Calif ornian specimens. The work was supported by an Australian 



332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Biological Resources Study grant, by an Australian Museum Postgraduate Award, 
and by a Commonwealth Postgraduate Study Award. 

Literature Cited 

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marine Isopoda, with notes on some previously incompletely known species. — Annals of the 

South African Museum 10:325a-358a, 359-442. 
. 1936. Isopods collected by R.I. M.S. "Investigator." — Records of the Indian Museum 38:147- 

191. 
. 1940. Contributions to the crustacean fauna of South Africa 12. Further additions to the 

Tanaidacea, Isopoda and Amphipoda, together with keys for the identification of hitherto 

recorded marine and freshwater species. — Annals of the South African Museum 32:381-515. 
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of 1938. — Proceedings of the Biological Society of Washington 89:653-666. 
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Group. — Bulletin of Marine Science 30:108-130. 
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Pacifique 2:155-173. 
. 1981. The Cirolanidae (Crustacea: Isopoda) of Australia. Diagnoses of Cirolana Leach, 

Metacirolana Nierstrasz, Neocirolana Hale, Anopsilana Paulian and Debouteville and three 

new genera — Natatolana, Politolana and Cartetolana. — Australian Journal of Marine and 

Freshwater Research 32:945-966. 
, and J. Ellis. 1982. Cirolana cranchii Leach, 1818 (Crustacea, Isopoda, Cirolanidae) rede- 
scribed. — Bulletin of the British Museum (Natural History), Miscellanea (in press). 
Chilton, C. 1924. Fauna of Chilka Lake. Tanaidacea and Isopoda. — Memoirs of the Indian Museum 

5:875-895. 
. 1926. The Tanaidacea and Isopoda of Tale Sap (Siam). — Records of the Indian Museum 

28:173-185. 
Edmondson, C. H. 1951. Some Central Pacific crustaceans. — Occasional Papers of the Bernice P. 

Bishop Museum, Honolulu 20:183-243. 
Hale, H. M. 1925. Review of Australian isopods of the cymothoid group Part 1. — Transactions of 

the Royal Society of South Australia 49:128-185. 
Hamsa, K. M. S. A., and P. Nannalwar. 1978. Description of isopod Cirolana parva parasitic on the 

eyeball of dolphin, Delphinus delphis with a key to the Indian species of the genus Cirolana. — 

Journal of the Bombay Natural History Society 75:516-519. 
Hansen, H. J. 1890. Cirolanidae et familiae nonnulae propinquae Musaei Hauniensis. — Kongeligt 

Videnskabernes Selskab Skrifter, 6, Raekke, Naturvidenskabelig og Mathematisk Afdeling 

5:237-426. 
Holdich, D. M., K. Harrison, and N. L. Bruce. 1981. Cirolanid isopod crustaceans from the Towns- 

ville region of Queensland, Australia, with descriptions of six new species. — Journal of Natural 

History 15:555-605. 
Jones, D. A. 1976. The systematics and ecology of some isopods of the genus Cirolana (Cirolanidae) 

from the Indian Ocean region. — Journal of Zoology, London 178:209-222. 
Kensley, B. 1978. Guide to the marine isopods of Southern Africa. — South African Museum, Cape 

Town, 173 pp. 
Menzies, R. J. 1962. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. — 

Pacific Naturalist 3:337-348. 
, and D. Frankenberg. 1966. Handbook on the common marine isopod crustaceans of Geor- 
gia. — University of Georgia Press, Athens, 93 pp. 
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the Fauna of Curasao and other Caribbean Islands 27:1-133. 
Miller, M. A. 1968. Isopoda and Tanaidacea from buoys in coastal waters of the continental United 



VOLUME 95, NUMBER 2 333 

States, Hawaii, and the Bahamas (Crustacea). — Proceedings of the United States National 
Museum 125 (3652): 1-53. 

Monod, T. 1930. Contribution a Fetude des "Cirolanidae." — Annales des Sciences Naturelles, Zool- 
ogie 10(13): 129-183. 

. 1931. Sur quelques crustaces aquatiques d'Afrique (Cameroun et Congo). — Revue de Zool- 

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. 1933. Mission Robert-Ph Dollfus en Egypte: Tanaidacea et Isopoda. — Memoires de I'lnstitute 

d'Egypte 21:161-264. 

. 1937. Missions A. Gruvel dans le Canal de Suez. I. Crustaces. — Memoires de I'lnstitute 

d'Egypte 34:1-19. 

. 1976. Remarques sur quelques Cirolanidae (Crustacea: Isopoda). — Bulletin du Museum Na- 
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Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 334-337 

DESCRIPTION OF THE MALE AND NOTES ON 

THE FEMALE OF ARGEIOPSIS INHACAE 

(CRUSTACEA: ISOPODA: BOPYRIDAE) 

Daniel L. Adkison, Richard W. Heard, and Guy T. Clark 

Abstract. — The male of Argeiopsis inhacae Kensley, 1974 is described for the 
first time from one specimen. The diagnosis for the male is: head fused with first 
pereomere; first antenna of 3 segments, second antenna of 5 segments, basal 
segment with posteriorly directed projection; midventral tubercles on pereo- 
meres; and pleon fused, segments laterally indicated, mid- ventral tubercles, pleo- 
pods, and uropods absent. The viability of the subfamily Argeiinae Markham, 
1977 is questioned. Based on a review of the literature and examination of the 
holotype A. inhacae and specimens from the Philippine Islands, the range of A. 
inhacae is extended from Mozambique to the Philippine Islands. 



The genus Argeiopsis Kensley, 1974 is based on a single species, Argeiopsis 
inhacae Kensley, 1974, described from a single ovigerous female collected from 
Mozambique on the banded coral shrimp Stenopus hispidus (Oliver). The present 
report is based on the examination of 2 females and a male of A. inhacae obtained 
by one of us (GTC) from an aquarium shop in Norfork, Virginia. Host shrimp 
were collected in the Philippine Islands and shipped to Norfork, Virginia as part 
of a consignment to a marine aquarium shop. Efforts to learn more precise col- 
lection data were unsuccessful. 

Argeiopsis inhacae Kensley, 1974 
Fig. 1 

Argeiopsis inhacae Kensley, 1974:259-261, fig. 1. — Markham, 1977:109-110. 

Material examined. — Infesting Stenopus hispidus. Inhaca Island, Mozambique; 
from right branchial chamber of Stenopus hispidus', 1 ? (holotype, ovigerous) 
SAM A 10979. — Philippine Islands; 1977 (aquarium trade, no other collection data 
known); from left branchial chamber of 5. hispidus; 1 ? (ovigerous), 1 S USNM 
172471. Philippine Islands; 1977 (aquarium trade, no other collection data); from 
right branchial chamber of S. hispidus; 1 9 (non-ovigerous, no S) USNM 172472. 

Description. — Female. Distortion slight. Head pentagonal, width less than 
twice length; anterior lamina present, medially indistinct, laterally curled dorsally. 
Small eyes on anterolateral margin of head. First antenna of 3 segments; second 
antenna of 4 segments. Maxilliped covered with scales; palp present, medially 
with fringe of long setae. Postero ventral lamina with 2 pairs of sickle shaped 
projections. 

Pereon broadest at pereomere 3; pereomere 3 nearly straight, pereomeres 4 to 
7 progressively directed more posteriorly. Narrow coxal plates on pereomeres 
1 to 4. Dorsal bosses present on pereomere 1 to 4. Pereomere 1 to 4 bilobed 
laterally, tergal area of pereomeres of expanded side enlarged; pereomere 1 on 
enlarged side with tergal area L- shaped or hooked anteriorly. Pereopods increas- 



VOLUME 95, NUMBER 2 



335 




Fig. 1. Argeiopsis inhacae, female a-d (USNM 172472), male e-h (USNM 172471): a, Dorsal 
view; b. Left oostegite 1, internal view; c, Pleon, dorsal view; d, Uropods and right pleopod 5, ventral 
view; e, Dorsal view; f, Right antennae; g. Left pereopod 1; h, Pleon, ventral view. 



ing in size posteriorly. Oostegites only laterally defining brood pouch. Oostegite 
1 with internal ridge unornamented; posterior plate developed into laterally di- 
rected point. 

Pleon of 6 distinct segments, length subequal to width. Lateral plates absent. 
Pleopods biramous, 5 pairs; larger on expanded side; pleopodal endopods de- 



336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

creasing in size posteriorly. Uropods biramous, uropodal exopod similar in size 
and shape to exopod of pleopod 5; endopods reduced, smaller than endopod of 
pleopod 5. 

Variation. — Variation was found in the relative development of coxal plates, 
dorsal bosses, pleopodal endopod, and uropodal endopods. The holotype has 
weakly developed coxal plates, dorsal bosses and uropodal endopods. On the 
other Philippine specimens, these structures are better developed. The uropodal 
endopod of the holotype is reduced to a small median projection. The uropodal 
endopods of the non-ovigerous female are about of the endopod of pleopod 5. 

Male. — Head narrower than first pereomere; head fused with first pereomere. 
Eyes present. First antenna of 3 segments; second antenna of 5 segments, prox- 
imal segment with posteriorly directed lobe; second antenna less than twice length 
of first; both antennae covered with scales. 

Pereon of 7 segments; pereomeres 1 and 7 narrowed, seventh narrowest; dorsal 
pigment spots on pereomere 2 to 7. Pereopods of similar size; covered with scales, 
scales with fringe of setae. Midventral tubercles on all pereomeres. 

Pleon fused, segments indicated laterally, first most prominent; 5 segments 
indicated on right side, 4 on left side. Pleopods, midventral tubercles and uropods 
absent. Anus on slight ventral lobe, flanked laterally with setae. 

Remarks. — Both the holotype and the ovigerous female from the Philippines 
have the cuticle pulled away from the body. This condition makes the determi- 
nation of the segmentation of the pleon and pleopods most difficult even when 
the specimens are stained. 

For female bopyrids with pleopods set on or near the lateral margin, it is 
difficult to determine whether one is dealing with a biramous pleopod or a uni- 
ramous pleopod and lateral plate. The pleopods of Argeiopsis inhacae are such 
a case, and have been considered both uniramous with lateral plate (Markham 
1977) and biramous without lateral plate (Kensley 1974). We believe that the 
pleopods should be considered biramous (lateral plate absent) since the point of 
pleopod attachment is on the ventral surface. We also feel that the presence of 
biramous uropods supports this conclusion because the uropods are similar in 
structure to the pleopods. 

The generic diagnosis presented below is amended from Kensley (1974) to 
include the male and changes necessary from the study of the 3 female specimens. 

Diagnosis. — Female: Body nearly circular; all regions and segments distinct; 
anterior lamina present, not greatly developed; dorsal bosses on pereomeres 1 to 
4; coxal plates narrow or reduced, present on pereomeres 1 to 4; oostegites only 
laterally defining brood pouch; pleon without lateral plates; pleopods 5 biramous 
pairs; uropod biramous, endopod reduced. Male: head fused with first pereomere; 
head narrower than first pereomere; pereon of 7 segments, sides nearly parallel; 
pleon with lateral indication of pleomeres; pleon narrower than pereon; pleopods, 
midventral tubercles, and uropods absent. 

The 5 important differences fisted by Markham (1977) distinguishing the Ar- 
geiinae from the Bopyrinae (head shape, body outline, pleomere fusion, lateral 
plate and uropod development, and number of pleopod rami) appear variable and 
insufficient to maintain Argeiinae as a separate subfamily. In addition, the genera 
Argeiopsis and Stegoalpheon Chopra, 1923 appear to have biramous pleopods. 
Argeiopsis has biramous pleopods and no lateral plates. Chopra (1923), when 



VOLUME 95, NUMBER 2 337 

describing Stegoalpheon kempi Chopra, 1923, reported the pleopods to be tri- 
ramous (lateral plates absent) but noted that the pleopods would be biramous 
were lateral plates present. Of the 5 genera in Argeiinae {Argeia Dana, 1853; 
Argeiopsis, Bopyrosa Nierstrasz and Brender a Brandis, 1923; Parargeia Hansen, 
1897 and Stegoalpheon), only Parargeia has a closed brood pouch and therefore 
appears more similar to the Pseudioninae than to the Bopyrinae. The 4 other 
genera have an open brood pouch and are therefore more similar to the Bopyri- 
nae. Since the location of the types of Bopyrosa phryxiformis Nierstrasz and 
Brender a Brandis, 1923 is unknown and the single known specimen appears to 
be immature, its placement is at best tentative (Markham 1977). Until members 
of Parargeia and Stegoalpheon can be examined, we tentatively continue to 
recognize Argeiinae as a distinct subfamily. 

Acknowledgments 

We wish to thank Elizabeth Louw of the South African Museum for loan of 
the holotype of A. inhacae and Brian Kensley of the National Museum of Natural 
History for comments and suggestions offered during a visit to the museum. This 
paper is a contribution of the Marine Environmental Sciences Consortium, 
P.O. Box 386, Dauphin Island, AL 36528. 

Literature Cited 

Chopra, B. 1923. Bopyrid isopods parasitic on Indian Decapoda Macrura. — Records of the Indian 

Museum 25:411-550. 
Dana, J. D. 1853. Crustacea. — United States Exploring Expedition during the years 1838-42 under 

the command of C. Wilkes 13(2):691-1018. 
Hansen, H. J. 1897. Reports on the dredging operations off the west coast of Central America . . . 

by the U.S. Fish Commission Steamer "Albatross" during 1891 . . . XXII. The Isopoda. — 

Bulletin of the Museum of Comparative Zoology, Harvard College 31(5):96-129. 
Kensley, B. 1974. Bopyrid Isopoda from southern Africa. — Crustaceana 26(3): 259-266. 
Markham, J. C. 1977. Description of a new Western Atlantic species oi Argeia Dana with a proposed 

new subfamily for this and related genera (Crustacea Isopoda, Bopyridae). — Zoologische Me- 

dedelingen 52(9): 107-123. 
Nierstrasz, H. F., and G. A. Brender a Brandis. 1923. Die Isopoden der Siboga-Expedition II. 

Isopoda Genuina. I. Epicaridea. — Siboga-Expeditie Monographic 32b:57-121, pis. 4-9. 

(DLA) Dauphin Island Sea Laboratory, Box 386, Dauphin Island, Alabama 
36528 (present address: Department of Biology, Tulane University, New Orleans, 
Louisiana 70118); (RWH) Gulf Coast Research Laboratory, Ocean Springs, Mis- 
sissippi 39564; (GTC) Department of Biology, Old Dominion University, Norfolk, 
Virginia 23508. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 338-346 

A DIAGNOSIS OF THE HOBBSI GROUP, WITH 

DESCRIPTIONS OF CAECIDOTEA TERES AE, N. SP., 

AND C. MACROPROPODA CHASE AND BLAIR 

(CRUSTACEA: ISOPODA: ASELLIDAE) 

Julian J. Lewis 

Abstract. — The Hobbsi Group has emerged as a major assemblage of both epi- 
gean and hypogean species found over much of the eastern United States. With 
the new diagnosis of the group, 17 species are now tentatively assigned. Caeci- 
dotea teresae, a phreatobite from southern Indiana, is described herein as an 
addition to the group, and C. ozarkana is synonymized with C. macropropoda. 



During his work on subterranean isopods, Steeves made the first attempt to 
divide some of the troglobitic species of the family Asellidae into assemblages 
which reflected their evolutionary histories: the Stygia (Steeves 1963), Hobbsi 
(Steeves 1964) and Cannulus groups (Steeves 1966). Of these, the Cannulus Group 
has received the most attention (Lewis 1981). However, with the addition of new 
species and the transfer of others, the Hobbsi Group is emerging as a dominant 
element among the North American asellids. 

The Hobbsi Group is especially interesting from an evolutionary standpoint, 
since unlike the Stygia and Cannulus groups, it contains both epigean and hy- 
pogean species. Within the group, morphological intergradations exist between 
eyeless, unpigmented troglobites, vestigially eyed and pigmented phreatobites, 
to epigean species with functional eyes and pigmentation. The following diagnosis 
expands that of Steeves (1964). 

The Hobbsi Group 

Diagnosis of male. — Eyes and pigmentation present or absent. Palmar margin 
of pereopod 1 propus with strong processes. Pleopod 1 longer than pleopod 2; 
lateral margin of exopod concave, with weak nonplumose setae, distal margin 
with elongate, plumose setae. Pleopod 2, exopod with numerous elongate, plu- 
mose setae along margin of distal segment; endopod, basal apophysis well de- 
fined; tip with low conical cannula, extending along the axis of the endopod, often 
obscured by other terminal processes. 

The following species are tentatively assigned to the Hobbsi group: Troglo- 
bites — Caecidotea acuticarpa, C. adenta, C.fustis, C. hobbsi, C. macropropoda, 
C. nickajackensis , C. packardi, C. reddelli, C. salemensis; Phreatobites — C. ken- 
deighi, C. spatulata, C. teresae, C. tridentata; and Epigean — C. brevicauda, C. 
dentadactyla, C. kenki, C. scrupulosa. 

Discussion. — The most unusual additions to the Hobbsi Group are undoubtedly 
the epigean species. Although Williams (1970) did not assign species to groups, 
in his cladogram of epigean asellids C. dentadactyla, C. kenki, C. scrupulosa, 
and C brevicauda formed a lineage related to C. montanus and C. nodula. Of 
these latter two species, C. nodula appears to have some affinities with the Hobb- 



VOLUME 95, NUMBER 2 339 

si Group, especially in the morphology of the first pleopod, but the torsion of the 
endopod tip is not apparent in other members of the group. Caecidotea montanas 
relationships with the Cannulus Group have been discussed elsewhere (Lewis 
1981). 

Unlike other epigean Caecidotea, which occur frequently in warm, lotic hab- 
itats, the epigean members of the Hobbsi Group occur mostly in cold water 
habitats. This is especially true of C. brevicauda, C. kenki, and C. scrupulosa, 
which are spring stream inhabitants and facultative cavernicoles . 

Three other species deserve mention, C. parva (included by Steeves 1964 in 
the original Hobbsi Group), C bisetus, and C. pilus. Although the endopod tips 
of these species are of the type found in members of the Hobbsi Group, the 
similarity ends there. The exopod of the first pleopod is ovate, not laterally con- 
cave, and weakly setose; the distal segment of the second pleopod bears only 1- 
5 setae; and the palmar margin of the gnathopod propus lacks processes. It is 
likely that in the future one or more of these species will be assigned to the genus 
Lirceolus. 

Caecidotea teresae, new species 
Figs. 1-2 

Caecidotea sp. #2.— Lewis, 1981:582-583, 585. 

Material examined. — INDIANA: Floyd Co., Indiana University Southeast 
campus, New Albany, drain tile near parking lot below Natural Sciences Building, 
19 Nov 1977, J. Lewis, T. Everitt (Lewis), 28 c?, 19 9. Same, 12 Dec 1978, J. 
Lewis, 5 c?, 6 9. Drain tile flowing into small campus pond, near service area, 
22 Mar 1980, T. Lewis, 1 d, 2 9. 

A 12 mm male, 19 Nov 1977, is the holotype (USNM 189468), the other spec- 
imens are paratypes (USNM 189469, USNM 189470, USNM 189471). All of the 
material examined has been deposited in the collections of the National Museum 
of Natural History, Smithsonian Institution. 

Description. — Eyeless, dorsum with light scattered pigmentation, darkest an- 
teriorly. Largest male to about 15.5 mm, female to about 9.5 mm. Body slender, 
linear, about 6.8x as long as wide in males, 5.1x in females. Margins of head, 
pereonites and telson moderately setose. Head about 1.5 x as wide as long, an- 
terior margin concave, postmandibular lobes moderately produced. Telson about 
1.3 X as long as wide, sides subparallel, caudomedial lobe moderately produced. 

Antenna 1 of male reaching midlength of last segment of antenna 2 peduncle, 
flagellum with up to about 13 segments, esthete formula 3-0-1-0-1; females with 
up to about 9 segments, 3-0-1. Antenna 2 about 0.6 length of body, last segment 
of peduncle about 1.3x as long as preceding segment, flagellum of about 95-100 
segments. 

Mandibles with 4-cuspate incisors and lacinia; distal segments of palp with 
plumose spine rows. Maxilla 1, outer lobe with 13 robust apical spines; inner lobe 
with 5 plumose setae apically. Maxilliped with 7 retinacula. 

Pereopod 1 of male, propus 1.5x as long as wide, palm with proximal triangular 
process (spine in less mature specimens), larger trapezoidal median process sep- 
arated by U-shaped cleft from shorter, rectangular, bicuspid distal process; dactyl 
flexor margin with weak spines. Female pereopod 1 propus about 1.5x as long 



340 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Caecidotea teresae, A from female paratype, B-H from male paratype: A, Habitus, dorsal; 
B, Pleopod 2; C, Same, endopod tip, posterior; D, Same, endopod tip, anterior; E, Uropod, dorsal; 
F, Maxilla 1, inner lobe; G, Same, outer lobe; H, Pereopod 1. 



as wide, processes absent, large proximal spine present; dactyl flexor margin with 
about 6 strong spines. Pereopod 4 more robust and spinose in males than females. 
Male pleopod 1 larger than pleopod 2; protopod about 0.7 length of exopod, 
with 8 retinacula. Exopod about 0.6 x as wide as long, lateral margin concave, 
with about 11 elongate plumose setae along distal margin. Pleopod 2, protopod 



VOLUME 95, NUMBER 2 



341 




Fig. 2. Caecidotea teresae, male paratype: A, Head and antennae, dorsal; B, Antenna 1, distal 
segments; C, Pereopod 4; D, Pleopod 3; E, Pleopod 1; F, Pleopod 4; G, Pleopod 5. 



with about 10 setae along medial margin. Exopod, proximal segment with up to 
8 lateral setae, distal segment with about 23 elongate, plumose setae. Endopod 
with distinct basal apophysis; tip of endopod terminating in 3 processes: (1) can- 
nula trapezoidal, low, wide, apically blunt, partly obscured by other processes; 
(2) mesial process slightly longer than cannula, slightly recurved, forming lateral 



342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

wall of endopodial groove; and (3) caudal process low, broadly rounded, heavily 
sclerotized. Female pleopod 2 pyriform, with about 13 elongate plumose setae 
along lateral margin, 2-3 setae on proximal-mesial margin. Pleopod 3 exopod, 
proximal segment about 0.7 length of distal segment, with about 12-13 setae on 
lateral margin; distal segment with about 19-20 elongate plumose setae on distal 
margin. Pleopod 4, about 5 setae on proximal lateral margin, 2 false sutures 
present. Pleopod 5, 3 setae on proximal lateral margin. 

Uropods showing pronounced sexual dimorphism; male uropod about 1.5x 
length of pleotelson; female uropod about 0.7 x length of pleotelson. 

Etymology. — This species is named for Teresa M. Lewis, in recognition of her 
collection of not only this species, but many other subterranean asellids from the 
eastern U.S. 

Relationships . — Caecidotea teresae has its closest morphological affinities with 
C. tridentata, C. salemensis, and an undescribed species from Illinois (Lewis and 
Bowman 1981). The relationship of Caecidotea teresae to these species was dis- 
cussed in detail previously (Lewis 1981) and will not be repeated here. 

Habitat and distribution. — Caecidotea teresae is known only from drain tiles 
on the New Albany campus of Indiana University Southeast. A small stream 
flows from a drain tile near the Natural Sciences Building, which at times is 
inhabited by a dense population of C. teresae. In addition to the phreatobitic 
isopods, an undescribed amphipod of the genus Stygobromus has also been taken 
from a drain tile near the campus service center. Both of these drain tile streams 
disappear during dry weather. 

Caecidotea macropropoda Chase and Blair 
Fig. 3 

Caecidotea macropropoda Chase and Blair, 1937:220-224. — Nicholas, 1960:131. 

Kenk, 1973:13. 
Caecidotea ozarkana Chase and Blair, 1937:220-224. Nicholas, 1960:131. 
Caecidotea macropropodus. — Mackin, 1940:17-18, — Mackin and Hubricht, 

1940:395. 
Asellus macropropodus. — Hubricht, 1950:17. — Dearolf, 1953:227. — Bresson, 

1955:51.— Mackin, 1959:875.— Black, 1971:6.— Fleming, 1973:294,298. 
Asellus ozarkanus. — Bresson, 1955:51. — Fleming, 1973:294, 298. 
Conasellus macropropodus. — Henry and Magniez, 1970:356. 
Conasellus ozarkanus. — Henry and Magniez, 1970:356. 
Asellus ozarkana. — Black, 1971:7. 

Material examined. — OKLAHOMA: Adair Co., stream in cave, 5 mi S. Kan- 
sas, A. P. Blair, 27 Dec 1935, 8 S (USNM 8625); cave (same as previous collec- 
tion?), A. P. Blair, 27 Dec 1935, 3 6\ Spring, 5 mi S. Kansas, A. P. Blair, 12 Jul 
1936, 1 (?, 3 ? (USNM 108660); Cave, 5 mi. S. Kansas, coll. unknown, 15 Sep 
1960, 3 c^, 3 9; Bat Cave, 5 mi. S. Kansas, J. Lewis, T. Lewis, 1 June 1981, 17 
cJ, 9 9; Cave behind old greenhouse, J. Lewis, T. Lewis, 1 June 1981, 2 c?, 5 
9; Cave behind Hardwicks house, 1 June 1981, 1 c^. 

Description of male. — Eyeless, unpigmented, length to about 15.0 mm. Body 
slender, linear, about 6x as long as wide; coxae visible in dorsal view. Head 
about 1.8x as long as wide, anterior margin concave, rostrum absent. Telson 
about 1.4x as long as wide, sides subparallel. 



VOLUME 95, NUMBER 2 



343 




Fig. 3. Caecidotea macropropoda, A-B, from male topotype of C. ozarkana, spring 5 miles south 
Kansas; C-H from male topotypes of C. macropropoda, Bat Cave, 5 miles south Kansas: A, Pleopod 
2 endopod tip, anterior; B, Same, posterior; C, Same, posterior; D, Pereopod 1, palmar margin of 
propodus; E, Antenna 1, distal segments, with abnormal esthete placement on 5th segment; F, Pleo- 
pod 1; G, Pleopod 2; H, Pleopod 4. Caecidotea stiladactyla, male, from small seep 9 miles southwest 
of Harrison, Boone Co., Arkansas (USNM 108666): I, Pleopod 2, endopod tip, anterior, recumbent 
position. 



Antenna 1 of about 13 segments, esthete formula 3-0-1. Antenna 2, last segment 
of peduncle about 1.5x length preceding segment, flagellum to about 105 seg- 
ments. 

Mandibles with 4-cuspate incisors and lacinia mobilis; distal segments of palp 



344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

with plumose spine rows. Maxilla 1, outer lobe with 13 robust spines, inner lobe 
with 5 robust, plumose setae. Maxilliped with 4-5 retinacula. 

Pereopod 1 propus about 1.5x as long as wide, with 3 processes: stout proximal 
spine, or spine-like process; median process large, blade shaped; distal process 
shorter, bicuspid, becoming lower and broader in larger individuals. Dactyl flexor 
margin with massive, lobed process. 

Pleopod 1 longer than pleopod 2, protopod with 6-7 retinacula; exopod about 
1.8x as long as wide, widened proximally with 1-2 proximomedial setae, lateral 
margin concave, bearing tiny spines, distal margin with about 10 non-plumose 
setae. Pleopod 2, exopod proximal segment with 4-5 small, lateral setae; distal 
segment with about 22 elongate, plumose marginal setae. Endopod with large, 
slightly decurved basal apophysis, tip with 3 processes: cannula decurved, tu- 
bular, flanked by broadly rounded, heavily sclerotized caudal process, and sim- 
ilar, but more slender, subterminal lateral process. Pleopod 3 with about 8 plu- 
mose setae along distal margin. Pleopod 4 with 2 false sutures, proximal lateral 
setae present. Pleopod 5 with poorly defined transverse suture. Uropods spatu- 
late, over 2x length of pleotelson. 

Remarks. — The report of 2 troglobitic asellids, Caecidotea ozarkana and C. 
macropropoda, from adjacent localities 5 miles south of Kansas, Oklahoma by 
Chase and Blair (1937) presented an unusual zoogeographic situation. Although 
not unprecedented, the syntopy (or near syntopy, as is the case here) of 2 trog- 
lobitic Caecidotea has been reported in only a few instances (Culver 1976; Lewis 
and Lewis 1981). Examination of specimens collected by A. P. Blair from the 
type localities of the 2 species confirmed the suspicion that C. ozarkana and C. 
macropropoda are conspecific. As C. ozarkana was apparently described from 
immature specimens taken from an unspecified spring, C. macropropoda is pro- 
posed here as the senior synonym. 

Relationships. — The general morphological affinities of Caecidotea macropro- 
poda appear to lie with the members of the Hobbsi Group. The strongly armed 
gnathopod and the shape and setation of the first pleopod are typical of other 
members of the group, but the structure of the second pleopod endopod tip is 
somewhat unusual. The cannula is recurved and appears to twist somewhat on 
its axis, in contrast to the usual low, conical cannula found in the Hobbsi Group. 
However, the basic shape of the cannula is conical and is obscured by other tip 
processes, as would be expected. The placement of Caecidotea macropropoda 
in the Hobbsi Group should be considered tentative. 

Caecidotea macropropoda is most similar morphologically to C. stiladactyla, 
which occurs in caves and springs in the part of Arkansas adjacent to the localities 
of C macropropoda. In both species the cannula appears movable, but in Cae- 
cidotea stiladactyla the cannula apparently extends and retracts from the body 
of the endopod. For comparison with C. macropropoda, the endopod tip of C. 
stiladactyla is illustrated in the retracted position (Fig. 3). Caecidotea macropro- 
poda and C. stiladactyla may be easily separated by the morphology of the fourth 
pleopod. In C. macropropoda, 2 false sutures are present, while in C. stiladactyla 
only a single, sigmoid suture is present. 

Distribution and habitat. — Dearolf (1953) reported Asellus macropropodus 
from 3 additional Oklahoma caves, besides the type-localities of C. ozarkana and 
C. macropropoda, plus a cave and an adjacent pool in Arkansas. These localities 



VOLUME 95, NUMBER 2 345 

remain unconfirmed. Besides the cave region of northeastern Oklahoma, which 
consists of an extension of the Springfield Plain section of the Ozark Plateau into 
the corner of the state, collections from the adjacent parts of Arkansas, Missouri, 
and Kansas have also been examined in search of new localities for C. macro- 
propoda. However, Caecidotea macropropoda still remains known only from the 
type-locality and the immediate vicinity. 

The type-locality of Caecidotea macropropoda, Bat Cave, is now also known 
as Christian School Study Cave. This cave was visited in June 1981 and found 
to be inhabited by a large population of isopods in a guano bog a short distance 
inside the lower entrance. The cave is also inhabited by a colony of Gray Bats, 
Myotis griscesens, which contribute the guano enrichment to the stream. The 
type-locality of C. ozarkana was not specified by Chase and Blair (1937), but 
several small springs and spring caves exist in the same valley as Bat Cave. 
Caecidotea macropropoda was also found at 2 of these small caves, along with 
numerous planarians, Dendrocoelopsis americana (Kenk 1973). Presumably, one 
of these small springs was the type-locality of C. ozarkana. 

Black (1971) presented some water quality data for Bat Cave. In comparison 
with other Oklahoma caves surveyed. Bat (Christian School Study) Cave had 
particularly high total dissolved solid (209 ppm), nitrate (17.0 ppm) and ortho- 
phosphate (4.2 ppm) levels, probably reflecting the guano enrichment of the 
stream. 

Acknowledgments 

I would like to thank Murray and Mary Looney for providing information on 
Oklahoma caves, field support, and other assistance during a collecting trip 
through the Ozarks. Thomas E. Bowman read the manuscript and provided the 
loan of specimens from the collections of the Smithsonian Institution. Travel 
funds were provided by a grant from the Graduate School of the University of 
Louisville. Finally, I thank Teresa M. Lewis for her assistance in making nu- 
merous collections in the Ozarks and Ouachitas. 

This paper is contribution number 202 (New Series) from the Department of 
Biology, University of Louisville. 

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Black, J. H. 1971. The cave life of Oklahoma.— Oklahoma Underground 4(1 & 2):2-56. 

Bresson, J. 1955. Aselles de sources et de grottes d'Eurasie et d'Amerique du Nord. — Archives de 

Zoologie et Generale 92(2): 45-77. 
Chase, H. D., and A. P. Blair. 1937. Two new blind isopods from northeastern Oklahoma. — American 

Midland Naturalist 18:220-224. 
Culver, D. C. 1976, The evolution of aquatic cave communities. — American Naturalist 110:945-957. 
Dearolf, K. 1953. The invertebrates of 75 caves in the United States. — Pennsylvania Academy of 

Science 27:225-241. 
Fleming, L. E. 1973. The evolution of the eastern North American isopods of the genus Asellus 

(Crustacea: Asellidae). — International Journal of Speleology 5:283-310. 
Henry, J. -P., and G. Magniez. 1970. Contribution a la systematique des Asellides (Crustacea: Isop- 

oda). — Annales de Speleologie 25(2):335-367. 
Hubricht, L. 1950. The invertebrate fauna of Ozark caves. — Bulletin of the National Speleological 

Society 12:16-17. 
Kenk, R. 1973. Freshwater triclads (Turbellaria) of North America, VI: The genus Dendrocoelop- 
sis. — Smithsonian Contributions to Zoology 135:1-16. 



346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Lewis, J. J. 1980. A comparison of Pseudobaicalasellus and Caecidotea, with a description of Cae- 
cidotea bowmani, n. sp. (Crustacea: Isopoda: Asellidae). — Proceedings of the Biological So- 
ciety of Washington 93(2):3 14-326. 

1981. Caecidotea salemensis and C. fastis, new subterranean asellids from the Salem Plateau 
(Crustacea: Isopoda: Asellidae). — Proceedings of the Biological Society of Washington 
94(2): 579-590. 

, and T. E. Bowman. 1981. The subterranean Caecidotea of Illinois (Crustacea: Isopoda: 

Asellidae). — Smithsonian Contributions to Zoology 335:1-66. 

, and T. M. Lewis. 1980. The distribution and ecology of two species of subterranean Cae- 
cidotea in Mammoth Cave National Park. — Cave Research Foundation Annual Report 
1980:23-27. 

Mackin, J. G. 1940. A key to the Oklahoma species of the family AseUidae. — Proceedings of the 
Oklahoma Academy of Science 20:17-18. 

. 1959. Malacostraca (Isopoda). Pp. 869-901 in Edmondson, W. T., ed., Freshwater Biology, 

2nd Ed. — John Wiley and Sons, Inc., New York. 

, and L. Hubricht. 1940. Descriptions of seven new species of Caecidotea (Isopoda: Asellidae) 

from the central United States. — Transactions of the American Microscopical Society 59:383- 
397. 

Nicholas, G. 1950. Checklist of macroscopic troglobitic organisms of the United States. — American 
Midland NaturaHst 64(1): 123-160. 

Steeves, H. R. III. 1963. The troglobitic asellids of the United States: the Stygius Group. — American 
Midland Naturalist 69(2): 470-481. 

. 1964. The troglobitic asellids of the United States: the Hobbsi Group. — American Midland 

NaturaHst 71(2):445-451. 

. 1966. Evolutionary aspects of the troglobitic asellids of the United States: the Hobbsi, Stygius 

and Cannulus Groups. — American Midland Naturalist 75(2): 392-403. 

Williams, W. D. 1970. A revision of North American epigean species of As ellus (Crustacea: Isopo- 
da). — Smithsonian Contributions to Zoology 49:1-80. 

Department of Biology, University of Louisville, Louisville, Kentucky 40292. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 347-351 

GONODACTYLUS INSULARIS, A NEW STOMATOPOD 

CRUSTACEAN FROM ENEWETAK ATOLL, 

PACIFIC OCEAN 

Raymond B. Manning and Marjorie L. Reaka 

Abstract. — The ninth species of the G. falcatus complex in the Indo-West 
Pacific region is recognized. Like other species of the complex, it can be distin- 
guished in the field by its color pattern. 



Until relatively recently, Gonodactylus falcatus (Forskal, 1775) was considered 
to be one of the two most widely distributed members of the genus in shallow 
water habitats in the Indo-West Pacific region, ranging from the Red Sea and 
East Africa eastward to Hawaii and Japan. Field studies conducted by R. L. 
Caldwell, University of California, H. Dingle, University of Iowa, and their col- 
leagues, including one of us (MLR), strongly suggested that G. falcatus actually 
was a complex of species sharing many morphological features as Well as habitat 
preferences, but differing in size, behavior, and color in life. Reexamination of 
type-specimens of taxa considered to be junior synonyms of G. falcatus resulted 
in the recognition of five species (Manning 1978), and two species have been 
described since then (Manning and Reaka 1981a, b). The other member of the 
complex, G. graphurus Miers, 1875, from Australasian localities, is distinctive 
morphologically (Manning 1978: fig. 2) and has not been confused with other mem- 
bers of the complex in the recent literature. 

We name here the ninth species of the complex, based upon specimens col- 
lected at Enewetak Atoll. All specimens have been deposited in the collections 
of the National Museum of Natural History, Washington, under USNM catalogue 
numbers. 

We thank C. W. Hart, Jr., Smithsonian Institution, and Austin B. Williams, 
Systematics Laboratory, National Marine Fisheries Service, for their comments 
on the manuscript. Philip Helfrich facilitated field work by one of us (MLR) at 
the Enewetak Marine Biological Laboratory in 1972 with a grant from that insti- 
tution. 

Gonodactylus insularis, new species 
Fig. 1 

Gonodactylus falcatus. — Manning, 1971:73, 74. — Reaka, 1975:56; 1976:484 
[discussion]; 1979:238, 248, 249, 252, figs. 2-4.— Dingle et al., 1973:55, 58, 59, 
62, table 2. 

Material. — Kidrenen [Keith or Grinem] Island, Enewetak, ir22'50"N, 
162°10'30"E, lagoon side, scattered small coral heads on bottom of coral sand at 
75 feet, C. A. Child, sta. 7-69, 23 September 1969: 1 female (holotype, USNM 
135632).— Enewetak, 1971(7), coral rubble, 0-1 meters, R. L. Caldwell: 2 females 
(paratypes, USNM 139883). — Enewetak, Aomon (Sally) Island, coral rubble, 1- 



348 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Gonodactylus insularis, female paratype, total length 31 mm: a, Rostral plate and ocular 
scales; b, Sixth abdominal somite and telson; c, Uropod, ventral view. 



3 meters, 1972, M. L. Reaka: 3 males, 10 females (paratypes, USNM 189383). — 
Sand Island, Enewetak, R. L. Caldwell et al., August 1971: 6 males, 5 females 
(USNM 189384). 

Description. — Rostral plate with length and width subequal or slightly longer 
than broad, median spine relatively short, anterior margins of basal part of plate 
at right angles to body line or sloping anteriorly, lateral margins divergent, an- 
terolateral angles acute but rounded. Ocular scales small, breadth of each no 
greater that that of rostral spine at base. Anterior 5 abdominal somites lacking 
transverse grooves, sixth somite with 6 carinae, variously inflated, each usually 
with strong posterior spine; median carinule absent. Small black spot usually 
present on each side of sixth somite between sub median and intermediate carinae. 
Abdominal width/carapace length index ranging from 882 in smaller specimens 
to 740 in larger ones. Telson appearing elongate because of slender marginal 
teeth, length and width subequal or width slightly greater. Dorsal carinae of telson 
slender or moderately inflated, median and accessory medians each usually with 
strong apical spine flanked ventrally by strong excavation. Knob distinctly bi- 
lobed. Three pairs of marginal teeth present, submedians with movable apices; 
apices of intermediate teeth slender, sharp, curved mesially; lateral teeth shorter, 
blunter. Anterior surface of telson with dark spot on each side anterior to each 
anterior submedian carina. Ventral surface of telson with sharp carina on each 
submedian tooth. Uropodal exopod with single line of marginal setae, exopod 
with 9-12, usually 11, graded movable spines on outer margin of proximal seg- 
ment; basal prolongation with one rounded lobe proximally on inner margin of 
outer spine. 

Color. — Body of both sexes similar, greenish or brownish, heavily speckled or 
mottled, lacking red posterior bands on abdominal somites in both sexes. Anten- 
nal scale variable in color, clear, red, yellow, orange, yellow-green, or blue-green, 
often (but not always) with white, blue- white, or blue patches or speckles, usually 
with pink or red edges; two specimens (both females) with base of scale red, and 
tip blue or blue-green. Meral spot yeflow with anterior reddish infusion, not 



VOLUME 95, NUMBER 2 



349 





Fig. 2. IGonodactylus insularis, female, total length 39 mm, Onatoa: a, Rostral plate and ocular 
scales; b, Sixth abdominal somite and telson; c, Uropod, ventral view. 



variable; smaller flanking spots as follows: dorsal posterior spot blue-green and 
black (not variable); dorsal anterior crescent variable, blue-green, brown, or 
clear; anterior spot variable, black, red, or brown; ventral spot green or brown, 
variable. Other maxillipeds variable in color, green, or blue-green, or usually 
yellow or yellow-green with pink front edges. Pereopods variable, sexes essen- 
tially similar, merus variably greenish or yellow or pink with white splotches, 
distal two segments red or pink; some individuals (both male and female) with 
bluish coloration on pereopods. Uropods similar in both sexes, not variable, 
endopod and exopod red or pink with white mottling or spots, setae red. 

Measurements in mm. — Total lengths of males 13 to 30, of females 10 to 31.5. 
Other measurements of female holotype: carapace length 7.3; rostral plate length 
2.4, width 2.4; fifth abdominal somite width 5.5; telson length 4.5, width 4.5. 

Reaka (1979:252, table 7) noted that at Enewetak this species settled at a length 
of 8 mm, was 13 mm long at sexual differentiation, reached reproductive maturity 
at 31 mm, and had a maximum size of 34 mm. 

Remarks. — This new species resembles three of the species of the G.falcatus 
complex that have a bilobed knob on the telson and that lack a median carinule 
on the sixth abdominal somite and for which color in life is known: G. aloha 
Manning and Reaka, 1981, from Hawaii, and G. mutatus Lanchester, 1903, and 
G. siamensis Manning and Reaka, 1981, both from Thailand. The most striking 
differences among these species are their color patterns in life. The diagnostic 
meral spot in G. insularis is basically yellow, as in these three other species, but 
the anterior infusion is reddish whereas it is brown in G. aloha, G. mutatus, and 
G. siamensis. Gonodactylus insularis lacks posterior red bands on any of the 
abdominal somites in either sex; these are also absent in G. mutatus, but are 
present in males of G. aloha and G. siamensis. The uropodal endopod in G. 
insularis is red or pink with white spots and red setae; in G. aloha they are 
pinkish-orange with yellow distally and green or pink setae. The uropod color has 
not been reported in G. mutatus. 



350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Gonodactylus insularis differs from both G. mutatus and G. siamensis in having 
narrower carinae with relatively long posterior spines on the median and acces- 
sory carinae on the telson, and much sharper, more slender intermediate marginal 
teeth on the telson. These features of the telson are so distinctive that we tentatively 
identify with this species two females, 27.5 and 39 mm long, from Onotoa, Gilbert 
Islands (Fig. 2). Gonodactylus insularis may well prove to frequent other oceanic 
islands in the Pacific. 

Nothing is known of the color pattern of G. glabrous Brooks, 1886, from the 
Philippines and Indonesia, but it is a larger species, 50 to 51 mm long, and the 
teeth and spines of the telson are all slender and sharp (Manning 1978:fig. 3). 

The abdominal width/carapace length indices of G. insularis, like those of G. 
siamensis, are somewhat variable, as follows: 



Carapace 
length, mm 


No. of 
specimens 


Enewetak 
(all but Sand Island) 

Index 


Sand Island 
Index 




Range 


Mean 


Range 




Mean 


3 


3 


794-824 


814 


882 




882 


4 


2 


800-806 


803 


821-875 




848 


5 


2 


800-820 


810 


784-849 




817 


6 


3 


764-797 


783 


— 




— 


7 


1 


753 


753 


821 




821 



It seems likely from the abdominal width/carapace length indices given above 
that the specimens from Sand Island represent a separate species. Samples from 
Sand Island, an apparently temporary emergent island between Enewetak (Fred) 
and Parry islands, on the southeastern quadrant of the atoll, were collected by 
R. L. Caldwell and colleagues in 1971. When researchers returned to Enewetak 
in 1972, Sand Island and its population of Gonodactylus no longer existed. 

In the 1971-1972 Annual Report of the Eniwetok Marine Biological Laboratory, 
R. L. Caldwell and H. Dingle reported that they found three species of gonodac- 
tyloids on Sand Island, one of which was referred to as Gonodactylus sp. Dingle 
et al. (1973:58) reported on their collections from 1971, and noted: ''G. falcatus, 
on the other hand, when we did find it, occupied rubble that was conspicuous for 
the attached overgrowing clumps of green algae." They also noted that coral 
rubble fragments with attached green algae were uncommon on Sand Island, 
possibly a reflection of the scarcity of the species they identified with G.falcatus. 

Without information on color in life of the population from Sand Island, we are 
reluctant to characterize it as a species distinct from G. insularis proper. In view 
of this, we tentatively identify the sample from Sand Island with G. insularis, but 
do not consider those specimens as paratypes. 

The holotype, the largest specimen available, appears to be conspecific with 
the remainder of the type-series. However, unlike the remainder of the speci- 
mens, which were collected in relatively shallow water, the holotype was col- 
lected at a depth of 75 feet. Whether the species actually occurs from the shallow 
intertidal to that depth or whether the holotype washed to that depth is unknown. 

Etymology. — The specific epithet is from the Latin, "insularis", of an island. 

Literature Cited 

Dingle, Hugh, R. C. Highsmith, K. E. Evans, and Roy L. Caldwell. 1973. Interspecific aggressive 
behavior in tropical reef stomatopods and its possible ecological significance. — Oecologia 
13:55-64. 



VOLUME 95, NUMBER 2 351 

Manning, Raymond B. 1971. Two new species of Gonodactylus (Crustacea, Stomatopoda), from 
Eniwetok Atoll, Pacific Ocean. — Proceedings of the Biological Society of Washington 84:73- 
80, figures 1,2. 

. 1978. Notes on some species of the Falcatus Group of Gonodactylus (Crustacea: Stomato- 
poda: Gonodactylidae). — Smithsonian Contributions to Zoology 258:1-15, figures 1-13. 

, and Marjorie L. Reaka. 1981a. Gonodactylus aloha, a new stomatopod crustacean from the 

Hawaiian Islands. — Journal of Crustacean Biology 1(2): 190-200, figures 1-3. 

, and . 1981b. Gonodactylus siamensis, a new stomatopod crustacean from Thailand. — 

Proceedings of the Biological Society of Washington 94(2): 479-482, figure 1. 

Reaka, Marjorie L. 1975. Molting in stomatopod crustaceans, I. Stages of the molt cycle, setagenesis, 
and morphology. — Journal of Morphology 146(l):55-80, figures 1-25. 

. 1976. Lunar and tidal periodicity of molting reproduction in stomatopod Crustacea: a selfish 

herd hypothesis. — Biological Bulletin 150:468-490, figure 1. 

. 1979. The evolutionary ecology of life history patterns in stomatopod Crustacea. Pp. 235- 

260 in S. E. Stancyk (editor). Reproductive ecology of marine invertebrates. — University of 
South Carolina Press: Columbia. 

(RBM) Department of Invertebrate Zoology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, D.C. 20560; (MLR) Department of 
Zoology, University of Maryland, College Park, Maryland 20742. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 352-353 

RISSOIDES, A NEW GENUS OF STOMATOPOD 

CRUSTACEAN FROM THE EAST ATLANTIC 

AND SOUTH AFRICA 

Raymond B. Manning and Ch. Lewinsohn 

Abstract. — A new genus is recognized for 5 East Atlantic species formerly 
assigned to Meiosquilla. The latter genus is restricted to 8 American species. 



As pointed out several times in recent studies of Atlanto-East Pacific stomato- 
pods (Manning 1969:102; 1975:365; 1977:117), the 5 East Atlantic and South Af- 
rican species assigned to the genus Meiosquilla Manning, 1968, differ from all 
American species in having 5 rather than 4 spines on the dactylus of the raptorial 
claw and in lacking elongate spines on the inner margin of the basal prolongation 
of the uropod. 

One of us (Manning 1962:507; 1969:103) suggested that some of the features 
exhibited by species of Meiosquilla, including the unarmed anterolateral angles 
of the carapace, the reduced carination of the carapace, thorax, and abdomen, 
and the movable apices of the submedian teeth of the telson, are all characteristic 
of squillid postlarvae. Thus, Meiosquilla sensu lato may have had a neotenic 
origin. That the American and East Atlantic/South African species differ in 2 
major features, number of teeth on the claw and ornamentation of the basal 
prolongation of the uropod, suggests that these groups of species were derived 
independently from different stocks and that they should not be assigned to the 
same genus. The type-species of Meiosquilla is the West Atlantic Squilla quad- 
ridens Bigelow, 1893, by original designation by Manning (1968:127). We recog- 
nize here a new genus for the 5 East Atlantic and South African species formerly 
assigned to Meiosquilla, which is here restricted to the 8 American species now 
assigned to it (Manning 1969, 1972). 

We thank C. W. Hart, Jr., Smithsonian Institution, and David K. Camp, Florida 
Department of Natural Resources, for their comments on the manuscript. 

Rissoides, new genus 

Definition. — Body smooth, compact, size small to moderate, total length to 8.5 
cm. Eye large, cornea bilobed, noticeably broader than stalk; ocular scales sep- 
arate. Carapace smooth, narrowed anteriorly, anterolateral angles unarmed; ca- 
rinae reduced, median and intermediates absent, at most reflected marginals and 
posterior part of each lateral carina present; cervical groove indistinct; posterior 
median margin evenly concave, posterolateral margins broadly rounded. Exposed 
thoracic somites with, at most, intermediate carinae, submedians absent; lateral 
process of fifth thoracic somite an inconspicuous diagonal or flattened lobe, a 
ventral spine present on each side; lateral processes of next 2 somites evenly 
rounded, not bilobed; ventral keel of eighth thoracic somite well developed. 4 
epipods present. Mandibular palp absent. Dactylus of raptorial claw with 5 teeth, 
outer margin evenly convex; propodus with outer part of upper (opposable) mar- 



VOLUME 95, NUMBER 2 353 

gin evenly pectinate, with 3 proximal movable teeth, middle smallest; dorsal ridge 
of carpus indistinct; ischiomeral articulation terminal. Endopods of walking legs 
slender, elongate. Abdomen with anterior 5 somites lacking submedian carinae; 
intermediates, laterals, and marginals present, usually unarmed anterior to fifth 
somite; sixth abdominal somite with armed submedian, intermediate, and lateral 
carinae. Telson broad, median carina present, supplemental dorsal carinae ab- 
sent; 3 pairs of marginal teeth present, sub medians with movable apices; pre- 
lateral lobes absent; postanal keel, if present, low. Basal prolongation of uropod 
with inner spine the longer, lacking elongate spines but with low tubercles or 
short denticles on inner margin; low, rounded lobe present on outer margin of 
inner spine. 

Type-species. — Squilla desmaresti Risso, 1816. 

Etymology. — We consider it appropriate to dedicate this genus to A. Risso, the 
nineteenth century carcinologist who named the type-species. The gender is mas- 
culine. 

Included species. — Five: Rissoides desmaresti (Risso, 1816); /?. pallidus (Gies- 
brecht, 1910); /?. af He anus (Manning, 1974); R. calypso (Manning, 1974); and R. 
barnardi (Manning, 1975). Original citations for all of these species are given in 
Manning (1977). 

Literature Cited 

Manning, Raymond B. 1962. Alima hyalina Leach, the pelagic larva of the stomatopod crustacean 
Squilla alba Bigelow. — Bulletin of Marine Science of the Gulf and Caribbean 12(3): 496-507, 
figures \-A. 

. 1968. A revision of the family Squillidae (Crustacea, Stomatopoda), with the description of 

eight new genera. — Bulletin of Marine Science 18(1): 105-142, figures 1-10. 

. 1969. Stomatopod Crustacea of the western Atlantic. — Studies in Tropical Oceanography, 

Miami 8:vii + 380, figures 1-91. 

. 1972. Stomatopod Crustacea. Eastern Pacific Expeditions of the New York Zoological So- 
ciety. — Zoologica, New York 56:95-113, figures 1-3. 

. 1975. A new species of Meiosquilla (Crustacea, Stomatopoda) from South Africa. — Annals 

of the South African Museum 67(9): 363-366, figure 1. 

. 1977. A monograph of the West African stomatopod Crustacea. — Atlantide Report 12:25- 

181, figures 1-57. 

(RBM) Department of Invertebrate Zoology (Crustacea), National Museum of 
Natural History, Smithsonian Institution, Washington, D.C. 20560; (ChL) De- 
partment of Zoology, Tel- Aviv University, Ramat-Aviv, Tel- Aviv, Israel. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 354-357 

NEW RECORDS OF PINNOTHERID CRABS 

FROM THE GULF OF CALIFORNIA 

(BRACHYURA: PINNOTHERIDAE) 

Mary K. Wicksten 

Abstract. — Pinnotheres margarita is illustrated for the first time. The range is 
extended northward in the Gulf of California. Pinnixa valerii is reported for the 
first time from western Mexico. Illustrations are given of P. valerii and the closely 
related species P. richardsoni. 



Pinnotherid crabs are commensals with other invertebrates: annelids, echiu- 
roids, mollusks, echinoderms, and ascidians. The species of the western coast of 
Mexico are poorly known, often reported from single specimens of a given species 
collected incidentally with other invertebrates. 

Recent collecting by Alex Kerstitch, Ernest Iverson, and Michel Hendrickx 
has resulted in the finding of many pinnotherid crabs. Among the specimens are 
two species not recorded for more than 45 years — Pinnotheres margarita and 
Pinnixa valerii. The new specimens have been deposited at the Allan Hancock 
Foundation, University of Southern California; and the Estacion Mazatlan. 

Pinnotheres margarita Smith 
Fig. 1 

Pinnotheres margarita Smith, 1869:245.— Smith, 1870:166.— Holmes, 1894:564.— 
Rathbun, 1918:91-93.— Glassell, 1934:301.— Schmitt, McCain, and Davidson, 
1973:56-57. 

Previous records. — Bay of Panama (type-locality), in pearl oyster (Pinctada 
mazatlanica (Hanley), as Margaritiphora fimbriata). — La Paz, Muleje Bay (Gulf 
of California, Mexico) (Rathbun 1918). 

Material examined. — Guaymas, Sonora (27°54'N, 110°53'W), 10 m, rocky bot- 
tom, commensal in Pinctada mazatlanica, 28 June 1981, A. Kerstitch, 1 female. — 
Punta Chivato, Baja California (27°08'N, lir54'W), 20 m among rock, sand and 
algae, commensal in P. mazatlanica, 28 June 1980, A. Kerstitch, female, oviger- 
ous.— SE side Bahia Concepcion, Baja California (26°43'N, 111°53'W), 18 Aug. 
1980, E. Iverson, taken while snorkeling, female. — Isla Carmen, Gulf of Califor- 
nia (25°58'N, liriO'W), 25 m, among rubble and small rocks, 10 July 1980, A. 
Kerstitch, female. — "Panama," 1866, F. H. Bradley, female. 

Measurements in millimeters. — Carapace widths 11.5, 14.1, 10.7, 8.5, and 3.3 
respectively; carapace lengths (in same order) 10.6, 12.3, 10.6, 8.2, and 3.1. 

Remarks. — Pinnotheres margarita is one of the largest pinnotherid crabs in the 
Gulf of California. However, the lack of illustrations has made identification of 
the species difficult. The holotype, reported to have been deposited at the Pea- 
body Museum of Natural History of Yale University, could not be located there 
or at the National Museum of Natural History, Smithsonian Institution. 



VOLUME 95, NUMBER 2 



355 





Fig. 1. Pinnotheres margarita, female from Punta Chivato, Baja California: a, Dorsal view; b, 
Third maxilliped; c, Frontal region; d, Chela; e, Abdomen. 



A small female, questionably identified as P. margarita, was found at the Pea- 
body Museum. I compared this juvenile with the females recently collected. The 
small crab has much the same shape of carapace, proportions of chelae and 
walking legs, and outline of third maxilliped as the larger females. It also agrees 
with the descriptions of Smith (1869, 1870) and Rathbun (1918). 

Although type material and previous illustrations are lacking, I believe that the 
females from the Gulf of California are indeed P. margarita. Smith and Rathbun 
both mentioned the inequality in lengths of the walking legs on the right and left 
sides in mature females. As described, the females are dull brown ("Hke a uniform 
coat of mud"), with short pubescence. The carapace is uneven, with a protuber- 
ant cardiac region and marked sutures. Reported size of the holotype (11.8 mm 
in length of the carapace, 13.4 mm wide) agrees with measurements for the series 
of crabs from the recent collections. Finally, the host {Pinctada mazatlanica) is 
the same. All of the crabs agree with the previous descriptions. 



356 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 2. a, Pinnixa valerii, male from Estero El Verde, Sinaloa; b, Pinnixa richardsoni, male 
holotype from Balboa, Canal Zone, Panama. 



Pinnixa valerii Rathbun 
Fig. 2a 

Pinnixa valerii Rathbun, 1931:262-263, figs. 1-2. — Schmitt, McCain, and David- 
son, 1973:124. 

Previous record. — Isla San Lucas, west coast of Costa Rica (type-locality), 15 
Jan. 1930, 1 male holotype, 1 female paratype (Rathbun 1931). 

Material examined.— Estero El Verde, Sinaloa (23°25'30"N, 106°33'30"W), 1 
m, among Ruppia sp., dredged, 11 Dec. 1979, M. E. Hendrickx and party. 

Measurements in millimeters. — Carapace width 9.3, carapace length 5.8 (spec- 
imen from Allan Hancock Foundation). 

Comments. — Pinnixa valerii is related closely to Pinnixa richardsoni Glassell, 
1936 (type-locality Balboa, Canal Zone, Panama). Both have laterally compressed 
chelae with thick tufts of setae along the ridges of the palm and fingers. The 
dactyls of the first pereopods appear twisted in dorsal aspect. Pinnixa valerii has 
been illustrated previously only by photographs (Rathbun 1931, figs. 1 and 2). 
Only the third maxilliped of P. richardsoni has been figured (Glassell 1936, pi. 
21, fig. 3). 



VOLUME 95, NUMBER 2 357 

I compared the types of the two species (P. valerii from the U.S. National 
Museum of Natural History and P. richardsoni from the San Diego Museum of 
Natural History). In P. richardsoni, the merus of the third pereopod is 1.9x as 
long as wide; in P. valerii, it is 2.7 x as long as wide. The carpus of the third 
pereopod in P. richardsoni bears a tubercle, absent in P. valerii. The first three 
segments of the abdomen of P. richardsoni are fused, not articulated as in P. 
valerii. In general, the legs of P. richardsoni (Fig. 2b) are much stouter than those 
of P. valerii. 

Some of the features given by Glassell in distinguishing between the two species 
vary from animal to animal. Tomentum occurs on the outer surface of the hands 
and carpus of the chelipeds of both species, differing merely in degree instead of 
presence or absence. The outer distal margin of the third maxilliped appears 
somewhat arched in two specimens of P. valerii, not angular. Both species have 
the same twisted shape of the propodus and dactyl of the first pereopods. 

So far, hosts for neither of these species have been recorded. Perhaps host 
specificity will help in identification of these closely related species. 

Pinnixa valerii was collected in an estuary at a salinity of 22%o. Future collec- 
tors might seek this species in back bays, swamps, or estuaries. 

Acknowledgments 

I thank Michel Hendrickx, Estacion Mazatlan, for collecting specimens and 
contributing toward preparation of the illustrations. Alex Kerstitch, University 
of Arizona; Ernest Iverson, University of Southern California; and James 
McLean, Los Angeles County Museum of Natural History, provided specimens 
and information on hosts. John Garth, University of Southern California, gave 
helpful criticism of the manuscript. The illustrations were prepared by Matilde 
Mendez (Fig. 1), Frances Runyan (Fig. 2a), and Paula Walker (Fig. 2b). This 
work was sponsored in part by a grant from Texas A&M University. 

Literature Cited 

Glassell, S. A. 1934. Affinities of the brachyuran fauna of the Gulf of California. — Journal of the 

Washington Academy of Sciences 24:296-302. 
. 1936. New porcellanids and pinnotherids from tropical North American waters. — Transac- 
tions of the San Diego Society of Natural History 8:277-304. 
Holmes, S. J. 1894. Notes on west American Crustacea. — Proceedings of the California Academy of 

Sciences 4, series 2:563-588. 
Rathbun, M. J. 1918. The grapsoid crabs of America. — Bulletin of the United States National Museum 

97:1-461. 
. 1931. A new species of pinnotherid crab from Costa Rica. — Journal of the Washington 

Academy of Sciences 21:262-263. 
Schmitt, W. L., J. C. McCain, and E. S. Davidson. 1973. Decapoda I, Brachyura I. Fam. Pinno- 

theridae. In Gruner, H.-E., and L. B. Holthuis, eds. Crustaceorum Catalogus. Den Haag, W. 

Junk B.V. 160 pp. 
Smith, S. I. 1869. Pinnotheres margarita Smith, sp. nov. In Verrill, A. E. On the parasitic habits of 

Crustacea. — American Naturalist 3:245. 
. 1870. Notes on American Crustacea. No. 1. Ocypodoidea. — Transactions of the Connecticut 

Academy of Sciences 2:113-176. 

Department of Biology, Texas A&M University, College Station, Texas 77843. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 358-370 

STUDIES OF NEOTROPICAL CADDISFLIES, XXXII: 

THE IMMATURE STAGES OF MACRONEMA VARIIPENNE 

FLINT & BUENO, WITH THE DIVISION OF MACRONEMA 

BY THE RESURRECTION OF MACROSTEMUM 

(TRICHOPTERA: HYDROPSYCHIDAE) 

Oliver S. Flint, Jr. and Joaquin Bueno-Soria 

Abstract. — The larva and pupa of Macronema variipenne are described and 
figured, and notes on their natural history given. A neotype is established for M. 
agnathum Miiller and it and Leptonema apicale Navas are synonymized with M. 
bicolor Ulmer, and M. pullatum Navas is synonymized with M. trigramma Na- 
vas. After considering all characteristics of variipenne and related species, Mac- 
ronema is divided into two genera: Macronema Pictet, with M. lineatum Pictet 
as type, is exclusively Neotropical; and Macrostemum Kolenati, with Macro- 
nema hyalinum Pictet as type, contains not only Neotropical species, but all the 
species presently assigned to Macronema in the Nearctic, Oriental, Ethiopian 
and Australasian Regions. 



The genus Macronema, as it has been recognized, is a large genus containing 
somewhat over 100 described species. The genus is most diverse in the tropics 
of the New World, Australia, Asia, and Africa, with a few species in North 
America and northeastern Asia. It is notably absent from Europe and most of 
northern Asia. 

Although many adults have been described, very few larvae have been carefully 
associated with their adult stage, and most of these are from the northern hemi- 
sphere. In spite of the lack of associated material, many larvae of the Macrone- 
matinae have been described, often incompletely, and then attributed to some 
genus or species purely on supposition. This has, as one can imagine, produced 
an extremely confused mess. Ulmer (1957, pp. 332-335) made a heroic attempt 
to straighten out this confusion, and, in general, succeeded. 

He united, under the name of Centromacronema auripenne (Ramb.), a series 
of descriptions of a very distinctive larval and pupal type from South America. 
This form was first made known through a series of works by Miiller (1880, 1881, 
1921 and variously referred to as Macronema, Macronema agnathum, Macro- 
nema species, third species, species C, and Macrostemum), and later fuUy sup- 
plemented by the works of Thienemann (1905 as Macronema), Marlier (1964 as 
Centromacronema), Roback (1966 as Hydropsychidae sp. 2), and Botosaneanu 
& Sykora (1973 as Centromacronema) . This larval type has often appeared in our 
collections in small numbers, but under circumstances that never permitted a firm 
association of stages. Finally we realized that these larvae were generally found 
on plants, often in attached, free-floating roots or leaves, or in and around the 
roots of emergent plants. With this information, Bueno was able to bring back to 
the laboratory and rear in an aquarium several adults from a series of these larvae. 
The adults that emerged turned out to be our recently described species, Mac- 



VOLUME 95, NUMBER 2 



359 






3 

Figs. 1^. Macronema bicolor, male: 1, Genitalia, lateral; 2, Same, dorsal; 3, Tip of aedeagus, 
dorsal; 4, Same, lateral. 

ronema variipenne, not a species of Centromacronema which also flies on the 
same stream. 

We take this opportunity to describe and figure the larvae and pupae of vari- 
ipenne, select and figure a male neotype of M. agnathum Miiller, synonymizing 
it and Leptonema apicale Navas with M. bicolor Ulmer, and finally to discuss 
the generic implications of this evidence, restricting Macronema to a group of 
exclusively Neotropical species and resurrecting Macrostemum for another group 
of worldwide distribution. 



Macronema bicolor Ulmer 
Figs. 1-6 

Macronema bicolor Ulmer, 1905a:75. Fischer, 1963:178. Flint, 1966:6. 
Macronema agnathum Miiller, 1921:530. Ulmer, 1957:338. Fischer, 1963:177. 

(New Synonymy) 
Leptonema apicale Navas, 1927:40. (New Synonymy) 

Because the identity of the larvae described by Miiller under the name Mac- 
ronema agnathum has been unknown, and this uncertainty has been in large part 
responsible for the confusion referred to above, we have made a concerted effort 
to locate material from Miiller. We have located some material in the Museum 



360 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



-^■S^-- 


^^^^^^^^''^^^ ^~^- 


1 


y^"^ 


^^^ 


1 


■-.. -^"C--- 


..^..-. .-'^^^Ljm. 


~ 




1 


■ © 




\ 



\ 




/ 



© 



Figs. 5-6. Macronema hicolor: 5, Wings of lectotype; 6, Wings of neotype of M. agnathum. 



VOLUME 95, NUMBER 2 



361 




Figs. 7-9. Habitats: 7, Rio Palma, above La Palma, Veracruz, Mexico; 8, Marginal vegetation; 9, 
Aroid plant removed, in whose roots larvae were found. 



of Comparative Zoology, Harvard University, and the Natural History Museum, 
Vienna. In this latter collection are an adult male and female and two pupal skins 
in excellent condition, all labeled in Miiller's hand "Macronema III," and the 
two adults also "19/8 87." Because the two adults agree with the brief diagnosis, 
and the pupae are in total agreement with the more extensive description and 
figures in Miiller (1921), and are from him personally, we believe this material to 
authentically represent Miiller's concept of a gnat hum and are therefore labelling 
the male "Neotype." 

The neotype is the same species as M. bicolor Ulmer with which it is here 
synonymized. Ulmer (1957, p. 341) also noted (courtesy of Mr. D. E. Kimmins) 
that the British Museum (Natural History) had examples labeled "III" and that 
these too were bicolor, but then decided that III referred to M. chloraemus 
Miiller, not agnathum. It appears that he was confused by the erratic usage of 
letters, numbers, and names by Miiller. 

The type male of Leptonema apicale Navas located in the Deutsches Ento- 
mologisches Institut, Akademie der Landwirtschaftswissenschaften, was bor- 
rowed (through the kindness of Dr. G. Petersen) and studied. It is a rather faded 
example that matches bicolor in all respects, with which it is herewith synony- 
mized. 

The figures of the male genitalia here presented (Figs. 1-4) were prepared from 
the neotype oi agnathum, as is the photograph of the wings (Fig. 6). The neotype 



362 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 10-12. Macronema variipenne, larva: 10, Lateral; 11, Head, dorsal; 12, Thorax, dorsal. 

is in excellent, unrubbed condition, whereas the types of bicolor and apicale are 
rubbed and appear rather different in coloration (Fig. 5). 



Macronema variipenne Flint & Bueno 
Figs. 7-24 

Macronema variipenne Flint & Bueno, 1979:528-532. 



VOLUME 95, NUMBER 2 



363 




13 







Figs. 13-19. Macronema variipenne, larva: 13, Maxillolabium and anterior region of genae, ven- 
tral; 14, Mandibles, ventral; 15, Labrum, dorsal; 16, Forepleuron and coxa, lateral; 17, Foreleg, 
lateral, with 2 margin hairs of femur enlarged; 18, Midleg, lateral; 19, Prosternite, ventral. 



This species is widespread through eastern and southern Mexico, south 
throughout Central and South America at least as far as Ecuador. We have taken 
adults commonly at lights at night as well as by net during the day. 

Larva. — Length to 21 mm, width 2.5 mm. Sclerites mostly golden-yellow with 
darker brown markings. 

Head: As wide as long. Color brown dorsomesally, golden-yellow laterally, 
posteriorly, and ventrally but becoming darkened ventromesally; frontoclypeus 
with a pair of pale spots laterally. Frontoclypeus with anterior margin concave, 
with prominent knobs anterolaterally; surface with only a few setae laterally. 
Genae with surface smooth, beset with short, bladelike setae in a broad band 



364 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






21 



Figs. 20-23. Macronema variipenne: 20, Larval anal proleg, lateral; 21, Pupal head, antero ventral; 
22, Pupal hook plates, with segment number and anterior or posterior positions indicated, plates 5A 
to 8A shown in dorsal aspect to left, in lateral aspect to right; 23, Apex of pupal abdomen, dorsal. 




VOLUME 95, NUMBER 2 



365 




Meso Meta 



1 



8 



Fig. 24. Macronema variipenne, schematic gill diagram of the lateral aspect of the meso- and 
metathorax and first 8 abdominal segments. S = a single long central stalk with numerous lateral 
filaments. D = 2 S-type gills arising from a common base. 



laterally and posteriorly (but not on pale area posteriorly); with scattered long 
setae. Genae without stridulatory grooves ventrally; ventral ecdysal line lacking 
on one side anteriorly, thus anterior ventral apotome is only half delimited. La- 
brum broad, rounded anterolaterally ; anterior half hairy; no anterolateral brushes. 
Mandibles short, broad, lacking mesal teeth and brushes; apicomesal region strik- 
ingly black internally. Submentum with anterior margin convex, with a few setae 
at anterolateral angles. 

Thorax: Pronotum brown on anterior half, yellow on posterior half; meso- and 
metanota golden-yellow with fuscous lateral and posterior margins. Pronotum 
with bladelike setae and long hairs on anterior half, posterior half smooth; meso- 
notum with a few shorter bladelike setae mostly anterolaterally and many long 
and short hairs; metanotum with only long and short hairs. Prosternite almost 
crescentic with an anteromesal projection; with a small, triangular sclerite pos- 
teromesally. Meso- and metasterna unornamented except for a few short setae. 
Foretrochantin tapering to a blunt point, with 1 bladelike seta; femur with a 
ventral row of 8 palmately divided setae; all segments with numerous hairs. Mid- 
and hindlegs very similar in structure: coxa, femur and trochanter very elongate; 
trochanter and segments distad, narrow and terete, with rows of short, peglike 
setae and long and short hairs; tarsus with a single apicodorsal bladelike seta; 
claw almost straight, with basal setae almost as long as claw. No thoracic gills. 

Abdomen: Gills only laterally as shown in Fig. 24. Lateral line lacking. Integ- 
ument with long and short hairs dorsally, smooth ventrally. Sternum 8 with scat- 
tered hairs including 2 pairs of long setae arising from a pair of small, indistinct 
sclerites. Sternum 9 very hairy, bearing a pair of large, pale, hairy sclerites which 
also bear a few short, bladelike setae anteriorly; a small narrow sclerite laterad. 
With 4 long, slender anal gills. Anal prolegs extremely long and slender; ventro- 
basal plate (attached to the abdomen) bearing a pair of longitudinal dark stripes; 
legs very long, slender, and hairy with a thin section of cuticle at midlength at 
which point the legs bend; apical brush reduced to 1 or 2 hairs; claw long, slender, 
curved. 

Pupa. — Length to 10 mm, width 3 mm. 

Head: Labrum semicircular, with rounded basolateral lobes; anterior third 
densely hairy, basolateral angles with 5-6 short setae, basolateral lobes with 5- 
6 setae. Mandibles membranous, triangular, anterior margin broadly truncate. 



366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Face with a few scattered setae; anterior tentorial pits well marked. Basal anten- 
nal segment with a group of setae; antennae very long, curled into several broad 
loops (not wrapped around the apex of the abdomen). 

Thorax: Meso- and metanota with a few scattered setae and a cluster of setae 
on the tegulae of mesonotum. Midlegs with tibiae and tarsi flattened and fringed. 

Abdomen: Segments 3-7 with long, terete lateral processes. Gills laterally be- 
tween processes and ventrally. Hook plates anteriorly on segments 3-8 (that of 
4 very small, those on 5-8 very high); posteriorly on 3 and 4. Segments 3-8 with 
a few pairs of setae posteriorly. Segments 8 and 9 with a dense fringe of black 
hairs laterally. No apical appendages, only a cluster of black hairs. 

Biology. — The larvae were found in the free-floating mats of roots, and in the 
tangled roots and stems of emergent plants. The retreats in the root mats were 
about 30 mm long by 7 mm wide, constructed of fragments of organic matter held 
together rather loosely by silk and all attached to the rootlets. These retreats 
were removed as carefully as possible and were preserved promptly, but the 
larvae generally crawled away quickly. The retreats were later cut open carefully, 
in an attempt to find some sort of capture net. Nothing of any type could be 
found, and the retreats seemed to be only a simple tube. We believe that the 
larvae construct no distinct net, either of the Macronema type or of the Hy- 
dropsychinae type. Perhaps they may construct an irregular mesh of threads 
outside the shelter, but even that seems unlikely. 

Pupal shelters have also been found in the same root mats. These are approx- 
imately the same size as the larval shelters, but are a little larger in diameter 
where the pupa is, and are more firmly constructed. Inside the shelter of silk and 
detritus, the larva constructs a perfect silken cocoon, free of the outer shelter 
except near each end. The ends are simply a loose tangle of threads. None of the 
pupae found in the wild was mature enough to permit species identification. 

In order to determine the species of these larvae, Bueno, in the summer of 
1978 coUected a series of larvae in the Rio Palma (Fig. 7), above the village of La 
Palma. This site is in the tropical wet forest of the coastal lowlands in the state 
of Veracruz, about 25 km east of Catemaco at an elevation of 30 m above sea 
level. The river is no more than 15 m wide with an average depth of about Va m, 
the water is clear, of good chemical and physical condition, and the bottom is 
sandy with much gravel and stone. At the sides there are often backwaters with 
emergent plants, and the marginal trees and vines often put into the water roots 
which greatly proliferate, frequently forming free-floating mats. The large leaves 
of the marginal trees and shrubs form a favored resting place for the adults during 
the day. 

The larvae, mostly in the 4th and 5th instars, were collected from the roots and 
stems of the emergent plants (Figs. 8-9), and then brought back alive to the 
laboratory in Mexico City, a distance of some 550 km. There they were kept 
alive in an aquarium in water with sand, sticks and debris afl from the Rio Palma. 
The aquarium was of about two liters capacity, and air was supplied by an air 
pump which at the same time caused the water to circulate in the aquarium. A 
supply of "Tetramin," a tropical fish food, was circulated with the water every 
day to provide food for the larvae. When most of the water had evaporated, it 
was replaced with water directly from the tap. 

The larvae mostly disappeared into the bottom sand and debris immediately. 



VOLUME 95, NUMBER 2 367 

However, a few constructed their retreats on the glass and could be observed in 
their tunnel. These larvae spent most of their time simply resting in their tube. 
Rarely they would crawl part way out and appear to be, perhaps, feeding on 
debris on the retreat. When the water level became low, the current diminished, 
and they spent much of their time making undulatory movements with the ab- 
domen. After a while these larvae disappeared, and in about 20-30 days adults, 
which were M. variipenne, emerged. 

We have examined the gut contents of several field preserved larvae. The guts 
were filled with rather uniform, roughly cubical, bits of plant tissue about 0.2 mm 
long on a side. 

In summary, it is our belief that the larvae: construct only a rather flimsy, 
tubular larval retreat without any type of capture net; construct the, retreat on 
plant structures in marginal situations or even buried in silt or sand in backwaters; 
that they are very tolerant of low water velocities and oxygen concentrations; 
and that they are "shredders," biting-out pieces of plant material. The pupae are 
sheltered in a slightly enlarged and strengthened part of the larval retreat; the 
pupa is surrounded by a silken cocoon only attached to the outer shelter near the 
ends; and that the ends of the cocoon are not firmly closed, but loosely plugged 
by an irregular tangle of silk. 

Generic Considerations 

Now, with the first unequivocal association of this form of larva with a species 
of the genus Macronema, not Centromacronema as it was believed to be (Ulmer 
1957; Marlier 1964), the entire question of generic relationships is thrown open. 

The descriptions of the immature stages of a number of species of ''Macro- 
nema ' have been in the literature for many years: Carolina Bks. (Ross, 1944), 
zebratum Hag. (Ross, 1944), transversum (Walk.) (Ross, 1944), radiatum McL. 
(Lepneva, 1964), capense Walk. (Scott, in press), ulmeri Bks. (MarHer, 1964, as 
siolii), and others described but not associated with any adult (as listed in Ulmer 
1957, pp. 344-345). All these descriptions are very similar and are summarized 
in Table 1 (under Macrostemum) where the contrasting characteristics of vari- 
ipenne are also given (under Macronema). I do not believe any trichopterist 
would consider that such totally discordant characteristics could belong to dif- 
ferent species in one genus, especially so if they considered the characteristics 
of other hydropsychid genera (as shown in Wiggins 1977, for example). No won- 
der Ulmer (1957, p. 339) concluded that the variipenne type larva could not be 
a Macronema, and assigned it to the closely related genus Centromacronema. 

Unfortunately, we still do not have any larvae definitely correlated with Centro- 
macronema, nor with some of the more distinctive species of Macronema from 
eastern Brazil. Such a lack, however, does not affect the problems within Mac- 
ronema, but does prevent a complete solution incorporating all the closely related 
genera. Recognizing the possibility of the future synonymy of Centromacronema 
with Macronema, and the possibility of having to carve more genera from Mac- 
ronema, we will consider the situation within Macronema, as these aforemen- 
tioned possibilites should not affect the stability of the names in consideration. 

Ulmer was the first to give notice in a publication (1907) of two distinct groups 
of species in Macronema. The first group of exclusively South American species 



368 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table L — Comparisons of certain characters of the larva and pupa of Macronema and Macroste- 
mum. 





Macronema 


Macrostemum 


Larva 






Labrum 


no brushes 


anterolateral brushes 


Mandibles 


short, no teeth 


elongate, toothed 


Gena, dorsally 


unmodified 


with a carina around central 
area 


Gena, ventrally 


no stridulatory grooves 


with stridulatory grooves 


Ventral ecdysal line 


lacking on 1 side of pregula 


separating pregula on both 
sides 


Coxa 


long 


short 


Foretibia and tarsus 


scattered hairs, ventral row of 
palmate setae 


with a large brush of hairs 


Mid- and hind tibiae and 


disproportionately short in 


typical proportions; with 


tarsi 


relation to femur; with rows 


ventral rows of feathered 




of short, pointed setae 


setae 


Thoracic gills 


lacking 


present 


Abdominal gills 


lateral line only 


dorsolateral, lateral, and 
ventrolateral rows 


Anal prolegs 


very long, jointed in middle 


normal, not jointed 


Retreat 


tubular 


branched 


Net 


none 


very regular, of fine mesh 


Food 


shredders, biting pieces from 


collectors, gathering suspended 




plant matter 


fine particulate matter 


Pupa 






Mandibles 


membranous 


sclerotized 


Labrum 


hairy on anterior third 


few scattered setae 


Antennae 


looped only 


coiled around abdomen 


Posterior hook plates 


on segments 3 and 4 


on 3 only 


Apical appendages 


lacking 


present 


Inner cocoon 


free of shelter except at ends 


united to shelter 


Cocoon ends 


loosely plugged 


closed 



(in which he included lineatum, argentilineatum, parvum, fulvum, bicolor, and 
percitans) he characterized by a minute discoidal cell, a very large median cell, 
a distinct, extra, costal crossvein, and the subcostal vein ending on the costa. The 
other group, consisting of all the other species in the genus, possessed a normal 
or large discoidal cell, a smaller median cell, no costal crossveins, and the &ub- 
costa was united to Ri just before the wing margin. However, Miiller had noticed 
as early as 1885 (but not pubHshed until 1921) a division of the genus based on 
the presence or absence of the apical tibial spurs on the foreleg and structure of 
the pupa (as in Table 1). Kolenati (1859) had proposed a new genus Macrostemum 
for M. hyalinum Pict. and M. auripenne Ramb. based on the absence of such 
spurs in the latter two species. M. hyalinum does have a pair of tibial spurs, but 
they are very small and easily overlooked in comparison to the large spurs of 
those species he placed in Macronema {lineatum although included, was appar- 



VOLUME 95, NUMBER 2 369 

ently unknown to him). Ulmer in 1957 selected hyalinum as the type of Macro s- 
temum (but considered it and Macronema synonymous), and did correctly at- 
tribute the species with the two small spurs to it and the ones lacking spurs to 
Macronema. In 1979 we independently recognized the same two groups on the 
basis of coloration and male genitalia. 

Now with the knowledge of the larval and pupal morphology and ecology, it 
is obvious that these differences between the two groups of species in the adult 
stage are much more fundamental. Therefore, we propose to recognize two gen- 
era. Macronema Pictet (1836) has as its type M. lineatum Pictet (monobasic) and 
in addition contains: amazonense Flint, argentilineatum Ulm., bicolor Ulm., 
bifidum Flint, burmeisteri Bks., chalybeoides Ulm., exophthalmum Flint, fragile 
Bks.,fraternum Bks., fulvum Ulm., gundlachi Bks., hageni Bks., immaculatum 
Mos., lachlani Bks., luteipenne Flint & Bueno, matthewsi Flint, muelleri Bks., 
paliferum Flint, parvum Ulm., pennyi Flint, percitans Walk., pertyi Bks., picteli 
Bks., reinburgi Nav., and variipenne Flint & Bueno. All species are limited to 
the Neotropical Realm. 

The salient characteristics distinguishing the adults of this genus from Mac- 
rostemum are: lack of tibial spurs on the forelegs; fore wing with Sc ending on 
the costa; forewing color due mostly to scales, basal % usually being green, 
bounded outwardly by a variably colored area and with costal cell filled with 
silvery scales; male with claspers undivided or nearly so, tenth tergum shorter 
and broader, often with lobes and processes, tip of the aedeagus also often bears 
lobes and processes. 

The larval and pupal characteristics are outlined in Table 1. 

The second genus is Macrostemum Kolenati (1859), with the type Macronema 
hyalinum Pictet (selected by Ulmer 1957). In addition to the type the genus con- 
tains the following species in the New World: arcuatum Erich., braueri Bks., 
Carolina Bks., digramma McL., erichsoni Bks., maculatum Perty, negrense 
Flint, par Nav., ramosa Nav., santaeritae Ulm., subequale Bks., surinamense 
Flint, transversum Walk., trigramma Nav. {pullatum Nav. is a NEW SYN- 
ONYM), triste Nav., tuberosum Ulm., ulmeri Bks., and zebratum Hag. In ad- 
dition, all species listed in the Fischer catalogs (1963, 1972) in Macronema from 
Africa, Asia, and Australia are transferred to Macrostemum. 

The adults differ from Macronema by a combination of the following charac- 
teristics: foretibia with 1 or 2 small apical spurs; Sc of forewing united with R^ 
apically, or ending in a fork whose ventral arm is united to Ri and is the stronger 
of the two arms; forewings with color due primarily to strongly contrasting colors 
of the membrane and the pattern is widespread over the wing; males with claspers 
distinctly 2-segmented, the tenth tergum is elongate and rather simple, and the 
tip of the aedeagus is generally bulbous without special structures. 

The larval and pupal structures of this genus are also outlined in Table 1 . 

Acknowledgments 

We express our appreciation to the Smithsonian Staff artists. Young T. Sohn 
(Figs. 11, 17, 18, 20-23) and L. Michael Druckenbrod (Figs. 10, 12), for their 
excellent illustrations. Drs. Richard C. Froeschner and Wayne N. Mathis pro- 
vided much useful discussion and criticism of the manuscript. We thank Jorge 
Padilla and other students of the U.N. A.M. for their valuable help in the field. 



370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Flint is most grateful to the Director of the Instituto de Biologia, Universidad 
Nacional Autonoma de Mexico, Dr. Jose Sarukhan Kermes, for travel funds and 
to the Secretary of the Smithsonian Institution, Dr. S. Dillon Ripley for Fluid 
Research Funds, which together made possible this work in Mexico. A CONA- 
CYT grant permitted Bueno to complete the study at the Smithsonian Institution. 

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Scott, K. M. F. In prep. On the Hydropsy chidae (Trichoptera) of Southern Africa, with keys to the 

African Genera of imagos, larvae and pupae and species lists. — Annals of the Cape Provincial 

Museums (Natural History). 
Thienemann, A. 1905. Trichopterenstudien. — Zeitschrift fiir wissenschaftliche Insektenbiologie 

1:285-291. 
Ulmer, G. 1905. Zur Kenntnis aussereuropaischer Trichopteren. — Stettiner Entomologische Zeitung 

66:1-119. 
. 1907. Monographic der Macronematinae. — Collections zoologiques du Baron Edm. de Selys 

Longchamps, pt. 2 of Fasc. 6:1-121. 
. 1957. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil III). — Archiv fiir Hydrobiol- 

ogie 23(suppl.): 109-470. 
Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichoptera). — University of 

Toronto Press, Toronto and Buffalo. 401 pp. 

(OSF) Department of Entomolgy, Smithsonian Institution, Washington, D.C. 
20560; (JB-S) Instituto de Biologia, U.N.A.M., Apartado Postal 70-153, Mexico 
20, D.F. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 371-376 

NOTES ON DISTRIBUTION OF SOME LATIN AMERICAN 

COTTON-STAINERS {DYSDERCUS: PYRRHOCORIDAE: 

HEMIPTERA) AND REMARKS ON THE BIOLOGY OF 

DYSDERCUS URBAHNI SCHMIDT 

Joachim Adis and Richard C. Froeschner 

Abstract. — New geographical distribution data are given for 11 Latin American 
species of Dysdercus, with new country and/or state records for 7 of them. Data 
on life history and food sources are given for D. urbahni Schmidt in a riverine 
inundation-forest in the Central Amazon, Brazil. 



Introduction 

Extensive taxonomic and distributional data for Latin American species of 
Dysdercus were given by Ballou (1906), Lima et al. (1962), Doesburg (1968), and 
Beccari and Gerini (1970). Now additional geographic distribution can be given 
based on specimens recently located or deposited in Museu Paraense Emilio 
Goeldi (MPEG), Belem, Brazil; Instituto Nacional de Pesquisas da Amazonia 
(INPA), Manaus, Brazil; and the Smithsonian Institution, National Museum of 
Natural History (NMNH), Washington, D.C., U.S.A. 

The first significant biological data for Dysdercus urbahni Schmidt were gath- 
ered during 1975-1976 ecological studies in a white water, inundation-forest 
("Varzea") near Manaus, Brazil. 

Notes on the Distribution of Some Latin American Dysdercus 

Present new geographic records, some based on identifications by P. H. van 
Doesburg in 1978, mostly complement the distribution data in Doesburg's (1968) 
revision of the New World species oi Dysdercus. 

Dysdercus mimus Group 

1) Dysdercus imitator Blote 

Previous records: Colombia, Venezuela, Ecuador, Peru, Bolivia (Doesburg 
1968). 

Present records: Brazil-Mato Grosso: Aripauna (5°07'S, 60°24'W), Reserva 
Humboldt, III-1977, 1 c?, W. L. Overal; Chapada dos Guimaraes (15°26'S, 
55°45'W), 3-II-1961, 1 9, J. e B. Bechyne Coll. (all in MPEG). 

2) Dysdercus mimus infuscatus Blote 

Previous records: British Guiana, Suriname (Doesburg 1968). 

Present records: Brazil-Amapa: Macapa, Praia do Araxa, 19-X-1978, 1 SIX 9, 
M. F. Torres (MEPG). — Amazonas: Manaus 010 km 232 (^highway Manaus- 
Itacoatiara km 232), 28-VI-77, \ S,B. Mesearenhas (INPA).— Maranhao: Im- 
peratriz (5°32'S, 47°29'W), 13-VM978, 1 c?, F. F. Ramos; Santa Ines, 05-X-1978, 
7c?c?/6 9$,F.F. Ramos/P. Celso. — Mato Grosso: Aripuana, Reserva Humboldt, 
11-1977, 1 d, W. L. Overal.— Para: Araguaia, Redencao, 25-XI-1978, 1 9 , W. 



372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Franca, Belem, Mocambo, 6-V-1977, 1 J; 01-III-1977, 1 c?, P. Waldir; 5-III-1977, 
1 9 , A. Y. Harada. Belem, Mocambo, 19-XII-1977, 2 ? $ , C. Moreira; XII-1977, 
1 ? , C. Moreira; 02-1-1978, 1 9 , S. A. Santos. Belem, Parque do Museu, X-1976, 
2SSII 9 , W. L. Overal. Belem, Utinga, 22-III-1977, 1 9 , P. Tadeu; 22-III-1977, 
1 (^, M. F. Torres; 24-III-1977, 1 cJ, M. F. Torres. Rodovia Belem-Brasilia, km 
90, F. S. Antonio, 14-11-1976, 1 (^/9 9 9, T. Pimentel; 14-11-1976, 3 SSI^ 9 9, 
M. F. Torres; 18-11-1976, 2 c?^, M. F. Torres; 26-11-1976, 1 c?, M. F. Torres. 
Rodovia Belem-Brasilia, km 90, F. Jaboti, 13-11-1976, 2 (?(J/2 9 9, T. Pimentel; 
13-11-1976, 1 9, M. F. Torres. Bragan9a (1°03'S, 46°46'W), 26-IV-1978, 1 cJ, W. 
L. Overal; 26-V-1978, 3 c^c^/4 9 9 , F. F. Ramos/M. F. Torres. Bujaru (1°31'S, 
48°0rW), 03-VIII-1978, 2 cJc?/2 9 9, P. Nolasco; 03-VII-1977, 1 9 , P. Nolasco; 
29-VI-1978, 1 c?, P. Waidir/L. Braak (MPEG). Capitao Poco, 25-11-1978, 2 SSI 
4 9 9, W. Franca/P. Tadeu; 26-11-1978, 1 SI5 9 9 , W. Fran9a/P. Tadeu; 27-11- 
1978, 1 SI2 9 9, W. Fran9a; 28-11-1978, 1 9 , W. Fran9a. Ilha das Oncas 
(^across Belem), 30-1-1977, 1 c^, W. L. Overal. Maraba (5°21'S, 49°07'W), 3- 
VII-1978, 1 (?/l 9, W. Franca. Peixe-Boi (ri2'S, 47°18'W), 24-1-1964, 1 9 , W. 
Franga; 15-IV-1977, 1 9 , P. Waldir; 16-IV-1977, 1 SSI5 9 9,V. Waldir; 12-IV- 
1977, 1 S P. Waldir; 17-IV-1977, 1 9, W. L. Overal; Peixe-Boi, 24-1-1964, 1 S, 
W. Franca; 22-IV-1978, 1 c^, M. F. Torres; 23-IV-1978, 1 c^/l 9, M. F. Torres; 
24-IV-1978, 1 (^, F. F. Ramos; 26-IV-1978, 1 9 , F. F. Ramos; 27-IV-1978, 1 9, 
N. de Souza. S. J. Pirabas, 27-VII-1978, 1 cJ/l 9, M. F. Torres. Santa Isabel 
(r41'S, 50°11'W), 29-1-1964, 1 c^/2 9 9, W. Franca. Tucurui (3°42'S, 49°27'W), 
7-VII-1978, 1 9 , P. Tadeu; 8-VII-1978, 4 9 9 , R. B. Neto; 21-X-1978, 1 c?, R. B. 
Neto; 14-III-1979, 1 S, margem esquerda, W. L. Overal (all in MPEG). 

These first Brazilian records for this subspecies extend its range coastwise and 
far inland up the valley of the Amazon River. 



Dysdercus flavolimbatus Group 

3) Dysdercus ruficeps Perty 

Previous records: Nicaragua; Costa Rica, Panama, Colombia, Ecuador, Peru, 
Bolivia, Brazil: Amazon (6 records, only one with exact locality). Para (4 records, 
one without exact locality), Rio de Janeiro (1 record; comp. Doesburg 1968, 
Beccari & Gerini 1970). 

Present records: Brazil-Para: Ananindeua (1°22'S, 48°23'W), III-1956, 1 9, C. 
Carvalho; Belem, Fazenda Velha, 14-V-1977, 1 9, M. F. Torres; Belem, Utinga, 
22-III-1977, 1 9, P. Tadeu; Benevides, VI-1964, 1 9 , W. Franga; Benevides 
(1°22'S, 48°15'W), VI-1964, 1 9 , W. Franga; Bujaru, Ol-VII-1977, 1 c?, P. Waldir/ 
L. Braack; Mosqueiro (1°10'S, 48°28'W), 17-VII-1978, 3 c^c^/1 9, M. F. Torres; 
Santa Isabel (1°41'S, 50°11'W), 17-XII-1973, 3 c?(?/3 9 9, B. Mascarenhas; 10-11- 
1974, 5 c? (?/2 9 9 , Jose Queiroz; (all in MPEG). 



4) Dysdercus flavolimbatus flavolimbatus Stal 

Previous records: Mexico, Guatemala, Panama: Rio Changena, Boca del Toro, 
2400 ft., 17-IX-1961, 1 ^, G. B. Fairchild (comp. Doesburg 1968). 

Present record: Panama- Chiriqui (Dist. Renacimiento): Santa Clara 4000 ft. 
(8°31'S, 82°39'W), 20-22- V-1977, 1 c?/2 9 9, B. C. Ratcliffe (INPA). 



VOLUME 95, NUMBER 2 373 

5) Dysdercus concinnus pehlkei Schmidt 

Previous records: Costa Rica, Venezuela, Colombia, Panama: La Chorrera, 
23-V-1944, K. E. Frick; Lino, 2 9 (comp. Doesburg 1968). 

Present record: Panama-Colon: Santa Rita Ridge (9°20'N, 79°48' W), 24-V-1977, 
1 (?, B. C. Ratcliffe (INPA). 

Dysdercus maurus group 

6) Dysdercus obscuratus flavipenuis Blote 

Previous records: Panama, Colombia, Ecuador, Venezuela (Doesburg 1968). 
Present record: Brazil-Mato Grosso: Chapada dos Guimaraes (15°26'S, 
55°45'W), 5-IM961, 1 $, J. e B. Bechyne (MPEG). 

7) Dysdercus honestus Blote 

Previous records: Peru, Colombia, Venezuela, Suriname, Paraguay, Bolivia, 
Brazil: Amapa (1 record), Bahia, Espirito Santo, Goias (5 records), Mato Grosso 
(1 record without exact locality), Minas Gerais, Para (3 records), Rio de Janeiro, 
Sta. Catharina (comp. Doesburg 1968). 

Present records: Brazil-Goias: Pirineus, 2-n-1962, 1 9; Paraiso, 9-n-1962, 1 
9, J. e B. Bechyne.— Maranhao: Buriticupu (4°13'S, 46°33'W), 30-IX-1978, 1 
(^, M. F. Torres.— Mato Grosso: Chapada dos Guimaraes (15°26'S, 55°45'W), 19- 
M961, 1 9, J. e B. Bechyne; 21-M961, 1 9, J. e B. Bechyne; 5-IM961, 1 9, J. 
e B. Bechyne (all in MPEG). Rio (=Res.) Humboldt, Estrada do Porto (=Cidade 
Humboldt; 5°07'S, 60°24'W), 1 c?, L. Albuquerque, nr. 0240 (INPA); Rio Hum- 
boldt, Estrada Dir. Aeroporto (-Cidade Humboldt), 26-M976, 1 S, (NMNH); 
Fazenda Ric. Franco, 6-in-1961, 1 cJ, J. e B. Bechyne; 7-m-1961, 1 c^/l 9, J. 
e B. Bechyne; 15-in-1961, 1 9 , J. e B. Bechyne.— Para: Tucurui (2°53'S, 52°0'W) 
14-m-1979, 1 9, W. L. Overal, "margem esquerda" (both in MPEG). 

Observation: The state record for Maranhao is the first record of D. honestus 
in the northeast of Brazil (comp. Doesburg 1968, fig. 174). 

8) Dysdercus ruficollis (Linnaeus) 

Previous records: British Guiana, Suriname, Argentina, Paraguay, Brazil: 
Amazonas, Bahia, Distrito Federal, Espirito Santo, Goias, Guanabara, Minas 
Gerais, Parana, Rio de Janeiro, Rio Grande do Sul, Sao Paulo, Sta. Catharina 
(comp. Doesburg 1968). 

Present records: Brazil-Maranhao: Imperatriz (5°32'S, 47°29'W), 13-VM978, 
7 c?(?/8 9 9, M. F. Torres/F. F. Ramos/R. B. Neto, (MPEG).— Rio de Janeiro: 
Manguinhos, 30-V-1967, 1 612 9 9, L. P. Albuquerque; IV-1967, 1 c^, L. P. 
Albuquerque (INPA); Petropolis, 5-in-1962, 1 9, J. e B. Bechyne (MPEG). 

Observation: The first state record for Maranhao indicates the presence of Z). 
ruficollis in the northeast of Brazil (comp. Doesburg 1968, fig. 210). 

Dysdercus albofasciatus group 

9) Dysdercus goyanus Doesburg 

Previous record (comp. Doesburg 1968): Brazil-Goias: Jatahy (latai). Clermont 
vend. (1 c^, holotype). 

Present record: Brazil-Mato Grosso: Cuiaba (15°35'S, 56°05'W), 22-in-1961, 1 
(^, J. e B. Bechyne (MPEG). 



374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Dysdercus fernaldi group 

10) Dysdercus urbahni Schmidt 

Previous records: Brazil: Amazonas, Para (comp. Doesburg 1968). 

Present records: Brazil- Amapa: Macapa, Pacoval, 07-XI-1978, 1 ^, W. Franca 
(MPEG).— Amazonas: Ilha de Curari (Rio Solimoes), lO-VII-1979, 1 SIX $/6 
nymphs (all in alcohol), J. Adis "em sementes de Pseudobombax munguba caidas 
no solo"; Iranduba, Rio Solimdes, 29-IX-1962, 3 6 6,5. Dellome, no. 2471; Estr. 
AM 1 (=Manaus-Itacoatiara) km 180, 21-X-1965, I 6,F. Antonio; Estr. AM 1 
km 244, 11-1-1977, 1 cJ, B. C. Ratcliffe; Urucury, Codajas, XII-1963, 1 6 , Ed- 
uardo, no. 2552.— Mato Grosso: Aripuana (=Cidade Humboldt; (5°07'S, 
60°24'W), Estrada do Porto, 1 6,L.F. Albuquerque/E. Rufino, no. 0180 (all in 
INPA). 

Observation: The state record for Mato Grosso indicates the presence of D. 
urbahni in the southern Amazon region, (comp. Doesburg 1968, fig. 242). For 
biology notes see p. 9. 

11) Dysdercus fernaldi fuscofasciatus Blote 

Previous records: Venezuela, British Guiana, Suriname, French Guiana, Bra- 
zil: Amazonas, Para (1 record without exact locality), Sao Paulo (comp. Doesburg 
1968). 

Present records: Brazil- Amazonas: Urucury, Codajas, (6°46'S, 64°35'W), XII- 
1963, 5 6 6/59'^, Eduardo, no. 2551 (INPA).— Para: Ilha das Oncas (=across 
Belem), 7-II-1977, I 6,W.h. Overal; 30-11-1977, 1 9, W. L. Overal (MPEG). 

Notes on the Biology of Dysdercus urbahni Schmidt 

The following information comprises the first biological observations recorded 
for Dysdercus urbahni. The species has been collected mainly along rivers (black 
water; e.g. Rio Negro; white water: e.g. Rio Solimoes) and on "terra firme" (e.g. 
Reserva Ducke, Manaus) in the Amazon area. The observations were made dur- 
ing ecological fieldwork in a white water, inundation forest on the Solimoes River 
near Manaus (Adis 1977 and 1979). Here the season can be divided into an 
inundation phase, when the forest may flood to a depth of 4 m, and a non-inun- 
dation phase. The inundation phase occurs March to August when the receding 
waters mark the end of the inundation phase and the beginning of the non-inun- 
dation phase. The latter begins with a dry period that lasts until December when 
the rainy period of the non-inundation phase begins the accumulation of waters 
for the beginning new inundation phase in March. 

Fruits of the tree Pseudobombax munguba were found to be the main source 
of nutrition for D. urbahni during the dry season of the non-inundation phase. 
These trees lose their leaves during the beginning of the inundation phase, and 
by June and July produce flowers and leaf buds, the fruit being formed and 
dropped from the trees between August and October — the end of the inundation 
phase until the beginning of the non-inundation phase. 

Adults of D. urbahni appeared on the forest floor during the first week of the 
non-inundation phase (the dry season); there they mated, usually near fallen 
"Munguba" fruits. The females laid their eggs in chapped fruits or on the soil 
beneath and beside open or still closed fruits. Apparently fruits in the canopy 



VOLUME 95, NUMBER 2 375 

were not attacked. After 5-6 days first instar nymphs could be found; new gen- 
eration adults began to appear after about 6 weeks [compare with the 42-55 days 
reported for D. delauneyi Lethierry by Ballou 1906]. One chapped fruit may 
contain as many as 1000 nymphs of all instar s, while the average was 400 nymphs 
per fruit (=61)! 

Dead adults of the former generation were found near "Munguba" fruits 3^ 
weeks after the forest dried up. At this time, the gregarious nymphs, migrating 
on the forest floor, found newly dropped and already chapped fruits and fed on 
the contained seeds. Field and laboratory experiments found that newly hatched 
nymphs and adults preferred ripe but still soft seeds over dry or hard seeds; and 
if no seeds were available they would feed on dead arthropods. Eight to ten 
weeks after the forest had dried up, nearly all nymphs had transformed to adults 
which frequented herbs, shrubs and young trees of the forest for another 2-3 
weeks but then disappeared until the end of the dry season in November. Within 
the rainy season of the non-inundation phase (December until March), adults 
were seen only sporadically. 

For the period from December to August, which also includes the inundation 
phase, the source of nutrition for these insects is unknown. Pseudobombax mun- 
guba is restricted to white water, inundation-forest ("Varzea"); hence at other 
localities, e.g., along blackwater rivers or on "terra firme" areas, different plants 
or "Munguba" species must provide reproductive sites and food sources for D. 
urbahni. 

Previous publications reported Bombacaceae to be visited by at least nine other 
Dysdercus species (comp. Doesburg 1968, pp. 184-189; Silva et al. 1968 pp. 53- 
56): D. fernaldi fuscofasciatus Blote {Bombax globosum; Suriname), D.fulvon- 
iger (De Geer) {Bombax globosum', Suriname), D. fulvoniger discolor Walker 
{Bombax malabaricum), D. honestus Blote ("Sumaiima," Brazil), D. longirostris 
Stal ("Sumauma," Brazil), D. maurus distant {Bombax globosum; Suriname), 
D. mimus (Say) {Bombax globosum ("Sumauma"); Belem, Brazil). D. mimus 
distant Blote (sucking on seeds of "Sumauma" which had fallen to the floor; 
Para, Brazil), D. ruficollis {Bombax ventricosa) and Dysdercus spp. {Bombax 
discolor). 

Acknowledgments 

We are grateful to W. L. Overal (MPEG, Belem) and Norman Penny (INPA, 
Manaus) for providing us with specimens. The manuscript was reviewed by Nor- 
man Penny and typed by Irmgard Adis, who joined the field excursions, and 
Silver B. West (Smithsonian Institution, Washington). We heartily thank Amy 
Levin (S.I., Washington) for the literature research by computer. 

Literature Cited 

Adis, J. 1977. Programma minimo para analises de ecossistemas: Arthropodos terrestres em florestas 

inundaveis da Amazonia Central. — Acta Amazonica 7(2): 223-229. 
. 1979. Vergleichende okologische Studien an der terrestrischen Arthropodenfauna zentral 

amazonischer Uberschwemmungswalder. — Ph.D. thesis, Univ. Ulm (West Germany), 99 pp. 
Ballou, H. A. 1906. Cotton Stainers.— West Indian Bull. 7:64-85. 
Beccari, F., and V. Gerini. 1970. Catalogo delle Specie appartenenti al genere Dysdercus Boisduval 

(Rhynchota, Pyrrhocoridae). — Revista di Agricolture subtropicale e tropicale 64:20-72. 



376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Doesburg Jr., P. H. van. 1968. A revision of the New World species oi Dysdercus Guerin Meneville 
(Heteroptera, Pyrrhocoridae). — Zoologische Verhandelingen 97:1-215. 

Lima, A. M. da Costa, N. Guitton, and O. V. Ferreira. 1962. Sobre as especies americanas do genero 
Dysdercus Boisuval (Hemiptera, Pyrrhocoridae, Pyrrhocorinae). — Memorias do Instituto Os- 
waldo Cruz 60(l):21-57. 

Silva, A. G. d'Araiijo, C. R. Congalves, D. M. Galyao, A. J. L. Gon9alves, J. Gomes, M. do 
Nascimento Silva, and L. de Simoni. 1968. Quarto catalogo dos insetos que vivem nas plantas 
do Brasil, seus parasitos e predadores. — II(1):622 pp., Rio de Janeiro. 



(J A) Max-Planck-Institut fiir Limnologie, AG Tropenokologie D-232 Plon (Hol- 
stein), West Germany in cooperation with Instituto Nacional de Pesquisas da 
Amazonia (INPA), Manaus, Amazonas, Brazil; (RCF) U.S. National Museum of 
Natural History, Smithsonian Institution, Washington, D.C. Present address: 
INPA, c.p. 478, 69000 Manaus, Amazonas, Brazil. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 377-385 

TWO NEW SPECIES OF THE FROG GENUS HYLODES 

FROM CAPARAO, MINAS GERAIS, BRASIL 

(AMPHIBIA: LEPTODACTYLIDAE) 

W. Ronald Heyer 

Abstract. — Two new species of the leptodactylid frog genus Hy lodes, H. babax 
and H. vanzolinii, are described from the Parque Nacional do Caparao, Minas 
Gerais, Brasil. The identities of Elosia {=Hylodes) glabra and H. lateristrigatus 
are reviewed. 



During a survey of the herpetofauna of the Parque Nacional do Caparao in 
November-December 1980, Dr. P. E. Vanzolini and I collected specimens of two 
species of Hy lodes. Study of these frogs in the laboratory reveals that both 
species are new to science. 

Examination of other Hylodes collected recently in the Atlantic Forests of 
Brasil, suggests that the current allocation of specific names to certain Hylodes 
species is in error. Reassignment of specific names is clarified preparatory to the 
description of the two new species from Caparao. 

Advertising calls were recorded on a Uher CR 134 cassette recorder with a 
Uher microphone or a Sony TCM 280 cassette recorder with a Sennheiser K3U 
microphone. Calls were analyzed on a Kay Sonagraph 6061 B, narrow filter, AGC 
in the off position. 

Current Status of Names 

Relatively few names have been proposed for members of the genus Hylodes. 
As currently understood, the following clusters of species can be recognized on 
the basis of external morphology. (1) The H. pulcher group contains Hylodes 
pulcher (B. Lutz, 1951), a very distinctive, moderate sized, slender, ranoid-like 
species. The distinctiveness of the species derives from its bright life colors; most 
individuals have bright blue and/or yellow dorsal spotting. (2) The H. mertensi 
group contains Hylodes mertensi (Bokermann, 1956), a large (56 mm SVL), ro- 
bust species with leathery dorsal skin. (3) The H. nasus group contains species 
that are moderate to large sized, of robust body form, with granular dorsal sur- 
faces, and lacking light dorsolateral stripes. Eight names have been proposed for 
members of this group: Elosia aspera Miiller, 1924; Hyla nasus Lichtenstein, 
1823; Hyla ranoides Spix, 1824; Elosia nasuta Tschudi, 1838; Elosia bufonium 
Girard, 1853; Hylodes truncatus Steindachner, 1864; Elosia nasus meridionalis 
Mertens, 1927; and Elosia perplicata Miranda- Ribeiro, 1926. (4) The H. lateri- 
strigatus group contains species that are of small to moderate size, body form 
slender, ranoid-like, dorsum smooth, and in most members with light dorsolateral 
stripes. The names proposed for this group are Elosia lateristrigata Baumann, 
1912; Elosia glabra Miranda- Ribeiro, 1926; Elosia magalhaesi Bokermann, 1964; 
Elosia ornata Bokermann, 1967; Hylodes regius Gouvea, 1979. Both new species 
belong to this latter cluster. 



378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



8n 



6- 



N _ 

X 



4- tf^frfffffff 



I 2 



TIME IN SECONDS 

Fig. L Advertising call oi Hy lodes lateristrigatus. From specimen USNM 208557, Brasil, Rio de 
Janeiro, near Teresopolis. Recorded 8 December 1977, air temperature 21.2°C. 



The remainder of this section discusses the status of the names in the H. 
lateristrigatus group and summarizes the advertising calls of those species for 
which calls are known to facilitate comparison with the two new species. 

Bokermann (1967b) discussed the taxonomic status of Elosia i=Hylodes) lat- 
eristrigata and glabra. Prior to Bokermann' s 1967 papers, only one species was 
recognized in the H. lateristrigatus group, i.e., the oldest available name, later- 
istrigatus. Bokermann (1967b) discerned two distinctive advertising calls asso- 
ciated with specimens from Friburgo in the Organ Mountains and Paranapiacaba, 
Sao Paulo, respectively. He allocated the name lateristrigatus to the Organ 
Mountain species. 

Recent collections from around Teresopolis (also in the Organ Mountains) doc- 
ument the presence of two syntopic members of the H. lateristrigatus group. The 
two species are similar morphologically and differ most markedly in size. The 
larger species (males 38-39 mm SVL) has a call of about 1.5 s duration which 
consists of about 13 notes; each note is a rising whistle with a dominant frequency 
of 3700-4300 Hz with about 9 notes per second (Fig. 1). The smaller species 
(males 33-34 mm SVL) has a call of about 1.1-1.3 s duration which usually 
consists of only 2-4 notes; each note is a rising and falling whistle with a dominant 
frequency of 4300-5500 Hz, with 3-4 notes per second (Fig. 2). Bokermann 
(1967b) assigned the name E. lateristrigata Baumann, 1912, to the species with 
the call shown in Fig. 2 (compare with Fig. 1 in his paper; differences are due to 
means of analysis, his figures emphasizing harmonic structure). 

The type-locality given by Baumann is simply "Orgel-Gebirge." Baumann' s 
(1912) description contains no measurements, although his Figure 1 on Plate 4 is 
drawn at natural size. I measured the illustration as 38.9 mm SVL, which agrees 
with data for the larger species at Teresopolis. This form always has sharply 
defined dorsolateral and lower lip-upper arm stripes, as shown in Baumann 's 
figure. Thus, H. lateristrigatus (Baumann) refers to the larger species, with the 
call shown in Fig. 1, contra Bokermann (1967b). 

The species that Bokermann (1967b) recorded from Paranapiacaba and referred 
to as glabra has a call duration of about 2 seconds with 15-30 notes per call, each 
note a rising whistle with a dominant frequency of 5600-6000 Hz (although his 
Fig. 2 shows a dominant frequency of about 4700-5500 Hz), and with 10-13 notes 



VOLUME 95, NUMBER 2 



379 



8-1 



6- 



4- 



^ 



TIME 



IN 



SECONDS 



Fig. 2. Advertising call of Hylodes sp. From specimen USNM field 6124, Brasil, Rio de Janeiro, 
near Teresopolis, calling from same stream as USNM 208557. Recorded 8 December 1977, air tem- 
perature 21. 2°C. 



per second. Bokermann examined the type of Elosia glabra Miranda- Ribeiro, but 
was not able to locate the specific site of "ribeirao da Passagem" within the 
general type locality of Itatiaia. However, Bokermann found a species of the H. 
lateristrigatus group on the lower slopes of Itatiaia at 700 m which compared well 
with the specimens from Paranapiacaba and the type of E. glabra. Two facts 
suggest that E. glabra does not refer to the species that Bokermann recorded 
from Paranapiacaba and collected from the lower slopes of Itatiaia. The "ribeirao 
da Passagem" is in the planalto of Itatiaia at 2200 m elevation at approximately 
22°25'S, 44°39'W (P. E. Vanzolini, in lift.). Second, the figures of E. glabra 
(Miranda- Ribeiro 1926, PL 4, Figs. 1, la, lb) do not appear to represent a species 
of the H. lateristrigatus group. The figures in Miranda- Ribeiro 's work are accu- 
rate representations of the specimens and species they represent, and show E. 
glabra as a completely uniform brown frog. There are two species of frogs in the 
upper elevations of Itatiaia that have occasional specimens with a uniform pat- 
tern: Hylodes pule her and Eleutherodactylus (Basanitia) nigriventris. The disks 
of the figure of E. glabra appear Basanitia-like', the snout shape is Hylodes-iike. 
The possibility that E. glabra refers to an upper elevation frog, rather than a 
lower elevation Hylodes is merely pointed out here and resolution of this problem 
is deferred until it can be dealt with in a revision of the entire genus. 

The conclusions drawn from the above comments are: (1) Hylodes lateristri- 
gatus is the Organ Mountain species with a fast call rate with more notes; (2) the 
species Bokermann referred to as Elosia lateristrigata currently has no name; 
(3) the species name glabra may not refer to the species for which Bokermann 
described and figured the call; and (4) the calls of H. lateristrigatus (in the sense 
used here) and the calls described by Bokermann (1967b) as glabra are distinctive 
and most probably represent two distinct species. 

Hylodes magalhaesi is a moderate sized species (male 30 mm SVL), some 
individuals of which have a distinctive belly pattern of discrete light spots on a 
dark background. The advertising call averages about 1 s duration, with about 26 
notes per call; each note has a dominant frequency centering on about 2000 Hz, 
and a rate of 20-25 notes per second (Bokermann 1964). 

Hylodes ornatus is a small species (males 25 mm SVL) (Bokermann 1967a). 



380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



6H ' ' ' * * * f f f 9 i i f t $ $ 9 i t t 9 t 9 $ t t t t f t ' 

N 

X 

4H 



2- 



I 2 

TIME IN SECONDS 

Fig. 3. Advertising call of Hylodes regius. From specimen USNM field 5316, Brasil, Rio de 
Janeiro, Brejo da Lapa, Itatiaia. Recorded 7 January 1977, air temperature 18-20°C. 



Most individuals have a distinctive dorsal pattern of two broad light dorsal stripes 
in addition to the narrow light dorsolateral stripes. Males have external vocal 
sacs indicating the species is vocal, but the call is presently unrecorded. 

Hylodes regius is a moderate sized species (males 33-36 mm SVL) with a 
distinctive dorsal pattern of small, irregular brilliant yellow spots on the dorsum 
(Gouvea 1979). The call is reported here for the first time. The call duration is 
about 1.7 s, with about 22-32 notes per call. Each note is a sharply rising whistle 
with a dominant frequency ranging from 5200-6300 Hz, and with 14-19 notes per 
second (Fig. 3). 

The two new species from Caparao are distinct from all taxa discussed above. 

Hylodes babax, new species 
Fig. 4 

Holotype. — MZUSP 57949, adult male, from Brasil: Minas Gerais; Parque Na- 
cional do Caparao, 1200 m, 20°26'S, 4r47'W. Collected by W. Ronald Heyer, 30 
November 1980. 

Paratopotype. — USNM 222553, male, same data as holotype. 

Diagnosis. — The dorsal coloration is sharply set off from the lateral coloration 
by a weakly developed light dorsolateral stripe. This pattern distinguishes Hy- 
lodes babax from the other new species from Caparao. Hylodes babax is small 
(male SVL 31-33 mm) and has a dark belly with light spots; the two species at 
Teresopolis are larger {lateristrigatus male SVL 38^0 mm; the second species 
male SVL 33-34 mm) and have light bellies with dark spots. Hylodes babax is 
larger than Hylodes ornatus (males SVL 25 mm) and lacks the distinctive dorsal 
pattern of a pair of elongate light dorsal bands characteristic of H. ornatus. 
Hylodes babax resembles H. magalhaesi from Campos do Jordao, Serra da Man- 
tiqueira, Sao Paulo, Brasil, in size and general coloration. In life, the under 
surfaces of the legs are brick red in H. babax, yellowish in H. magalhaesi. The 
advertising calls of H. babax and magalhaesi are strikingly different (compare 
Fig. 4 of Bokermann 1964, with Fig. 5 here) . 

Description of holotype. — Snout rounded-truncate from above, protruding in 
profile; canthus rostralis angular-obtuse; loreal weakly concave in cross section; 



VOLUME 95, NUMBER 2 



381 





Fig. 4. Holotype oi Hy lodes babax (MZUSP 57949, a male): dorsal and ventral views. 



tympanum distinct, diameter about % eye diameter; vomerine teeth in two small 
transverse patches, medial and just posterior to choanae; vocal slits and well 
developed lateral vocal pouches present; first finger long, just longer than second; 
thumb lacking nuptial asperities; dorsal texture essentially smooth, with small 
pebble like granulations; weak supratympanic fold, no other body folds obvious; 
no body glands; belly smooth, ventral femur granular; fingers and toes disked, 
disks about Vi again as broad as toe width just behind disks, toe disks slightly 
larger than finger disks, each disk with a pair of dorsal scutes; fingers and toes 
with considerable fringe; subarticular tubercles present but not prominent; ovate 
inner metatarsal tubercle not quite twice as large as semicircular outer metatarsal 
tubercle; well developed tarsal fold extending about % distance of tarsus, con- 
tinuous distally with outer toe fringe of toe 5; no metatarsal fold; outer tarsus and 
sole of foot smooth. 

SVL 30.6; head length 11.6; head width 9.7; eye-nostril distance 2.5; femur 
15.1; tibia 17.2; foot 15.9 mm. 

Dorsal pattern of variegated brown and tan markings with irregular dark brown 
interocular mark; sides almost black, demarcated from dorsal coloration by thin 
light pinstripe extending from tip of snout over the eye to the upper groin; light 
lip stripe, broader but less distinctive anteriorly; upper limbs brown with irregular 
dark transverse bars, upper forearm with light pin stripe on anterior aspect of 
arm from shoulder to inner elbow; throat black with very few, somewhat sym- 
metrically arranged light dots; belly black with contrasting white irregular spots, 
posterior surface of thigh with indistinct dark mottle; ventral surface of thigh with 
pale straw colored patch, other ventral limb surfaces black with little pale areas. 

In life the upper lip stripe tannish bronze; the belly with contrasting black and 
white markings, and the lower surface of the thigh brick (not brilliant) red. 



382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



8n 



6- 



t4 _ 

I 



2- 



TIME IN SECONDS 

Fig. 5. Advertising call of Hylodes babax. From specimen USNM 222553, recorded 30 November 
1980, 1540 h, air temperature 21.6°C. 



Description of paratopotype. — The paratopotype resembles the holotype in 
most aspects with the following exceptions: dorsal coloration not as dark, weak 
bronze colored dorsolateral fold demarcates dorsal and lateral patterns; belly with 
small and less distinct light spots; SVL 32.6 mm. 

Advertising call. — Call duration 0.23-0.48 s, mean 0.36 s; 4-8 notes per call, 
mode 6; each note a rising whistle, typically first note lower in pitch (mean 
dominant frequencies 4290-5030 Hz) than remaining notes (mean dominant fre- 
quencies 4690-5420 Hz); notes produced at a mean rate of 16.2 notes per second 
(Fig. 5). 

Etymology. — The name babax is Greek for chatterer, in allusion both to the 
insistent diurnal calling of this species (as in many other Hylodes species) and to 
the fact that of the two species of the H. lateristrigatus group from Caparao, this 
is the only one known to call. 

Hylodes vanzolinii, new species 
Fig. 6 

Holotype. — MZUSP 57950, male, from Brasil: Minas Gerais; Parque Nacional 
do Caparao, 2300 m, 20°26'S, 41°47'W. Collected by W. Ronald Heyer and P. E. 
Vanzolini, 5 December 1980. 

Paratopotypes. — MZUSP 52923, collected by Lynn C. Branch, November 
1977; USNM 222554-5, same data as holotype. 

Diagnosis. — The dorsal color pattern is not sharply set off from the lateral 
coloration. This pattern is distinct from all other members of the H. lateristrigatus 
group with the possible exception oi glabra (see comments in previous section), 
which in the original figure is completely uniform dorsally and ventrally. Hylodes 
vanzolinii has small bright yellow dorsal spots in life (white in preservative) and 
a variegated belly, differing from the figures oi E. glabra. 

Description of holotype. — Snout rounded-truncate from above, protruding in 
profile; canthus rostralis angularly obtuse; loreal slightly concave in cross section; 
tympanum about Vi diameter of eye; vomerine teeth in two small slightly trans- 
verse patches between posterior extent of choanae; no vocal slits or vocal sac; 
first finger long, just longer than second; thumb lacking nuptial asperities; dorsal 



VOLUME 95, NUMBER 2 



383 





Fig. 6. Holotype oi Hy lodes vanzolinii (MZUSP 57950, a male): dorsal and ventral views. 



texture smooth; very weak supratympanic fold, otherwise no other body folds or 
glandular structures; belly smooth, under surface of thighs granular; finger and 
toe tips with disks, disks about V2 again as broad as digit immediately behind 
disk, finger and toe disks about equal size, upper surface of disks with a pair of 
scutes; fingers with lateral ridge, toes with extensive lateral fringe; subarticular 
tubercles moderate; inner ovate metatarsal tubercle much larger than rounded 
outer metatarsal; extensive tarsal fold extending about % distance of tarsus, con- 
tinuous distally with toe fringe on outer side of first toe; no metatarsal fold; outer 
tarsus and sole of foot smooth. 

SVL 29.0; head length 10.4; head width 9.5; eye-nostril distance 2.0; femur 
16.0; tibia 16.9; foot 15.8 mm. 

Dorsum almost black with nondescript grayish mottling and with a few small 
light dots; dorsal pattern not distinct from lateral pattern except on head with an 
irregular whitish gray stripe from tip of snout to just over tympanum demarcating 
dorsal pattern from uniform dark lateral head pattern; light upper lip stripe in- 
complete, weaker in front of eye than behind; dorsal limb surfaces almost uni- 
formly dark brown, front of forearm with very weak, incomplete light pin-stripe; 
throat mostly brown with a symmetrical series of medial coalescing spots; belly 
variegated brown and white in about equal ratio; posterior and ventral surfaces 
of thighs uniform brown. 

In life upper lip stripe dull white; dorsum dark green-black with yellowish 
flecks; belly white and tan; posterior surface of thigh uniform tan. 

Description of paratopotypes. — USNM 222554, a 37.3 mm female, resembles 
the holotype in pattern with the following differences: the upper lip has a series 
of light dots, not a stripe; there is no light stripe separating the dorsal color from 
the side color on the head; the throat is mostly white with some dark markings. 
MZUSP 52923 is uniformly dark dorsally and laterally except for a few small light 
dorsal spots and a faint light upper lip stripe mostly in front of the eye; the belly 
is dark with small, distinct light spots. 



384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Advertising call. — The fact that no Hy lodes vanzolinii were heard calhng, to- 
gether with the lack of vocal slits in the males, strongly suggests that this species 
lacks an advertisement call. 

Etymology. — It is a pleasure to name this species for P. E. Vanzolini. First, I 
wish to commemorate an enjoyable field trip during which he shared many in- 
sights concerning the Atlantic Forest with me. Second, I wish to pay tribute to 
Dr. Vanzolini' s studies on the montane lizard fauna of eastern Brasil where he 
developed the concept of stranded species. Without his studies, explanation of 
the history of this new montane frog would not be possible. 

Discussion 

Hylodes vanzolinii replaces H. babax altitudinally in the Parque Nacional do 
Caparao, with the latter occurring along streams in the Atlantic Forest vegetation 
and H. vanzolinii occurring along the exposed streams at elevations above the 
Atlantic Forest vegetation (1900 m and above). The distributions of the species 
are so far known only from the collections reported here. 

Hylodes babax is similar to several other members of the H. lateristrigatus 
group and on the basis of color pattern is predicted to be related to one of these 
species than to the altitudinally adjacent H. vanzolinii. 

Hylodes vanzolinii and H. regius (Itatiaia, Serra da Mantiqueira) both have 
small yellow spots on the dorsum. Hylodes vanzolinii differs from H. regius in 
the dorsolateral stripes and in lacking a voice. These two species do not seem to 
be closely related. 

As far as is known, there are only two Hylodes species that lack advertising 
calls, H. pulcher and vanzolinii. Both of these species are montane (in the coastal 
mountain system of Brasil at elevations above 1700-1900 meters) species that 
have striking dorsal colors in life. The relationships between H. pulcher and 
vanzolinii are probably not close, however, as the color patterns and sizes are 
very distinctive. 

There are two major zoogeographic patterns that account for montane distri- 
butions in eastern Brasil. The first is one of relictual distribution of stranded 
species associated with drier and cooler climates that today are restricted to the 
upper elevations of mountains, but that were much more widespread during gla- 
cial maxima periods of the Pleistocene (Vanzolini 1982; Vanzolini and Ramos 
1977). The montane species that exhibit this zoogeographic pattern are closely 
related to each other. The second pattern is one of in situ differentiation, where 
the present montane species were independently derived from lowland forms. 
Hylodes vanzolinii appears to fit into this second zoogeographic pattern. Detailed 
resolution of relationships awaits biochemical and morphological analyses. 

Acknowledgments 

P. E. Vanzolini (Museu de Zoologia da Universidade de Sao Paulo, MZUSP) 
obtained the proper Instituto Brasileiro de Desenvolvimento Florestal permits 
and invited me to join in the exploration of the herpetofauna of the Parque Na- 
cional do Caparao. George R. Zug and Ronald I. Crombie (National Museum of 
Natural History, Smithsonian Institution, USNM) critically read the manuscript. 
George Steyskal (USNM) advised on scientific name usage and formation. 



VOLUME 95, NUMBER 2 385 

The following programs and institutions supported this research: Funda9ao de 
Amparo a Pesquisa do Estado de Sao Paulo; Museu de Zoologia da Universidade 
de Sao Paulo; and the International Environmental Science Neotropical Lowland 
Research Program, Smithsonian Institution. 

Literature Cited 

Baumann, F. 1912. Brasilianische Batrachier des Berner Naturhistorischen Museums. — Zoologischen 
Jahrbiichern, Abteilung fiir Systematik, Geographic und Biologie der Tiere 33:87-172 + plates 
4&5. 

Bokermann, W. C. A. 1964. Una nueva e specie de Elosia de la Serra da Mantiqucira, Brasil (Am- 
phibia, Leptodactylidac). — Neotropica 10:102-107. 

. 1967a. Una nueva especie de Elosia de Itatiaia, Brasil (Amphibia, Leptodactylidac). — Neo- 
tropica 13:135-137. 

. 1967b. Notas sobre cantos nupciais de anfibios brasileiros. IL O canto de "Elosia lateristri- 

gata" e "Elosia glabra" (Anura). — Re vista Brasileira de Biologia 27:229-231. 

Gouvea, E. 1979. Uma nova especie de elosiineo da Serra do Itatiaia (Amphibia, Anura, Leptodac- 
tylidac). — Re vista Brasileira de Biologia 39:855-859. 

Miranda-Ribeiro, A. 1926. Notas para servirem ao estudo das Gymnobatrachios (Anura) Brasileiros — 
Tomo primeiro. — Archivos do Museu Nacional do Rio de Janeiro 27:1-227 + 22 plates. 

VanzoHni, P. E. 1982. A new Gymnodactylus from Minas Gerais, Brasil, with remarks on the genus, 
on the area and on montane endemisms in Brasil (Sauria, Gekkonidae). — Papeis Avulsos de 
Zoologia 34:403-413. 

, and A. M. M. Ramos. 1977. A new species of Colobodactylus, with notes on the distribution 

of a group of stranded microteiid lizards (Sauria, Teiidae). — Papeis Avulsos de Zoologia 31:19- 
47. 

Amphibians and Reptiles, Department of Vertebrate Zoology, Smithsonian In- 
stitution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 386-391 

A NEW CAVE PLATYMANTIS (AMPHIBIA: RANIDAE) 
FROM THE PHILIPPINE ISLANDS 

Walter C. Brown and Angel C. Alcala 

Abstract. — A new cave-dwelling frog, Platymantis spelaeus (Ranidae), is de- 
scribed from southern Negros Island in the Philippines. This is the second cave- 
dweller and the twelfth species of the genus known from this archipelago. 



Recognized species of the genus Platymantis include two from the Fiji Islands, 
nine from the Solomons, nine from the Bismarcks, four or five from New Guinea 
and small associated islands, one from the Palaus, and eleven from the Philip- 
pines. With the exception of one species (as presently defined) which ranges from 
New Guinea to the Solomons, all are endemic to the regions indicated. 

In the western Philippines the genus is absent from Palawan and associated 
small islands. In the rest of Philippines, three species {P. corrugatus, P. dorsalis 
and P. guentheri) are widespread, being recorded from Mindanao in the south 
through Luzon in the north. In addition, P. ingeri occurs in Mindanao and other 
islands in the southern part, P. hazelae has been recorded from Negros and 
Luzon Islands, and the very closely related P. polillensis is known from Polillo 
Island. The remaining five species are known from limited localities on single 
islands. Platymantis cornutus and P. subterrestris are recorded from Luzon Is- 
land, P. lawtoni and P. levigatus from Tablas Island to the south of Luzon and 
east of Mindoro, and P. insulatus from the very small island, Gigante South, off 
the northeast coast of Panay. 

Typically these frogs inhabit the rain forest, being either arboreal or in habitats 
of the duff or under rocks and logs on the forest floor. Only one species, P. 
insulatus (Brown and Alcala, 1970), has previously been recorded as cave-dwell- 
ing. Now a second such species is described in this paper. The type series is from 
two limestone caves in a forested area in southern Negros Island. 

Measurements of specimens were made using a Helios dial caliper. Head length 
is measured from the tip of the snout to the posterior edge of the tympanum and 
head breadth at the widest point near the angle of the jaws. The diameter of the 
eye is from the anterior to the posterior edge of the socket. Finger and toe lengths 
are to the proximal edge of the proximal subarticular tubercle. Other measure- 
ments are probably not subject to much variation in method. 

Platymantis spelaeus, new species 
Fig. 1 

Holotype. — California Academy of Sciences number 153469, mature female, 
collected in a limestone cave, Tiyabanan Barrio, Basay, southern Negros Ori- 
ental, Negros Island, Philippines, by Angel Alcala and C. A. Ross, 18 March 
1981. 

Paratypes. — California Academy of Sciences 153470-83; United States Na- 
tional Museum 221838-39; Australian Museum, Sydney R98394; Field Museum 



VOLUME 95, NUMBER 2 



387 




Fig. 1. Platymantis spelaeus, showing the lack of ridges on the dorsum, the large tympanum, and 
characteristic features of the color pattern. 



Natural History 213331; British Museum (Natural History) 1981.9; Museum of 
Comparative Zoology, Harvard A 100300; Silliman University, Philippines 2288- 
92; American Museum Natural History 109454, all from two caves in southern 
Negros Island. 

Diagnosis. — This Platymantis is distinguished from other Philippine species by 
the following combination of characters: (1) Large size (snout-vent length 41,5- 
60.5 mm for 17 adults); (2) The only slightly dilated finger disks, about the same 
diameter as those of the toes; (3) Tympanum relatively large, its diameter about 
62-82% of that of the eye; (4) Skin of the dorsum rough (shagreened) , neither 
smooth nor with prominent tubercles or elongate ridges. 

Description. ^A large Platymantis, snout-vent length (SVL) 41.5^6.9 mm for 
5 males and 52.8-60.5 mm for 11 females (Table 1); head breadth 107-111% of 
head length; snout broadly rounded, its length 40^4% of head breadth and 41- 
49% of head length; upper jaw protruding; canthus rostralis rounded; lores mod- 
erately oblique, concave; diameter of eye 27-37% of head breadth and 63-88% 
of snout length; tympanum large, its diameter 63-82% of eye diameter, 105-147% 
of interorbital distance, and about twice the diameter of the third finger disk; 
prominent fold dorsal and posterior to tympanum; dorsum shagreened (Fig. 1) 
but lacking ridges; venter smooth; fingers slender, without webs; tips of fingers 
with rounded disks having a circummarginal groove but lacking a transverse 
ventral groove (Fig. 2); diameter of disk of third finger 20-25% of length of third 
finger (measured to base of second subarticular tubercle); disk of second finger 



388 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Snout-vent lengths for Philippine species oi Platymantis (number in sample in paren- 
theses). 







Snout-vent length (mm) 






,^,/ 


Sex 




Species 


Males 


undetermined 


Females 


P. cornutus 




30.7 (1) 




P. corrugatus 


28.5-35.0 (15) 




33.6-50.2 (15) 


P. dorsalis 


21.5-30.5 (25) 




25.5-42.4 (30) 


P. guentheri 


27.4-35.5 (24) 




38.3-49.1 (20) 


P. hazelae 


20.6-27.6 (15) 




20.7-31.7(15) 


P. ingeri 


23.0-30.5 (16) 




26.9-34.5 (23) 


P. insulatus 


37.8-41.7(3) 




40.2-45.5 (3) 


P. lawtoni 


— 




39.0(1) 


P. levigatus 


30.3 (1) 




38.5 (1) 


P. polillensis 


20.2-22.6 (2) 




— 


P. spelaeus 


41.5^6.9(6) 




52.8-60.5(11) 


P. subterrestris 


24.0-26.3 (2) 


27.2(1) 





nearly as large as that of third finger (Table 2); first finger slightly longer than 
second when adpressed; fingers with prominent rounded subarticular tubercles; 
a prominent palmer tubercle at base of each finger; three large metacarpal tuber- 
cles; hind limbs relatively long, length of tibia 52-63% of snout-vent length and 
127-148% of head breadth; toes with small web at base; tips dilated into rounded 
disks with circum marginal grooves; diameter of disk of third toe 87-1.07% of 
diameter of disk of third finger; subarticular tubercles prominent, round-pointed; 

Table 2. — Proportions in samples of some of the larger Philippine species oi Platymantis. 



p. guentheri 
(n = 20) 



P. insulatus 
(n = 5) 



P. levigatus 
(n = 2) 



P. spelaeus 
(n = 19) 



Eye diameter 
Head breadth 
Eye diameter 
Snout length 
Snout length 
Head breadth 
Tympanum diameter 
Eye diameter 
Tympanum diameter 
Breadth of 3rd finger disk 
Interorbital breadth 
Eye diameter 
Breadth of 3rd finger disk 
Length of 3rd finger 
Breadth of 2nd finger disk 
Breadth of 3rd finger disk 
Breadth of 3rd toe disk 
Breadth of 3rd finger disk 



0.33-0.39 


0.37-0.41 


0.29-0.30 


0.27-0.37 


1.04-1.09 


0.99-1.06 


0.97-0.98 


0.63-0.88 


0.30-0.35 


0.35-0.39 


0.29-0.31 


0.40-0.49 


0.37-0.48 


0.48-0.61 


0.33-0.40 


0.63-0.82 


0.63-0.87 


1.22-1.71 


1.40-1.50 


1.89-2.60 


0.45-0.67 


0.44-0.49 


0.60-0.61 


0.47-0.67 


0.36-0.51 


0.27-0.31 


0.17-0.20 


0.20-0.25 


0.73-0.90 


0.63-0.74 


0.91-1.00 


0.76-0.95 


0.47-0.56 


0.62-0.74 


0.85-0.90 


0.87-1.07 



VOLUME 95, NUMBER 2 



389 




Fig. 2. P. spelaeus (SU 2288): palmar view of hand. 



an elongate outer metatarsal tubercle and a rounded inner one; solar tubercles 
lacking (Fig. 3). 

Measurements {in mm) of holotype . — Snout-vent length 60.5; head length 22.5 
head breadth 24.4; snout-length 10.0; eye diameter 8.8; tympanum diameter 5.5 
interorbital distance 4.4; 3rd finger length 10.8; diameter of 3rd finger disk 2.7 
diameter of 3rd toe disk 2.5; hind limb length 95 ±; tibia length 31.5. 

Color. — In preservative the dorsum and the head are mottled grayish-olive and 
black with a short, broad, blackish "W" in the forehmb region. The lateral sur- 
faces are more grayish with the darker mottling reduced. The lips are dark barred. 
The dark transverse bands on the limbs are about as wide as or wider than the 
light bands (Fig. 1). The venter is creamy white with some faint gray mottling. 

In life the dorsum is olive-green to brown with darker mottling. The upper 
surfaces of the thighs are dark barred and the inner surface either orange or 
lavender. The venter is cream colored with or without brown flecks. 

Comparisons. — This is the largest species of the genus known from the Phil- 
ippines (Table 1). Eight of the previously described species are small, arboreal 
forms with the SVL in mature males under 36 mm and with females rarely at- 
taining a SVL greater than 40 mm {P. dorsalis does). Only one mature female is 
known for both P. lawtoni and P. levigatus, in which the SVL is 39 and 38.5, 
respectively, and it is possible that a SVL of 40 mm may be exceeded in those 
species. Only males off. insulatus overlap P. spelaeus in SVL and only females 



390 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. P. spelaeus (SU 2288): plantar view of foot. 



of P. corrugatus, P. dorsalis, P. guentheri and P. insulatus approach females of 
P. spelaeus in size although they do not overlap (Table 1). 

Most Philippine species also have broadly dilated digital disks, especially on 
the fore limbs (Inger 1954; Brown and Alcala 1963, 1970, 1974). Only in P. cor- 



VOLUME 95, NUMBER 2 391 

rugatus, p. dorsalis and P. levigatus are the finger tips only slightly dilated as is 
characteristic of P. spelaeus. 

The shagreened skin of the dorsum (not smooth or with prominent tubercles 
or ridges) is also characteristic of P. insulatus and P. lawtoni. 

Platymantis spelaeus, insulatus and dorsalis are the most apt to be confused 
in terms of general appearance. The somewhat smaller P. dorsalis has prominent, 
short ridges on the dorsum, and the general color is more brown to reddish. 
Platymantis insulatus is closer to P. spelaeus in size, texture of the skin on the 
dorsum, and general color pattern, but the tips of the fingers are much more 
broadly dilated and the tympanum is smaller (Table 2). The species P. spelaeus 
and P. insulatus are probably closely related. 

Ecological notes. — All specimens were collected near the entrances of two 
moist limestone caves, in crevices or pockets in the walls or on the floor. Sixty 
large (2.5-3.0 mm diameter) unpigmented eggs were found in the ovaries of a 
large gravid female (55 mm SVL). 

Acknowledgments 

We gratefully acknowledge the assistance of Charles A. Ross of the U.S. Na- 
tional Museum of Natural History, Smithsonian Institution in the field work and 
W. Ronald Heyer of the same institution for the loan of specimens. We also thank 
Richard G. Zweifel, American Museum of Natural History, for reading the manu- 
script. 

Literature Cited 

Brown, Walter C, and Angel C. Alcala. 1963. A new frog of the genus Cornufer (Ranidae) with 
notes on other amphibians known from Bohol Island, Philippines. — Copeia 1963:672-675. 

, and . 1970. A new species of the genus Platymantis (Ranidae) with a list of amphibians 

known from South Gigante Island, Philippines. — Occasional Papers of the California Academy 
of Sciences 84:1-8. 

, and . 1974. New frogs of the genus Platymantis (Ranidae) from the Philippines. — 

Occasional Papers of the California Academy of Sciences 113:1-12. 

Inger, Robert F. 1954. Systematics and zoogeography of the Philippine Amphibia. — Fieldiana: Zo- 
ology 33:183-531. 

(WCB) Department of Herpetology, California Academy of Sciences, Golden 
Gate Park, San Francisco, California 94118 and Menlo College, Menlo Park, 
California 94025; (AC A) Department of Biology, Silliman University, Manila, 
Philippines. 



PROC. BIOL. SOC. WASH. 
95(2), 1982, pp. 392-397 

A NEW SPECIES OF SPHAERODACTYLUS 

(REPTILIA: SAURIA: GEKKONIDAE) 

FROM EASTERN CUBA 

Orlando H. Garrido and Albert Schwartz 

Abstract. — A new species of gecko, Sphaerodactylus celicara, of the notatus- 
group, is described from extreme northeastern Cuba. Comparisons are made with 
its most closely related geographical congeners, 5. notatus and S. bromeliarum. 



A striking feature of the distribution of Sphaerodactylus in Cuba is that the 
easternmost province, Oriente, harbors the greatest diversity of species, repre- 
senting several species-groups of these small lizards. One of these, S. notatus, 
belongs to a complex of geckos whose center of diversity is Hispaniola (and 
secondarily Puerto Rico). On Cuba, there is only one species {S. bromeliarum 
Peters and Schwartz) that is closely related to S. notatus, in contrast to 14 «o- 
tatus -gvon^ species on Hispaniola. 

There are some peculiarities in scutellation and pattern in S. notatus on Cuba. 
For instance, keeling on the throat does not occur in specimens from the Isla de 
la Juventud (=Isla de Pinos) or, on the main island, from Pinar del Rio to Ca- 
magiiey provinces. Throat keeling occurs more or less variably (in some individ- 
uals from the same locality and in varying degrees) in Oriente specimens. A 
second variable feature is the occurrence in females of a dark scapular patch with 
2, rarely 1, included pale ocelli. In western and central Cuba, the female scapular 
pattern is absent. In Oriente, on the other hand, 26% of the females have a 
scapular patch and ocelli. Since absence of this patch, the presence of which is 
characteristic of S. bromeliarum, is one of the distinctive characteristics of the 
Cuban subspecies S. n. atactus Schwartz, these eastern specimens have seemed 
anomalous. 

The senior author sent a series of 14 specimens from the extreme northeastern 
region of Cuba to the junior author. Many of these are larger lizards than typical 
S. n. atactus and differ in scutellation; in addition, they to a large extent (but not 
totally) clarify the problem of occurrence of a female scapular patch and ocelli. 
The males normally have both the upperside of the head and throat vividly and 
distinctly spotted or dotted with very dark brown. Data from these specimens 
were combined with data from other geckos from this same general region, using 
the presence of a scapular patch and ocelli and scutellar counts as criteria, and 
we now have a clearer picture of the variation in notatus -Vikt geckos in this 
particular region. We have examined a total of 239 S. notatus from Cuba, in- 
cluding 182 specimens from Oriente alone (on 118 of which full counts were 
taken) and 37 from the Isla de la Juventud and the Archipielago de los Canarreos, 
and we feel secure in naming this northeastern population as a distinct species. 

Sphaerodactylus celicara, new species 

Holotype. — IZ 5613, adult female, from Asuncion, Maisi, Baracoa, Oriente 
Province, Cuba, collected 10 May 1979 by Orlando H. Garrido. 



VOLUME 95, NUMBER 2 393 

Paratypes. —IZ 5611-12, IZ 5616-19, ASFS V44969-70, same data as holotype; 
ASFS V44968, La Maquina, Gran Tierra, Maisi, Oriente Province, Cuba, 10 May 
1979, M. L. Jaume; IZ 5378, Hotel Asuncion, Grand Tierra, Maisi, Oriente Prov- 
ince, Cuba; 17 September 1978, P. Espinosa, H. Saragua, A. Vega; IZ 4725, 3 
km W Yumuri, Oriente Province, Cuba, May 1976, Gilbert; IZ 5414-15, Zapote 
de Mandinga, Baracoa, Oriente Province, Cuba, 10 May 1979, O. H. Garrido. 

Associated specimens. — All are from Oriente Province, Cuba: IZ 3455-57, 
Hoyos de Sabanilla; IZ 4214-16, La Poa, Sabanilla; IZ 3453, IZ 3537, IZ 3585, 
base of Monte Iberia; IZ 3648, Jaguani, Baracoa: IZ 106, IZ 113, La Tinta, barrio 
be Jauco, Baracoa; IZ 4096, La Yagruma, Maisi, Baracoa; IZ 4258, Punta de 
Maisi; MCZ 13595-96, Jauco, seacoast at Cabo Maisi; MCZ 11215, Cueva de 
Majana, Baracoa. 

Definition. — A species of the notatus-group oi Sphaerodactylus characterized 
by the combination of: 1) moderate size, males to .32 mm, females to 31 mm 
snout-vent length (SVL); 2) high number of dorsal scales (21-30) between axilla 
and groin, each dorsal scale with 11-14 hair-bearing organs, each with one hair, 
around the free edge and apex of the scale; 3) high number of scales (40-51) 
around midbody; 4) scapular patch and ocelli almost always present in females, 
only ocelli usually present in males; 5) heads of adults of both sexes heavily 
dotted or marbled with dark brown, both dorsally, and ventrally on the throat, 
in strong contrast to the brown color of the remainder of the body; 6) females 
typically bilineate dorsally, males more or less unicolor to salt-and-pepper, 
brown; 7) internasals 0-2, with a high incidence (35%) of internasals absent (=0). 

Description of holotype . — Adult female, SVL 31 mm, tail length 34 mm. Scale 
counts: dorsal scales keeled, tectiform, imbricate, between axilla and groin 29; 
ventral scales between axilla and groin 24; scales around midbody 47; fourth toe 
lamellae 11; 3/3 supralabials to mid-eye; no internasals; 2 postnasals. Throat, 
chest, and ventral scales smooth, imbricate, cycloid. Dorsal color (preserved) 
dark brown with dorsolateral pair of slightly paler lines (Fig. 1), strongly outlined 
with black, beginning at well developed black scapular patch which includes two 
pale (grayish) ocelli; head pattern basically dark trilineate, with a median dark 
line, narrow on snout and expanding on occiput and continuing to scapular patch, 
somewhat paler centrally, and on each side a narrow dark canthal line to the eye, 
continuing posteriorly from eye as broad dark line to lateral margins of scapular 
patch, the two forming a more or less square brownish area surrounding the 
scapular patch. The lateral pair of cephalic lines bordered below by pale lines 
that continue clearly above the forelimb insertion; pale area between lateral lines 
and central dark line on head containing a dark postocular line extending to 
occiput. Sides of neck brownish, dotted with darker brown; upper lip mottled 
with strongly contrasting dark brown and pale ground color, the mottling ex- 
tending onto the throat as strongly contrasting dark brown dots and lines, this 
pattern becoming diffuse on the chest which is stippled with dark brown. Ventral 
scales edged with dark brown, resulting in a generally dark venter; tail brown 
with obscure darker brown marbling; upperside of all limbs dark brown, stippled 
or spotted with darker brown. 

Variation. — The holotype and 30 paratypes and associated specimens show the 
following variation. Largest males (IZ 5611, IZ 5617) 32 mm SVL (range 25-32 
mm), largest females (IZ 5613, IZ 5619) 31 mm (22-31 mm); dorsals between 
axilla and groin 21-30 {x = 25.8), ventrals between axilla and groin 22-34 (26.1); 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






Fig. I. Sphaerodactylus celicara, holotype (IZ 5613). Upper figure, dorsal view; middle figure, 
lateral view of head pattern; lower figure, ventral view of throat pattern. 



midbody scales 40-51 (45.2); fourth toe lamellae 9-13 (10.6; mode 11); suprala- 
bials 3/3 (29 individuals), 3/4 (1), 4/4 (1); internasals (11 individuals), 1 (17), 2 
(3); male escutcheon 4-7 x 10-26; throat, chest, and ventral scales smooth in 23 
specimens, throat scales keeled in 7 specimens; upper side of head lined or heavily 
dotted in 7 males, head markings absent in 2; throats heavily dotted in 7 males, 
these markings absent in 2; scapular patch and ocelli present in 21 females, absent 
in one. 

Of the four male paratypes, three agree well with the definition of the species. 
They have the dorsum salt-and-pepper or apparently unicolor tan with strongly 
contrasting dark brown head mottling or marking on a pale ground. The dark 
dorsal head mottling continues onto the throat in these specimens as very heavy 
dark mottling comparable to that of the upper surface of the head. Venters have 



VOLUME 95, NUMBER 2 



395 



20 
15 
10 

5 

N 

5 

10 

15"- 



m 




2ir 



22 



24 



26 

SVL 



28 



30 



32 



Fig. 2. Histogram of snout-vent length (in mm) of two samples of S. notatus (open rectangles) 
and S. celicara (dark rectangles); males are shown above the horizontal line, females below it. Each 
individual is indicated by an interval of 0.2 mm. 



dark-edged scales. The exceptional male (IZ 5612, SVL 31 mm) lacks any dorsal 
or ventral pattern. Although there is no dark scapular patch in males, two (IZ 
561 1, IZ 5617) have a faint pair of pale ocelli; the other two males (IZ 5612, ASFS 
V44968) lack them. 

Scapular patches and oceUi are present in all but one of the 10 female paratypes, 
as well as in all 10 associated females and one juvenile. The dorsal bilineate 
pattern is usually quite distinct (even in small lizards — IZ 5378, SVL 23 mm) and 
is especially vivid in some (ASFS V44969). In some smaller Uzards the tail has 
a tiny black tip, with a broader white band proximally, which is in turn bounded 
by a narrower black band. The head pattern is trilineate, with interspace dark 
lines between the major head lines in all specimens; there is no blurring or dis- 
tortion of the three major cephalic lines in large specimens. Scapular patches 
vary from large and extensive (ASFS V44970) to relatively small and constricted 
(IZ 5614), but they and the included ocelli are always present. Dark throat mark- 
ings are almost always present and obvious, varying from heavy in the holotype 
to less bold and lineate in some specimens. The distinct pale-and-dark neck 
streaking is also a constant feature, although the more lateral nuchal lines may 
disintegrate into a linear series of very dark spots or blotches (IZ 5618). 

The associated specimens agree very well with our concept of S. celicara. 
There are five males, ten females, and two juveniles (SVL 16 and 20 mm). Only 
one male (IZ 3537) lacks a dotted or spotted head and throat, and another male 
(MCZ 13595, SVL 25 mm) not only has a dotted head and throat but also has a 
scapular patch and two oceUi, a most unusual condition. In the females, scapular 
patches and oceUi are present in all but one (IZ 3453, adult, SVL 29 mm). The 
juveniles show the female head and shoulder pattern. 



396 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Extremes and means of four characters of S. notatus and 5'. celicara, plus data on 
underbody keeling. S. notatus has been divided into two samples: Sample 1 from Pinar del Rio to 
Camagiiey provinces, Sample 3 from Oriente Province. 







s. 


notatus 






Sample 1 


Sample 2 


5. celicara 


n 




57 


118 


30 


Snout-vent length (mm) 


S 


20-30 


20-30 


25-32 






(25.8) 


(25.0) 


(28.7) 




9 


20-30 


20-32 


22-31 






(25.5) 


(25.1) 


(27.2) 


Dorsal scales 




18-28 


18-27 


21-30 






(21.6 ± 0.7) 


(22.1 ± 0.4) 


(25.8 ± 0.9) 


Midbody scales 




31^9 


33-47 


40-51 






(37.4 ± 1.4) 


(39.5 ± 0.6) 


(45.2 ± 1.2) 


4th toe lamellae 




7-11 


7-12 


9-13 






(9.3; M, = 10) 


(9.7; Mo = 10) 


(10.8; Mo = 11) 


Ventral keeling 










Throat 




none 


2 (weak), 6 distinct 


7 (absent in 23) 


Chest 




none 


1 (+ throat) 


none 


Ventrals 




none 


109 


none 



Comparisons. — Sphaerodactylus bromeliarum is similar in general appearance 
but is not lineate dorsally, has keeled ventral scales, has a prominently black- 
and-white banded tail tip in adults, and has smaller scales (midbody scales 56-58 
versus 40-51 in S. celicara). The only known adult (female) of S. bromeliarum 
has a SVL of 24 mm, much smaller than S. celicara. 

It is with S. notatus that S. celicara most requires comparison. We have 
grouped data on 57 specimens from Pinar del Rio to Camagiiey provinces (Sample 
1) and 118 specimens from Oriente Province (Sample 2) for the purposes of dis- 
cussion (Table 1, Fig. 2). Both samples of 5. notatus average smaller than both 
sexes of S. celicara in SVL. Samples 1 (21.6 ± 0.7) and 2 (22.1 ± 0.4) do not 
differ significantly in dorsal scale counts, but they do differ significantly from S. 
celicara (25.8 ± 0.9). The means of midbody scale counts in Samples 1 and 2 
(37.4 ±1.4 and 39.5 ± 0.6) differ significantly from each other and they are also 
significantly different from the mean in S. celicara (45.2 ± 1.2). There are fewer 
toe lamellae in the two samples of 5". notatus than in S. celicara (Table 1). 

Although the two samples of S. notatus as well as S. celicara modally have 1 
internal scale, the frequency of internasals (= nasals in contact) is much greater 
(35%) in S. celicara than in either sample (11% in each) of S. notatus. Throat, 
chest, and ventral keeling is summarized in Table 1. 

The head pattern of male S. notatus atactus changes with age from a juvenile 
(=female) trilineate pattern to a dotted one and finally to an unpatterned one. Six 
males (50%) have head dotting, four (33%) have the head somewhat dotted, and 
two (17%) lack head dotting. In Sample 2, 16 males (37%) have the head either 
dotted or trilineate, two (5%) have the head somewhat dotted, and 25 (58%) lack 
head dotting. Head dotting is present in seven (78%) of nine male S. celicara. In 
throat spotting. Sample 1 has six of 12 males (50%) with spotted throats. Sample 



VOLUME 95, NUMBER 2 397 

2 has nine of 35 males (20%) with the throat spotted, whereas seven of nine (78%) 
male S. celicara have the throat spotted. 

The female scapular pattern is absent in all 38 females in Sample 1 but occurs 
in 18 of 70 females (26%) in Sample 2. In S. celicara, 21 of 22 females (95%) have 
scapular patches and ocelH. 

Remarks. — We have no doubt that S. celicara is indeed a distinct biological 
entity. Whether it would be more appropriate to consider it a subspecies of S. 
notatus rather than a separate species is a problem. Sphaerodactylus notatus and 
S. celicara are not known to be sympatric, but the ranges of the two approach 
fairly closely. The area from which S. celicara is known is remote and difficult 
to reach, and the region between the known ranges of *S. celicara and S. notatus 
remains virtually unknown herpetologically. 

Most of the type-series of S. celicara has been taken in or near the Hotel 
Asuncion, on the walls and in the lobby. One male was secured in a curujey 
(Bromeliaceae) at La Maquina, and another was clinging to a rock. Like S. notatus, 
S. celicara apparently can adapt easily to edificarian situations, but the species 
may occupy a wide variety of niches. 

Etymology. — The name celicara is a noun in apposition derived from the Greek 
kelis, meaning spot or stain, and kara (an indeclinable neuter noun) meaning 
head, in allusion to the spotted heads in both sexes. 

Acknowledgments 

In addition to specimens in the Instituto de Zoologia (IZ) La Habana, and the 
Albert Schwartz Field Series (ASFS), we have examined material in the American 
Museum of Natural History (AMNH) and the Museum of Comparative Zoology 
(MCZ) through the cooperation of the curators and their assistants: Richard G. 
Zweifel, George W. Foley, Ernest E. Williams, and Jose P. Rosado. The senior 
author had assistance in the field from Luis de Armas, Lorenzo Zayas, Jorge de 
la Cruz, Luis R. Hernandez, Jose Espinosa, A. Vega, H. Sarazua, and Miguel 
L. Jaume. The senior author also wishes to extend thanks to Fernando Gonzalez, 
director of the Instituto de Zoologia, Academia de Ciencias de Cuba, for organ- 
izing the trip to Baracoa, and to the authorities of "Poder Popular" for providing 
transporation and accommodations in the environs of Baracoa. Jan Krescek sup- 
plied us with a transparency of an individual from Monte Iberia. The drawings 
of S. celicara are the work of Alvis Gineika, to whom we are also grateful. 

(OHG) Instituto de Zoologia, Academia de Ciencias, Capitolio Nacional, La 
Habana, Cuba; (AS) Miami-Dade Community College, North Campus, Miami, 
Florida 33167. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 398-402 

CYRTOCARA LIEMI, A PREVIOUSLY UNDESCRIBED 

PAEDOPHAGOUS CICHLID FISH (TELEOSTEI: 

CICHLIDAE) FROM LAKE MALAWI, AFRICA 

Kenneth R. McKaye and Catherine Mackenzie 

Abstract. — A new Lake Malawi paedophagous cichlid, Cyrtocara lie mi, is de- 
scribed. Cyrtocara lie mi is probably most closely related to C. orthognathus. 
The ecology and behavior of C. liemi is discussed briefly. 



The cichlid species flocks of the Great Lakes of East Africa are the most 
spectacular examples of speciation and adaptive radiation in any vertebrate fam- 
ily. Each of the lakes, Malawi, Tanganyika and Victoria, contains more species 
of fish than any lake outside the Rift Valley region. Lake Malawi has about 300 
described species of fish, 90% of which are cichlids, and 100-200 are believed 
stiU to be described (pers. obs.; Marsh et al. 1981). CichHds are weU known for 
their great diversity of feeding adaptations (Fryer and lies 1972) which allows 
them to exploit a wide array of foods, including fish eggs, fish embryos and fish 
larvae. The existence of paedophagous species, i.e. those which primarily exploit 
fish embryos and larvae, was first reported for the Lake Victoria cichlid flock 
(Greenwood 1959, 1967, 1974). At least three species of paedophagous cichlids 
occur in Lake Malawi, Cyrtocara orthognathus (Trewavas, 1935) and two un- 
described cichlids (McKaye and Kocher, in press). This report describes the one of 
those paedophagous species of which we have sufficient material. The counts and 
measurements employed follow Barel et al. (1977). 

Cyrtocara liemi, new species 
Fig. 1 _. 

Holotype. — National Museum of Natural History (USNM) 227497, adult male, 
175 mm standard length (SL) from W. Thumbi Island, Lake Malawi, Malawi 
(34°40'E, 14°01'S), collected by K. McKaye, T. Kocher and M. Oliver, Field No. 
MKO 80-49, July 1980. 

Paratypes. — USNM 227498 (4 specimens, 136.2 mm, 154.1 mm, 166.5 mm, 
175.2 mm SL, north side of W. Thumbi Island); 227499 (2 specimens 191.3 mm, 
174.2 mm SL, south side of W. Thumbi Island); 227500 (1 specimen, 178.3 mm 
SL, Chikale Beach, Nkhata Bay, 34°17'E ir35'S); 227501 (1 specimen, 124.3 mm 
SL, swamp east of Chembe village, Nankumba Peninsula, 34°5rE, 14°0rS); 
227502 (1 specimen, 131.8 mm SL); 227503 (1 specimen, 130.4 mm SL, Otter 
Point 34°48'E, 14°02'S); 227504 (2 specimens, 60.7 mm, 62.4 mm SL, south east 
corner of W. Thumbi Island). 

Diagnosis. — Cyrtocara liemi is similar to and probably most closely related to 
another paedophagous cichlid C. orthognathus (Fig. 2). The two species can be 
easily distinguished by the profile of the head, particularly gape inclination, which 
in C. liemi is 40-60° (i = 43.4) but in C. orthognathus is 60-80° {x = 66.6), and 



VOLUME 95, NUMBER 2 



399 




Fig. 1. Holotype of Cyrtocara liemi; USNM 227497. 

the premaxillary pedicel inclination, which ranges from 20-30° (f = 23.7) in C. 
liemi and 0-10° {x = 1.1) in C. orthognathus. Both species belong to a distinct 
group with a single oblique stripe within the genus Cyrtocara and are restricted 
to Lake Malawi. 

Description. — This description is based on the holotype (Fig. 1) and twelve 
paratypes, ten adults 124.3-191.3 mm SL and two juveniles 60.7 mm and 62.4 
mm SL. The principal morphometric ratios are given in Table 1. 

The portion of the head anterior to the interorbital region is concave in profile 
in larger specimens and nearly straight in smaller specimens. Premaxillary pedicel 
inclination 20-30°. Inclination of the dorsal head profile 28^0°. Snout 1.2-1.6 
times longer than broad. Cephalic laterosensory pores and canals not hypertro- 
phied. 

Posterior margin of maxilla does not extend to the vertical through the anterior 
margin of orbit. Jaws narrowly rounded when viewed from above. Lips not no- 
ticeably thickened. Lower jaw projects slightly. Gape inclination 40-60°. Lower 
jaw 1.15-1.98 times longer than broad. 





Fig. 2. Syntype of Cyrtocara orthognathus from British Museum; BM(NH) 1936.6.14. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Principal morphometric ratios of Cyrtocara liemi. % 
percent of standard length, Ma = mean of adult specimens, Mj 
Ra = range of adult specimens, Rj = range of juvenile specimens. 



percent of head length, * = 
mean of juvenile specimens. 



Character 


Ma 


Ra 


M, 


Ri 


Standard length mm 


157.9 


124.3-191.3 


61.6 


60.7-62.4 


Body depth * 


38.5 


36.1^2.1 


37.1 


34.6-39.5 


Head length mm 


51.3 


39.5-61.0 


22.1 


21.3- 


-22.9 


Head length * 


32.5 


31.4-33.8 


34.1 


32.4-35.9 


Head breadth % 


37.9 


36.4-40.3 


40.1 


39.3-40.8 


Interorbital width % 


21.0 


19.7-22.1 


19.3 


17.9- 


-20.7 


Snout length % 


37.1 


34.7-39.1 


31.1 


28.8- 


-33.3 


Orbit length % 


24.6 


22.5-26.5 


29.0 


28.2- 


-29.7 


Cheek depth % 


19.8 


17.0-22.0 


12.2 


12.2- 


-12.2 


Post-orbital head length % 


40.5 


38.2^2.9 


36.4 


36.1- 


-36.7 


Upper jaw length % 


34.4 


32.6-36.2 


33.5 


31.5- 


-35.4 


Premaxillary pedicels % 


27.0 


25.4-30.4 


31.5 


29.1- 


-33.2 


Lower jaw length % 


44.2 


42.7^5.9 


44.8 


43.2-46.3 


Predorsal length * 


33.8 


33.1-34.6 


36.4 


32.3-40.5 


Dorsal fin base * 


57.6 


53.9-59.1 


57.5 


55.5- 


-59.5 


Caudal peduncle length * 


15.3 


13.9-17.5 


16.7 


16.5- 


-16.8 



Caudal peduncle 13.9-16.8% SL. 1.15-1.5 times longer than deep. 

Dorsal fin with 27 (1), 28 (10) or 29 (2) elements, comprising 16 (1), 17 (10), 18 
(2), spines and 10 (2), 11 (9) or 12 (2) segmented rays; last spine 11.9-15.3% SL. 
Anal fin with 10 (1), 12 (3) or 13 (9) elements, comprising 3 spines and 7 (1), 9 (3) 
or 10 (9) segmented rays. Pectorals 27.8-32.5% SL comprising 11-14 segmented 
rays. Pelvic fins 23.9-31.1% SL, comprising 5 segmented rays. Caudal fin emar- 
ginate, lobes pointed, subequal, scaled over entire surface. 

Scales ctenoid. Lateral line with 31-34 pored lateral line scales (mode 31), 
upper part 23-26 (mode 25), lower part 6-9 (mode 8) Cheek with 3^ rows (mode 
3); uppermost scale often significantly larger. 5 to 7 scales between dorsal-fin 
origin and lateral line (mode 5). 7 to 9 scales (mode 8) between pectoral and 
pelvic-fin bases, scales gradually decreasing in size from pectoral fin to pelvic fin. 
Sixteen scales around caudal peduncle. 

Four gill rakers on epibranchial + 1 (in angle) + 10(1), 11 (11), 12(1) on cer- 
atobranchial. Anterior 4 on ceratobranchial often short, peg-like, remainder lon- 
ger, some club-like and bent laterally. 

Outer tooth row of upper jaw with a total of 42-60 teeth in adults; two juvenile 
specimens with 38 and 39. Outer rows of teeth of both upper and lower jaws 
composed of somewhat recurved, slightly movable teeth, embedded in fleshy 
gums. Teeth broad, long, and unequally bicuspid anteriorly, becoming shorter, 
more slender, and unicuspid posteriorly. Anterior teeth pale, distal portion rusty 
brown, color decreasing in extent and intensity in more posterior teeth; posterior- 
most teeth uniformly pale. In bicuspid teeth, lateral margin of major cusp convex, 
medial margin concave; inner margin of minor cusp more oblique than outer. 
Major cusp usually, but not always, anterior to minor cusp. Teeth of larger spec- 
imens uniformly unicuspid. Inner tooth rows irregular, 2-3 rows on upper and 
lower jaws, teeth unicuspid, occasionally weakly tricuspid with middle cusp ex- 
tended. 



VOLUME 95, NUMBER 2 



401 




Fig. 3. Lower pharyngeal bone of Cyrtocara liemi in occlusal view. 



Lower pharyngeal bone (n = 7) (Fig. 3) triangular in outline, 1.13-1.5 times 
broader than long, its length 10.9-28.4% of head length, breadth 57.1-72.7% of 
head breadth. Suture between two halves straight. Dentigerous area 1.2-1.6 times 
broader than long. Teeth variable. Postero-median teeth large but decreasing in 
size toward end of row, weakly bicuspid, with posterior cusp longest and colored 
rusty brown at tip and curved forward slightly. Lateral teeth compressed, bicus- 
pid or unicuspid, paler in color. Teeth closely packed posterolaterally, more 
openly packed medially, 29-35 teeth in posterior row, 5-11 in median row, 4-8 
in oblique rows, and 19-26 in lateral rows. 

Coloration in preservation. — Both sexes with head and dorsal surface dark 
grey-brown, paling posteriorly and ventrally. Sides with single oblique stripe, 
divided into 3 overlapping sections. Anterior section extending posteriorly to 
scale 13-18 of lateral line, beginning above the upper part of the lateral line and 
crossing over it at scale 7-8. Middle section extending from scale 7-9 to scale 
19-28, between upper and lower lateral line sections. Posterior section beginning 
at scale 16-19 and extending along upper side of lower lateral line to base of 
caudal fin where it dips slightly below the lower lateral line. Fins translucent 
brown-grey, with pale maculae. Juveniles pale yellow-brown. Oblique stripe com- 
posed of discontinuous segments. 10 vertical bars distinguishable. Fins colorless 
to grey, translucent. 

Distribution. — This species is not common, but it does occur in the Cape Ma- 
clear (Nankumba Peninsula) region of Lake Malawi as well as in Nkhata Bay. It 



402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

was caught and observed by SCUBA divers (McKaye and Kocher, in press) only 
in areas where brooding cichHd females congregated. 

Relationships . — An analysis of the phylogenetic relationship of C liemi must 
await revision of the genus Cyrtocara, which includes the group of obliquely- 
striped species of which the paedophages C. liemi and C. orthognathus are mem- 
bers. These fishes with a single oblique stripe have been grouped by Trewavas 
(1935) in her key and may represent a monophyletic group. Cyrtocara liemi and 
C. orthognathus occur sympatrically at Cape Maclear and at Nkhata Bay. Not 
only do they differ significantly in morphology, they also differ in behavior. 

Etymology. — -Named after Karel Liem in recognition of his pioneering studies 
and his insight into the feeding mechanism of cichlid fishes. 

Ecology. — Cyrtocara liemi is a paedophagous cichlid which acquires eggs, em- 
bryos, and fry from the mouths of mouthbrooding cichlids. Cyrtocara liemi at- 
tacks females from 0.5 to 2 m below and hits them in the hyoid region of the 
head. The only items found in their stomachs were eggs, embryos and larval 
cichlids. A more detailed description of their behavior and ecology along with 
two other paedophagous cichlids is discussed elsewhere (McKaye and Kocher, 
in press). 

Acknowledgments 

We thank L. Knapp and R. Vari of the Smithsonian Institution for their en- 
couragement and guidance, and T. Kocher, D. Lewis and M. Oliver for discussion 
of the material. The junior author wishes to thank L. Buss for his generosity and 
support throughout this project. We especially thank the government of Malawi 
for providing the facilities to make this research possible, and the Cape Maclear 
fisheries technicians, R. D. Makwinje, W. M. Menyani, and O. K. Mhone for 
help in making our collections. Financial support was from the National Science 
Foundation DEB 79-12338, Smithsonian Sorting Center, World Wildlife Fund 
U.S.A., and Peabody Museum, Yale University. 

Literature Cited 

Barel, C. D. N., M. J. P. van Oijen, F. Witte, and E. L. M. Witte-Maas. 1977. An introduction to 
the taxonomy and morphology of the haplochromine Cichlidae from Lake Victoria. — Nether- 
land Journal of Zoology 27(4):339-389. 

Fryer, G., and T. D. lies. 1972. The cichlid fishes of the Great Lakes of Africa. — Oliver & Boyd, 
Edinburg, 641 pp. 

Greenwood, P. H. 1959. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), 
Part 3. — Bulletin British Museum of Natural History (Zoology) 5:179-218. 

. 1967. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), Part 6. — 

Bulletin British Museum of Natural History (Zoology) 25:139-242. 

. 1974. The cichlid fishes of Lake Victoria, East Africa: The biology and evolution of a species 

flock. — Bulletin British Museum of Natural History (Zoology) Supplement 6:1-134. 

Marsh, A. C, A. J. Ribbink, and B. A. Marsh. 1981. Sibling species complexes in sympatric pop- 
ulations of Petrotilapia Trewavas (Cichlidae, Lake Malawi). — Journal Linnean Society (Zo- 
ology) 71:253-264. 

McKaye, K. R., and T. Kocher. Head ramming behavior in three paedophagous cichlids in Lake 
Malawi, Africa. — Animal Behavior (in press). 

Trewavas, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa. — Annals and Magazine of Natural 
History 16:65-118. 

(KRM) Duke University Marine Laboratory, Beaufort, North Carolina 28516, 
and (CM) Ecologia, I.N.P.A. EP 478, Manaus 69000, Brazil. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 403-407 

HYPSELOGNATHUS HORRIDUS, A NEW SPECIES OF 
PIPEFISH (SYNGNATHIDAE) FROM SOUTH AUSTRALIA 

C. E. Dawson and C. J. M. Glover 

Abstract. — An unusual pipefish, characterized by spinigerous head and body 
surfaces, absence of scutella, and presence of caudal fin and caudal brood pouch, 
is described from depths of 42-55 m off Anxious Bay, South Australia. Provi- 
sionally referred to the Australian endemic genus Hypselognathus, this species 
differs from its only congener {H. rostratus) in having spiny rather than essentially 
smooth head and body surfaces, in lacking scutella, in having more total subdorsal 
rings (11-11.5 versus 8-9.5) and a higher HL in SL ratio (8.0-9.0 versus 5.9-6.6), 
as well as in other morphological, proportional and meristic features. 



Current studies on Australian pipefishes indicate the presence of about 90 valid 
species and several yet undescribed forms. Most of the latter have similar or 
closely related congeners and descriptions are delayed pending collection of more 
comparative material. Specimens reported here differ so strikingly from all known 
pipefishes that we see no need for postponing their description. The occurrence 
of this species and the recently described Kimblaeus bassensis Dawson (1980) in 
moderate depths (40-70 m) of southern Australian coasts suggests that continued 
sampling in this region may educe further additions to the pipefish fauna of Aus- 
tralia. 

Measurements are, in part, referred to standard length (SL) and head length 
(HL); counts of the holotype are marked *; as employed here, the term "venter" 
refers to the ventral surface of head or body; other methods are those of Dawson 
(1977). Specimens are deposited in the South Australian Museum (SAM) and Gulf 
Coast Research Laboratory Museum (GCRL). Drawings are by Mrs. Yasue Mat- 
thews (GCRL). 

Hypselognathus horridus, new species 
Figs. 1 and 2 

Holotype.— SAM F.4676 (154.0 mm SL, adult male). South Australia, Great 
Australian Bight, off Anxious Bay, 54.9 m (30 fm.), trawl, 25 Feb. 1981, P. C. 
Halsey. 

Paratypes. — GCRL 18057 (226.5 mm SL, adult female), South Australia, Great 
Australian Bight, ca. 32°24'S, 133°30'E, 42 m, trawl, 5 May 1973, SA Fish. Dept. 
SAM F.4681 (228.0 mm SL, adult female). South Australia, Great Australian 
Bight, ca. 33°24'S, 134°37'E, 14.8 km S of Point Weyland, 54.9 m, trawl, 18 Sept. 
1981, KarlOlsen. 

Diagnosis. — Head and body surfaces (except membranes) spinigerous, scutella 
absent, HL 8.0-9.0 in SL, snout length 1.6-1.8 in HL, total rings 70-72. 

Description.— Rings 27* + 43-45*, subdorsal rings 6.75-6.0* + 4.25-5.5* = 
11.0-11.5*, dorsal-fin rays 31*, pectoral-fin rays 10* (2), 11 (2), 12 (2), anal-fin 
rays 3*, caudal-fin rays 10*. Measurements of holotype are followed (in paren- 



404 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




'l£^^3^s;^ 







/!r^ •-,'.: 




^rp 



•'/r /r //// // .// // // // // //////// 



^// I; II II II II II I II / // // // // /////// // V / -., 



f^" " I I ! 

'/tin J JL-Lji,. 



ij II II II II II II 



! II II II II II 



'I II I II I 1 II I II II 1/ // '/ // //7^ 

! ! / " "' ! II II " II II if if // // /////^ 
' / , / I I I II a II II II / // / ///■ 



// //' 



'' yt. 








Fig. \. Hypselognathus horridus. Upper pair: Lateral and dorsal aspects of head and anterior 
trunk rings. Lower pair, Top: Posterior trunk and anterior tail rings, together with dorsal and anal 
fins and anterior portion of brood pouch. Bottom: Posterior tail rings and caudal fin. From holotype 
(SAM F.4676). 



theses) by those of shorter and longer paratypes, respectively: SL 154.0 (226.5, 
228), HL 17.1 (28.5, 28.4), snout length 9.3 (17.3, 17.3), least snout depth 2.1 
(3.4, 3.2), length of dorsal-fin base 16.4 (24.3, 24.9), anal ring depth 2.6 (4.5, 4.2), 
trunk depth 3.2 (6.8, — ), pectoral-fin length 3.2 (4.2, — ), length of pectoral-fin 
base 1.6 (2.9, 2.8), caudal-fin length 7.3 (— , 11.1). 

Median dorsal snout ridge (Fig. 1) low, irregularly denticulate to spinulose, 
ending just before vertical through anterior margin of orbit, not confluent with 
anterior continuations of supraorbital ridges; snout compressed laterally, its least 
depth 4.4-5.4 in snout length; nares 2-pored bilaterally, the anterior pore on a 



VOLUME 95, NUMBER 2 



405 




Fig. 2. Hypselognathus horridus. Top and middle: SAM F.4676 (154 mm SL, male, holotype). 
Bottom: GCRL 18057 (226.5 mm SL, female, paratype). 



short tubule; preorbital bone rather narrow, its width less than diameter of pupil. 
Opercle with radiating denticulate striae, none clearly enlarged or ridge-like; dor- 
sal margins of orbits elevated slightly; interorbital essentially flat, a little de- 
pressed between dorsal margins of orbits; dorsum of head more or less rounded 
behind orbits; nuchal ridge obsolete; pre nuchal and frontal ridges vestigial; pec- 
toral-fin base protruding a little laterad, without obvious ridges. 

Superior trunk and tail ridges discontinuous near rear of dorsal-fin base; lateral 
trunk and tail ridges discontinuous, terminating on last trunk ring; inferior trunk 
and tail ridges continuous (Fig. 1). Principal body ridges distinct, somewhat el- 
evated and angled a little laterad on posterior 16-20 tail rings; ridge margins 
irregularly denticulate, usually with a slightly enlarged hook-like spine on pos- 
terior third of each ring; anterior and posterior margins of rings straight to irreg- 
ularly emarginate; scutella absent. Most surfaces (except eye, gill membranes, 
fins and pouch folds) more or less covered with minute spines, their distribution 
irregular but numbers somewhat reduced on posterior tail rings. Dorsum a little 
convex on trunk and anterior half of tail, gradually becoming flat and depressed 
between somewhat elevated superior ridges on posterior tail rings; venter of trunk 
V-shaped, the median ridge not clearly enlarged; 12th- 19th trunk rings of holo- 
type swollen or enlarged in dorsal and lateral aspects (Fig. 2); llth-27th trunk 
rings of adult females deeper (ca. 22%) than preceding rings; sides and venter of 
posterior tail rings more or less flat, depressed distally between elevated principal 
ridges. 



406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Dorsal-fin base not elevated; superior trunk ridge not arched strongly dorsad 
below dorsal-fin base; pectoral fin more or less rounded; caudal-fin membranes 
broad above and below, narrower between the long median fin-rays; tips of cau- 
dal-fin rays somewhat flattened and spatulate. 

Brood pouch developed below the anterior 18 tail rings of the holotype; pouch 
plates moderately enlarged, angled somewhat laterad; membranous pouch folds 
present. Pouch closure, uncertain due to absence of eggs or larvae, probably the 
everted type of Herald (1959). 

Color of holotype in alcohol mainly grayish, shading to brown on distal half of 
tail. Sides and dorsum of head and body, and venter of tail behind brood pouch, 
peppered with brown to black microchromatophores, somewhat larger on sides 
and most abundant on posterior half of tail; anterior and posterior margins of 
rings often with fine dark edging. Lower portions of gill membranes, gular mem- 
brane, and venter of trunk mainly pale. Brood-pouch folds pale, with moderately 
broad, irregular, brownish shading along mesial margins. Caudal-fin rays largely 
brownish, the membranes shaded diffusely with brown; other fins hyaline. Col- 
oration similar in a recently preserved female paratype (SAM F.4681) but the 
venter of the deepened posterior portion of the trunk is shaded with pink. 

Etymology. — Named horridus, from the Latin, meaning bristly or rough. 

Relationships. — The combination of spinigerous surfaces, absence of scutella, 
presence of caudal fin and a brood pouch, with plates and folds, located under 
the tail distinguishes the species described here from all known pipefishes. Its 
relationships are unclear and it may eventually warrant separate generic treat- 
ment. In general morphology, this fish appears most closely related to the poorly 
defined Hypselognathus Whitley (1948) and it is provisionally referred to this 
otherwise monotypic, Australian endemic, genus (type species Histiogamphelus 
rostratus Waite and Hale, 1921) until additional material is available for study. 

Comparisons. — Hypselognathus rostratus and H. horridus agree in the config- 
uration of principal body ridges (Fig. 1), in the presence of dorsal, pectoral, anal 
and 10-rayed caudal fins, and both share an elongate, laterally compressed, snout. 
In general agreement with H. horridus, specimens of H. rostratus may have a 
small spine or vestige thereof on margins of principal ridges of trunk rings, prin- 
cipal ridges of the posterior tail rings are somewhat elevated and angled laterad, 
and some swollen trunk rings are typically present. However, H. horridus lacks 
the small round scutella present in H. rostratus, fails to have the supraorbital 
ridges confluent with the median dorsal snout ridge (confluent in rostratus), and 
has spinigerous head and inter-ridge body surfaces (smooth in rostratus). Com- 
pared to 8 specimens of H. rostratus (152-305 mm SL), H. horridus has more 
trunk rings and total rings (respectively, 27 and 70-72 versus 24-25 and 65-69), 
fewer pectoral-fin rays (10-12 versus 12-14), more subdorsal trunk rings and total 
subdorsal rings (respectively, 6.75-6 and 11-11.5 versus 2.5-0.75 and 8-9.5), a 
higher HL in SL ratio (8.0-9.0 versus 5.9-6.6), and a lower snout depth in snout 
length ratio (4.4-5.4 versus 6.2-13.0). 

Among other pipefishes, spinigerous head and body surfaces, somewhat similar 
to those of H. horridus, occur only in some species of Solegnathus Swainson 
(e.g., S. spinosissimus Giinther). However, species of Solegnathus differ from 
Hypselognathus horridus in lacking the caudal fin and brood-pouch plates and 
folds and in having well-developed scutella. 



VOLUME 95, NUMBER 2 407 

Remarks. — Each of the three known specimens of Hypselognathus horridus 
was taken with one or more specimens of Solegnathus robustus McCuUoch. 

Literature Cited 

Dawson, C. E. 1977. Review of the pipefish genus Corythoichthys with description of three new 
species.— Copeia 1977:295-338. 

. 1980. Kimblaeus, a new pipefish genus (Syngnathiformes: Syngnathidae) from Australia, with 

a key to genera of pipefishes with continuous superior ridges. — Australian Journal of Marine 
and Freshwater Research 31:517-523. 

Herald, E. S. 1959. From pipefish to seahorse — a study of phylogenetic relationships. — Proceedings 
of the California Academy of Sciences 42:181-227. 

Waite, E. R., and H. M. Hale. 1921. Review of the lophobranchiate fishes (Pipe-fishes and Sea- 
horses) of South Australia. — Records of the South Australian Museum 1:293-324. 

Whitley, G. P. 1948. Studies in ichthyology. No. 13. — Records of the Australian Museum 22:70-94. 

(CED) Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi 
39564; (CJMG) South Austrahan Museum, Adelaide, S.A. 5000, Australia. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 408-411 

LATERAL LINE OF DIPLOSPINUS MULTISTRIATUS 
(TELEOSTEL GEMPYLIDAE) 

Izumi Nakamura 

Abstract. — The existence of the lateral line in Diplospinus multistriatus has 
been questioned. The lateral line of D. multistriatus is described here and com- 
pared with that of closely related Paradiplospinus gracilis. Most of the (upper) 
lateral line of both species runs nearer the ventral contour of the body than the 
dorsal contour. This type of lateral-line conformation is found in some trichiurid 
genera, but not in other gempylid genera. Lateral-line scales of both species are 
tube-shaped with an irregular longitudinal slit. This type of lateral-line scale is 
not found in any other genera of Gempylidae or Trichiuridae. 



Since Diplospinus multistriatus was described by Maul (1948) from Madeira, 
considerable numbers of specimens have been collected widely in tropical and 
temperate waters of the Atlantic, Pacific and Indian oceans (Parin and Bekker 
1972:161, Fig. 17). Maul (1948) studied the holotype with six paratypes and wrote, 
"the skin is smooth and there is no trace of a lateral line." Later, Tucker (1956) 
examined one of the paratypes and reported, "there are traces of an apparent 
and highly probable lateral line," but he did not mention this further. No other 
authors have paid special attention to the lateral line of D. multistriatus. 

While examining specimens of D. multistriatus at the National Museum of 
Natural History (USNM), Smithsonian Institution, traces of a lateral line were 
recognized. As there are no undamaged specimens of either D. multistriatus or 
Paradiplospinus gracilis, these observations are based on many specimens com- 
pounded to reconstruct the lateral line. 

Material examined. — Diplospinus multistriatus Maul: USNM 215428, 2 speci- 
mens (16.5, 55.2 mm SL), 21°30'N, 158°30'W, 19 Jan. 1970; USNM 215449, 6 
(42.1-188.4), 2r30'N, 158°30'W, 23 Sep. 1970; USNM 100492, 1 (48.0), 29°00'N, 
76°23'W, 28 Feb. 1914; USNM 226993, 3 (98.2-162.6), 27°45'N, 9ri8.5'W, 23 
Feb. 1964; USNM 215397-8, 3 (134.7-203.0), 2r30'N, 158°30'W, 28 Feb. 1971; 
USNM 215395, 1 (141.0), 21°30'N, 158°30'W, 17 Sep. 1970; USNM 215433, 2 
(142.5, 152.4), 21°30'N, 158°30'W, 16 Sep. 1970; USNM 194458, 1 (175.1), 
29°40'N, 69°05'W, 29 Mar. 1957; USNM 219967, 2 (175.6, 213.3), 33°04'N, 
39°29'W, 27 Apr. 1979; USNM 215391, 1 (188.4), 2r30'N, 158°30'W, 15 Sep. 
1970; USNM 215394, 1 (203.8), 2r30'N, 158°30'W, 14 Dec. 1970; USNM 215454, 
1 (232.8), 2r30'N, 158°30'W, 18 Sep. 1971. 

Paradiplospinus gracilis (Brauer): USNM 208104, 1 specimen (47.7 mm SL), 
49°06'S, 120°15'W, 19 Dec. 1965; USNM 208446, 9 (141.0-335.0), 40°18'S, 
39°04'W, 8 Mar. 1971; USNM 208448, 6 (161.7-320.0), 39°47'S, 43°38'W, 7 Mar. 
1971; USNM 208449, 9 (182.0-349.0), 38°20'S, 54°33'W, 5 Mar. 1971; USNM 
226992, 1 (355.0), 40°08'S, 82°47'W, 2 Oct. 1966. 

Identification 

Diplospinus multistriatus and Paradiplospinus gracilis are similar in external 
appearance. Specimens obtained by usual collecting methods are almost always 



VOLUME 95, NUMBER 2 



409 



Table 1 . — Some distinguishing characters (counts compared with those of some authors) of Diplo- 
spinus multistriatus and Paradiplospinus gracilis. 

Characters 



z 



Species 



Diplospinus multistriatus 



Paradiplospinus gracilis 



Position of anus 



Anal fin 

Lateral line 
Dorsal-fin rays 



Anal-fin rays 



Vertebrae number 



midway between tip of snout and tip 
of caudal fin; in front of first anal 
spine by distance equal to head 
length (Fig. lA) 

anterior part very low, with almost no 
fin membrane (Fig. lA) 

easily removed; double ? (Fig. lA) 

XXXI-XXXIV, 35-^0 (this study) 
XXX-XXXIV, 36-42 (Parin and 

Bekker 1972) 
XXXI-XXXIII, 37-41 (Karrer 1975) 
XXXII-XXXVI, 37^1 (Parin et al. 

1978) 
XXXII-XXXIII, 40 (Tucker 1956) 

II, 28-33 (this study) 

II, 29-32 (Parin and Bekker 1972) 

II, 28-34 (Karrer 1975) 

II, 28-33 (Parin et al. 1978) 

22-24 -f 34-37 = 57-61 (this study) 

59-61 (Karrer 1975) 

34 + 24 = 58 (Tucker 1956) 



nearer tip of caudal fin than to tip of 
snout; in front of first anal spine by 
distance equal to snout length (Fig. 
IB) 

anterior part fairly high, with fin 

membrane (Fig. IB) 
usually intact; single (Fig. IB) 

XXXVI-XXXIX, 28-33 (this study) 
XXXVI-XXXIX, 28-33 (Parin and 

Bekker 1972) 
XXXVI-XXXVII, 28-30 (Karrer 1975) 
XXXVI-XXXVII, 28-32 (Bussing 

1965) 
XXXVI-XXXVII, 28-32 (DeWitt and 

Hureau 1979) 

II, 26-31 (this study) 

II, 25-30 (Parin and Bekker 1972) 

II, 26-28 (Karrer 1975) 

II, 25-28 (Bussing 1965) 

IL 25-28 (DeWitt and Hureau 1979) 

32-34 + 32-34 = 65-66 (this study) 
61 (Karrer 1975) 

63-67 (DeWitt and Hureau 1979) 
63-66 (Bussing 1965) 
38^0 + 26-27 = 65-66 (Andriashev 
1960) 



more or less damaged and the vertical-fin rays are folded with damaged fin mem- 
branes. Some distinguishing characters of the species are summarized in Table 
1. Useful characters for distinguishing the species are the number of dorsal spines 
and the number of dorsal soft rays (the total number of both is not useful) and 
the position of the anus. The number of anal-fin rays overlaps, so it is not a very 
useful character. The shapes of the anal fin and the lateral line are often hard to 
see. The number of vertebrae (total) clearly differentiates the species. 



Lateral Line 

The lateral line of D. multistriatus is easily removed, but careful examination 
with a microscope can find remains somewhere on the bodies of even fairly 
damaged specimens. The remains are found most often on the shoulder region 
and next most often on the mid-portion of the body. The lateral-line system based 
on several specimens compounded is shown in Fig. lA. The upper lateral line is 
recognized certainly. The lower lateral line was discerned only partly in three out 
of 24 specimens examined, and it is not certain whether the lower lateral line 
connects with the upper lateral line or where its anterior and posterior termina- 
tions may be. From the upper margin of the opercle the upper lateral fine of/). 
multistriatus descends gently to below the middle of the body at the anus, there- 
after running near the ventral contour of the body to the caudal region. The shape 



410 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L A, A reconstruction of the lateral-line system of Diplospinus multistriatus based on several 
specimens; B, A reconstruction of the lateral-line system of Paradiplospinus gracilis based on several 
specimens. Vertical broken lines show the position of the anus, a-g shows the details of lateral-line 
scales, a-d from the positions shown in D. multistriatus (A) and e-g from P. gracilis (B). a-c: USNM 
215454, 232.8 mm SL; d: USNM 215428, 55.2 mm SL; e: USNM 208446, 255.5 mm SL; f: USNM 
226992, 355.0 mm SL; g: USNM 208449, 190.5 mm SL. Scales indicate 1 mm. 



of the lateral-line scales is slightly different in various parts of the body as shown 
in Fig. la-d. Each scale is basically tube-shaped with an irregular longitudinal 
external slit. 

The lateral line of P. gracilis is conspicuous, tough, and not usually lost. Most 
of the lateral line remains intact in even fairly damaged specimens. The lateral- 
line system, based on several specimens, is shown in Fig. IB. The lateral line of 
P. gracilis is single, descending gently from the upper margin of the opercle to 
slightly below the middle of the body at the anus, thereafter running slightly 
nearer the ventral contour of the body to the caudal region (Fig. IB). The shape 
of the lateral-line scales is somewhat different in various parts of the body, as 
shown in Fig. le-g. Each scale is basically a short tube with an irregular external 
slit. Scales in the middle of the lateral line (Fig. le) are much more elongate than 
those in the anterior and posterior parts (Fig. If, g). Lateral-line scales are gen- 
erally stouter in P. gracilis than in D. multistriatus. 

The course of the (upper) lateral line is similar in both species. Careful obser- 
vation, however, reveals that most of the (upper) lateral line is situated only 
slightly below mid-body in P. gracilis and far below mid-body in D. multistriatus. 
Excluding the lower lateral line of D. multistriatus from consideration, similar 
lateral lines running below mid-body are found in some trichiurid genera, such 
as Trichiurus, Lepturacanthus , Eupleurogrammus and Tentoriceps, but not in 
any other gempylid genera, which have the single lateral line or the upper part 
of a double lateral line running nearer to the dorsal countour than the ventral 
contour. This may suggest a close relationship of gempylids and trichiurids. 



VOLUME 95, NUMBER 2 411 

The basic structure of the lateral-line scales of D. multistriatus and P. gracilis 
are similar. This type of lateral-line scale (tube-shaped scale with an irregular 
longitudinal external slit which has relatively small openings at both ends of each 
scale) is not found in any other genera of Gempylidae and Trichiuridae. 

The lateral line seems to become fully formed at about 50 mm SL in both D. 
multistriatus and P. gracilis. The size at which the lateral-line development starts 
could not be ascertained, though some larval materials of both species were 
examined in this study. Specimens of D. multistriatus about 20 mm SL do not 
show any traces of the lateral line (Strasburg 1964; Yevseyenko and Serebryakov 
1973) and a specimen of P. gracilis 32.2 mm SL does not show any traces of it 
(Bussing 1965). 

Acknowledgments 

I am grateful to the Smithsonian Institution for providing me a postdoctoral 
fellowship in 1981, which made it possible for me to do this study. Thanks are 
also due to my supervisor, Robert H. Gibbs, Jr. and Joseph L. Russo of NMFS 
Systematics Laboratory for their critical review. Technical assistance rendered 
by Reiko Nakamura is acknowledged. 

Literature Cited 

Andriashev, A. P. 1960. Families of fishes new to the Antarctic. I. Paradiplospinus antarcticus gen. 
et sp. (Pisces, Trichiuridae). — Zoologicheskii Zhurnal 39(2): 244-249 (in Russian). 

Bussing, W. A. 1965. Studids of the midwater fishes of the Peru-Chile Trench. — American Geo- 
physical Union, Antarctic Research Series 5:185-227. 

DeWitt, H. H., and J.-C. Hureau. 1979. Fishes collected during "Hero" Cruise 72-2 in the Palmer 
Archipelago, Antarctica, with the description of two new genera and three new species. — 
Bulletin du Museum National d'Histoire Naturelle, Paris 4*^ serie 1:775-820. 

Karrer, C. 1975. Uber Fische aus dem Siidostatlantik (Teil 2). — Mitteilungen aus dem Zoologischen 
Museum in Berlin 51(l):63-82. 

Maul, G. E. 1948. Quatro peixes novos dos mares da Madeira. — Boletim do Museum Municipal do 
Funchal (3) Art. 6:41-55 (in Portuguese and Enghsh). 

Parin, N. V., and V. E. Bekker. 1972. Materials on taxonomy and distribution of some trichiuroid 
fishes (Pisces, Trichiuroidae: Scombrolabracidae, Gempylidae, Trichiuridae). — Transactions 
of the P. P. Shirshov Institute of Oceanography 93:110-204 (in Russian). 

, Y. I. Sazonov, and S. V. Mikhailin. 1978. Glubokovodnye Pelagicheskie Ryby V Clorkh Ips 

"Fiolent" V Gvineiskom Zalive i Prilegayoshchikh Raionakh. — Trudy Instituta Okeanologii 
im P. P. Shirshova 3:169-183 (Translated from Russian for the Smithsonian Institution and the 
National Science Foundation, Washington, D. C. by Esduck Cairo: Deep-water pelagic fishes 
in the collections of R/V "Fiolent" in the Gulf of Guinea and adjacent regions. 1981). 

Strasburg, D. W. 1964. Postlarval scombroid fishes of the genera Acanthocybium, Nealotus, and 
Diplospinus from the Central Pacific Ocean. — Pacific Science 18(2): 174-185. 

Tucker, D. W. 1956. Studies on the trichiuroid fishes — 3. A preHminary revision of the family Trichi- 
uridae. — Bulletin of the British Museum (Natural History) Zoology 4(3):73-130. 

Yevseyenko, S. A., and V. P. Serebryakov. 1973. Larvae oi Diplospinus multistriatus (Pisces, Gem- 
pylidae) from the northwestern Atlantic. — Journal of Ichthyology 14(l):92-98. 

Fisheries Research Station, Kyoto University, Maizuru, Kyoto 625, Japan, and 
Division of Fishes, Department of Vertebrate Zoology, National Museum of Nat- 
ural History, Smithsonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

95(2), 1982, pp. 412-420 

VARIATION OF A MUSCLE IN HUMMINGBIRDS AND 
SWIFTS AND ITS SYSTEMATIC IMPLICATIONS 

Richard L. Zusi and Gregory Dean Bentz 

Abstract. — Historically, features of the muscle tensor propatagiaHs pars brevis 
have been used to argue that swifts and hummingbirds comprise a single mono- 
phyletic order, the Apodiformes, and that this order is most closely related to the 
"pico-passeriforms." We describe variations of this muscle in swifts and hum- 
mingbirds as well as in other orders and conclude that the new evidence does not 
support these claims. The variations observed, however, do show morphological 
trends that help to clarify relationships within both swifts and hummingbirds. 



The phylogenetic relationships of hummingbirds and swifts to each other and 
to other avian orders are among the major unsolved ornithological problems in 
systematics. Subfamilial and generic relationships within swifts are fairly well 
understood (see Brooke 1970), but internal relationships of hummingbirds are still 
poorly known. Although we do not claim to have solved any of these problems, 
we have found that variation in a single muscle, M. tensor patagii brevis, bears 
on all of them. We report this variation and its systematic impHcations as a 
stimulus for further study of these problems, and as partial evidence toward their 
eventual solution. 

The tensor patagii brevis muscle (TPB) extends from the shoulder to the fore- 
arm in birds and apparently can serve either to flex the forearm or to support the 
prepatagial membrane of the extended wing. This muscle exhibits marked vari- 
ation among birds, especially in relation to its tendon or tendons of insertion. 
Garrod (1876) drew attention to this variation and to its taxonomic implications, 
and Fiirbringer (1888) devoted six plates and considerable discussion to this mus- 
cle. Since then it has been further described in major systematic and anatomical 
works (e.g. Buri 1900; Beddard 1898; George and Berger 1966). In this paper we 
describe in some detail the variation of TPB in swifts and hummingbirds, and we 
test Lowe's (1939:329) contentions that the fleshy beUy and tendon of insertion 
of TPB in hummingbirds is almost identical with that of the swift, that both are 
fundamentally passerine in design, and that the arrangement in non-passerines is 
quite different. 

Hummingbirds 

In hummingbirds the origin of TPB is consistently by a tendon from the head 
of the coracoid. The nearly parallel fibers of the short, wide belly pass distally 
and end on the surface of the extensor metacarpi radialis muscle (EMR) and on 
a short internal aponeurosis that fuses with the aponeurosis of origin of EMR. At 
this point of fusion the aponeurosis of TPB forms a tendon (the humeral tendon) 
that extends across the belly of EMR and inserts on the humerus. In humming- 
birds another aponeurosis or tendon (the distal tendon) passes distally from the 



VOLUME 95, NUMBER 2 413 

same point of fusion along the surface of EMR toward the wrist. Within hum- 
mingbirds, four different patterns of insertion exist. 

Type 1 (Fig. If) is characteristic of the Phaethornithinae or hermits (see Ap- 
pendix for species examined). In these forms the humeral tendon of TPB is par- 
tially covered by fibers of EMR that arise from the process of origin of that 
muscle. The humeral tendon of TPB and the aponeurosis of origin of EMR are 
tightly bound together where they cross within the belly of EMR, thus forming 
a firm base for the attachment of overlying muscle fibers. The humeral tendon of 
TPB emerges from the belly of EMR, passes superficial to the distal-most fibers 
of origin of EMR and inserts on a tubercle of the humerus distal to the process 
of EMR. (Such terms as ectepicondylar process, lateral epicondyle, and dorsal 
supracondylar process are sometimes used for the points of attachment of EMR 
and TPB on the humerus. Until a better understanding of the homologies of 
projections on the distal end of the humerus is obtained throughout birds we 
prefer to avoid these terms and refer simply to the process of EMR and the 
tubercle of TPB.) 

The distal "tendon" of TPB in hermits is so transparent that we initially con- 
sidered it absent. Closer examination revealed a short tendon from the junction 
of the bellies of TPB and EMR that quickly fans out into a broad aponeurotic 
sheet and invests most of the distal half of the belly of EMR. Proximally it is free 
of a superficial dorsal aponeurosis of EMR, but distally the two aponeuroses fuse. 

Eutoxeres is the only phaethornithine exception to the Type 1 format. In this 
genus the humeral tendon of TPB is visible for its entire length. Although not 
covered by fibers of EMR, it serves as a point of origin for some of the superficial 
fibers of that muscle. The humeral tendon and the aponeurosis of origin of EMR 
are fused where they cross as in other hermits. 

Type 2 (Fig. Ig) occurs in some of the Trochilinae: Anthracothorax, Doryfera, 
Androdon, Eulampis, Sericotes, Chrysolampis, Polytmus, Topaza, Heliothryx, 
Colibri, and Florisuga. In these forms the humeral tendon of TPB is clearly 
visible on the surface of the proximal end of EMR. Unlike Type 1 the hunieral 
tendon is essentially free of the aponeurosis of origin of EMR, attaching only to 
its cranial edge. The insertion of the humeral tendon is on a tubercle of the 
humerus. As in other trochiline hummingbirds a distinct distal tendon of TPB is 
formed. This tendon broadens distally and fuses with the dorsal surface of a 
superficial aponeurosis of EMR that contributes to its tendon of insertion near 
the wrist. This distal tendon of TPB lies on the dorsal surface of EMR so that the 
tendon appears to divide the belly of EMR unequally into cranial (smaller) and 
caudal (larger) portions. This is not the case with Type 3 birds. 

Type 3 (Fig. Ih) is found in all trochiUne genera listed in the Appendix except 
those of Types 2 and 4. The distal tendon of TPB is well developed and, at least 
proximally, free of the belly of EMR. It passes along the cranial edge of the belly 
of EMR rather than along the dorsal surface. Only near the distal end of the 
radius does it fuse with the craniodorsal portion of the tendon of insertion of 
EMR. The relations of the humeral tendon of TPB are like Type 2. This is the 
type illustrated by Garrod (1876). 

Type 4 (Fig. li) is restricted to trochiline hummingbirds of the genera Aces- 
trura, Chaetocercus, Calliphlox, and possibly others. Here the relations of the 



414 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




a 




TPB 



EMR 



PPB 





g 



h 





Fig. L Diagrams showing M. tensor propatagialis pars brevis and M. extensor metacarpi 
radialis (right, dorsal view): a, Piciformes and some Passeriformes; b, Most Passeriformes (dotted 
outHne represents Hirundinidae and Lonchurae); c, Hemiprocnidae (Type A); d, Cypseloidinae (Type 
B); e, Apodinae (Type C); f, Trochilidae (Type 1); g, Trochilidae (Type 2); h, Trochilidae (Type 3); i, 
Trochilidae (Type 4). EMR = M. extensor metacarpi radialis; PPB = M. pectoralis pars propatagialis 
brevis; TPB = M. tensor propatagialis pars brevis. Long arrow indicates humeral tendon of TPB; short 
arrow indicates distal tendon of TPB. 



distal tendon of TPB are similar to Type 3 , but a difference exists in the humeral 
tendon. In these forms, that tendon lies directly superficial to and almost parallel 
with the aponeurosis of origin of EMR, to which it is firmly fused. The short 
tendon of TPB inserts on the process of origin of EMR rather than on a separate 
tubercle. 



VOLUME 95, NUMBER 2 415 

Swifts 

The condition of TPB in swifts is significantly different from that of humming- 
birds. According to Cohn (1968) the muscle arises from the head of the coracoid 
and from the adjacent dorsal arm of the furcula in larger swifts, and only from 
the coracoid in some smaller swifts. Furthermore, M. pectoralis pars propatagialis 
longus and M. pectoralis pars propatagialis brevis are present in swifts but neither 
is present in hummingbirds. In swifts the latter muscle is represented by a short 
tendon that extends from M. pectoralis near the deltoid crest to the tendon of 
insertion of TPB. In those instances where the tendon of insertion of TPB is 
absent, the propatagialis brevis joins TPB at its juncture with EMR. In all of the 
swifts we examined there was no trace of a distal tendon of TPB. Within the 
swifts the TPB differs in other ways as well, and we recognize three basic types. 

Type A (Fig. Ic) is found in the crested swifts (Hemiprocnidae). In these forms 
the belly of TPB ends on a short tendon that receives the pectoralis pars pro- 
patagialis brevis tendon and continues to the cranial surface of EMR. The humeral 
tendon then passes across the surface of EMR to insert on a tubercle of the 
humerus. 

Type B (Fig. Id) is found in the Cypseloidinae. It is similar to Type A except 
that the belly of TPB reaches the surface of EMR without first forming a tendon 
of insertion. Instead, the belly tapers to a narrow, semitendinous insertion on 
EMR. At this juncture pars propatagialis brevis attaches and a well-defined hu- 
meral tendon of TPB arises and crosses EMR to insert on a tubercle of the 
humerus. 

Type C (Fig. le) is seen in the Apodinae (sensu Morony et al. 1975; includes 
CoUocaliini, Chaeturini, and Apodini). Type C resembles Type B except that the 
belly of TPB makes broad contact with EMR rather than tapering to its insertion. 
In addition, the humeral tendon of TPB almost parallels that of EMR and inserts 
directly on the process of origin of EMR, much like Type 4 in hummingbirds. 
Hirundapus giganteus and H. caudacutus are somewhat different. In these 
species the humeral tendon of TPB attaches slightly distal to the process of EMR 
or on its distal edge. However, the extent of separation between that tendon 
and the aponeurosis of origin of EMR is not nearly as great as it is in Types A 
and B. 

The use oi Hirundapus giganteus to illustrate the TPB of swifts by Lowe (1939) 
was unfortunate because that species is atypical of either subfamily. Although 
used to support the presence of a humeral tendon of TPB in swifts, it probably 
represents a stage in the loss of that tendon. 

Discussion 

Separate humeral tendon of TPB. — The contention that passerine birds differ 
from most other birds in having a humeral tendon of TPB that is separate from 
rather than fused with the aponeurosis of origin of EMR (Fig. la, b) originated 
with Garrod (1876). Lowe (1939) used this feature to ally swifts and hummingbirds 
with the Passeriformes (which he broadened to include also the Capitonidae, 
Indicatoridae, and Picidae). Not all pas serif orms (sensu strictu) have a separate 
tendon; we found it fused in Eurylaimidae and Rhinocryptidae. Garrod (1876) 
said it was fused in Menuridae and Atrichornithidae but Raikow (pers. comm.) 
found it separate in Atrichornis clamosus. In the Caprimulgidae we found the 



416 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






► 







g 



Fig. 2. Selected humeri (left, caudal view) showing positions of the process of M. extensor meta- 
carpi radialis (opposite black triangle) and of tubercle of M. tensor propatagialis pars brevis (white 
triangle): a, Coccothraustes vespertinus; b, Cordeiles minor; c, Hemiprocne comata; d, Hemiprocne 
longipennis; e, Nephoecetes niger; f, Hirundapus caudacutus; g, Apus apus; h, Glaucis hirsuta; i, 
Coeligena wilsoni; j, Chaetocercus jourdanii. Some sizes have been adjusted to make them roughly 
comparable, but all are not drawn to same humerus length because that measure varies greatly in 
relation to body size. 



tendon to be fused in Caprimulgus but completely separate in Cordeiles. Thus 
separate tendons, while probably derived within birds, have evidently evolved 
independently in at least the Caprimulgidae and Passeriformes. 

Comparison among passerines, caprimulgids, crested swifts, and swifts strong- 
ly suggests that the degree of separation of the humeral attachments of TPB and 
EMR results mainly from proximal displacement of EMR. The humeral attach- 
ment of TPB also moves proximally (in an evolutionary sense), but it lags behind, 
creating a gap between the attachments. Eventually it may reach the level of the 
process of EMR. Such stages can be represented, as in Fig. 2, by: a, a passerine; 
b, Cordeiles', c, Hemiprocne comata and mystacea; d, H. longipennis; e, Cyp- 
seloidinae; f, Apodinae (Hirundapus); and g, all other Apodinae. In this morpho- 
logical series, separate tendons in b-e appear to be derived relative to the fused 
tendons of most non-passerines, and the fused tendons of f and g derived relative 
to the separate tendons of b-e. A somewhat parallel trend occurs in hummingbirds 



VOLUME 95, NUMBER 2 417 

although there is considerable varation among closely related species. Types 1 
and 2 tend to have the process of EMR and the tubercle of TPB widely separated, 
while in Type 3 they are usually closer together (Fig. 2h and i). In hummingbirds, 
as in swifts, loss or fusion of the humeral tendon of TPB (Type 4) probably 
represents a derived condition (Fig. 2j). These differences are caused mainly by 
proximal migration of the TPB tubercle because the process of EMR is located 
far proximally in all hummingbirds. (Any quantitative comparison of the proximal 
shift of these processes should compare their distance from the distal end of the 
humerus to some measure of body size rather than to humeral length because the 
humerus itself has become relatively shorter, to differing degrees, in the evolution 
of swifts and hummingbirds.) 

Attachment of TPB on EMR. — The broad attachment of the belly of TPB on 
that of EMR in both swifts and hummingbirds is stated to be unique to those 
families and is used to support their placement in the order Apodiformes. How- 
ever, other birds show various approaches to the condition in swifts and hum- 
mingbirds. Several passerines have only a short tendon between the bellies of 
TPB and EMR (swallows, pers. obs.; many Lonchurae [Bentz 1979]). In Colius 
the fleshy belly of TPB extends distally as far as EMR on the dorsal surfaces of 
two tendons, and in some pigeons the belly of TPB attaches on a broad tendon 
almost to EMR. Thus we see that an approach toward attachment of the belly of 
TPB on EMR has been made in at least three orders apart from swifts and hum- 
mingbirds. 

Within the Apodidae a broad fleshy attachment is characteristic of the Apod- 
inae, as described by Lowe (1939), who dissected only members of that subfamily. 
The Cypseloidinae, however, have a narrow and tendinous attachment of TPB 
(noted by us in Cypseloides, Nephoecetes, and Streptoprocne) and in the Hem- 
iprocnidae there is a short tendon. Thus, if we assume that the presence of one 
or more tendons between the bellies of TPB and EMR as found in most birds 
was the ancestral condition for swifts, we have a morphological series within the 
Apodiformes from primitive (tendon) to intermediate (tapered semitendinous bel- 
ly) to advanced (broad attachment of the befly). 

All hummingbirds display a broad attachment resembling that of the most spe- 
cialized swifts. In the morphology of the humeral tendon, however, the more 
primitive stages in hummingbirds resemble the stage seen in the less specialized 
swifts. Unless the ancestral hummingbirds underwent a reversal (from stage g to 
h in Fig. 2) of the evolutionary trend in the humeral tendon seen in swifts, fol- 
lowed by another reversal to parallel the trend in swifts (compare h-j and e-g in 
Fig. 2), we conclude that the resemblance of the belly of TPB in hummingbirds 
and the Apodinae is the result of either parallel or convergent evolution. 

Distal tendon of TPB. — The early illustration of Patagona gigas in Garrod 
(1876) and subsequent work based largely on dissections of trochiline humming- 
birds of Type 3 (Beddard 1898; Cohn 1968) give the impression that the presence 
of a distal tendon of TPB is found in all hummingbirds and is unique to that 
family. We have seen that the "tendon" is a diffuse, expansive, and essentially 
transparent sheet in hermits. This structure might have become stronger and 
better defined (as in Type 2), and further specialized into a strong tendon (Types 
3 and 4) as an expression of its increasing role in supplementing extension of the 
hand by EMR. We think it unlikely that evolution proceeded in the opposite 
direction because reduction of a well-defined distal tendon without a change in 



418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

its function would probably result in loss of the tendon rather than in production 
of a broadly investing, diaphanous sheet. We hypothesize that Types 1, 2, and 
3 represent progressive evolutionary stages in the specialization of the distal 
tendon of TPB. 

Systematic conclusions. — The direct connection of the belly of TPB with that 
of EMR in swifts and hummingbirds proves to represent an autapomorphy in 
each group. Whether these apomorphies were derived from a common ancestor 
(in which a tendon was present) or arose twice in different lineages cannot be 
determined from this muscle alone. Thus this aspect of the muscle gives no 
definite evidence for monophyly of the Apodiformes. 

The alleged passerine nature of the humeral tendon of TPB in swifts and hum- 
mingbirds must be discounted because some nonpasserines have a similar form 
of the tendon. Furthermore, separation of the humeral tendons of TPB and EMR 
is related to a proximal shift in position of origin of EMR, which has occurred in 
unrelated orders or families. Once again, this feature of TPB is not by itself a 
good indicator of the ordinal affinities of either swifts or hummingbirds. 

Within the Apodidae, our interpretation of the evolution of specialization of 
the belly and of the humeral tendon of TPB corresponds with Brooke's (1970) 
concept of a more primitive subfamily, Cypseloidinae, and a more advanced 
Apodinae. Our data are not useful in determining whether the Collocaliini, Chae- 
turini, and Apodini should be regarded as tribes of a single subfamily (Brooke 
1970) or as two subfamilies (ColHns 1976). Unfortunately, Brooke followed Lowe 
(1939) in regarding the Hemiprocnidae as more nearly passerine, and therefore 
more advanced, than the Apodidae. The characters used by Lowe are, in fact, 
found in many other orders besides the Passeriformes, and are thus primitive 
relative to the corresponding derived states in the Apodidae. The Hemiprocnidae 
should precede rather than follow the Apodidae in a linear classification. 

Within the hummingbirds, our interpretation of the variation in TPB would 
support a hypothesis of a primitive phaethornithine group (hermits) and a more 
advanced trochiline group. Within the trochilines those genera with a dorsal distal 
tendon (Type 2) are more likely to be primitive (see Appendix). The presence of 
superficial muscle fibers of EMR covering part of the humeral tendon of TPB in 
most hermits (Fig. If) might best be regarded as a specialization of an otherwise 
primitive condition in hermits. 

Acknowledgments 

We thank David M. Niles for the loan of specimens from the Delaware Museum 
of Natural History, and Edwin O. WilHs for providing us with anatomical spec- 
imens of Ramphodon naevius. This study was partially funded by Smithsonian 
Research Award SQ3372400 to R. L. Zusi. 

Literature Cited 

Beddard, F. E. 1898. The structure and classification of birds. — Longmans, Green and Company, 

London, 548 pp. 
Bentz, G. D. 1979. The appendicular myology and phylogenetic relationships of the Ploceidae and 

Estrildidae (Aves: Passeriformes). — Bulletin of Carnegie Museum of Natural History 15:1-25. 
Brooke, R. K. 1970. Taxonomic and evolutionary notes on the subfamilies, tribes, genera, and 

subgenera of the swifts (Aves: Apodidae). — Durban Museum Novitates 9(2): 13-24. 
Buri, R. O. 1900. Zur Anatomic des Fliigels von Micropus melba und einigen anderen Coracomithes, 



VOLUME 95, NUMBER 2 419 

zugleich Beitrag zur Kenntnis der systematischen Steilung der Cypselidae. — Jenaische Zeit- 

schrift fiir Naturwissenschaft 33:361-610. 
Cohn, J. 1968. The convergent flight mechanism of swifts (Apodi) and hummingbirds (Trochili) 

(Aves). — Unpublished Ph.D. Dissertation, University of Michigan, Ann Arbor, Michigan. 
Collins, C. T. 1976. A review of the Lower Miocene swifts (Aves: Apodidae). In S. L. Olson, ed. 

Collected Papers in Avian Paleontology Honoring the 90th Birthday of Alexander Wetmore. — 

Smithsonian Contributions to Paleobiology No. 27. 
Fiirbringer, M. 1888. Untersuchungen zur Morphologic und Systematik der Vogel, zugleich ein Bei- 
trag zur Anatomic der Stiitz- und Bewegungsorgane. — 2 vols. — Van Holkema, Amsterdam, 

1751 pp. 
Garrod, A. H. 1876. On some anatomical peculiarities which bear upon the major division of the 

passerine birds, Part 1. — Proceedings of the Zoological Society of London: 506-5 19. 
George, J. C, and A. J. Berger. 1966. Avian myology.— Academic Press, New York and London, 

xii + 500 pp. 
Lowe, P. R. 1939. On the systematic position of the swifts (Suborder Cypseli) and hummingbirds 

(Suborder Trochili), with special reference to the Order Passeriformes. — Transactions of the 

Zoological Society of London 24:307-349. 
Morony, J., W. Bock, and J. Farrand, Jr. 1975. Reference list of the birds of the world. — American 

Museum of Natural History, New York, 207 pp. 

(RLZ) National Museum of Natural History, Smithsonian Institution, Wash- 
ington, D.C. 20560; (GDB) Mount Vernon College, Washington, D.C. 20007. 

Appendix 

Listed below are the species dissected by us for M. tensor propatagialis pars 
brevis and M. extensor metacarpi radialis. We follow the terminology of Morony 
et al. (1975). 

Trochilidae 

Type 1. — Ramphodon naevius, Glaucis hirsuta, Threnetes ruckeri, Phaethornis yaruqui, P. malaris, 
P. ruber, P. longuemareus, Eutoxeres aquila, E. condamini. 

Type 2. — Androdon aequatorialis, Doryfera johannae , D. ludovicae, Florisuga mellivora, Colibri 
delphinae, C. thalassinus, C. coruscans, Anthracothorax nigricollis, A. dominicus, A. viridis, A. 
mango, Eulampis jugularis, Sericotes holosericeus, Chrysolampis mosquitus, Polytmus guainumbi, 
Topaza pella, T. pyra, Heliothrix barroti. 

Type 3. — Campylopterus duidae, Orthorhyncus cristatus, Stephanoxis lalandi, Lophornis ornata, 
L. pavonina, Popelaria sp., Chlorostilbon swainsonii, C. maugaeus, Cynanthus latirostris, Cyano- 
phaia bicolor, Thalurania furcata, Panterpe insignis, Damophila Julie, Lepidopyga coeruleogularis, 
Hylocharis chrysura, Chrysuronia oenone, Goldmania violiceps, Trochilus polytmus, Leucochloris 
albicollis, Leucippus fallax, Amazilia amabilis, A. viridigaster, A. tzacatl, Elvira cupreiceps, Chal- 
ybura buffonii, Lampornis castaneoventris, Adelomyia melanogenys, Heliodoxa rubinoides, H. xan- 
thogonys, H. branickii, Eugenes fulgens, Sternoclyta cyanopectus, Oreotrochilus estella, Patagona 
gigas, Aglaeactis cupripennis, Lafresnaya lafresnayi, Pterophanes cyanopterus, Coeligena coeligena, 
C. violifer, Ensifera ensifera, Sephanoides sephanoides, Heliangelus amethysticollis, Eriocnemus 
luciani, Haplophaedia aureliae, H. lugens, Ocreatus underwoodii, Lesbia victoriae, Sappho spar- 
ganura, Metallura tyrianthina, Aglaiocercus kingi, Heliomaster longirostris, Philodice evelynae, Cal- 
othorax lucifer, Archilochus alexandri, Mellisuga minima, Calypte costae, Stellula calliope, Myrtis 
fanny, Selasphorus rufus. 

Type 4. — Calliphlox amethystina, Acestrura mulsant, Chaetocercus jourdanii. 

Hemiprocnidae and Apodidae 

Type A. — Hemiprocne comata. 

Type B. — Cypseloidinae: Cypseloides rutilus, Nephoecetes niger, Streptoprocne zonaris, S. semi- 
collaris. 

Type C. — Apodinae: Collocalia brevirostris, C. whiteheadi, C. esculenta, Hirundapus caudacutus, 



420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

H. giganteus, Chaetura martinica, C. pelagica, Aeronautes montivagus, A. saxatilis, Tachornis 
phoenicobia, T. squamata, Cypsiurus parvus, Apus pallidas, A. pacificus. 

Other families 

Columbidae: Columba nigrirostris, Columbina minuta; Cuculidae: Coccyzus erythropthalmus; Po- 
dargidae: Batrachostomus septimus; Nyctibiidae: Nyctibius griseus; Caprimulgidae: Chordeiles pu- 
sillus, Caprimulgus europaeus', Coiiidae: Colius striatus; Trogonidae: Trogon viridis; Alcedinidae: 
Chloroceryle aenea; Todidae: Todus mexicanus; Momotidae: Electron platyrhynchum; Meropidae: 
Merops apiaster; Coraciidae: Coracias garrulus; Upupidae: Upupa epops; Phoeniculidae: Phoenicu- 
lus purpureas; Bucerotidae: Tockus erythrorhynchus; Galbulidae: Galbula ruficauda; Bucconidae: 
Chelidoptera tenebrosa; Capitonidae: Megalaima haemacephala; Indicatoridae: Indicator archipe- 
lagicus; Ramphastidae: Baillonius bailloni; Picidae: Jynx torquilla, Melanerpes striatus; Eurylaimi- 
dae: Smithornis capensis, Eurylaimus ochromalus, Calyptomena whiteheadi; Furnariidae: Certhiaxis 
subcristata; Formicariidae: Thamnophilus punctatus , Myrmotherula hauxwelli; Rhinocryptidae: Sce- 
lorchilus rubecula, Scytalopus latebricola; Cotingidae: Pachyramphus cinnamomeus; Pipridae: 
Chiroxiphia linearis; Tyrannidae: Ochthoeca rufipectoralis, Tyrannus dominicensis, Todirostrum 
cinereum, Sublegatus arenarum; Alaudidae: Lullula arborea; Hirundinidae: Progne dominicensis, 
Riparia riparia; Laniidae: Lanius cristatus; Muscicapidae: Niltava grandis; Nectariniidae: Nectarinia 
jugularis; Meliphagidae: Melidectes fuscus; Parulidae: Basileuterus coronatus; Sturnidae: Acridoth- 
eres tristis; Corvidae: Garrulus glandarius . 



ERRATA FOR VOLUME 94(4) 



page 1229 (Table 2, line 4) 
"Second maxilliped," should read "Second maxilliped exopod," 

page 1229 (Table 2, line 6) 

"Third maxilliped," should read "Third maxilliped exopod," 

page 1231 (Table 3, line 10) 

"Second maxilliped," should read "Second maxilliped exopod," 

page 1231 (Table 3, line 12) 

"Third maxilliped," should read "Third maxilliped exopod," 



PROC. BIOL. SOC. WASH. 

95(2), 1982, p. 421 



JASCOTTELLA, NOM. NOV. FOR MAMILLA 

SCOTT, 1974 (MICROPROBLEMATICA) 

NON FABRICIUS, 1823 (MOLLUSC A) 

Richard W. Huddleston and Drew Haman 



Scott (1974) erected the genus Mamilla (type-species M. hemispherica Scott, 
1974) for problematic microfossils of possible foraminiferal affinity from the Has- 
1am Formation, Vancouver Island. 

The generic name Mamilla previously had been used twice, Mamilla Fabricius, 
1823 and Mamilla Wagner, 1907, both in the Mollusca. The use of this generic 
name has priority in Fabricius (1823). Baker (1954) replaced Mamilla Wagner, 
1907 {non Fabricius, 1823) with Weinlandella. Mamilla Scott, however, remains 
a junior homonym and requires a new name (International Code of Zoological 
Nomenclature, Article 53). The new name Jascottella is proposed to replace 
Mamilla Scott, non Fabricius. (Repeated attempts to contact Dr. Scott in con- 
formity with ICZN Appendix A, 3 have been unsuccessful.) 

Jascottella is constructed in honor of J. A. Scott for his recognition of this new 
genus. 

Acknowledgments 

We thank Chevron Oil Field Research Company for their assistance and per- 
mission to publish. 

Literature Cited 

Baker, H. B. 1954. New subgeneric names in Helicinidae. — The Nautilus 67:139-140. 
Fabricius, O. 1823. Pp. 51-114 in Fortegnelse over afg. Biskop Fabriciusses eiterladte Naturalier. 
Scott, J. A. B. 1974. The Foraminifera of the Haslam, Qualicum, and Trent river Formations, Van- 
couver Island, British Columbia. — Bulletin of Canadian Petroleum Geology 22(2): 119-176. 

Chevron Oil Field Research Company, P.O. Box 446, La Habra, California 
90631. 



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CONTENTS 

Atrophecaecum lobacetabulare, n. sp. (Digenea: Cryptogonimidae: Acanthostominae) with 
discussion of the generic status of Paracanthostomum Fischthal and Kuntz, 1965, and 
Ateuchocephala Coil and Kuntz, 1960 Daniel R. Brooks and Janine N. Caira 223 

A partial revision of the genus