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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 102 
1989 



Vol. 102(1) published 29 March 1989 
Vol. 102(2) published 28 June 1989 



Vol. 102(3) published 18 October 1989 
Vol. 102(4) published 19 December 1989 



WASHINGTON 
PRINTED FOR THE SOCIETY 



EDITOR 
C. Brian Robbins 



ASSOCIATE EDITORS 

Classical Languages Invertebrates 

George C. Steyskal Stephen D. Cairns 

Frank D. Ferrari 
Raymond B. Manning 

Plants Vertebrates 

David B. Lellinger G. David Johnson 

Insects 
Wayne N. Mathis 



All correspondence should be addressed to the 

Biological Society of Washington, Smithsonian Institution 

Washington, D.C. 20560 



Allen Press Inc. 
Lawrence, Kansas 66044 



OFFICERS AND COUNCIL 

of the 

BIOLOGICAL SOCIETY OF WASHINGTON 

FOR 1988-1989 



OFFICERS 

President 
KRISTIAN FAUCHALD 

President-Elect 
LESLIE W. KNAPP 

Secretary 
G. DAVID JOHNSON 

Treasurer 
DON E. WILSON 



COUNCIL 

Elected Members 
GARY R. GRAVES MEREDITH L. JONES 

W. RONALD HEYER RAYMOND B. MANNING 

W. DUANE HOPE WAYNE N. MATHIS 



TABLE OF CONTENTS 

Volume 102 

Alvarez, Fernando. Smalleyus tricristatus, new genus, new species, and Pseudothelphusa 
parabelliana, new species (Brachyura: Pseudothelphusidae) from Los Tuxtlas, Vera- 
cruz, Mexico 45-49 

Anderson, William D., Jr. and Phillip C. Heemstra. Ellerkeldia, a junior synonym of 
Hypoplectrod.es, with redescription of the type species of the genera (Pisces: Serranidae: 
Anthiinae) 1001-1017 

Barnard, J. L. Rectification of Halirages regis and H. huxleyanus (Crustacea: Amphip- 
oda), from marine Antarctica, with description of a new genus, Austroregia 701-715 

Barnard, J. L. and James Darwin Thomas. Four species of Synopiidae from the Carib- 
bean region (Crustacea: Amphipoda) 362-374 

Barnard, J. L. and James Darwin Thomas. A new species, Ampelisca burkei, (Crustacea, 
Amphipoda) from Florida 375-384 

Barr, Douglas J. and Susumu Ohtsuka. Pseudocyclops lepidotus, a new species of demersal 
copepod (Calanoida: Pseudocyclopidae) from the northwestern Pacific 331-338 

Bieri, Robert. Krohnittellidae and Bathybelidae, new families in the Phylum Chaetog- 
natha; the rejection of the family Tokiokaispadellidae and the generea Tokiokaispa- 
della, Zahonya, and Aberrospadella 973-976 

Boeger, Walter A., Delane C. Kritsky, and Joaber Pereira, Jr. Neotrpical Monogena. 14. 
Revision of Callorhynchocotyle Suriano and Incorvaia, 1982 (Hexabothriidae) with 
the description of C. amatoi 264-27 1 

Bowman, Thomas E. The high frequency of errors in format in manuscripts submitted 
to Proceedings of the Biological Society of Washington 1067-1068 

Bowman, Thomas E. and Robert Bieri. Paraspadella anops, new species, from Sagittarius 
Cave, Grand Bahama Island, the second troglobitic chaetognath 586-589 

Brinkhurst, Ralph O. Varichaetradrilus angustipenis (Brinkhurst and Cook, 1966), new 
combination for Limnodrilus angustipenis (Oligochaeta: Tubificidae) 311-312 

Brownell, Robert L., Jr. and James G. Mead. Taxonomic status of the delphinid (Mam- 
malia: Cetacea) Tursiol panope Philippi, 1895 532-534 

Browning, M. Ralph. The type specimens of Hekstra's owls 515-519 

Bullock, Robert C. Rhyssoplax baliensis, a new species of chiton from Indonesia (Mol- 
lusca: Polyplacophora: Chitonidae) 808-8 1 9 

Campbell, Jonathan A. A new species of colubrid snake of the genus Coniophanes from 
the highlands of Chiapas, Mexico 1 036-1 044 

Campbell, Jonathan A., William W. Lamar, and David M. Hillis. A new species of 
diminutive Eleutherodactylus (Leptodactylidae) from Oaxaca, Mexico 491-499 

Carvacho, Alberto. Cancer johngarthi, n. sp. and Cancer porteri (Bell) (Crustacea, De- 
capoda): comparisons and hypothesis 6 1 3-6 1 9 

Chace, Fenner A., Jr. The holotype of Heterocarpus alexandri A. Milne-Edwards (Crus- 
tacea: Decapoda: Pandalidae) 84-88 

Child, C. Allan. Pycnogonida of the Western Pacific Islands VI. Sericosura cochleifovea, 
a new hydrothermal vent species from the Marianas back-arc basin 732-737 

Chirino-Galvez, Luis A. and Raymond B. Manning. A new deep-sea crab of the genus 

Chaceon from Chile (Crustacea, Decapoda, Geryonidae) 401-404 

Colbath, G. Kent. Revision of the family Lysaretidae, and recognition of the family 
Oenonidae Kinberg, 1865 (Eunicida: Polychaeta) 116-123 

Colbath, G. Kent. A revision of Arabella mutans (Chamberlin, 1919) and related species 
(Polychaeta: Arabellidae) 283-299 

Cutler, Edward B. and Norma J. Cutler. A revision of the genus Aspidosiphon (Sipuncula: 
Aspidosiphonidae) 826-865 

Desbruyeres, Daniel and Lucien Laubier. Paralvinella hessleri, new species of Alvinel- 
lidae (Polychaeta) from the Mariana back-arc basin hydrothermal vents 761-767 



Erseus, Christer. Phallodrilus hessleri, new species (Oligochaeta: Tubificidae), from abys- 
sal depths in the western Pacific Ocean 131-133 

Erseus, Christer. Phallodrilus vescus, new species (Oligochaeta, Tubificidae) from the 
Gulf of Mexico _ 134-136 

Erseus, Christer. Four new west Atlantic species of Tubificoides (Oligochaeta, 
Tubificidae) 877-886 

Erseus, Christer and Michael R. Milligan. Three new species of Tubificidae (Oligochaeta) 
from an oil seepage area on the continental slope of the northern Gulf of Mexico 887-893 

Etnier, David A. and James D. Williams. Etheostoma {Nothonotus) wapiti (Osteichthyes: 
Percidae), a new darter from the southern bend of the Tennessee River system in 
Alabama and Tennessee 987-1000 

Fauchald, Kristian. The Second Annual Riser Lecture: Eclecticism and the study of 
Polychaetes _ 742-752 

Fautin, Daphne G. and Robert R. Hessler. Marianactis bythios, a new genus and species 
of actinostolid sea anemone (Coelenterata: Actiniaria) from the Mariana vents 815-825 

Fitzpack, J. F., Jr. and Craig A. Busack. Hobbseus yalobushensis, a new crawfish from 
central Mississippi (Decapoda: Cambaridae) 637-643 

Flint, Oliver S., Jr. and D. G. Denning. Studies of neotropical caddisflies, XL: new 
species of Smicridea (Smicridea) from middle America and the West Indies (Tri- 
choptera: Hydropsychidae) _ 418^433 

Formas, J. Ramon. A new species of Eupsophus (Amphibia: Anura: Leptodactylidae) 
from southern Chile 568-576 

Froeschner, Richard C. A small collection of Heteroptera from the Galapagos Islands, 
with the description of the new species Niesthrea ashlocki and a list of Niesthrea 
species (Rhopalidae) 609-612 

Froeschner, Richard C. Three new species of Colombian lace bugs of the genera Le- 
podictya and Leptopharsa (Heteroptera: Tingidae) 968-972 

Gardner, Alfred L. and G. Ken Creighton. A new generic name for Tate's (1933) Mi- 
crotarsus group of South American mouse opossums (Marsupialia: Didelphidae) 3-7 

Giere, Olav. The First Annual Riser Lecture: Meiofauna and Microbes— the interactive 
relations of annelid hosts with their symbiotic bacteria 109-1 15 

Gillet, Patrick. Axiothella crozetensis, a new species of maldanid polychaete from Crozet 
Islands (Indian Ocean) 8 6 6-8 6 9 

Goeke, Gary D. Stenorhynchus yangi, a new western Atlantic species of Arrow Crab 
(Crustacea, Brachyura, Majidae) and a redescription of S. seticornis 620-636 

Grabe, Stephen A. Some aspects of the biology of Rhopalophthalmus tatersallae Pillai, 
1961 (Crustacea, Mysidacea) and extension of range into the Khor Al Sabiya, Kuwait 
(Arabian Gulf) 726-731 

Grygier, Mark J. Three new species of Myzostoma (Myzostomida) 793-804 

Hardy, Laurence M. The karyotype of Exilliboa placata Bogert (Tropidopheidae), and 
comparisons with the family Boidae (Reptilia: Serpentes) 1045-1049 

Hensley , Dannie A. and Kunio Amaoka. A redescription of Pseudorhombus megalops, 
with comments on Cephalopsetta ventrocellata (Osteichthyes: Pleuronectiformes: 
Paralychthyidae) 577-585 

Hershler, Robert. Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa Riv- 
er (exclusive of Ash Meadows) drainages, Death Valley System, California-Nevada.. 176-248 

Heyer, W. Ronald and Alan J. Wolf. Physalaemus crombiei (Amphibia: Leptodactyli- 
dae), a new frog species from Espirito Santo, Brazil with comments on the P. signifer 
group 500-506 

Hobbs, Horton H., Jr. and Daniel J. Peters. New records of entocytherid ostracods 
infesting burrowing crayfishes, with the description of a new species, Ascetocythere 
stockeri...„ 324-330 

Hobbs, Horton H., Jr. and Henry W. Robison. On the crayfish genus Fallicambarus 
(Decapoda: Cambaridae) in Arkansas, with notes on the fodiens complex and descrip- 
tions of two new species 651-697 

Holsinger, John R. Allocrangonyctidae and Pseudocrangonyctidae, two new families of 



holarctic subterranean amphipod crustaceans (Gammaridea), with comments on their 

phylogenetic and zoogeographic relationships 947-959 

Holt, Perry C. A new species of the cambarincolid genus Sathodrilus from Missouri, 

with the proposal of a replacement name for Adenodrilus Holt, 1977 (Clitellata: 

Branchiobdellida) 738-741 

Humes, Arthur G. Acontiophorus excavatus, new species (Copepoda: Siphonostoma- 

toida) associated with the soft coral Dendronephthya (Alcyonacea) in the Indo- 

Pacific 916-923 

Jara, Carlos G. Aegla denticulata lacustris, new subspecies, from Lake Rupanco, Chile 

(Crustacea: Decapoda: Anomura: Aeglidae) 385-393 

Johnson, G. David and Edward B. Brothers. Acanthemblemaria paula, a new diminutive 

chaenopsid (Pisces: Blennioidei) from Belize, with comments on life history 1018-1030 

Kensley, Brian. New genera in the thalassinidean families Calocarididae and Axiidae... 960-967 

Keppner, Edwin J. Four new species of free-living marine nematodes in the genus 

Pareurystomina (Nematoda: Enoplida) with observations on other members of the 

genus 249-263 

Kornicker, Louis S. The adult male of the troglobitic ostracode Spelaeoecia bermudensis 

Angel and Iliffe, 1987, from an anchialine cave in Bermuda (Crustacea: Ostracoda: 

Halocypridoidea) 3 1 3-3 2 3 

Kornicker, Louis and Thomas M. Iliffe. Dantya ferox, a new species of mydocopid 

ostracode from Niue, central South Pacific (Crustacea: Ostracoda: Sarsiellidae) 901-91 1 

Lowry, James K. and Gary C. B. Poore. First ingolfiellids from the southwest Pacific 

(Crustacea: Amphipoda) with a discussion of their systematics 933-946 

Manning, Raymond B. Sanquerus, a replacement name for Posidon Herklots, 1851 

(Crustacea, Decapoda, Portunidae) 698-700 

Manning, Raymond B. and David K. Camp. Additional records for an Atlantic reef 

lobster, Enoplometopus antillensis Lutken, 1865 (Crustacea, Decapoda, 

Enoplometopidae) 411-417 

Manning, Raymond B. and L. B. Holthuis. Two new genera and nine new species of 

geryonid crabs (Crustacea, Decapoda, Geryonidae) 50-77 

Manning, Raymond B., Marcos Siqueira Taveres, and Elaine Figueiredo Albuquerque. 

Chaceon ramosae, a new deep-water crab from Brazil (Crustacea: Decapoda: 

Geryonidae) 646-650 

Martin, Joel W. and Denton Belk. Eulimnadia ovilunata and E. ovisimilis, new species 

of clam shrimps (Crustacea, Branchiopoda, Spinicaudata) from South America 894-900 

Mathis, Wayne N. A review of the beach flies of the Caribbean and Gulf of Mexico 

(Diptera: Canacidae) 590-608 

Mathis, Wayne N. and Jin Zuyin. A review of the shore-fly genus Polytrichophora 

Cresson from Asia (Diptera: Ephydridae) 434-446 

McCranie, James R., Jay M. Savage, and Larry David Wilson. Description of two new 

species of the Eleutherodactylus milesi group (Amphibia: Anura: Leptodactylidae) 

from northern Honduras 483—490 

McLelland, Jerry A. Eukrohnia calliops, a new species of Chaetognatha from the northern 

Gulf of Mexico with notes on related species 33—44 

Miller, John E. and David L. Pawson. Hansenothuria benti, new genus, new species 

(Echinodermata: Holothuroidea) from the troptical western Atlantic: a bathyal, epi- 

benthic holothurian with swimming abilities 977-986 

Miller, Scott E. Paleontological type specimens in the Santa Barbara Museum of Natural 

History 1-2 

Monniot, Claude and Francoise Monniot. Ascidians collected around the Galapagos 

Islands using the Johnson- Sea-Link research submersible 14-32 

Munroe, Thomas A. and Madhu N. Mahadeva. Symphurus callopterus (Cynoglossidae, 

Pleuronectiformes), a new deepwater tonguefish from the eastern Pacific 458-467 

Ohwada, Takashi. Redescription of Nephtys squamosa Ehlers (Polychaeta: 

Nephtyidae) 124-130 

Olson, Storrs L. Two overlooked holotypes of the Hawaiian flycatcher Chasiempis 

described by Leonhard Stejneger (Aves: Myagrinae) 555-558 



Ortiz, Juan Carlos, Hector Ibarra- Vidal, and J. Ramon Formas. A new species of Eupso- 
phus (Anura: Leptodactylidae) from Contulmo, Nahuelbuta Range, southern Chile... 1031-1035 

Ota, Hidetoshi and Ronald I. Crombie. A new lizard of the genus Lepidodactylus (Rep- 
tilia: Gekkonidae) from Batan Island, Philippines 559-567 

Perkins, Philip D. Adelphydraena, new genus, and two new species from Venezuela, and 
remarks on phylogenetic relationships within the subtribe Hydraenina (Coleoptera: 
Hydraenidae) 447-457 

Pettibone, Marian H. New species of scale-worms (Polychaeta: Polynoidae) from the 
hydrothermal rift-area of the Mariana back-arc basin in the western central Pacific... 137-153 

Pettibone, Marian H. Polynoidae and Sigalionidae (Polychaeta) from the Guaymas 
Basin, with descriptions of two new species, and additional records from hydrothermal 
vents of the Galapagos Rift, 21°N, and seep-sites in the Gulf of Mexico (Florida and 
Louisiana) 154-168 

Pettibone, Marian H. A new species of Benhamipolynoe (Polychaeta: Polynoidae: Lep- 
idastheniinae) from Australia, associated with the unattached stylasterid coral Co- 
nopora adeta 300-304 

Pettibone, Marian H. Two new species of Harmothoinae (Polychaeta: Polynoidae) from 
the East Pacific Rise, collected by Alvin dives 2000 and 2003 305-310 

Rickart, Eric A., Lawrence R. Heaney, and Mark J. Rosenfeld. Chromosomes of ten 
species of Philippine fruit bats (Chiroptera: Pteropodidae) 520-531 

Rodriguez, Gilberto and Horton H. Hobbs, Jr. Freshwater crabs associated with caves 
in southern Mexico and Belize, with descriptions of three new species (Crustacea: 
Decapoda) 394-400 

Romero, Raul Castro and Hernan Baeza Kuroki. Neobrachiella anisotremi (Copepoda: 
Lernaeopodidae), a new species parasitic on an inshore fish, Anisotremns scapularis, 
off the Chilean coast 1 06- 1 08 

Romero, Raul Castro and Hernan Baeza K. Lamelliform structures on the proboscis of 
Peniculus and Metapeniculus (Copepoda: Pennellidae) „ 9 1 2-9 1 5 

Roper, Clyde F. E. and C. C. Lu. Systematic status of Lepidoteuthis, Pholidoteuthis, and 

Tetronychoteuthis (Cephalopoda: Oegopsida) 805-807 

Rossman, Douglas A., Ernest A. Liner, Carlos H. Trevino, and Alan H. Chaney. Re- 
description of the garter snake Thamnophis exsul Rossman, 1969 (Serpentes: 
Colubridae) _ 507-514 

Ruff, R. Eugene and Betsy Brown. A new species of Euchone (Polychaeta: Sabellidae) 
from the northwest Atlantic with comments on ontogenetic variability 753-760 

Russell, David E. A new species of Odontosyllis (Polychaeta: Syllidae) from Twin Cays, 
Belize _ 768-771 

San Martin, Guillermo and Carmen Alos. Paraprocerastea crocantinae, a new genus 
and species (Polychaeta: Syllidae: Autolytinae) from the Spanish Mediterranean 870-876 

Schotte, Marilyn. Two new species of wood-boring Limnoria (Crustacea: Isopoda) from 
New Zealand, L. hicksi and L. reniculus 716-725 

Sheavly, Seba B. and Harold G. Marshall. Plytoplankton composition in a borrow pit 
lake in Virginia 2 7 2-2 7 9 

Soils- Weiss, Vivianne and Kristian Fauchald. Orbiniidae (Annelida: Polychaeta) from 
mangrove root-mats in Belize, with a revision of protoariciin genera 772-792 

Stauffer, J. R., Jr. and J. M. Boltz. Description of a rock-dwelling cichlid (Teleostei: 
Cichlidae) from Lake Malawi, Africa 8-13 

Steadman, David W. New species and records of birds (Aves: Megapodiidae, Colum- 
bidae) from an archaeological site on Lifuka, Tongo „ 537-552 

Storer, Robert W. Geographic variation in the Yellow-rumped Tanager (Aves: Thrau- 

pinae) 553-554 

Thomas, James Darwin and J. L. Barnard. Gammaropsis arawakia, a new species of 
marine Amphipoda (Crustacea) from Jamaica 89-94 

Vari, Richard P. and Ann Williams Vari. Systematics of the Steindachnerina hypostoma 
complex (Pieces, Ostariophysi, Curimatidae), with the description of three new 
species 468^482 

Wagele, Johann Wolfgang and Niel L Bruce. Natatolana pastorei (Giambiagi, 1925) 



(Crustacea, Isopoda, Cirolanidae) from the Straits of Magellan, South America: Re- 
description and notes on functional morphology 

Waren, Anders and Robert Moolenbeek. A new eulimid gastropod, Trochostilifer eu- 
cidaricola, parasitic on the pencil urchin Eucidaris tribuloides from the southern 
Caribbean 

Wicksten, Mary K. Synalpheus arostris and Philocheras lapillus, two new species of 
caridean shrimp (Crustacea) from the tropical eastern Pacific 

Wicksten, Mary K. Hippolyte zostericola (Crustacea: Decapoda) in the eastern Pacific. 

Williams, Austin B. and P. J. B. Scott. Upogebia corallifora, a new species of coral- 
boring shrimp from the West Indies (Decapoda: Upogebiidae) 

Wilson, George D. F., Oleg G. Kussakin, and Galina S. Vasina. A revision of the genus 
Microprotus Richardson with descriptions of two new species, M. acutispinatus and 
M. lobispinatus (Asellota, Isopoda, Crustacea) 

Zullo, Victor A. and Anna V. Buising. An unusual species of the Balanus amphitrite 
Darwin complex (Cirripedia, Balanidae) from the ancestral Colorado River delta in 
western Arizona and southeastern California _ 



95-105 



169-175 

78-83 
644-645 

405^110 



339-361 



924-932 



INDEX TO NEW TAXA 



Volume 102 

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

COELENTERATA 
Anthozoa 

Marianactis 816 

bythios 818 

PLATYHELMINTHES 

Trematoda 

Callorhynchocotyle amatoi 268 

callorhynchi n.c 268 

NEMATODA 

Pareurystomina alima _ 255 

americana 251 

parafloridensis 254 

vaughtae 253 

MOLLUSCA 

Polyplacophora 

Rhyssoplax baliensis 808 

Gastropoda 

Pyrgulopsis aardahli 169 

amargosae 181 

owensensis 187 

perturbata _ 189 

wongi 196 

Trochostilifer eucidaricola 169 

Tryonia margae 202 

protea 207 

robusta 208 

rowlandsi 21 1 

salina 215 

SIPUNCULOIDEA 

Aspidosiphon (Akrikos) 839 

MYZOSTOMIDA 

Myzostoma armatae 793 

attenuatum 796 

divisor 799 

ANNELIDA 

Polychaeta 

Arabella monroi 292 

panamensis 296 

Axiothella crozetensis 866 

Bathykurila guaymasensis 159 



Benhamipolynoe cairnsi 301 

Branchinologluma burkensis 144 

Euchone bansei 753 

Eunoe alvinella 308 

Harmothoe hollisi 305 

LEPIDASTHENIINAE 300 

Lepidonotopodium minutum 142 

Levensteiniella raisae 139 

Macellicephaloides alvini 162 

Odontosyllis twincayensis 768 

Opisthotrochopodus marianus 149 

Paralvinella hessleri 761 

Paraprocerastea 872 

crocantinae 874 

Pararicia belizensis 789 

Pettibonella 785 

multiuncinata 785 

Protoaricia pigmentata 783 

Oligochaeta 

Limnodriloides olearius 887 

Phallodrilus hessleri 131 

vescus 134 

Tubificoides 

bruneli _ 882 

crinitus 883 

inops 880 

panamensis 878 

paracrinitus 889 

pequegnatae 891 

Branchiobdellida 

Sathodrilus nigrofluvius 738 

ARTHROPODA 

Pycnogonida 

Odontothelphusa monodontis 396 

Potamocarcinus leptomelus 394 

Sericosura cochleifovea 732 

Typhlopseudothelphusa hyba 398 

Crustacea 

Acontiophorus excavatus 916 

Aegla denticulata lacustris 385 

ALLOCRANGONYCTIDAE. 947 

Ampelisca burkei 375 

Ascelocythere stockeri 325 

Austroregia 703 

Balanus canabus 926 

Callistocaris _ 961 

Cancer johngarthi _ „ 613 

Chaceon „ 51 

atopus 53 

bicolor. 55 

chilensis 401 

crosnieri 57 

eldorado 61 

inglei 62 



mediterraneus 66 

notialis 69 

ramosae 646 

sanctaehelenae 70 

Dantya ferox 902 

Eulimnadia ovilunata 894 

ovisimilis 898 

Fallicambarus (Creaserinus) gilpini 684 

Fallicambarus (Fallicambarus) petilicarpus 6 6 1 

Gammaropsis arawakia 89 

Garosyrrhoe bigarra n.c 370 

Hobbseus yalobushensis 637 

Ingolfiella australiana 933 

bassiana _ 938 

Limnoria (Limnoria) hicksi _ 717 

reniculus 720 

Lophaxius. 962 

rathbunae 963 

Microprotus acutispinatus 354 

lobispinatus 349 

Neobrachiella anisotremi _ _ 106 

Philocheras lapillus 80 

Posthonocaris 964 

PSEUDOCRANGONYCTIDAE 953 

Pseudocyclops lepidotus 331 

Pseudothelphusa parabelliana 46 

Sakaiocaris _ 964 

Sanquerus, new name 698 

Smalleyus 45 

tricristatus 46 

Stenorhynchus yangi 631 

Synalpheus arostris _ 78 

Upogebia corallifora 405 

Zariquieyon 72 

inflatus _ 73 

Insecta 

Adelphydraena 448 

orchymonti 450 

spangleri 448 

Leptodictya fuscepes 968 

Leptopharsa madrigali 969 

reflexa 970 

Niesthrea ashlocki 609 

Nocticanace panamensis 599 

wirthi 598 

Smicridea (Smicridea) anomala 419 

aurimacula 421 

bulara 423 

cartiensis 421 

circinata 429 

corralita 423 

filicata 426 

holzenthali 431 

latipala 431 

mirama 426 

multidens 423 

tobada 419 



CHAETOGNATHA 

BATHYBELIDAE 973 

Eukrohnia calliops 34 

KROHNITTELLIDAE 973 

Paraspadella anops 586 

ECHINODERMATA 

Holothuroidea 

Hansenothuria 979 

benti 983 

TUNICATA 

Ascidiacea 

Ascidia fusca 24 

Ciona pomponiae 17 

P 'olyclinum johnsoni 15 

Situla rineharti 19 

Styela psoliformis 25 

CHORDATA 

Pisces 

Acanthemblemaria paula 1019 

Etheostoma wapiti 989 

Pseudotropheus xanstomachus 8 

Steindachnerina gracilis .-. 474 

planiventris 479 

quasimodoi _ 476 

Symphurus callopterus 460 

Amphibia 

Eleutherodactylus chrysozetetes 483 

cruzi 485 

polymniae 492 

Eupsophus contulmoensis 1031 

emiliopugini 568 

Physalaemus crombei 500 

Reptilia 

Coniophanes alverezi 1036 

Lepidodactylus balioburius 560 

Aves 

Megapodius alimentum 538 

Mammalia 

Gracilinanus 4 

PLANTAE 

Algae 

Rhodogorgon 1053 

carriebowensis _ 1057 

ramosissima 1059 



ERRATA 

Hobbs, H. H., & H. W. Robison. 1989. 102(3):65 1-697. For Fig. 2 (p. 657) and Fig. 7 (p. 673), the same 
maps were inadvertently printed. The correct map and caption for Fig. 2 is shown below. 



MISSOURI 




Fig. 2. Distribution of Fallicambarus (F.) dissitus (encircled stars), F. (F.) jeanae (encircled dots), F. (F.) 
petilicarpus (triangle) and F. (F.) strawni (dots) in Arkansas. (Some localities listed in text too close to others to 
be shown.) 



AJ/+ 



PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY 

OF 

WASHINGTON J 




VOLUME 102 NUMBER 1 
29 MARCH 1989 



ISSN 0006-324X 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

1988-1989 

Officers 



President: Kristian Fauchald 
President-elect: Leslie W. Knapp 

Gary R. Graves 
W. Ronald Heyer 
W. Duane Hope 



Elected Council 



Secretary: G. David Johnson 
Treasurer: Don E. Wilson 



Meredith L. Jones 
Raymond B. Manning 
Wayne N. Mathis 



Custodian of Publications: David L. Pawson 



PROCEEDINGS 



Editor: C. Brian Robbins 



Associate Editors 



Classical Languages: George C. Steyskal 

Plants: David B. Lellinger 
Insects: Wayne N. Mathis 



Invertebrates: Stephen D. Cairns 
Frank D. Ferrari 
Raymond B. Manning 

Vertebrates: G. David Johnson 



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THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. 



PROC. BIOL. SOC. WASH. 
102(1), 1989, pp. 1-2 

PALEONTOLOGICAL TYPE SPECIMENS IN THE 
SANTA BARBARA MUSEUM OF NATURAL HISTORY 

Scott E. Miller 

Abstract.— In accordance with Recommendation 72G of the International 
Code of Zoological Nomenclature, which recommends that institutions publish 
lists of the type specimens in their collections, this paper lists primary paleon- 
tological type specimens in the Santa Barbara Museum of Natural History, 
Santa Barbara, California (SBMNH). This list does not include paratypes and 
plesiotypes, which include those of Bremner (1933), Chaney and Mason (1 933), 
and Orr (1968). 



Kingdom Plantae 
Family Rosaceae 

Pyrus hoffmannii Chaney and Mason, 
1933:61, pi. 7, figs. 9, 10, 13; syntypes flow- 
er, fruit, and leaf, SBMNH Pal. 277-279 
(SBMNH Pal. Bot. Types 26-28); Late 
Pleistocene, "Carpinteria Formation"; Car- 
pinteria Asphalt Deposit, Santa Barbara 
County, California. Synonymized under 
Amelanchier pallida Greene by Miller 
(1979). 



Stone Quarry (depth 50 feet), Santa Bar- 
bara, Santa Barbara County, California. 

Class Mammalia 
Family Otariidae 

Imagotaria downsi Mitchell, 1968:1844, 
figs. 1-14; holotype partial skeleton, 
SBMNH Pal. 342; Early Pliocene or Late 
Miocene, Sisquoc Formation; Quarry of 
Great Lakes Carbon Company, near Lom- 
poc, Santa Barbara County, California. 



Kingdom Animalia 

Class Aves 

Family Pseudodontornithidae 

Osteodontornis orri Howard, 1957:3, figs. 
2-8; holotype nearly complete skeleton in 
2 shale slabs, SBMNH Pal. 309; Miocene, 
Monterey Formation; Flagstone quarry of 
G. Antolini and Sons, west side of Tepus- 
quet Creek, Santa Barbara County, Califor- 
nia. 

Family Phalacrocoracidae 

Phalacrocorax rogersi Howard, 1932:118, 
fig. 19; holotype coracoid, SBMNH Pal. 1 3 1 
(SBMNH Pal. Orn. 32. 1); Early Pleistocene, 
Santa Barbara Formation; Veronica Springs 



Literature Cited 

Bremner, C. S. J. 1933. Geology of San Miguel Island, 
Santa Barbara County, California. — Santa Bar- 
bara Museum of Natural History Occasional Pa- 
per 2:1-23. 

Chaney, R. W., & H. L. Mason. 1933. A Pleistocene 
flora from the asphalt deposits at Carpinteria, 
California.— Carnegie Institute of Washington 
Publication 415:45-79. 

Howard, H. 1932. A new species of cormorant from 
Pliocene deposits near Santa Barbara, Califor- 
nia. -Condor 34:1 18-120. 

. 1957. A gigantic "toothed" marine bird from 

the Miocene of California. — Santa Barbara Mu- 
seum of Natural History, Department of Ge- 
ology Bulletin 1:1-23. 

Miller, S. E. 1979. Reevaluation of Pyrus hoffmannii 
(Rosaceae) from the Pleistocene Carpinteria As- 
phalt Deposit, California. — Madrono 26:190- 
191. 



2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Mitchell, E. D., Jr. 1968. The Mio-Pliocene pinniped Santa Barbara Museum of Natural His- 

Imagotaria. -Journal of the Fisheries Research tory ^ 2559 Puesta Del Sol Road, Santa Bar- 

„ » B ^aL C Z a t 25 ' AS TJ 90 \ T1 A bara, California 93105; Present address: 

Orr, P. C. 1968. Prehistory of Santa Rosa Island.- ' __ ,' A __ . . 

Santa Barbara Museum of Natural History, San- Blsho P Museum, Box 1 9000-A, Honolulu, 

ta Barbara, California, xxi + 253 pp. Hawaii 968 17. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 3-7 

A NEW GENERIC NAME FOR TATE'S (1933) 

MICROTARSUS GROUP OF SOUTH AMERICAN 

MOUSE OPOSSUMS 

(MARSUPIALIA: DIDELPHIDAE) 

Alfred L. Gardner and G. Ken Creighton 

Abstract. —Until recently, all mouse opossums were considered as belonging 
to the genus Marmosa, as revised by Tate (1 933). Today, however, we recognize 
that Marmosa (s.l.) contains several distinctive species groups whose relation- 
ships are best expressed by dividing this complex into five genera as follows: 
Marmosa (s.s.), Marmosops, Micoureus, Thylamys, and a complex of species 
for which we propose a new name Gracilinanus because no genus-group name 
is available. 



Systematists have differed in their inter- 
pretation of the generic and subgeneric tax- 
onomy of the smaller mouse- sized marsu- 
pials of the family Didelphidae. Thomas 
(1888) grouped the known taxa of mouse 
opossums under the subgenus Micoureus in 
the genus Didelphys. Matschie (1916) dis- 
tributed these taxa among five subgenera of 
Didelphis: Marmosa, Grymaeomys, Mar- 
mosops, Thylamys, and Caluromys. Ca- 
brera (1919) grouped mouse opossums into 
two subgenera {Marmosa and Thylamys) 
under the genus Marmosa. Tate's (1933) re- 
vision was the most comprehensive. Tate 
(1933:22) believed that the genus Marmosa 
was a "'good,' natural genus of didelphids" 
that included all Neotropical mouse opos- 
sums. Although Tate acknowledged natural 
subunits in his concept of Marmosa, he did 
not recognize subgenera and, instead, used 
the following names to indicate five infor- 
mal groups: cinerea, murina, noctivaga, mi- 
crotarsus, and elegans. He also expressed 
the opinion (p. 22) that "there is always the 
chance that subgenera may be later con- 
verted into full genera, and thus, in the case 
of Marmosa the undoubted unity of the ge- 
nus be obscured." 

Over the ensuing half-century, however, 
new information has made the division of 



Marmosa (sensu Tate 1933) into several 
genera a desirable and necessary step to bet- 
ter reflect relationships and natural assem- 
blages. Gilmore (1941) again divided Mar- 
mosa into the subgenera Marmosa and 
Thylamys but did not list species. Cabrera 
(1958) used Marmosa and Thylamys as sub- 
genera as he had earlier (Cabrera 1919) and 
followed the species associations of Tate 
(1933) except that he divided Tate's micro- 
tarsus group, assigning the microtarsus sec- 
tion to Thylamys and the lepida section to 
Marmosa. Marshall (1981) listed the names 
Marmosa ("murine opossums"), Micoureus 
("large murine opossums"), and Thylamys 
("small murine opossums"), as genera 
among other didelphids and attributed this 
arrangement to a manuscript by Reig et al. 
subsequently published in 1985. Reig et al. 
(1985) assigned Tate's (1933) murina and 
noctivaga groups to the genus Marmosa 
along with the more recently described taxa 
M. andersoni, M. cracens (incorrectly spelled 
crascens), and M. xerophila. They allocated 
Tate's (1933) cinerea group to Micoureus. 
They also assigned the 14 species listed by 
Kirsch & Calaby (1977) along with Mar- 
mosa lepida, M. emiliae, and M. contrerasi, 
to Thylamys. Creighton (1984) also recog- 
nized five groups in his revision of the genus 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Marmosa (s.L). He placed four of these (ci- 
nerea, murina, noctivaga, and microtarsus) 
in the subgenus Marmosa and the fifth (ele- 
gans) in the subgenus Thy I amy s. 

Despite Creighton's (1984) conservative 
treatment of Marmosa (s.L), each of the 
groups he recognized corresponds to a genus 
in the sense used by Kirsh & Calaby (1977) 
and Reig et al. (1985, 1987). Creighton's 
elegans-group equals Thylamys and corre- 
sponds closely to Tate's elegans group, but 
is more restrictive than construed by Ca- 
brera (1958), Kirsch & Calaby (1977), and 
Reig et al. (1985, 1987). We recognize Thy- 
lamys Gray, 1843 as a genus that includes 
the species T elegans (genotype), T. ma- 
crura (Olfers, 1818:205; Didelphis grisea 
Desmarest, 1827, is a junior synonym), T 
pallidior, T pusillus, and T. velutinus. 
Creighton's cinerea -group corresponds to 
that of Tate (1933) as used by Reig et al. 
(1985, 1987) under the generic name Mi- 
coureus Lesson, 1 842 (Micoures Reig et al. 
1987, is an incorrect subsequent spelling), 
to which we assign the species Micoureus 
cinereus (genotype), M. constantiae, M. re- 
gina, and M. alstoni. Creighton's murina- 
group is similar to Tate's (1933) group by 
the same name except that Creighton also 
included M. lepida. The earliest available 
name for Creighton's murina-group is Mar- 
mosa Gray, 1821, which we use here as a 
genus in a narrower sense than treated by 
Reig et al. (1985, 1987). As we understand 
it, Marmosa (s.s.) includes Marmosa an- 
dersoni, M. canescens, M. lepida, M. mex- 
icana, M. murina (genotype), M. robinsoni, 
M. rubra, M. tyleriana, and M. xerophila. 
Reig et al. (1985, 1987) combined Tate's 
noctivaga and murina groups under Mar- 
mosa. However, Creighton (1984) showed 
that Tate's (1933) noctivaga group, with the 
addition of M. parvidens (which Tate had 
included in his microtarsus group), and the 
more recently described taxa M. cracens and 
M. handleyi, is a natural unit of closely re- 
lated species for which the name Marmo- 
sops Matschie, 1916, is available. Marmo- 



sops includes the species M. cracens, M. 
dorothea, M. fuscatus, M. handleyi, M. im- 
pavidus, M. incanus (genotype), M. invictus, 
M. noctivagus, and M. parvidens. This is 
similar to Reig et al.'s (1987:7) composition 
of Marmosops, which they ranked as a sub- 
genus of Marmosa. 

Creighton's (1984) fifth taxon, the micro- 
tarsus-group, is nearly identical to Tate's 
microtarsus section with the addition of the 
more recently described M. agricolai. How- 
ever, Kirsch & Calaby (1977) and Reig et 
al. (1985, 1987) followed Cabrera (1958) in 
combining Tate's (1933) elegans group and 
microtarsus section under Thylamys, along 
with some other taxa that Creighton (1984) 
showed belong elsewhere. Creighton includ- 
ed nine species in his microtarsus '-group, 
although here we recognize only six. No ge- 
nus-group name is available for this species 
complex for which we propose the name: 

Gracilinanus, new genus 

Type species. —Didelphys microtarsus 
Wagner, 1842:359; type locality "Ypane- 
ma" (=Bacaetava), Sao Paulo, Brazil. 

Etymology. —From the Latin gracilis 
(thin, slender) and Greek nanos (dwarf) 
combined to reflect the small size and slen- 
der or gracile form of the species included 
in this genus. 

Distribution. — Gracilinanus contains six 
living species whose composite geographic 
range is below 3000 m in the tropical and 
subtropical zone east of the Andean Cor- 
dillera of South America. 

Diagnosis and description.— A genus of 
small mouse opossums (head and body, 
85.0-130.0 mm; tail, 90.0-150.0 mm). The 
ratio of tail length to head-and-body length 
is always greater than 1.3 and usually less 
than 1.5. Dorsal pelage ranges from bright 
reddish brown to dull brownish gray. The 
tail has small (more than 40 per cm) round- 
ed to square scales, arranged in annular rows. 
Triplet interscalar hairs are subequal in 
length with the middle hair about three scale 
rows long. The diameter of the middle hair 



VOLUME 102, NUMBER 1 



is about twice that of the lateral hairs, but 
not as stout and petiolate as in Marmosops. 
The tail is weakly bicolored in some species 
(G. agilis and G. marica) and unicolored 
fuscous in others (G. aceramarcae, G. dryas, 
and G. microtarsus). Claws on the manus 
do not extend beyond terminal digital pads. 
The central palmar surfaces lack the gran- 
ular appearance of those of Thylamys and 
Monodelphis. All palmar and plantar tu- 
bercles (description by Creighton 1984) are 
present and separated by at least a double 
row of granules. Sparse granules are present 
on the central plantar surface. Granules on 
the proximal ventral surfaces of the digits 
are fused into transverse bars. The skull lacks 
postorbital processes; however, the su- 
praorbital margin of the frontals may be 
beaded in larger individuals of some species. 
The lambdoidal crest is weakly developed 
or absent except in the largest individuals. 
The hard palate is highly fenestrated, usu- 
ally with three pairs of medial fenestrae 
(medial, posteromedial, and mesolateral— 
described in Creighton 1984). The postero- 
lateral fenestrae are moderate in size, usu- 
ally about a third to half the breadth of the 
last upper molar in length. The nasals are 
moderately expanded laterally at the max- 
illo-frontal suture. The auditory bullae are 
relatively large compared with those of 
Marmosa, Micoureus, and Marmosops, but 
proportionately smaller than in Thylamys. 
A slender anteromedial process of the ali- 
sphenoid portion of the auditory bulla is 
always present, although frequently dam- 
aged during specimen preparation. The sec- 
ond upper premolar is always larger than 
the third. The incisors increase slightly in 
size from 12 through 15. The lower canines 
are relatively short compared with those of 
Marmosa, Micoureus, and Thylamys, but 
are not as short or premolariform as in Mar- 
mosops. 

Species of Gracilinanus are distinguish- 
able from those of Marmosops by the shape 
and arrangement of tail scales and bristles, 
the non-premolariform lower canine, and 



the presence of pectoral mammae. They can 
be separated from species of Thylamys by 
the absence of seasonal fat deposits in the 
tail (thus non-incrassate), absence of dense- 
ly granular central palmar and plantar sur- 
faces, and relatively longer digits and broad- 
er interdigital pads on manus and pes. 
Gracilinanus can be distinguished from 
Marmosa and Micoureus by the lack of 
postorbital processes on the frontals and by 
the annular arrangement of minute scales 
on the tail. 
Included species. — 

Gracilinanus aceramarcae (Marmosa ac- 
eramarcae Tate, 1931:12; type locality 
"Rio Aceramarca, tributary of Rio Un- 
duavi, Yungas," La Paz, Bolivia). 

Gracilinanus agilis (Grymaeomys agilis 
Burmeister, 1854:139; type locality "La- 
goa Santa," Minas Gerais, Brazil). 

Synonyms: 

Marmosa beatrix Thomas, 1 9 1 0: 502; type 
locality "Ipu," Ceara, Brazil. 

Marmosa muscula Shamel, 1930a: 83; 
type locality "Kilometro 182" (=Ria- 
cho Pilaga, 1 miles northwest of Km 
182), Formosa, Argentina. 

Marmosa formosa Shamel, 1930b:311; 
renaming of M. muscula Shamel, 
1 930tf, preoccupied by Didelphis (Mar- 
mosa) musculus Cabanis, 1848. 

Marmosa agilis chacoensis Tate, 1931:10; 
type locality "Sapucay," Paraguay. 

Marmosa agilis buenavistae Tate, 1931: 
10; type locality "Buena vista, Depart- 
ment of Santa Cruz, Bolivia." 

Marmosa agilis peruana Tate, 1931:11; 
type locality "Tingo Maria, Rio Hua- 
llaga," Huanuco, Peru." 

Marmosa unduaviensis Tate, 1931:11; 
type locality "Pitiguaya, Rio Unduavi, 
Yungas," La Paz, Bolivia. 

Marmosa blaseri Miranda-Ribeiro, 1936: 
373; type locality "S. Bento," Goias, 
Brazil. 

Thylamys rondoni Miranda-Ribeiro, 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



1936:387; type locality "salto do Se- 
potuba e S. Joao da Serra do Norte," 
Mato Grosso, Brazil. 
Gracilinanus dryas {Marmosa dryas Thom- 
as, 1898:456; type locality "Culata, Me- 
rida, Venezuela"). 
Gracilinanus emiliae {Marmosa emiliae 
Thomas, 1909:379; type locality "Para, 
Brazil"). 

Synonym: 

Marmosa agricolai Moojen, 1943:2; type 
locality "Crato, Ceara," Brazil. 

Gracilinanus marica (Marmosa marica 
Thomas, 1898:455; type locality "R. Al- 
barregas, Merida, Venezuela"). 

Gracilinanus microtarsus (Didelphys micro- 
tarsus Wagner, 1842:359; type locality 
"Ypanema," Sao Paulo, Brazil). 

Synonyms: 

Marmosa microtarsus guahybae Tate, 
1931:10; type locality "Island of Gua- 
hyba near Porto Alegre, Rio Grande do 
Sul," Brazil. 

Marmosa herhardti Miranda-Ribeiro, 
1936:382; type locality "Humboldt," 
Santa Catarina, Brazil. 

Literature Cited 

Burmeister, H. 1854. Systematische Uebersicht der 
Thiere Brasiliens, welche wahrend einer Reise 
durch die Provinzen von Rio de Janeiro und 
Minas Geraes gesammelt oder beobachtat wur- 
den von Dr. Hermann Burmeister. I. Sauge- 
thiere (Mammalia). G. Reimer, Berlin, x + 342 
pp. 

Cabanis, J. 1848. Saeugethiere. Pp. 766-786 in R. 
Schomburgk, ed., Versuch einer Fauna und Flora 
von Britisch-Guiana, in Reisen in Britisch- 
Guiana in den jahren 1840-1844. J. J. Weber, 
Leipzig, 3:vii + 531-1260. 

Cabrera, A. 1919. Genera Mammalium. Museo Na- 
tional de Ciencias Naturales, Madrid, 177 pp., 
17 pis. 

. 1958. Catalogo de los mamiferos de America 

del Sur. — Revista del Museo Argentino de Cien- 
cias Naturales "Bernardino Rivadavia," Cien- 
cias Zoologicas 4(l):iv + 307 pp. 

Creighton, G. K. 1984. Systematic studies on opos- 
sums (Didelphidae) and rodents (Cricetidae). 



Unpublished Ph.D. dissertation, University of 
Michigan, xi + 220 pp. 

Desmarest, A. G. 1827. Sarigue. Pp. 392-399, Vol. 
47 in F. G. Cuvier, ed., Dictionnaire des sciences 
naturelles, dans lequel on traite methodique- 
ment des differens etres de la nature considered 
soit en eux-memes, d'apres l'etat actuel de nos 
connoissances, soit relativement a Futilite qu'en 
peuvent retirer la medecine, l'agriculture, le 
commerce et les arts. F. G. Levrault, Strasbourg; 
Le Normant, Paris, 1816-1845, 61 Vols., 12 
Vols. pis. 

Gilmore, R. M. 1941. Zoology. Pp. 3 1 4-3 1 9 in J. C. 
Bugher, J. Boshell-Manrique, M. Roca-Garcia, 
and R. M. Gilmore, eds., The susceptibility to 
yellow fever of the vertebrates of eastern Co- 
lombia.— American Journal of Tropical Medi- 
cine 21:309-333. 

Gray, J. E. 1821. On the natural arrangement ofver- 
tebrose animals. —London Medical Repository 
15:297-311. 

. 1843. List of the specimens of Mammalia in 

the collection of the British Museum. British 
Museum (Natural History), London, xxviii + 
216 pp. 

Kirsch, J. A. W., & J. H. Calaby. 1977. The species 
of living marsupials: An annotated list. Pp. 9- 
26 in B. Stonehouse and D. Gilmore, eds., The 
biology of marsupials, University Park Press, 
Baltimore, viii + 486 pp. 

Lesson, R. P. 1842. Nouveau tableau do Regne An- 
imal. Mammiferes. Arthus-Bertrand, Paris, 204 
pp. 

Marshall, L. G. 1981. The families and genera of 
Marsupialia. — Fieldiana: Geology, new series 8: 
1-65. 

Matschie, P. 1916. Bemerkungen iiber die Gattung 
Didelphis L.— Sitzungsberichte der Gesellschaft 
Naturforschender Freunde, Berlin 1916(1): 259- 
272. 

Miranda-Ribeiro, A. 1936. Didelphia ou Mamma- 
lia— ovovivipara. — Revista do Museu Paulista, 
Sao Paulo 20:245-427. 

Moojen, J. 1943. Algunos mamiferos colecionados 
do nordeste do Brasil com a descriaao de duas 
especies novas e notas de campo.— Boletim do 
Museu Nacional, Rio de Janeiro, Zoologia 5:1- 
14. 

Olfers, I. von. 1818. Bemerkungen zu Illiger's Ue- 
berblik der Saugthiere nach ihrer Vertheilung 
uber die Welttheile, rucksichtlich Sudamerican- 
ischen Arten [chapter 10]. Pp. 192-237 in W. 
L. Eschwege, Journal von Brasilien, oder ver- 
mischte Nachrichten aus Brasilien, auf 
wissenschaftlichen Reisen gesammelt. Vol. 15, 
heft 2 in F. T. Bertuch, ed., Neue Bibliotek des 
wichtigsten Reisen-beschreibungen zur Erwei- 
terung der Erd- und Volkerkunde; in Verbin- 



VOLUME 102, NUMBER 1 



dung mit einigen anderen Gelehrten gesammelt 
und herausgegeben .... Gr. H. S. pr. Landes- 
industries-comptoirs, Weimar. 

Reig, O. A., J. A. W. Kirsch, & L. G. Marshall. 1985. 
New conclusions on the relationships of the 
opossum-like marsupials, whith [sic] an anno- 
tated classification of the Didelphimorphia. — 
Ameghiniana 21(2-4):335-343. 

, , & . 1987. Systematic relation- 
ships of the living and Neocenozoic American 
"opossum-like" marsupials (suborder Didel- 
phimorphia), with comments on the classifica- 
tion of these and of the Cretaceous and Paleo- 
gene New World and European metatherians. 
Pp. 1-90 in Possums and opossums: Studies in 
evolution. Surrey Beatty & Sons PTY Limited, 
New South Wales, frontispiece, lxxii + 400 pp., 
4 pis. 

Shamel, H. H. 1930a. A new murine opossum from 
Argentina.— Journal of the Washington Acad- 
emy of Sciences 20:83. 

. 1930b. A new name for Marmosa muscula 

Shamel. —Journal of Mammalogy 11:311. 

Tate, G. H. H. 1931. Brief diagnoses of twenty-six 
apparently new forms of Marmosa (Marsupi- 
alia) from South America.— American Museum 
Novitates 493:1-14. 

. 1933. A systematic revision of the marsupial 

genus Marmosa.— Bulletin of the American 
Museum of Natural History 66:1-250, 26 pis., 
1 table (9 sections, pocketed). 



Thomas, O. 1888. Catalogue of the Marsupialia and 
Monotremata in the collection of the British 
Museum (Natural History). British Museum 
(Natural History), London, xiv + 401 pp., 33 
pis. 

. 1 898. On seven new mammals from Ecuador 

and Venezuela. — Annals and Magazine of Nat- 
ural History, Series 7 1:451^457. 

. 1 909. New species o/Oecomys and Marmosa 

from Amazonia. —Annals and Magazine of Nat- 
ural History, Series 8 3:378-380. 

. 1910. On mammals collected in Ceara, N. E. 

Brazil by Fraulein Dr. S net hlage.— Annals and 
Magazine of Natural History, Series 8 6:500- 
503. 

Wagner, A. 1 842. Diagnosen neuer Arten Brasilischer 
Saugethiere.— Archiv fur Naturgeschichte 8(1): 
359-360. 

(ALG) Biological Survey Field Station, 
National Ecology Research Center, U.S. Fish 
and Wildlife Service, National Museum of 
Natural History, Washington, D.C. 20560; 
(GKC) Division of Mammals, National 
Museum of Natural History, Washington, 
D.C. 20560. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 8-13 

DESCRIPTION OF A ROCK-DWELLING CICHLID 

(TELEOSTEI: CICHLIDAE) FROM 

LAKE MALAWI, AFRICA 

J. R. Stauffer, Jr. and J. M. Boltz 

Abstract. —A new species of cichlid fish of the Pseudotropheus zebra complex, 
P. xanstomachus, is described from Lake Malawi, Africa. The species is en- 
demic to the Maleri Islands. The species is morphologically similar to the 
sympatric Pseudotropheus barlow McKaye & Stauffer, but the species are dis- 
tinguishable in both coloration and shape. 



The rock-dwelling cichlids ("mbuna") 
which inhabit the rocky shores and rock 
outcroppings of Lake Malawi are repre- 
sented by many species which are endemic 
to particular islands. Ribbink et al. (1983) 
published a survey of this group and Lewis 
et al. (1986) published a guide to the fishes 
of the Lake Malawi National Park, which 
discussed the distribution of many of the 
mbuna. The purpose of this paper is to de- 
scribe a new species closely related to Pseu- 
dotropheus zebra (Boulenger) endemic to a 
group of islands collectively referred to as 
the Maleri Islands, located in Lake Malawi, 
Malawi, Africa. Ribbink et al. (1983) rec- 
ognized this form and referred to it as P. 
zebra "yellow throat." 

Methods and materials. —Standard length 
(SL) is used throughout. External counts and 
measurements follow Barel et al. (1977). 
Scale counts in the lateral line series do not 
include scales in the overlapping portion of 
the lower lateral line. Except for gill raker 
meristics, which were recorded from the 
right side, all counts and measurements were 
made on the left side of the fish. Vertebral 
counts were made from radiographs. All 
specimens were collected at Nakantenga Is- 
land (34°39'E, 13°55'S), Lake Malawi, Af- 
rica (Apr 1984 and Aug 1987). 

The new species is compared with P. bar- 
lowi by color descriptions of fresh speci- 
mens, meristic differences, and shape dif- 



ferences. Body shape differences were 
compared using sheared principal compo- 
nent analysis (Humphries et al. 1981, Book- 
stein et al. 1985). This analysis quantifies 
shape differences between the two popula- 
tions independent of size of the individuals 
(Reyment et al. 1984). 



Pseudotropheus xanstomachus, new species 
Figs. 1, 2; Tables 1, 2 

Pseudotropheus zebra, Ribbink et al., 1983: 
162 (in part). 

Holotype. —National Museum of Natural 
History (USNM) 297268, adult male, 63.3 
mm, Nakantenga Island, Lake Malawi, 3- 
7 m, 4 Aug 1987. 

Paratypes. -USNM 297269, 11 (47.0- 
70.8 mm), data as for holotype; USNM 
297270, 4 (63.5-78.5 mm), Nakantenga Is- 
land, Lake Malawi, 12 m, 19 Apr 1984. 

Diagnosis. —This species fits the descrip- 
tion of the genus Pseudotropheus as given 
by Regan (1921) in that: 1) its jaws have 
several rows of teeth, with the ones in the 
outer row bicuspid and those in the inner 
rows tricuspid; 2) it has 16-18 dorsal-fin 
spines, 7-8 dorsal-fin rays, 3 anal-fin spines, 
and 6-7 anal-fin rays. However, as noted 
by Ribbink et al. (1983), Fryer (1957:350) 
regarded the genus Pseudotropheus as being 
"rather ill-defined." Pseudotropheus xan- 



VOLUME 102, NUMBER 1 




Fig. 1. Holotype (USNM 297268) of Pseudotropheus xanstomachus. 



stomachus differs from other members of 
the group in coloration. It differs from the 
only described sympatric member of the 
group in the number of anal-fin rays, shape 
of the lower pharyngeal bone, the length of 
the dentigerous surface of the lower pha- 
ryngeal bone, and in body shape. 
Description. — Morphometric ratios and 



meristics are presented in Table 1. Body 
moderately compressed and elongate; jaws 
isognathous (Fig. 1). Teeth on lower jaw in 
3-4 rows, those on premaxilla in 4 rows; 
teeth in outer rows bicuspid, with occasion- 
al conical lateral tooth in some individuals; 
teeth in inner rows tricuspid; 12 teeth in 
outer row of left lower jaw of holotype, 9- 





mm 



P. barlowi 




R xanstomachus 



Fig. 2. 
machus. 



Lower pharyngeal bones of the holotypes of Pseudotropheus barlowi and Pseudotropheus xansto- 



10 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1 . — Morphometric characters and meristics of Pseudotropheus xanstomachus. Range includes holotype 
and 15 paratypes. 





Holotype 


Mean 


SD 


Range 


Standard length, mm 


63.3 


62.9 


7.1 


47.0-78.5 


Head length, mm 


20.5 


20.4 


2.4 


15.3-26.0 


Percent Standard Length 










Head length 


32.4 


32.4 


0.7 


31.5-33.4 


Snout to dorsal 


32.2 


33.3 


1.2 


30.9-35.0 


Snout to pelvic 


43.4 


43.1 


1.3 


40.8^45.2 


Greatest Body depth 


36.7 


34.2 


1.4 


31.8-36.5 


Caudal peduncle length 


12.8 


13.5 


1.3 


11.2-15.5 


Least caudal peduncle depth 


13.0 


12.9 


0.3 


12.3-13.4 


Pectoral-fin length 


26.5 


25.6 


1.8 


22.4-28.2 


Pelvic-fin length 


31.8 


27.7 


4.5 


21.6-36.4 


Dorsal-fin base length 


60.7 


60.4 


1.1 


58.6-62.2 


Percent Head Length 










Horizontal eye diameter 


30.7 


31.3 


1.1 


28.5-32.8 


Vertical eye diameter 


30.2 


30.9 


1.3 


28.1-33.3 


Snout length 


33.7 


35.1 


1.4 


33.2-37.7 


Postorbital head length 


40.5 


40.0 


0.8 


38.5-41.2 


Preorbital depth 


21.5 


20.3 


1.2 


18.2-22.6 


Lower jaw length 


34.6 


33.6 


1.2 


32.0-36.5 


Interorbital width 


28.3 


28.3 


2.4 


25.3-33.6 


Cheek depth 


30.2 


30.0 


2.8 


24.5-34.4 


Head depth 


98.5 


95.1 


3.0 


91.1-100.1 


Counts 










Lateral line scales 


30 


29.9 


0.5 


29-31 


Scale rows on cheek 


5 


5.4 


0.5 


5-6 


Dorsal-fin spines 


17 


17.1 


0.5 


16-18 


Dorsal-fin rays 


8 


7.9 


0.4 


7-8 


Pectoral-fin rays 


13 


13.3 


1.0 


10-14 


Anal-fin rays 


7 


6.9 


0.3 


6-7 


Gillrakers on first ceratobranchial 


12 


10.8 


0.9 


9-12 


Gillrakers on first epibranchial 


3 


2.5 


0.5 


2-3 


Teeth in outer row of left lower jaw 


12 


10.7 


1.1 


9-12 


Abdominal vertebrae 


14 


13.6 


0.8 


12-15 


Caudal vertebrae 


14 


14.7 


0.5 


14-15 



1 2 in paratyes. Pectoral fins with 1 3 rays in 
holotype and 10-14 in paratypes; anal fin 
with 7 rays in holotype and 6-7 in para- 
types; caudal fin emarginate. Vertebrae of 
holotype 14 + 14 (abdominal + caudal); 
paratypes 12-14 + 14-15. Lower pharyn- 
geal bone of holotype triangular in outline, 
length of dentigerous surface 59% of lower 
pharyngeal bone length; pharyngeal teeth in 
left posterior row 23, those in left median 
row, 13. Holotype with 5 scale rows on 
cheek, paratypes with 5-6; pored scales along 



lateral line of holotype 30, and of paratypes 
29-31; pored scales posterior to hypural 
plate 2, with exception of two paratypes 
which had scales missing from this area. 
Gill rakers simple, first gill arch of holotype 
with 12 on ceratobranchial (9-12 in para- 
types); 3 on epibranchial of holotype and 
2-3 in paratypes; and 1 between epibran- 
chial and ceratobranchial. 

Body coloration in freshly collected males 
light blue with six black vertical bars; head 
dark blue with one light blue interorbital 



VOLUME 102, NUMBER 1 



11 



Table 2.— Sheared principal components (shape factors) of the untransformed morphometric data for Pseu- 
dotropheus xanstomachus (n = 16) and Pseudotropheus barlowi (n = 10). 





Size 


Sheared 




PC 2 


PC 3 


Standard length 


0.215 


-0.021 


-0.082 


Head length 


0.207 


-0.011 


-0.108 


Horizontal eye diameter 


0.183 


0.095 


-0.154 


Vertical eye diameter 


0.189 


0.048 


-0.048 


Snout length 


0.231 


-0.088 


-0.225 


Postorbital head length 


0.193 


-0.107 


-0.048 


Preorbital depth 


0.282 


-0.117 


0.042 


Lower jaw length 


0.210 


0.070 


-0.048 


Interorbital width 


0.275 


-0.065 


-0.014 


Snout to dorsal fin origin 


0.212 


0.052 


-0.149 


Snout to pelvic fin origin 


0.261 


-0.064 


-0.137 


Body depth 


0.231 


0.136 


-0.050 


Cheek depth 


0.169 


-0.594 


0.447 


Head depth 


0.229 


0.010 


-0.067 


Caudal peduncle length 


0.219 


-0.459 


0.206 


Least caudal peduncle depth 


0.215 


0.048 


-0.050 


Pectoral-fin length 


0.258 


0.155 


-0.158 


Pelvic-fin length 


0.311 


0.521 


0.728 


Dorsal-fin base length 


0.215 


-0.009 


-0.069 



bar, yellow gular and branchiostegal rays; 
dorsal fin medium blue with grey flecks and 
light blue marginal band; pectoral fins black; 
pelvic fins black anteriorly fading to yellow- 
brown posteriorly; anal fin blue; males with 
three to five yellow ocelli on anal fin. Body 
coloration of females similar, but some in- 
dividuals have light brown body coloration 
with brownish cast to fins. 

The description of the coloration and the 
color plate of the males in Ribbink et al. 
(1983:162; plate 2a) agree with the above 
description, except that they noted between 
1-5 ocelli on the anal fin. Ribbink et al. 
(1983:162) note that the females are "dark 
brown sometimes almost black with darker 
bars . . . ." The differences in color of the 
females may be associated with breeding 
coloration and may be influenced by the 
time of the year when the fish were ob- 
served. However, all descriptions of the fe- 
male coloration note a yellow chin and gular 
region. 

Etymology.— The specific epithet is de- 
rived from the Greek "xanthos" and "sto- 



machus" meaning yellow throat, which char- 
acterizes the yellow gular region present in 
both sexes. 

Discussion. —Pseudotropheus xanstoma- 
chus is morphologically similar to the P. 
zebra complex in that it has a terminal 
mouth, has three or four rows of teeth, of 
which the outer rows are bicuspid, with the 
occasional conical lateral tooth, and the in- 
ner rows are tricuspid (see Ribbink et al. 
1983). The only sympatric species of this 
complex which is described is P. barlowi, 
from which P. xanstomachus differs in col- 
oration. P. barlowi males are bright gold, 
and the females are uniformly brown 
(McKaye & Stauffer 1986). Differences in 
color patterns among cichlids. especially the 
male's, generally are recognized to be suf- 
ficient to delimit valid species (Greenwood 
1981, Hoogerhoud & Witte 1981). 

In addition to coloration differences, P. 
xanstomachus has six or seven anal rays 
while P. barlowi has eight. The shape of the 
lower pharyngeal bone between the two 
species also differs. The angle of the suture 



12 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





0.125 -I 














0.100 ■ 












SHRD_PC3 


0.075 ■ 

0.050 ■ 

0.025 ■ 

0.000 ■ 

-0.025 ■ 

-0.050 ■ 

-0.075 ■ 

-0.100 ■ 

-0.125 - 

-0.150 - 




♦ 


— h- 




♦> 

H 1 



o 



oo' 



+ 



+ 



-0.20 -0.15 -0.10 -0.05 0.00 0.05 0.10 0.15 0.20 

SHRD_PC2 

Fig. 3. Plot of the sheared principal components of Pseudotropheus xanstomachus (solid) and Pseudotropheus 
barlowi (open). 



of the two posterior halves of the lower pha- 
ryngeal bone is more acute and the resultant 
depression is greater in P. barlowi (Fig. 2). 
Moreover, the length of the dentigerous sur- 
face of the lower pharyngeal bone of P. bar- 
lowi was 76% (range = 74-78%) of the length 
of the bone (n = 3; USNM 274782; USNM 
274783), while the length of the dentigerous 
surface of the lower pharyngeal bone of P. 
xanstomachus was 62% (n = 4; range = 57- 
67%). 

An attempt was made to determine if there 
were differences in body shape between the 
two described sympatric species of the P. 
zebra complex using a sheared principal 
component analysis. Data for P. barlowi 
were obtained from the original data sheets 
used by McKaye & Stauffer (1986). Sheared 
principal component analysis demonstrated 
that there was no overlap between species 
when the first sheared principal component 
was plotted against the second sheared prin- 
cipal component (Fig. 3). The first principal 
component is interpreted as a size compo- 
nent and the sheared components as shape, 



independent of size (see Humphries et al. 
1981, Bookstein et al. 1985). Thus, these 
data were calculated using the untrans- 
formed values of the morphometries rather 
than percent standard length or percent head 
length. Those morphometries which have 
the highest loadings on the first sheared 
principal component are cheek depth, pel- 
vic-fin length, and caudal peduncle length, 
while those which have the highest loadings 
on the second sheared principal component 
are pelvic-fin length, cheek depth, and snout 
length (Table 2). 

All of the specimens reported herein were 
captured at a depth between 7-12 m at Na- 
kantenga Island. Ribbink et al. (1983) re- 
ported this form from all three Maleri Is- 
lands: Namkoma, Maleri, and Nakantenga. 
They stated that it was most common be- 
tween depths of 2-8 m, and was rare in 
water deeper than 12 m. Conversely, 
McKaye & Stauffer (1986) stated that P. 
barlowi occurred primarily at depths greater 
than 10 m and hypothesized that the oc- 
currence of these brightly colored forms in 



VOLUME 102, NUMBER 1 



13 



deeper water may be an adaptation related 
to avoidance of surface predators (i.e., birds 
and otters). Therefore, even though P. xan- 
stomachus and P. barlowi are indigenous to 
the same islands, they are in effect allotopic. 

Acknowledgments 

We thank the government of Malawi for 
providing the facilities to make this research 
possible. We thank L. Knapp of the Smith- 
sonian Institution who arranged for ship- 
ment of specimens from Malawi to the 
USNM. We benefited from discussions with 
K. R. McKaye. We are especially thankful 
to D. Swofford who provided the SAS pro- 
gram to conduct the sheared principal com- 
ponent analysis. The original art work was 
completed by M. Katz. 

Literature Cited 

Barel, C. D. N., M. J. O. Van Oijen, F. Witte, & E. L. 
M. Witte-Mass. 1977. An introduction to the 
taxonomy and morphology of the haplochro- 
mine Cichlidae from Lake Victoria. Part A. 
Text.— Netherlands Journal of Zoology 27:333- 
389. 

Bookstein, F., B. Chernoff, R. Elder, J. Humphries, G. 
Smith, & R. Strauss. 1985. Morphometries in 
evolutionary biology. Special Publication 1 5. The 
Academy of Natural Sciences of Philadelphia. 
Philadelphia, Pennsylvania. 277 pp. 

Fryer, G. 1957. A new species of Gephyrochromis 
(Pisces: Cichlidae) from Lake Nyasa, with notes 



on its ecology and affinities. — Revue de Zool- 
ogie et de Botanique Africaines 55:347-352. 

Greenwood, P. H. 1981. The haplochromine fishes 
of East African lakes. Cornell University Press, 
Ithaca, New York. 839 pp. 

Hoogerhoud, R. J. C, & F. Witte. 1981. Revision of 
species from the "Haplochromis" empodisma 
group. Revision of the haplochromine species 
(Teleostei, Cichlidae), from Lake Victoria, Part 
II.— Netherlands Journal of Zoology 31:232- 
274. 

Humphries, J. M., F. L. Bookstein, B. Chernoff, G. R. 
Smith, R. L. Elder, & S. G. Poss. 1981. Multi- 
variate discrimination by shape in relation to 
size. — Systematic Zoology 30:291-308. 

Lewis, D., P. Reinthal, & J. Trendall. 1986. A guide 
to the fishes of Lake Malawi National Park. 
World Wildlife Fund, Gland, Switzerland. 72 
pp. 

McKaye, K. R., & J. R. Stauffer, Jr. 1986. Description 
of a gold cichlid (Teleostei: Cichlidae) from Lake 
Malawi, Africa. -Copeia 1986:870-875. 

Regan, C. T. 1921. The cichlid fishes of Lake Nyara. — 
Proceedings of the Zoological Society of London 
1921:675-727. 

Reyment, R. A., R. E. Blackith, & N. A. Campbell. 
1984. Multivariate morphometries. Academic 
Press, New York, New York. 233 pp. 

Ribbink, A. J., B. A. Marsh, A. C. Marsh, A. C. Rib- 
bink, & B. J. Sharp. 1983. A preliminary sur- 
vey of the cichlid fishes of the rocky habitats in 
Lake Malawi.— South African Journal of Sci- 
ence 18:149-310. 

School of Forest Resources, The Penn- 
sylvania State University, University Park, 
Pennsylvania 16802. 



^ 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 14-32 

ASCIDIANS COLLECTED AROUND THE GALAPAGOS 

ISLANDS USING THE JOHNSON-SEA-LINK 

RESEARCH SUBMERSIBLE 

Claude Monniot and Francoise Monniot 

Abstract. — Ascidians were collected for the first time by a manned sub- 
mersible on ocean-bottoms between 300 and 800 meters around the Galapagos 
Islands. Solitary forms dominate; among the eight species collected, five are 
new ones: Polyclinum johnsoni, Ciona pomponiae, Situla rineharti, Ascidia 
fusca, and Styela psoliformis. They belong to surprisingly diverse families and 
several have deep-sea characteristics. 



The fauna of the eastern part of the trop- 
ical Pacific Ocean is poorly known, espe- 
cially the ascidians. Tokioka (1972) report- 
ed on a small collection of ascidians from 
the coast of Costa Rica, and recently Millar 
(1988) identified some specimens from the 
Galapagos and Ecuador. Nothing else is re- 
ported about ascidians between Baja Cali- 
fornia and Chile. 

In November 1986, an expedition to the 
Galapagos Islands collected deep-water or- 
ganisms for pharmacological studies. Over 
600 animals were collected by the R/V Sew- 
ard Johnson and Johnson-Sea-Link- 1 
manned submersible (Harbor Branch 
Oceanographic Institution, Fort Pierce, FL) 
using a manipulator arm equipped with a 
claw, a suction tube, and a grab sampler. 
Among this collection there are eight species 
of ascidians, five of which are new species. 

The material is particularly interesting 
because the specimens were collected un- 
damaged on rocky bottoms, where a dredge 
or any blunt device monitored from a sur- 
face boat cannot operate. The animals are 
unusually large compared to the size of the 
ascidians collected in shallow water by 
SCUBA during the same cruise and relative 
to the deep-water ascidians on soft bottoms 
elsewhere. 

Observation from the submersible during 
several dives suggests that ascidians are not 



evenly distributed on the bottom, but live 
in patches in restricted areas. They grow 
attached to vertical walls, in small caves, or 
on large cobbles; they are directly in contact 
with loose sediment. 

The solitary forms seem more abundant 
in deep water than colonial ones, a situation 
opposite to that in shallow water areas out- 
side of the Galapagos Islands. This may be 
due to the collecting method: solitary ascid- 
ians, especially large ones, are easily seen 
from the submersible and can be collected, 
but colonial ascidians normally live in crusts 
on the undersides of rocks or pebbles, or in 
deep crevices, and so escape notice. The col- 
onies reported here were attached to cob- 
bles. 

Of the eight species collected, two are co- 
lonial. They both belong to the family Poly- 
clinidae (order Aplousobranchiata). One is 
a new species of the genus Polyclinum, 
named P. johnsoni after the submersible and 
ship. The other species is probably Aplidium 
californicum (Ritter & Forsyth 1917), one 
of the most common Polyclinidae on the 
shore nearby, but the samples are too small 
and not fully enough developed to verify 
their identity. The other ascidians belong to 
surprisingly diverse families: one Cionidae, 
two Ascidiidae, and one Octacnemidae 
among the order Phlebobranchiata; and one 
Styelidae and one Pyuridae in the Stolido- 



VOLUME 102, NUMBER 1 



15 



branchiata. Several of these solitary ascid- 
ians have deep-sea characteristics: Ciona 
pomponiae, n. sp. has a particularly solid, 
noncontractile tunic, and a very small gut 
compared to the branchial sac; Situla rine- 
harti, n. sp. belongs to a typical bathyal fam- 
ily (Octacnemidae); Styela psoliformis, n. 
sp. has the same characteristics as other 
deep-water species of that genus. 

The presence of a deep-water ascidian 
fauna around the Galapagos, different from 
the shore species collected by SCUBA, is 
not surprising. The bottom temperature at 
500-700 m was between 7°C and 5.5°C. The 
water near the bottom was rich in suspend- 
ed particles, but it is not known whether the 
particles were plankton except for the nu- 
merous crustacean larvae observed. 

Type specimens are deposited at the 
Smithsonian Institution, National Museum 
of Natural History (USNM). Sorted sam- 
ples received a Sea Pharm number (SP), in- 
dicated here for each specimen. 

Polyclinidae 

Polyclinum johnsoni, new species 

Fig. 1A, B, C, D 

Holotype. -USNM 18249; SP 25-XI-86- 
3-17. 

Description. — Two colonies were collect- 
ed close to Punta Espinosa, Fernandina Is- 
land, at 306 m depth, attached to a sponge 
underneath a large piece of rock. The colony 
retained as a taxonomic voucher is a cush- 
ion 2.5 cm in diameter. Color in life pale 
yellow with translucent tunic through which 
yellow zooids were clearly visible. Branchial 
sac and internal papillae obscured by an or- 
ange-yellow pigment. Zooids situated on 
either side of cloacal canals that converge 
towards the center of the colony where col- 
ony's sole common cloacal aperture opens. 

The zooids have a large size range. Tho- 
rax large compared to the abdomen and the 
post-abdomen (Fig. 1A). Oral siphon star- 
shaped with six triangular pointed lobes 
above a strong sphincter that closes the si- 



phon into a narrowed tube. Oral tentacles 
vary in number with the zooids and are in- 
serted in at least three circles, with three or 
four orders of length, the largest being dor- 
sal. At least 40 oral tentacles can be counted 
in the largest zooids. The peripharyngeal 
band lined with two high laminae and not 
curved at the level of the dorsal tubercle. 
Cloacal aperture close to the oral siphon and 
provided with a sphincter which may make 
the aperture into a tube when contracted. 
Dorsal lappet (Fig. 1 A) well developed and 
inserted on a large base; not wide but its 
length, which varies according to the loca- 
tion of the zooid in the colony, can reach at 
least the length of the thorax; has two lon- 
gitudinal muscular bands (Fig. 1A). There 
is a rounded button under the cloacal ap- 
erture, as is usual in the genus. Thoracic 
musculature is weak; longitudinal fibers 
from the oral siphon stop very quickly and 
do not pass the level of the first row of stig- 
mata; there is an average of eight muscles 
on each side. 

Branchial sac, in both the live and pre- 
served animals, visible by transparency; the 
transverse vessels underlain by muscular fi- 
bers and bear high papillae (Fig. IB) con- 
taining an orange-yellow opaque pigment. 
The number of rows of stigmata is variable 
according to size of zooids, and divisions of 
stigmatal rows commonly seen. In the col- 
ony studied there is a maximum of 16-17 
rows of stigmata. In this example the fourth 
row is divided as follows: 25 stigmata on 
the right and 23 on the left side with 16 
papillae on the right and 15 on the left on 
the fourth transverse vessel. Branchial sac 
not perforated along the endostyle or under 
the rapheal papillae. Dorsal languets short 
and wide, only slightly displaced to the left 
side. The whole branchial sac has a rect- 
angular shape with a right angle at the base 
of the endostyle (Fig. 1A). 

Abdomen slightly twisted; esophagus 
short; the stomach placed anteriorly is sud- 
denly cut in its pyloric part (Fig. 1C). There 
is a ring-like post-stomach; intestine not 



16 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Polyclinum johnsoni: A, Zooid; B, Detail of the branchial sac; C, Digestive tract; D, Larva. Ciona 
pomponiae: E, Animal without tunic, right side; F, Neural region; G, Gonads on the digestive loop; H, Extremities 
of gonoducts. 



VOLUME 102, NUMBER 1 



17 



further differentiated into regions (Fig. 1C). 
Anus opens at the level of the 7th or 8th 
row of stigmata and has two lips. 

Post-abdomen egg-shaped, linked to the 
abdomen by a thin peduncle (Fig. 1 A); ovary 
is anterior followed by a cluster of testes. 
The heart is terminal. One or two vascular 
appendices extend from the post-abdomen 
into the test. 

Larvae (Fig. 1 D) incubated in the cloacal 
cavity. The zooids contain 1-3 larvae. The 
trunks measure 550 /um. They have 3 ad- 
hesive papillae, 7 pairs of thick epidermal 
papillae arranged on a line on each side of 
the body except for one pair which is lightly 
displaced inside, and there are two areas of 
epidermal vesicles on each side, one dorsal 
and the other ventral (Fig. ID). 

Remarks. — This species differs from all 
other Polyclinum with more than 15 stig- 
mata rows in having higher papillae on the 
transverse bars, a large number of stigmata 
per row, and a peculiar distribution of the 
pigments in the colony. The genus Polycli- 
num is very homogeneous and the anatom- 
ical differences between the species are slight, 
so it is not surprising that deep-water species 
do not differ substantially from shallow- 
water species. 

There is only one Polyclinum species 
known to live deeper than P. johnsoni, n. 
sp.; P. neptunium Hartmeyer, 1912, col- 
lected at 106 m and 318 m off the Cape of 
Good Hope. 

The species is named after the submers- 
ible from which it was collected. 

Cionidae 

Ciona pomponiae, new species 

Figs. IE, F, G, H, 2 A, B 

Holotype. -USNM 18247, SP 21-XI-86- 
2-1. 

Description. —The unique specimen mea- 
sures 15 cm in length; attached by the left 
posterior side; oral siphon terminal; cloacal 
aperture located l h the way down the dorsal 



side (Fig. 2A). Both siphons sessile. Test soft, 
gelatinous, colorless in life and free of for- 
eign particles; is at least 5 mm thick and 
does not appear to consist of several layers 
as in Ciona intestinalis. Body contracted in- 
side tunic and no longer attached to it, even 
at the level of the siphons; size of body re- 
duced to 6.9 cm. Tunic much less retractile 
than in other Ciona species. 

Musculature on each side (Fig. IE) forms 
four muscular ribbons initiated from the oral 
siphon and two ribbons from the cloacal 
siphon. The two most dorsal muscles of the 
oral siphon and the one most ventral muscle 
from the cloacal siphon gather to form one 
ribbon in the posterior part of the body. The 
two ventral muscular bands issuing from 
the oral siphon become narrow before 
reaching the posterior part of the body and 
two muscles reach the bottom of the bran- 
chial sac (Fig. IE). There is a network of 
transverse fibers. This small number of 
muscular bands is peculiar to this species. 

There are few oral tentacles (16 large ones 
and several very small) placed without or- 
der on a crest. The length of some tentacles 
allows them to protrude through the oral 
aperture. The peripharyngeal band consists 
of two unequal laminae; the anterior thin 
and high, the posterior low and thick. There 
is a marked dorsal V in which the neural 
duct opens as a simple slit (Fig. IF). Neural 
gland close to the nerve ganglion. Dorsal 
languets have an increased length poste- 
riorly, measuring 6 mm in the anterior v h 
and 18 mm near the esophagus. 

Branchial sac (Fig. 2B) thin, the trabec- 
ule connecting it to the body wall long. 
Transverse vessels of several orders regu- 
larly distributed; those of the first order wide 
and protrude into the branchial cavity, those 
of the third or fourth orders are thin laminae 
protruding into the branchial cavity. They 
bear large papillae that are flattened longi- 
tudinally and form hooks above the longi- 
tudinal bars. Longitudinal bars thin; stig- 
mata elongated, generally numbering six to 



18 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 










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Fig. 2. Ciona pomponiae: A, Group of in situ individuals (from a submersible videotape); B, Part of branchial 
sac. C: Ascidia clementea part of the branchial sac with dorsal lamina. Situla rineharti: D, Part of the branchial 
sac; E, Habitus left side; F, The same without tunic. (Scales: A = 10 cm; B, C, D = 1 mm; E, F = 1 cm.) 



VOLUME 102, NUMBER 1 



19 



eight in a mesh; most crossed by a high 
parastigmatic vessel bearing papillae (Fig. 
2B); often divided under this vessel. If the 
division is complete, the stigmata on each 
side of the vessel are shorter than the others, 
and a very thin new parastigmatic vessel 
without papillae can occur above them. 

Digestive tract clearly situated under the 
branchial sac; enlarged stomach has internal 
plications visible by transparency; most 
posterior fold prolonged and ends in a small 
ampulla at its junction with the intestine; 
intestinal wall thin and gut contents can be 
seen through it. Rectum elongated and ends 
in a non-lobated narrow anus; posterior in- 
testine encircled by a well developed pyloric 
gland. 

The pear-shaped ovary is located under 
the intestinal loop (Fig. IE, G). Testis dif- 
fuse, consisting of numerous acini which coat 
the intestinal wall; ducts join to form a com- 
mon sperm duct at the level of the anterior 
part of the ovary; i.e., oviduct and common 
sperm duct originate at the same point. 
Gonoducts contain large numbers of eggs 
(400 /urn in diameter) and spermatozoa, and 
pass beyond the anus; oviduct opens in a 
flat horseshoe-shaped papillae (Fig. 1H). 
Male papilla has several small apertures; 
there is no pigment spot at this level. 

Remarks. — This species is very uncom- 
mon for a Ciona. Although very few species 
occur in shallow waters, the genus is better 
differentiated deeper on the continental 
slope. In the northeast Atlantic is found 
Ciona gelatinosa Bonnevie, 1896 and Ciona 
imperfecta Monniot & Monniot, 1977, and 
in the northeast Pacific Ciona mollis Ritter, 
1907. 

Ciona mollis, known from California at 
201 1 m depth off San Nicolas Island, differs 
from the new species in several important 
characters: C. mollis's size is smaller (3 cm 
maximum) and its tunic is very soft. There 
are 6 muscle bands on each side of the body, 
all of them reaching the posterior part where 
they fuse. There is circular musculature only 
on the siphons. The oral tentacles are very 



numerous, more than 200, while the present 
specimen, much larger, has only 16. This 
difference is perhaps not as significant as the 
reduction of the number of oral tentacles 
even with increased size in Ascidia translu- 
cida according to Millar (1960) and in 
Cnemidocarpa verrucosa according to Mon- 
niot (1978). In C. mollis, furthermore, the 
digestive tract does not have the same shape 
as in this specimen, C. mollis's intestine 
curves immediately after the stomach, and 
its gonoducts do not open near the anus. 
Ciona mollis seems closer to the Atlantic 
species C gelatinosa Bonnevie 1896, re- 
described by C. Monniot ( 1 970), in the mus- 
culature (7 bands), the shape of the gut, and 
the location of the genital papillae. 

The species is named after Shirley Pom- 
poni, co-chief scientist of the cruise. 



Octacnemidae 
Situla rineharti, new species 
Figs. 2D, E, F, 3A, B, C, D 

Holotype. —USNM 18244; paratype 
USNM 18245; SP 14-XI-86-1-2 and 18- 
XI-86-2-11. 

Description. —Among several samples 
collected, two specimens have been exam- 
ined: one was designated as the holotype, 
collected north of Floreana (Santa Maria Is- 
land) at 790 m depth, and another (para- 
type) collected south of Genovesa (Tower 
Island) at 695 m depth. All specimens col- 
lected are approximately the same size, al- 
most spherical, with a cartilaginous white 
tunic that is translucent more than trans- 
parent (Fig. 2E). Holotype ovoid, attached 
by the small ventral end (7 x 5 x 5.2 cm). 
The part of the tunic attached to the sub- 
stratum torn but represents a narrow area 
of attachment; in other specimens there is 
no particular differentiation of the attach- 
ment area. Oral aperture a gaping transverse 
slit, made of two large lips located at half 
the height of the body (Figs. 2F, 3A). Dorsal 
lip protrudes more than ventral one. Cloacal 



20 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



aperture not obvious; it is a simple hole at 
the upper part of the body slightly displaced 
to the dorsal side (Fig. 3A). Tunic on the 
two lips of the oral siphon is thin, cartilag- 
inous, and glossy, radially wrinkled, and dif- 
ferent from the thick test (at least 3 mm) 
around the rest of the body. Border between 
the two kinds of tunic marked by a groove 
at the limit of the lips. 

Removed from the tunic, the body ap- 
pears composed of three parts: the oral si- 
phon formed of two well developed lips, a 
branchial cavity widely spread separated 
from the oral siphon by a ring of tentacles, 
and a "nucleus" containing the gut and go- 
nads. 

The musculature is complex (Fig. 3A, B). 
The muscles of the oral siphon, independent 
of the body musculature, include radial fi- 
bers regularly distributed on both lips; these 
end at the circle of oral tentacles and the 
peripharyngeal groove. The oral siphon's 
circular fibers are also regularly placed, 
spread over both lips and gathered at the 
lip corners. At each of those corners is a fan 
of fibers belonging to the radial muscula- 
ture, connecting the corners to the large 
muscle located at the level of the ring of oral 
tentacles. 

Musculature of the cloacal siphon con- 
sists of radial and circular fibers, weakly de- 
veloped and anastomosed. It is limited to 
the immediate vicinity of the siphon. 

A large, strong ribbon of muscle fibers 
encircles the body above the ring of oral 
tentacles, and has a horseshoe shape opened 
on each side of the endo style at the level 
where the body attaches to the substratum 
(Fig. 3A). A muscular string binds the rib- 
bon's two free extremities. There are also 
two areas of transverse muscles dorsally, and 
ventrally two bundles more or less parallel 
to the endostyle, connected to the "string" 
linking the posterior extremities of the large 
horseshoe muscle ribbon. 

Oral tentacles (Fig. 3C, D) situated on a 
rim located posterior to the horseshoe mus- 
cle-band. Dorsally the rim is low; its height 



increases ventrally where it is lined by a flat 
muscular band. Tentacles have a unique leaf- 
shaped structure, narrowed at their inser- 
tion on the rim. On the anterior side of each 
tentacle is an ampulla which stains very 
deeply with hemalum. The tentacles are of 
two regularly alternating sizes (Fig. 3C). 

The peripharyngeal groove is far from the 
ring of tentacles in the dorsal half of the 
body but comes close to it in the ventral 
part. Dorsally, a series of irregular crests 
separate the peripharyngeal groove and the 
tentacles and they may be considered as the 
anterior rim of the peripharyngeal groove. 
Dorsally the peripharyngeal groove is deep- 
ly incurved (Fig. 3D) to reach the dorsal 
tubercle. The latter is rather far from the 
neural ganglion; its aperture is a simple hole. 
The neural gland is located on the left pos- 
terior side of the neural ganglion. 

There is no differentiated dorsal lamina. 
The restricted space between the dorsal tu- 
bercle and the opening of the esophagus is 
marked by transverse elevations continuous 
with more or less regular internal transverse 
crests on the branchial sac; these elevations 
can be considered as transverse vessels. 
There is a short endostyle originating near 
the oral tentacles and ending far from the 
entrance of the esophagus (Fig. 3A, B). At 
its posterior extremity, on the left, is a ret- 
ropharyngeal groove which initially consists 
of a single rim but divides in two wide and 
flat ridges. 

The entire surface placed between the 
peripharyngeal band and the branchial stig- 
mata is covered with small papillae. Bran- 
chial cavity cup-shaped with the bottom 
close to the digestive tract. The perfora- 
tions, limited to two small areas in the flat 
bottom of the cup on each side of the re- 
tropharyngeal band, lie in thick tissue. The 
perforations are oval (Fig. 2D) and generally 
grouped in pairs; they are not ciliated, and 
are larger than in the other known species 
of Octacnemidae. 

The digestive tract forms a simple loop 
(Fig. 3B); esophagus opens by a simple hole 



VOLUME 102, NUMBER 1 



21 




Fig. 3. Situla rineharti: A, Right side of the animal without tunic; B, Postero-dorsal side; C, Oral tentacles; 
D, Peripharyngeal groove indentation, tentacles and neural ganglion. Ascidia clementea: E, Left side of the body, 
tunic removed; F, Dorsal part of the tentacle ring and peripharyngeal groove; G, Intestinal loop with ovary and 
oviduct. 



22 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



in the branchial sac; stomach is wide and 
has a glandular wall which appears some- 
what wrinkled but has no internal plications 
(Fig. 3A, B); its external oblique ridges may 
be due to contraction. The intestine is thin 
and transparent, ending in a plain-edged 
anus placed beside the esophageal entrance. 
The digestive tract and gonads are embed- 
ded in a thick mesenchyme. The retro- 
pharyngeal groove lies beside the middle of 
the gut loop and indicates that the stomach 
is on the right side. 

The gonads (Fig. 3 A, B) consist of a mas- 
sive and protrusive ovary located between 
the two branches of the digestive loop; testis 
is composed of numerous acini; it is flat and 
covers the ventral part of the ovary and each 
side of the intestine. The oviduct opens on 
the medial side of the intestine a short dis- 
tance before the anus. The spermduct, full 
of spermatozoa, opens next to the oviducal 
aperture on a small papilla. The heart lies 
alongside the intestine. 

Remarks. — Octacnemidae having an oral 
aperture with two large lips belong to four 
genera: Dicopia Sluiter, 1905; Megalodico- 
pia Oka, 1918; Situla Vinogradova, 1969; 
and Cibacapsa Monniot & Monniot, 1983. 
Cibacapsa is characterized by the total ab- 
sence of a branchial sac. The distinctions 
between the other genera are less obvious. 
The difference between Dicopia and Meg- 
alodicopia is the presence in the latter of a 
peduncle, which in our opinion is not a dif- 
ference of much importance. Dicopia differs 
from Situla by the presence in Dicopia of a 
true branchial cavity with a conical perfo- 
rated area and oral tentacles forming a ring 
of a smaller diameter than that of the bran- 
chial cavity. In Situla the branchial cavity 
opens widely and the perforated area is in 
a flat disk around the digestive tract. The 
Galapagos species is closer to Situla. The 
main differences between the species of Di- 
copia and Situla are summarized in Table 
1 . The new species is the only one having a 
large imperforate space between the 
peripharyngeal groove and the branchial tis- 



sue, and its oral tentacles have a very orig- 
inal structure. 

Ritter (1907) has incompletely described 
a species of Octacnemidae collected off San 
Diego, California: Benthascidia michael- 
seni. His account suggests he did not un- 
derstand the anatomy of the animal very 
well. The two specimens of that species pre- 
served at the Smithsonian Institution are 
dissected and their organs isolated, and so 
it is impossible to reconstitute their original 
disposition in the animal. In any case Rit- 
ter' s material cannot be of the same species 
as the new Galapagos one. 

The species is named after Kenneth Rine- 
hart, co-chief scientist of the cruise. 

Ascidiidae 

Ascidia clementea, Ritter, 1907 

Figs. 2C, 3E, F, G 

SP 23-XI-86-3-27. 

Description. — One specimen was collect- 
ed north of Isabella Island at 335 m depth. 
The ovoid body is attached to the substra- 
tum by the posterior part of the left side. 
The maximum length of the animal is 6.8 
cm. Oral aperture terminal; cloacal aperture 
located at the anterior x h, the distance be- 
tween the siphons being 2.9 cm. The tunic 
in life is yellowish, covered with small cy- 
lindro-conical papillae. Papillae and grooves 
of the tunic are pigmented with ochre; both 
apertures are 6-lobed. The test is partially 
covered with epibionts: foraminifera, poly- 
chaete tubes, bryozoans and molluscan egg- 
capsules. One of the cheilostome bryozoans 
consists of thecal rows, each zooid having 
a long ventral extension penetrating deeply 
into the ascidian's tunic; the bryozoan col- 
ony is raised above the tunic surface by these 
extensions. Thickness of the tunic averages 
2 mm; its internal side is soft and translu- 
cent. 

Body wall thin and transparent; the mus- 
culature forms 2 bands of transverse parallel 
fibers— a well developed dorsal band, and 
a smaller ventral band— and longitudinal 



VOLUME 102, NUMBER 1 



23 



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24 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



muscle fibers extend from the oral siphon. 
On the right side all the muscles spread and 
intermingle to make a loose network. On 
the left side the longitudinal fibers extend 
only a short way from the oral siphon, and 
most of the left body wall is devoid of mus- 
culature. 

Seventy oral tentacles arise from an ele- 
vated rim that has a muscular ring. The 
tentacles are of at least four orders of size, 
regularly alternated although the smallest 
ones are not developed everywhere. Peri- 
pharyngeal groove made of two unequal 
laminae, the anterior one being larger, is 
close to the tentacles, and the space between 
is covered with thin papillae (Fig. 3F). It 
does not make a deep ' V dorsally; the dorsal 
tubercle is low, J-shaped. Neural ganglion 
distant from the dorsal tubercle; neural duct 
is twice the length of the neural ganglion. 
Rapheal band consists of two laminae which 
measure five times the length of the dorsal 
tubercle. Posteriorly, its edge is higher and 
carries teeth that coincide with the trans- 
verse vessels (Fig. 2C). After passing around 
the esophageal entrance it becomes irregu- 
lar. On the right side close to the esophagus 
the transverse vessels end without differ- 
entiation. 

Branchial sac thin and flat (Fig. 2C). At 
the level of the top of the intestinal loop 
there are 1 00 longitudinal vessels. Main pa- 
pillae are located at the junctions of the lon- 
gitudinal and transverse vessels, and almost 
everywhere in the branchial sac there are 
smaller intermediate papillae, which are not 
linked to the presence of parastigmatic ves- 
sels. These vessels exist only where there is 
duplication of a stigmata row. There are 
four to six stigmata in a mesh, except on 
the right side, where the meshes located near 
the esophagus have more than 20 stigmata. 

The gut forms an open loop with an ac- 
centuated secondary curve (Fig. 3E). The 
stomach appears plicated externally (Fig. 3E, 
G). The intestine is deeply embedded in a 
mass of large renal vesicles which spread 
over the body wall. The anus is attached to 
the body wall and has two slightly scalloped 



lips. The specimen was not sexually mature. 
The ovary, a network of tubes inside the 
intestinal loop, does not spread out onto the 
intestine (Fig. 3G). We have not found the 
testicular acini that Ritter (1907) shows 
against the stomach. 

Remarks. —A. clementea has not been re- 
ported since its description in 1907 based 
on eight specimens: one collected by the Al- 
batross near San Clemente Island at 654- 
704 fathoms (station 4405), and seven spec- 
imens at 2 1 .8 miles to the south of San Nic- 
olas Island, 1100 fathoms (station 4425). 
There are a few small differences from Rit- 
ter' s description: he does not mention the 
tunic papillae and counts eight lobes at the 
oral siphon instead of six. His figure 33 
shows more than 75 small oral tentacles all 
of equal size. He describes "deep narrow 
plications" of the branchial sac, but his fig. 
34 does not show this. In his discussion, he 
recognizes that these plications do not al- 
ways exist. All the other characters of our 
specimen are identical to his description. 

Ascidia fusca, new species 
Fig. 4A, B, C, D 

Holotype. -USNM 18248; SP 20-XI-86- 
1-25. 

Description. —The sole specimen was tak- 
en at 375 m depth southeast of San Salvador 
(James) Island. Maximum length 5.6 cm; it 
was lying on its left side. Apertures distant 
from each other (4.6 cm); have an undeter- 
mined number of external large soft swell- 
ings. Tunic dark brown in life (thus the 
species name fusca) and its surface wrin- 
kled. Some foraminifera and sedimentary 
particles adhere to the settlement surface; 
elsewhere on the tunic are small epibionts, 
mostly bryozoans. The test is soft and its 
internal layer is colorless. 

Musculature strong and lies entirely on 
the right side; is interrupted before it reach- 
es the endostyle and dorsal lamina; com- 
prises one ventral and one dorsal field, which 
intermix in the middle of the right side. The 
fields' fibers are close together and nearly 



VOLUME 102, NUMBER 1 



25 



joined; siphonal musculature weak. On the 
right side of the oral aperture an array of 
about 1 bands blends into both transverse 
fields and disappears in the first l A of the 
body. 

Oral tentacles arise from posterior part of 
a velum linked by a muscular ring; there are 
about 40 large ones and as many very small 
ones alternating and irregularly developed; 
bases crooked; peripharyngeal groove made 
of two unequal laminae: anterior high and 
thin and the posterior forming a flat ribbon. 
There is a large space between the tentacle 
ring and the peripharyngeal groove which 
is indented into a deep dorsal V (Fig. 4B). 
Dorsal tubercle is urn-shaped, opening with 
a simple aperture (Fig. 4B). Neural ganglion 
far from the dorsal tubercle at a distance of 
3 times its length (0.6 cm in the type spec- 
imen). Dorsal lamina has 2 rims which ex- 
tend to the posterior part of the neural gan- 
glion and then form a single lamina that is 
thin and high. This carries digitiform pa- 
pillae which prolong the transverse sinuses. 
The lamina becomes progressively lower and 
somewhat irregular as it nears the esopha- 
gus, where it forms a low, toothed crest. At 
this level on the right side, one of two trans- 
verse vessels ends in a large lanceolate pa- 
pilla. 

Branchial sac flat; at the level of the top 
of the intestinal loop there are 70 longitu- 
dinal vessels on the right and 55 on the left, 
the left side being much more narrow than 
the right side. Longitudinal vessels very thin 
and carry fingerlike main papillae where they 
intersect transverse vessels. There are nei- 
ther secondary papillae nor parastigmatic 
vessels. The meshes are elongate and con- 
tain 2 to 3 stigmata. 

Gut (Fig. 4A, C, D) small; stomach, clear- 
ly enlarged, has some plications only visible 
on its medial side. Intestine inserted inside 
the wall of the mantle. The 2 lips of the anus 
have many small lobes. In this specimen a 
translucent swelling is located close to the 
anus. It has no discernable internal structure 
and we do not know if it is a specific char- 
acter or an artefact. 



The ovary (Fig. 4C, D) consists of a mass 
of tubes projecting internally, with edges 
growing partly over the inside of the intes- 
tine. The testicular lobules lie on the 2 sides 
of the gut. The gonoducts open together next 
to the anus. The renal vesicles are small, 
almost invisible. 

Remarks. —Ascidia fusca differs from 
A.vermiformis (Ritter 1913), another deep- 
water Ascidia in the East Pacific, by the for- 
mer's external shape, the length of its cloacal 
siphon and the long posterior extension of 
its branchial sac. As in several of the rare 
Ascidia known from the continental and is- 
land slopes, Ascidia fusca has a relatively 
small gut compared to the branchial sac, an 
open primary gut loop, and a flat branchial 
sac. 

The species name refers to the dark tunic 
of the animal. 

Styelidae 

Styela psoliformis, new species 

Fig. 4E, F, G 

Holotype. -USNM 18246, SP 16-XI- 
86-1. 

Description. —The sole specimen was 
found in a cavity of a basaltic cobble near 
Espanola island at 567 m depth. The visible 
part of the still attached body is an oval disk, 
14 x 11 mm, white, covered with low, 
round, plate-like papillae which have a vas- 
cular dilation full of blood granules. Around 
the disk the tunic is thinner and also has 
vascular ampullae; it strongly attaches the 
animal to the substratum. Both apertures 
protrude a distance of 6 mm, but they are 
located in depressions, so they do not ex- 
tend beyond the body surface after contrac- 
tion. Tunic of the central disk opaque and 
internally pearly. The rest of the body has 
a thinner tunic that is soft and transparent. 
Inside the tunic the body is flattened against 
the substrate. The dorsal part of the body 
wall is opaque and muscular, the ventral 
side thin and transparent without muscu- 
lature. At the boundary between the dorsal 
and ventral side the mantle has a ring of 



26 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Ascidia fusca: A, Left side of the body, tunic removed; B, Dorsal tubercle; C, External side of the gut 
and gonads; D, Internal side of the gut and gonads. Styela psoliformis: E, Ventral side of the body, tunic removed; 
F, Internal side of the body, branchial side removed; G, Gonad. 



tissue rich in tan blood lacunae clearly seen velum. There are at least four different sizes 

by transparency. which regularly alternate. The oral tentacles 

About 30 oral tentacles are attached to a are wide and transversally flattened. The 

thin crest located posteriorly to a large oral peripharyngeal groove has 2 equally pro- 



VOLUME 102, NUMBER 1 



27 



truding rims. It is far from the tentacles on 
the ventral part, nearer on the sides, and 
dorsally makes a little but distinct V; it is 
only curved around branchial folds number 
1 and 2. The opening of the dorsal tubercle 
has the shape of a C opened on the left. The 
neural ganglion is in contact with it. The 
dorsal lamina is high and smooth. 

Branchial sac has 4 folds on each side in 
the anterior part only; fold number 4 dis- 
appears in the posterior % of the branchial 
sac. Longitudinal vessels very numerous and 
difficult to count; there are as many as 15 
vessels between the folds and more than 20 
on folds 1 and 3. Fold 4 has more than 15 
longitudinal vessels anteriorly. The folds are 
high and the longitudinal vessels close to 
each other, with only one or two elongated 
stigmata per mesh. Parastigmatic vessels are 
generally present. 

Gut a narrow loop (Fig. 4E, F). Stomach 
and intestine weakly linked to the ventral 
side of the body wall but rectum strongly 
united to dorsal side. The globular stomach 
has 14 obvious plications. A very small cae- 
cum is located near the gastric aperture of 
the pyloric gland. Intestine long; the short 
rectum opens by a wide, lobed anus. 

There are two gonads on each side (Fig. 
4E, F), weakly linked to the body wall; each 
is made of a double series of testes (Fig. 4G). 
The central ovary of each gonad spreads 
onto the internal part of these lobes, which 
makes this structure intermediate between 
the genera Styela and Cnemidocarpa. The 
gonoducts open together at the margin of 
the dorsal disk of the body wall. 

Endocarps are evenly distributed on the 
ventral side of the body (Fig. 4F). They are 
very numerous and protrude along the ring 
of tissue rich in tan blood lacunae which 
marks the boundary between dorsal and 
ventral halves of the body. The dorsal part 
is devoid of endocarps. The cloacal aperture 
is circled by thin cloacal tentacles arising 
from a crest (Fig. 4F). 

Remarks. —This species has several char- 
acters which are similar to Styela milled 
Ritter, 1907, collected frequently from 900 



to 4000 m depth along the Pacific coast from 
California to Chile and also recorded in the 
south Atlantic and the Indian Oceans. The 
oral tentacles, dorsal tubercle, dorsal lami- 
na, branchial sac, and gut are similar in both 
species. In contrast, S. milleri has only one 
gonad on each side and they are different in 
structure from those of S. psoliformis. The 
new species has the same appearance as S. 
similis C. Monniot, 1969 from the conti- 
nental slope of Europe and West Africa, and 
these species have many anatomical char- 
acters in common due to their similar body 
shape. S. similis also has only one gonad on 
each side, but its structure differs. 

The gonad of S. psoliformis is morpho- 
logically intermediate between typical Sty- 
ela gonads and the very peculiar gonad of 
Cnemidocarpa pfefferi (Michaelsen, 1904) 
shown in Millar 1960; that large (5 cm), 
round species lives on the slope of South 
Georgia and South Shetland Islands. It has 
two gonads on each side, each consisting of 
a lobed ovary with each lobe encircled by 
testes. The structure of the gonad of S. pso- 
liformis resembles that of species of Pyura, 
but this is only a convergence: all other char- 
acteristics of the present species are typical 
of the Styelidae family. 

The animal looks like a psolid holothu- 
rian after which it is named. 

Pyuridae 

Halocynthia hispida Herdman, 1881 

Fig. 5 

Synonymy: see Kott, 1985:342, fig. 169. pi. 

7F, Millar (1988). 
SP 24-XI-86-3-9 and 27-XI-86-1-9. 

Description. —This species has two dif- 
ferent types of morphology on the Gala- 
pagos Islands: a hard form with a thick tunic 
collected in shallow water by SCUBA and 
a larger form with a soft hairy tunic collected 
from deep water by the submersible. The 
two forms also differ in internal structure, 
and only a very careful analysis of all char- 
acters has convinced us that they belong to 
the same species. A reduction of the organs 



28 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



linked to increased depth and decreased food 
supply has been described in ascidians 
(Monniot & Monniot 1978), but the case of 
H. hispida in the Galapagos is extreme. 

The shallow-water form was collected at 
Bainbridge Rocks, east of San Salvador Is- 
land, at a depth of 17 m. It was firmly at- 
tached by its ventral side; both apertures 
are erect, at almost the same level. The spec- 
imen measures 10 x 7 x 5 cm, the aper- 
tures separated by 4 cm. The side facing the 
light is dark red in life. The tunic is wrinkled 
and covered with small smooth spines that 
become larger near the siphons, the largest 
grouped on tubercles. Around the apertures 
there are large branched spines 3 to 4 mm 
in length. The thickness of the cartilaginous 
tunic reaches 5 mm. 

The body wall is 2-3 mm thick and fleshy. 
The muscles are well developed: the exter- 
nal circular ones make a continuous layer, 
the internal radial muscles are gathered to- 
gether in strong bands. The oral tentacles 
are located behind a strong sphincter; they 
number about 20 in three orders and are 
only weakly pinnate. The peripharyngeal 
groove has 2 equal rims and shows a clear 
dorsal V. The dorsal tubercle is large, com- 
plex and protruding (Fig. 5D). The dorsal 
lamina has a double series of languets. 

The middle level of the branchial sac has 
10 high folds on each side. Anteriorly and 
posteriorly, additional ventral folds appear, 
so the maximum number of folds in this 
specimen is 13 on the right and 12 on the 
left side. The distance between 2 longitu- 
dinal vessels is the same on the folds as 
between them, so it is difficult to distinguish 
the base of a fold. In the ventral part of the 
branchial sac there are only 2 or 3 longi- 
tudinal vessels between folds, the meshes 
are larger there and they may have a dozen 
stigmata. 

The gut (Fig. 5C) is voluminous and oc- 
cupies most of the left side. The esophagus 
is long and the stomach is covered with a 
hepatic gland of two parts: the anterior la- 
mellar and the posterior cauliflower-like. 



The intestine is attached to the body wall 
and has a smooth edged bilobed anus. 

The gonads are very numerous ( 1 3 on the 
right and 10 on the left) and so massed and 
interpenetrated (Fig. 5C) that they can only 
be counted by their genital papillae. On the 
left side they cover the descending limb of 
the intestine. Each gonad consists of an ovary 
surrounded by lobes of testes. 

There are numerous endocarps on the 
body wall, inside the gut loop and even pro- 
truding from the anterior part of the stom- 
ach (Fig. 5C). 

The deep-water form is rounded; the larg- 
est specimen measures 7x5x4 cm with 
a distance of 4 cm between the apertures. 
The color in life is tan-brown with some 
amounts of red near the siphons. The entire 
tunic bears short branched spines with much 
larger ones on the siphons; between the 
branched spines there are smaller simple 
ones, often placed in rings. The test is thin 
(1 mm) and flexible. Internal organs are vis- 
ible through the thin, transparent body wall. 
The external layer of circular muscles is thin. 
The internal radial muscles are gathered to- 
gether in weak ribbons. 

When the tunic is removed, the oral ap- 
erture has 4 low lobes with undulated edges. 
There is a small plain velum at the base of 
the oral siphon. The oral tentacles are not 
numerous (36) and occur in at least 4 orders. 
Between the largest tentacles on the tentacle 
crest there are numerous small digitiform 
ones. The largest tentacles measure up to 
13 mm but only have about 20 primary 
branchings, which are themselves thin and 
poorly branched in turn. The peripharyn- 
geal groove has 2 equal laminae; it is curved 
around each branchial fold and makes a large 
dorsal V. The dorsal tubercle is round and 
has a C-shaped aperture open on the ante- 
rior left side (Fig. 5A). The neural ganglion 
is elongated (1 5 mm), and thin. Nerves from 
it are clearly visible: 2 anterior and 2 pos- 
terior ones. The visceral nerve follows the 
dorsal lamina and is accompanied by a neat 
dorsal band; it originates on the right pos- 




Fig. 5. Halocynthia hispida: A, Deep-water form, internal side of the body, branchial sac removed; B, Detail 
of the right gonad; C, Shallow-water form, internal side of the body; D, Dorsal tubercle of the shallow-water 
specimen. 



30 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tenor side of the neural ganglion. In its an- 
terior part the dorsal lamina consists of a 
double row of languets; the left line has true 
languets and is more developed than the 
right line, which is made of prolongations 
of the right transverse branchial vessels. 

The branchial sac has 10 folds on each 
side. The branchial formula is: 

R.E.2(8)2(13)2(15)3(17)2(18)2(20)2(22)2 
(2 1)3(20)3(1 6)2.DL 

L.DL.2(1 7)2(2 1)2(22)3(22)2(2 1)2(1 9)3(1 5) 
2(13)2(1 1)1(10)O.R 

The 10th (most ventral) fold on the left is 
only present in the anterior part of the bran- 
chial sac. The folds are very high and largely 
cover each other. The longitudinal vessels 
are raised, and there are only two or three 
between the folds. We counted 24 stigmata 
in a median mesh between two folds and 8 
to 1 in meshes close to the base of the folds. 
It is only at the crest of the fold that the 
number of stigmata in a mesh go down to 
five or six. We have not observed spiral 
stigmata at the crests of folds. The stigmata 
are not very elongated; they are crossed by 
parastigmatic vessels only when a division 
begins in a transverse row. Between the folds 
the transverse vessels are inflated into am- 
pullae attaining several millimeters, much 
like endocarps. 

Gut short; a long esophagus leads to a 
stomach with a 2-part liver (Fig. 5A), the 
anterior one with parallel ridges and the 
posterior with papillae on crests. The two 
parts of the gland are separated by a nar- 
rowing, but they remain united. At the nar- 
rowing level there are parallel riges, each 
dividing in two in the anterior part (Fig. 
5 A). At the pyloric end of the stomach the 
hepatic gland stops abruptly. The intestine 
is short and translucent. Rectum long; at- 
tached along the dorsal lamina, and the anus 
has two lobed lips. 

On each side there is a cluster of four 
parallel ovaries united at their proximal 



ends. Each cylindrical ovary is surrounded 
by ramified testes (Fig. 5B). Each ovary and 
its testes are wrapped in a common mem- 
brane. The testes give off long thin ducts 
which join at the atrial surface of the ovary 
to form a common spermduct. The com- 
mon spermduct consists of canals coming 
from both sides of the ovary. The sperm- 
duct opens just behind the oviduct on a 
common protruding papilla. The four left 
ovaries originate inside the gut loop and 
open onto the intestine (Fig. 5A). 

The entire body wall is provided with en- 
docarps. The cloacal siphon has a short ve- 
lum similar to the oral siphon. We have not 
seen cloacal tentacles. 

Remarks. —This species is recorded from 
the Galapagos by Millar (1988) and the 
specimens he describes are intermediate be- 
tween our two forms. If we examine point 
by point the differences between our shallow 
and deep populations, we find that they are 
almost all quantitative ones: thickness of 
the tunic and body wall, gonad number, size 
of the gut, and number of branchial folds. 
The difference in the number of oral ten- 
tacles has no taxonomic significance. The 
most important contrast is the reduction of 
the gut in deep-water specimens, and its po- 
sition at the posterior part of the body. The 
gonads are also posteriorly displaced in deep- 
water animals. 

The thinning of the tissues in deep-water 
animals is probably linked to decreased food 
supply. The species of Halocynthia have a 
highly evolved branchial sac which is very 
efficient in filtering phytoplankton, but the 
deep samples certainly have less food avail- 
able. In fact, the branchial sac is the only 
organ not reduced; on the contrary, the 
number of stigmata per mesh increases with 
depth. The reduction in size of the gut is 
correlated with this nutritional phenome- 
non; it is a common characteristic for as- 
cidians living on the slope such as in Ascidia 
(see below). A similar but less accentuated 
situation has been described for Ascidia 



VOLUME 102, NUMBER 1 



31 



challengeri and Corella eumyota (Monniot 
& Monniot 1983) on the slope in the Ant- 
arctic Ocean and at Kerguelen. 

The shifting of the gut and gonads pos- 
teriorly may be a consequence of the loss of 
tunic rigidity. In 1 973 we reported that deep- 
sea ascidians show a tendency to equilibrate 
their visceral masses to keep their balance 
on the substratum. In other species of Halo- 
cynthia the disposition of the gonads on the 
right side seems linked to the tunic rigidity: 
H. ritteri from Japan has a hard tunic and 
gonads located in the middle of the right 
side, while H. papillosa, H. aurantium and 
H. pyriformis, with thin tunics, have gonads 
located posteriorly. 

Except for two Arctic species, the tropical 
and temperate Halocynthia are found in very 
shallow water not deeper than 30 meters. Is 
the presence of deep specimens in the Ga- 
lapagos exceptional? We cannot answer be- 
cause the deep rocky bottoms have not been 
sampled enough, but this species represents 
the most extreme case of ecological dimor- 
phism known in an ascidian. 

Conclusions 

The bathyal ascidian fauna of the Gala- 
pagos does not provide any evidence in sup- 
port of an archaic fauna. All species belong 
to genera and more advanced families which 
are well represented in other oceans and 
which show adaptations here to deep-sea 
life. 

The origin of the fauna is unknown. Rel- 
atively few collections have been made along 
the South American Pacific coast; we cannot 
determine if the new species described here 
are endemic or if they have been carried 
north by the Humboldt Current. The Ga- 
lapagos ascidian fauna differs greatly from 
the assemblage of Antarctic ascidians known 
from the south coast of Chile. Among the 
eight species recorded from the Galapagos, 
only three were previously known: Apli- 
dium californicum, a tropical shallow- water 



shallow-water species probably at its depth 
limit here, Halocynthia hispida, widely dis- 
tributed and known in shallow water from 
both the Galapagos and Australia; and As- 
cidia clementea, a north Pacific species. Sit- 
ula and Ciona are genera which are well 
represented in bathyal areas in other parts 
of the world. It seems unlikely that the 
Humboldt Current contributes to the oc- 
currence or distribution of ascidian fauna 
around the Galapagos islands. 

Observations from the Johnson- Sea- Link 
have shown that ascidians are a very small 
part of the bathyal biomass which is mainly 
comprised of sponges, cnidarians and echi- 
noderms. Other filter-feeding animals, such 
as bivalves or bryozoans, were also very 
seldom seen. Except for Situla, which is 
phytophagous and carnivorous, the other 
ascidians we report have the same diet as 
sponges: small suspended particles and dis- 
solved substances. Sponges in Galapagos 
deep-water habitats are large, abundant, and 
diversified, but ascidians remain scarce even 
though they are relatively large. 

Acknowledgments 

The R/V Seward Johnson and Johnson- 
Sea-Link- 1 cruise was funded by Harbor 
Branch Oceanographic Institution, the U.S. 
National Cancer Institute, Natural Products 
Branch (contract no. N01-CM-67919), and 
SeaPharm Inc. Francoise Monniot is par- 
ticularly thankful to the chief scientist of the 
cruise, K. L. Rinehart and to Shirley A. 
Pomponi who gave her the opportunity to 
participate in the submersible dives and in 
SCUBA collecting. In gratitude two of the 
species reported here are named after them. 

Literature Cited 

Bonnevie, K. 1896. Ascidiae simplices og Asciidae 
compositae fra Nordhavs-Expeditionen.— 
Norske Nordhavs-Expeditionen 1876-78 23:1- 
16. 

Kott, P. 1969. Antarctic Ascidiacea. — Antarctic Re- 
search Series 13:1-239. 



32 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



. 1985. The Australian Ascidiacea. Part I, 

Phlebobranchiata and Stolidobranchiata.— 
Memoirs of the Queensland Museum 23: 1-440. 

Hartmeyer, R. 1912. Die Ascidien der deutschen 
Tiefsee-Expedition. — Deutschen Tiefsee-Expe- 
dition 7:223-392. 

Herdman, W. A. 1881. Preliminary report on the 
Tunicata of the Challenger expedition. — Pro- 
ceedings of the Royal Society of Edinburgh, part 
3 Cynthiadae 11:233-240. 

Millar, R. H. 1960. Ascidiacea.— Discovery Reports 
30:1-160. 

. 1988. Ascidians collected during the South- 
east Pacific Biological Oceanographic Program 
(SEPBOP). -Journal of Natural History 22:225- 
240. 

Monniot, C. 1969. Ascidies recoltees par la "Tha- 
lassa" sur la pente du plateau continental du 
golfe de Gascogne: (3-12 Aout 1967).— Bulletin 
du Museum National d'Histoire Naturelle, Paris 
(2),41(1): 155-1 86. 

. 1970. Ascidies recoltees par la "Thalassa" 

sur la pente du plateau continental du golfe de 
Gascogne (18-25 Octobre 1968). -Bulletin du 
Museum National d'Histoire Naturelle, Paris 
(2),41(5):1 131-1 145. 

. 1972. Dicopia antirrhinum n. sp. ascidie de 

la pente du plateau continental du golfe de Gas- 
cogne. Interpretation nouvelle de la famille des 
Octacnemidae. — Cahiers de Biologie Marine 13: 
9-20. 

. 1978. Ascidies phlebobranches et stolidob- 

ranches du sud de l'Ocean Indien.— Annales de 
FInstitut Oceanographique, Paris 54(2): 1 7 1-224. 

, & F. Monniot. 1973. Ascidies abyssales re- 
coltees au cours de la campagne oceanogra- 
phique Biacores par le "Jean Charcot".— Bul- 
letin du Museum National d'Histoire Naturelle, 
Paris (3), 121 Zoologie 93:289-475. 

, & . 1977a. Quelques ascidies abyssales 

du Sud-Ouest de l'Ocean Indien. — Comite Na- 
tional Francais de Recherches Antarctiques 42: 
305-327. 

, & . 1977b. Tuniciers benthiques pro- 

fonds du Nord-Est Atlantique. Resultats des 
campagnes Biogas. — Bulletin du Museum Na- 
tional d'Histoire Naturelle, Paris (3), 466 Zoo- 
logie 323:695-720. 

, & . 1978. Recent work on the deep- 
sea tunicates. Oceanography and marine biol- 
ogy, annual review, 16:181-228. 



-,& 



-. 1983. Ascidies antarctiques et sub- 



antarctiques. Morphologie et biogeographie.— 
Memoires du Museum National d'Histoire Na- 
turelle, Paris (A), Zoologie 125:1-168. 

Oka, A. 1913. Zur Kenntnis der zwei aberranten As- 
cidiengattungen Dicopia Sluit. und Hexacroby- 
lus Sluit.— Zoologischer Anzeiger 43(1): 1-10. 

. 1918. Megalodicopia hians n. g., n. sp., eine 

sehr merkwiirdige Ascidie aus dem japanischen 
Meere.— Annotationes Zoologicae Japonense 9: 
399-407. 

Ritter, W. E. 1907. The ascidians collected by the 
United States Fisheries Bureau Steamer Alba- 
tross on the coast of California during the sum- 
mer of 1904. University of California, Publi- 
cations in Zoology, Berkeley 4(1): 1-52. 

. 1913. The simple ascidians from the North- 
eastern Pacific in the collection of the United 
States National Museum. — Proceedings of the 
United States National Museum 45:427-505. 

, & R. A. Forsyth. 1917. Ascidians of the lit- 
toral zone of southern California. University of 
California Publications in Zoology 16:439-5 12. 

Sluiter, C. P. 1905. Die Tunicaten der Siboga-Ex- 
pedition. Supplement zu der I. Abteilung: die 
socialen und holosomen Ascidien.— Siboga Ex- 
pedition Monographie 56A: 129-1 39. 

Tokioka, T. 1953. Ascidians of Sagami Bay collected 
by His Majesty the Emperor of Japan.— Tokyo: 
Iwanami Shoten 1-315. 

. 1 972. On a small collection of ascidians from 

the Pacific coast of Costa Rica. Publications of 
the Seto Marine Biological Laboratory 19(6): 
383-404. 

Vinogradova, N. G. 1969. On the finding of a new 
aberrant ascidian in the ultrabyssal of the Ku- 
rile-Kamchatka trench.— Bulletin of the Mos- 
kovit Society of Natural History, Biological Part 
3:27-42. 

. 1975. On the discovery of two new species 

of an aberrant deep-water ascidiacean genus Sit- 
ula in the South-Sandwich trench.— Transac- 
tions of the P. P. Shirshov Institute of Oceano- 
logia 103:289-303. 

Museum national d'Histoire naturelle, 
Laboratoire de Biologie des Invertebres 
Marins et Malacologie, 55 rue Buffbn, 75005 
Paris, France. 



PROC. BIOL. SOC. WASH. 
102(1), 1989, pp. 33^4 

EUKROHNIA CALLIOPS, A NEW SPECIES OF 

CHAETOGNATHA FROM THE NORTHERN GULF OF 

MEXICO WITH NOTES ON RELATED SPECIES 

Jerry A. McLelland 

Abstract. —Eukrohnia calliops, a new species of Chaetognatha, was discov- 
ered in deep-water plankton collections made at four stations in the northern 
Gulf of Mexico in August 1987, and in February 1988. The new species appears 
to occur in bottom water over the upper continental slope at depths between 
200 and 700 m. Its large pigmented eyes easily distinguish it from other Eu- 
krohnia species occurring in the Gulf of Mexico. It differs from the closely 
related E. kitoui Kuroda, 1981, a species from Japanese waters, chiefly by the 
size and shape of the eyes. Thirteen specimens of E. calliops are studied and 
compared with specimens of E. kitoui, E. proboscidea, and E. fowleri, all of 
which possess pigmented eyes. The first record of E. proboscidea in the Gulf 
of Mexico is documented and affinities within the proposed "proboscidea group" 
are discussed. Distribution records of four species of Eukrohnia reported from 
the Gulf of Mexico prior to this study are presented with a key separating the 
six species now known. 



Members of the chaetognath genus Eu- 
krohnia typically inhabit deep-water plank- 
ton communities throughout the world (Al- 
varifio 1965, 1969 for zoogeography). Of 
the nine currently described species of Eu- 
krohnia (Casanova 1986), only three are 
known to possess pigmented eyes: E. fowleri 
Ritter-Zahony, 1909, E. proboscidea Fur- 
nestin & Ducret, 1965, and E. kitoui Ku- 
roda, 1981. Eukrohnia fowleri has a well 
known cosmopolitan distribution, while the 
latter two species appear restricted to deep 
coastal waters of the Atlantic Ocean and the 
Pacific coast of Japan, respectively. 

One hundred and eighty three specimens 
of an undescribed species of Eukrohnia with 
large pigmented eyes were discovered in 
plankton samples collected at four study 
areas in the northern Gulf of Mexico in Au- 
gust 1987 and February 1988 by personnel 
of the Gulf Coast Research Laboratory 
aboard the R/V Tommy Munro (Fig. 1, Ta- 
ble 1). Quantitative samples were obtained 
by towing horizontally at depths of 200-500 



meters using a Niskin double-trip system 
with 1 m diameter, 333 fxm. mesh nets. 
Qualitative samples were obtained near the 
bottom with 0.3 m diameter, 183 jum mesh 
nets attached to crab traps and allowed to 
passively filter the bottom current. Only 
material from the February 1988 cruise was 
used in this work because specimens in the 
August 1987 samples were in poor condi- 
tion. 



Class Sagittoidea Claus and Grobben, 1905 

Order Phragmophora Tokioka, 1965 

Family Eukrohniidae Tokioka, 1965 

Genus Eukrohnia Ritter-Zahony, 1 909 

Diagnosis. —One pair of lateral fins 
emerging at or near ventral ganglion; one 
pair of tooth rows; apical gland cell com- 
plex, or papillae tuft, at anteroventral point 
of head; ventral transverse musculature in 
trunk segment extending from neck to at 
least posterior edge of ventral ganglion. 



34 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




24 l 



23' 



2000 m 



GULF OF MEXICO 



**,, ^:' 



1000' 





CARIBBEAN SEA 



■i i i '1 



24^ 



1 I 



93° 92° 91° 90° 89° 88° 87° 86° 85° 84° 83° 82° 81° 80° 79 c 



Fig. 1 . Map showing locations of five study areas along the continental slope in the northern Gulf of Mexico 
from which deep-water plankton collections were made in August 1987. and February' 1988. 



Eukrohnia calliops, new species 
Figs. 2, 3, Tables 1-4 

Material— Holotype, 21.5 mm (USNM 
119293), R/V Tommy Munro, Area 3, 
(29°21'N, 86°57'W) northern Gulf of Mex- 
ico, 200-500 m, 16 Feb 1988, 1910 CDT, 
11 paratypes as follows: 1 specimen, 18.2 
mm (USNM 1 19296), from type locality; 2 
specimens 18.4, 15.0 mm (USNM 119294), 
from Area 1 (29°00'N, 88°26'W), 677 m, 28 
Feb 1988, 1930 CDT; 5 specimens, 14.4, 
12.6, 11.7, 10.4, 8.4 mm (USNM 119295), 
from Area 2 (29°18'N, 87°32'W), 200-500 
m, 13 Feb 1988, 1035 CDT; 1 specimen, 
16.6 mm (GCRL 1131), from Area 4 
(27°50'N, 85°24'W), 400-485 m, 25 Feb 



1988, 2345 CDT; 3 specimens, 16.2, 13.0, 
11.3 mm (GCRL 1132), from Area 4 
(27°50'N, 85°24'W), 677 m, 26 Feb 1988, 
1910 CDT. 

Description. —Based on type material with 
five stages of sexual maturity described in 
Table 2. Body firm, translucent, with well- 
developed dorsal and ventral longitudinal 
muscle bands; widest at trunk-caudal sep- 
tum, tapering slightly toward the head; 
maximum length recorded 21.5 mm. Tail 
length 21.7-29.3% of total body length. 
Head wider than long with distinct constric- 
tion at neck. Corona ciliata not observed. 
Collarette (alveolar tissue) absent. 

Eyes large, bulbous, sub-oval to tear-drop 
shaped (Figs. 3C, 5B); widely separated on 



VOLUME 102, NUMBER 1 



35 



Table 1.— Station and hydrographic data for Eukrohnia species in the northern Gulf of Mexico. 





Position 


Date 


Time 


Bottom 

depth 

(m) 


Sampling 

depth 

(m) 


Length 
of tow 
(min) 


Bottom 

salinity 

(PPt) 


Bottom 

temp. 

(°C) 


Number o: 
specimens 


r 


Station 


Ec. 


Ef. 


Ep. 


Area 1 


29°02'N, 088°27'W 


08/02/87 


2007 


494 


200-400* 


33 


36.0 


8.4 


3 










29°00'N, 088°26'W 


02/28/88 


1930 


677 


677** 


25 hr 


36.0 


6.5 


2 








Area 2 


29°18'N, 087°34'W 


08/04/87 


1910 


494 


200-400* 


44 


38.0 


8.6 


131 










29°18'N, 087°32'W 


02/13/88 


1035 


677 


200-500 


30 


34.0 


6.3 


5 










29°18'N, 087°32'W 


02/14/88 


0730 


677 


677** 


23 hr 


34.0 


6.3 


5 








Area 3 


29°21'N, 086°57'W 


08/06/87 


0838 


677 


200-500 


54 


39.0 


6.4 


13 










29°21'N, 086°57'W 


02/16/88 


1910 


677 


200-500 


94 


36.0 


6.7 


2 


3 







29°21'N, 086°57'W 


02/17/88 


1840 


677 


677** 


25 hr 


36.0 


6.7 


7 








Area 4 


27°54'N, 085°16'W 


08/08/87 


1150 


494 


200-400 


31 


38.0 


7.7 


1 










27°50'N, 085°24'W 


02/25/88 


2345 


494 


400-485 


27 


38.0 


8.4 


1 










27°50'N, 085°24'W 


02/26/88 


1910 


677 


677** 


24 hr 


38.0 


6.5 


3 








Area 5 


27°00'N, 084°57'W 


02/24/88 


0720 


677 


677** 


24 hr 


38.0 


6.6 





4 


1 



* Net accidentally dragged bottom. 

** Passive sampling by 0.3 m diameter nets attached to crab traps. 
Ec. = E. calliops, Ef. = E. fowled, Ep. = E. proboscidea. 



dorsal surface of the head. Pigment cups 
lunate to hemispherical, dark-brown to black 
in color; oriented parallel to longitudinal 
body axis with apex of cup on inner median 
edge of eyes. 

Apical gland cell complex bilobate, pro- 
truding anteriorly. Two less conspicuous 
gland cell complexes, a smaller anterior one 
and a larger posterior one, both with small, 
densely packed papillae, situated anterior to 
mouth between vestibular ridges (Fig. 3A). 

Hooks 12-13 on each side of head; long, 
slender, and somewhat straight, with sharp- 
ly bent tips (Fig. 3D, E); no denticulate hooks 
observed. Anterior teeth absent. Posterior 
teeth 9-21 (17-21 in stage III and IV in- 
dividuals); thin, pointed, set close together 
(Fig. 3B); located midway between hooks 
and apical gland cell complex. Up to 20 
vestibular palps in one or two rows on ves- 
tibular ridge, beginning ahead of and run- 
ning parallel to entire length of tooth row; 
those adjacent to teeth elongate, highly 
transparent with cup-like terminal ends (Fig. 
3B). 

Gut extending from mouth to anus, flat- 
tened laterally into thin ribbon, appears im- 



pregnated with fine darkened granular ma- 
terial; usually with distinctly thickened area 
directly beneath ventral ganglion (Fig. 2B). 
Intestinal diverticulae absent. 

Ventral ganglion located at anterior fourth 
of body on ventral surface of trunk; small, 
rectangular to oval in shape, with fine nerve 
fibers emerging at corners. Ventral trans- 
verse musculature extending from neck re- 
gion past posterior edge of ventral ganglion 
by a distance of up to one-fourth the length 
of ganglion (Fig. 2B). 

Lateral fins extending along body from 
posterior third of tail segment to about an- 
terior third of ventral ganglion; broadly 
rounded at posterior end and tapering an- 
teriorly to delicate ridge; posterior portions 
entirely rayed to about level of trunk-caudal 
septum, rays continuing anteriorly along 
distal edge for about half fin length; inner 
and anterior portions containing laminar 
material and fine fibrils. Tail fin distinc- 
tively large, broadly triangular, completely 
rayed. 

Ovaries short, club shaped, 10-17 ova per 
ovary; ova arranged in two rows when 
viewed laterally (Fig. 2C). Seminal vesicles 



36 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




B 




Fig. 2. Eukrohnia calliops from the northern Gulf 
of Mexico. A, ventral view of a mature specimen; B, 
lateral view of ventral ganglion area with cut-away 
showing thickened area of gut tube underlying trans- 
verse musculature; C, lateral view of ovary at maturity 
stage III; D, detail of mature seminal vesicle. 



oval to elongate, about a third their length 
overlapped by posterior edges of lateral fins; 
separated from tail fin by about a fourth 
their length (Fig. 2D). 

Etymology. —The specific name is de- 
rived from the Greek words "kallos" (beau- 
ty) and "ops" (pertaining to the eyes), and 
is a noun in apposition. 

Comparative material examined (Table 
3).- 

Eukrohnia kitoui, 10 specimens, A.1-25A 
mm; Tokyo Bay, Japan (35°08'N, 
139°46'E), 13 Feb 1981, 0-240 m; from 
Kazunori Kuroda, Tokai Regional Fish- 
eries Research Laboratory, Tokyo, Japan. 

Eukrohnia proboscidea, 2 specimens, one 
18.0 mm; origin unknown; from Jean- 
Paul Casanova, Laboratorie de Biologie 
animal (Plancton), Marseille, France. 

Eukrohnia proboscidea, 1 specimen, 10.6 
mm; Gulf of Mexico (27°00'N, 84°57'W), 
14 Feb 1988, 677 m; from Area 5, R/V 
Tommy Munro (Table 1). 

Eukrohnia fowleri, 7 specimens co-occur- 
ring with E. calliops and E. proboscidea 
in the northern Gulf of Mexico (Table 1). 

Results and Discussion 

Affinities. — Morphological characteris- 
tics of the four species of Eukrohnia pos- 
sessing eye pigmentation are compared in 
Table 4 which combines personal obser- 
vations with published data. 

Eukrohnia calliops is most closely related 
to E. kitoui (Figs. 4B; 5C, D). The two species 
are very similar in all respects including eye 
pigment configuration. In fact, they were at 
first considered synonymous until a detailed 
comparison was made between non-con- 
tracted E. calliops specimens collected in 
February 1988 and material of similar con- 
dition from Tokyo Bay, Japan. Results of 
this comparison appear in Table 3 and Fig. 
6. Eukrohnia calliops has greater numbers 
of posterior teeth and higher tail segment 
percentages in specimens of comparable size. 
Obvious differences between the two species 
are eye size and the appearance of the pig- 



VOLUME 102, NUMBER 1 



37 




Fig. 3. Head details of Eukrohnia calliops, holotype: A, ventral view of head; B, vestibular ridge showing 
rows of palps and teeth; C, dorsal view of eyes; D, mature hook; E, hook tip variations. (Abbreviations for Fig. 
3A, B: gc = gland cell complex, m = mouth, pt = posterior teeth, vp = vestibular palps.) 



ment cup within the eye. The pigment cups 
of E. calliops are darker in color, larger in 
relation to the total area of the eye, and have 
a greater three-dimensional aspect causing 
them to "bulge" from the surface of the head. 
The larger eye size of E. calliops is dem- 
onstrated in Fig. 6 which compares ratios 
of eye length to total body length (EL/TL 
x 100) between the two species. Eukrohnia 
calliops was found to differ from E. kitoui 
at the 95% level of significance using a t-test 
on the mean ratios with variances pooled. 
Eukrohnia proboscidea (Figs. 4C; 5E, F), 
originally described from tropical African 



waters by Furnestin & Ducret (1965), is ap- 
parently a close relative of E. calliops and 
E. kitoui as demonstrated by their morpho- 
logical similarities (Table 4) and similar 
habitats. As such, the three might be loosely 
separated from the rest of the genus to form 
the "proboscidea group." This grouping 
would further subdivide the "fowleri group" 
proposed by Casanova (1986:827), an ar- 
tificial arrangement of five species of Eu- 
krohnia based on the presence of similar 
light sensitive cells in the eyes, pigmenta- 
tion in the digestive tube, and numbers and 
coloration of hooks. 



38 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 2.— Maturity stages for Eukrohnia calliops in February 1988 collections from the Gulf of Mexico. 



Stage 



TL (mm) 



Male gonads 



Female gonads 



I 8.4-12.6 Testes are small rods originating in anterio- 
lateral corners of tail segment and not ex- 
tending over half the length of the seg- 
ment; spermatids not enlarged; seminal 
vesicles not formed. 
II 1 1.7-14.4 Testes elongated to full length of caudal seg- 
ment; spermatids enlarging; seminal vesi- 
cles emerging but not enlarged. 

III > 15.0 Testes completely fill tail segment; sperm 

cells enlarged; seminal vesicles enlarged, 
fully formed. 

IV > 15.0 Tail segment partially empty; seminal vesi- 

cles ruptured. 

V > 18.0 Tail segment empty (spent). 



Ovaries short, club-shaped; no ova 
formed but oocyte clusters present. 



Ovaries longer (4% TL); ova small, 
uniform in size, arranged in 2 rows 
or clustered. 

Ovaries longer and thicker (5-7% TL); 
some ova enlarged (3-5 per ovary), 
arranged in 2 rows. 

Ovaries at maximum length (10-11% 
TL); all ova enlarged (10-14 per 
ovary), arranged in 2 rows. 

Ovaries contracted, empty (spent) or 
with 1 or 2 large ova; oviducts swol- 
len. 



The pigment of E. proboscidea, which is 
situated in the posterior portion of the eye, 
differs considerably from that of E. calliops 
and E. kitoui. It is light-brown in color with 
a distinct black edge along the inner border 
which lends a "U"-shaped appearance when 
viewed dorsally. 

An interesting affinity within the "pro- 
boscidea group" is the posterior extent from 
the neck of the ventral transverse muscu- 
lature. Casanova (1986:831) found that for 
E. proboscidea the musculature exceeded the 
position of the ventral ganglion by half the 
length of the ganglion. He considered this 
to be a unique feature of that species and 
stated that in all other members of the genus 
Eukrohnia, the musculature extends only to 
the edge of the ganglion. The overreaching 
of the musculature by a fourth the length of 
the ganglion in E. calliops, while not as great 
an extent as that of E. proboscidea, does 
demonstrate a similarity. In the largest spec- 
imen of the E. kitoui material (25.4 mm), 
the extent of transverse musculature was 
similar to that of E. calliops; however, this 
characteristic was reported by Kuroda (1981: 
181) as "extending from the neck to the 
posterior edge of the ventral ganglion." 



The hooks of the "proboscidea group" 
species are nearly identical. The hook tips 
(Fig. 3E), abruptly deflected inward at 45- 
90°, are similar to those of E. hamata and 
E. bathypelagica. Denticulate hooks, a com- 
mon characteristic in juveniles of Eukroh- 
nia and Heterokrohnia (Kapp & Hagen 
1985:58), were not found on small speci- 
mens of the "proboscidea group" in this 
study nor were any reported by Kuroda 
(1981) for E. kitoui or by Furnestin (1965) 
for E. proboscidea. Owre (1973:957), how- 
ever, reported that the ventral-most hooks 
on 13.0 and 16.5 mm specimens of E. pro- 
boscidea from the Caribbean Sea were "ser- 
rated with 10-18 small hooks." 

The single specimen of E. proboscidea ob- 
tained from the bottom sample at Area 5 
represents a new record for the Gulf of Mex- 
ico. At 10.6 mm, it is the smallest specimen 
yet recorded for the species. Meristic counts 
were similar to those reported by Owre 
(1973) for a 13.0 mm specimen from the 
Caribbean Sea but the Gulf specimen had 
fewer number of hooks and teeth than the 
smallest specimen (13.3 mm) of the type 
material from tropical African waters (Du- 
cret 1965). The Gulf of Mexico specimen 



VOLUME 102, NUMBER 1 



39 



Table 3.— Meristic values of Eukrohnia comparison material. 



TL mm 


% Tail 


Hooks 


Teeth 


Eye L mm 


Mat. St. 




Source 


Eukrohnia 


calliops, n. sp. 


from the Gulf of Mexico 










21.5 


24.9 


12 


21 


0.26 


IV-V 


A3, 


200-500 m 


18.4 


27.0 


12 


17 


0.28 


III 


Al, 


bottom 


18.2 


25.6 


12 


15-17 


0.24 


V 


A3, 


200-500 m 


16.6 


26.6 


12 


17-19 


0.22 


III 


A4, 


400-440 m 


16.2 


24.8 


12-13 


16 


0.24 


IV 


A4, 


bottom 


15.0 


29.3 


NA 


NA 


0.27 


IV 


Al, 


bottom 


14.4 


23.3 


13 


12 


0.24 


II 


A2, 


200-500 m 


13.0 


26.4 


12 


13 


0.20 


II 


A4, 


bottom 


12.6 


21.7 


12 


12-13 


0.20 


I 


A2, 


200-500 m 


11.7 


25.3 


12 


7-11 


0.19 


II 


A2, 


200-500 m 


11.3 


31.0 


13 


15 


NA 


II 


A4, 


bottom 


10.4 


22.3 


11-12 


10 


0.18 


I 


A2, 


200-500 m 


8.4 


26.7 


11 


9-10 


0.14 


I 


A2, 


200-500 m 


Eukrohnia 


kitoui from Tokyo Bay, Japan (35°08'N, 


139°46'E), 0-240 


m 






25.4 


24.6 


13 


16-17 


0.20 


IV 


Kazunori Kuroda 


18.4 


22.2 


12 


14 


0.17 


III 


Kazunori Kuroda 


16.6 


23.6 


10-11 


14 


0.14 


III 


Kazunori Kuroda 


15.0 


22.5 


11 


13 


0.14 


II 


Kazunori Kuroda 


13.4 


22.8 


11 


9-10 


0.12 


II 


Kazunori Kuroda 


10.4 


25.4 


10 


7 


0.10 


II 


Kazunori Kuroda 


7.9 


23.2 


10 


5 


0.08 


I 


Kazunori Kuroda 


7.0 


25.3 


9 


4 


0.07 


I 


Kazunori Kuroda 


5.0 


26.2 


8 


3 


0.06 


I 


Kazunori Kuroda 


4.7 


27.4 


7 


1 


0.07 


I 


Kazunori Kuroda 


Eukrohnia 


proboscidea 














18.0 


23.0 


12 


13 




II 


J. P 


. Casanova 


NA 


NA 


12 


15 




II 


J. P 


. Casanova 


10.6 


24.2 


10 


8 




I 


A5, 


Gulf of Mex. 


Eukrohnia 


fowleri from the Gulf of Mexico 












28.5 


23.0 


13 


27 




II 


A5, 


bottom 


28.5 


21.3 


11-13 


27 




II 


A5, 


bottom 


26.6 


22.0 


11 


20 




II 


A3, 


200-500 m 


14.4 


23.3 


11 


12 




I 


A3, 


200-500 m 


14.0 


24.0 


1 1 (2)* 


12 




I 


A3, 


200-500 m 


11.8 


20.0 


12(4)* 


10 




I 


A5, 


bottom 


10.7 


20.9 


12(4)* 


12 




I 


A5, 


bottom 



* Denticulate hooks (number). 



lacked denticulations on the ventral-most 
hooks. 

Eukrohnia fowleri (Figs. 4D; 5G, H), a 
cosmopolitan, mesoplanktonic species, dif- 
fers from the "proboscidea" group in nearly- 
all morphological features and armature 
meristics. The eye pigment is a dark, vari- 
ably shaped fleck in the center of the eye. 
The hooks are robust, red in color, and have 



straight tips. The ventral-most three or four 
hooks of small specimens are strongly den- 
ticulate (Table 3). 

Ecology and distribution. -Based on avail- 
able data, the three species of the "probos- 
cidea group" appear to occupy similar hab- 
itats in deep-water areas adjacent to coastal 
regions, although E. proboscidea has so far 
been found at greater depths, with occur- 



40 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




B 



« 





Fig. 4. Dorsal views of heads of the four species of Eukrohnia possessing pigmented eyes. A, E. calliops, 
holotype; B, E. kitoui Kuroda, 1981; C, E. proboscidea Furnestin & Ducret, 1965; D, E. fowled Ritter-Zahony, 
1909. Scales: A, B, and D = 0.5 mm; C = 0.2 mm. 



rences below 1000 m, off the west African 
coast (Ducret 1965) and in samples collect- 
ed between 750 and 1 100 m in the Carib- 
bean Sea (Owre 1973, Michel et al. 1976). 
Eukrohnia kitoui and E. calliops were col- 
lected at similar depths (0-400 and 200- 
677 m) in Tokyo Bay and the northern Gulf 
of Mexico, respectively, near undersea can- 
yon areas where upwelling of water from 
deeper regions occurs. 

It is worthy to note that at the Area 5 
station (Table 1), no specimens of E. cal- 
liops were present, but in the February bot- 
tom net samples, one immature specimen 
of E. proboscidea was found, indicating the 



northward extension of a deep-dwelling Ca- 
ribbean population of that species. More ex- 
tensive sampling near the bottom in these 
areas may identify boundaries between these 
two species in the Gulf of Mexico. 

In the August 1987 cruise, the nets acci- 
dentally dragged bottom at the Area 2 sta- 
tion near the Desoto Canyon, resulting in a 
sample containing a large number (131) of 
E. calliops (Table 1). This suggests a pos- 
sible hypo-planktonic distribution for the 
species in the Gulf of Mexico. 

The genus Eukrohnia was not mentioned 
in any of the major studies of the chaeto- 
gnath fauna of coastal areas of the Gulf of 



VOLUME 102, NUMBER 1 



41 



Table 4.— Characteristics of the four species of Eukrohnia with pigmented eyes from descriptions published 
by Ducret ( 1 965), Alvarino ( 1 967), Kuroda (1981), Michel ( 1 984), and from personal observations by the author. 





E. fowler i 


E. proboscidea Furnestin & 






Character 


Ritter-Zahony, 1909 


Ducret, 1965 


E. kitoui Kuroda, 1981 


E. calliops, n. sp. 


Total length 


40 


25 


24.5 


21.5 


(mm) 










% Tail length 


22-25 


21.5-31 


22-24 


24-29 


Eye shape 


Elliptical 


Elongate 


Circular to sub- 
oval 


Oval to tear-drop 
shaped 


Eye pigment 


Variably shaped 


Elongate to 


Semi-circular in 


Large, lunate, in cen- 




rectangular fleck 


U-shaped in pos- 


center of eye, 


ter of eye, dark 




in center of eye, 


terior part of eye, 


dark brown to 


brown to black 




black 


light-brown 


black 




Apical gland 


Single mass of cells, 


Bilobate, protrud- 


Bilobate, protrud- 


Bilobate, protruding 




not protruding 


ing anteriorly 


ing anteriorly 


anteriorly 


Hooks 


11-13, robust, 


10-13, slender, 


8-13, slightly 


11-13, slightly 




strongly curved, 


gently curved, 


curved, light- 


curved, light- 




reddish, tips 


light-brown, 


brown, transpar- 


brown, transpar- 




straight 


transparent, tips 


ent, tips curved 


ent, tips curved 






curved sharply 


sharply inward 


sharply inward 






inward 






Posterior teeth 


Up to 30 


Up to 25 


Up to 17 


Up to 21 


Transverse 


Level with poste- 


Past ventral gan- 


Usually level with 


Past ventral ganglion 


musculature 


rior edge of ven- 


gion by up to x h 


posterior edge of 


by up to x k length 


extent 


tral ganglion 


length of gan- 
glion 


ventral ganglion 


of ganglion 


Epidermal col- 


Thick band around 


Absent 


Absent 


Absent 


larette tissue 


body at ventral 
ganglion 








Lateral fins 


Extending from an- 


Extending from 


Extending from 


Extending from ante- 




terior Vi of ven- 


base of ventral 


slightly anterior 


rior third of ven- 




tral ganglion to 


ganglion to mid- 


to ventral gan- 


tral ganglion to 




middle of tail 


dle of tail seg- 


glion to posterior 


posterior third of 




segment 


ment 


third of tail seg- 
ment 


tail segment 


Mature ova- 


Short (13% TL); 


Short (15% TL); 


Short (13.5% TL); 


Short (7.5-10.5% 


ries 


ova large, in two 


ova in 2 rows, 


ova in 2 rows, 


TL); ova in two 




rows 


about 14 per 
ovary 


10-15 per ovary 


rows, 10-14 per 
ovary 


Seminal vesi- 


Ovoid; adjacent to 


Ovoid, elongate; 


Ovoid; adjacent to 


Ovoid, elongate; ad- 


cles 


posterior fins and 


adjacent to inser- 


insertion of later- 


jacent to insertion 




separated from 


tion of lateral fins 


al fins with ante- 


of lateral fins with 




tail fin a distance 


with anterior l h 


rior x h over- 


anterior l h over- 




greater than their 


overlapped by 


lapped by 


lapped by posterior 




length 


posterior edge of 


posterior edge of 


edge of fins, sepa- 






fins, separated 


fins, separated 


rated from tail fin 






from tail fin by 


from tail fin by 


by about Vi their 






about V2 their 


about V2 their 


length 






length 


length 





42 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





F 




H 



* 



o 



Fig. 5. Eyes of the four species of Eukrohnia possessing pigmented eyes. A, C, E, and G show relative 
distances between eyes and B, D, F, and H show detail of individual eyes. A and B, E. calliops, holotype; C 
and D, E. kitoui Kuroda, 1981; E and F, E. proboscidea Furnestin & Ducret, 1964; G and H, E. fowled Ritter- 
Zahony, 1909. All scales = 0.1 mm. 



VOLUME 102, NUMBER 1 



43 



r = 0.989 

V = 139.35 x -3.645 



20 - 


E 

E 












■ 




/ 


15 - 


I 
i- 
O 

z 

> 
o 
o 
m 






/ a 

/ ■ 


■ 


■ 
■ 


/ 


■ 


■ 


10 - 




a/ 
o / 


s 


/ ■ 










■ £ calhops 


b - 




o/a 














' = 0.865 


. 


















y = 79.23 x -2.931 


1 








EYE LENGTH mm. 
















1 1 


1 


1 i 










1 



Fig. 6. Comparative relationships of body length 
to eye length for Eukrohnia kitoui and E. calliops. 



Mexico (Pierce 1951, 1962; Suarez-Caabro 
1955; Vega-Rodriguez 1965; McLelland 
1984) because sufficient depths were either 
not present or not sampled. Deep-water 
studies from areas further removed from 
the continental slopes that did include four 
species of Eukrohnia are presented in Table 
5. Based on personal observations and on 
characteristics of mature specimens report- 
ed in the literature, the following key sep- 
arating the species of Eukrohnia occurring 
in the Gulf of Mexico is presented. 



Key to the Species of Eukrohnia of the 
Gulf of Mexico 

la. Eyes with pigment 2 

lb. Eyes non-pigmented 4 

2a. Apical gland bilobate and protrud- 
ing, causing head to appear point- 
ed. Hook tips bent inward at 45 to 
90 degree angles. Transverse mus- 
culature extending past posterior 
edge of ventral ganglion 3 

2b. Apical gland not prominent, a sin- 
gle lobed mass. Hook tips straight, 
transverse musculature even with 
posterior edge of ganglion E. fowled 

3a. Eye pigment small, elongate or 
"U"-shaped, in posterior region of 
eye E. proboscidea 

3b. Eye pigment large, lunate, encom- 
passing most of the median por- 
tion of the eye . . . . E. calliops, n. sp. 

4a. Number of hooks greater than 1 1 , 
hook tips straight E. bathyantarctica 

4b. Number of hooks less than 1 1 , 

hook tips bent inward 5 

5a. Hooks stout, nearly straight; tail 
less than 25% of body length . . . 
E. hamata 

5b. Hooks long, gently curved; tail 
greater than 25% of body length 
E. bathypelagica 



Table 5.— Distribution records of the genus Eukrohnia in the Gulf of Mexico. 



Species 


Area 


Position 


Sampling 

depth 

(m) 


Abundance 


Source 


E. hamata 


Central Gulf 


27°15'N, 029°09'W 


495-749 


Common 


Pierce (1954) 


Mobius, 1875 












E. fowleri Ritter- 


Central Gulf 


27°21'N, 088°42'W 


1100 


Common 


Every (1968) 


Zahony, 1909 


Central Gulf 


24°39'N, 093°09'W 


2000 


1 specimen 


Fagetti(1968) 




SW Gulf 


21°55'N, 095°25'W 


509-1000 


1 specimen 


Fagetti(1968) 


E. bathyantarctica 


Central Gulf 


24°39'N, 093°09'W 


2000 


2 specimens 


Fagetti(1968) 


David, 1958 


Central Gulf 


27°08'N, 089°55'W 


1032-2000 


1 specimen 


Fagetti(1968) 




Central Gulf 


26°16'N, 087°52'W 


2002-2400 


2 specimens 


Fagetti(1968) 




Central Gulf 


23°41'N, 090°08'W 


1971-2800 


1 specimen 


Fagetti(1968) 




SW Gulf 


21°55'N, 095°25'W 


509-1900 


2 specimens 


Fagetti(1968) 




Central Gulf 


25°59'N, 086°11'W 


580-2000 


15 specimens 


Owre(1973) 


E. bathypelagica 


Central Gulf 


25°59'N, 086°11'W 


580 


8 specimens 


Owre(1973) 


Alvarino, 1962 













44 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Acknowledgments 

This work was developed with materials 
collected under Grant No. NA87WC-H- 
06142, Marine Fisheries Initiative (MAR- 
FIN), NMFS, NOAA, U.S. Department of 
Commerce. I wish to thank Harriet Perry 
for providing the MARFIN plankton col- 
lections, Dr. Richard Heard and Sara Le- 
croy for their review and suggestions on the 
manuscript, and Dr. Robin Overstreet, Mary 
Ann Pavlov, and Debbie Jackson for their 
help in producing the photographs. All of 
the above are from the Gulf Coast Research 
Laboratory. Dr. Kazunori Kuroda of the 
Tokai Regional Fisheries Laboratory (To- 
kyo, Japan) and Dr. Jean-Paul Casanova of 
the Laboratorie de Biologie animal-planc- 
ton (Marseille, France) kindly provided me 
with some of the specimens of Eukrohnia 
used in this study. 

Literature Cited 

Alvarino, A. 1965. Chaetognaths. Pp. 115-194 in 
Harold Barnes, ed., Oceanography and marine 
biology, annual revue, Vol. 3. George Allen & 
Unwin Ltd., London. 

. 1967. The Chaetognatha of the NAGA ex- 
pedition (1959-1961) in the South China Sea 
and the Gulf of Thailand. Part 1— systemat- 
ics.— NAGA Reports, University of California, 
Scripps Institute of Oceanography 4(2): 1-1 97. 

. 1969. Los quetognatos del Atlantico. Distri- 

bucion y notas esenciales de sistematica. Tra- 
bajos del Instituto Espafiol de Oceanografia 37, 
290 pp. 

Casanova, J. P. 1986. Deux nouvelles especes & Eu- 
krohnia (Chaetognathes) de FAtlantique sud- 
tropical africain.— Bulletin du Museum Natio- 
nal d'Histoire Naturelle, Paris (4)8(A no. 4): 8 1 9- 
833. 

Ducret, F. 1965. Les especes du genre Eukrohnia 
dans les eaux equatoriales et tropicales afri- 
caines. — Cahiers O.R.S.T.O.M.-Oceanogra- 
phie 3(2):63-78. 

Every, M. G. 1968. The taxonomy and areal distri- 
bution of the Chaetognatha in the oceanic Gulf 
of Mexico. M.S. Thesis, Texas A&M Univer- 
sity, College Station, 67 pp. 

Fagetti, G. E. 1968. New record of Eukrohnia bathy- 



antarctica David, 1958, from the Gulf of Mex- 
ico and Caribbean Sea.— Bulletin of Marine Sci- 
ence 18(2):383-387. 

Furnestin, M. L. 1965. Variations morphologiques 
des crochets au cours du developpement dans 
le genre Eukrohnia.— Revue des Travaux. In- 
stitut des Peches Maritimes 29(3):275-284. 

, & F. Ducret. 1965. Eukrohnia proboscidea, 

nouvelle espece de Chaetognathe. — Revue des 
Travaux. Institut des Peches Maritimes 29(3): 
271-273. 

Kapp, H., & W. Hagen. 1985. Two new species of 
Heterokrohnia (Chaetognatha) from Antarctic 
waters.— Polar Biology 4:53-59. 

Kuroda, K. 1981. A new chaetognath, Eukrohnia 
kitoui n. sp., from the entrance to Tokyo Bay.— 
Publications of the Seto Marine Biological Lab- 
oratory 26 (1/3): 177-1 85. 

McLelland, J. A. 1 984. Observations on chaetognath 
distributions in the northeastern Gulf of Mexico 
during the summer of 1974.— Northeast Gulf 
Science 7(l):49-59. 

Michel, H. B. 1984. Chaetognatha of the Caribbean 
Sea and adjacent areas. NOAA Technical Re- 
port NMFS 15, United States Department of 
Commerce, National Technical Information 
Service, Springfield, Virginia, 33 pp. 

, M. Foyo, & D. A. Haagensen. 1976. Carib- 
bean Zooplankton. Office of Naval Research, 
Department of the Navy, U.S. Government 
Printing Office, Washington, D.C., 712 pp. 

Owre, H. B. 1973. A new chaetognath genus and 
species, with remarks on the taxonomy and dis- 
tribution of others.— Bulletin of Marine Science 
23(4):948-963. 

Pierce, E. L. 1951. The Chaetognatha of the west 
coast of Florida.— Biological Bulletin (Woods 
Hole) 100:206-228. 

. 1954. Notes on Chaetognatha of the Gulf of 

Mexico.— Fishery Bulletin of the U.S. Fish & 
Wildlife Service 55:327-329. 

. 1962. Chaetognatha from the Texas coast.— 

Publications of the Institute of Marine Science, 
University of Texas 8:147-152. 

Suarez-Caabro, J. A. 1955. Quetognatos de los mares 
Cubanos. — Memorias de la Sociedad Cubana de 
Historia Natural 22:125-180. 

Vega-Rodriguez, F. 1965. Distribution de Chaetog- 
natha en Veracruz, Ver.— Anales del Instituto 
de Biologia, Universidad Nacional Autonoma 
de Mexico 36(l/2):229-247. 

Gulf Coast Research Laboratory, P.O. Box 
7000, Ocean Springs, Mississippi 39564. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 45-49 

SMALLEYUS TRICRISTATUS, NEW GENUS, 

NEW SPECIES, AND PSEUDOTHELPHUSA PARABELLIANA, 

NEW SPECIES (BRACHYURA: PSEUDOTHELPHUSIDAE) 

FROM LOS TUXTLAS, VERACRUZ, MEXICO. 

Fernando Alvarez 

Abstract.— Owq new genus and species, Smalleyus tricristatus, and a new 
species of Pseudothelphusa, P. parabelliana, are described from the Los Tuxtlas 
region in the State of Veracruz, Mexico. The new species described here belong 
to the Tribe Pseudothelphusini. The distribution patterns of both genera are 
discussed. 



The Los Tuxtlas region of Veracruz lies 
adjacent to the coast of the Gulf of Mexico, 
and comprises the area between 1 8°10'N to 
18°40'N and 94°45'W to 95°25'W. This re- 
gion encompasses a small mountain range 
of volcanic origin, where the highest vol- 
canic cone is more than 1700 m high. In 
this area four genera and six species of pseu- 
dothelphusid crabs co-occur: Odontothel- 
phusa maxillipes (Rathbun, 1898), Te- 
huana veracruzana (Rodriguez & Smalley, 
1 969), T. poglayenorum (Pretzmann, 1 980), 
T. diabolis (Pretzmann, 1980), Smalleyus 
tricristatus, and Pseudothelphusa parabel- 
liana (described below). Topographic het- 
erogeneity, the presence of tropical rainfor- 
est, and the fact that this is an isolated 
mountain range situated on the coastal plain, 
may account for much of the observed di- 
versity. Future collections in southern Ve- 
racruz and Tabasco will confirm if, in fact, 
pseudothelphusid crabs are absent from 
these coastal lowlands. This suggests that 
Los Tuxtlas may have served as a refuge in 
the past and that its brachyuran fauna has 
been preserved. The sites where collections 
were made are at altitudes that range be- 
tween 200 and 1000 m. All the organisms 
are deposited in the Carcinologic Collec- 
tion, Instituto de Biologia, Universidad Na- 
cional Autonoma de Mexico (IBUNAM). 



Smalleyus, new genus 

Diagnosis.— In cephalic view, proximal 
half of gonopod thinner than distal half. In 
mesial view, middle portion, along longi- 
tudinal axis, narrower than rest. In caudal 
view, lateral process also exhibiting con- 
striction in middle portion. In lateral view, 
proximal half straight with constant thick- 
ness; distal half slightly curved caudally. 

Apex of gonopod bearing 3 lobes: mesial, 
cephalolateral, and lateral. Mesial lobe with 
basal % of crest disposed at 90° angle to 
longitudinal axis of gonopod; distal portion 
sloping proximally; small field of spines 
along mesial surface of crest; apical margin 
of crest serrate, spine size increasing distal- 
ly. In cephalic view, mesial process with 
cephalic serrate projection, lying oblique to 
longitudinal axis of gonopod. Cephalolater- 
al lobe with median ridge in apical cavity, 
cephalic blade-like projection oriented 
cephalolaterally. Apical median ridge 
rounded, bearing spines. Cephalolateral 
projection with 3 strong teeth. Lateral lobe 
forming lateral wall of apical cavity. Field 
of spines along lateral crest and lateral sur- 
face; spine size increasing laterally and ce- 
phalically. Field of terminal pore setae sit- 
uated between median ridge and lateral crest. 
Sperm channel opening situated caudal to 
median ridge. 



46 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Type species. —Smalleyus tricristatus. 
Etymology.— This genus is named after 
Dr. Alfred E. Smaliey. Gender is masculine. 

Smalleyus tricristatus, new species 
Fig. 1 

Material examined. — Camino Izquierdo, 
Sierra de Santa Marta, Veracruz (18°26'N, 
94°57'W), Jul 1985, collector Mardocheo 
Palma: male holotype, c.b. 23.4 mm, c.l. 
15.1 mm; 2 females, c.b. 27.2, 23.2 mm, 
c.l. 17.0, 14.9mm,IBUNAMEM-7030.The 
large female had 1 9 juveniles in the abdo- 
men. 

Gonopod description. —Same as of genus. 

Description.— Carapace slightly convex. 
Superior frontal border formed by sloping 
of carapace; inclined towards median re- 
gion. Inferior frontal border well marked, 
formed by diffuse tubercles; sinuous in fron- 
tal view, arched in dorsal view. Median 
groove short, deep between front and post- 
frontal lobes. Cervical grooves straight. 
Notch on anterolateral margin between or- 
bit and cervical groove. Anterolateral mar- 
gin with 17-19 small blunt denticles be- 
tween cervical groove and epibranchial 
region. Pterygostomian region, around third 
maxillipeds, covered with setae. Third max- 
illiped with ischium/exopod ratio of 0.71. 
Five rows of spines on dactylus of walking 
legs. Strong heterochely in males, in females 
obvious size differences. Cutting margins of 
propodus and dactylus of major chelae with 
alternating large and small teeth, when 
closed large teeth opposing small ones on 
opposite finger. 

Type. —The holotypic male is deposited 
in the Carcinologic Collection, Instituto de 
Biologia, Universidad Nacional Autonoma 
de Mexico (Catalog No. EM-7029). 

Etymology. —From the Latin "tri" (three) 
and "crista" (crest), referring to the three 
crests on the apex of the male's gonopod. 

Remarks. —Smalleyus belongs to the 
Tribe Pseudothelphusini because it exhibits 
a curved sperm channel, a reniform mesial 



process, and a discernible lateral lobe. A 
new genus is created due to the presence of: 
a mesial process whose shape is not com- 
pletely reniform as in Pseudothelphusa or 
Tehuana and bearing spines along its su- 
perior margin; a cephalic serrate projection 
of the mesial process; three ridges (crests) 
on the apex of the gonopod, the median one 
partly filling what would otherwise be the 
apex cavity; a cephalolateral lobe, which 
seems to be a continuation of the median 
apical ridge; a lateral lobe, which corre- 
sponds to the lateral lobe in the genus Pseu- 
dothelphusa, but in this crab occupying a 
more caudal position; and three spine fields, 
one on each one of the three apical ridges. 
The relationships of this genus with the oth- 
er genera of the tribe are not clear. However, 
this aberrant form, together with Spirothel- 
phusa, Lobithelphusa, Odontothelphusa, and 
Disparithelphusa constitute a belt of mono- 
typic genera dispersed along the Isthmus of 
Tehuantepec. Even though these taxa be- 
long to three different tribes, the factors in- 
fluencing the origin of these five genera may 
be similar. In this case, the isthmus repre- 
sents a barrier that these crabs could not 
cross (Rodriguez 1987). Epithelphusa with 
two species, occurring in the same area, has 
more distinct affinities. Rodriguez (1982) 
places the genus linking the tribes Hypo- 
lobocerini and Pseudothelphusini. The af- 
finities of the monotypic genera mentioned 
are obscure, with the exception of Odon- 
tothelphusa which derives from the genus 
Potamocarcinus. 

Pseudothelphusa parabelliana, new species 
Fig. 2 

Material examined. — Cerro El Vigia, Los 
Tuxtlas Biological Station, Veracruz 
(18°35'N, 95°03'W), 7 Jul 1986: male ho- 
lotype, c.b. 24.5 mm, c.l. 15.8 mm; 1 male 
paratype, c.b. 21.4 mm, c.l. 13.0 mm; 2 
females, c.b. 34.1, 19.2, c.l. 21.3, 1 1.9 mm, 
IBUNAM EM-703 1 . Same locality, 30 Aug 
1984, collector Mardocheo Palma: 1 male 



VOLUME 102, NUMBER 1 



47 




Fig. 1. Left gonopod and major chela of Smal ley us tricristatus: a, Total mesial view; b, Cephalic view of 
apex; c, Lateral view of apex; d, Caudal view of apex; e, Apical view; f, Right chela. 



paratype, c.b. 23.9 mm, c.l. 14.8, IBUNAM 
EM-7032; 5 females c.b. 32.3, 29.2, 27.7, 
22.7, 20.9 mm, c.l. 18.8, 17.6, 16.8, 14.0, 
13.6 mm, IBUNAM EM-7033. Same lo- 
cality, 7 Jul 1986, 1 female with 8 juveniles 



in abdomen, c.b. 28.0 mm, c.l. 16.9 mm, 
IBUNAM EM-7034. 

Description.— Carapace slightly convex. 
Superior frontal border absent, but front 
limited by edge. Front inclined in dorsal 



48 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Left gonopod of Pseudothelphusa parabelliana: a, Total mesial view; b, Cephalic view of apex; c, 
Lateral view of apex; d, Caudal view of apex. 



view. Inferior frontal border smooth, some- 
what bilobed in dorsal and frontal views. 
Median groove narrow, deep, dividing su- 
perior edge of front and separating post- 



frontal lobes. Cervical grooves shallow, 
wide, straight; not reaching anterolateral 
margin. Anterolateral margin with slender 
notch lateral to orbit. Anterolateral margin 



VOLUME 102, NUMBER 1 



49 



with 16-19 denticles between cervical 
groove and epibranchial region. Ratio is- 
chium/exopod of third maxilliped 0.91. 
Pterygostomian region covered with setae. 
Major chelae right, fingers not gaping. 

Gonopod description.— -Well developed 
reniform mesial process. Marginal process 
reduced to acute spine. Mesial crest round- 
ed. Lateral lobe completely closing apical 
cavity cephalically and ending in 2 tips dis- 
tally; inferior sharper than superior. Lateral 
lobe with mesial surface triangular. Ter- 
minal pore setae 19. Setal field restricted to 
lateral portion of cavity. Mesial crest higher 
than lateral. Opening of sperm channel cau- 
dal. 

Type.— The holotypic male is deposited 
in the Carcinologic Collection, Instituto de 
Biologia, Universidad Nacional Autonoma 
de Mexico (Catalog No. EM-7028). 

Etymology. —From the Latin "para" (be- 
side) and "belliana" a freshwater crab spe- 
cific name. 

Remarks.— Before this species was de- 
scribed the genus Pseudothelphusa was 
known only from the central part and Pa- 
cific slope of Mexico. The presence of P. 
parabelliana now widens this distribution 
to the 95°W meridian. This species seems 
to be closely related to P. belliana Rathbun, 
1898, which occurs in the states of Guer- 
rero, Michoacan, and Mexico. The gonopod 
of P. parabelliana has a lateral lobe with 
two acute distal tips, a straight mesial crest, 
and the mesial process is much larger in 
relation to the marginal process. In P. bel- 
liana the lateral lobe bears only one distal 
tip, the mesial crest is broadly rounded, and 
the mesial process is slightly larger than the 



marginal process. The occurrence of P. 
parabelliana in the Gulf coast of Mexico 
agrees well with the morphocline proposed 
recently by Rodriguez (1987). He places on 
a west-east axis, across central Mexico, a 
succession of species (including Tehuana 
veracruzana, P. guerreroensis Rathbun, 
1 933, P. belliana, P. americana de Saussure, 
1857, and P. peyotensis Rodriguez & Smal- 
ley, 1969) in which a progressive reduction 
of the lateral lobe can be seen. 

Acknowledgments 

I wish to thank Mardocheo Palma for col- 
lecting most of the crabs included in this 
study, and Dr. Alfred E. Smalley for pro- 
viding much helpful advice and reviewing 
the manuscript. This study was supported 
by a grant from the Mesoamerican Ecology 
Institute of Tulane University. 

Literature Cited 

Pretzmann, G. 1980. Von Dr. Ivo Poglayen-Neuwall 
1975 in Mittelamerika gesammelte Krabben.— 
Annalen Naturhistorisches Museum Wien 83: 
651-666. 

Rodriguez, G. 1982. Les crabes d'eau douce d'Ame- 
rique. Famille des Pseudothelphusidae. — Faune 
Tropicale 22:1-223. 

. 1987. Centers of radiation of freshwater crabs 

in the Neotropics. Pp. 51-67 in R. H. Gore & 
K. L. Heck, eds., Crustacean issues 3: Biogeog- 
raphy of the Crustacea. A. A. Balkema. Rotter- 
dam. 

Department of Biology, Tulane Univer- 
sity, New Orleans, Louisiana, 701 18. Pres- 
ent address: Department of Zoology, Uni- 
versity of Maryland, College Park, Maryland 
20742. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 50-77 

TWO NEW GENERA AND NINE NEW SPECIES OF 

GERYONID CRABS 
(CRUSTACEA, DECAPODA, GERYONIDAE) 

Raymond B. Manning and L. B. Holthuis 

Abstract. —The family Geryonidae Colosi, 1923 is restricted to three genera: 
Geryon Kroyer, 1837, containing two species, G. trispinosus (Herbst, 1803), 
and G. longipes A. Milne Edwrds, 1882; Chaceon, new genus, containing 21 
species, eight of which are newly described; and Zariquieyon, new genus, con- 
taining one new Mediterranean species, Z. inflatus. The new species of Chaceon 
and their ranges are: C. atopus, Saint Helena Island; C. bicolor, central Pacific 
Ocean; C. crosnieri, Madagascar; C. eldorado, northern South America; C. 
inglei, northeastern Atlantic; C. mediterraneus, western Mediterranean Sea; C. 
notialis, Uruguay and Argentina; and C. sanctaehelenae, Saint Helena Island. 



In addition to results published elsewhere 
(see Manning & Holthuis 1984, 1986, 1987, 
1988), our studies on deep-sea crabs of the 
family Geryonidae have revealed the exis- 
tence of nine other undescribed species from 
various localities that can be assigned to two 
new genera. These taxa are named herein. 

Specimens have been deposited in the 
British Museum (Natural History), London 
(BMNH), the Museum National d'Histoire 
Naturelle, Paris (MNHN), the Rijksmu- 
seum van Natuurlijke Historie, Leiden 
(RMNH), the National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton (USNM), and the Zoological Museum, 
Copenhagen (ZMC). 

We use the following abbreviations: cb, 
carapace breadth, measured between the tips 
of the posterior pair of anterolateral teeth; 
cl, carapace length, measured on the mid- 
line; fm, fathoms; leg, collector; m, meter(s); 
mm, millimeter(s); P5, fifth leg (fourth 
walking leg); sta, station. 

Family Geryonidae Colosi, 1923 

Geryonidae Colosi, 1923:249. 

Diagnosis. — Carapace hexagonal, with 
three or five anterolateral teeth on each side. 



Suborbital margin with inner tooth only. 
Chelipeds unequal, merus with subdistal 
dorsal spine, carpus with strong inner spine. 
Chelae portunid-like. Walking legs with na- 
ked dactyli, lacking segments expanded for 
swimming. Abdomens of male and female 
with seven free- segments. 

Remarks. —The status of the family Ger- 
yonidae and of the genera assigned to it have 
long posed problems for students of the 
group, as we already have pointed out 
(Manning & Holthuis 1981:109). That the 
family Geryonidae deserves recognition as 
a family distinct from the Xanthidae s.l. and 
the Goneplacidae s.l. and that its aflinities 
are with the Portunidae no longer seem to 
be in question. 

Several genera of uncertain affinities have 
been placed in the family Geryonidae (see 
Guinot 1971:1077-1078), e.g., Paragalene 
Kossmann, 1878, Bathyplax A. Milne Ed- 
wards, 1880, Platypilumnus Alcock, 1894, 
Progeryon Bouvier, 1922, and Platychelo- 
nion Crosnier & Guinot, 1969. 

In spite of Guinot's (1969:692) observa- 
tion, "Les deux genres monospecifiques 
Progeryon and Platypilumnus nous parais- 
sent etroitement apparentes a Geryon, in- 
separables de celui-ci," we do not believe 



VOLUME 102, NUMBER 1 



51 



that these or any of the genera other than 
Geryon and the two new genera established 
here belong in the Geryonidae. We believe 
that for the moment all other Recent genera 
formerly assigned to the Geryonidae should 
be placed in the Xanthidae s.l. Several of 
them, including Paragalene and Progeryon, 
may well belong in a new family. 

Members of all of the genera here ex- 
cluded from the Geryonidae lack the por- 
tunid-like chelipeds and the characteristic 
subdistal dorsal spine on the merus of the 
chelipeds found in the geryonids proper as 
well as in some portunids, and they all can 
be distinguished from geryonids as restrict- 
ed here by these features alone. Further, all 
of the geryonids as restricted here have na- 
ked dactyli on the walking legs and lack black 
fingers on the chelae, other features that will 
distinguish them from members of Para- 
galene and Progeryon. 

As restricted here, the Geryonidae com- 
prise three genera and 24 species: Geryon 
Kroyer, with 2 species; Chaceon, new genus, 
with 21 species; and Zariquieyon, new ge- 
nus, with 1 species. 

Geryon Kroyer, 1837 

Geryon Kroyer, 1837:10, 20, 21 [type 
species, by original designation and 
monotypy, Geryon tridens Kroyer, 1837, 
a subjective junior synonym of Cancer 
trispinosus Herbst, 1803]. 

Chalaepus Gerstaecker, 1856:118 [type 
species, by monotypy, Cancer trispinosus 
Herbst, 1803]. 

Definition. — Geryonid crabs with three 
anterolateral teeth on each side of the car- 
apace. Carapace length half to two-thirds 
width. Branchial regions not markedly in- 
flated. Frontal teeth poorly developed, small. 
Orbits shallow, rounded. 

Included species.— Two, both from the 
northeastern Atlantic Ocean: 

Geryon longipes A. Milne Edwards (1882: 

16, 39). 
Geryon trispinosus (Herbst) (1803:43) 



(= Geryon tridens Kroyer, 1 837, see Man- 
ning & Holthuis 1987). 

Remarks. —Geryon differs from both 
Chaceon and Zariquieyon in having three 
rather than five anterolateral teeth on each 
side of the carapace, and the two submedian 
frontal teeth are smaller than in the repre- 
sentatives of either of the other two genera. 
Geryon agrees with Chaceon and differs from 
Zariquieyon in having the orbits shallow and 
rounded rather than rectangular, and Ger- 
yon and Chaceon lack the swollen branchial 
regions that are characteristic of the only 
known species of Zariquieyon. 

Chaceon, new genus 

Type species.— Geryon fenneri Manning 
& Holthuis, 1984. 

Name. — The name is given in honor of 
our colleague Fenner A. Chace, Jr. The suf- 
fix -on is used to make the name similar to 
Geryon. This suffix should not be consid- 
ered to be the Greek neuter ending -on. The 
gender of the name Chaceon is masculine, 
like that of Geryon. 

Definition. —Geryonid crabs with five an- 
terolateral teeth on each side of the cara- 
pace. Carapace length half to two-thirds 
width. Branchial regions not markedly in- 
flated. Frontal teeth well-developed, large. 
Orbits shallow, rounded. 

Included species. —Twenty-one, of which 
eight are described herein. Nominal species 
include the following: 

Chaceon ajfinis (A. Milne Edwards & Bou- 
vier, 1894:41), northeastern Atlantic. 

Chaceon chuni (Macpherson, 1 983:23), Na- 
mibia and South Africa. 

Chaceon erytheiae (Macpherson. 1984:86), 
Valdivia Bank, southeastern Atlantic. 

Chaceon fenneri (Manning & Holthuis, 
1984:666), northwestern Atlantic. 

Chaceon gordonae (Ingle, 1985:90), West 
Africa. 

Chaceon granulatus (Sakai. 1978:11), Ja- 
pan. 



52 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




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Chaceon inghami (Manning & Holthuis, 
1986:367), Bermuda. 

Chaceon macphersoni (Manning & Hol- 
thuis, 1988:83), southwestern Indian 
Ocean and South Africa. 

Chaceon maritae (Manning & Holthuis, 
1981:112), West Africa. 



Chaceon paulensis (Chun, 1903:531), cen- 
tral Indian Ocean. 

Chaceon quinquedens (Smith, 1879:35), 
northwestern Atlantic. 

Two other species originally described in 
Geryon, Geryon incertus Miers, 1886, and 



VOLUME 102, NUMBER 1 



53 



Geryon ischurodous Stebbing, 1923, were 
incorrectly assigned to the genus. Manning 
& Holthuis (1986:369) showed that G. in- 
certus is a synonym of the portunid Bathy- 
nectes longispina Stimpson, 1871, and 
Manning & Holthuis (1988:78) showed that 
G. ischurodous is a species of the goneplacid 
genus Carcinoplax. 

Remarks. —Accounts of two other species, 
one from Chile, and one from Brazil are in 
preparation. These 2 species, the 1 1 known 
species, and the 8 new species described be- 
low raise the known number of species to 
21. 

Members of Chaceon agree with Zari- 
quieyon and differ from Geryon in having 
five anterolateral teeth on each side of the 
carapace. 

Chaceon atopus, new species 
Figs. 1-2 

Material— Off Rupert's Bay, Saint He- 
lena Island [15°58'S, 5°43'W], 130 fm (238 
m), 5 Aug 1983, leg A. J. Edwards: one male 
(holotype, RMNH). 

Diagnosis. —A large Chaceon, cl 90 mm, 
cb more than 1 20 mm in only known spec- 
imen, with well-developed anterolateral 
teeth on the carapace in adults and with 
laterally compressed dactyli on the walking 
legs. Carapace 1.4 times broader than long, 
very inflated, strongly convex from front to 
back, appearing smooth. Median pair of 
frontal teeth long and sharp, separated by a 
narrow, V-shaped emargination, medians 
extending further forward than laterals; out- 
er margin of lateral frontal tooth with slight 
convex projection, indicating position of in- 
ner orbital angle. Anterolateral teeth well- 
developed, all sharp, fourth smallest but 
sharp and distinct. Distance from first to 
second tooth about the same as from third 
to fourth, distance from first to third tooth 
less than distance between third to fifth. 
Carapace surface finely granular postero- 
laterally. Suborbital tooth strong and sharp- 
ly pointed, visible in dorsal view, extending 
about to level of lateral frontal tooth; sub- 
orbital margin evenly curved, tuberculate. 




Fig. 2. Chaceon atopus, male holotype, cb 1 24 mm, 
Saint Helena: a, Ventral view of orbit; b, P5 dactylus, 
posterior view; c, P5 dactylus, dorsal view. 



Cheliped merus with sharp spine subdistally 
and with distal dorsal spine; carpus rough- 
ened dorsally, with distal outer spine, den- 
ticulate anterior margin, and strong and 
slender distal spine; propodus with distal 
dorsal spine broken in left chela, right chela 
with distal angled projection. Meri of walk- 
ing legs with distinct distal dorsal spine. 
Dactyli of walking legs laterally com- 
pressed, height at midlength greater than 
width. P5: merus 6.5 times longer than high, 
length 0.58 cb; carpus with line of erect spi- 
nules dorsally; propodus longer than dac- 
tylus, length 5.8 times height. 

Size.— Unique male holotype, cl 90 mm, 
cb 124 mm. 

Remarks. — This species resembles C. 
gordonae, from the Cape Verde Islands and 
Sierra Leone, in many features, including 
the large size, the well-developed frontal and 
anterolateral teeth on the carapace, and the 
compressed dactyli of the walking legs. It 
differs from C. gordonae in having much 



54 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Chaceon bicolor, male paratype, cb 158 mm, Passe de Saint Vincent: a, Dorsal view; th Carapace. 



VOLUME 102, NUMBER 1 



55 



longer walking legs, with the merus more 
than 6 times longer than high, and in its 
habitat. Chaceon gordonae, in which the P5 
merus is less than 6 times longer than high, 
generally occurs in depths in excess of 2000 
m, whereas the present new species occurs 
in less than 250 m. 

This species can be distinguished on sight 
from C sanctaehelenae, described below, 
by the long, slender walking legs, the distal 
meral spine on the walking legs, and the 
well-developed anterolateral spines of the 
carapace. 

Name. — The specific name is from the 
Greek, atopos, out of place, strange, our re- 
action when we learned this specimen was 
specifically distinct from the holotype of C. 
sanctaehelenae, described below. 

Distribution. —Known only from Saint 
Helena Island, in 238 m. 

Chaceon bicolor^ new species 
Figs. 3-4 

Geryon affinis. — Griffin & Brown, 1976:256, 

figs. 7-9.-Sakai, 1978:9, figs. 18-19, pi. 

.2+. fig. D (color). [Not Geryon affinis A. 

Milne Edwards & Bouvier, 1894.] 
Geryon quinquedens.— Intes, 1978:7, figs. 

5B, 8. -Guinot& Richer de Forges, 1981: 

249. -King, 1984:186. [Not Geryon 

quinquedens Smith, 1879.] 
Geryon. -Intes, 1978: fig. lO.-Guinot & 

Richer de Forges, 1981:249. 

Previous records.— Emperor Seamount 
Chain: North of Nintoku Seamount, 
42°20'N, 170°50'E, 800 m; Jingu Seamount 
[38°50'N, 171°15'E], 890-930 m; Kinmei 
Seamount, 35°34'N, 171°41'E, 600-640 m, 
and 500-700 m; south of Kinmei Sea- 
mount, 34°42'N, 171°48'E, 980-1100 m 
(Sakai 1978). 

Southwestern Pacific: in depths greater 
than 600 m (King 1984). 

Loyalty Islands [21°00'S, 167WE]: (Intes 
1978). 

New Caledonia [21°30'S, 165°30'E]: 600- 





Fig. 4. Chaceon bicolor, male paratype, cb 1 48 mm, 
Passe de Saint Vincent: a, Ventral view of orbit; b, P5 
dactylus, posterior view; c, P5 dactylus, dorsal view. 



1000 m (Intes 1978; Guinot & Richter de 
Forges 1981). 

New South Wales, Australia: Southeast 
of Grafton, 29°49'S, 153°42'E to 29°59'S, 
153°38'E, 369 m; east of Broken Bay, 
33°40'S, 151°53'E to 33°22'S, 152°09'E, 450 
m, 33°40'S, 151°53'Eto 33°35'S, 151°58'E, 
450-810 m, and 33°38'S, 151°57'E to 
33°34'S, 152°01'E, 774-792 m; north of 
Sydney, 33°43'S, 151°55'E to 33°37'S, 
152°02'E, 675 m; off Sydney, 33°52'S, 
1 52°50'E to 33°48'S, 1 52°54'E, 765 m (Grif- 
fin & Brown 1976). 

Material. —Emperor Seamount Chain: 
Kinmei Seamount, 35°17.50'N, 171°25.98'E, 
1 50 fm (275 m), Townsend Cromwell cruise 



56 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Cromwell cruise 82-05, sta 57, 1 1 Nov 1982: 
three females (USNM 205974). 

New Caledonia: 23°44'S, 166°58'E, 1490- 
1620 m, BIOCAL, Jean Charcot sta CP 57, 
1 Sep 1985: one juvenile male, one female 
(MNHN), one female (USNM 205975).- 
23°07'S, 166°51'E. BIOCAL, Jean Charcot 
sta CP 32, 825 m, 29 Aug 1985: one male 
(MNHN). -Passe de Saint Vincent [22°02'S, 
165°57'E], 800 m, traps, leg B. Richer de 
Forges: four males (3 MNHN, 1 USNM 
205976). 

Australia: Southeast of Newcastle, New 
South Wales, 33°1 1-09'S, 152°24-25'E, 732 
m, leg D. E. Brown on FRV Kapala, sta 
K77-23-10, 7 Dec 1977: one male (USNM 
205978).-East of Broken Bay, New South 
Wales, 33°35-33'S, 152°00-02'E, 823 m, leg 
D. E. Brown on FRV Kapala, sta K77-23- 
12, 8 Dec 1977: one male, one female 
(USNM 205977). 

The holotype is the male, cl 144 mm, cb 
165 mm, from New Caledonia, Jean Char- 
cot sta CP 32 (MNHN). All of the other 
specimens are paratypes. 

Diagnosis.— A. very large Chaceon, cl to 
more than 140 mm, cb to 180 mm in adults, 
with small, blunt anterolateral teeth on the 
carapace in adults and with laterally-com- 
pressed dactyli on the walking legs. Cara- 
pace 1.1-1.2 times broader than long in 
adults, 1.4 times broader than long in ju- 
veniles. Median pair of frontal teeth nar- 
rower than laterals, separated by U-shaped 
emargination, medians extending further 
forward than laterals. Second and fourth an- 
terolateral teeth reduced, fourth often ob- 
solete. Distance from first to second an- 
terolateral tooth less than that from third 
to fourth, distance from first to third sub- 
equal to that from third to fifth. Carapace 
with distinct granulation mesial to fifth tooth 
and on protogastric, cardiac, and branchial 
regions; hepatic region smooth; protogastric 
region inflated in large specimens, especially 
in females. Suborbital tooth short and blunt 
in males, longer and sharper in females and 
juveniles, scarcely or not at all visible in 



dorsal view, suborbital margin evenly 
curved, tuberculate. Cheliped lightly tuber- 
culate dorsally; upper margin of merus with 
sharp subdistal spine; carpus lacking outer 
spine in adult, blunt projection present in 
some specimens, with distinct spine in ju- 
venile; propodus unarmed distally. Meri of 
posterior walking legs with distinct distal 
dorsal angled projection, spined in juve- 
niles. Dactyli of walking legs laterally com- 
pressed, height at midlength greater than 
width. P5: merus 4.3-5.1 (mean 4.6) times 
longer than high in females, 5.2-5.6 (mean 
5.4) times longer than high in males, and 
7.0-7.5 times longer than high in juveniles, 
usually with blunt distal dorsal projection 
in adults, distinct distal spine in juveniles; 
merus length 0.49-0.56 cb in females, 0.6 1— 
0.64 cb in males; carpus smoothly tuber- 
culate dorsally; propodus distinctly longer 
than dactylus, 3.6-4.6 (mean 4.1) times 
longer than high in adults, 5.6-6.0 times 
longer than high in juveniles. 

Size.— Males, cl 116-147 mm, cb 134- 
165 mm; females, cl 21-110 mm, cb 30- 
122.5 mm; juvenile, cl 7 mm, cb 10 mm. 
The largest specimens recorded in the lit- 
erature are: male, cl 141 mm, cb 155 mm 
(Sakai 1978); a specimen with cb 176 mm 
(Griffin & Brown 1976); a specimen with cb 
180 mm (Lutes 1978). 

Color. —Anterior part of carapace pre- 
dominantly purple, branchial regions tan, 
legs yellowish. Sakai (1978: pi. 2, fig. D) 
gave a figure in color. 

Remarks. — Chaceon bicolor differs from 
all other species of the genus in color pat- 
tern, with the anterior part of the body pur- 
plish rather than reddish or tan, set off from 
the tan branchial regions. Remnants of this 
distinctive color pattern are visible in one 
of the juveniles from deep water off New 
Caledonia. In addition to color pattern, C. 
bicolor also differs from the only other 
species known to occur in the central Pa- 
cific, C granulatus, in having compressed 
rather than depressed dactyli on the walking 
legs; also, the hepatic region of the carapace 



VOLUME 102, NUMBER 1 



57 



in C. granulatus is coarsely granular, where- 
as it is smooth in C. bicolor. 

The juvenile specimens taken in 1490- 
1620 m off New Caledonia differ from the 
adults in many features: the teeth of the 
carapace are much larger and sharper, there 
is a sharp outer spine on the carpus of the 
cheliped and a sharp distal spine on the me- 
rus of each walking leg, and the legs are 
much longer and slenderer. As mentioned 
above, one specimen shows traces of the 
typical color pattern of the adults, suggest- 
ing that these small specimens are juveniles 
that occur in much deeper water than do 
the adults. 

Adult females differ from males in having 
much sharper anterolateral teeth on the car- 
apace, sharper suborbital spines, and much 
shorter legs, with less trace of a distal dorsal 
projection on the merus. Also, the carapace 
of females is more strongly arched from front 
to back and the protogastric regions are no- 
ticeably more inflated. 

Our specimens were taken in depths of 
275, 732, 800, 823, 825, and 1490-1620 m; 
the juveniles were collected at the latter 
depth. Depth records in the literature in- 
clude: 369 to 774-792 m (Griffin & Brown 
1976); 500-700 to 980-1 100 m (Sakai 1978); 
600-1000 m (Intes 1978, Guinot & Richer 
de Forges 198 1); and in depths greater than 
600 m (King 1984). 

Name. — The specific name is from the 
Latin, the prefix bi- and color, referring to 
the purple and tan color pattern of this 
species in life. 

Distribution. —Widely distributed in the 
central Pacific, from the Emperor Seamount 
Chain (42°20'N) to eastern Australia, off 
Sydney (33°52'S), in depths between 275 and 
about 1600 (1490-1620) m. 

Chaceon crosnieri, new species 
Figs. 5-6 

Material. -Madagascar: 22°21'S, 43°05.5'E, 
450-420 m, sta 27, 15 Jan 1986: two males, 
three females (MNHN).-22°13.5'S, 



43°05'E, 530 m, mud, chalutage 62, leg Rudo 
von Cosel, 19 Oct 1986: one female 
(MNHN).-22°26.1'S, 43°04.6'E, 520 m, 
mud, chalutage 65, leg Rudo von Cosel, 
1986: one male, four females (MNHN).— 
22°12.5'S, 43°02.5'E, 560 m, mud, chalu- 
tage 75, leg Rudo von Cosel, 23 Oct 1986: 
two females (MNHN).-22°24.7'S, 
43°04.2'E, 535 m, mud, chalutage 91, leg 
Rudo von Cosel, 4 Nov 1986: one male, 
two females (USNM 205979).-22°24.6'S, 
43°03.7'E, 590 m, mud, chalutage 93, leg 
Rudo von Cosel, 7 Nov 1986: one male, 
one female (MNHN).-22°S, 43°E, 790 m, 
mud, chalutage 102, leg Rudo von Cosel, 
24 Nov 1986: one male, one female 
(MNHN).-22°24.7'S, 43°03.7'E, 630 m, 
mud, chalutage 115, leg Rudo von Cosel, 
28 Nov 1986: one female (MNHN). 

The holotype is the male, cl 1 1 6 mm, cb 
140 mm, from chalutage 93; all other spec- 
imens are paratypes. 

Diagnosis.— A very large Chaceon, cl to 
127 mm, cb to 155 mm in adults, with low, 
blunt anterolateral teeth on the carapace in 
adults and with laterally compressed dactyli 
on the walking legs. Carapace 1.2 times 
broader than long, very inflated dorsally, 
especially at protogastric regions Median 
pair of frontal teeth narrower than laterals, 
not set ahead of laterals, both pairs extend- 
ing about to same level. All five anterolat- 
eral teeth reduced, distance from first to sec- 
ond equal to distance from third to fourth, 
distance from first to third equal to distance 
from third to fifth. Carapace relatively 
smooth, with small granules and pits on 
branchial, cardiac, and gastric regions, he- 
patic regions smooth. Suborbital tooth sharp 
in females, very short and blunt in males, 
scarcely visible in dorsal view, suborbital 
margin concave, tuberculate. Cheliped 
lightly granular, not conspicuously rough- 
ened, merus with sharp subdistal spine; car- 
pus irregular dorsally, with sharp inner 
tooth, lacking any trace of outer tooth; prop- 
odus with some small tubercles dorsally, 
lacking distal spine. Meri of walking legs. 




Fig. 5. Chaceon crosnieri, male holotype, cb 140 mm, Madagascar: a, Dorsal view; b, Carapace. Female 
paratype, cb 127 mm, Madagascar: c, Carapace, lateral view. 



VOLUME 102, NUMBER 1 



59 



especially posterior ones, with distal, dorsal 
spine. Dactyli of walking legs laterally com- 
pressed, height at midlength greater than 
width. P5: merus 4. 1-5.0 (mean 4.7 in males, 
4.5 in females) times longer than high, with 
distal dorsal spine, length 0.47-0.55 cb 
(mean 0.5 1) in females, 0.57-0.63 cb (mean 
0.59) in males; carpus lacking erect spinules 
dorsally; propodus length 3.6-4.4 times 
height (mean 3.9 in males, 4.0 in females), 
usually longer than dactylus, subequal to 
dactylus in large females. 

Size. -Males, cl 83-127 mm, cb 101-155 
mm; females, cl 82-127 mm, cb 97-148 
mm. 

Color.— In preservative, some males show 
tips of fingers of chela with red band proxi- 
mally, apices whitish. Color in life un- 
known. 

Remarks. — Chaceon crosnieri differs from 
C. qffinis in that the carapace is much more 
inflated, the outer orbital and the suborbital 
teeth are stronger, the subdistal tooth on the 
merus of the cheliped is stronger and the 
cheliped is smoother dorsally, the carpus of 
the walking legs lacks dorsal spinules, and 
the merus of the walking legs, especially the 
posterior ones, has a strong distal dorsal 
spine. This new species differs from C. chuni 
in being much larger, cb to 155 mm, the 
carapace is much more inflated, especially 
at the protogastric region, the gap between 
the first and second anterolateral tooth of 
the carapace is larger (equal to distance from 
third to fourth in C. crosnieri, less in C. 
chuni), the frontal teeth of the carapace are 
stronger, the suborbital spine is smaller, and 
the carpus of the cheliped lacks an outer 
spine. 

Chaceon crosnieri is a much smoother 
species than C bicolor, with much shorter, 
stouter legs, the suborbital spine is much 
lower and blunter, and the distal projection 
on the merus of the walking legs is much 
less developed in larger specimens. 

A male of C. paulensis in the USNM, cl 
82 mm, cb 113 mm, from 38°24.92'S, 
77°25.15'E, in 1050-1110 m off Amster- 
dam Island in the southern Indian Ocean, 





Fig. 6. Chaceon crosnieri, male paratype, cb 155 
mm, Madagascar: a, Ventral view of orbit; b, P5 dac- 
tylus, posterior view; c, P5 dactylus, dorsal view. 



is available for comparison. It shares the 
compressed dactyli of the walking legs with 
C crosnieri, and little else. It differs in many 
features, especially in having strongly de- 
veloped, sharp anterolateral teeth on the 
carapace and on the meri of the walking legs, 
and in having a large, sharp outer spine on 
the carpus of the cheliped. 

Chaceon crosnieri was taken together with 
C. macphersoni at sta 27 and at chalutages 
62, 65, 91, and 93. The two species can be 
distinguished immediately, as C macpher- 
soni is much rougher dorsally and has the 
dactyli of the walking legs depressed, not 
laterally compressed. 

Our specimens were taken in depths of 
420-450 to 790 m, with five samples being 
taken in depths between 520 and 590 m. 

Name.— We are pleased to dedicate this 



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Fig. 7. Chaceon eldorado, male holotype, cb 109 mm, Venezuela: a, Dorsal view; b, Carapace. 



VOLUME 102, NUMBER 1 



61 



species to our colleague Alain Crosnier, Mu- 
seum National d'Histoire Naturelle, Paris, 
who made special efforts to save specimens 
of geryonids whenever they were encoun- 
tered in the field. 

Distribution. —Known only from locali- 
ties around Madagascar, in depths between 
420-450 and 790 m. 

Chaceon eldorado, new species 
Figs. 7-8 

Geryon quinquedens. — Takeda, 1983:15, 18, 
31, 164, color fig. on p. 164. [Not Geryon 
quinquedens Smith, 1879.] 

Previous records. —Off Suriname and 
French Guiana, 310-790 m (Takeda 1983). 

Material. -Colombia: 12°06'N, 72°55'W, 
350-500 fm (641-915 m), Oregon sta 4912, 
31 May 1964: one male (USNM 205980). 

Venezuela: 12°55'N, 70°16'W, 340 fm 
(622 m), Oregon II sta 1 1307, 26 Nov 1970: 
one female (USNM 205981).- 1 1°53'N, 
69°25'W, 350 fm (641 m), Oregon sta 4413, 
3 Oct 1963: one male (USNM 205982).- 
1 1°36'N, 62°46'W, 290 fm (531 m), Oregon 
sta 2777, 19 Apr 1960: two males (USNM 
205983). 

French Guiana: 7°37'N, 53°32'W, 395 fm 
(723 m), Oregon II sta 10616, 13 May 1969: 
one male (USNM 205984). 

The male, cl 89.5 mm, cb 109 mm, from 
Oregon sta 4413, is the holotype (USNM 
205982); the other specimens are paratypes. 

Diagnosis. —A moderately large Cha- 
ceon, cl to 89.5 mm, cb to 109 mm in ma- 
terial examined, with low, blunt anterolat- 
eral teeth on the carapace in adults and with 
dorsoventrally depressed dactyli on the 
walking legs. Carapace 1.2-1.3 times broad- 
er than long, inflated dorsally, especially at 
protogastric regions. Median pair of frontal 
teeth about as wide as and extending farther 
forward than laterals. Second and fourth an- 
terolateral teeth usually reduced, much 
smaller and lower than remainder. Distance 
from first to second tooth equal to or less 




Fig. 8. Chaceon eldorado, male holotype, cb 109 
mm: a, Ventral view of orbit; b, P5 dactylus, posterior 
view; c, P5 dactylus, dorsal view. 



than that from third to fourth tooth, dis- 
tance from first to third tooth less than that 
from third to fifth. Carapace granular pos- 
terolaterally and on cardiac and protogastric 
regions, hepatic region smooth. Suborbital 
tooth short, broad, visible in dorsal view, 
lower margin of orbit evenly curved, tuber- 
culate. Cheliped lightly granular; merus with 
sharp subdistal spine; carpus with sharp in- 
ner tooth and distinct angled projection or 



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low tooth on outer margin; propodus un- 
armed distally. Meri of walking legs, espe- 
cially posterior ones, with distal dorsal tu- 
bercle, lacking distinct distal spine. Dactyli 
of walking legs dorsoventrally depressed, 
height at midlength less than width. P5: 
merus 5.3-5.9 (mean 5.5) times longer than 
high, length 0.56 cb in female, 0.61-0.67 
(mean 0.63) cb in males; carpus with line 
of erect spinules dorsally; propodus length 
4.5-5. 1 (mean 4.8) times height, longer than 
dactylus. 

Size. -Males, cl 46-89.5 mm, cb 56-109 
mm; only female examined, cl 48 mm, cb 
64 mm. 

Color. -Takeda's (1983:164) colored fig- 
ure shows a dark red crab. 

Remarks. — This species resembles C. 
quinquedens in having depressed dactyli on 
the walking legs, but differs in that the legs 
are shorter (P5 propodus 5-7 times longer 
than high) and lack a strong distal spine on 
the merus, the propodus of the chela lacks 
a distal dorsal spine, and the carapace is 
more granular. 

The specimens from stations 4912 (male, 
cl 47 m), 11,307 (female, cl 48 mm), and 
10,616 (male, cl 46 mm) have much stron- 
ger anterolateral spines on the carapace than 
those from stations 2777 (males, cl 53.5 and 
55 mm) and 4413 (female, cl 89.5 mm). 

Takeda (1983) reported this species from 
depths between 310 and 790 m. Our spec- 
imens were taken in depths of 531, 622, 
641, 641-915, and 723 m. 

Name. —The name refers to the mythical 
land of gold, El Dorado, thought by six- 
teenth century Spanish explorers to be in 
northern South America. 

Distribution. —Northern coast of South 
America, from Colombia to French Guiana, 
in depths between 531 and 641-915 m. 

Chaceon inglei, new species 
Figs. 9-1 1 

Geryonaffinis.— Hansen, 1908:19, pi. l,fig. 
la, b. [Not Geryon affinis A. Milne Ed- 
wards & Bouvier, 1894.] 



Geryon gordonae Ingle, 1985:90, figs. 3, 4 
[part, not material from Sierra Leone and 
Cape Verde Islands]. 

Previous records. —Northeastern Atlan- 
tic: 61°33'N, 19°0'W, 1089 fm (1993 m) and 
61°30'N, 22°30'W, 975 fm (1784 m) (Han- 
sen 1908).-56°49'-56°48'N, 09°51'- 
09°57'W, 2000 m; 51 6.8'-51°6.9'N, 
13°16.7'-13°24.4'W, 1817-1930 m;51°05.3'- 
51°06.5'N, 13°04.5'-12°59.5'W, 1925-1960 
m; 49°39.5'N, 12°36.9'W, 1857-1910 m; 
49°38.6'N, 12°40.9'W, 1860-1875 m; 
49°32.6'^9°33.5'N, 13°7.1'-13°5.9'W, 1630- 
1640 m; 49°30.1'-49°27.7'N, 13°19.9'- 
13°17.2'W, 1794-1785 m; and 49°27.3'- 
49°30.1'N, 13°21.1'-13°26.8'W, 2045-2110 
m (Ingle 1985). 

Material. —North Atlantic: 61°33'N, 
19°00'W, 1089 fm (1993 m), Ingolf sta 65: 
one male (ZMQ.-61°30'N, 22°30'W, 975 fin 
(1784 m), Ingolf sta. 67: one ovigerous female 
(ZMC).-56°49'-56°48'N, 09°51'-09°57'W, 
2000 m, Challenger sta 12, leg R. W. Ingle: 
one female (BM 1978.102).-55°07.7'N, 
15°11.2'W to 55°10.3'N, 15°09.3'W, 2215- 
2233 m, COB NORATLANTIQUE, PR 
0006, sta CH04, Blake trawl, 10 Aug 1969: 
two males (USNM 205986). -49°27. 3'- 
49°30.1'N, 13°21.1'-13°26.8'W, 2045-2110 
m, Challenger sta 50518, 7 Jun 1979: one 
male (BMNH).-48°45'N, 11°19.8'W to 
48°46.6'N, 11°21.8'W, 1830-1870 m, Thal- 
assa sta 2448, 1973: one female (MNHN 
B.15835).-48°34'N, 10°51.6'Wto 48°32.9'N, 
10°49'W, 1975-2070 m, Thalassa sta 2453, 
1973: one female, two juveniles (MNHN 
B.15834).-47°34.6'N, 08°38.8'W, 2245 m, 
BIOGAS IV, Jean Charcot sta CP 01, 25 Feb 
1974: one male, one female (MNHN 
B.7215).-47°34.5'N, 08°34.2'W, 2180 m, 
CENTOB BIOGAS, sta 01 1, CV 08: one fe- 
male (MNHN B.17207).-47°34.1'N, 
08°40.5'W, 2175 m, CENTOB BIOGAS XI, 
sta 1,CP 37: one female (MNHN B. 17583). - 
47°34'N, 8°41'W, 2100 m, CENTOB, EPI IV, 
sta 1, CP 40, 3 Sep 1985: one juvenile female 
(MNHN B.17585).-47°33.2'N, 08°38.5'W, 
2177 m, CENTOB BIOGAS, PR 08, CP: 



VOLUME 102, NUMBER 1 



63 




Fig. 9. Chaceon inglei, male paratype, cb 6 1 mm, North Atlantic: a, Dorsal view; b, Carapace, enlarged. 



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Fig. 10. Chaceon inglei, male paratype, cb 34 mm, North Atlantic: a, Carapace; b, Ventral view of orbit; c, 
Cheliped, dorsal view; d, Chela, outer view; e, P5;f, P5 dactylus, dorsal view; g, P5 dactylus, posterior view. 



one female (MNHN B.l 582 1).-47°32.8'N, 
08°33.5'W, 2115 m, BIOGAS VI, CP 26: 
two males, one female (USNM 205987).- 
47°32.7'N, 08°34.2'W, 2034 m, BIOGAS 
III, CV 23: one ovigerous female (MNHN 
B.l 7208). -47°31.4'N, 8°42.7'W, 2100 m, 
CENTOB, EPI IV, sta 1, CP 41, 4 Sep 1985: 
one male (MNHN B.17586).-47°31'N, 
08°16'W, 2200 m, BIOGAS I, CV 06: 2 
females (MNHN B.15820).-47°28.1'N, 
08°25.1'Wto 47°29.6'N, 08°22.6'W, 2149- 
2047 m, COB NORATLANTIQUE, 
CH004, PR 124, B0 19, sta 48, Blake Trawl, 
3 1 Oct 1 969: one male, one female (MNHN 
B.15822).-42°32'N, 08°39.4'W, 2100 m, 
CENTOB, EPI I, sta 1, CP 39, 30 Mar 1984: 
one female (MNHN B.17584).-39°04.5'N, 
32°43.5'W, 2120 m, Blake trawl, BIA- 



CORES 1971, sta 131, 24 Oct 1971: one 
male, one female (MNHN B. 15836). 

The female from Challenger sta 12, cl 43 
mm, cb 62 mm, is the holotype (BM 
1978.102). The other specimens are para- 
types. 

Diagnosis.— A small Chaceon, cl to 54 
mm, cb to 72 mm in adults, with sharp, 
well-developed anterolateral spines on the 
carapace in adults and with laterally com- 
pressed dactyli on the walking legs. Cara- 
pace 1.3-1.5 times broader than long. Me- 
dian pair of frontal teeth strong, sharp, 
separated by U-shaped sinus, medians nar- 
rower and extending farther forward than 
laterals. Second lateral tooth small, fourth 
usually obsolete or even totally absent, fifth 
usually largest. Distance from first to second 



VOLUME 102, NUMBER 1 



65 




Fig. 1 1 . Chaceon inglei, North Atlantic: a, Male paratype, cb 1 8 mm, dorsal view; b, Male paratype, cb ca. 
28 mm, dorsal view. 



tooth subequal to or less than distance from 
third to fourth tooth, distance from first to 
third tooth subequal to or less than distance 
from third to fifth tooth. Suborbital tooth 
large, extending at least to level of outer 
frontal tooth, visible in dorsal view, sub- 
orbital margin evenly concave. Cheliped 
roughened dorsally; merus with sharp sub- 
distal and smaller distal spine; carpus with 
long inner spine and shorter outer spine; 
propodus with distal dorsal spine or angled 
projection. Meri of walking legs, especially 
fourth and fifth, with distal, dorsal spine. 
Dactyli of walking legs laterally com- 
pressed, height at midlength greater than 
width. P5: merus more than 6 times longer 
than high, 6.5-7.2 (mean 6.8) times in males, 
6.3-7.5 (mean 6.7) times in females. 8.0- 
9.0 times in juveniles, with distal dorsal 
spine, length 0.52-0.61 cb (mean 0.56 cb) 
in females, 0.53-0.66 cb (mean 0.60 cb) in 
males; carpus with line of erect spinules dor- 
sally; propodus 5.1-6.7 (mean 5.7 in both 
sexes) times longer than high, longer than 
dactylus. 



Size.— Males, cl 22-54 mm, cb 34-72 
mm; females, cl 15-45 mm, cb 24-62 mm; 
ovigerous females, cl 43 and 44 mm, cb 6 1 
mm; juveniles cl 9 mm. Hansen (1908) re- 
ported a male with cl 40 mm, an ovigerous 
female with cl 42 mm. Ingle (1985) studied 
males with cl 1 1.5-49.5 mm, females with 
cl 7-43 mm. 

Remarks. — The long anterolateral teeth 
of the carapace, the long, slender legs, with 
the merus of the fifth leg more than 6 times 
longer than high, the sharp outer spine on 
the carpus of the claw and the merus of the 
walking legs, all will serve to differentiate 
this species from C. affinis, the only other 
species of Chaceon known to occur in the 
northeastern Atlantic off the coast of Eu- 
rope. 

Chaceon affinis, which, like this species, 
has compressed dactyli on the walking legs, 
differs from C inglei of the same size in the 
features mentioned above and in having 
smaller and blunter frontal teeth on the car- 
apace and a much smaller suborbital spine. 
A small male, cl 30 mm, cb 42 mm, of C 



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Fig. 12. Chaceon mediterraneus, male holotype, cb 65 mm, Mediterranean, dorsal view. 



affinis available for comparison (USNM) has 
a much shorter fifth leg, with the merus less 
than 6 times as long as high and the prop- 
odus less than 5 times as long as high. The 
merus shows no trace of the distal, dorsal 
spine that is so prominent in members of 
C inglei. 

Chaceon gordonae, a species that occurs 
off West Africa, is a much larger species, cb 
to at least 161 mm (male from the Cape 
Verde Islands, MNHN B.6432) rather than 
less than 75 mm, that has much shorter legs, 
with the merus less than 6 times longer than 
high. In C gordonae the fourth anterolateral 
tooth of the carapace is present, whereas it 
usually is obsolete in C inglei. 

This species differs from Chaceon medi- 
terraneus, described below, in having com- 
pressed rather than depressed dactyli on the 
walking legs. That species also inhabits rel- 
atively deep water. 

Small specimens that lack the second and 
fourth anterolateral teeth of the carapace 
could be mistaken for Geryon longipes; both 
species occur together in the northeastern 
Atlantic. The frontal teeth of G. longipes are 



much smaller than those of C. inglei at the 
same carapace length. 

This species occurs in relatively deep 
water, with all records coming from depths 
in excess of 1600 m. The specimens re- 
ported by Hansen (1908) were taken in 1 7 84 
and 1993 m, and Ingle (1985) saw material 
from depths ranging from 1630-1640 to 
2045-2110 m. Our specimens were taken 
in depths between 1784 and 2245 m, with 
most lots coming from depths in excess of 
2000 m. 

Name. —We are pleased to dedicate this 
species to our colleague R. W. Ingle, British 
Museum (Natural History). 

Distribution. —Northeastern Atlantic, 
from about 61°N, south of Iceland, to 39°N, 
off the Azores, in depths between 1 630-1 640 
m to 2245 m. 

Chaceon mediterraneus, new species 
Figs. 12-13 

Geryon gordonae. —Delia Croce, Drago, & 
Flocchini, 1988:6, 7, 8 [not Geryon gor- 
donae Ingle, 1985]. 



VOLUME 102, NUMBER 1 



67 




Fig. 1 3. Chaceon mediterraneus, male holotype, cb 
65 mm, Mediterranean: a, Ventral view of orbit; b, P5 
dactylus, posterior view; c, P5 dactylus, dorsal view. 



Previous records. — Mediterranean Sea, 
north of Sardinia, 1990-2008 m (Delia 
Croce, Drago, & Flocchini, 1988). 

Material— Mediterranean Sea, 37°56.7'N, 
07°31.6'E, 2830 m, POLYMED sta CV 04, 
18 May 1970: one male (holotype, MNHN 
B.15824). Off Sardinia, 1990-2016 m; N. 
Drago: one male (USNM); one male, one 
female (BMNH). 

Diagnosis. —A moderately large Cha- 
ceon, cl to 78 mm, cb to 103 mm, with well- 
developed anterolateral teeth on the cara- 
pace and with dorsoventrally depressed 
dactyli on the walking legs. Carapace 1.3 
times broader than long. Median pair of 
frontal teeth broad and short, separated by 
V-shaped emargination, medians extending 
further forward than laterals. Orbit broadly 
concave, with low swelling on margin of 
inner orbital tooth. First, third, and fifth 
anterolateral spines stronger than remain- 
der, fifth strongest, slender; gap between first 
and second tooth very small. Distance from 
first to second tooth subequal to distance 
from third to fourth tooth, distance from 
first to third tooth less than distance from 



third to fifth tooth. Raised areas of carapace 
pitted and eroded, but not tuberculate. Sub- 
orbital tooth strong, sharp, extending about 
to level of lateral frontal tooth, suborbital 
margin tuberculate, evenly rounded. Che- 
liped sparsely tuberculate dorsally; upper 
margin of merus with sharp subdistal spine 
and distal angled lobe; carpus with strong 
inner spine and distinct but smaller outer 
spine; propodus with distinct distal spine. 
Meri of walking legs broad, upper margin 
distinctly convex, with distal, dorsal spine. 
Dactyli of walking legs dorsoventrally de- 
pressed, width near midlength slightly larg- 
er than height. P5: merus 4.5-5.4 times 
longer than high, with distal dorsal spine, 
length about half cb; carpus with line of 
sharp spinules dorsally; propodus 4.1-4.3 
times longer than high, slightly longer than 
dactylus. 

Size. -Males, cl 50-78 mm, cb 65-103 
mm; female, cl 59 mm, cb 82 mm. 

Remarks. —This species resembles C. 
maritae and differs from both C. affinis and 
C. inglei in having relatively broad, de- 
pressed dactyli on the walking legs. It differs 
from C. maritae in having much stronger 
anterolateral spines on the carapace (even 
when specimens of the same size are com- 
pared), in having distal dorsal spines on the 
meri of the walking legs, and in having slen- 
derer walking legs. The merus of the fifth 
leg is more than 5 times longer than high in 
C. mediterraneus, about 4 times longer than 
high in C. maritae. 

Although the dactyli of the walking legs 
of C. mediterraneus are not as conspicu- 
ously flattened as in C maritae, they are 
distinctly broader than the dactyli in C. gor- 
donae or C. inglei. 

It is more than a little surprising to find 
an undescribed Chaceon from the Mediter- 
ranean. Even more surprising is that this 
species was collected together with the new 
genus and species described below. 

The only eastern and central Atlantic 
species with strong anterolateral spines on 
the carapace in adults are C atopus, C gor- 
donae, and C inglei, and all of them have 



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Fig. 14. Chaceon notialis, ovigerous female paratype, cb 110 mm, Argentina: a, Dorsal view; b, Carapace. 



VOLUME 102, NUMBER 1 



69 



distinctly compressed dactyli on the walk- 
ing legs. 

Name.— Derived from the type locality 
in the Mediterranean Sea. 

Distribution. —Known only from the 
western Mediterranean Sea, in 1990-2830 
m. 

Chaceon notialis, new species 
Figs. 14-15 

Geryon quinquedens.—Juanico, 1973:145, 
fig. on p. 147. — Scelzo & Valentini, 1974: 
561, figs. 1-2 [part, not specimens from 
Brazil]. -Boschi, 1976:66, 67; 1979: 
I40.-Boschi et al., 1981:247.-Barea & 
Defeo, 1985:189, fig. 1; 1986:38, fig. 1. 
[Not Geryon quinquedens Smith, 1879.] 

Previous records.— Uruguay and Argen- 
tina: 34°40'-36°45'S, 250-800 m (Barea & 
Defeo 1985).-34°40'-39°04'S, 250-800 m 
(Barea & Defeo 1986).-33°56'-36°25'S, 
52°35'-54°51'W (Juanico 1973). -Uru- 
guay: 33°38'S, 50°38'W, 790 m; 34°48'S, 
52°02'W, 400 m; 35°04'S, 52°06'W, 800 m; 
35°04'S, 52°15'W, 600 m (Scelzo & Val- 
entini 1974). -Argentina: 36°24'S, 53°58'W, 
120 m; 37°45'S, 54°55'W, 300 m; and 
38°55'S, 55°35'W, 170 m (Scelzo & Val- 
entini 1974).— Buenos Aires Province, 36°- 
41°S (Boschi, 1976, 1979).-35°40'S, 
52°47'W, 260 m (Boschi et al. 1981). 

Material. -Argentina: 37°45'S, 54°55'W, 
280-320 m, R/V Cruz del Sur, 17-18 May 
1973: one ovigerous female, paratype 
(USNM 205702).-38°55'S, 55°35'W, 170 
m, R/V Cruz del Sur, 16 Apr 1973: one 
male, holotype (USNM 205701). 

Diagnosis. —A large Chaceon, cl to about 
120 mm, cb to 135 mm in adults, with low, 
blunt anterolateral spines on the carapace 
and with dorsoventrally depressed dactyli 
on the walking legs. Carapace 1.1-1.2 times 
broader than long. Median pair of frontal 
teeth sharp, projecting forward, separated 
by U-shaped sinus, extending farther for- 
ward than laterals. Anterolateral teeth low, 





Fig. 15. Chaceon notialis, male holotype, cb 104 
mm, Argentina: a, Ventral view of orbit; b, P5 dactylus, 
posterior view; c, P5 dactylus, dorsal view. 



blunt, fourth tooth very small, almost ob- 
solete. Distance from first to second an- 
terolateral tooth less than distance from third 
to fourth, distance from first to third an- 
terolateral tooth less than distance from third 
to fifth. Carapace surface strongly granular 
posterolaterally. Suborbital tooth low, blunt, 
falling short of level of lateral frontal tooth, 
suborbital margin evenly curved. Cheliped 
with blunt tooth subdistally and with distal 
angled lobe on merus; carpus roughened 
dorsally, lacking distal outer spine, anterior 
margin denticulate; propodus lacking distal 
spine or angled projection. Meri of walking 
legs with at most a distal dorsal tubercle, 
lacking a distinct distal dorsal spine. Dactyli 
of walking legs depressed, width at mid- 
length much greater than height. P5: merus 
less than 5 times longer than high, length 
0.60 cb; carpus with line of low but sharp 
spinules dorsally; propodus length less than 
4 times height, slightly longer than dactylus. 



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Size. — Male, cl 94 mm, cb 104 mm; ovig- 
enous female, cl 99 mm, cb 110 mm. Jua- 
nico (1973) reported a male with cl 61 mm, 
cb 73 mm; Boschi et al. (1981) studied a 
male cl 30 mm. Scelzo & Valentini (1974) 
studied 1 8 specimens, measuring as follows: 
males, cl 88-116 mm, cb 97-124 mm; fe- 
males, cl 72-99 mm, cb 82-1 10 mm. Barea 
& Defeo (1985) reported catching males with 
cb 32-1 35 mm, females with cb 48-124 mm. 

Color.— Scelzo & Valentini (1974) re- 
ported that this species is reddish in color. 

Remarks. — Chaceon notialis resembles 
both C. quinquedens, from the northeastern 
Atlantic, and C. maritae, from West Africa, 
in having depressed dactyli on the walking 
legs, but it differs from both in numerous 
features. It is a much shorter-legged species 
than C. quinquedens: the merus of the fifth 
leg is about three-fifths the width of the car- 
apace as opposed to three-fourths the car- 
apace width in C. quinquedens, and the 
propodus of the fifth leg is less than 4 times 
as long as high instead of 5-7 times as long 
as high. In C. quinquedens there is a sharp 
outer spine on the carpus and a distal dorsal 
spine on the propodus of the cheliped, and 
each walking leg has a distinct distal dorsal 
spine. These spines are not present on the 
cheliped of G. notialis, and in this latter 
species the meri of the walking legs are armed 
with at most a distal dorsal tubercle. The 
surface of the carapace also is much coarser 
in C. notialis, especially posterolateral^. 

Chaceon notialis differs from C. maritae 
as follows: the frontal teeth are sharper and 
are not as close together; the anterolateral 
teeth of the carapace all are sharper, and the 
fourth tooth is much more distinct; the sub- 
orbital teeth are sharper and longer; and the 
walking legs are slightly longer. The merus 
of the fifth leg is about 5 rather than 4 times 
as long as high, and is about three-fifths 
rather than two-thirds of the carapace width. 

Barea & Defeo (1985, 1986) provided de- 
tails of the occurrence and biology of this 
species on joint fishing grounds off Uruguay 
and Argentina. Boschi (1976, 1979) char- 



acterized it as a cold-temperate species of 
the Buenos Aires Province (36-41° South 
latitude). 

The specimen from off Brazil identified 
with Geryon quinquedens by Rathbun (1937: 
270) and material from Brazil identified by 
Scelzo & Valentini (1974:560) with the same 
species is referable to a new species, the de- 
scription of which is in preparation. Scelzo 
& Valentini reported that their Brazilian 
specimens were cream-colored, whereas 
their material from Uruguay and Argentina 
was reddish. Further, the length/height ratio 
of the propodus of the walking legs of their 
specimens from Brazil was 4.3-4.5, longer 
than that of C. notialis. The range in the 
length/height ratio observed by Scelzo & 
Valentini for their specimens from Uruguay 
and Argentina, with one exception, is as we 
found in our material of C. notialis. 

Our specimens were taken in depths of 
170 and 280-320 m. Boschi et al. (1981) 
recorded a specimen from 260 m. Scelzo & 
Valentini (1974) reported specimens from 
depths of 120, 170, 300, 400, 600, 790, and 
800 m, and Barea & Defeo (1985, 1986) 
reported this species from fishing grounds 
in depths between 250 and 800 m. 

Name. — The specific name is from the 
Latin, notialis, southern. 

Distribution. —Known from the coasts of 
Uruguay and Argentina, from about 33°S to 
about 41°S, in depths between 120 and 
800 m. 

Chaceon sanctaehelenae, new species 
Figs. 16-17 

Material— Sandy Bay, St. Helena Island 
[15°58'S, 5°43'W], trap, 8 Oct 1968, leg F. 
N. Martin: one male (holotype, USNM 
125510). 

Diagnosis. —A large Chaceon, cl 1 1 3 mm, 
cb 134 mm in adult, with low, blunt an- 
terolateral teeth on the carapace in adults 
and with laterally compressed dactyli on the 
walking legs. Carapace about 1.2 times 
broader than long. Median pair of frontal 



VOLUME 102, NUMBER 1 



71 




^^^^^.^^ 






2v: 







Fig. 16. Chaceon sanctaehelenae, male holotype, cb 134 mm, Saint Helena: a, Dorsal view; b, Carapace. 



teeth broad, obtuse, separated by shallow as low lobes in adults. Distance from first 

U-shaped emargination, medians extending to second anterolateral tooth subequal to 

farther forward than laterals. Second and distance from third to fourth tooth, distance 

fourth anterolateral teeth obsolete, present from first to third tooth subequal to distance 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 17. Chaceon sanctaehelenae, male holotype, 
cb 1 34 mm, Saint Helena: a, Ventral view of orbit; b, 
P5 dactylus, posterior view; c, P5 dactylus, dorsal view. 



from third to fifth. Carapace surface finely 
granular, hepatic regions largely smooth. 
Suborbital tooth short, blunt, scarcely vis- 
ible in dorsal view, falling short of level of 
lateral frontal tooth, suborbital margin 
evenly rounded. Cheliped with blunt tooth 
subdistally on merus; carpus roughened 
dorsally, lacking outer spine, anterior mar- 
gin smooth; propodus lacking distal angled 
projection. Meri of walking legs lacking dis- 
tal, dorsal spine. Dactyli of walking legs 
compressed, height at midlength greater than 
width. P5: merus 4.5 times longer than high, 
length 0.6 cb; propodus slightly longer than 
dactylus, length 3.3 times height. 

Size. — Male, cl 113 mm, cb 134 mm. 

Remarks. —This new species differs from 
both C. affinis and C. atopus in having much 
lower and blunter frontal and anterolateral 
teeth on the carapace; in C. affinis the sec- 
ond and fourth teeth are well-developed in 



large specimens, but in C. sanctaehelenae 
the second and fourth teeth are almost com- 
pletely obsolete. In addition, in C. sanctae- 
helenae the orbit appears to be shallower, 
the suborbital tooth is shorter and more ob- 
tuse, and the carpus of the walking legs is 
much smoother dorsally than in C affinis. 
This new species further differs from C. ato- 
pus in having much shorter walking legs (P5 
merus 4.5 times rather than 6.5 times longer 
than high), and in having the distance from 
the first to third anterolateral teeth on the 
carapace subequal to the distance from the 
third to the fifth tooth; in C. atopus the dis- 
tance from the first to third tooth is shorter. 

Chaceon sanctaehelenae differs from C. 
fenneri in some of these same features. All 
marginal teeth of the carapace are lower than 
in C. fenneri', in that species the frontal teeth 
are much sharper and more prominent, and 
the first, third, and fifth anterolateral teeth 
are much more prominent. The carapace is 
smoother than in C. fenneri, with lower and 
fewer granules, and the dorsal ridge on the 
carpus of the walking legs is much smooth- 
er. 

Chaceon chuni, from the southeastern At- 
lantic, differs in having a larger suborbital 
spine and in having a distinct distal dorsal 
spine on the merus of each walking leg. 

Name. — The specific name is derived 
from the type locality. 

Distribution. —Known only from the type 
locality. 

Zariquieyon, new genus 

Type species.— Zariquieyon inflatus, new 
species. 

Name. —The name is dedicated to the late 
Spanish carcinologist, Ricardo Zariquiey 
Alvarez, whose studies of Mediterranean 
decapods added so much to our knowledge 
of the group. As in Chaceon, the ending 
-on is added to make the name similar to 
Geryon. The gender is masculine. 

Definition. — Geryonid crabs with five an- 
terolateral spines on each side of the cara- 



VOLUME 102, NUMBER 1 



73 



pace. Carapace length about seven-tenths 
width. Branchial regions markedly inflated. 
Frontal teeth well-developed, large. Orbits 
deep, rectangular. 

Included species.— One, Zariquieyon in- 
flatus, n. sp. 

Remarks. — Zariquieyon shares with 
Chaceon and Geryon the portunid-like che- 
lipeds, with the subdistal dorsal spine on 
the merus, the strong inner spine on the 
carpus, the portunid-like chela, the single 
inner suborbital tooth, the slender walking 
legs with their naked dactyli, and the seven- 
segmented abdomen in both sexes. Zari- 
quieyon differs from those genera in having 
a comparatively broader carapace, with the 
posterolateral margins convex, strongly in- 
flated branchial regions, and rectangular 
rather than rounded orbits. It also shows far 
more pebbling dorsally, visible only under 
magnification, than any species of Chaceon 
or Geryon. 




Fig. 18. Zariquieyon inflatus, female holotype, cb 
35 mm, Mediterranean, dorsal view. 



Zariquieyon inflatus, new species 
Figs. 18-19 

Material— Mediterranean Sea, 37°56.7°N, 
07°31.6'E, 2830 m, POLYMED sta CV 04, 
18 May 1970: one male, two females 
(MNHN; one female USNM 205985). 

The holotype is the female, cl 22 mm, cb 
35 mm, in the collection of the Museum 
National d'Histoire Naturelle, Paris; the 
other specimens are paratypes. 

Diagnosis. —A very small geryonid, cl 19- 
22 mm, cb 27-35 mm, with strongly de- 
veloped anterolateral teeth on the carapace 
in adults and with laterally compressed dac- 
tyli on the walking legs. Body, except ab- 
domen, covered with fine granules, coarser 
and larger on posterior branchial regions. 
Regions of carapace well-marked, proto- 
gastric region, area adjacent to fifth antero- 
lateral spine, and branchial regions very in- 
flated. Intestinal region with two 
prominences. Posterolateral margins 
strongly convex. Carapace 1.4-1.6 (mean 
1.5) times broader than long. Median pair 



of frontal teeth slender, separated by 
V-shaped emargination. Second and fourth 
anterolateral teeth reduced, fourth smallest; 
other teeth progressively increasing in size 
posteriorly, fifth much the largest. Distance 
from first to second tooth short, about the 
same as from third to fourth tooth, distance 
from first to third tooth much less than dis- 
tance from third to fifth. Orbits rectangular. 
Suborbital tooth strong, extending to level 
of outer frontal tooth, suborbital margin 
denticulate. Cheliped finely granular dor- 
sally; merus with sharp subdistal spine, dis- 
tal angled projection spinulose; carpus peb- 
bled dorsally, with long, sharp inner spine 
and distinct outer spine or crest of spinules; 
propodus pebbled dorsally, with distal dor- 
sal projection. Meri of walking legs with dis- 
tinct distal dorsal spine. Dactyli of walking 
legs laterally compressed, height near mid- 
length greater than width. P5: merus 6.3- 
6.6 times longer than high, pebbled dorsal- 
ly, with distal dorsal spine, length 0.45 times 
cb; carpus with line of spinules dorsally; 



74 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 19. Zariquieyon inflatus, female holotype, cb 35 mm, Mediterranean: a, Dorsal view; b, Ventral view 
of orbit; c, Third maxilliped; d, Carpus of cheliped; e, P5;f, P5 dactylus, dorsal view. 



propodus length about 5 times height, much 
longer than dactylus. 

Size. — Male, cl 19 mm, cb 28 mm; fe- 
males, cl 20-22 mm, cb 27-35 mm. 

Color. —Not recorded. 

Remarks.— This species was taken to- 
gether with Chaceon mediterraneus at a 
depth of 2830 m in the western Mediter- 
ranean basin. We know of no deeper record 
for a brachyuran crab from the Mediterra- 
nean. 

Name.— The specific name is from the 
Latin, inflatus, swollen. 

Distribution. —Known only from the type 
locality, in 2830 m. 

Remarks 

The genus Chaceon has proven to be much 
more speciose than even we imagined when 
we began studying its members, and much 



additional work remains to be done on the 
genus. The geographic ranges of most species 
remain to be determined; in general, ranges 
of each species are restricted latitudinally, 
although some species, like C. bicolor, have 
relatively wide ranges. Chaceon paulensis 
from the southern Indian Ocean should be 
redescribed, and the status of the species 
reported from off Travancore, India, in 224- 
284 fm (410-520 m) by Alcock (1899:85) 
needs to be determined. A particularly in- 
teresting problem that remains to be solved 
is whether the types of Chaceon quinque- 
dens, one of which is an ovigerous female 
only 22 mm long (Smith, 1879:36), are con- 
specific with the large, commercially im- 
portant species now identified as C. quin- 
quedens. 

For most species there is no information 
available on growth changes, especially as 
it pertains to proportions of the walking legs 



VOLUME 102, NUMBER 1 



75 



Table 1.— Characteristics of known species of Chaceon. Abbreviations are as follows: ? (not known); Color: 
B (bicolor, tan and purple), R (red), T (tan), W (whitish); Size (maximum cb in mm); P5 dactylus: C (laterally 
compressed), D (dorsoventrally depressed); Distal spine on P5 merus: + (large, well developed), - (small, 
reduced or absent); Anterolateral teeth of carapace: + (large, well-developed in adults), - (small, reduced or 
absent). 





Color 


Size 


Dactylus 
P5 


Merus 
P5 


Carapace 
spines 


Northwestern Atlantic 












C. eldorado 
C. fenneri 
C. inghami 
C. quinquedens 


R 
T 
R 
R 


109 
190 
110 
178 


D 
C 
C 
D 


+ 
+ 


+ 


Southwestern Atlantic 












C. notialis 
n. sp. Brazil 


R 
T 


135 
158 


D 
D 


— 


— 


Northeastern Atlantic 












C. qfflnis 
C. inglei 
C. mediterraneus 


T 
R 
? 


210 

72 
103 


C 
C 
D 


+ 
+ 


+ 
+ 


Central South Atlantic 












C. atopus 

C. sanctaehelenae 


? 
? 


124 
134 


C 
C 


+ 


+ 


Southeastern Atlantic 












C. chuni 
C. erytheiae 
C. gordonae 
C. maritae 


T 
RT 

9 

T 


114 
125 
161 

172 


C 
C 
C 
D 


+ 
+ 
+ 


+ 


Western Indian Ocean 












C. crosnieri 
C. macphersoni 
C. paulensis 


? 
? 
R 


155 
126 
113 


C 
D 
C 


+ 
+ 
+ 


+ 
+ 


Western and Central Pacific 












C. bicolor 
C. granulatus 


B 
T 


180 
146 


C 
D 


— 


— 


Eastern Pacific 












n. sp. Chile 


W 


175 


C 


- 


- 



and spination of the carapace and walking Chaceon may be difficult to distinguish in 
legs. In the case of C. bicolor, for example, the absence of comparative material, we 



legs. In the case of C. bicolor, for example 
we have identified a small, spiny, long-legged 
specimen from deep water with the much 
larger adult found in the same area, but the 
possibility remains that more than one 
species is involved. 

Certainly additional species remain to be 
recognized, and we believe that a key to the 
species now known would be premature and 



the absence of comparative material, we 
provide here, by broad geographic region, a 
summary of the species found in each region 
and their major characteristics (Table 1). 

Acknowledgments 

Many people contributed to this study. 
We thank all of those who provided us with 
material or access to their collections: E. E. 



perhaps even confusing. As the species of Boschi, Instituto Nacional de Investigation 



76 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



y Desarollo Pesquero, Mar del Plata, Ar- 
gentina; Diane E. Brown, Australian Mu- 
seum, Sydney; Alain Crosnier and D. Gui- 
not, Museum National d'Histoire Naturelle, 
Paris; Reginald M. Gooding, then of the 
NOAA/NMFS Southwest Fisheries Center, 
Honolulu Laboratory; R. W. Ingle, British 
Museum (Natural History), London; En- 
rique Macpherson, Instituto de Ciencias del 
Mar, Barcelona; Torben Wolff, Zoological 
Museum, Copenhagen; and Nicolino Dra- 
go, Universita di Genova, Italy, who al- 
lowed us to add reference to his material 
from the Mediterranean in galley. 

Our studies on geryonid crabs have been 
supported in part by the Food and Agri- 
culture Organization of the United Nations 
(FAO); that support is gratefully acknowl- 
edged. This paper also was supported in part 
by the Smithsonian Institution, through its 
Research Opportunities Fund. 

We thank Roy K. Kropp for taking the 
photographs used in Figs. 14 and 16. Fig. 1 
was taken by the staff photographer of the 
Rijksmuseum van Natuurlijke Historie, 
Leiden, and Figs. 3 and 5 were taken by the 
staff photographer of the Museum National 
d'Histoire Naturelle, Paris. Lilly King Man- 
ning drew the line drawings and prepared 
all of the figures for publication. 

We thank Fenner A. Chace, Jr., Enrique 
Macpherson, and Marilyn Schotte for read- 
ing a draft of the manuscript. 

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(RBM) Department of Invertebrate Zo- 
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tuurlijke Historie, Postbus 9517, 2300 RA 
Leiden, The Netherlands. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 78-83 

SYNALPHEUS AROSTRIS AND PHILOCHERAS LAPILLUS, 

TWO NEW SPECIES OF CARIDEAN SHRIMP 

(CRUSTACEA) FROM THE TROPICAL 

EASTERN PACIFIC 

Mary K. Wicksten 

Abstract.— Two new species of shrimp (Decapoda: Caridea) are described 
from the eastern Pacific. Synalpheus arostris is described from specimens from 
western Colombia. Philocheras lapillus is the first species of its genus to be 
reported in the area. 



During examination of specimens from 
western Colombia and the Galapagos Is- 
lands, I discovered two undescribed species 
of shrimp. The new species of Philocheras 
is the first report of the genus from the trop- 
ical eastern Pacific. I thank Gabriel Ramos, 
Universidad del Valle (UV); Raymond 
Manning, United States Museum of Natural 
History (USNM); and Janet Haig, Allan 
Hancock Foundation, University of South- 
ern California (AHF) for their interest and 
assistance during this study. The figures are 
by Gabriel Ramos and Joseph Goy, Texas 
A&M University. This project was aided by 
a Visiting Fellowship of the Smithsonian 
Institution. 

Family Alpheidae 

Synalpheus arostris, new species 
Fig. 1 

Type material. — Holotype: female, total 
length 10.5 mm. Morro de Los Agujeros, 
Bahia de Malaga, Colombia (3°55'N, 
77°20'W), rocky intertidal zone, 28 Nov 
1981, collector not reported, AHF no. 8111. 
Paratype: female, same site and date, UV. 

Description.— Rostrum lacking, a slight 
raised area present between orbital teeth. 
Orbital teeth reaching less than 0.5 x length 
of first segment of antennular peduncle, 
longer than broad. 

Visible part of first antennular segment 



1.3 x as long as second antennular article, 
third article shorter. Stylocerite reaching just 
beyond first antennular segment. Scapho- 
cerite with lateral tooth reaching beyond end 
of antennular peduncle and exceeding car- 
pocerite. Squamous part of scaphocerite 
reaching or almost reaching end of anten- 
nular peduncle. Inferior tooth of basicerite 
reaching as far forward as stylocerite, su- 
perior tooth well developed and acute. 

Large chela 3 x as long as wide, with fin- 
gers occupying about 0.3 x entire length. 
Superior margin of palm bearing blunt knob 
proximal to dactylus. Merus 2x long as 
broad, superior margin blunt. 

Small chela 2.4 x long as wide, with fin- 
gers occupying about 0.4 x entire length. 
Merus almost 3 x long as wide, bearing 3 
teeth on superior margin. 

Carpal articles of second leg with ratio 1 0: 
3:3:3:4. 

Merus of third leg 4 x long as broad, with- 
out spines. Carpus without teeth or spines. 
Propodus 7 x long as wide, straight, bearing 
7-8 spines on inferior margin and a pair of 
terminal spines. Dactyl biunguiculate, 0.3 x 
propodus, relatively straight; superior and 
inferior ungui slender and almost equal in 
size and shape. Fourth and fifth legs similar 
to third. 

Telson only slightly longer than width of 
anterior margin, tapering to posterior mar- 
gin. Two to three dorsolateral spines per 



VOLUME 102, NUMBER 1 



79 




Fig. 1 . Synalpheus arostris, A, Holotype in lateral view; B, Anterior end in dorsal view; C, Telson; D, Small 
cheliped; E, Large cheliped; F, Third pereopod. 



side, pair long spines at distolateral margin, 
distolateral margin produced into small 
tooth. Posterior margin of telson rounded. 
Discussion. — Coutiere (1909) established 
six groups in the genus Synalpheus, based 
on features of the rostrum, orbital teeth, se- 
tae of the smaller chela, ungui of the dactyls, 
and other features. Banner & Banner (1975) 
reexamined these groups and found that 
perhaps only two of them were of taxonom- 
ic validity— the others contained such a 
range of variation or overlapped other 
groups to such an extent that the group cer- 
tainly was artificial. The new species illus- 
trates the difficulties in finding relationships 
among the species of Synalpheus; lacking a 



rostrum, it cannot easily be compared with 
many other species. 

The short orbital teeth and equal, slender 
ungui of the dactyls of S. arostris are found 
in species of Coutiere's "Brevicarpus" group, 
known only from American waters. Three 
species, S. minus (Say), S. obtusifrons Chace, 
and S. brevicarpus Coutiere are found in the 
Caribbean region. All of these species have 
short rostra. Synalpheus minus has a sty- 
locerite that greatly exceeds the first article 
of the antennular peduncle, but the spine of 
the scaphocerite does not reach the end of 
the carpocerite. Synalpheus obtusifrons has 
a rounded stylocerite and a short scapho- 
cerite that does not reach the end of the 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Philocheras lapillus, paratype: A, Entire animal in lateral view; B, Carapace in dorsal view; C, Carapace 
in lateral view; D, Telson and uropods. 



antennular peduncle. The eastern Pacific 
species, S. digueti Coutiere, is common in 
rocky intertidal areas. Aside from lacking a 
rostrum, S. arostris can be distinguished 
from S. digueti by having a shorter stylo- 
cerite, reaching barely beyond the end of 
the first article of the antennular peduncle 
rather than to the middle of the second ar- 
ticle; and having a longer spine on the 
scaphocerite, reaching proportionally far- 
ther beyond the blade than in S. digueti. 
Only S. arostris has three teeth on the su- 
perior margin of the merus of the small che- 
liped. 

Synalpheus arostris can be distinguished 
easily from S. lockingtoni Coutiere by the 
lack of the rostrum (seen in both specimens 
of the new species). Should one encounter 
a specimen of S. lockingtoni missing a ros- 
trum, the species can be distinguished by 
the sharp spine at the anterior margin of the 



palm of the major chela, above the articu- 
lation of the dactyl, in S. arostris. In S. lock- 
ingtoni, the anterior margin of the major 
chela ends in a knob or tubercle. The pos- 
terior margins of the telson of S. arostris are 
sharply triangular, those of S. lockingtoni 
are weakly pointed. Adult S. lockingtoni can 
be larger than the specimens of S. arostris: 
Schmitt (1921) gives a length of 30 mm for 
a large female S. lockingtoni. 

Family Crangonidae 

Philocheras lapillus, new species 

Figs. 2-4 

Type material. — Holotype male, total 
length 13.7 mm. Off Gardner Bay, Hood 
Island (Isla Espafiola), Galapagos (Archi- 
pelago de Colon) 1°22'S, 90°40'W), 46-65 
m, rock, 31 Jan 1934, Velero III sta. 201- 
34, USNM 234435. 



VOLUME 102, NUMBER 1 



81 





Fig. 3. Philocheras lapillus: A, Antennule; B, Scaphocerite; C, Mandible; D, First maxilla; E, Second maxilla: 
F, First maxilliped; G, Second maxilliped; H, Third maxilliped. 



Paratypes. — Tagus Cove, Albemarle Is- 
land (Isla Isabela) (0°17'S, 91°23'W), 37 m, 
rock and nullipores, 11 Jan 1934, Velero HI 
sta. 149-34, one specimen. Stephens Bay, 
Chatham Island (Isla San Cristobal), (0°48'S, 
89°31'W), 59 m, fine sand and coralline al- 
gae, Velero III sta. 170-34, 21 Jan 1934, 
three specimens. Off Cartago Bay, Albe- 
marle Island (0°35'S, 90°54' W), 59 m, mud, 
25 Jan 1934, Velero III sta. 185-34, five 
specimens. Off Cartago Bay, Albemarle Is- 
land (0°34'S, 90°53'W), 59 m, sand and nul- 
lipores, 25 Jan 1934, Velero III sla. 186-34, 
two specimens. Off Gardner Bay, Hood Is- 
land (same station as holotype), Velero III 
sta. 201-34 (four specimens USNM, two 
specimens AHF). All specimens USNM ex- 
cept as noted. 

Description. —Rostrum exceeding or 
nearly exceeding end of cornea, distal end 
wide. Anterior margin of carapace with 
teeth, one suborbital tooth, pair near base 
of second antenna and two pterygostomial 
teeth. One large tooth on dorsal midline. 

Abdominal pleura rounded. Sixth ab- 
dominal segment about 1 .5 x length of fifth. 



slightly shorter than length of telson. Telson 
narrow, with two pair dorsolateral, three pair 
terminal spines and sharp posteromesial tip. 

Basal article of antennular peduncle not 
reaching as far as end of cornea. Stylocerite 
not as long as basal article of antennular 
peduncle, broad and quadrangular. Last two 
articles of antennular peduncle short. 

Scaphocerite longer than antennular pe- 
duncle, outer margin straight, blade longer 
than spine. Basicerite short. 

Third maxilliped exceeding scaphocerite. 
Ultimate segment longer than penultimate, 
0.6 x antepenultimate. Exopod present. 
Other mouthparts as figured. 

First pereopod stout, subchelate. Dactyl 
about 0.3 x propodus length, subchelar spine 
narrow and simple. Carpus 0.25 x propo- 
dus, with one or no spines. Merus shorter 
than chela, with two teeth on distal end near 
articulation with carpus and one large tooth 
on lower margin. Ischium and basis short, 
without teeth. No exopods or epipods on 
pereopods. 

Second pereopod short, chelate. Dactyl 
0.5 x propodus. carpus about equal to chela, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Philocheras lapillus: A. First pereopod; B, Second pereopod; C, Third pereopod; D. Fourth pereopod; 
E, Fifth pereopod: F. Second pleopod of female. 



0.5 x merus. Merus longer than ischium. 
Third pereopod slender, thread-like. Dactyl 
thin, 0.7 x propodus. Propodus 0.6 x car- 
pus, carpus 1.8 x merus, merus 0.6 x is- 
chium. Fourth and fifth pereopods similar. 
Dactyl long and simple, 0.8 x propodus. 
Carpus 0.8 x propodus, merus 1.2 x carpus, 
ischium 0.8 x merus. 

Second pleopod with appendix interna, 
male with appendix masculina. Uropods 
long, outer uropod shorter than inner. Outer 
uropod with small posterolateral tooth. 

Color in life. —Carapace almost all white, 
body Van Dyke brown, speckled. (Field note 
for specimen at sta. 149-34, by Waldo L. 
Schmitt.) 

Discussion. —I follow the revision of 
Chace (1984) in considering Philocheras to 
be distinct from Pontophilus. Philocheras 
lapillus most closely resembles P. gorei 
(Dardeau, 1980) from the Gulf of Mexico 



and Georgia. In P. gorei, however, there are 
always teeth on the distal margins of the 
carpus of the first pereopod. The outer fla- 
gellum of the antennular peduncle of P. la- 
pillus is much thicker than that of P. gorei. 
The tip of the rostrum of P. lapillus is round- 
ed, not spatulate. Pontophilus lapillus also 
may be larger than P. gorei: the carapace 
length of the holotype is 3.3 mm, that of the 
type of P. gorei is 2.5 mm. 

The habitat and living color of P. lapillus 
suggest that the animal's mode of life is sim- 
ilar to that of species of Crangon and Lis- 
socrangon along the coast of California. 
These species have a speckled, cryptic color 
pattern, which provides camouflage when 
the shrimp dig into sand (Kuris & Carlton 
1977; Ricketts et al. 1985). 

The species epithet means "pebble," 
which the resting animal resembles. The 
name is a noun in apposition. 



VOLUME 102, NUMBER 1 



83 



Literature Cited 

Banner, D. M., & A. H. Banner. 1975. The alpheid 
shrimp of Australia. Part 2: the genus Synal- 
pheus. — Records of the Australian Museum 
29(12):267-389. 

Chace, F. A., Jr. 1984. The caridean shrimps (Crus- 
tacea: Decapoda) of the Albatross Philippine Ex- 
pedition, 1907-1910, Part 2: Families Glypho- 
crangonidae and Crangonidae. — Smithsonian 
Contributions to Zoology No. 397:1-63. 

Coutiere, H. 1 909. The American species of snapping 
shrimps of the genus Synalpheus. —Proceedings 
of the United States National Museum 36(1659): 
1-93. 

Dardeau, M. 1980. A new species of Pontophilus 
(Crustacea: Natantia: Crangonidae) from the 
Gulf of Mexico and the western Atlantic — Pro- 



ceedings of the Biological Society of Washington 
93(3):563-572. 

Kuris, A. M., & J. T. Carlton. 1977. Description of 
a new species, Crangon handi, and new genus, 
Lissocrangon, of crangonid shrimps (Crustacea: 
Caridea) from the California coast, with notes 
on adaptation in body shape and coloration.— 
Biological Bulletin 153:540-559. 

Ricketts, E. F., J. Calvin, J. W. Hedgpeth, & D. W. 
Phillips. 1985. Between Pacific tides, 5th ed. 
Stanford University Press, Stanford, Califor- 
nia, 652 pp. 

Schmitt, W. L. 1921. Marine decapod Crustacea of 
California. — University of California Publica- 
tions in Zoology 23:1-470. 

Department of Biology, Texas A&M Uni- 
versity, College Station, Texas 77843. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 84-88 

THE HOLOTYPE OF HETEROCARPUS ALEXANDRI 

A. MILNE-EDWARDS 
(CRUSTACEA: DECAPODA: PANDALIDAE) 

Fenner A. Chace, Jr. 

Abstract. — The ovigerous female holotype of Heterocarpus alexandri from 
off Havana, Cuba, is described and reillustrated. A male of presumably the 
same species is recorded from the Bahamas, and probably two related species 
are recorded and diagnosed from the Pacific region: H. nesisi Burukovsky from 
Baja California and an undescribed species previously recorded as H. alexandri 
from Hawaii. The systematic status of H. alexandri is discussed but not con- 
summated. 



This paper was prompted by the discov- 
ery in the collection of the Museum of Com- 
parative Zoology, Harvard University, of 
the holotype of Heterocarpus alexandri, 
which had been rather inadequately de- 
scribed and was feared lost. This opportu- 
nity was made especially significant by the 
revelation that A. Milne-Edwards' (1883) 
original illustration (there was no written 
description) was erroneous in several im- 
portant characters, thereby precluding as- 
surance of the identification of the species 
from other material. Although I have of- 
fered comparisons of H. alexandri with 
specimens of related species from the Pacific 
and have discussed the possible systematic 
status of those species, my objective in this 
contribution has been to record the typical 
features of//, alexandri for future reference, 
not to define its hierarchical position in the 
family. 

Family Pandalidae 

Genus Heterocarpus A. Milne-Edwards 

Heterocarpus alexandri A. Milne-Edwards, 

1883 

Fig. 1 

Heterocarpus Alexandri A. Milne-Edwards, 
1883 [unnumbered plate] [type locality: 
Blake dredging no. 2; north of Havana; 



23°14'00"N, 82°25 WW; 805 fms (1472 
m)].-DeMan, 1920:108, 153, 154. 
Heterocarpus alexandri. —Faxon, 1896: 
161. -Chace, 1985:19, 20, fig. 136. -Bu- 
rukovsky, 1986:64, 67, 69. 

Description of holotype.— Ovigerous fe- 
male with postorbital carapace length of 1 2.8 
mm. Rostrum (Fig. la) incomplete, existent 
portion nearly three-fourths as long as post- 
orbital carapace, overreaching antennal 
scale, with sharp lateral carina arising from 
orbital margin, armed dorsally with eight 
strong teeth, four on carapace posterior to 
orbital margin, posteriormost arising slight- 
ly anterior to midlength of carapace, bearing 
denticle on posterior slope, armed ventrally 
with four teeth. Carapace with middorsal 
carina extending over anterior two-thirds of 
length, posterior one-third smoothly round- 
ed; antennal spine flaring sinuously laterad 
from origin below orbit, slightly smaller than 
anteriorly directed branchiostegal spine, 
latter buttressed by submarginal carina ex- 
tending to near midlength of carapace; low, 
broad elevation, accentuated by rather deep 
dorsal and ventral depressions, extending 
posteriorly from orbital region and indis- 
tinctly joining somewhat more distinct su- 
prabranchial ridge reaching nearly to pos- 
terior margin of carapace. 



VOLUME 102, NUMBER 1 



85 




Fig. 1. Hetewcarpus alexandri, ovigerous female holotype: a, Carapace and anterior appendages, lateral 
aspect; b, Same, dorsal aspect; c, Abdomen, lateral aspect; d, third abdominal somite, dorsal aspect; e, Sixth 
abdominal somite, telson, and uropods, dorsal aspect; f, Juncture of sixth abdominal somite and telson, lateral 
aspect; g, Right second pereopod. (Magnifications: a, c, x3; b, d, e, g, *6;/ x 12.) 



Abdomen (Fig. lc) with tergum of first 
somite rather severely distorted, second so- 
mite with distinct, anteriorly convex, pos- 
teriorly setose transverse groove crossing 
tergum posterior to midlength, third somite 
with sharply defined middorsal boss on ter- 
gum (Fig. Id); sixth somite fully one and 
three-fourths times as long as fifth somite, 
slightly more than twice as long as high, 



dorsal surface shallowly convex and bluntly 
distinct from lateral surface (Fig. 1 e), armed 
posteroventrally with small but discrete 
tooth (Fig. 1/); pleura of four anterior so- 
mites with irregularly convex margins, pleu- 
ron of fifth somite with sharp posteroventral 
tooth. Telson (Fig. \e) deeply sulcate mid- 
dorsally on anterior half of length, armed 
with five pairs of dorsolateral spines, in- 



86 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



eluding pair lateral to long, intermediate 
posterior spines. 

Eye with subspherical cornea slightly 
broader than stalk, without ocellus (Fig. 1 b). 
Antennule with stylocerite narrowly taper- 
ing to sharp point near level of second third 
of second antennular segment. Antennal 
scale with lateral margin nearly straight, ter- 
minating in strong distolateral tooth not 
reaching level of distal oblique margin of 
blade. Third maxilliped overreaching an- 
tennal scale by length of terminal segment, 
exopod reaching to about midlength of an- 
tepenultimate segment. 

Pereopods with epipods on four anterior 
pairs. First pereopods missing or incom- 
plete. Right second pereopod (Fig. \g) over- 
reaching antennal scale by length of chela 
and about one-third of carpus, carpus com- 
posed of 18 articles; left second pereopod 
missing. All three pairs of posterior pereo- 
pods missing. 

Uropod (Fig. 1 e) with branches subequal 
in length, slightly overreaching telson, not 
including posterior spines. 

Eggs measuring about 0.4 x 0.6 mm. 

Remarks.— The unique holotype differs 
from A. Milne-Edwards' original illustra- 
tion in having the margin of the pleuron of 
the fourth abdominal somite rounded in- 
stead of posteroventrally acute, the telson 
armed with five rather than four pairs of 
dorsolateral spines, the blade of the anten- 
nal scale oblique rather than transversely 
truncate, and the carpus of the right second 
pereopod composed of 18 rather than 25 
articles. 

The specimen assigned to H. alexandri 
by Faxon (1896:161) from Blake dredge no. 
196; off Martinique; 1030 fms (1884 m) ap- 
parently has been temporarily misplaced in 
the MCZ collections, but there is a male of 
that species in the Smithsonian collections, 
taken by the Albatross at station 2629; 
mouth of Exuma Sound, Bahamas; 
23°48'40' , N, 75°10'40"W; 1169 fms (2138 
m); 8 Mar 1886. That specimen has a post- 
orbital carapace length of 1 1.0 mm; the an- 



terior part of the rostrum is missing beyond 
the seventh dorsal and the posterior ventral 
teeth. It differs otherwise from the holotype 
in having the submarginal carina that but- 
tresses the branchiostegal spine less sharp; 
the tergum of the first abdominal somite 
traversed by a distinct, anteriorly convex 
groove similar to the one on the second so- 
mite; the margins of the dorsal boss on the 
third somite curving laterad more strongly 
at the anterior end; the sixth somite nearly 
twice as long as the fifth and about two and 
one-half times as long as high; the telson 
armed with five pairs of dorsolateral spines 
on the left side but only four on the right, 
including the pair lateral to the intermediate 
posterior spines; the stylocerite not over- 
reaching the acute lateral projection of the 
first antennular segment; the blade of the 
antennal scale slightly narrower beyond the 
base of the distolateral spine than in the 
holotype (but not quite as distinctly so as 
indicated in Chace (1985:fig. 13&)); the first 
pereopod overreaching the antennal scale 
by about one-half the length of the chela; 
and the appendix interna on the second 
pleopod overreaching the appendix mas- 
culina by at least one-fourth of its length. 
In virtually all other characters, the male 
agrees with the ovigerous female holotype. 
The Hawaiian specimen recorded as H. 
alexandri by Rathbun (1906:918) from Al- 
batross station 4181; near Kauai Island; 811- 
671 fms (1483-1227 m) is an ovigerous fe- 
male with a postorbital carapace length of 
9.7 mm. It almost certainly represents a dis- 
tinct species. The rostrum reaches only about 
as far as the distal end of the antennular 
peduncle, does not reach the level of the 
distal end of the antennal scale, and has a 
rostral formula of 4 + 6/4; although the car- 
apace is very like that of H. alexandri, the 
transverse grooves on the first and second 
abdominal somites are much less distinct, 
that on the second being - virtually non- 
existent, and there is no suggestion of a boss 
on the tergum of the third somite; the sixth 
somite is two and one-fourth times as long 



VOLUME 102, NUMBER 1 



87 



as the fifth and fully two and one-half times 
as long as high; the telson is deeply sulcate 
anteriorly, but the posterior one-half is 
missing. 

Another Smithsonian specimen, identi- 
fied by Waldo L. Schmitt as H. alexandri, 
from "D. 5682," which almost certainly 
refers to an Albatross station by that num- 
ber in San Lucas Bay, Baja California; 22° 
48'20"N, 109°52'40' , W; 491 fms (898 m); 
24 Mar 1 9 1 1 , is a large female with a post- 
orbital carapace length of 31.5 mm; the 
rostrum far overreaches the antennal scale 
and has a rostral formula of 4 + 7/7; the 
middorsal postrostral carina reaches poster- 
iorly to about the posterior one-sixth of the 
postorbital carapace length, and the lateral 
postorbital ridge is very like the one in the 
holotype of H. alexandri, but the branchio- 
stegal spine is situated closer to the antennal 
spine and is not supported by a long, cari- 
nate buttress; the two anterior abdominal 
somites lack transverse grooves; the third 
abdominal somite is produced posteriorly 
in the midline, but it is without a dorsal 
boss on the tergum, except for a vestige sug- 
gested by a short, mesially convex furrow 
on each side of the midline; the sixth ab- 
dominal somite is somewhat transversely 
convex dorsally, nearly two and one-fourth 
times as long as the fifth, and one and two- 
thirds times as long as high; the telson is 
rather deeply sulcate dorsally, especially an- 
teriorly, armed with five pairs of dorsolat- 
eral spines, including the pair lateral to the 
intermediate posterior spines; the eyestalk 
widens distally to a nearly hemispherical 
cornea; the pereopods have well-developed 
epipods on the four anterior pairs; the first 
pair slightly overreaching the antennal scale; 
the right second pereopod reaches at least 
to the distal one-fourth of the antennal scale, 
the carpus is composed of five articles; the 
left second pereopod overreaches the an- 
tennal scale by the length of the chela and 
the distal carpal article, and the carpus com- 
posed of 15 articles; the third pereopod 
overreaching the antennal scale by the 



lengths of the dactyl and propodus, the dac- 
tyl bearing four spines on the flexor margin 
proximal to the distal tooth; the uropod with 
the lateral branch reaching to the level of 
the posterior margin of the telson, not in- 
cluding the posterior spines. 

It seems probable that the specimen from 
the southern tip of Baja California may be 
identifiable as H. nesisi Burukovsky, 1986, 
previously known only from the male ho- 
lotype, with an apparent postorbital cara- 
pace length of about 27 mm, found about 
1 600 kms southwest of the Albatross station 
at 13°34'N, 120°33'W, on an apparent sea- 
mount in 800 m in an area where total depths 
are near 4300 m. Although the Hawaiian 
specimen agrees with the one from Baja Cal- 
ifornia and disagrees with H. alexandri in 
the indistinct or nonexistent grooves on the 
first and second abdominal tergites and the 
absence of a dorsal boss on the third tergite, 
it differs so clearly from H. nesisi in its much 
smaller size, shorter rostrum, and much 
longer sixth abdominal somite as to suggest 
that it represents an undescribed species. 

There is little doubt that Crosnier (1 986a: 
368) had ample reason to remove H. laevis 
A. Milne-Edwards, 1883, from the genus 
Heterocarpus and to transfer it and its Pa- 
cific analogue Plesionika chacei Crosnier, 
1 986 (renamed P. fenneri Crosnier, 1 986b), 
to the genus Plesionika because of their sim- 
ilarity to P. bifurca Alcock & Anderson, 
1894, and P. spinidorsalis (Rathbun, 1906). 
Similarly, the three species discussed above 
probably belong to the same genus as the 
four assigned to Plesionika by Crosnier 
(1986a), because they lack strong postor- 
bital or postantennal carinae and have the 
carapace transversely convex rather than 
carinate in the dorsal midline of the pos- 
terior one-fourth or more of its length. I am 
not proposing that change here, however, 
because these seven species seem to form a 
rather homogeneous group quite different 
from Plesionika uniproducta Bate, 1888 
(=Acanthephyra ensis A. Milne-Edwards, 
1881), the type species of Plesionika. The 



88 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



family Pandalidae displays cosiderable 
morphological diversity, but some of the 
genera are so obscurely differentiated as to 
engender the possibility that a group of 
species clustered around Heterocarpus lae- 
vis and H. alexandri might eventually con- 
stitute a valid separate genus. The danger 
of prematurely accepting potential generic 
characters in the family, however, is man- 
ifested by the occurrence of a well-defined 
boss on the tergum of the third abdominal 
somite in those two Atlantic species and the 
virtually complete absence of such a boss 
in the Pacific species that are obviously 
closely related to them: Plesionika fenneri, 
Heterocarpus nesisi, and the Hawaiian spec- 
imen assigned to H. alexandri by Rathbun 
(1906). 

Acknowledgments 

But for the serendipity provided by Ardis 
Johnston of the Museum of Comparative 
Zoology in response to my inquiry about 
the specimen identified by Faxon (1896:161) 
as Heterocarpus alexandri, the holotype of 
that species would have remained "lost" in- 
definitely. I am most grateful to her for that 
welcome information and for making the 
type specimen available to me. I also thank 
Alain Crosnier of the Museum national 
d'Histoire naturelle in Paris and my Smith- 
sonian colleague Brian Kensley for reading 
the manuscript. 

Literature Cited 

Alcock, A., & A. R. Anderson. 1 894. Natural history 
notes from H. M. Indian Marine Survey Steam- 
er "Investigator," Commander C. F. Oldham, 
R.N., commanding, series 2(14); An account of 
a recent collection of deep sea Crustacea from 
the Bay of Bengal and Laccadive Sea.— Journal 
of the Asiatic Society of Bengal, new series, 63: 
141-185, pi. 9. 

Bate, C. S. 1888. Report of the Crustacea Macrura 
collected by H.M.S. Challenger during the years 
1 873-76. — Report on the scientific results of the 
voyage of H.M.S. Challenger during the years 
1873-76 24:i-xc, 1-942, figs. 1-76, 157 pi. 



Burukovsky, R. N. 1986. [A new shrimp species from 
the genus Heterocarpus (Crustacea: Decapoda: 
Pandalidae) and a brief review of the genus.] — 
Byulleten Moskovskogo Obshchestva Ispytate- 
ley Prirody Otdel Bioloogicheskiy 91(5):62— 73, 
figs. 1-4. [Russian with English summary.] 

Chace, F. A., Jr. 1985. The caridean shrimps (Crus- 
tacea: Decapoda) of \hz Albatross Philippine Ex- 
pedition, 1907-1910, part 3: Families Thalas- 
socarididae and Pandalidae. — Smithsonian 
Contributions to Zoology 41 1:1-143, 62 fig. 

Crosnier, A. 1986a. Crevettes de la famille des Pan- 
dalidae recoltees durant ces dernieres annees en 
Polynesie francaise. Description de Plesionika 
chacei et P. carsini spp. nov.— Bulletin du Mu- 
seum National d'Histoire Naturelle, ser. 4, 8, 
sect. A(2):36 1-377, figs. 1-4. 

. 1986b. Plesionika fenneri, nouveau nom pour 

Plesionika chacei Crosnier, 1986. — Bulletin du 
Museum National d'Histoire Naturelle, ser. 4, 
8, sect. A(3):691. 

de Man, J. G. 1920. The Decapoda of the Siboga 
Expedition, IV: Families Pasiphaeidae, Stylo- 
dactylidae, Hoplophoridae, Nematocarcinidae, 
Thalassocaridae, Pandalidae, Psalidopodidae, 
Gnathophyllidae, Processidae, Glyphocrango- 
nidae and Crangonidae. — Siboga-Expeditie 
Monographic 39a: 1-3 18, pi. 1-25. 

Faxon, W. 1896. Reports on the results of dredging, 
under the supervision of Alexander Agassiz, in 
the Gulf of Mexico and the Caribbean Sea, and 
on the east coast of the United States 1877 to 
1880, to the U.S. Coast Survey Steamer "Blake," 
Lieut-Commander C. D. Sigsbee, U.S.N., and 
Commander J. R. Bartlett, U.S.N., command- 
ing. 37. Supplementary notes on the Crusta- 
cea.— Bulletin of the Museum of Comparative 
Zoology at Harvard College 30(3): 153-1 66. 

Milne-Edwards, A. 1881. Description de quelques 
Crustaces Macroures provenant des grandes 
profondeurs de la mer des Antilles.— Annales 
des Science Naturelles, Zoologie, ser. 6, 11(4): 
1-16. 

. 1883. Recueil de figures de Crustaces nou- 

veaux ou peu connus, Paris, 3 pp, 44 pis. 

Rathbun, M. J. 1906. The Brachyura and Macrura 
of the Hawaiian Islands. —Bulletin of the United 
States Fish Commission (1903) 23(3):827-930 
[reprint with index pages i-viii], figs. 1-79, pi. 
1-24. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
102(1), 1989, pp. 89-94 

GAMMAROPSIS ARAWAKIA, A NEW SPECIES OF 

MARINE AMPHIPODA (CRUSTACEA) 

FROM JAMAICA 

James Darwin Thomas and J. L. Barnard 

Abstract.— One of the dominant amphipods in algal turf communities at 
Discovery Bay, Jamaica, is a new species, Gammaropsis arawakia, character- 
ized by long peduncle of uropod 3, small coxa 1 , 3-articulate accessory flagellum, 
extremely setose gnathopod 2, one pair of dorsal teeth on urosomite 1, pointed 
ocular lobe, and oblique upper margin of cephalic sinus for antenna 2. 



Gammaropsis arawakia is very abundant 
in the algal turf that now (1986) covers the 
heavily disturbed reef at Discovery Bay, Ja- 
maica. The living coral was damaged by the 
hurricane of 1980 and any regenerants al- 
most completely exterminated by algal turf 
in 1984-1986. This turf developed follow- 
ing the death of the herbivorous urchins 
Diadema antillarum (information from Di- 
rector Dr. Jeremy Woodley). A nearby load- 
ing dock for bauxite may also be a factor in 
the disturbance, although we are informed 
that the coral damage and turf development 
are common to most of the north coast of 
Jamaica. Discovery Bay is also character- 
ized by the presence of large, cold fresh- 
water seeps. Other amphipod genera pres- 
ent in this turf mat are Lembos, Elasmopus 
and Hyale. 

Isaeidae 

Gammaropsis arawakia, new species 

Figs. 1-3 

Etymology. —Named for the Arawak, 
former Indian tribe of the central Carib- 
bean. 

Diagnosis. —Ocular lobe projecting mod- 
erately, bearing sharp anterior cusp, lower 
antennal sinus with oblique dorsal margin; 
eyes of medium size, with irregular mass of 
purple pigment surrounded sparsely by clear 
ommatidia. Accessory flagellum 3-articu- 



late, articles 1-2 long, apical article minute. 
Epistome sharply produced into long an- 
terior spike. Mandibular palp article 3 cla- 
vate, with subdistal anterior notch, heavily 
setose. Inner plate of maxilla 1 naked, outer 
plate with 9 spines. Plates of maxilla 2 broad, 
inner with oblique facial row of 6 setae. 

Anterior coxae short, coxa 1 less than half 
as wide as coxa 2, rounded-quadrate an- 
teroventrally, coxa 3 narrower than coxa 2, 
coxa 5 as long as coxa 4. Carpus of gnatho- 
pod 1 longer than propodus, dactyl strongly 
overreaching oblique palm. Article 2 of 
gnathopod 2 densely setose mediodistally, 
carpus short, propodus greatly enlarged, 
palm oblique, convex, defined by tooth 
lacking spine, densely setose medially and 
anteriorly, these setae projecting dorsally and 
medially. Locking spines of pereopods 3-4 
absent, of pereopods 5-7 forming unequal 
pair. Pereopods 5-7 of similar structure but 
increasingly longer, article 2 very narrow 
and smooth posteriorly. 

Epimera 1-3 similar, each with small pos- 
teroventral tooth and setule-notch, poste- 
rior margins weakly convex. Urosomite 1 
with pair of weak dorsal teeth side by side. 
Uropod 1 with medium interramal tooth, 
lacking on uropod 2; each ramus of uropods 
1-2 with elongate apical spine. Peduncle of 
uropod 3 elongate, rami lanceolate but al- 
most rod-like, subequal to each other but 
longer than peduncle, outer ramus with 



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91 



elongate apical spine-setae about as long as 
peduncle of uropod 2. Telson ordinary. 

Description of male "a" 2.09 mm.— Left 
mandibular incisor with four teeth, lacinia 
mobilis with four teeth, main rakers three. 
Apex of palp on maxilla 1 with four spines, 
one apicolateral spine-seta, 3 subdistal fa- 
cial setae; inner plate short, softly triangu- 
lar. Inner plate of maxilliped with 3 medial 
setae, 7 apical setae, three stout apical spines, 
one ventral facial locking spine and one 
ventral seta; outer plate with four thick ser- 
rate medial spines, two similar but thinner 
apical spines and 2 apical setae, ventral sur- 
face with 1 1 scattered setae; dactyl elongate, 
nail of medium length, with three accessory 
setules (Figs. 1-3). 

Apices of rami on uropods 1-2 as in en- 
larged illustration, with medial elongate 
spine and short lateral spine, except inner 
ramus of uropod 2 with long and short spines 
reversed. 

Description of female "b" 2.47 mm.— 
Oostegites 2 and 5 short, slender, with 2 and 
6 setae each, oostegites 3 and 4 twice as long 
and wide, with 1 setae each; gills 2-3 and 
6-7 sac-like, slender, 3 times as long as 
broad, gills 4 and 5 about 4 times as long 
as broad, scarcely broader than others. 

Holotype. -USNM No. 1 95 1 75, male "a" 
2.09 mm. 

Type locality. —Jamaica station IB, Dis- 
covery Bay, backreef, 1 m, 6 Sep 1986, on 
Acropora palmata rubble mound near chan- 
nel marker, formaldehyde wash of algal turf, 
coll. J. D. Thomas and J. Clark. 

Material— Type locality, female "b" 2.47 
mm, female "c" 2.14 mm, male "d" 2.03 
mm and 15 other specimens. Numerous 



specimens from many other similar samples 
also deposited in Smithsonian crustacean 
collections. 

Relationship.— There are 105 species of 
Gammaropsis so that it was a laborious task 
to distinguish our species from the score of 
species that it resembles. To avoid a lengthy 
discussion we have eliminated those species 
not fitting significant parts in the following 
diagnosis: 

Ocular lobe pointed, short; eyes small; 
antennae of medium length; coxae 1-5 short 
and almost evenly extended; carpus of 
gnathopod 1 long; carpus of gnathopod 2 
short, propodus evenly rectangular, palm 
shorter than posterior margin of propodus, 
regular, poorly sculptured, defined, medial 
surface of hand heavily setose; epimeron 3 
with point; urosome with only one pair of 
dorsal teeth; peduncle and rami of uropod 
3 elongate, armaments of rami very sparse 
and simple. 

Gammaropsis (=Eurystheus, Kermys- 
theus, Podoceropsis, Megamphopus, Se- 
gamphopus, Pseudeurystheus, see Barnard 
1973) contains a diversity of species not yet 
organized cladistically. Note that Paranae- 
nia Chilton (1884) is being revived by Bar- 
nard & Karaman (1989). Gammaropsis ara- 
wakia resembles Segamphopus and 
Pseudeurystheus in the elongate carpus of 
gnathopod 1 and such species as G. exser- 
tipes and G. spinosa in the elongate peduncle 
of uropod 3. It also resembles several species 
with densely setose gnathopod 2 such as G. 
setifera, G. tonichi, and G. tawahi. The nar- 
row coxa 1 suggests certain species of Me- 
gamphopus but most members of that "ge- 
nus" have other anomalies in gnathopods 



Fig. 1. Gammaropsis arawakia, unattributed figures = holotype male "a" 2.09 mm; d = male "d" 2.03 mm. 
Capital letters denote main parts in the following list; lower case letters to right of capital letters or in body of 
figure indicate modifications as follows; lower case letters to left of capital letters indicate specimens described 
in captions: B, body; D, dactyl of thoracic leg; E, prebuccal lateral; G, gnathopod; H, head; M, mandible; N, 
mandibular molar; O, outer ramus; P, pereopod; R, uropod; S, maxilliped; T, telson; V, palp; W, pleon; X, 
maxilla; r, right; s, setae removed; t, left. 



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Fig. 2. Gammaropsis arawakia, holotype male "a" 2.09 mm. Letter codes see Fig. 1. 



VOLUME 102, NUMBER 1 



93 




Fig. 3. Gammaropsis arawakia, holotype male "a" 2.09 mm. Letter codes see Fig. 1. 



or other coxae. We have ignored species of 
"Podoceropsis" and " ' Kermystheus" which 
have severely reduced accessory flagella. 

Gammaropsis tawahi Barnard (1972), 
from New Zealand, has an elongate pedun- 
cle on uropod 3 and densely setose gnatho- 
pod 2 but the palm bears a tooth near the 
hinge, the apical armaments on uropods 1- 
3 are all short, mandibular palp article 3 
lacks a notch and the antennae are short- 
ened and sparsely setose. 

Our species is close to G. averus Reid 
(1951) from West Africa, but differs in the 
broader hand of gnathopod 2 and less sculp- 
tured palm. 

Our species differs: From G. sutherlandi 
Nelson (1981), North Carolina, in the small 
even coxae 1-5, strongly subchelate gnatho- 
pod 2 and simpler spination of uropod 3; 
From G. hirsuta Reid (1951), West Africa, 
in the more sculptured palm and long prop- 
odus of male gnathopod 2 and the longer 
carpus of gnathopod 1 ; From G. tonichi Bar- 
nard (1969), Pacific Mexico, in the presence 
of only one pair of teeth on the urosome, 



and the lack of radical transformations oc- 
curring in male gnathopod 2 and coxa 7; 
From G. spinosa Shoemaker (1942), Pacific 
Mexico, in the small coxae 1-5 and medium 
(versus very short) antennae; From G. chil- 
toni and G. haswelli Thomson (1897), New 
Zealand, in the sharp ocular lobes, small 
coxae 5-6 and the ordinary, not grotesque, 
male gnathopod 2; From the pan-tropical 
G. atlantica Stebbing (1888) in the normal 
and small eyes and longer carpus of gnatho- 
pod 1; From G. thomsoni Stebbing (1888), 
New Zealand, in the more simplified ar- 
maments on the more elongate rami of uro- 
pod 3; From G. exsertipes Stebbing (1888), 
Kerguelen, in the ordinary condition of cox- 
ae 1-5 (versus enlarged), male gnathopod 2 
(versus grotesque or elephantine), epimeron 
3, regular length of antennae and sharp ocu- 
lar lobes; From G. anomala Chevreux 
(1926), warm eastern Atlantic, in the nor- 
mal gnathopod 2 and pereopod 5 which in 
the Lusitanian species are respectively 
weakly palmate and elephantine. 

The following species also have many 



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similarities to ours but differ in two or more 
of the characters in the diagnosis presented 
just above: G. di I at at a and G. pseudoden- 
ticulata Ledoyer (1982) from Madagascar; 
G. pacifica and G. setifera Schellenberg 
(1938) from the middle Pacific Ocean; G. 
crenulata and G. ulrici Krapp-Schickel & 
Myers (1979) from the Mediterranean Sea; 
G. palmata (Stebbing & Robertson, 1891) 
from Europe; G. ellisi Conlan (1983) from 
cold-temperate eastern Pacific; and G. lo- 
bata Chevreux (1920) from Europe. 

Acknowledgments 

We thank Janice Clark for her assistance 
in the field in Jamaica and Kimberly R. 
Cleary for her assistance in our laboratories 
at the Smithsonian Institution. Dr. Jeremy 
Woodley, Director of the Discovery Bay 
Marine Laboratory, and his staff gave us 
considerable help in our survey studies. 

Literature Cited 

Barnard, J. L. 1969. A biological survey of Bahia de 
Los Angeles Gulf of California, Mexico, IV. 
Benthic Amphipoda (Crustacea). — Transac- 
tions of the San Diego Society of Natural His- 
tory 15:175-228. 

. 1972. The marine fauna of New Zealand: 

Algae-living littoral Gammaridea (Crustacea 
Amphipoda).— New Zealand Oceanographic 
Institute Memoir 62:1-216. 

. 1973. Revision of Corophiidae and related 

families (Amphipoda). — Smithsonian Contri- 
butions to Zoology 151:1-27. 

— , & G. S. Karaman. 1989. The families and 

genera of Gammaridean Amphipoda (except 
marine Gammaroids). — Smithsonian Contri- 
butions to Zoology (In press). 

Chevreux, E. 1920. Sur quelques amphipodes nou- 
veaux ou peu connus provenant des cotes de 
Bretagne. — Bulletin de la Societe Zoologique de 
France 45:75-87. 

. 1926. Amphipodes I.— Gammariens {suite). 

Voyage de la goelette Melita aux Canaries et au 
Senegal 1889-1 890. -Bulletin de la Societe 
Zoologique de France 50:365-398. 

Chilton, C. 1 884. Additions to the sessile-eyed Crus- 



tacea of New Zealand.— Transactions and Pro- 
ceedings of the New Zealand Institute 16:249- 
265. 

Conlan, K. 1983. The amphipod superfamily Coro- 
phioidea in the northeastern Pacific region 3. 
Family Isaeidae: Systematics and distributional 
ecology.— Publications in Natural Sciences, Na- 
tional Museum of Natural Sciences, Canada 4: 
1-75. 

Krapp-Schickel, G., & A. A. Myers. 1979. The Med- 
iterranean species of Gammawpsis Liljeborg 
(Crustacea, Amphipoda).— Bollettino del Mu- 
seo Civico di Storia Naturale, Verona 6:441- 
467. 

Ledoyer, M. 1982. Crustaces amphipodes gamma- 
riens families des Acanthonotozomatidae a 
Gammaridae.— Faune de Madagascar 59(1): 1- 
598. 

Nelson, W. 1981. A new species of the marine am- 
phipod genus Gammawpsis from the south- 
eastern United States (Photidae).— Proceedings 
of the Biological Society of Washington 93: 1223- 
1229, 3 figs. 

Reid, D. M. 1951. Report on the Amphipoda (Gam- 
maridea and Caprellidea) of the coast of tropical 
West Africa. -Atlantide Report 2:189-291. 

Schellenberg, A. 1938. Litorale Amphipoden des tro- 
pischen Pazifiks. — Kungliga Svenska Veten- 
skapsakademiens Handlingar (3) 16(6): 105 pp. 

Shoemaker, C. R. 1942. Amphipod crustaceans col- 
lected on the Presidential Cruise of 1938. — 
Smithsonian Miscellaneous Collections 101(11): 
1-52. 

Stebbing, T. R. R. 1888. Report on the Amphipoda 
collected by H. M.S. Challenger during the years 
1873-76.— Report on the Scientific Results of 
the Voyage of H.M.S. Challenger During the 
Years 1873-1876, Zoology 29:xxiv and 1737 
pp. 

, & D. Robertson. 1891. On four new British 

Amphipoda.— Transactions of the Zoological 
Society of London 13:31-42. 

Thomson, G. M. 1897. On two new gammarids from 
New Zealand.— Annals and Magazine of Nat- 
ural History, ser. 6, 20:447^451. 

(JDT) The Reef Foundation, P.O. Box 
170, Big Pine Key, Florida 33043; (JLB) 
Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, NHB- 
163, Smithsonian Institution, Washington, 
D.C. 20560. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 95-105 

NATATOLANA PASTOREI (GIAMBIAGI, 1925) 

(CRUSTACEA, ISOPODA, CIROLANIDAE) FROM THE 

STRAITS OF MAGELLAN, SOUTH AMERICA: 

REDESCRIPTION AND NOTES ON 

FUNCTIONAL MORPHOLOGY 

Johann Wolfgang Wagele and Niel L. Bruce 

Abstract. — Cirolanids caught with baited traps in the Straits of Magellan and 
identified as Natatolana past or ei (Giambiagi) are described. Observations on 
living specimens suggest that several features, such as the broadened merus of 
the anterior pereopods, the arrangement of spines and setae are adaptations to 
digging and respiration in sand. 



Giambiagi (1925) described briefly from 
"San Sebastian, Tierra del Fuega," south of 
the eastern entrance of the Straits of Ma- 
gellan, a cirolanid named Cirolana pastorei. 
No other figures than those published by 
Giambiagi exist; they are not adequate to 
distinguish this species from the many sim- 
ilar cirolanids known today. Another ciro- 
lanid from the Straits of Magellan was men- 
tioned by Pfeffer (1887) in a list of species 
collected during a German expedition to 
South Georgia (1882-1883). Of this Ciro- 
lana magellanica no illustrations or de- 
scriptions were ever published. The name 
was therefore listed as nomen nudum by 
Bruce (1986). 

During an expedition to Antarctica (1 985- 
1986) with R/V Polar stern a large number 
of cirolanids were caught in a baited trap in 
the Straits of Magellan. Most of these ani- 
mals were kept in aquaria for bionomic 
studies. Later the same species was found 
in collections of the Zoologisches Museum 
Hamburg, in vials labelled "Cirolana ma- 
gellanica, cf. types," which possibly contain 
the specimens studied by Pfeffer (1887). 
Further material was kindly sent to the first 
author by Dr. Gruner from the Zoologisches 
Museum Berlin, also with the label "Ciro- 
lana cf. magellanica. " 

The species clearly belongs to the genus 



Natatolana Bruce, 1981 and is related to a 
group of large species, the Natatolana valida 
group, that have an exclusively southern 
hemisphere distribution (Bruce 1986). The 
species was compared to drawings of the 
types of Natatolana rossi, N. hirtipes and all 
subantarctic species and found to be dis- 
tinct. The species here described is identical 
with the Cirolana pastorei of Giambiagi 
(1925); no differences from the original de- 
scription or the type specimens could be 
found. The species is not identical with the 
Cirolana pastorei from South Georgia men- 
tioned by Monod (1931). 



Natatolana pastorei (Giambiagi) 
Figs. 1-7 

Cirolana pastorei Giambiagi, 1925:4-7, pi. 
1, pi. 3, fig. 3, not Cirolana pastorei sensu 
Monod, 1931:22. 

Cirolana magellanica Pfeffer, 1887:58, no- 
men nudum.— Bruce, 1986:218. 

Material examined. —Male, length 33 
mm, partly dissected, Magellan Straits, 
52°17.49'S, 69°8.96'W, 35 m; 15 Nov 1984; 
four adult specimens, 27-36 mm, from the 
same locality, Zoological Museum Ham- 
burg, K-33324. Further material: Zoologi- 
cal Museum Hamburg, K- 192 19, Magellan 



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Fig. 1 . Natatolana pastor ei: A, Male in dorsal view; B, Ventral aspect of cephalothorax; C, Posterior segments, 
ventral. Arrow: respiratory current passing through filter of pereopods 5-7. Abbreviations.— Al, A2: antenna 1,2; 
La: labrum; Md: mandible; MdP: palp of Md; Mxl, Mx2: maxilla 1,2; Mxp: maxilliped; PI -7: pereopods 1- 
7; Plpl-5: pleopods 1-5; Tel: telson; Urp: uropod. 



Straits, labelled "Cirolana magellanica, cf. 
types," leg. Kpt. R. Paessler 1886: 44 spec- 
imens, 13-29 mm, one male of 30.5 mm, 
no mature females; also included: 5 speci- 



mens, 13-15 mm, leg. Prof. Carlos Grethe, 
Magellan Str., 19 Nov 1908; further mate- 
rial collected by Kpt. Paessler: K-23268 
(about 52°20'S, 69°W), K-23287 (51°54'S, 



VOLUME 102, NUMBER 1 



97 




Fig. 2. Natatolana pastorei, male in lateral view; arrows: respiratory current passing through setal combs of 
basipodites, antennule with detail of flagellar setation. For abbreviations see Fig. 1. 



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73°42'W), K-23281 (about 47°S, Golfo de 
Penas), K-23278 (Hale Cove, Patagonia); 
with number K-23242 two samples are de- 
posited: leg. Dr. Gassmann, Magellan Str., 
1 Nov 1895, and leg. Umbach, off Punta 
Arenas, Chile (53°10'S, 70°54'W), 1 Jan 
1903. 

Zoological Museum Berlin, Nr. 8118, 
Magellan Str., leg. Pohl: three small speci- 
mens (10-18 mm); one further specimen 
was sent to the Copenhagen Museum, prob- 
ably for H. J. Hansen. 

Museo Argentino de Ciencias Naturales 
"Bernardino Rivadavia," Buenos Aires, Nr. 
12690: nine syntypes, San Sebastian, Tierra 
del Fuego. 

Description. —Body about 2.5 times as 
long as broad. Cephalothorax and eyes as 
in Figs. 1 & 2; beneath eye a deep groove 
for antennae. Coxae one to three with curved 
carina running from frontal area of coxa to 
posterolateral point; coxae four to seven with 
more pronounced, less curved medial ca- 
rina; coxa seven reaching beyond epimeral 
plate of pleonite 1 (Fig. 2). Epimeral plate 
of pleonite 3 narrower than preceding plate, 
posteriorly acute. Outline of telson as in Fig. 
7, distal margin with six to seven sensory 
spines on each side and between them rows 
of four to five setae. 

Antennule peduncle of 4 articles (3 in the 
sense of Wagele 1983), article 3 longest (Fig. 
2); article 4 short, with three plumose setae; 
flagellum not much longer than peduncle, 
with 17 articles, articles short and broad, 
with rows of aesthetascs and each 2nd or 
3rd article with a group of short simple se- 
tae, last article with simple setae and one 
plumose seta (Fig. 2). Antenna peduncle ar- 
ticle 2 very short, article 5 longest; fourth 
article with lateral row of seven simple se- 
tae; flagellum long, extending to pereonite 
3, of about 30 articles; last articles more 
slender and relatively longer than basal ar- 
ticles (Fig. 3). 

Mandible palp setation as in Fig. 3. Lat- 
eral endite of maxillule gnathal surface with 
13 stout spines, medial endite short, with 



three stout apical teeth and subapically a 
smaller, slender trifurcate spine (Fig. 2). 
Maxilla of three lobes; proximal endite with 
two rows of setae, first row with 1 1 , second 
with 17 setae; medial (second) endite with 
15 long setae and a second row with 11 
shorter setae; distal lobe bearing 5 setae. 
Maxilliped endite with two coupling hooks; 
all palp articles bearing large numbers of 
simple setae (Fig. 4). 

Pereopod 1 merus posterior margin with 
14 acute spines; carpus posterodistal margin 
with 2 long and 2 short acute spines: pro- 
podal palm with 4 large acute and 5 small 
acute spines and stout spine opposite base 
of dactylus. Pereopods 2 and 3 similar to 1 
but spines of merus and carpus larger, pe- 
reopod propodal palm with 3 spines, is- 
chium posterodistal margin with additional 
spines. Pereopod 3 with only 2 small spines 
on propodal palm. Pereopods 4-7 essen- 
tially similar but basis widening and all ar- 
ticles increasing in length towards posterior. 
Pereopod 7 basis about 1.8 times as long as 
greatest width, posterior margin shallowly 
convex, anterior margin feebly sinuate; is- 
chium, merus and carpus with clusters of 
spines at distal angles and along posterior 
margins; propodus with three groups of 2- 
4 spines on posterior margin and a group 
of 4 spines opposite base of dactylus. 

Pleopods conform to the generic condi- 
tion; pleopod 2 of male with appendix mas- 
culina, apex narrowed, slightly angled, 
slightly extending beyond apex of endopod. 
Uropod endopod medial margin convex 
with six spines and a further apical spine, 
lateral margin with two large and two small 
spines, apex narrow. Exopod lanceolate, lat- 
eral margin with eight spines and a further 
small apical spine, medial with four. 

Variations of spine armature of tail fan 
in adult specimens (25-36 mm): Distal mar- 
gin of telson with 6 to 8 spines on each side 
(generally 6), lateral margin of uropod ex- 
opod with 9 to 11 (usually 10, including 
apical spine), medial margin always with 4 
(including apical spine of this side), lateral 



VOLUME 102, NUMBER 1 



99 




Fig. 3. Natatolana pastorei, male: Last articles of antenna and apical part of 5th peduncular article (inset, 
bottom) of immature adult; Right mandible (r. Md) with detail of setal lobe. For abbreviations see Fig. 1 . 



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P1 (detail) 



Fig. 4. Natatolana pastorei, male: Maxilliped with detail of endite; several setae and sensory spines cut off. 
For abbreviations see Fig. 1 . 



VOLUME 102, NUMBER 1 



101 




Fig. 5. Natatolana pastorei, male: Pereopods 2-5, several setae and sensory spines cut off. For abbreviations 
see Fig. 1. 



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i 1.25 mm , 



Fig. 6. Natatolana pastorei, male: Several setae and sensory spines cut off; setae of pleopods shown as simple 
setae; dashed line (Pip 2): range of setae. For abbreviations see Fig. 1. 



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103 




Fig. 7. Natatolana pastorei, male: Setae of pleopods shown as simple setae, dashed line indicating length of 
setae; detail of Pip 5 showing apex of endopod; detail of uropod showing a sensory spine. For abbreviations see 
Fig. 1. 



margin of endopod always with 4, medial 
with 4 to 7 spines (including apical spine). 

Color. —Purple brown, chromatophore 
pattern shown in Fig. 1. 

Size. —Immature adults 27-34 mm, males 
30.5-33 mm. 

Remarks. —Natatolana pastorei belongs 
to the TV. valida group of species character- 
ized by Bruce ( 1 986). It can be distinguished 



from all of those species listed by having a 
slightly wider basis to pereopod 7. narrow 
apices on the uropod rami, and also differ- 
ences in the spination of the uropod rami. 
Other large Natatolana species include those 
found in the subantarctic region such as N. 
meridionalis Hodgson, N. nitida Hale or N. 
obtusata Vanhoffen (these species were list- 
ed by Bruce, 1986). All of these species are 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



characterized by large penial processes, 
broad or elongate appendix masculina and 
acute upturned posterolateral margins on 
pleonite 4. 

The present material differs from the ma- 
terial discovered in South Georgia and men- 
tioned by Monod (1931) as "Cirolana pas- 
torei." An examination of Monod's material 
(collection of the Senckenberg Institute, 
Frankfurt) revealed the following differ- 
ences: Lateral margin of uropod endopod 
with six spines (instead of four in N. pas- 
torei); telson narrower; antennule shorter 
with only 12 flagellar articles; groove be- 
neath the eyes not present; eyes longer and 
less wide (number of ommatidia in N. pas- 
tor ei (17.5 mm): longitudinal line with 11- 
12, vertical line with 12; in the specimen 
from South Georgia: longitudinal line with 
14-15, vertical line with 8-9). So the pop- 
ulation from South Georgia must belong to 
a different species. 

Distribution. —Known from Straits of 
Magellan and between Hale Cove, Argen- 
tina to Punta Arenas, Chile, and from San 
Sebastian, Argentina. Occurs in shallow 
sublittoral sediments. 

Observations on biology and functional 
morphology. —About 20 specimens were 
observed in aquaria kept at 1-2°C together 
with other Antarctic isopods. At this low 
temperature the animals were inactive for 
many weeks, only sporadically feeding on 
pieces of krill and fish. All specimens bur- 
rowed in sand; when resting only the tips of 
the antennae and sometimes the eyes could 
be seen. In coarse sand burrowing is diffi- 
cult, as the grains keep rolling back into the 
groove, but in fine sand the specimens dis- 
appear within 1-2 min. The sand is pushed 
with the first pereopods laterally and cau- 
dally, the broad merus with its setae being 
a very effective broom; the posterior pe- 
reopods push the sand caudally, their effec- 
tiveness is increased due to the lateral spines 
(Fig. 1, bottom). The pleopods help when 
the animals dig fast; with their beat a strong 
current sweeps away the grains loosened by 



the pereopods. In a depth of 2-5 cm the 
animals stop digging. After feeding the an- 
imals can obviously remain inactive for 
about two months; in aquaria the animals 
were fed only once a week and mostly the 
food was not consumed. Though the low 
temperature affects the metabolism, in na- 
ture this species is probably also able to sur- 
vive several weeks without food, living as 
an opportunistic carrion feeder. 

Some isopods living in sand possess a 
protective operculum to cover the respira- 
tory pleopods (Chaetiliidae, Bathynatali- 
idae, Serolidae, Anthuridae). The cirolanid 
genera Conilera and Oncilorpheus also have 
such structures, but operculate pleopods are 
absent from other cirolanid genera. In Na- 
tatolana the setae of the pereopods are ar- 
ranged in such a way, that sand grains are 
kept away from the thin pleopods. While 
on the ventral side gravity will keep many 
grains in their place, the sides of the animal 
are protected against sand; the marginal se- 
tae of the pereopods cover the slits between 
the basipodites, where the respiratory cur- 
rent passes (Fig. 2). Directly in front of the 
pleopods, the setae of the frontal margins 
of the basis and ischium of P5-7 form a fan- 
like filter, to intercept grains and detritus. 

The pereopods are not used for swim- 
ming, animals swim with their dorsum 
downwards, propelled only by the beating 
pleopods. The broadened merus of PI -3, 
flattened basis with the marginal setae of 
P5-7, the arrangement of the sensory spines 
are adaptations to digging in fine sand. 

Acknowledgments 

We are grateful to Prof. G. Hartmann 
(Hamburg), Dr. M. Tiirkay (Frankfurt), and 
Prof. H. E. Gruner (Berlin, G.D.R.) for the 
loan of specimens. Mrs. S. Willig kindly fed 
and cultured the living animals. Mr. A. 
Pompecki helped to prepare the plates. 

This study was supported by the German 
Science Foundation (DFG Nr. Wa-530) to 
J.-W. Wagele, and a Queen's Fellowship in 
Marine Science to N. L. Bruce. 



VOLUME 102, NUMBER 1 



105 



Literature Cited 

Bruce, N. L. 1981. Cirolanidae (Crustacea: Isopoda) 
of Australia: Diagnoses of Cirolana Leach, Me- 
tacirolana Nierstrasz, Neocirolana Hale, An- 
opsilana Paulian & Deboutteville, and three new 
genera— Natatolana, Politolana and Carteto- 
lana. —Australian Journal of Marine and Fresh- 
water Research 32:945-966. 

. 1986. Cirolanidae (Crustacea: Isopoda) of 

Australia. — Records of the Australian Museum 
Supplement 6:1-240. 

Giambiagi, D. 1925. Resultados de la primera ex- 
pedition a Tierra del Fuego (1921). Crustaceos 
Isopodos.— Anales de la Sociedad Cientifica Ar- 
gentina, Buenos Aires 1925:229-245. 

Monod. T. 1931. Tanaidaces et isopodes subantarc- 



tiques de la collection Kohl-Larsen du Senck- 

enberg Museum.— Senckenbergiana 13:10-30. 
Pfeffer, G. 1887. Die Krebse von Siid-Georgien nach 

der Ausbeute der Deutschen Station 1882/83. 

I.— Jahrbuch der Hamburger Wissenschaftli- 

chen Anstalten 4:41-150. 
Wagele, J. W. 1983. On the homology of antennal 

articles in Isopoda. — Crustaceana 45:31-37. 

(JWW) Arbeitsgruppe Zoomorphologie, 
Fachbereich 7, Universitat Oldenburg, 
P.O.B. 2503, 2900 Oldenburg, FRG; (NLB) 
Australian Museum, Division of Inverte- 
brate Zoology, P.O.B. A285, Sydney South, 
N.S.W. 2000, Australia. 



PROC. BIOL. SOC. WASH. 
102(1), 1989, pp. 106-108 

NEOBRACHIELLA ANISOTREMI 

(COPEPODA: LERNAEOPODIDAE), A NEW SPECIES 

PARASITIC ON AN INSHORE FISH, 

ANISOTREMUS SCAPULARIS, OFF THE 

CHILEAN COAST 

Raul Castro Romero and Hernan Baeza Kuroki 

Abstract. —Neobrachiella anisotremi, parasitic on Anisotremus scapularis, an 
inshore fish of the northern coast of Chile, is described and illustrated. The 
new species can be distinguished from all the species in the genus by a com- 
bination of characters including second maxilla, trunk-cephalothorax length 
ratio, and specially the posterior processes. 



The existing literature contains very little 
information on the genus Neobrachiella Ka- 
bata, 1979, in Chilean waters. Only five 
species have been recorded. Kroyer (1863) 
gave a sketchy decription of Anchor ella 
(=Neobrachiella) appendiculata, without 
recording its host. Ho (1983) described Neo- 
brachiella amphipacifica, a species occur- 
ring in both the North and South Pacific. In 
the latter it was found on Psichrolutes sio 
Nelson, a deep water fish, in northern Chile. 
Castro & Baeza (1 986) added to the list Neo- 
brachiella exilis (Shiino, 1956) parasitic on 
Mugil cephalus L., 1758; N. chevreuxii (van 
Beneden, 1891), from Paralonchurus peru- 
anus (Steindachner, 1785) and N paralich- 
hyos, Castro & Baeza, 1986. Recently we 
decribed four other species collected from 
sciaenid hosts from the Chilean coast (Cas- 
tro & Baeza 1987). 

In the course of our studies of the parasitic 
copepods of Chilean fishes, we discovered 
another species of Neobrachiella, parasitic 
on an inshore fish, Anisotremus scapularis 
(Tschudi). This species is described and il- 
lustrated below. Terminology used is that 
proposed by Kabata (1979). 

Neobrachiella anisotremi, new species 
Figs. 1-7 

Material examined.— ¥ our ovigerous fe- 
males (8 Jan 1982); one female (12 Mar 



1982) and three females (14 Jul 1982). Types 
deposited in National Museum of Natural 
History (USNM), Smithsonian Institution. 
Holotype USNM 210517; Paratype USNM 
210158. 

Habitat.— Branchial arches of A. scapu- 
laris, collected by beach seining. 

Locality. — Isla Santa Maria, Antofagasta 
(23°27'S; 70°25'W). 

Description. —Female (Fig. 1). Cephalo- 
thorax subcylindrical, arching with slightly 
expanded cephalic region, mean length 2636 
l*m (range 2242-2908 ixm), width 455 /im 
(range 364-545 /im); dorsal shield well de- 
limited, subrectangular. Trunk much longer 
than wide, mean length 2909 jum (range 
2121-3636 Mm), width 1 136 /an (range 909- 
1333 ixm); its length varying from 75 to 1 23% 
of that of cephalothorax; posterior extrem- 
ity with two pairs of processes (Fig. 7 A, B); 
first pair ventral to egg sacs, mean length 
736 nm (range 606-805 jum); its length 20- 
38% that of trunk; second pair dorsal to egg 
sacs and shorter, mean length 570 jttm (range 
483-606); its length 73-83% that of first; 
genital process of varying length, mean 186 
^m (range 113-145 jum); poorly developed 
in some specimens. Egg sacs subcylindrical, 
multiseriate, mean length 2651 ^m (range 
1818-3515 /mi); their length 71-1 12% that 
of trunk. 

First antenna (Fig. 2A, B) apparently 4- 



VOLUME 102, NUMBER 1 



107 




Figs. l~4. Neobrachiella anisotremi: 1 , Female, lat- 
eral view; 2 A, First antenna; 2B, First antenna, detail 
of apical armament; 3A, Second antenna; 3B, Second 
antenna, detail of apical armament; 4, Mandible. 




Figs. 5-7. Neobrachiella anisotremi: 5, First max- 
illa, ventral; 6A, Maxilliped; 6B, Maxilliped, detail of 
myxal area and claw; 7A, Posterior end of trunk, fe- 
male; 7B, Lateral view of same, showing processes and 
egg sacs. 



segmented; first and second segments in- 
distinctly separated, second armed with 
spine on distoventral margin; third short, 
unarmed; fourth segment with apical ar- 
mament as follows: tubercles 1 and 3 well 
developed, 4a centrally situated, digitiform 
seta; 6a single slender seta, and 5a complex, 
consisting of a bifid seta and an additional 
simple process; gibber absent. Second an- 
tenna (Fig. 3A, B) strongly sclerotized, its 
long axis running through sympod and ex- 
opod; sympod bearing short robust spine on 
its distoventral border; exopod bulbous, 
apex spinulose on dorsal side; endopod 
2-segmented, distal segment with apical spi- 
niform process (reduced hook one, see Ka- 
bata 1979) and two setae (spinulation on 
ventral surface of distal segment can be seen 
on distended cuticle in Fig. 3B). Mandible 
(Fig. 4) with dental formula P1,P1,S1,P1, 
S1,B3. First maxilla (Fig. 5) with papilli- 



form exopod situated ventrally, bearing 
single seta; endopod armed with two sub- 
cylindrical, setiferous papillae, ventral 
papilla stouter than dorsal; no spinulation 
observed. Second maxilla (Fig. 1) fused only 
at tip, slender, less than half length of ceph- 
alothorax; mean length 105 1 fim. (range 909- 
1273 ixm); without conspicuous collars (bul- 
la with short manubrium and anchor of nar- 
row diameter). 

Maxilliped (Fig. 6A, B) corpus strong, 
slightly tapering, with spiniform process and 
denticulate pad in myxal area; patch of spi- 
nules near maxilla with subchela; shaft of 
subchela with barb and spinulose pad on 
distomedial part, claw gently curved, with 
two accessory teeth. 

Male. —Unknown. 

Etymology. —The specific name anisotre- 
mi, is derived from the generic name of the 
host. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Discussion.— To establish the identity of 
Neobrachiella anisotremi it must be com- 
pared with all the species of the genus pos- 
sessing two pairs of posterior processes. A 
comparison within this group must be based 
on a combination of gross morphological 
characters (cephalothorax length; trunk 
length; relationships between the length of 
posterior process and other parts of the 
body), since descriptions of the appendages 
are not available for all species. Using these 
criteria, the present species can be distin- 
guished from all of them by a combination 
of characters. It resembles most closely N. 
gulosa (Wilson, 1915) but can be differen- 
tiated from this species by the posterior pro- 
cesses. 

Wilson's species has ventral processes as 
long as the trunk, whereas in N. anisotremi 
they are only 20-38% of the trunk length. 
Slight differences exist also in the structure 
of the second maxilla, that of N. gulosa, in 
contrast to N. anisotremi, having a promi- 
nent swelling accommodating the excretory 
glands. 

Another difference can be found in the 
cephalothorax- trunk length ratio. In N. gu- 
losa the cephalothorax is considerably long- 
er than the trunk, whereas in anisotremi the 
trunk is longer than the cephalothorax. 

A unique feature of N. anisotremi is the 
presence of a short spine on the distolateral 
border of the sympod of the second antenna. 

The four recently described species, para- 
sitic on host fishes of the genus Sciaena (N. 
oralis, N. auriculata, N.fasciata, and N. dis- 
par), which are included in the same group 
as the present one, can be differentiated by 



a combination of characters that clearly dis- 
tinguish between each one and N. anisotre- 
mi. Their differences are given in the key to 
the species of Neobrachiella (Castro & Baeza 
1987). 

Acknowledgments 

The authors wish to thank Dr. Z. Kabata 
for his review of the manuscript and spe- 
cially for inviting the first author to spend 
some time at the Pacific Biological station, 
Nanaimo, B.C., Canada where this paper 
was completed. They are also grateful to the 
Universidad de Antofagasta for making the 
journey to Canada possible. 

Literature Cited 

Castro, R., & Baeza, H. 1986. Some species of Neo- 
brachiella Kabata, 1979 (Copepoda, Lernaeo- 
podidae), parasitic on Chilean fishes, with de- 
scription of Neobrachiella paralichthyos sp. nov. 
from Paralichthys adspersus (Steindachner). — 
Crustaceana 5 1(3): 24 5-25 3. 

, & . 1987. Four new species of Neo- 
brachiella (Copepoda: Lernaeopodidae), para- 
sitic on Sciaena genus (Teleostei: Sciaenidae) in 
the South Pacific— Estudios Oceanologicos 6(1): 
1-24. 

Ho, J. S. 1982. Copepod parasites of Psychrolutes 
(Pisces: Scorpaeniformes) from deep waters in 
the eastern Pacific— Parasitology 85:451-458. 

Kabata, Z. 1979. Parasitic copepoda of British fish- 
es.— The Ray Society, London, No. 152:1-469. 

Kroyer, H. 1863. Bidrag til Kundskab om snyltek- 
rebsene.— Naturhistorisk Tiddsskrift, ser. 3(2): 
75-426, pis. 1-18. 

Universidad de Antofagasta, Instituto de 
Investigaciones Oceanologicas, Casilla 170, 
Antofagasta, Chile. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 109-115 

THE FIRST ANNUAL RISER LECTURE: 

MEIOFAUNA AND MICROBES -THE INTERACTIVE 

RELATIONS OF ANNELID HOSTS WITH THEIR 

SYMBIOTIC BACTERIA 

Olav Giere 

Abstract.— The gutless marine tubificid Phallodrilus leukodermatus (Oligo- 
chaeta) harbors two kinds of chemoautotrophic sulfur bacteria beneath its 
cuticle. The annelid hosts live predominantly around the redox potential dis- 
continuity layer in calcareous sediments, providing their bacterial endosym- 
bionts with access to both oxygen and reduced sulfur compounds. The bacteria 
produce energy by oxidation of these sulfur compounds, and use this energy 
for the fixation of C0 2 and production of carbohydrates. The worms utilize 
the exudates of the bacteria and also ingest a portion of the bacterial population 
in the epidermal cytoplasm. Further interactions in this apparently obligate 
symbiosis are discussed. The transmittance of the bacteria through the host 
generations by "infection" of freshly deposited eggs is correlated with a modified 
reproductive biology. The studies on this symbiosis reveal a well-coordinated 
balance between the annelid hosts and their extracellular bacterial partners. 



Serendipity let us find a new species of 
small (2-3 cm long) white annelid worms 
in the calcareous sands of Bermudian reefs. 
These tubificid oligochaetes, described as 
Phallodrilus leukodermatus (Giere, 1979), 
were unique among known annelids. They 
lacked mouth, gut and anus and had no ex- 
cretory organs. But the most striking feature 
was a thick, subcuticular 'mantle' of extra- 
cellular bacteria living between extensions 
of the epidermal cells (Fig. 1). 

This bacterial association, present in each 
worm, even in the youngest, apparently 



The Riser Lecture Series. — In 1985 the annual Riser 
Lecture was initiated by members, alumni and friends 
of the Marine Science Center, Northeastern University 
at Nahant, Massachusetts. The occasion was the official 
retirement of Professor Nathan W. Riser. As teacher, 
biologist and founder of the facility, "Pete" Riser en- 
dowed the laboratory with a legacy— the importance 
of considering the whole organism regardless of one's 
special focus. We dedicate these annual lectures to that 
principle. 



consists of two different gram -negative pro- 
karyotes (Fig. 2), a smaller, rod-shaped form 
(maximally 1.9 x 0.4 nm) with a thick cell 
envelope, and a larger oval one (about 3 x 
2 jum) characterized by a thinner body wall 
and, more conspicuously, by the rich supply 
of vesicles in its cytoplasm (Giere et al. 
1984). 

This peculiar bacterial layer raises ques- 
tions about the nutrition of these worms and 
about the structural implications of com- 
plete reduction of the gut in worms from 
sands that otherwise are richly populated by 
other meiofauna, including oligochaetes with 
normal guts. Also the microdistribution of 
these exceptional worms is quite unusual 
(Giere et al. 1982). We found rich popula- 
tions in an area where, despite strong cur- 
rents, anoxic horizons occurred at 5 cm sed- 
iment depth. Only a few worms were present 
in the upper, oxygenated strata, while the 
bulk of the population always occurred in 
the deeper layers where they were exposed 
to considerable amounts of toxic hydrogen 



110 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Kr .'-csrm m 



VOLUME 102, NUMBER 1 



111 



sulfide (around 100 /jM S = ) with oxygen 
present only in minimal concentrations. 

What promotes this exceptional distri- 
bution? How are the bacteria involved in 
this scenario? The answers require integra- 
tion of morphological studies with ecolog- 
ical, physiological and microbiological 
analyses, i.e., consideration of the whole 
animal's biology. These studies were per- 
formed by a multidisciplinary team during 
several research periods at the Bermuda Bi- 
ological Station for Research and in mem- 
bers' respective home laboratories. 

Structural and ecological similarities sug- 
gest that the symbiotic prokaryotes are sul- 
fur bacteria similar to those found in the 
giant vestimentiferan Riftia (Cavanaugh et 
al. 1981) from hydrothermal vents. The 
bacteria obtain energy by oxidation of re- 
duced-sulfur compounds, enabling them to 
fix ambient C0 2 with subsequent produc- 
tion of carbohydrates. 

For Phallodrilus leukodermatus, confir- 
mation of this pathway results from phys- 
iological analyses: Ribulosebiphosphate- 
carboxylase in high activities in the bacteria 
enables a notable incorporation of C0 2 with 
a most rapid transfer into the malate pool 
(Felbeck et al. 1983). The high activity of 
enzymes like ATP-sulfurylase and sulfite 
oxidase, involved in the sulfur metabolism, 
indicates more specifically that sulfur com- 
pounds become metabolized by the sym- 
biotic bacteria. Moreover, it can be shown 
that the products of this metabolism get 
stored in the numerous bacterial vesicles: 



Energy-dispersive X-ray microanalysis 
demonstrates high sulfur peaks in mem- 
brane-bound bacterial vesicles, suggesting 
deposition of polysulfanes or polythionates 
as easily available reserve products (Steudel 
1985, Steudel et al. 1987). 

Sulfur also is found in the internal tissues 
of the worms where it accumulates in the 
"chloragosomes," annelid structures known 
to store xenobiotic substances. Thus, sulfur 
is transported from the ambient pore water 
(perhaps via the symbiotic bacteria?) to the 
internal worm tissues. Sulfate, the end prod- 
uct of oxidative sulfur metabolism, is ex- 
creted by the worms into the ambient sea 
water (Giere et al. 1988). 

Bacteria oxidizing sulfur compounds are 
micro-aerophilic, gram-negative microor- 
ganisms thriving in those horizons where 
they have access to both reduced sulfur and 
oxygen, i.e., around the redox potential dis- 
continuity layer. Microdistribution of the 
worm population presents ecological con- 
firmation of these sulfur-based chemoau- 
totrophic processes in the annelid sym- 
bionts. The bulk of the worm population is 
always concentrated around this transition- 
al zone between the oxygenated surface lay- 
er and the anoxic depth with its poisonous 
hydrogen sulfide. In this zone the prokary- 
otic symbionts can synthesize their reserve 
substances. In addition to reduced- sulfur 
compounds, the bacterial cytoplasm 
abounds with unwalled globules (Fig. 2) filled 
with polyhydroxybutyric acid. Thus, in in- 
cident light worms from this zone have well- 



Figs. 1-3. Phallodrilus leukodermatus: 1, Cross section through the gutless worm with "mantle" of subcu- 
ticular bacteria, scale bar = 50 jwm; chl = chloragocytes, coe = coelomic cavity, cu = cuticle, ep = epidermis, 
mu = musculature, ne = ventral nerve cord, scu = subcuticular space filled with bacteria; 2, The two kinds of 
bacteria endosymbionts; note difference in thickness of cell envelope, scale bar = 0.5 fim; lba = large bacterium, 
phb = vesicle filled with polyhydroxybutyric acid, sba = small bacterium, sve = sulfur vesicle; 3, Typical zonation 
pattern of subcuticular bacteria with free, active bacteria in peripheral position and enclosed, lytic bacteria in 
innermost position, scale bar = 5 nm; blys = bacteria in lysis, zba = zone of (free) bacteria, zint = intermediate 
zone bacteria become enclosed by epidermal strands, zlys = zone of bacterial lysis: bacteria are enclosed in 
vacuoles of epidermal cells and become digested (Fig. 1: light microscopy; Figs. 2 + 3: transmission electron 
microscopy). 



112 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



fed bacteria and appear shiny white due to 
the rich content of light-refractile sub- 
stances in the bacterial cytoplasm, which 
can be almost 6% of the worm's dry weight 
(Giere et al. 1988). When kept in pure, ox- 
ygenated seawater, the worms (through their 
bacteria) continue to take up C0 2 for a while 
from the environment, using up energy 
stored in the reserve substances. However, 
this uptake decreases soon, the worms be- 
come (after about 24 h) transparent pale- 
grayish, indicating rapid loss of their re- 
serves, which in these "pale worms" is only 
1.7% of their dry weight (Giere et al. 1988). 

These results emphasize the biological in- 
teractions between the oligochaete host and 
its prokaryotic symbionts and lead to ques- 
tions for the biological function of the bac- 
teria/host system. The presence of bacteria 
in all members of the population and the 
ubiquity of bacterial incorporation in gut- 
less oligochaete species — several species 
mainly from the Caribbean, but also from 
Australian reefs have been examined— sug- 
gests an obligate symbiosis with mutual 
benefits. 

What are the benefits to the microorgan- 
isms of this incorporation into the worms' 
bodies? 

-In a changing sedimentary system with 
steep, overlapping gradients of oxygen, sul- 
fide, thiosulfate and CO z , the highly motile 
worms are able to quickly adjust position 
relative to environmental fluctuations, 
maintaining an optimal combination of 
gradients required for growth of sulfur-ox- 
idizing bacteria. 

-It is conceivable that the prokaryotes take 
up organic substances excreted by their host 
which, in turn, has lost its excretory or- 
gans. The highly vesiculate epidermal cy- 
toplasm of the worms and the multitude 
of membrane-bound secretory granules 
found extracellularly in their subcuticular 
lumen suggest this physiological pathway. 

-The favorable position of the bacteria un- 
derneath the cuticle affords both shelter 
and easy access to nutrients from the am- 



bient pore water through the highly perme- 
able cuticle. 

On the other hand, what benefits might 
the oligochaetes derive from close associa- 
tion with bacteria? 

-Since the bacterial oxidation of reduced 
sulfur combines production of energy with 
detoxification of poisonous hydrogen sul- 
fide, the microorganisms enhance the 
chances of the worms to invade a biotope 
of high ecological potential, rich in dis- 
solved nutrients, but with little competi- 
tion or predation. Our physiological ex- 
periments demonstrate that the worms can 
actively and selectively take up dissolved 
hexoses (particularly glucose and galac- 
tose) and acidic free amino acids (partic- 
ularly glutamic acid and aspartate). These 
purely transcuticular absorption processes 
seem so intensive that, compared to cor- 
responding figures from pogonophorans of 
the same size, they "contribute signifi- 
cantly" to the nutrition of P. leukoder- 
matus (Liebezeit et al. 1983, Liebezeit, pers. 
comm.). 

-Exuded bacterial products could serve as 
a valuable and ever-present source of fur- 
ther dissolved organic matter. 

-Digestion of a certain portion of the inter- 
nal bacterial population would provide an 
additional trophic pathway. Comparison 
of bacteria in the deeper layer of the epi- 
dermis with those in the subcuticular areas 
reveals a zonation pattern (Fig. 3): Periph- 
eral bacteria are extracellular in a subcu- 
ticular lumen, where they accumulate re- 
serve substances and multiply intensively 
by division. In the middle zone they tend 
to be surrounded by strands of the epider- 
mal cells, the innermost bacteria are al- 
ways found enclosed in intracellular vac- 
uoles, apparently "engulfed" by the 
epidermal cells, and clearly undergoing lysis 
(Giere and Langheld 1987). 

This coordinated system, combining bac- 
terial growth and multiplication with lysis 
and digestion, is similar to those described 



VOLUME 102, NUMBER 1 



113 







Figs. 4-5. Phallodrilus leukodermatus: 4, Cross section through genital pad in clitellar region, scale bar = 
50 Mm; cl = clitellar glands in epidermis, gepa = genital pad filled with bacteria, ne = ventral nerve cord; 5, 
Cross section through young embryo with some free subcuticular bacteria (both kinds) and many lytic bacteria 
enclosed in host cells, scale bar = 1 yum; balys = bacteria enclosed in embryonic cells, mostly in lysis, fba = free 
bacteria in peripheral layer (Fig. 4: light microscopy; Fig. 5: transmission electron microscopy). 



for some thyasirid bivalves (Southward 
1986), for the vestimentiferans Riftia 
pachyptila (Bosch & Grasse 1984a, b) and 
Tevnia sp. (Bassaglia 1986) and is compa- 
rable to the processes found in the root nod- 
ules of Leguminosae (Sprent 1979). 

Thus, it seems that by their symbiosis 
with sulfur-oxidizing bacteria the worms 
have acquired a tripartite nutritional basis 
via transepidermal uptake of dissolved or- 
ganic matter, utilization of bacterial meta- 
bolic products, and digestion of bacterial 
cells. This flexible trophic pattern seems to 
allow for the complete loss of digestive and 
excretory organs. Moreover, the worms 
gained access to a new habitat, the sulfidic 
sediments. Examination of four additional 
species within the approximately 40 species 
of gutless marine tubificids showed that in 
each case associations with subcuticular 
bacteria are established, suggesting that this 
symbiosis is geographically as ubiquitous as 
it is evolutionarily favorable. 



An intricate, mutualistic and well-estab- 
lished symbiosis as seen in P. leukoder- 
matus might be predicted to have evolved 
a fairly secure method of transmitting the 
bacterial partners through the generations 
of their hosts rather than depending on more 
or less randomly repeated de-novo-infec- 
tions of the worms. A regular transmission 
would require specific adaptative trends in 
the reproductive biology of these oligo- 
chaetes (Giere & Langheld 1987). 

In fact, investigation of fully mature 
worms reveals formation of a ventral "gen- 
ital pad" (Fig. 4), a blister-like structure 
packed with masses of fully active bacteria 
ensheathed only by a thin cuticular enve- 
lope. In a few cases deposition of single eggs 
was observed. They are covered by a sticky 
mucus, but not enclosed in a cocoon as 
would be typical for oligochaetes. These eggs 
were already infected by the characteristic 
bacteria. These first bacteria are mostly in- 
tracellular, within the cleavage cells, and 



114 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



clearly in a lytic stage (Fig. 5). Only a few 
peripheral bacteria maintain their subcutic- 
ular, extracellular position and have a 
healthy appearance. Probably the latter stock 
is the source of the rich supply of extracel- 
lular, active prokaryotes in adult worms. In 
fortuitous sections of freshly laid eggs, still 
adhering to the body, the bacteria appear to 
be just about to enter the egg membrane 
through deep invaginations. Since we found 
free bacteria only during egg deposition and 
the earliest developmental stages, we con- 
clude that prokaryotic symbionts are re- 
leased by rupture or, more probably, by lysis 
of the genital pad adjacent to the oviducal 
pore. Subsequently, they are transmitted by 
invasion of the freshly deposited egg. 

Although some microbiological and zoo- 
logical details are not yet fully understood, 
we envisage the following scenario: While 
in early stages a defense mechanism initially 
causes intracellular lysis of the bacterial in- 
truders, a balance between production of 
extracellular bacteria and their intracellular 
digestion becomes established in the grow- 
ing worms, resulting in a coordinated sym- 
biotic relationship between these aberrant 
marine tubificids and chemoautotrophic 
sulfur bacteria. 



Acknowledgments 

As pointed out, this lecture is based on a 
series of studies performed by a group of 
colleagues from various institutes and dis- 
ciplines. I owe sincere thanks for intensive 
collaboration and stimulating discussions 
to R. Dawson, now Chesapeake Biological 
Laboratory, University of Maryland, Sol- 
omons, Maryland, USA; H. Felbeck, Scripps 
Institution of Oceanography, La Jolla, Cal- 
ifornia, USA; H. Jannasch and C. Wirsen, 
both Woods Hole Oceanographic Institu- 
tion, Woods Hole, Massachusetts, USA; C. 
Langheld, G. Liebezeit and C. Schmidt, all 
University of Hamburg, Hamburg, FRG. 



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Bassaglia, Y. 1986. Le trophosome d'un Vestimen- 
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cenologie Biologique Paris, 42 pp. + 18 pp. of 
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Bosch, C, & P.-P. Grasse. 1984a. Cycle partiel des 
bacteries chimiautotrophes symbiotiques et leurs 
rapports avec les bacteriocytes chez Riftia 
pachyptila Jones (Pogonophore Vestimentifere). 
I. Le trophosome et les bacteriocytes.— Comptes 
Rendus de l'Academie des Sciences, Paris 299 
(ser. 3):37 1-376. 

. 1984b. Cycle partiel des bacteries chimiau- 
totrophes symbiotiques et leurs rapports avec 
les bacteriocytes chez Riftia pachyptila (Pogono- 
phore Vestimentifere). II. L'evolution des bac- 
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Comptes Rendus de l'Academie des Sciences, 
Paris 299 (ser. 3):413-419. 

Cavanaugh, C. M., S. L. Gardiner, M. L. Jones, H. W. 
Jannasch, & J. B. Waterbury. 1981. Prokary- 
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Riftia pachyptila Jones: Possible chemoauto- 
trophic symbionts.— Science 213 (4505):340- 
342. 

Felbeck, H., G. Liebezeit, R. Dawson, & O. Giere. 
1983. C0 2 fixation in tissues of marine oligo- 
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planus) containing symbiotic, chemoautotro- 
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Giere, O. 1979. Studies on marine Oligochaeta from 
Bermuda, with emphasis on new Phallodrilus- 
species (Tubificidae).— Cahiers de Biologie Ma- 
rine 20:301-314. 

; H. Felbeck, R. Dawson, & G. Liebezeit. 1 984. 

The gutless oligochaete Phallodrilus leukoder- 
matus Giere, a tubificid of structural, ecological 
and physiological relevance. — Hydrobiologia 
115:83-89. 

, & C. Langheld. 1987. Structural organisation, 

transfer and biological fate of endosymbiotic 
bacteria in gutless oligochaetes. — Marine Biol- 
ogy 93:641-650. 

, G. Liebezeit, & R. Dawson. 1982. Habitat 

conditions and distribution pattern of the gut- 
less oligochaete Phallodrilus leukodermatus.— 
Marine Ecology Progress Series 8:291-299. 

, C. O. Wirsen, C. Schmidt, & H. W. Jannasch. 

1988. Contrasting effects of sulfide and thio- 
sulfate on symbiotic C0 2 -assimilation of Phal- 
lodrilus leukodermatus (Annelida). —Marine Bi- 
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Liebezeit, G., H. Felbeck, R. Dawson, & O. Giere. 
1983. Transepidermal uptake of dissolved car- 



VOLUME 102, NUMBER 1 



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bohydrates by the gutless marine oligochaete 
Phallodrilus leukodermatus (Annelida).— 
Oceanis 9:205-211. 

Southward, E. C. 1986. Gill symbionts in thyasirid 
and other bivalve molluscs. —Journal of the Ma- 
rine Biological Association of the United King- 
dom 66:889-914. 

Sprent, J. I. 1979. The biology of nitrogen-fixing or- 
ganisms. McGraw-Hill Book Company, Lon- 
don, 196 pp. 

Steudel, R. 1 985. Neue Entwicklungen in der Chemie 
des Schwefels und des Selens. — Nova Acta Leo- 
poldina, N.F. 59, No. 264:231-246. 



,G.Holdt,T.Gobel,&W.Hazeu. 1987. Chro- 

matographische Trennung hoherer Polythionate 

S n 6 2_ (n = 3 22) und deren Nachweis in 

Kulturen von Thiobacillusferwoxidans; molek- 
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lausscheidungen.— Angewandte Chemie 99: 143- 
146. 

Zoologisches Institut und Zoologisches 
Museum, Universitat Hamburg, Martin- 
Luther-King-Platz 3, D-2000 Hamburg 13, 
Federal Republic of Germany. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 116-123 

REVISION OF THE FAMILY LYSARETIDAE, AND 

RECOGNITION OF THE FAMILY 

OENONIDAE KINBERG, 1865 

(EUNICIDA: POLYCHAETA) 

G. Kent Colbath 

Abstract. —As recognized by previous authors, the family Lysaretidae Kin- 
berg, 1865 includes genera with dissimilar jaw morphology and composition. 
Lysaretidae is restricted here to include only Lysarete Kinberg, characterized 
by jaws mineralized with calcite and relatively short carriers with transverse 
musculature. Oenone Savigny, Halla Costa and Tainokia Knox & Green are 
removed into the family Oenonidae Kinberg, 1865. These genera are charac- 
terized by heavily sclerotized jaws which are not mineralized, and by long 
carriers with oblique-longitudinal musculature. 



Hartman (1944) published a family-level 
revision of the eunicoid polychaetes which 
was widely accepted. Drawing on the work 
of Kinberg (1865) and Ehlers (1868), she 
placed heavy emphasis on the configuration 
of the jaws as a taxonomic character. 

Hartman recognized three basic maxil- 
lary jaw patterns within the superfamily Eu- 
nicea: the labidognath pattern ascribed to 
the Eunicidae, Onuphidae, and Lumbriner- 
idae; the prionognath pattern in the 
Arabellidae and Lysaretidae; and a third 
pattern (later named ctenognath by Kielan- 
Jaworowska 1966) found only in the Dor- 
villeidae. Labidognath jaws are character- 
ized by having two relatively short, broad 
carriers, in contrast to the two long, slender 
carriers with a ventral median ligament (or 
"third carrier") which characterize priono- 
gnath jaw apparatuses. In the ctenognath 
condition the maxillae consist of two to four 
rows of numerous separate denticles. 

During a study of the composition and 
fossilization potential of poly chaete jaws, I 
divided eunicoid jaws into three groups 
based on their composition (Colbath 1986). 
Labidognath jaws are mineralized with cal- 
cium carbonate in the form of aragonite in 
the Onuphidae and Eunicidae, and calcite 
in the Lumbrineridae. Labidognath jaws are 



also weakly sclerotized, and the outer pro- 
tein wall is readily leached in 2% KOH 
(Hartmann-Schroder 1967, Colbath 1986). 
In contrast, prionognath and ctenognath 
jaws are amorphous to x-rays, do not react 
with HC1, and are relatively resistant to 
leaching in KOH (Colbath 1986). 

The jaws of species of Lysarete Kinberg, 
1865 are mineralized with calcite (Colbath 
1986), and exhibit other labidognath mor- 
phological characters. Other genera previ- 
ously included within the Lysaretidae have 
jaws of the prionognath type, prompting a 
revision of the family, and recognition of 
the family Oenonidae Kinberg, 1865. 

History of Study 

Early classifications of the eunicoid poly- 
chaetes were based exclusively on features 
of the external anatomy, and produced ar- 
bitrary groupings of genera which are dis- 
similar internally (Hartman 1944:1). First 
to emphasize the importance of jaw parts 
in classification was Kinberg (1865). Kin- 
berg recognized ten families which he di- 
vided into four major groups (labeled in- 
formally as A-D). He defined group C by 
the presence of mandibles with separated 
right and left halves (an ontogenetically 
variable feature), and included only La- 



VOLUME 102, NUMBER 1 



117 



randa Kinberg, 1865, which was considered 
unrecognizable by Hartman (1 944: 10, 1948: 
98). Kinberg's other groups correspond to 
the labidognath (A), prionognath (B), and 
ctenognath (D) jaw types presently recog- 
nized. Kinberg considered the Lysaretae (sic) 
and Oenonidea (sic) distinct, including the 
former in group A and the latter in group B. 

Ehlers (1868:280-282) also emphasized 
the importance of jaw configuration in clas- 
sifying these polychaetes, but did not for- 
malize the distinctions. Instead, he recog- 
nized a single family Eunicea (sic), which 
he informally divided into the Eunicea lab- 
idognatha and Eunicea prionognatha. Eh- 
lers (1868) did not stress the morphology of 
maxillary carriers as strongly as did Kin- 
berg, but rather subdivided the eunicoids 
according to whether the anterior maxillae 
are arranged in a semicircle (labidognatha) 
or in parallel rows (prionognatha). 

In contrast to Kinberg, Ehlers (1868) 
placed the genus Lysarete close to Aglau- 
rides Ehlers (=Oenone Savigny) in the 
prionognath subdivision. He later (Ehlers 
1887:107-108) acknowledged the close 
similarity between the jaws of Lysarete and 
those of Lumbriconereis Grube (=Lumbri- 
neris Blainville), but did not emend his ear- 
lier taxonomic treatment. 

Hartman (1944:2) essentially formalized 
the 1868 scheme of Ehlers using the family- 
level nomenclature of Kinberg. She retained 
six of the ten families recognized by Kin- 
berg, placing the Oenonidae into synonymy 
with the Lysaretidae. 

Fauchald (1970:118) removed Iphitime 
Marenzeller from the Lysaretidae, and 
erected the family Iphitimidae. He later 
(Fauchald 1977) erected the order Eunicida 
to include the seven families within the su- 
perfamily Eunicea, plus two additional fam- 
ilies (Histriobdellidae, Ichthyotomidae) 
which were not assigned to a superfamily. 

Materials and Methods 

All lysaretids in the collection of the U.S. 
Museum of Natural History (USNM) were 



examined for the present study, and selected 
specimens were dissected for examination 
of the jaws. Additional specimens of Ly- 
sarete were obtained on loan from the Flor- 
ida Department of Natural Resources 
(FSBC). The holotype of L. brasiliensis Kin- 
berg, 1865 was obtained from the Swedish 
National Museum, Stockholm. 

One specimen of Lysarete brasiliensis with 
jaws in excellent condition (USNM 55884) 
was used for the chemical and mineralogical 
analyses, and for scanning electron micros- 
copy. The maxillary apparatus was re- 
moved, dehydrated in an alcohol series, and 
prepared for SEM by critical point drying. 
The specimen was initially coated with car- 
bon only, and chemically analyzed using 
x-ray dispersion. The specimen was then 
coated with gold palladium for routine scan- 
ning electron microscopy. The mandibles 
were removed, photographed with a Wild 
400 wide-field optical microscope, and ana- 
lyzed with the Gandalfi x-ray spectrometer 
in the Department of Mineralogy, Smith- 
sonian Institution. 

Taxonomic Characters 

Jaw morphology and composition. —X-ray 
dispersion analysis of the maxillae of Ly- 
sarete brasiliensis indicates the presence of 
abundant calcium carbonate. The mandi- 
bles are mineralized with calcite, as are jaws 
in the lumbrinerids (Colbath 1986). Calci- 
fication is also evident in L. australiensis 
Benham, 1915, in that the blades of the 
mandibles are white, and the maxillae are 
dark brown rather than black. The presence 
of calcite is inferred by comparison with L. 
brasiliensis. 

In contrast to Lysarete, jaws in species of 
Oenone Savigny, Halla Costa and Tainokia 
Knox & Green (commonly included in the 
Lysaretidae) are not mineralized, but rather 
are composed almost exclusively of sclero- 
protein. As a consequence, both maxillae 
and mandibles are jet black in color. The 
distinction between heavily sclerotized and 
calcified jaws is apparent upon inspection, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



and was described by Benham (1915:233, 
236-237). Benham made no comment about 
the utility of this distinction above the 
species level, however. 

Other aspects of jaw morphology are cor- 
related with the difference in mineraliza- 
tion. In Lysarete the maxillary carriers are 
relatively short (approximately equal to the 
length of the maxillae) and broad (Fig. lb), 
with no ventral ligament (= "third carrier"). 
A thick band of transverse muscles joins the 
carriers dorsally (Fig. 2f ), allowing them to 
be pulled closed in the "trap" motion de- 
scribed by Wolf (1980) for other labido- 
gnath jaws. The MI (maxilla I) elements in 
the maxillary apparatus are the same length 
(Fig. 2c), which allows the anterior tips to 
come together in a pincer-like fashion as 
part of the trap motion. The posterior dorsal 
surfaces of the Mis exhibit an interlocking 
ridge and furrow system (Fig. 2e), which 
may help to lock the apparatus in a closed 
position (Wolf 1980). 

In species of Oenone, Halla and Tainokia 
the carriers are narrow and much longer than 
the combined length of the maxillae, and 
are not equiped with a transverse muscle 
band. Instead, a series of long, oblique-lon- 
gitudinal muscles are attached along the 
length of the carriers. A ventral ligament is 
present. The right and left MI elements may 
be of different lengths (Crossland 1924; 
Hartman 1944; Kielan-Jaworowska 1966), 
and no basal ridge and furrow system is 
present to lock the jaws in a closed position. 
Such jaws are presumably incapable of per- 
forming a trap motion, as Wolf (1980) also 
concluded for the arabellids. 

Soft-part morphology.— The genera tra- 
ditionally included within the Lysaretidae 
share an elongated post-setal lobe and 
prominent dorsal cirrus on posterior para- 
podia. The morphology of the head, how- 
ever, is distinctly different in each of the 
four genera under discussion. Although the 
difference between Lysarete and the other 
genera may not merit consideration as a 
family-level character, the anterior mor- 



phology is none-the-less a useful key char- 
acter. 

In Lysarete, three antennae emerge from 
the posterior margin of the prostomium, and 
may lie flat in a V-shaped notch (Fig. la). 
In adult specimens the notch extends pos- 
teriorly across the peristomial rings, and in 
some cases across the first one or two seti- 
gers. Halla has three antennae which may 
lie back in a semicircular notch in the two 
peristomial rings (Fig. le). Tainokia has a 
single antennae, and a slight invagination 
which extends posteriorly only partially 
across the first of two peristomial rings 
(Knox & Green 1972, fig. 1). In Oenone, 
three antennae are tucked into a nuchal fold 
(Fig. Id), no notch is present, and dorsally 
the peristomium has only one distinct ring. 

Setae and acicula.— All of the worms 
considered here have simple geniculate se- 
tae. In addition, bifid hooks have been re- 
ported in Oenone and Halla. In Lysarete, 
Oenone, and Tainokia the geniculate setae 
are smooth (Knox & Green 1972, figs. 6, 7; 
pers. observ.). In Halla, however, the dorsal 
setae are denticulate, and the ventral setae 
are marked by distinctive oblique striae (Fig. 
2a, b; Fauvel 1923:427, fig. 169 g, h). 

Both described species of Lysarete have 
black acicula. In Oenone fulgida (Savigny, 
1818) and Halla parthenopeia (Chiaje, 1 828) 
the acicula are amber, while Knox & Green 
(1972:433) described the acicula in Taino- 
kia iridescens as colorless. Black acicula are 
thus useful for distinguishing Lysarete from 
the other three genera, supporting the fam- 
ily-level distinction based on jaw morphol- 
ogy. 

Systematics 

Order Eunicida Fauchald, 1977 
Family Lysaretidae Kinberg, 1865 

Emended diagnosis. — Eunicoid poly- 
chaetes with jaws mineralized with calcite. 
Mandibles flat with semicircular growth 
rings anteriorly (Fig. 2d). Maxillary carriers 
no longer than combined length of maxillae, 



VOLUME 102, NUMBER 1 



119 








Fig. 1 . a-c, Lysarete brasiliensis, holotype, Brazil; a, Dorsal view, anterior end, illustrating V-shaped notch 
in peristomium; b, Dorsal view, MI elements and carriers, muscle attachment stippled; c, Labial view, MIIL- 
MVL. d, Oenonefulgida, USNM 1 7729, Dry Tortugas, dorsal view, anterior end, note tips of antennae protruding 
from nuchal fold, e, Halla parthenopeia, USNM 5 147, Bay of Naples, dorsal view, anterior end, note semicircular 
notch in peristomium. All scale bars = 1 mm. 



articulated by prominent transverse mus- 
cular bundle, no ventral ligament. Maxillary 
jaws symmetrical. MI elements of identical 
length, distally falcate with prominent sec- 
ondary fang, additional small denticles may 
be present at base. Three short antennae 
arise from base of prostomium. Elongate 
post-setal lobe and flattened dorsal cirrus 
(or notopodium) developed in posterior 
parapodia. Simple geniculate setae present, 
bifid hooks absent. 

Comparison.— The Lysaretidae are dis- 
tinguished from the Oenonidae based on 



jaw morphology and mineralogy. Lysarete, 
the only genus presently recognized within 
the family, is further distinguished from the 
Oenonidae in having black acicula, and a 
V-shaped notch in the dorsal surface of the 
peristomium. 

The Lysaretidae are similar to the Lum- 
brineridae in having: 1) jaws mineralized 
with calcite; 2) flat mandibles with a semi- 
circular blade; 3) symmetrical maxillary 
jaws; 4) transverse musculature on the car- 
riers. The Lysaretidae are distinguished from 
the Lumbrineridae by: 1) well-developed 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. a, b, Halla parthenopeia, USNM 5147, Bay of Naples; a, SEM dentate setae from dorsal bundle, 
posterior parapodium. scale bar = 10 /zm; b, SEM striate setae from ventral bundle, scale bar = 50 fim. c-g, 
Lysarete brasiliensis, USNM 55884, Gulf of Mexico; c, SEM maxillary apparatus, dorsal view, scale bar = 1 
mm; d, Light photomicrograph mandibles, ventral view, scale bar = 1 mm; e, SEM MI elements, oblique-lateral 
view, note ridges and grooves on bases of Mis, scale bar = 0.5 mm; f, SEM carriers, oblique-lateral view, carriers 
canted at an angle to Mis, note transverse muscle band, scale bar = 0.2 mm; g, SEM anterior maxillary jaws, 
oblique-lateral view, scale bar =0.2 mm. 



dorsal cirri; 2) a prominent secondary fang 
on the MI elements of the maxillae; 3) the 
absence of hooded hooks. Kuwaita Moham- 
mad, 1973 is intermediate between the Ly- 
saretidae and Lumbrineridae in having an- 



tennae on the prostomium, but clearly 
belongs in the Lumbrineridae based on the 
3 criteria above (Mohammad 1973:34, 36, 
figs. 6, 7). 

Distribution.— Lysarete includes one 



VOLUME 102, NUMBER 1 



121 



species recorded from the east coast of North 
and South America in low latitudes, and a 
second species from a single locality east of 
Australia. Larymna Kinberg, 1865 may also 
belong in the Lysaretidae based on jaw mor- 
phology (Hartman 1948:99). The type and 
only specimen (from Mozambique) is so 
poorly preserved that, according to Hart- 
man, attribution cannot be made below the 
family level, and Larymna should be re- 
garded as unrecognizable. 

Genus Lysarete Kinberg, 1865 

Type species.— L. brasiliensis Kinberg, 
1865:570. 

Lysarete brasiliensis Kinberg, 1865 
Figs, la-c, 2c-g 

Lysarete brasiliensis Kinberg 1865:570; 
1910:49, pi. 17, fig. 30.-Ehlers 1887:107- 
108, pi. 33, figs. l-8.-Rioja 1944:130- 
131, figs. 47-50. -Hartman 1951:65, pi. 
14, figs. 7, 8.-Orensanz 1975:106-108, 
fig. 8. -Gardiner 1976:213, figs. 29b-e.- 
Gilbert 1984:43-6-43-7, figs. 43-3, 43-4. 

lOenone brevimaxillata. — Treadwell 1931: 
1-3, figs. 4-8. 

Remarks.— In her re-appraisal of Kin- 
berg's type material, Hartman (1948:8) did 
not consider the holotype of Lysarete bra- 
siliensis, but instead referred to an earlier 
revision by Ehlers (1887:107-108, pi. 33). 
Ehlers did not figure the Brazilian type of 
L. brasiliensis, but rather figured a specimen 
collected from Florida. The posthumous 
publication of Kinberg' s plates (Kinberg 
1910) did not include illustrations of the 
jaws of the holotype. The maxillae are there- 
fore illustrated here (Fig. lb, c). 

On the holotype, maxillary jaws III and 
IV each have a prominent anterior denticle, 
with two progressively smaller denticles 
posteriorly. The jaws in a specimen from 
Argentina (Rioja 1944, fig. 50) are similar, 
except that the Mill elements have four 
denticles instead of three. 

Most specimens examined from the east 



coast of the United States have five to six 
denticles on the Mill and four to seven den- 
ticles on the MIV elements (Fig. 2g). The 
anterior three denticles on the Mill element 
are sub-equal in length, unlike those of the 
type. 

One specimen from Florida (FSBC I 
31008) has Mill jaws like the type, how- 
ever, and a specimen from Argentina illus- 
trated by Orensanz (1 975, fig. 8-3) has Mills 
and IVs indistinguishable from most of the 
North American specimens. There is thus 
some overlap in jaw morphology between 
the North and South American populations. 
The number and shape of denticles on an- 
terior jaws is variable in eunicoid poly- 
chaetes, and the two populations are ten- 
tatively considered conspecific in spite of 
their disjunct geographic ranges. 

The type and only specimen of Oenone 
brevimaxillata Treadwell, 1931 was never 
deposited (H. S. Feinberg, AMNH, pers. 
comm. 1985), and is presumed lost. Tread- 
well's (1931, fig. 8) illustration of the max- 
illae is schematic, and insufficient for de- 
tailed comparison. 

Family Oenonidae Kinberg, 1865 

Emended diagnosis. —Jaws jet black, 
composed primarily of scleroprotein, no 
crystalline mineral phases present. Mandi- 
bles H-shaped. Maxillary carriers much 
longer than anterior jaws, attached to bun- 
dles of oblique-longitudinal muscles, ven- 
tral ligament present. Maxillae commonly 
asymmetrical, Mis differ slightly to mark- 
edly in length. One or three antennae arise 
from base of prostomium. Elongate post- 
setal lobe and prominent dorsal cirri de- 
veloped on posterior parapodia. Geniculate 
setae present, bifid hooks present or absent. 

Key to the genera of Oenonidae 

la. One or 3 antennae recline in notch 
in peristomium, 2 peristomial rings 
dorsally, bifid hooks present or ab- 
sent 2 

lb. Three antennae tucked into nuchal 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



fold, 1 peristomial ring dorsally, 
bifid hooks present Oenone 

2a. One antenna, peristomial notch 
weakly developed, bifid hooks ab- 
sent, setae smooth Tainokia 

2b. Three antennae, distinct semicir- 
cular peristomial notch, bifid hooks 
present (small individuals only), 
dorsal setae dentate, ventral setae 
striated Halla 

Genus Halla A. Costa, 1 844 

Type species.— Halla parthenopeia 
(Chiaj e) Costa, 1844:63.— Lysidice parthe- 
nopeia Chiaje, 1828:175. 

Remarks. -Fauvel (1923:427, fig. 169f) 
described Halla parthenopeia as having bi- 
fid hooks, in addition to the setae figured 
here (Fig. 2a, b). Hooks are absent in para- 
podia from the large specimen of H. par- 
thenopeia examined here (USNM 5 1 47), nor 
were hooks recorded from the large speci- 
men figured by Ehlers (1868:28, figs. 27, 
28). The type of Halla okudai Imajima 
(1967:437) is also a large worm, and lacks 
hooks (Imajima & Hartman 1964:267-268). 
Individuals of Halla which attain large size 
may lose their bifid hooks. A similar con- 
dition prevails in several large species with- 
in the Eunicidae (K. Fauchald, pers. comm. 
1985). 

Distribution. — All western hemisphere 
specimens deposited in the USNM collec- 
tion as Halla proved to be misidentified 
members of Oenone or Lysarete. Halla may 
be restricted to the Old World, with docu- 
mented reports from the Mediterranean 
(Fauvel 1923) and Japan (Imajima 1967). 

Acknowledgments 

I thank Kristian Fauchald for extensive 
discussions about the taxonomy of eunicoid 
polychaetes, and for critically reading the 
manuscript. Thomas H. Perkins arranged 
the loan of material from the Florida De- 
partment of Natural Resources. H. S. Fein- 
berg, American Museum of Natural His- 



tory, provided information concerning the 
type of Oenone brevimaxillata Treadwell. 
This research was supported by a post-doc- 
toral fellowship, Smithsonian Institution. 

Literature Cited 

Benham, W. B. 1915. Report on the Polychaeta ob- 
tained by the F. I. S. "Endeavour" on the coasts 
of New South Wales, Victoria, Tasmania and 
South Australia, part I.— Biological Results of 
the Fishing Experiments carried on by the F. I. 

5. "Endeavour," 1909-14:173-273. 

Chiaje, S. delle. 1 828. Memorie sulla storia a notomia 
degli animali senza vertebre del regno di Napoli, 
3. Napoli, 232 pp. 

Colbath, G. K. 1986. Jaw mineralogy in eunicean 
polychaetes (Annelida).— Micropaleontology 32: 
186-189. 

Costa, A. 1 844. Cerni intorno alle osservazioni zool- 
ogiche fatte durante intre mesi vernali del 
1844.— Annales Accademia aspirantes Natu- 
rales Napoli 2:40-68. 

Crossland, C. 1924. Polychaeta of tropical East Af- 
rica, the Red Sea and Cape Verde Islands. The 
Lumbriconereidae and Staurocephalidae.— 
Proceedings of the Zoological Society of London 
1924:1-106. 

Ehlers, E. 1864-1868. Die Bomstenwurmer (Anne- 
lida, Chaetopoda). Leipzig, 748 pp. 

. 1887. Reports on the results of dredging in 

the U.S. Coast steamer 'Blake.' XXXI. Report 
on the annelids.— Memoirs of the Museum of 
Comparative Zoology, Harvard 15:1-335. 

Fauchald, K. 1 970. Polychaetous annelids of the fam- 
ilies Eunicidae, Lumbrineridae, Iphitimidae, 
Arabellidae, Lysaretidae and Dorvilleidae from 
western Mexico.— Allan Hancock Monographs 
in Marine Biology 5:1-335. 

. 1977. The polychaete worms . — Natural His- 
tory Museum of Los Angeles, Science Series 28: 
1-188. 

Fauvel, P. 1923. Faune de France 5, polychetes er- 
rantes.— Office Central de Faunistique, Paris, 
488 pp. 

Gardiner, S. L. 1976. Errant polychaete annelids from 
North Carolina.— Journal of the Elisha Mitchell 
Scientific Society 91:77-220. 

Gilbert, K. M. 1984. Family Lysaretidae Kinberg, 
1865. Pp. 43-1-43-7 in J. M. Uebelacker & P. 
G. Johnson, eds., Taxonomic guide to the poly- 
chaetes of the Northern Gulf of Mexico, Volume 

6. Barry A. Vittor and Associates, Mobile, Al- 
abama. 

Hartman, O. 1944. Polychaetous annelids. Part V. 
Eunicea.— Allan Hancock Pacific Expedition 10: 
1-339. 



VOLUME 102, NUMBER 1 



123 



. 1948. The marine annelids erected by Kin- 
berg, with notes on some other types in the 
Swedish State Museum.— Arkiv For Zoologi 
42A: 1-137. 

. 1951. The littoral marine annelids of the Gulf 

of Mexico. — Publications of the Institute for 
Marine Science, University of Texas 2:7-124. 

Hartmann-Schroder, G. 1967. Feinbau und Funktion 
des Kieferapparates der Euniciden am Beispiel 
von Eunice (Palola) siciliensis Grube (Poly- 
chaeta). — Mitteilungen aus dem Hamburg Zool- 
ogischen Museum und Institut 64:5-27. 

Imajima,M. 1967. Errant polychaetous annelids from 
Tsukumo Bay and vicinity of Noto Peninsula, 
Japan. — Bulletin of the National Science Mu- 
seum, Tokyo 10:403-441. 

, & O. Hartman. 1964. The polychaetous an- 
nelids of Japan.— Allan Hancock Foundation 
Occasional Papers 26:1-452. 

Kielan-Jaworowska, Z. 1966. Poly chaete jaw appa- 
ratuses from the Ordovician and Silurian of Po- 
land and a comparison with modern forms.— 
Paleontologica Polonica 16:1-152. 

Kinberg, J. G. H. 1865. Annulata nova.— Ofversigt 
af Svenska Kongliga Vetenskaps-Akademiens 
Forhandlingar 21:559-574. 

. 1910. Kongliga Svenska Fregatten Eugenies 



Resa omkrig jorden. Vetenskapliga Iakttagelser. 
II. Zoologi. 3. Annulater, Uppsala, 78 pp. 

Knox, G. A., & K M. Green. 1972. The polychaetes 
of New Zealand part 3: Lysaretidae.— Journal 
of the Royal Society of New Zealand 2:43 1^*34. 

Mohammad, M.-B. 1973. New species and records 
of polychaete annelids from Kuwait, Arabian 
Gulf.— Zoological Journal of the Linnean So- 
ciety 52:23^14. 

Orensanz, J. M. 1975. Los anelidos poliquetos de la 
provincia biogeographica Argentina. VII. Eu- 
nicidae y Lysaretidae. — Physis 34:85-1 1 1. 

Rioja, E. 1944. Estudios anelidologicos. X. Estudio 
de algunos poliquetos del Museo Argentino de 
Ciencias Naturales.— Anales del Instituto de 
Biologia, Mexico 15:115-138. 

Treadwell, A. L. 1931. New species of polychaetous 
annelids from California, Mexico, Puerto Rico, 
and Jamaica.— American Museum Novitates 
(N.Y.) 1023:1-7. 

Wolf, G. 1980. Morphologische Untersuchungen an 
den Kieferapparaten einiger rezenter und fos- 
siler Eunicoidea (Polychaeta). — Senckenber- 
giana Maritima 12:1-182. 

2195 Deborah Way, Upland, California 
91786, USA. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 124-130 

REDESCRIPTION OF NEPHTYS SQUAMOSA EHLERS 
(POLYCHAETA: NEPHTYIDAE) 

Takashi Ohwada 

Abstract.— Confusion about the identity of Nephtys squamosa Ehlers, 1887 
requires redescription of the species based on the syntypes. A lectotype is 
designated. Comparisons are made with similar species. 



Nephtys squamosa was originally de- 
scribed by Ehlers (1 887) as having scale-like 
structures on both dorsum and ventrum, a 
character unique to the family Nephtyidae. 
Many subsequent authors have described 
N. squamosa on the basis of specimens from 
temperate and tropical waters of the Atlan- 
tic and the East Pacific (Augener 1918; 
Monro 1933; Fauvel 1936; Hartman 1940, 
1950; Pettibone 1963; Nonato & Luna 1970; 
Fauchald 1972, 1976; Gardiner 1976; Tay- 
lor 1984; Lana 1986). Although all agree on 
the presence of the scale-like lamellae, dis- 
crepancies exist among descriptions in the 
parapodial morphology. None of these de- 
scriptions after that of Augener (1 9 1 8) men- 
tioned the presence of infrapreacicular la- 
mellae on neuropodia, which is clearly stated 
in the original description (Ehlers 1887). 

Types of N. squamosa have been exam- 
ined in the present study to clarify its iden- 
tity. The presence of infrapreacicular la- 
mellae has been confirmed on neuropodia, 
and suprapreacicular lamellae have been also 
found on notopodia. The present exami- 
nation has further shown the presence of 
tiny, slender dorsal cirri on the first noto- 
podia, overlooked by previous workers. 

Nephtys squamosa Ehlers, 1887 
Figs. 1-3 

Nephthys squamosa Ehlers, 1887:128-131, 
pi. 37, figs. 7-10. 

Material examined. —Original specimens 
collected from Florida during the U.S. Coast 



Survey 1868-1869, L. F. Pourtales, assist.: 
off French Reef, Cast No. 6, 103 fms, 3 Apr 
1869, paralectotype (MCZ 735). -off French 
Reef, Cast No. 4, 50 fms, 3 Apr 1869, para- 
lectotype (MCZ 740).— off Alligator Reef, 
Cast No. 9, 113 fms, 8 May 1869, paralec- 
totype (MCZ 836). — off American Shoal, 55 
fms, lectotype (MCZ 1202). 

Measurements. —Lectotype complete, 
48.4 mm long, 3.4 mm wide including para- 
podia with 120 setigers. Paralectotypes 
(MCZ 735, 740, 836) incomplete posterior- 
ly, respectively, 21.7 mm long, 3.5 mm wide 
with 47 setigers, 54.4 mm long, 2.9 mm 
wide with 105 setigers, 30.7 mm long, 2.4 
mm wide with 96 setigers. 

Description. —Body creamy white to 
brown, lacking pigmentation in alcohol. Eyes 
absent. 

Prostomium pentagonal with anterior 
margin convex, thin, spatulate (Fig. la, b); 
lateral margins rounded, broadest between 
second antennae (Fig. 2a), posterior margin 
with V-shaped middorsal projection ex- 
tending to posterior border of setiger 1 . An- 
terior part between first antennae translu- 
cent, marked by an intricate pattern of 
opaque tissue (Figs, lb, 2a). First antennae 
small, flattened, attached obliquely, ap- 
pearing as raised lobes originating some dis- 
tance from and fused with frontal prosto- 
mial margin (Figs, lb, 2a). Second antennae 
large, basally flattened and distally conical, 
slightly constricted at bases, much larger 
than 1 st antennae, directed laterally at mid- 
dle of ventral surface of prostomium, well 



VOLUME 102, NUMBER 1 



125 




Fig. 1. Nephtys squamosa (setae omitted): a, c, d, lectotype (MCZ 1202), b, paralectotype (MCZ 735): a, 
Anterior end, dorsal view, ventrally dissected and somewhat flattened, dorsal cirrus missing and preacicular 
lobe twisted outside on 1 st right notopodium (acL-acicular lobe; prL-preacicular lobe); b, Same, ventral view, 
3rd right ventral cirrus missing (notopodia omitted); c, 72nd to 76th setiger, dorsal view (neuropodia omitted); 
d, 74th to 78th setiger, ventral view (notopodia omitted). 



behind 1 st antennae and slightly anterior to 
1st neuropodia (Figs, la, b; 2a, b). Mouth 
covered with two thick lateral lips and thick 
rugose midventral lip at level of setiger 1 
(Fig. lb); rugose area extending to setiger 5. 
Setiger 1 modified (Figs, la, 2b); noto- 



podia reduced, with triangularly pointed 
acicular lobes, short preacicular and post- 
acicular lobes (preacicular lobes concealed 
behind acicular lobes and usually not rec- 
ognizable from above); dorsal cirri (barely 
visible unless stained) tiny, slender, point- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Nephtys squamosa: a, c, paralectotype (MCZ 735), b, lectotype (MCZ 1202): a, Anterior half of 
prostomium, dorsal view (setae omitted); b, 1st left parapodium, dorsal view, preacicular lobe obscured by 
acicular lobe (acL-acicular lobe; dc-dorsal cirrus; tips of setae omitted); c, 12th right neuropodium, dorsal view 
(setae omitted). 



ed, arising from outer side of acicular lobes 
near their base between preacicular and 
postacicular setal fascicles (Fig. 2b). Neu- 
ropodia of setiger 1 on anterior margin, lat- 
eral to prostomium, with slender rudimen- 
tary acicular lobes projecting anteriorly 
(Figs, la, 2b); acicular lobes supported by 
thin acicula whose tip curves inward. Ven- 
tral cirri large, similar to or smaller than 
2nd antennae, digitate but tips may be con- 
ical, lateral to neuropodia and continuous 
with anterior setigerous margins, directed 
laterally or bent backward (Figs, la, 2b). 
Notosetae of setiger 1 including preacicular 
barred (laddered) capillaries and very finely 
serrated slender postacicular capillaries; 
neurosetae long, thin, smooth capillaries, 
surrounding acicular lobes (Fig. 2b). 

Parapodia similar throughout from se- 
tiger 2; both rami with preacicular and post- 
acicular setal fascicles; branchiae (interra- 
mal cirri) may be lacking on setiger 2. Both 
notopodial and neuropodial acicular lobes 
conical (Fig. 3a-c), becoming acutely coni- 



cal and pointed posteriorly (Fig. 3e). No- 
topodial presetal lamellae thin, rather broad 
and slightly bilobed, distally rounded, ex- 
tending from lower margins of notopodia, 
as long as acicular lobes on anterior para- 
podia (Fig. 3a-c), posteriorly reduced and 
much shorter than acicular lobes (Fig. 3e). 
Small lamellae dorsal to notopodial presetal 
lobes, thinner than presetal lobes except on 
middle parapodia, rounded on anterior no- 
topodia (Fig. 3a, b), posteriorly reduced and 
becoming much thinner and smaller (Fig. 
3c, e), barely recognizable on posterior se- 
tigers (Fig. 3e). Preacicular setae appearing 
mostly beyond dorsal margins of presetal 
lamellae and usually partially covering small 
suprapreacicular lamellae (Fig. 3a-c, e). 
Neuropodial presetal lamellae thin, larger 
than notopodial presetal lamellae, extend- 
ing from upper margins of neuropodia rath- 
er triangularly on middle parapodia (Fig. 
3b, c), distally rounded on all but especially 
on anterior and posterior parapodia (Fig. 
3a, e), as long as acicular lobes on anterior 



VOLUME 102, NUMBER 1 



127 



0.5 mm 




Fig. 3. Nephtys squamosa (tips of setae omitted): a, b, paralectotype (MCZ 735), c-e, lectotype (MCZ 1202): 
a, Left parapodium, setiger 6, anterior view; b, Same, setiger 32, anterior view; c, Same, setiger 40, anterior 
view; d, Same, posterior view (preacicular setae omitted); e, Right parapodium, setiger 79, anterior view. 



neuropodia (Fig. 3a, b), posteriorly reduced 
and much shorter than acicular lobes (Fig. 
3e). Lamellae ventral to neuropodial pre- 
setal lobes, broad, thin, shorter than presetal 
lobes, slightly bilobed on anterior parapodia 



(Fig. 3a, b), posteriorly reduced and becom- 
ing narrower and much thinner, thinner than 
presetal lobes, hardly recognizable on pos- 
terior setigers (Fig. 3e). Preacicular setae ap- 
pearing mainly beyond ventral margins of 



128 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



presetal lamellae and usually partially cov- 
ering infrapreacicular lamellae (Fig. 3a-c, 
e). Notopodial postsetal lamellae thin, lig- 
ulate, extending dorsolaterally from seg- 
mental wall above and longer than acicular 
lobes, distally rather rounded on anterior 
setigers (Fig. 3a), posteriorly becoming more 
or less pointed (Fig. 3b-e). Neuropodial 
postsetal lamellae large, thin, extending from 
behind and longer than acicular lobes (Fig. 
3a-e), posteriorly extending much further 
(Fig. 3c, e), but degree of extension variable 
between specimens (Fig. 3b, c), with rather 
pointed tips except on anterior parapodia 
where distally rounded (Fig. 3a). Small, short 
lobe on superior edge of neuropodia be- 
tween presetal and postsetal lamellae (Figs. 
2c, 3a, d, e) except on setiger 1, distinct on 
anterior neuropodia but posteriorly reduced 
and barely recognizable. Dorsal cirri flat- 
tened, thin, triangularly elongate with rather 
pointed tips, smaller than notopodial post- 
setal lamellae, extending laterally from bas- 
al part of branchiae behind acicular lobes 
(Fig. 3a-e). Ventral cirri digitiform on an- 
terior several parapodia (Figs, lb, 3a), 
thereafter foliaceous, constricted at bases 
(Figs, lb, 3b-d). 

Branchiae first present on setigers 2-3 
usually as processes slightly curved out- 
ward, completely recurved within a few se- 
tigers (Fig. 3a), thereafter gradually increas- 
ing in size and fully developed by setigers 
25-27 (Fig. 3c-e), then (in lectotype) de- 
creasing gradually in size after setigers 89- 
92, and not recurved after setigers 106-107, 
still present as very small rudimentary ridge 
on setiger 116, completely absent on last 4 
setigers. 

Scale-like dorsal lamellae broad, folia- 
ceous (Fig. lc), originating from dorsolat- 
eral wall of body (Fig. 3b-e), extending 
mainly posterolateral^ over notopodia, 
partially overlapping successive segments in 
shingle-like arrangement (Fig. lc), begin- 
ning on setigers 5-9, becoming larger more 
posteriorly but not fully developed until 



middle setigers (Fig. 3c, d), gradually de- 
creasing in size (in lectotype) over posterior 
several setigers but still recognizable on se- 
tiger 117, absent on last 3 setigers. Similar 
but much smaller lamellae present on ven- 
trolateral surface of body (Figs. Id, 3c, d), 
foliaceous, extending somewhat postero- 
laterally, only slightly covering neuropodia 
and successive segments (Fig. Id), begin- 
ning on setiger 2, increasing in size more 
posteriorly but not fully developed until 
middle setigers (Fig. 3c, d), gradually re- 
duced (in lectotype) over posterior several 
setigers but still recognizable on setiger 117, 
absent on last 3 setigers. 

Both preacicular and postacicular setae 
occurring in spreading fascicles, with post- 
acicular fascicles wider than preacicular ones 
(Fig. 3a-c, e); preacicular setae much shorter 
and fewer than postacicular setae. Most 
preacicular setae consisting of barred cap- 
illaries; a few, similarly long, very finely ser- 
rated capillaries at upper and lower end of 
preacicular fascicles. Postacicular setae con- 
sisting of finely serrated long capillaries. No 
lyrate seta found. Acicula single in rami, 
completely embedded in conical acicular 
lobes (Fig. 3b-e). 

Proboscis not everted in specimens ex- 
amined. Proboscis in lectotype previously 
dissected, damaged, incomplete, with ter- 
minal circlet of 20 bifid papillae and simple 
middorsal papilla, with at least 20 rows of 
5-6 cirriform papillae subterminally, and 
more distal cirriform middorsal papilla; 
middorsal papilla not noticeably larger nor 
longer than others. Pair of horny jaws pres- 
ent inside proboscis. Pharyngeal trepans ab- 
sent. 

Remarks.— In the original description, 
Ehlers (1887:130) stated that dorsal cirri 
were about half as long as notopodial post- 
setal lamellae. This is the case in the middle 
parapodia of the lectotype (Fig. 3c, d). In 
two paralectotypes (MCZ 735, 836), how- 
ever, notopodial postsetal lamellae do not 
extend beyond dorsal cirri (Fig. 3a, b). The 



VOLUME 102, NUMBER 1 



129 



relative length of notopodial postsetal la- 
mellae to dorsal cirri is thought to vary 
greatly between specimens. 

There are obvious problems with descrip- 
tions of Nephtys squamosa. Although the 
presence of suprapreacicular lamellae on 
notopodia has been clearly recognized in the 
lectotype and paralectotypes in the present 
study, neither the original description (En- 
ters 1887) nor any subsequent description 
mentioned this structure, whereas Ehlers 
described infrapreacicular lamellae on neu- 
ropodia (p. 130, Taf. 37, fig. 8). Examina- 
tion of the lectotype and paralectotypes also 
revealed some other structures such as tiny, 
slender dorsal cirri on 1st notopodia and 
small, short lobes on the superior edge of 
neuropodia between presetal and postsetal 
lamellae, which had been overlooked by 
previous workers. 

Descriptions subsequent to Augener 
(1918) did not mention the presence of in- 
frapreacicular lamellae on neuropodia. 
Hartman's figure (1940: pi. 41, fig. 98) of 
the 25th parapodium of N squamosa from 
the Gulf of California showed only preacic- 
ular lamellae both on the notopodium and 
the neuropodium, and she stated (p. 237) 
that the prostomium of her specimens dif- 
fered from that originally described for the 
species. Neither is an additional preacicular 
lamella shown on either the notopodium or 
the neuropodium in Taylor's figure (1984: 
fig. 3 5 -8b) of the 20th parapodium of N 
squamosa from the Gulf of Mexico. Taylor 
also stated (p. 1 1) that dorsal cirri were lack- 
ing on the 1 st notopodia of his specimens, 
whereas dorsal cirri, although tiny and slen- 
der, are present on 1st notopodia of the lec- 
totype and paralectotypes. Additional pre- 
acicular lamellae are neither described nor 
figured also in Lana's description (1986:139, 
figs. 4, 5) of N. squamosa from Brazil. Since 
additional preacicular lamellae, especially 
suprapreacicular lamellae on notopodia, are 
difficult to observe in some specimens, the 
lack of description may not necessarily im- 



ply absence in specimens concerned. Never- 
theless, it seems to be necessary to reex- 
amine these specimens for certain 
identification. 

Nephtys squamosa is closely related to N. 
acrochaeta Hartman, 1950, N. imbricata 
Grube, 1857 and N. serratifolia Ehlers, 1897 
in the possession of scale-like dorsal la- 
mellae. Nephtys squamosa, however, differs 
from N. acrochaeta in that branchiae are 
present from setigers 2-3 in the former and 
9-10(Hartman 1950, Fauchald 1976) in the 
latter. Neuropodial postsetal lamellae are 
lacinated in N acrochaeta (Hartman 1950) 
and N serratifolia (Ehlers 1897), whereas 
they are entire in N. squamosa. Capillary 
setae with a large spur are present in post- 
acicular fascicles both in A r . acrochaeta 
(Hartman 1950) and N. serratifolia (Hart- 
man-Schroder 1 965), but absent in N squa- 
mosa. Nephtys imbricata is 'too incomplete- 
ly known for certain identification' 
(Hartman 1950:111), but according to the 
original description (Ehlers 1887), N. squa- 
mosa differs from N. imbricata in parapo- 
dial morphology. I have examined types of 
both species, however further study will be 
needed to fully characterize N. imbricata. 

Acknowledgments 

I am grateful to Dr. Ardis B. Johnston, 
Museum of Comparative Zoology, Harvard 
University (MCZ) for the loan of the spec- 
imens. The manuscript benefited from the 
helpful suggestions by Dr. Mary E. Petersen, 
Zoological Museum, University of Copen- 
hagen, Denmark and the critical review by 
Dr. Thomas H. Perkins, Florida Depart- 
ment of Natural Resources. 

Literature Cited 

Augener, H. 1918. Polychaeta. — Beitrage zur Kennt- 
nis des Meeresfauna West-Afrikas 2:67-625. 6 
pis. 

Ehlers, E. 1887. Report on the annelids of the dredg- 
ing expedition of the U.S. coast survey steamer 
Blake. — Memoirs of the Museum of Compar- 



130 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ative Zoology of Harvard College 15:vi + 335 
pp., 60 pis. 

. 1897. Polychaeten.— Hamburger Magalla- 

nische Sammelreise. Hamburg, Friedrichsen & 
Co. 148 pp., 9 pis. 

Fauchald, K. 1972. Benthic polychaetous annelids 
from deep water off western Mexico and adja- 
cent areas in the eastern Pacific Ocean.— Allan 
Hancock Monographs in Marine Biology No. 7: 
1-575. 

. 1976. Some Nephtyidae (Polychaeta) from 

Ubatuba, Brasil.— Bulletin of the Southern Cal- 
ifornia Academy of Sciences 75:16-19. 

Fauvel, P. 1936. Contribution a la fauna des Anne- 
lides polychetes du Moroc. — Memoires de la So- 
ciete des Sciences Naturelles du Moroc 43:1- 
143. 

Gardiner, S. L. 1 976. Errant polychaete annelids from 
North Carolina.— Journal of the Elisha Mitchell 
Scientific Society 91:77-220. 

Grube, E. 1857. Annulata Orstediana Enumeratio 
Annulatorum, quae in itinere per Indiam occi- 
dentalem et Americam centralem annis 1845— 
1848 suscepto legit cl. A. S. Orsted, adjectis spe- 
ciebus nonnullis a cl. H. Kroyero in itinere ad 
Americam meridionalem collectis.— Videnska- 
belige Meddelelser fra Dansk naturhistorisk 
Forening i Kjobenhavn 1857:1-29. 

Hartman, O. 1940. Polychaetous annelids. Part 2. 
Chrysopetalidae to Goniadidae.— Allan Han- 
cock Pacific Expeditions 7:173-287. 

1950. Polychaetous annelids. Goniadidae, 
Glyceridae, Nephtyidae.— Allan Hancock Pa- 
cific Expeditions 15:1-181. 

Hartmann-Schroder, G. 1965. Zur Kenntnis des Sub- 



litorals der chilenischen Kiiste unter besonderer 
Beriicksichtigung der Polychaeten und Ostra- 
coden. Teil II. Die Polychaeten des Sublito- 
rals. — Mitteilungen aus dem Hamburgischen 
Zoologischen Museum und Institut 62 (Supple- 
ment): 5 9-305. 

Lana, P. C. 1986. Nephtyidae (Annelida; Polychaeta) 
do litoral do estado do Parana (Brazil).— Neri- 
tica 1:135-154. 

Monro, C. C. A. 1933. The Polychaeta Errantia col- 
lected by Dr. C. Crossland at Colon, in the Pan- 
ama region, and the Galapagos Islands during 
the expedition of the S. Y. 'St. George'.— Pro- 
ceeding of the Zoological Society of London 
1933:1-96. 

Nonato, E. F., & J. A. C. Luna. 1970. Anelideos 
poliquetas do nordeste do Brasil. 1 . Poliquetas 
bentonicos da costa de Alagoas e Sergipe.— 
Boletim do Institute Oceanografico, Sao Paulo 
19:57-130. 

Pettibone, M. H. 1963. Marine polychaete worms of 
the New England Region. 1. Aphroditidae 
through Trochochaetidae. — United States Na- 
tional Museum Bulletin No. 227:1-356. 

Taylor, J. L. 1984. Family Nephtyidae Grube, 1850. 
Pp. 1-20 in J. M. Uebelacker & P. G. Johnson, 
eds., Taxonomic guide to the polychaetes of the 
Northern Gulf of Mexico, Vol. 5, Chapt. 35. 
Barry A. Vittor & Associates, Inc., Mobile, Al- 
abama. 

Ocean Research Institute, University of 
Tokyo, 15-1, 1-Chome, Minamidai, Na- 
kano-ku, Tokyo 164 Japan. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 131-133 

PHALLODRILUS HESSLERI, NEW SPECIES 

(OLIGOCHAETA: TUBIFICIDAE), FROM ABYSSAL 

DEPTHS IN THE WESTERN PACIFIC OCEAN 

Christer Erseus 

Abstract.— Phallodrilus hessleri is described from abyssal clay at 5916 m 
depth east of the Philippines. It is characterized by (1) slender, sharply pointed 
somatic setae with very thin (or absent?) upper teeth, (2) somewhat sigmoid 
penial setae in segment XI, 2 per bundle, and (3) very small, spindle-shaped 
atria. The species represents the second deepest record of marine oligochaetes 
in the world. 



In a collection of benthic invertebrates 
from a deep-sea area near the Philippine 
Trench in the western Pacific Ocean, de- 
posited in the National Museum of Natural 
History (USNM), Washington, D.C., the 
author found one well preserved individual 
of a new species of Phallodrilus Pierantoni, 
1902 from a station almost 6000 m deep. 
Oligochaetes from such great depths are in- 
deed rare. It was therefore considered ap- 
propriate to describe the species on the basis 
of this specimen. 

The worm was stained in paracarmine and 
mounted whole in Canada balsam. 

Phallodrilus hessleri, new species 
Fig. 1 

Holotype. -USNM 104128, whole- 
mounted specimen from E of the Philippine 
Trench, West Pacific Ocean, 9°06.0'N, 
127°43.6'E, 5916 m, abyssal red clay (13 
Mar 1975). 

Etymology.— Named for Dr. Robert R. 
Hessler (Scripps Institution of Oceanogra- 
phy, La Jolla, California), who collected the 
sample. 

Description. —Length more than 2.5 mm, 
more than 16 segments (worm not com- 
plete). Width at XI (compressed) 0.22 mm. 
Prostomium large, rounded. Body wall 
bearing scattered, very small particles. Cli- 
tellum extending over V^X-XII. Somatic se- 



tae (Fig. 1 A, B) bifid, with teeth almost par- 
allel, upper tooth shorter and much thinner 
than lower, sometimes not visible (or ab- 
sent?; cf. right bundle in Fig. IB). Bifids 50- 
85 nm long, 1.5-2.5 ^m thick at node, three 
to four per bundle anteriorly, two (or three) 
per bundle in postclitellar segments. Penial 
setae (Fig. 1C, D, ps) two per bundle, some- 
what sigmoid, sharply single-pointed, 65- 
75 Mm long, about 3 ium thick at node, di- 
rected in an almost antero-posterior direc- 
tion. Male pores inconspicuous, paired ven- 
trally and posteriorly in XL Spermathecal 
pores paired, in line with ventral setae(?) in 
anteriormost X. 

Pharyngeal glands in IV-VII. Male gen- 
italia (Fig. ID) paired. Vas deferens 8-10 
jttm wide, coiled, much longer than atrium, 
entering apical end of latter. Atrium pear- 
shaped, about 60 /im long, 28-33 jtim wide, 
with very thin outer lining and ciliated inner 
epithelium; latter densely granulated in ec- 
tal, wide part of atrium. Prostate glands 
compact, round, one at each end of atrium. 
Atrium opening to exterior through simple 
pore; penis or pseudopenis absent. Sper- 
mathecae (Fig. ID, s) not seen in their full 
length (hidden behind gut), but consisting 
of short, narrow duct, and slender ampulla. 

Remarks. — The diagnostic characters of 
this species are (1) the slender, sharply 
pointed somatic setae with very thin upper 



1 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

B.. Z rs D 




25 pm 




J(B-C) 



100 pm 



Fig. 1 . Phallodrilus hessleri; A, Free-hand drawing of tip of somatic seta; B, Two bundles of somatic setae 
(from postclitellar segments); C, Penial setae; D, Horizontal view of spermathecae (only parts visible) and male 
genitalia in segments X-XI. Abbreviations: a, atrium; pr 1, anterior prostate gland; pr 2, posterior prostate 
gland; ps, penial seta; s, spermatheca; sf, sperm funnel; vd, vas deferens. 



teeth (or sometimes absent?), (2) the bisetal 
penial bundles, and (3) the small atria. Oth- 
er species within the genus with similar fea- 
tures are all deep-sea forms, viz. P. profun- 
dus Cook, 1970 (N Atlantic), P. remus 
Erseus, 1979 (NW Atlantic), P. altus Erseus, 
1980 (Beaufort Sea, Arctic), and P. segon- 
zaci Erseus, 1986 (S Indian Ocean), and they 
are probably the closest relatives of P. hess- 
leri. However, in all these other species, the 
penial setae are clearly longer than the so- 
matic setae (setae about equally long in hess- 
leri). The new species is further distin- 
guished from all of them except P. remus 
by its very short atria (atria not very large, 
but elongate in the others), from P. remus 
by the shape of its penial setae (those of 
remus oar- shaped, with wide inner ends), 
and from P. profundus and P. segonzaci by 
its lack of modified genital setae in segments 
IX and X. 

Distribution and habitat.— Known only 
from the type-locality E of the Philippines, 
W Pacific Ocean. Abyssal clay, 5916 m. 

Discussion 

More than 30 species of Tubificidae have 
been described from the deep sea, all within 
the last 20 years (Cook 1970a, b; Erseus 



1979a, b, 1980, 1982a, b, 1983a-c, 1984, 
1986, 1988; see also review by Erseus 
1985). Most of them are bathyal forms, but 
four are known from depths greater than 
4000 m: the North Atlantic Bathydrilus at- 
lanticus Erseus, 1979 (depth range 1600- 
4632 m; Erseus 1 983), the two Indian Ocean 
species Phallodrilus segonzaci and P. stilus 
Erseus, 1986, both from a single station at 
4910m (Erseus 1986), and the North Pacific 
B. hadalis Erseus, 1979, which was collected 
at 7298 m in the Aleutian Trench (Erseus 
1979a). Phallodrilus hessleri (from 5916 m) 
thus represents the second deepest record 
of marine oligochaetes in the world. 

Acknowledgments 

I am indebted to Dr. K. Fauchald for 
bringing my attention to the Philippine 
Trench collection in the USNM, and for the 
loan of the specimen studied; to Dr. R. R. 
Hessler for providing station data; and to 
Ms. Barbro Lofnertz and Mrs. Aino Falck- 
Wahlstrom for technical assistance. 

Literature Cited 

Cook, D.C. 1970a. Peloscolex dukei n. sp. and P. 
aculeatus n. sp. (Oligochaeta, Tubificidae) from 
the N.W. Atlantic, the latter being from abyssal 



VOLUME 102, NUMBER 1 



133 



depths.— Transactions of the American Micro- 
scopical Society 88(1969):492-497. 

. 1970b. Bathyal and abyssal Tubificidae (An- 
nelida, Oligochaeta) from the Gay Head-Ber- 
muda transect, with descriptions of new genera 
and species.— Deep-Sea Research 17:973-981. 

Erseus, C. 1979a. Taxonomic revision of the marine 
genera Bathydrilus Cook and Macroseta Erseus 
(Oligochaeta, Tubificidae), with descriptions of 
six new species and subspecies.— Zoologica 
Scripta 8:139-151. 

. 1979b. Taxonomic revision of the marine 

genus Phallodrilus Pierantoni (Oligochaeta, 
Tubificidae), with descriptions of thirteen new 
species.— Zoologica Scripta 8:187-208. 

. 1980. New species of Phallodrilus (Oligo- 
chaeta, Tubificidae) from the Arctic deep sea 
and Norwegian fjords. — Sarsia 65:57-60. 

. 1 982a. Atlantidrilus, a new genus of deep-sea 

Tubificidae (Oligochaeta).— Sarsia 67:43-46. 

. 1982b. Taxonomic revision of the marine 

genus Limnodriloides (Oligochaeta: Tubifici- 
dae).— Verhandlungen des Naturwissenschaft- 
lichen Vereins in Hamburg (Neue Folge) 25: 
207-277. 

. 1983a. New records of Adelodrilus (Oligo- 
chaeta, Tubificidae), with descriptions of two 
new species from the Northwest Atlantic— Hy- 
drobiologia 106:73-83. 

. 1983b. Deep-sea Phallodrilus and Bathydri- 



lus (Oligochaeta, Tubificidae) from the Atlantic 
Ocean, with descriptions often new species.— 
Cahiers de Biologie Marine 24:125-146. 
-. 1983c. A new bathyal species of Coralliodri- 
lus (Oligochaeta, Tubificidae) from the S.E. At- 
lantic—Proceedings of the Biological Society of 
Washington 96:273-275. 

-. 1984. New and little-known species of deep- 
sea Tubificidae (Oligochaeta) from the North- 
west Atlantic— Zoologica Scripta 13:101-106. 
-. 1985. Distribution and biogeography of Oli- 
gochaeta. Pp. 365-367 in L. Laubier and C. 
Monniot, eds., Peuplements profonds du Golfe 
de Gascogne. Institut Francais de Recherche pour 
l'Exploitation de la Mer (IFREMER), Brest. 
-. 1986. Two new abyssal species of Phallodri- 
lus (Oligochaeta, Tubificidae) from the South 
Indian Ocean.— Bulletin du Museum National 
d'Histoire Naturelle (4 e ser.) 8 (section A, n° 3): 
567-572. 

-. 1988. Deep-sea Tubificidae (Oligochaeta) 
from the Gulf of Mexico.— Proceedings of the 
Biological Society of Washington 1 1 :67-7 1 . 



Zoo-Tax, Swedish Museum of Natural 
History, Stockholm, and (postal address): 
Department of Zoology, University of G6- 
teborg, Box 25059, S-400 31 Goteborg, 
Sweden. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 134-136 

PHALLODRILUS VESCUS, NEW SPECIES 

(OLIGOCHAETA, TUBIFICIDAE) FROM THE 

GULF OF MEXICO 

Christer Erseus 

Abstract.— Phallodrilus vescus (subfamily Phallodrilinae) is described from 
the continental slope of the northern part of the Gulf of Mexico. The species 
is distinguished from the closely related P. davisi Erseus, 1984 by (1) its fewer 
setae, (2) its very small anterior prostate glands, (3) its very short penial papillae, 
and (4) the markedly narrow inner part of its spermathecal ampullae. 



Species of Phallodrilinae from the north- 
ern Gulf of Mexico continental slope, orig- 
inating from a project at LGL Ecological 
Research Associates, Inc. (Bryan, Texas), 
were recently described by Erseus (1988). 
Subsequently, further material was sent to 
this author for identification. Three worms 
proved to represent an additional new 
species of Phallodrilus Pierantoni, 1902, 
which is described herein. 

The specimens, weakly stained in Rose 
Bengal, were mounted whole in Canada bal- 
sam by the author. They are deposited in 
the United States National Museum of Nat- 
ural History (USNM), Smithsonian Insti- 
tution, Washington, D.C. 

Phallodrilus vescus, new species 
Fig. 1 

Holotype. -USNM 118716, from south 
ofLouisiana, 27°49'36"N, 90°07'06' / W, 841 
m (12 Apr 1984). 

Paratypes. -USNM 1 1 87 1 7, 1 1 87 1 8, one 
from type locality, and one from 
27°49'24"N, 90°07'00"W, 840 m (12 Apr 
1984). 

Etymology.— The epithet vescus is Latin 
for "thin, slender," a feature of the new 
species. 

Description. —Length (two complete, fixed 
specimens) 3.4—5.6 mm, 31-38 segments. 
Width at XI (compressed, whole-mounted 



specimens) 0.08-0.13 mm. Prostomium 
rounded triangular, pygidium blunt-ended, 
clitellum extending over V2X-XII. Setae (Fig. 
1A) bifid, with upper tooth slightly thinner 
and shorter than lower. Setae 35-50 jum long, 
1-1.5 nm thick, three per bundle anteriorly, 
two per bundle in postclitellar segments. 
Ventral setae absent from XL Male pores 
paired in line with ventral setae, posterior 
to middle of segment X. 

Pharyngeal glands poorly developed. Male 
genitalia (Fig. 1 B) paired. Vas deferens about 
5 fim wide, long and coiled, entering apical 
end of atrium. Atrium comma-shaped, 
about 30 jum long, about 15 ^m wide, with 
thin outer (muscular) lining and thick, gran- 
ulated inner epithelium; ciliation not seen. 
Anterior prostate gland small, inconspic- 
uous, located near junction between vas def- 
erens and atrium. Posterior prostate gland 
large, conspicuous, attached to ectal part of 
atrium. Atrium terminating in small penial 
papilla, about as long as wide (10-15 ^m), 
enclosed in simple penial sac. Spermathecae 
(Fig. IB, s) slender, 65-90 fim long, con- 
sisting of short, 11-13 fim wide ducts, and 
11-18 /j.m wide ampullae; latter with inner 
end very narrow in (at least) two of the spec- 
imens studied. Lumen of ampullae incon- 
spicuously and irregularly incised in a few 
places. A few spermatozoa, arranged in a 
bundle, present in each ampulla. 

Remarks. — This species is distinguished 



VOLUME 102, NUMBER 1 



135 




Fig. 1 . Phallodrilus vescus: A, Free-hand drawing of somatic seta; B, Lateral view of spermatheca and male 
genitalia in segments X-XL Abbreviations: a, atrium; p, penial papilla; pr 1, anterior prostate gland; pr 2, 
posterior prostate gland; s, spermatheca; vd, vas deferens. 



from the closely related P. davisi, described 
by Erseus (1 984) from sublittoral depths (78- 
80 m) offMassachusetts, by: (1) having only 
two setae per bundle in postclitellar seg- 
ments (up to four setae in davisi), (2) its very 
small anterior prostate glands (in davisi, 
these glands are conspicuous and in fact 
larger than the posterior prostates), (3) its 
very short penial papillae (penes slender and 
conspicuously hollow in davisi), and (4) the 
narrow inner end of its spermathecal am- 
pullae (these ends rounded, not markedly 
narrow in davisi). 

The genitalia of P. vescus bear resem- 
blance to those of P. const rictus Erseus, 1988, 
which also occurs on the upper continental 
slope in the northern Gulf of Mexico (Erseus 
1988). The latter species, however, pos- 
sesses a very thick muscular lining on the 
atria, copulatory sacs and spermathecae; 
penial setae in segment XI; and a very dis- 
tinct constriction on the male ducts between 
the atria and the copulatory sacs; these fea- 
tures are all absent in P. vescus. 

The spermathecal ampullae are markedly 



incised or compartmented in P. davisi (Er- 
seus 1984:fig. 6B) and P. constrictus (Erseus 
1988:fig. 1C). A similar, but less developed, 
compartmentalization is present in the 
spermathecae of P. vescus (Fig. IB). 

Distribution and habitat. — N Gulf of 
Mexico. Upper continental slope sediment, 
840-841 m. 

Acknowledgments 

The LGL specimens reported on in this 
publication were collected through funding 
by the U.S. Department of Interior, Min- 
erals Management Service, Gulf of Mexico 
Regional OCS Office under Contract Num- 
ber 14-12-0001-30046 and 14-12-0001- 
30212. I am indebted to Dr. Linda H. Pe- 
quegnat (LGL) for placing the material at 
my disposal, and to Ms. Barbro Lofnertz 
and Mrs. Aino Falck-Wahlstrom for tech- 
nical assistance. 

Literature Cited 

Erseus, C. 1984. Taxonomy of some species of Phal- 
lodrilus (Oligochaeta: Tubificidae) from the 



1 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Northwest Atlantic, with description of four new Zoo-Tax, Swedish Museum of Natural 

species.-Proceedingsofthe Biological Society History, Stockholm, and (postal address): 

"T^'Teep^s^Tubmddae (Oligochaeta) department of Zoology, University of Go- 

fromtheGulfofMexico.-Proceedingsofthe teborg, Box 25059, S-400 31 Goteborg, 

Biological Society of Washington 101:67-71. Sweden . 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 137-153 

NEW SPECIES OF SCALE-WORMS 

(POLYCHAETA: POLYNOIDAE) FROM THE 

HYDROTHERMAL RIFT-AREA OF THE 

MARIANA BACK- ARC BASIN IN THE 

WESTERN CENTRAL PACIFIC 

Marian H. Pettibone 

Abstract. — Polynoid polychaetes collected by the Mariana Back- Arc Basin 
Expedition in the western Central Pacific in 1987 are described. They include 
four new species belonging to three subfamilies and four genera of Polynoidae: 
Macellicephalinae: Levensteiniella raisae; Lepidonotopodiinae: Lepidonoto- 
podium minutum; and Branchinotogluminae: Branchinotogluma burkensis and 
Opisthotrochopodus marianus. Included also is a brief summary of the subfam- 
ilies, genera, and species associated with deep-sea hydrothermal vents, deep 
seep-sites, trenches, abyssal basins, experimental deep wood panels, and caves. 



In her revision of the genera and species 
that had been grouped in Macellicephalinae 
Hartmann-Schroder, 1971, containing 
mostly abyssal species, Pettibone (1976) 
separated Macellicephalinae and five ad- 
ditional subfamilies: Macellicephaloidinae 
(four species), Macelloidinae, Bathyedithi- 
nae, Polaruschakovinae (two species), and 
Bathymacellinae. In the remaining Macel- 
licephalinae, in addition to Macellicephala 
(seven species), six new genera were added: 
Bathykermadeca, Bathyeliasona (three 
species), Bathyvitiazia, Bathyfauvelia, 
Bathycatalina, and Bathykurila. 

Subsequently, additional new subfami- 
lies, genera, and species have been added. 
Pettibone (1979) emended and added to 
Macellicephalinae Bruunilla natalensis 
(Hartman, 1971). Levenstein (1975) added 
two new species of Macellicephala from the 
deep-sea trenches in the Antarctic: M. tri- 
cornis and M. grandelytris. The latter species 
was later transferred to Bathyfauvelia by 
Levenstein (1978a:77). Levenstein (1975) 
also added the new species Macellicepha- 
loides sandvichensis. Levenstein (1978b) 
added the new species Macellicephala alia 
from the Palau trench and the new genus 



Bathymariana for B. zebra from the Ryuku 
trench. The latter genus was placed in 
Bathyedithinae by Levenstein (1984). 

From the deep Canadian basin in the Arc- 
tic Ocean, Levenstein (1981) added the new 
genus Bathypolaria for B. carinata in Ma- 
cellicephalinae; the new species Bathyedi- 
thia tuberculata in Bathyedithinae; and the 
new genera Bathymiranda for B. microceph- 
ala and Bathycanadia for B. diaphana in 
Polaruschakovinae. In the same subfamily, 
Loshamn (1981) added the new genus Di- 
placonotum for D. paucidentatum (Eliason, 
1962) from the Skagerrak. 

From the Japanese trench, Levenstein 
(1982a) added a new species of Macelli- 
cephaloides, M. villosa. From the Tasman 
trench, Levenstein (1982b) included a key 
to the genera of Macellicephalinae and added 
two new genera: Bathytasmania for B. in- 
solita, and Bathynotalia for B. perplexa. The 
latter genus and species were referred to 
Harmothoinae by Pettibone (1985a: 130), 
based on the figure of the prostomium 
showing long lateral antennae (not men- 
tioned in the text): the prostomium is har- 
mothoid, with cephalic peaks; prostomial 
lateral antennae are lacking in Macelli- 



138 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



cephalinae. From the Kurile-Kamchatka 
trench, Levenstein (1983) added a new 
species of Macellicephaloides, M. improvi- 
sa, and included a key to the seven species 
of the genus. 

Levenstein (1984) summarized the dis- 
tributional patterns for the 19 genera and 
38 species of polynoid polychaetes belong- 
ing to the six deep-sea subfamilies covered 
in Pettibone (1976), along with the addi- 
tional genera and species indicated above. 
They were widely distributed on the abyssal 
plains and in the deep-sea trenches. The 
Macellicephalinae had the greatest number 
of genera (10) and species (21). Three 
subfamilies were represented by fewer gen- 
era and species: Macellicephaloidinae (one 
genus, seven species), Bathyedithinae (two 
genera, three species), and Polaruschako- 
vinae (four genera, five species). Two 
subfamilies were monotypic: Macelloidinae 
and Bathymacellinae. These groups were 
defined as ancient or primary abyssal be- 
cause all the representatives lived only in 
abyssal depths except for a few that ascend- 
ed secondarily. Members of the Macelli- 
cephalinae showed the greatest vertical range 
and panoceanic distributions, mostly in the 
Northern Hemisphere. This group appeared 
to be the most ancient of the abyssal poly- 
noids. 

Since the summary of the group by Lev- 
enstein (1984), additional new subfamilies, 
genera, and species have been added, chiefly 
in connections with studies on the hydro- 
thermal vents in the northeastern Pacific, 
deep seep-sites in the Gulf of Mexico, and 
deep wood panel and cave studies in the 
northwestern Atlantic. 

From the hydrothermal rift-area off west- 
ern Mexico at 2 1°N, Pettibone (1983) added 
the new subfamily Lepidonotopodiinae for 
Lepidonotop odium flmbriatum. From the 
Galapagos vent, the new subfamily Bran- 
chipolynoinae for Branchipolynoe symmy- 
tilida, commensal with the deep-sea mussel, 
was added by Pettibone (1984a). Also from 
the vents, Pettibone (1 984b) added two new 



species of Lepidonotopodium: L. riftense and 
L. williamsae and included a key to the 
species of the genus. 

From the North Atlantic, Pettibone 
(1985a) added to Macellicephalinae two new 
genera: Pelagomacellicephala for P. iliffei, 
from a cave in the Bahamas, and Bathy- 
bahamas for B. charleneae from off the Ba- 
hamas, associated with experimental wood 
panels, and a new species of Bathykerma- 
deca, B. turnerae, from off the Virgin Is- 
lands, also associated with wood panels. 

Pettibone ( 1 985b) added the new subfam- 
ily Branchiplicatinae for Branchiplicatus 
cupreus from the vents off western Mexico 
at 21°N. From the vents off the Galapagos 
and western Mexico, Pettibone (1985c) 
added the third new subfamily having ar- 
borescent branchiae, Branchinotogluminae 
for two new genera: Branchinotogluma, with 
three new species: B. hessleri, B. sandersi, 
and B. grasslei; and Opisthotrochopodus for 
O. alvinus. A key to the three branchiate 
subfamilies was included. 

Pettibone (1985d) added to Macelli- 
cephalinae the new genus Levensteiniella for 
L. kincaidi, and the new species Macelli- 
cephala galapagensis from the Galapagos, 
and from the Santa Catalina Channel, the 
new genus Natopolynoe for N. kensmithi. 
Included also was a summary of the poly- 
noid polychaetes collected by DSRV Alvin 
in the Galapagos Rift in 1979 and East Pa- 
cific Rise at 21 °N in 1982, along with a key 
to the subfamilies, genera, and species. 

From the deep seep-sites at the Florida 
escarpment in the Gulf of Mexico, Petti- 
bone (1986) added a new species of Bran- 
chipolynoe, B. seepensis, commensal in the 
mantle cavities of the Gulf of Mexico mus- 
sels, along with a key to the two species of 
the genus. 

From the hydrothermal vents in the 
Northeast Pacific Explorer and Juan de Fuca 
Ridges, Pettibone (1988) added two new 
species: Lepidonotopodium piscesae in the 
Lepidonotopodiinae, and Opisthotrochop- 
odus tunnicliffeae in Branchinotogluminae. 



VOLUME 102, NUMBER 1 



139 



A new species of Macellicephala from the 
Antarctic, M. australis, was added by Wu 
and Wang (1987). 

Thus to the summary by Levenstein 
(1984), four new subfamilies have been 
added (total 10), nine new genera (total 28), 
and 20 new species (total 58). Four of the 
new genera were added to Macellicephali- 
nae (total 14) and seven new species (total 
28). Two of the new species were referred 
to Macellicephala, one to Bathykermadeca. 

Additional polynoids were collected by 
the Mariana Back-Arc Basin Expedition 
during April and May 1987, where active 
hydrothermal vents were found in the west- 
ern Pacific off the Mariana Islands about 
18°N and 144°E. The polynoid polychaetes 
were collected during dives of DSRV Alvin 
and sent to me for study by Robert Hessler, 
along with information on the collecting 
sites. Four species of polynoids were col- 
lected on two Alvin dives: dive 1836, re- 
ferred to as Snail Pits in Burke Field (1 
species), and dive 1843, designated as Alice 
Springs (3 species). The polynoids agree with 
some of the subfamilies and genera that were 
previously described from the hydrother- 
mal vents in the eastern Pacific. 

Burke Field (Alvin dive 1836, 27 Apr, 
18°10.9'N, 144°43.2'E, 3660 m) was an ex- 
tensive area of low temperature vents at the 
southern end of an active spreading zone 
and dominated by pillow basalts. The Snail 
Pits vent had a water temperature of 1 5°C, 
with small, dense aggregations of "hairy" 
snails that clogged the vent openings. The 
hot vent water was crystal clear. The single 
polynoid collected is described below under 
Branchinotogluminae: Branchinotogluma 
burkensis, new species. 

Alice Springs {Alvin dive 1843, 4 May, 
18°12.6'N, 144°42.4'E, 3640 m) was an ex- 
tensive area dominated by pillow basalts 
with sharp cliffs up to 40 m high. Some low 
temperature vent sites (10-25°C) were char- 
acterized by cloudy, particle-filled water and 
dense concentrations of white anemones. 
Some high temperature vents (up to 250°C) 



had dense aggregations of "hairy" snails, 
crabs, and shrimps, with emerging crystal 
clear water. The three species of polynoids 
collected are described below under three 
subfamilies: Macellicephalinae: Levenstei- 
niella raisae, new species (1 specimen): Lep- 
idonotopodinae: Lepidonotop odium minu- 
tum, new species (9 specimens); and 
Branchinotogluminae: Opisthotrochopodus 
marianus, new species (12 specimens). 

The types are deposited in the National 
Museum of Natural History, Smithsonian 
Institution (USNM). 

Subfamily Macellicephalinae 

Hartmann-Schroder, 1971 

emended Pettibone, 1976 

Levensteiniella Pettibone, 1985 

Levensteiniella was erected for the single 
species L. kincaidi from off western Mexico 
at 21°N and the Galapagos Rift (Pettibone 
1985d:741). An additional species is added 
from the Mariana Back- Arc Basin. 

Levensteiniella raisae, new species 
Figs. 1, 2 

Material examined. —Western Central 
Pacific in Mariana Back- Arc Basin, Alvin 
dive 1843, 4 May 1987, Alice Springs, 
18°12.6'N, 144°42.4'E, 3640 m, R. Hessler, 
collector, holotype (USNM 118362). 

Description. —Length of holotype 2 1 mm, 
width 1 1 mm with setae, segments 27, last 
one very small (Fig. ID). Body flattened, 
tapering anteriorly and more so posteriorly, 
with parapodia about as long as body width. 
Elytra 1 1 pairs, on segments 2, 4, 5, 7, con- 
tinuing on alternate segments to 21, large, 
covering dorsum, oval to subreniform; most 
of surface covered with unusual type of 
macro- and micro-tubercle-papillae, with 
chitinous bases and bulbous papillar tips, 
larger ones on posterior border; bases cov- 
ered with brownish foreign material and 
bacterial "hairs" (Fig. 1G). Elytrophores 
large and prominent (Figs. 1A, C, 2 A, D). 
Dorsal cirri on segments lacking elytra, with 
cylindrical cirrophores on posterodorsal 



140 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Levensteiniella raisae, holotype, USNM 1 18362: A, Dorsal view of anterior end; B, Ventral view of 
anterior end, bases of palps only shown; C, Dorsal view of left half of segments 8 (cirrigerous) and 9 (elytragerous); 
D, Dorsal view of posterior end, including parapodia of segments 24-27, last one very small; E, Ventral view 
of left side of segments 11-13 showing ventral papillae on segments 1 1 and 12, tip of long papilla on segment 
12 hidden from view; F, Dorsal row of border papillae and jaws of pharynx (cut open), inner view; G, Left 
elytron, with detail of macro- and micro-tubercle-papillae (not to scale). Scales =1.0 mm for A-E; 0.5 mm for 
F; 2.0 mm for G. 



faces of notopodia; long styles extending be- 
yond neurosetae, with filamentous tips; dor- 
sal tubercles prominent, nodular to truncate 
(Figs. 1C, D, 2B, C). Dorsal transverse cil- 
iated ridges, up to five per segment, con- 
tinuing onto bases of elytrophores and dor- 
sal tubercles (Fig. 1A, C, D). 



Prostomium oval, deeply bilobed: lobes 
projecting anteriorly, with frontal filaments; 
ceratophore of median antenna short, oval, 
inserted in anterior notch, with style short, 
tapered, shorter than palps; palps stout, long, 
tapered; lateral antennae and eyes lacking 
(pair of tannish spots resembling "eyes") 



VOLUME 102, NUMBER 1 



141 




Fig. 2. Levensteiniella raisae, holotype, USNM 118362: A, Right elytragerous parapodium of segment 2, 
anterior view, acicula dotted; B, Right cirrigerous parapodium of segment 3, posterior view; C, Same, from 
segment 8; D, Right elytragerous parapodium of segment 9, anterior view, acicula dotted; E, Notosetae; F, 
Supraacicular neuroseta, with detail of parts; G, Subacicular neurosetae. Scales =1.0 mm for A-D; 0. 1 mm for 
E-G. 



(Fig. 1A). Tentacular segment not distinct 
dorsally; tentaculophores lateral to prosto- 
mium, lacking setae; each with pair of ten- 
tacular cirri, dorsal one about as long as 
palps, ventral one shorter, both with long 
slender tips; large bilobed facial tubercle 
medial to palps (Fig. 1A, B). 

Second or buccal segment bearing first pair 
of elytrophores, biramous parapodia, with 
notopodium shorter than neuropodium; 
ventral buccal cirri similar to tentacular cir- 
ri, longer than following ventral cirri (Figs. 
1 A, B, 2A). Pharynx not extended (cut open); 
seven pairs of border papillae, none elon- 
gated; two pairs of light amber-colored, en- 
tire jaws (not denticled) (Fig. IF). 

Both rami of biramous parapodia well de- 



veloped, notopodium on dorsoposterior side 
of larger neuropodium, rounded basally with 
projecting acicular process on lower side; 
neuropodium with conical presetal lobe with 
projecting acicular process, postsetal lobe 
shorter, rounded, deeply cut on dorsal side 
(Fig. 2B-D). Notosetae very numerous, 
forming radiating bundle, short to longer, 
much stouter than neurosetae, straight to 
slightly curved, tapering to bare blunt tips, 
with widely spaced spines along one side, 
variable in number (1-18) (Fig. 2E); tips 
often covered with reddish-brown foreign 
material and bacterial "hairs." Neurosetae 
slender, numerous, forming fan-shaped 
bundle; supraacicular neurosetae very slen- 
der, with longer spinous regions, consisting 



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Fig. 3. Lepidonotopodium minutum, A-F, holotype, USNM 1 18363; G, paratype, USNM 1 18364: A, Dorsal 
view of anterior end, right dorsal and left ventral tentacular cirri missing; B, Right 1st elytron from segment 2, 
with detail of micropapillae; C, Right 2nd elytron from segment 4; D, Right 6th elytron from segment 1 1, with 
detail of micropapillae; E, Right 10th elytron from segment 19; F, Right 11th elytron from segment 21; G, 
Ventral view of extended pharynx. Scales = 0.5 mm for A; 1.0 mm for B-F; 0.2 mm for G. 



of two rows of long, widely spaced spines 
along borders, and with shorter spines near 
tapered tips (Fig. 2F); subacicular neuro- 
setae with shorter spinous regions, stouter, 
with minute, close-set spines along one bor- 
der, and with slightly hooked bare tips (Fig. 
2G). Ventral cirri short, tapered, attached 
on middle of neuropodia (Fig. 2C, D). 

Anus dorsal, medial to parapodia of last 
two segments (26, 27); pygidium small, rect- 
angular, with pair of long anal cirri (Fig. IE). 
Two pairs of long ventral papillae on seg- 
ments 11 and 12, their tips sometimes ex- 
tending dorsally between parapodia (Fig. 
IE). 

Etymology. — The species, as well as the 
genus, is named for Raisa J. Levenstein in 
recognition of her numerous contributions 
to the study of the deep-sea polychaetes. 

Remarks.— Levensteiniella raisae differs 
from L. kincaidi in the form of the elytra, 
where the surfaces are nearly covered with 
unusual macro- and micro-tubercle-pa- 
pillae; in L. kincaidi the elytra are more 
delicate with only scattered micropapillae. 
The notosetae differ in having more distinct 
and fewer spines along the border, rather 
than more numerous spinous rows in L. 
kincaidi. 



Subfamily Lepidonotopodiinae 

Pettibone, 1983 

Lepidonotopodium Pettibone, 1983 

Lepidonotopodium was erected for L.fim- 
briatum Pettibone, 1983 from the East Pa- 
cific Rise at 21°N. Two additional species 
were added by Pettibone (1984b) from the 
Galapagos Rift, as well as 21°N: L. william- 
sae and L. riftense, and a fourth species from 
the North East Pacific Explorer and Juan de 
Fuca Ridges by Pettibone (1988), L. pisce- 
sae. An additional species is added from the 
Mariana Back-Arc Basin. 

Lepidonotopodium minutum, new species 
Figs. 3, 4 

Material— Western Central Pacific in 
Mariana Back- Arc Basin, Alvin dive 1843, 
4 May 1987, Alice Springs, 18°12.6'N, 
144°42.4'E, 3640 m, R. Hessler, collector, 
holotype (USNM 1 18363) and 8 paratypes 
(USNM 118364, 118365). 

Description. —Holotype 7.5 mm long, 3.5 
mm wide with setae, with 23 segments; 
slightly smaller paratypes 6-6.5 mm long, 
3-4 mm wide, with 22-23 segments; small- 
est paratype 5.2-5.5 mm long, 3-3.2 mm 
wide, with 21-22 segments. Body short, 



VOLUME 102, NUMBER 1 



143 




Fig. 4. Lepidonotopodium minutum, holotype, USNM 118363: A, Right elytragerous parapodium from 
segment 2, anterior view; B, Right cirrigerous parapodium from segment 3, posterior view; C, Right elytragerous 
parapodium from segment 9, anterior view; D, Right cirrigerous parapodium from segment 10, posterior view; 
E, Right elytragerous parapodium from segment 1 1 , posterior view, notopodium and neuropodium separated, 
acicula dotted; F, Upper, middle and lower notosetae; G, Short anterior notosetae; H, Upper and lower neurosetae. 
Scales = 0.2 mm for A-E; 0.1 mm for F-H. 



suboval in outline, flattened dorsoventrally 
and slightly tapered anteriorly and poste- 
riorly. Elytra 1 1 pairs, on segments 2, 4, 5, 
7, continuing on alternate segments to 21, 
attached eccentrically on prominent elyt- 
rophores (Figs. 3A, 4A, C, E). Elytra oval 
to subreniform, imbricated, covering dor- 
sum, variable in size, last pair very small, 
opaque, smooth, with branching "veins" 
emanating from place of attachment to elyt- 
rophores. Surface of first pair of elytra and 
posterior half on rest of elytra covered with 
scattered micropapillae, globular with short 
to longer tapered or cylindrical bases (Fig. 
3B-F). Elytral surface and borders covered 
with foreign material and bacterial "hairs." 
Dorsal cirri witih cylindrical cirrophores at- 
tached on dorsoposterior sides of notopo- 
dia; styles tapered, extending beyond tips of 
neurosetae: dorsal tubercles elongated, in- 
flated (Fig. 4B, D). 



Prostomium deeply bilobed, anterior 
lobes subtriangular with small frontal fila- 
ments; ceratophore of median antenna in 
anterior notch, short, bulbous, with short, 
subulate style; palps cylindrical, with ta- 
pered tips, nearly two tirnes longer than pro- 
stomium; lateral antennae and eyes lacking 
(Fig. 3A). Tentacular segment not visible 
dorsally; tentaculophores lateral to prosto- 
mium, achaetous, each with pair of tenta- 
cular cirri, dorsal ones nearly as long as palps, 
ventral ones slightly shorter, both with long 
tapering tips (Fig. 3A). Without distinct fa- 
cial tubercle. 

Second or buccal segment with first pair 
of elytrophores, biramous parapodia, and 
ventral or buccal cirri attached basally on 
prominent cirrophores lateral to ventral 
mouth, with styles similar to tentacular cir- 
ri, longer than following ventral cirri (Figs. 
3 A, 4A). Opening of extended pharnyx en- 



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Table 1.— Comparison of Lepidonotopodium rif- 
tense and L. minutum. 





L. riftense 


L. minutum 


Borders of 


with short pa- 


lack papillae 


notopodi- 


pillae 




al bracts 






Papillae of 


7-9 pairs, none 


7 pairs, unequal, 


pharynx 


elongated 


some elongated 


Jaws of 


with numerous 


smooth, without 


pharynx 


teeth 


teeth 


Long ven- 


2 pairs on seg- 


not present 


tral pa- 


ments 1 1 & 




pillae 


12 (on some 
specimens) 





circled with seven pairs of unequal-sized pa- 
pillae; dorsal with three medial ones long, 
tapered, and two short lateral pairs; ventral 
with three middle ones and lateral pair short, 
next to lateral pair long, tapered; two pairs 
of hooked jaws, without teeth or serrations 
(Fig. 3G). 

Biramous parapodia with shorter noto- 
podia on anterodorsal sides of longer neu- 
ropodia (Fig. 4A-D). Notopodia subcorn- 
eal, with projecting acicular lobe, hidden 
anteriorly by very numerous notosetae; well- 
developed flaring bracts enclosing bases of 
notosetae; ventro-anterior and dorso-pos- 
terior bracts attached basally to acicular lobe 
(Fig. 4 A, C, E). Neuropodia with conical 
presetal acicular lobe and shorter, rounded 
postsetal lobe (Fig. 4B-E); distal borders with 
bacterial "hairs." Notosetae very numer- 
ous, forming thick radiating bundle, stouter 
than neurosetae, with widely spaced spines 
along one side; notosetae arranged in four 
groups: upper (about 8), middle (11), lower 
(8) (Fig. 4F), and short anterior group (Fig. 
4G). Notosetae covered with foreign ma- 
terial and bacterial "hairs." Neurosetae nu- 
merous, forming fan-shaped bundle; neu- 
rosetae with two rows of numerous spines 
along one side and slightly hooked bare tips; 
upper neurosetae longer, with longer spi- 
nous regions (Fig. 4H). Ventral cirri at- 
tached on middle of posterior face of neu- 
ropoda, short, tapered (Fig. 4B-E). 



Segmental or nephridial papillae not ob- 
vious. Without elongated ventral papillae 
on segments 11 and 12. Pygidium small, 
rectangular, between posterior parapodia, 
with pair of long anal cirri. 

Etymology.— The specific name minu- 
tum refers to the relatively small size of the 
species. 

Remarks. —Lepidonotopodium minutum 
is closest to L. riftense. Both are of small 
size with relatively few segments (up to 7 
mm in length and 23 segments). The elytra 
lack macrotubercles and micro-tubercles but 
have clavate micropapillae, differing from 
the other species of Lepidonotopodium. The 
notosetae have widely spaced spines along 
one side (also found in L. fimbriatum). The 
two species may be separated by the char- 
acters indicated in Table 1 . 

Subfamily Branchinotogluminae 

Pettibone, 1985 

Branchinotogluma Pettibone, 1985 

Branchinotogluma was established by 
Pettibone (1985c) for three species from the 
hydrothermal vents off western Mexico at 
21°N and the Galapagos: B. hessleri, B. san- 
dersi, and B. grasslei, with B. hessleri as the 
type species. B. grasslei and B. sandersi were 
also reported from the North Pacific Ex- 
plorer and Juan de Fuca Ridges by Petti- 
bone (1988). An additional species is added 
from the Mariana Back-Arc Basin. 

Branchinotogluma burkensis, new species 
Figs. 5, 6 

Material— Western Central Pacific in 
Mariana Back-Arc Basin, Alvin dive 1836, 
27 Apr 1987, Burke Field, Snail Pits, 
18°10.9'N, 144°43.2'E, 3660 m, R. Hessler, 
collector, holotype (USNM 118366). 

Description. —Length of holotype 30 mm, 
width with setae 1 6 mm, segments 2 1 , with 
additional small parapodium on right side 
(Fig. 5B). Body flattened, tapering anteriorly 
and posteriorly, with parapodia longer than 
body width. No color except for golden-col- 
ored setae. Dorsum with transverse ciliated 



VOLUME 102. NUMBER 1 



145 




Fig. 5. Branchinotogluma burkensis. holoiype. L'SNM 118366: A. Dorsal view of anterior end: B. Dorsal 
view of posterior end including segments 18-21: C. Ventral view of left side of segments 11-13. showing small 
segmental ventral papillae: D. Right 1st elytron from segment 2: E. Right 7 th elytron from segment 13: F. Dorsal 
view of left side of cirrigerous segment 14. only base of style of dorsal cirrus shown: G. One of jaws removed 
from body. H. Right elytragerous parapodium from segment 2. anterior view, acicula dotted. Scales = 2.0 mm 
for A-E: 0.5 mm for F. G: 1.0 mm for H. 



bands. 2— I per segment, extending onto 
elytrophores. dorsal tubercles, and bran- 
chial bases (Pig. 5A. B. F). 

Elytra 10 pairs, attached on large bulbous 
elytrophores on segments 2. 4. 5. ~. con- 
tinuing on alternate segments to 19 (Pigs. 
5 A. B. H. 6B). Elytra large, overlapping, 
covering dorsum, round to oval, delicate, 
with branched "veins."" without tubercles or 
papillae (Pig. 5D. E). Dorsal cirri on non- 
elytragerous segments, with cylindrical cir- 
rophores on posterodorsal sides of noto- 



podia. and long smooth styles with slender 
tips extending beyond tips of neurosetae: 
dorsal tubercles nodular, projecting (Pigs. 
5B. F. 6 A. O. Branchiae arborescent, in two 
groups: on dorsal bases of notopodia and 
on lateral sides of elytrophores and dorsal 
tubercles (Figs. 5F. 6B). Branchiae begin- 
ning on segment 3 as two small groups (Pig. 
6A). becoming larger (Figs. 5F. 6B. O. con- 
tinuing to posterior end and becoming very 
small (Fig. 5B>. 

Bilobed prostomium with prominent cy- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Branchinotogluma burkensis, holotype. USNM 118366: A. Right cirrigerous parapodium from seg- 
ment 3. posterior view: B. Right ehlragerous parapodium from segment 9. anterior view, acicula dotted: C. 
Right cirrigerous parapodium from segment 10. posterior view; D, notosetae: E. Tip of long supraacicular 
neuroseta. with detail of parts: F. Subacicular neuroseta. with detail of parts. Scales =1.0 mm for A-C; 0.1 mm 
for D-F. 



lindrical anterior lobes with terminal fila- 
ments; median antenna with bulbous cer- 
atophore in anterior notch and short style 
tapered to slender tip; palps long, tapered. 



smooth, about three times length of prosto- 
mium; without eyes, but non-ocular paired 
oval shaded areas present (Fig. 5A). Ten- 
tacular segment not distinct dorsally; ten- 



VOLUME 102, NUMBER 1 



147 



taculophores lateral to prostomium, achae- 
tous, each with pair of dorsal and ventral 
tentacular cirri about as long as palps, dorsal 
tentacular cirri slightly longer than ventral 
ones (Fig. 5A). Second or buccal segment 
with first pair of large elytrophores, bira- 
mous parapodia, and long, slender buccal 
cirri similar to tentacular cirri, longer than 
following ventral cirri; notopodium with 
prominent bract fused with acicular lobe (not 
distinct, as usual in genus) and small bundle 
of stout notosetae; neuropodium similar to 
following (Fig. 5A, H). Pharynx not everted 
(cut out), with five border papillae around 
opening: three dorsal and two ventral; two 
pairs of prominent dark reddish amber-col- 
ored jaws, minutely denticled on inner bor- 
der (about 50) (Fig. 5G). 

Biramous parapodia with notopodia 
shorter than neuropodia, both rami with 
golden-colored setae (Figs. 5H, 6A-C). No- 
topodia conical, with prominent projecting 
acicular processes, without notopodial bracts 
except on segment 2 with bract fused to acic- 
ular process. Notosetae numerous, forming 
radiating bundles, not as long as but much 
stouter than neurosetae, straight, tapered, 
acicular, shorter ones smooth, longer ones 
with some spines near distal tips (Fig. 6D). 
Larger and longer neuropodium with sub- 
conical presetal acicular lobe with project- 
ing acicular process and shorter rounded 
postsetal lobe, lobes deeply separated dor- 
sally and ventrally. Neurosetae very nu- 
merous, forming fan-shaped bundles, slen- 
der, with slightly hooked tips. Supraacicular 
neurosetae with two rows of widely spaced 
prominent spines and finely spinous hood 
on slightly hooked tip (Fig. 6E).Subacicular 
neurosetae finely spinous along border, with 
slightly hooked tip (Fig. 6F). Ventral cirri 
short, tapering, attached on middle of neu- 
ropodia (Fig. 6A-C). 

Posterior end not modified or com- 
pressed. Pygidium small lobe between para- 
podia of last segment (21), with dorsal anus 
and pair of anal cirri (Fig. 5B). Segmental 
ventral papillae seven pairs, small squarish, 



on ventral bases of parapodia of segments 
1 1-17, none elongated (Fig. 5C). 

Etymology. —The species is named bur- 
kensis for the collecting area, Burke Field 
on the Mariana Back-Arc Basin. 

Remarks.— B. burkensis is closest to B. 
grasslei Pettibone (1 985c). Both species have 
a pair of anal cirri and posterior segments 
that are not compressed or modified; bran- 
chiae are present on all segments from 3 on; 
and both lack elongated ventral papillae. B. 
burkensis has seven pairs of small segmental 
ventral papillae on segments 11-17, com- 
pared to five pairs on segments 11-15 in B. 
grasslei. In B. burkensis, the notopodial bract 
on segment 2 is fused with the acicular lobe 
and not distinct, as in B. grasslei, where the 
notopodial bract encloses the acicular lobe 
and notosetae (Pettibone 1985c, fig. 6A). 
The long notosetae in B. burkensis have dis- 
tal spines, whereas they are smooth in B. 
grasslei. 

Opisthotrochopodus Pettibone, 1985, 
emended 

Opisthotrochopodus was established by 
Pettibone (1985c) for O. alvinus from the 
hydrothermal vents off western Mexico at 
21°N and the Galapagos. O. tunnicliffeae, 
from the hydrothermal vents of the Ex- 
plorer and Juan de Fuca Ridges, was added 
by Pettibone (1988). A third species is added 
from the Mariana Back-Arc Basin and the 
genus is emended. 

Diagnosis. — Body short, with 21 seg- 
ments, first achaetous. Elytra 10 pairs, on 
segments 2, 4, 5, continuing on alternate 
segments to 19. Dorsal cirri with short cir- 
rophores and long style, and dorsal tuber- 
cles, in line with elytrophores, on segments 
3, 6, continuing on alternate segments to 20 
and 2 1 . Arborescent branchiae two pairs per 
segment, attached to lateral sides of elyt- 
rophores, dorsal tubercles, and dorsal side 
of notopodia, beginning on segment 3 and 
continuing to segment 15-18. Prostomium 
bilobed, with triangular anterior lobes bear- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. Opisthotrochopodus marianus, A-I, paratype, USNM 1 18369; J, holotype, USNM 1 18367: A, Dorsal 
view of anterior end; B, Ventral view of segments 12-18; C, Right 1st elytron from segment 2; D, Right 2nd 
elytron from segment 4; E, Right 4th elytron from segment 7; F, Right 6th elytron from segment 1 1; G, Right 
9th elytron from segment 17; H, Right 10th elytron from segment 19; I, Ventral view of extended pharynx; J, 
Dorsal jaws, inner view. Scales = 0.5 mm for A, B, I; 0.5 mm for C-H; 0.1 mm for J. 



ing frontal filaments, with ceratophore of 
median antenna in anterior notch, with 
paired ventral palps, without lateral anten- 
nae or eyes. First or tentacular segment not 
visible dorsally; tentaculophores lateral to 
prostomium, achaetous, each with dorsal 
and ventral tentacular cirri. Second or buc- 
cal segment with first pair of elytra, bira- 
mous parapodia, and ventral or buccal cirri 
attached to basal parts of parapodia lateral 
to mouth; styles longer than following ven- 
tral cirri. Parapodia biramous, with noto- 
podia shorter than neuropodia. Notopodia 
of elytragerous segments 2-17 with promi- 
nent rounded bracts enclosing conical acic- 
ular lobes and notosetae, only on segment 
2 or lacking. Neuropodia with longer con- 
ical presetal and shorter rounded postsetal 
lobes. Notosetae stouter than neurosetae, 
straigjit, acicular, smooth or with spines. 
Neurosetae long, slender, finely spinous, 
with slightly hooked tips. Ventral cirri short, 
tapered. Posterior 3 to 6 segments modified, 
including wheel organs on segment 20 or 
lateral lamellae on segment 2 1 . Pharynx with 
5 to 8 border papillae; 2 pairs of dorsal and 
ventral hooked jaws, with or without den- 



ticles. Elongated ventral papillae on seg- 
ment 12 and rounded ventral lamellae on 
some following segments. Pygidium with or 
without pair of anal cirri. 

Opisthotrochopodus marianus, new species 
Figs. 7-9 

Material.— West Central Pacific, Mar- 
iana Back- Arc Basin, Alvin dive 1843,4 May 
1987, Alice Springs, 18°12.6'N, 144°22.4'E, 
3640 m, holotype (USNM 118367), para- 
type (USNM 118369), paratype (USNM 
118368), 9 paratypes (3 small, USNM 
118370). 

Description. —Holotype 6 mm long, 3 mm 
wide, with 21 segments. Two adult para- 
types 5-5.5 mm long, 3 mm wide, with 21 
segments. Two young paratypes 2.5-3 mm 
long, 2-3 mm wide, with 13-14 segments 
plus small growing zone. Body flattened, ta- 
pering slightly anteriorly, anterior parapo- 
dia (segments 2-13) longer, tapering grad- 
ually (segments 14-18), narrowing and 
projecting posteriorly with extra long setae 
(segments 19-21) (Figs. 7B, 9H, I). Dorsum 
with transverse ciliated bands, 1-2 per seg- 



VOLUME 102, NUMBER 1 



149 




Fig. 8. Opisthotrochopodus marianus, paratype, USNM 118369: A, Right elytragerous parapodium from 
segment 2, anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 3, posterior view, style 
of dorsal cirrus broken off; C, Left elytragerous parapodium from segment 9, anterior view, acicula dotted; D, 
Left cirrigerous parapodium from segment 10, posterior view; E, Short and long notosetae; F, Supraacicular 
neuroseta; G, Subacicular neurosetae; H, Right elytragerous parapodium from segment 15, anterior view; I, 
Right cirrigerous parapodium from segment 16, posterior view; J, Right elytragerous parapodium from segment 
17, posterior view; K, same, anterior view. Scales = 0.3 mm for A-D, H-J; 0.1 mm for E-G; 0.2 mm for K. 



ment, extending onto elytrophores and dor- 
sal tubercles (Figs. 7A, 9H). 

Elytra and prominent bulbous elytro- 
phores 10 pairs (Figs. 7 A, 8 A, C, H, J, K, 
9H). Elytra large, overlapping, covering 
dorsum, except for posterior modified par- 
apodia, round to oval, larger on middle ely- 
tra, very small on segments 17 and 19, del- 
icate, showing branched "veins" emanating 
from scars of attachment to elytrophores, 
without tubercles or papillae (Figs. 7C-H, 
9H). Dorsal cirri with cylindrical cirro- 
phores on posterodorsal sides of notopodia, 
long smooth styles with slender tips ex- 
tending beyond setae; dorsal tubercles elon- 
gate, tapered, with long cilia (Fig. 8B, D, I). 
Branchiae delicate, arborescent, on lateral 
bases of elytrophores and dorsal tubercles 
and dorsal bases of notopodia, beginning on 



segment 3 as two small groups (Fig. 8B), 
becoming larger (Fig. 8C, D), and continu- 
ing to segment 1 5 as single small group (Fig. 
8H). 

Prostomium bilobed, anterior lobes tri- 
angular with slender frontal filaments; me- 
dian antenna with bulbous ceratophore in 
anterior notch, with slender short style; palps 
stout, tapered, about two times longer than 
prostomium; without eyes (Fig. 7A). Ten- 
taculophores lateral to prostomium, achae- 
tous, with 2 pairs of tentacular cirri, dorsal 
pair longer than palps, ventral pair shorter 
(Fig. 7A). Second or buccal segment with 
first pair of prominent elytrophores, bira- 
mous parapodia, similar to but smaller than 
following: ventral buccal cirri similar to ten- 
tacular cirri, longer than following ventral 
cirri; notopodium without notopodial bract 



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Fig. 9. Opisthotrochopodus marianus, A-G, paratype. USNM 118369; H-I, paratope. USNM 118368: A, 
Right cirrigerous parapodium from segment 18. anterior view, acicula dotted: B. Right elytragerous parapodium 
from segment 19. anterior view, acicula dotted, elytrophore not shown; C, Right cirrigerous parapodium from 
segment 20. anterior view, acicula dotted: D. Tips of long notosetae from same; E, Neuroseta from same; F, 
Left cirrigerous parapodium from segment 2 1 . anterior view, acicula dotted: G. Same, cirrigerous parapodium 
from segment 2 1 . anterior view, acicula dotted: G. Same, enlarged: H. Dorsal view of posterior end including 
segments 19-21. acicula dotted: I. same, ventral view. Scales = 0.3 mm for A-C, F; 0.1 mm for D, E. G; 0.2 
mm for H. I. 



(Figs. 7 A, 8A). Notosetae stout, similar to 
those of following segments: lower neuro- 
setae with capillary tips, rest similar to up- 
per neurosetae of following segments. Ex- 
tended pharynx encircled with four pairs of 
papillae, dorsal with two medial ones larger 
and lateral ones smaller; ventral with four 
small papillae: two pairs of curved reddish 
jaws occupying most of opening, without 
denticles on inner border (Fig. 71. J). 

Parapodia of segments 2-16 biramous. 
with notopodia shorter than neuropodia, 
both rami with light yellow amber-colored 



setae (Fig. 8A-D, H, I). Notopodia rather 
long, conical, without notopodial bracts en- 
closing acicular lobe and notosetae (as in O. 
alvinus). Notosetae numerous, stouter than 
neurosetae. short, smooth and longer, 
smooth or with faint spinous rows, tapered 
to blunt tips (Fig. 8E). Larger neuropodia 
with presetal lobe long, subconical. with 
projecting acicular process, postsetal lobe 
shorter, rounded. Neurosetae numerous, 
long, slender, wider basally. with slightly 
hooked tips. Supraacicular neurosetae more 
slender, with longer spinous regions (Fig. 



VOLUME 102, NUMBER 1 



151 



Table 2.— Comparison of three species of Opisthotrochopodus. 





O. alvinus Pettibone, 


O. tunnicliffeae Pettibone, 






1985c (Figs. 7-9) 


1988 (Figs. 6-9) 


O. marinus n. sp. (Figs. 7-9) 


Notopodial bracts on 


segments 2-17 (Figs. 


segment 2 only (Fig. 


without 


elytragerous parapo- 


7A, G) 


6A, C) 




dia 








Branchiae on segments 


3-17 


3-18 


3-15 


Pharynx: 








Papillae 


5 small: 3 dorsal, 2 


same 


8: 4 dorsal, 2 medial 




ventral 




larger; 4 ventral small 


Jaws 


with minute denticles 


same 


without denticles (Fig. 




(Fig. 8H, I) 




71, J) 


Segment 20: 








Notopodia 


achaetous (Fig. 


short notosetae (Figs. 


long notosetae 




9B, F-L) 


8B, C, 9A-F) 


(Fig. 9C-E) 


Neuropodia 


with "wheel" organ, 


with "balloon" organ 


long, slender, acicular 




incl. stout, acicular 


including stout har- 


lobe & short capillary 




& slender spinous 


poon & slender spi- 


neurosetae 




neurosetae 


nous neurosetae 




Ventral cirrus 


without 


present 


without 


Segment 2 1 : 








Lamellae 


small notopodial 


without (Fig. 9C) 


large lateral neuro- 




& neuropodial 




podia (Fig. 9G-I) 




(Fig. 9M, N) 






Neurosetae 


few, short 


same 


long capillary 


Ventral cirrus 


without 


present 


without 



8F). Subacicular neurosetae decreasing in 
length ventrally, finely spinous more basal- 
ly, with longer spines distally; few shorter 
lower ones with capillary tips (Fig. 8G). 
Ventral cirri short, tapered, attached on 
middle of neuropodia (Fig. 8B-D, H, I). 

Elongated ventral papillae on segment 1 2, 
extending posteriorly to segment 15 (Fig. 
7B); 6 pairs of short, flat ventral lamellae 
on segments 13-18, ciliated on upper side 
(Figs. 7B, 8H, J, K). 

Parapodia from segment 15 on more or 
less modified. Parapodium 15 with only 
small branchia on lateral side of elytro- 
phore; notopodium almost as long as neu- 
ropodium; supraacicular neurosetae with 
capillary tips (Fig. 8H). Parapodium 16, and 
following parapodia, without branchiae; all 
neurosetae with capillary tips (Fig. 81). Par- 
apodium 1 7 with small neuropodium short- 
er than ventral cirrus, with bundle of long 



slender capillary neurosetae; notopodium 
small, with three shorter and seven longer 
stout notosetae extending to tip of neuro- 
podium (Fig. 8J, K). 

Parapodium 18 with small notopodial 
acicular lobe fused to cirrophore of dorsal 
cirrus, ciliated on lower side, with few (4) 
short stout notosetae on upper side; conical 
neuropodium shorter than ventral cirrus, 
ciliated on upper side, with small bundle of 
long capillary neurosetae (Fig. 9A). 

Parapodium 19 with elytrophore and 
small elytron, inflated oval notopodium with 
short acicular lobe and few (3-4) short stout 
notosetae near tip, ciliated on underside; 
neuropodium similar to preceding parapo- 
dium, with small bundle of long capillary 
neurosetae (Fig. 9B, H, I). 

Parapodium 20 directed posteriorly, with 
small notopodial acicular lobe fused to cir- 
rophore of dorsal cirrus; neuropodium with 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



long slender acicular lobe nearly as long as 
dorsal cirrus, with short ventral cirrus (Fig. 
9C, H, I). Long notosetae wider subdistally, 
spinous, tapering to slender tips (Fig. 9D). 
Shorter neurosetae slender, wider subdis- 
tally, spinous, tapering to capillary tips (Fig. 
9E). 

Parapodium 2 1 directed posteriorly, with 
notopodial acicular lobe fused to cirrophore 
of dorsal cirrus, without notosetae; neuro- 
podial conical acicular lobe with slender 
capillary neurosetae and large, oval, delicate 
lateral lamella; without ventral cirrus (Fig. 
9F-I). Pygidium small rounded lobe with 
dorsal anus medial to parapodia of segment 
2 1 , and pair of long anal cirri (Fig. 9H, I). 

Etymology.— The specific name mari- 
anus is based on the collecting site of the 
Mariana Back-Arc Basin. 

Remarks. — O. marianus can be separated 
from the other two species of Opisthotro- 
chopodus according to Table 2. 

Acknowledgments 

My thanks go to the members of the Mar- 
iana Back-Arc Basin Expedition, the crew 
of the Alvin, and especially Robert R. Hes- 
sler and Michel A. Boudrias, who sent me 
the polynoid specimens and furnished sta- 
tion data and information on the collecting 
area. The manuscript benefited from the re- 
views by James A. Blake and Jerry D. Ku- 
denov. 



Literature Cited 

Levenstein, R. J. 1975. [The polychaetous annelids 
of the deep-sea trenches of the Atlantic sector 
of the Antarctic Ocean.] — Trudy Institut 
Okeanologii P.P. Shirshov Academii Nauk SSSR 
103:119-142 [in Russian, English summary]. 

. 1 978a. [Annelida (Polychaeta) from the deep 

waters of the Pacific region of Antarctica.] — 
Trudy Institut Okeanologii P.P. Shirshov Aca- 
demii Nauk SSSR 113:73-87 [in Russian, En- 
glish summary]. 

. 1978b. [Polychaetes of the family Polynoidae 

(Polychaeta) from the deep-water trenches of the 
western part of the Pacific.]— Trudy Institut 



Okeanologii P.P. Shirshov Academii Nauk SSSR 
112:162-174 [in Russian, English summary]. 

. 1981. [Some pecularities in the distribution 

of polychaetes of the family Polynoidae in the 
Canadian basin of the Arctic Ocean.]— Trudy 
Institut Okeanologii P.P. Shirshov Academii 
Nauk SSSR 115:26-36 [in Russian, English 
summary]. 

. 1982a. [On the polychaete fauna of the family 

Polynoidae from the trench of Japan.]— Trudy 
Institut Okeanologii P.P. Shirshov Academii 
Nauk SSSR 117:59-62 [in Russian, English 
summary]. 

. 1982b. [New genera of the subfamily Ma- 

cellicephalinae (Polychaeta, Polynoidae) from 
the Tasman Trench]— Zoologichesky Zhurnal 
6 1(9): 129 1-1 296 [in Russian, English summa- 
ry]. 

. 1983. [Macellicephaloides improvisa sp. n. 

(Polychaeta, Polynoidae) from the Kurile-Kam- 
chatka Trench]— Zoologichesky Zhurnal 62(9): 
1419-1421 [in Russian, English summary]. 

. 1984. On the ways of formation of the deep- 
sea polychaete fauna of the family Polynoidae. 
Pp. 72-85, in P.A. Hutchings, ed., Proceedings 
of the First International Polychaete Confer- 
ence, Sydney, Australia, 1983. The Linnean So- 
ciety of New South Wales. 

Loshamn, A.-A. 1981. Descriptions of five polynoid 
species (Polychaeta) from the coasts of Norway 
and Sweden, including three new species, one 
new genus and one new generic name.— Zoo- 
logica Scripta 10(1):5— 13. 

Pettibone, M. H. 1976. Revision of the genus Ma- 
cellicephala Mcintosh and the subfamily Ma- 
cellicephalinae Hartmann-Schroder (Polychae- 
ta: Polynoidae). —Smithsonian Contributions to 
Zoology 229:1-71. 

. 1979. Redescription of Bruunilla natalensis 

Hartman (Polychaeta: Polynoidae), originally 
referred to Fauveliopsidae.— Proceedings of the 
Biological Society of Washington 92(2):384-388. 

. 1983. A new scale worm (Polychaeta: Poly- 
noidae) from the hydrothermal rift area off 
Western Mexico at 2 1°N.— Proceedings of the 
Biological Society of Washington 96(3):392-399. 

. 1984a. A new scale-worm commensal with 

deep-sea mussels on the Galapagos hydrother- 
mal vent (Polychaeta: Polynoidae).— Proceed- 
ings of the Biological Society of Washington 
97(l):226-239. 

. 1984b. Two new species of Lepidonotopo- 

dium (Polychaeta: Polynoidae: Lepidonotopo- 
dinae) from hydrothermal vents off the Gala- 
pagos and East Pacific Rise at 21°N.— 
Proceedings of the Biological Society of Wash- 
ington 97(4):847-863. 



VOLUME 102, NUMBER 1 



153 



-. 1 985a. Polychaete worms from a cave in the 
Bahamas and from experimental wood panels 
in deep water of the North Atlantic (Polynoidae: 
Macellicephalinae, Harmothoinae). — Proceed- 
ings of the Biological Society of Washington 
98(1): 127-149. 

-. 1985b. An additional new scale worm (Po- 
lychaeta: Polynoidae) from the hydrothermal rift- 
area off Western Mexico at 2 1°N. — Proceedings 
of the Biological Society of Washington 98(1): 
157-164. 

-. 1985c. Additional branchiate scale-worms 
(Polychaeta: Polynoidae) from Galapagos hy- 
drothermal vent and rift-area off Western Mex- 
ico at 2 1°N. — Proceedings of the Biological So- 
ciety of Washington 98(2):447-469. 
-. 1985d. New genera and species of deep-sea 
Macellicephalinae and Harmothoinae (Poly- 
chaeta: Polynoidae) from the hydrothermal rift 
areas of the Galapagos and western Mexico at 
21°N and from the Santa Catalina Channel.— 
Proceedings of the Biological Society of Wash- 
ington 98(3):740-757. 



. 1986. A new scale-worm commensal with 

deep-sea mussels in the seep-sites at the Florida 
Escarpment in the eastern Gulf of Mexico (Poly- 
chaeta: Polynoidae: Branchipolynoinae). — Pro- 
ceedings of the Biological Society of Washington 
99(3):444-451. 

. 1 988. New species and new records of scaled 

polychaetes (Polychaeta: Polynoidae) from hy- 
drothermal vents of the Northeast Pacific Ex- 
plorer and Juan de Fuca Ridges. — Proceedings 
of the Biological Society of Washington 101(1): 
192-208. 

Wu, B. & Wang, Y.-H. 1987. Two new species of 
Polychaeta from South Ocean.— Acta Zootax- 
onomica Sinica 1 2( 1 ):23-29 [in Chinese and En- 
glish]. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 154-168 

POLYNOIDAE AND SIGALIONIDAE (POLYCHAETA) FROM 

THE GUAYMAS BASIN, WITH DESCRIPTIONS OF 

TWO NEW SPECIES, AND ADDITIONAL RECORDS FROM 

HYDROTHERMAL VENTS OF THE GALAPAGOS 

RIFT, 21°N, AND SEEP-SITES IN THE GULF OF 

MEXICO (FLORIDA AND LOUISIANA) 

Marian H. Pettibone 

Abstract.— Eight species of scaled polychaetes are reported from hydrother- 
mal vents of the Guaymas Basin in the Gulf of California: Neoleanira racemosa 
in the Sigalionidae and seven species of Polynoidae, including two new species, 
Bathykurila guaymasensis, in Macellicephalinae, and Macellicephaloides alvini, 
in Macellicephaloidinae. The latter genus and subfamily are reviewed, and a 
Key to the seven species in the genus is provided. Included are additional 
records of some species of Polynoidae from hydrothermal vents from the Ga- 
lapagos and 21°N in the eastern Pacific, as well as seep-sites in the Gulf of 
Mexico Florida Escarpment and off Louisiana. 



A small collection of scaled polychaetes 
from the Guaymas Basin in the Gulf of Cal- 
ifornia or the Sea of Cortez was sent to me 
for study by J. F. Grassle. Hot vents and 
hydrocarbon seeps were discovered in this 
area in 1980 and sampled in 1982 during 
dives of the submersible DSRV Alvin (Lons- 
dale 1984, Grassle 1986). The area, a con- 
tinuation of the East Pacific Rise into the 
Gulf of California, is covered by a thick 
layer of pelagic muddy sulfide sediment. The 
study site consists of mounds of hydrother- 
mal precipitates in the spreading center 
troughs at about 2000 meters. The fauna 
included large numbers of the giant vesti- 
mentiferans, Riftia, giant clams, such as Ca- 
lyptogena pacifica, nuculanid bivalves, al- 
vinellid polychaetes, as Paralvinella grasslei, 
and fluffy mats of the large bacterium Beg- 
giatoa. 

Eight species of scaled polychaetes are re- 
ported herein from the Guaymas Basin, sev- 
en of them belonging to the Polynoidae and 
one to the Sigalionidae: Neoleanira race- 
mosa (Fauchald), previously reported from 
the Guaymas Basin by Fauchald (1972, as 



Sthenolepis). Of the polynoid species, five 
were previously described by me from other 
vent areas: one, Branchiplicatus cupreus, 
from the tropical eastern Pacific at 21°N; 
one, Lepidonotopodium riftense, the same 
area and the Galapagos Rift; and three, 
Branchinotogluma grasslei, B. sandersi, and 
Levensteiniella kincaidi, from the above two 
areas and also the Explorer and Juan de 
Fuca Ridges. The other two polynoid species 
are described below as new species: Bathy- 
kurila guaymasensis, in Macellicephalinae, 
and Macellicephaloides alvini, in Macelli- 
cephaloidinae. The specimens were collect- 
ed during Alvin dives in 1982 and 1985, in 
washings from rock with worms, Bead Ex- 
periment washings, washings from Rock 
Chimney, washings from Riftia, Calypto- 
gena, and alvinellid polychaetes, from Box 
cores taken on the slopes of the mounds, 
some with numerous nuculanid bivalves. 

Some additional records of previously de- 
scribed vent fauna are included in this re- 
port, such as Branchipolynoe symmytilida, 
removed from some Galapagos mussels and 
sent to me in July 1985 by J. F. Grassle. 



VOLUME 102, NUMBER 1 



155 



Also, specimens of Branchipolynoe seepen- 
sis were removed by Ruth Turner from Gulf 
of Mexico mussels collected in the seep-sites 
at the Florida Escarpment and sent to me 
in October 1986, by Barbara Hecker. An 
additional specimen of this species was col- 
lected in May 1987, by the US Navy sub- 
marine NR- 1 and R/V Gyre and sent to me 
by T. H. Perkins. Additional vent polynoids 
from the Galapagos and the eastern Pacific 
Rise at 21°N, collected during Alvin dives 
in 1979 and 1982, chiefly among fauna in 
washings from mussels and crab traps, were 
sent to the Smithsonian Sorting Center for 
further sorting and transferred to me in De- 
cember 1987. They supplement my pre- 
vious studies on this group. 

Types and additional specimens are de- 
posited in the collections of the Department 
of Invertebrate Zoology, National Museum 
of Natural History, Smithsonian Institution 
(USNM). 

Family Polynoidae 

Subfamily Branchipolynoinae Pettibone, 

1984 

Genus Branchipolynoe Pettibone, 1984, 

emended Pettibone, 1986 
Branchipolynoe symmytilida Pettibone 

Branchipolynoe symmytilida Pettibone, 
1984a:227, figs. l-8.-Kenk & Wilson, 
1985:264.-Desbruyeresetal. 1985:104, 
113, 114,-Grassle, 1986:326, 327, 336. 

Material. —East Central Pacific, from 
dives of the Alvin on three vent areas along 
the Galapagos Rift in 1979, associated with 
deep-sea mussels, Branchimodiolus ther- 
mophilics Kenk & Wilson: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 880, 21 Jan, 
2493 m, from 9 mussels, 9 specimens 
(USNM 98577). Dive 888, 13 Feb, 2493 m, 
from mussel, 1 specimen (USNM 98578). 
Dive 986, 3 Dec, 2494 m, mussel bucket 
washings, 1 specimen (USNM 118168). 
Dive 989, 6 Dec, 2482 m, from small mus- 
sels, 5 specimens (USNM 118171). Dive 
991,8 Dec, 2490 m, washings from 68 mus- 



sels in mesh-lined milk crates, 5 specimens, 
(USNM 118170). Garden of Eden, 
00°47.69'N, 86°07.74'W: Dive 883, 25 Jan, 
2482 m, from mussel, 1 specimen (USNM 
98579). Dive 884, 25 Jan, 2482 m, from 6 
mussels, 7 specimens (USNM 98580) and 
3 young specimens (USNM 1 18172). Dive 
993, 10 Dec, 2518 m, slurp sample from 
base of vestimentiferans, 1 specimen 
(USNM 1 18166). Rose Garden, 00°48.25'N, 
86°13.48'W: Dive 892, 17 Feb, 2454 m, 
from 3 mussels, 3 specimens (USNM 
98581). Dive 894, 19 Feb, 2457 m, from 
12 mussels, 12 specimens (USNM 98582). 
Dive 983, 30 Nov, 2457 m, mussel bucket 
residue, 2 young specimens (USNM 
118167). Dive 984, 1 Dec, 2451 m, mussel 
washings left in clam bucket overnight, 1 3 
specimens (USNM 118169); residue from 
retrieved box containing vestimentiferans, 
1 small specimen (USNM 118173). 

Remarks. —Branchipolynoe symmytilida 
has been found living commensally in the 
mantle cavities of the giant deep-sea vent 
mussels, Branchimodiolus thermophilus 
Kenk & Wilson (Pettibone 1984a, Kenk & 
Wilson 1985). Of the 37 polynoids taken 
from 36 mussels (USNM 98577-98582), 1 
had a length of 40 mm; 14, lengths of 29- 
33 mm; 9, lengths of 21-25 mm; 10, lengths 
of 15-20 mm; 2, lengths of 1 1 mm; and 1, 
length of 6 mm. The two polynoids taken 
from a single mussel had lengths of 25 and 
1 5 mm. The additional 3 1 specimens from 
8 Alvin dives, taken mostly from mussel 
washings (USNM 1 18166-73), consisted of 
smaller worms: adults up to 1 1 mm in length, 
3 mm in width, with 20 segments and 10 
pairs of elytra; juveniles 3-4 mm in length, 
1.5-2.2 mm in width, with 14-17 segments; 
and very young specimens 1.2-2 mm in 
length, 1 mm in width, with 11-12 seg- 
ments. 

According to observations by Desbru- 
yeres et al. (1985:104) on B. symmytilida, 
gut contents included pieces of mussel gills, 
coccoid and filamentous bacteria-like cells, 
and a mixture of crustacean molts, uniden- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tifiable worm setae, diatom frustules, and 
amorphous organic matter consistent with 
a mixed diet including living tissue and 
pseudofeces of mussels. Special features are 
their red blood, segmental arborescent 
branchiae, 10 pairs of elytra, and 20-21 seg- 
ments. 

Distribution.— East Central Pacific along 
Galapagos Rift, associated with Galapagos 
deep-sea mussels, in 2451-2518 m. Re- 
ported for 1 3°N by Desbruyeres et al. ( 1 98 5 : 
114). 

Branchipolynoe seepensis Pettibone 

Branchipolynoe seepensis Pettibone, 1986: 

445, figs. 1-4. 
Branchipolynoe sp. Hecker, 1985:470. 

Material. —Gulf of Mexico, from dives of 
the Alvin in Oct 1986, Florida Escarpment, 
26°01'46"N, 84°54'36"W, 3270 m: Dive 
1753, commensal with mussels, 15 speci- 
mens (USNM 10151). Dive 1755, with 
mussels, 19 specimens (USNM 101503). 
Dive 1756, in washings and from broken 
mussels, 25 specimens (USNM 101502). 
Dive 1758, with mussels, 2 juveniles 
(USNM 101504). 

Gulf of Mexico off Louisiana, 27°47'N, 
91°30'W, 1097 m, Fla. Dept. Nat. Res., 5 
Mar, EJ 87-127, 1987, Bush Hill Box 1, 
among mussels, collected by US Navy sub- 
marine NR-1, recovered on surface by 
R/V Gyre, cruise 87-G-2, one specimen 
(USNM 102844). 

Remarks. — The additional 6 1 specimens 
from 4 Alvin dives (USNM 10151-4), as- 
sociated with the unnamed Gulf of Mexico 
giant mussels (to be described and named 
by Ruth Turner and Barry Wilson) include 
adults and juveniles; some are larger than 
previously reported. Larger adults are 31- 
40 mm long, 14-17 mm wide, with 21 seg- 
ments; smaller adults, 9-17 mm long, 4-8 
wide, with 21 segments; larger juveniles, 5- 
9 mm long, 3-4.5 mm wide, with 17-19 
segments; smaller juveniles, 3-4.5 mm long, 



2-2.5 mm wide, with 15-16 segments; and 
the smallest juvenile, 2 mm long, 1.5 mm 
wide, with 14 segments. The single speci- 
men from Bush Hill off Louisiana (USNM 
102844) is 20 mm long, 8 mm wide, with 
2 1 segments; it differs by having long ven- 
tral papillae on segment 12 only, not on 
both segments 11 and 12. The smaller ju- 
veniles lack both pairs of ventral papillae. 
Distribution. —Gulf of Mexico, near hy- 
persaline seeps at base of Florida Escarp- 
ment, associated with Gulf of Mexico mus- 
sels, in 3266-3270 m and off Louisiana, in 
1097 m. 

Subfamily Lepidonotopodiinae Pettibone, 

1983 

Genus Lepidonot op odium Pettibone, 1983 

Lepidonotopodium riftense Pettibone 

Lepidonot op odium riftense Pettibone, 
1984b:852, figs. 2-4. 

Material. —East Central Pacific, from 
dives of the Alvin on 3 vent areas along the 
Galapagos Rift in 1979: Mussel Bed, 
00°47.89'N. 86°09.21'W: Dive 986, 3 Dec, 
2494 m, mussel bucket washings, one spec- 
imen (USNM 118180). Garden of Eden, 
00°47.09'N.86°07.74'W: Dive 993, 10 Dec, 
2518 m. slurp sample from base of vesti- 
mentiferans, three specimens (USNM 
118181). Rose Garden, 00°48.25'N, 
86°13.48'W: Dive 896-4. 21 Feb, 2460 m, 
six specimens (USNM 118184). Dive 983, 
30 Nov, 2457 m, mussel bucket washings 
and crab trap residue, 1 3 specimens (USNM 
118183). Dive 990, 7 Dec, 2451 m, wash- 
ings from crab trap. 1 specimen (USNM 
118185). 

East Pacific Rise off western Mexico, 
20°50'N, 109°06'W, Oasis Alvin dives in 
1982: Dive 1220, 26 Apr, 2617 m, Riftia 
and Calyptogena washings, nine specimens 
(USNM 118 182). Dive 1229. 14 May, 2615 
m. Riftia, Calyptogena and Alvinella wash- 
ings, eight specimens (USNM 118186). 

Gulf of California. Guaymas Basin, dive 



VOLUME 102, NUMBER 1 



157 



ofthe^/v/winjan 1982, 27°01'N, 111°25'W, 
2020 m. Dive 1 1 70, rock sample, two small 
specimens (USNM 118708). Dives of the 
Alvin in 1985, 27°00'N, 111°24'W, 2000- 
2009 m: Dive 1603, 25 Jul, Angel Rock, 
bacterial mat, Riftia washings, three spec- 
imens (USNM 1 18707). Dive 1607, 29 Jul, 
Lutz 2, bacterial mat, washings from rocks 
worms, one specimen (USNM 118706). 
Dive 1609, 1 Aug, Lutz 2, sediment, bottom 
of bucket of Bead Exp. no. 1 , one minute 
specimen (USNM 118705). Dive 1615, 7 
Aug, smoker in hydrothermal area, Riftia 
washings, 16 specimens (USNM 118709). 

Remarks. —Some specimens from the 
Guaymas Basin are larger than previously 
reported: up to 19 mm long, 9 mm wide, 
with 26 segments, compared to 1 3 mm long, 
7 mm wide, with 25 segments. 

Distribution.— Gulf of California, East 
Pacific Rise at 21°N, and East Central 
Pacific in the Galapagos Rift, in 2020- 
2633 m. 

Lepidonotopodium williamsae Pettibone 

Lepidonotopodium williamsae Pettibone, 
1984b:857, figs. 5-7. 

Material. —East Central Pacific, from 
dives of the Alvin on two vent areas along 
the Galapagos Rift in 1979: Mussel Bed, 
00°47.89'N, 86°21'W: Dive 991, 8 Dec, 2490 
m, washings from mussels in mesh-lined 
milk crates and slurp samples from mussel 
wall, eight specimens (USNM 1 181 18-90). 
Rose Garden, 00°48.25'N, 86°13.48'W: 
Dive 990, 7 Dec, 2451 m, residue from re- 
trieval box containing vestimentiferans, two 
specimens (USNM 1 18187). 

East Pacific Rise off western Mexico, 
20°50'N, 109°06'W, Oasis Alvin dive in 
1982: Dive 1220, 26 Apr, 2617 m, Riftia 
and Calyptogena washings, three specimens 
(USNM 118188). 

Distribution. —Hydrothermal vents of 
Tropical East Pacific, Galapagos Rift and 
East Pacific Rise at 21°N, in 2451-2633 m. 



Subfamily Branchinotogluminae Pettibone, 

1985 

Genus Branchinotogluma Pettibone, 1985 

Branchinotogluma grasslei Pettibone 

Branchinotogluma grasslei Pettibone, 
1985b:457, figs. 5, 6; 1988: 215. 

Material. — East Central Pacific, Galapa- 
gos Rift, Alvin dives in 1979: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec, 
2490 m, two slurp samples from mussel wall, 
three specimens (USNM 118151). Rose 
Garden, 00°48.25'N, 86°13.48'W: Dive 983, 
30 Nov, 2457 m, mussel bucket washings, 
three specimens (USNM 118152). 

East Pacific Rise off western Mexico, Oa- 
sis Alvin dive in 1982, 20°50'N, 109°06'W: 
Dive 1220, 26 Apr, 2617 m, Riftia and Ca- 
lyptogena washings, one specimen (USNM 
118150). 

Gulf of California, Alvin dive in Guaymas 
Basin in Jan 1982, 27°01'N, 1 1 1°25'W: Dive 
1172, 2010 m, Box core no. 1, on slope of 
mound in area of sulfides, sediment covered 
with white material, one specimen (USNM 
118699). 

Distribution. —Hydrothermal vents of 
Tropical East Pacific, Galapagos Rift and 
East Pacific Rise at 2 1°N; Gulf of California; 
Northeast Pacific, Explorer and Juan de Fuca 
Ridges, in 1495-2633 m. 

Branchinotogluma sandersi Pettibone 

Branchinotogluma sandersi Pettibone, 
1985b:453, figs. 3, 4; 1988: 217. 

Material.— East Central Pacific, Galapa- 
gos Rift, Alvin dives in 1979: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec, 
2490 m, two slurp samples from mussel wall, 
two specimens (USNM 118157). Rose Gar- 
den, 00°48.25'N, 86°13.48'W: Dive 983, 30 
Nov, 2457 m, mussel bucket washings, one 
specimen (USNM 1181 56). Dive 988,5 Dec, 
2450 m, slurp sample from base of vesti- 
mentiferans, one specimen (USNM 
118158). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



East Pacific Rise off western Mexico, Oa- 
sis Alvin dive in 1982, 20°50'N, 109°06'W: 
Dive 1220, 26 Apr, 2617 m, Riftia and Ca- 
lyptogena washings, three specimens 
(USNM 118159). 

Gulf of California, Alvin dive in Guaymas 
Basin in 1985, 27°00'N, 111°24'W: Dive 
1599, 13 Jul, 2009 m, Lutz 2, bacterial mat, 
washing from Rock Chimney, one specimen 
(USNM 118700). 

Distribution. — Hydrothermal vents of 
Tropical East Pacific, Galapagos Rift and 
East Pacific Rise at 2 1°N; Gulf of California; 
Northeast Pacific, Explorer and Juan de Fuca 
Ridges, in 1592-2633 m. 

Branchinotogluma hessleri Pettibone 

Branchinotogluma hessleri Pettibone, 
1985b:450, figs. 1, 2. 

Material— East Central Pacific, Galapa- 
gos Rift, dives of Alvin in 1 979: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec, 
2490 m, two slurp samples from mussel wall, 
three specimens (USNM 118164). Garden 
of Eden, 00°47.69°N, 86°07.74'W: Dive 993, 
10 Dec, 2518 m, slurp sample from base of 
vestimentiferans, four specimens (USNM 
118162). Rose Garden, 00°48.25'N, 
86°13.48'W: Dive 983, 30 Nov, 2457 m, 
mussel bucket washings, three specimens 
(USNM 118161). Dive 988, 5 Dec, 2450 
m, slurp sample from base of vestimenti- 
ferans, one specimen (USNM 118163). 

East Pacific Rise off western Mexico, Oa- 
sis Alvin dives in 1982, 20°50'N, 109°06'W: 
Dive 1220, 26 Apr, 2617 m, Riftia and Ca- 
lyptogena washings, 1 3 specimens (USNM 
1181 60). Dive 1 229, 1 4 May, 2615 m, Rif- 
tia, Calyptogena and Alvinella washings, 2 
specimens (USNM 118165). 

Distribution. —Hydrothermal vents of 
Tropical East Pacific, Galapagos Rift and 
East Pacific Rise at 21°N, in 2450-2633 m. 

Opisthotrochopodus Pettibone, 1985 
Opisthotrochopodus alvinus Pettibone 

Opisthotrochopodus alvinus Pettibone, 
1985b:459, figs. 7-9. 



Material.— East Central Pacific, Galapa- 
gos Rift, dives of Alvin in 1979: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec, 
2490 m, two slurp samples from mussel wall, 
one specimen (USNM 118179). Garden of 
Eden, 00°47.69'N, 86°07.74'W: Dive 993, 
10 Dec, 25 18 m, slurp sample from base of 
vestimentiferans, 2 specimens (USNM 
118177). Rose Garden, 00°48.25'N, 
86°13.48'W: Dive 983, 30 Nov, 2457 m, 
mussel bucket washings, 1 1 specimens 
(USNM 1181 74), crab trap residue, 2 spec- 
imens (USNM 118176). 

East Pacific Rise off western Mexico, Oa- 
sis Alvin dive in 1982, 20°50'N, 109°06'W: 
Dive 1220, 26 Apr, 2617 m, Riftia and Ca- 
lyptogena washings two specimens (USNM 
118178). 

Distribution. —Hydrothermal vents of 
Tropical East Pacific, Galapagos Rift and 
East Pacific Rise at 21°N, in 2451-2633 m. 

Branchinotogluma sp. A 
[Young of B. hessleri or O. alvinus] 

Branchinotogluma sp. A Pettibone, 1985b: 
466. 

Material. —East Central Pacific, from 
dives of Alvin along Galapagos Rift in 1979: 
Mussel Bed: Dive 991,8 Dec, 2490 m, slurp 
sample from mussel wall, nine young spec- 
imens (USNM 118154). Garden of Eden: 
Dive 993, 10 Dec, 2518 m, slurp sample 
from base of vestimentiferans, two young 
specimens (USNM 1 18155). Rose Garden: 
Dive 983, 30 Nov, 2457 m, mussel bucket 
washings, 10 young specimens (USNM 
118153). 

Remarks. —Adults of the above two 
species were collected on the same dives. 

Branchinotogluma sp. B 
[Young of B. grass lei or B. sandersi] 

Branchinotogluma sp. B Pettibone, 1985b: 
466. 

Material. —East Central Pacific, from 
dives of Alvin along Galapagos Rift in 1 979: 
Rose Garden: Dive 983, 30 Nov, 2457 m, 



VOLUME 102, NUMBER 1 



159 



mussel bucket washings, 1 1 young speci- 
mens (USNM 118149). 

Gulf of California, Alvin dive in Guaymas 
Basin in 1985, 27°00'N, 1 1 1°24'W: Dive 
1603, 25 Jul, 2004 m, Angel Rock, bacterial 
mat, Riftia washings, one small specimen 
(USNM 118701). 

Remarks. —Adults of the above two 
species were collected on the same dives. 

Subfamily Branchiplicatinae Pettibone, 

1985 

Genus Branchiplicatus Pettibone, 1985 

Branchiplicatus cupreus Pettibone 

Branchiplicatus cupreus Pettibone, 1985a: 
151, figs. 1-4. 

Material. —Gulf of California, Alvin dive 
in Guaymas Basin in 1982, 27°01'N, 
1 1 1°25'W: Dive 1 169-001, 1 1-20 Jan, 2020 
m, rock sample, one specimen (USNM 
118702). Alvin dive in 1985, 27°00'N, 
111°24'W: Dive 1615, 7 Aug, 2000 m, 
smoker 1, hydrothermal area, Riftia wash- 
ings, one specimen (USNM 118703). 

Distribution. —Hydrothermal vents in 
Gulf of California and East Pacific Rise at 
21°N, in 2000-2633 m. 

Subfamily Macellicephalinae Hartmann- 

Schroder, 1971, 

emended Pettibone, 1976 

Genus Levensteiniella Pettibone, 1985 

Levensteiniella kincaidi Pettibone 

Levensteiniella kincaidi Pettibone, 1985c: 
471, figs. 1-3; 1988:215. 

Material.— East Central Pacific, Galapa- 
gos Rift, dives of Alvin in 1 979: Mussel Bed, 
00°47.89'N, 86°09.21'W: Dive 986, 3 Dec, 
2494 m, mussel bucket washings, three 
specimens (USNM 118196). Dive 991, 8 
Dec, 2490 m, slurp samples from mussel 
wall, four young specimens (USNM 
118194). Garden of Eden, 00°47.69'W, 
86°07.74'W: Dive 993, 10 Dec, 2518 m, 
slurp sample from base of vestimentiferans, 
29 young specimens (USNM 118191). Rose 
Garden, 00°48.25'W, 86°1 3.48'W: Dive 983, 



30 Nov, mussel bucket washings, 20 young 
specimens (USNM 118197). Dive 988, 5 
Dec, 2450 m, slurp sample from base of 
vestimentiferans, one specimen (USNM 
118192). 

East Pacific Rise off western Mexico, 
20°50'N, 109°06'W, Oasis Alvin dives in 
1982: Dive 1220, 26 Apr, 2617 m, Riftia 
and Calyptogena washings, 1 specimens (4 
young, USNM 1 18195). Dive 1229, 14 May, 
2616 m, Riftia, Calyptogena and Alvinella 
washings, small specimen (USNM 118193). 

Distribution. —Hydrothermal vents of 
Tropical East Pacific: Galapagos Rift and 
East Pacific Rise at 2 1°N; Northeast Pacific, 
Explorer and Juan de Fuca Ridges, in 1818- 
2633 m. 



Genus Bathykurila Pettibone, 1976 

Bathykurila guaymasensis, new species 

Figs. 1, 2 

Material. —Gulf of California, Alvin dive 
in Guaymas Basin in Jan 1982, 27°01'N, 
1 1 1°25'W: Dive 1 176, 2020 m, Box core 2, 
taken over vent, with numerous nuculanid 
bivalves, 2d, holotype (USNM 1 1 8694) and 
paratype (USNM 1 18695); sec. B, paratype 
(USNM 1 18696). Alvin dive in 8 Aug 1985, 
27WN, 111°24'W: Dive 1614, 2004 m, 
Angel Rock, bacterial mat, Box core 5, sub- 
core B, 0-5 cm, paratype (USNSM 1 1 8697), 
subcore, C, 0-3 cm, paratype (USNM 
118698). 

Description.— Length of holotype 8 mm, 
width with setae 5 mm, segments 1 5; length 
of two larger paratypes 7 mm, width 5 mm, 
segments 14; length of two smaller para- 
types 3-4 mm, width 3-3.5 mm, segments 
15. Body fusiform, flattened, with parapo- 
dia and setae very long, projecting, longer 
than body width (Fig. 1A, B). 

Elytra (mostly missing) and large, bul- 
bous elytrophores (Figs. 1A, B, 2A) seven 
pairs, on segments 2, 4, 5, 7, 9, 11, and 13. 
Elytra large, oval, covering dorsum, deli- 
cate, with larger conical to rounded tuber- 
cles variable in size on posterior and lateral 
borders, with smaller ones scattered on sur- 



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Fig. 1. Bathykurila guaymasensis, A, B, holotype, USNM 1 18694; C, paratype, USNM 1 18698; D, paratype, 
USNM 118695: A, Dorsal view of anterior end, acicula in tentaculophores of 1st segment dotted; B, Dorsal 
view of posterior end (segments 12-15); C, Ventral view of posterior end (segments 10-14), anal cirri and most 
of ventral cirri broken off; D, Border papillae on dorsal side of pharynx (dissected). Scales = 0.5 mm for A, B; 
1 .0 mm for C; 0. 1 mm for D. 



face (Fig. 2F): larger tubercles not usual type, 
not chitinous, not sharply set off from sur- 
face. Dorsal tubercles on cirrigerous seg- 
ments large, bulbous to nodular (Figs. 1A, 
B,2B). 

Prostomium oval, deeply bilobed; trian- 
gular anterior continuations of prostomium 
with small frontal filaments; median anten- 
na with small ceratophore in anterior notch, 
with short subulate style; without lateral an- 
tennae and eyes; palps stout, tapered, about 
two times longer than prostomium (Fig. 1 A). 
First or tentacular segment fused to prosto- 
mium, not visible dorsally; tentaculophores 
lateral to prostomium, each with prominent 
acicular lobe on inner side, without setae, 
and pair of subulate tentacular cirri, shorter 
than palps; upper lip large, bilobed (Fig. 1 A). 
Second or buccal segment with first pair of 
elytrophores, biramous parapodia and ven- 
tral or buccal cirri lateral to ventral mouth, 
longer than following ventral cirri (Fig. 1 A). 



Parapodia biramous, with notopodia 
shorter and smaller than neuroodia (Fig. 2A, 
B.) Notopodium rounded basally, with pro- 
jecting acicular lobe on lower side; neuro- 
podium with conical projecting presetal 
acicular lobe and shorter, rounded postsetal 
lobe. Notosetae numerous, short to longer, 
forming radiating bundles, stouter than 
neurosetae, with row of widely spaced teeth 
(4-11) along one side and blunt tips (Fig. 
2C). Neurosetae very numerous, forming 
fan- shaped bundles; supraacicular neuro- 
setae with longer spinous regions and two 
rows of long spines, tapering to bluntly 
pointed bare tips (Fig. 2D); subacicular neu- 
rosetae with shorter spinous regions, close- 
set short spines, and tapered bare tips (Fig. 
2E). Dorsal cirri with cylindrical cirro- 
phores on posterior sides of notopodia, 
shorter than notopodia, with styles wider 
basally and long slender tips extending to 
tips of setae or beyond (Figs. 1A, B, 2B). 



VOLUME 102, NUMBER 1 



161 




D 

Fig. 2. Bathykurila guaymasensis, holotype, USNM 118694: A, Right elytragerous parapodium, anterior 
view, acicula dotted; B, Right cirrigerous parapodium, posterior view; C, Notosetae; D, Supraacicular neuroseta; 
E, Subacicular neuroseta; F, Right elytron. Scales = 0.5 mm for A, B; 0.1 mm for C-E; 0.5 mm for F. 



Ventral cirri attached on middle of neuro- 
podia, short, tapered (Fig. 2 A, B). 

Pair of long ventral papillae usually pres- 
ent on segment 1 1 (present on four of five 
specimens), sometimes extending far be- 
yond posterior end, with yellowish secretion 
inside (Fig. 1C). Posterior 2 segments (14, 
15) smaller, enclosed in elytragerous seg- 
ment 13 (Fig. IB); segment 14 with dorsal 
cirri; segment 1 5 smaller, without dorsal cirri 
(Fig. IB), or segment absent (Fig. 1C); py- 
gidium long, slender, wedged between par- 
apodia of segment 15, with pair of slender, 
short anal cirri (Fig. IB). 

Pharynx (dissected) with opening encir- 
cled by eight pairs of border papillae (Fig. 
ID), and two pairs of heavy, hooked jaws. 

Etymology. —The species is named for the 
collecting site, the Guaymas Basin. 

Remarks.— Bathykurila guaymasensis is 
close to B. zenkevitchi (Uschakov 1955, fig. 
la-c; Pettibone 1976, fig. 24a-d), the type 
species of Bathykurila, from the Kurile- 
Kamchatka Trench in the northwest Pacific, 
in 8100 m. They differ as indicated in Ta- 
ble 1. 



Subfamily Macellicephaloidinae 

Pettibone, 1976 

Genus Macellicephaloides Uschakov, 1955 

Type species.— Macellicephaloides gran- 
dicirrus Uschakov, 1955 (as grandicirra). 
Gender: masculine (according to the Code, 
Art. 30.a.i, names ending in -oides should 
be masculine, not feminine, as indicated by 
Pettibone 1976:42). 

Table 1 . —Differences between Bathykurila zenkevit- 
chi (Uschakov) and Bathykurila guaymasensis, n. sp. 



Character 


B. zenkevitchi 


B. guaymasensis 


Tentacular 


about as long as 


shorter than palps 


cirri 


palps 




Notopodia 


as long as or 


shorter than neu- 




longer than 


ropodia 




neuropodia 




Cirrophore 


longer than no- 


shorter than no- 


of dorsal 


topodia 


topodia 


cirri 






Ventral pa- 


4 pairs on seg- 


1 pair on segment 


pillae 


ments 10-13 


11 


Length 


15-21 mm 


3-8 mm 


Width 


12 mm 


3-5 mm 


Segments 


15 


14-15 



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Fig. 3. Macellicephaloides alvini, holotype, USNM 1 18704: A, Dorsal view of anterior end, styles of median 
antenna, tentacular cirri, and dorsal cirri of segment 3 missing, right parapodia of segments 2 and 3 damaged 
or not developed; B, Ventral view of anterior end, ventral cirri missing; C, Dorsal view of posterior end (segments 
13-17), elytra and styles of dorsal cirri missing. Scale =1.0 mm for A-C. 



Macellicephaloides alvini, new species 
Figs. 3, 4 

Material. —Gulf of California, Alvin dive 
in Guaymas Basin, F. Grassle, collector, 
Dive 1614, 6 Aug 1985, 27°00'N, 1 1 1°24'W: 
2004 m, Angel Rock, bacterial mat, Box 
core 5, subcore D, holotype (USNM 
118704). 

Description. —Length of holotype 7.5 mm, 
width with setae 4 mm, segments 17. Body 
flattened dorsoventrally, tapering slightly 
anteriorly and more so posteriorly, with 
parapodia of 5 posterior segments directed 
posteriorly (Fig. 3C). Integument smooth, 
not papillate. Elytra (all missing) and large, 
bulbous elytrophores eight pairs, on seg- 
ments 2, 4, 5, 7, 9, 11, 13, and 1 5 (Figs. 3A, 
C, 4A). Dorsal tubercles on cirrigerous seg- 
ments indistinct. 

Prostomium (poor condition) withdrawn 
and fused to tentacular segment, oval, 
slightly bilobed; ceratophore of median an- 
tenna in middle of prostomium, style miss- 
ing; palps on short palpophores, rather short, 
tapering; without lateral antennae and eyes 
(Fig. 3 A, B). Tentacular segment distinct 
dorsally, ventrally forming upper, lateral, 
and posterior lips of ventral mouth; tenta- 



culophores of dorsal and ventral cirri short, 
lateral to prostomium, styles missing (Fig. 
3 A, B). Segment 2 with first pair of elytro- 
phores and elongate parapodia; ventral cirri 
missing; deep depression in middle of ven- 
tral surface, covered by large, oval flap at- 
tached to segment 3 (Fig. 3 A, B). 

Parapodia subbiramous; notopodium 
bulbous basally, with projecting, stout, am- 
ber-colored notoaciculum, without noto- 
setae (Fig. 4A-C); neuropodium elongated, 
with projecting acicular lobe; neurosetae 
numerous, delicate, slender, slightly wider 
basally, tapering to slender tips, with two 
rows of delicate spines (Fig. 4A, B, D). Pos- 
terior five segments with notoacicula stou- 
ter, projecting beyond neuropodia (Fig. 3C). 
Dorsal cirri with rather long, thick, cylin- 
drical cirrophores on posterior dorsal sides 
of notopodia; styles missing (Fig. 4B). Ven- 
tral cirri missing. 

Pygidium rounded, enclosed in parapo- 
dia of last segment (Fig. 3C). Pharynx (dis- 
sected) large, muscular; anterior end encir- 
cled with papillae, number and arrangement 
not evident but with longer lateral pair (not 
extra long as in M. vitiazi); two pairs of long 
V-shaped hooked jaws; dorsal pair fused 
medially, with additional median piece (Fig. 



VOLUME 102, NUMBER 1 



163 




Fig. 4. Macellicephaloides alxini, holotype, USNM 1 18704: A, Right elytragerous parapodium from segment 

7, anterior view, acicula dotted, elytron and ventral cirri missing; B, Right cirrigerous parapodium of segment 

8, posterior view, styles of dorsal and ventral cirri missing; C, Notoaciculum from same; D, Neuroseta from 
same, with detail of parts: E, Dorsal jaws, outer view; F, Same, inner view; G, Ventral jaws, outer view; H, 
Same, inner view. Scales = 0.5 mm for A, B; 0.1 mm for C-H. 



4E, F); ventral pair denticulate on inner side 
(Fig. 4G, H). 

Etymology.— The species is named for the 
collecting submersible, DSRV Ahin. 

Remarks on the genus Macellicephaloides 
Uschakov, 1955, and the subfamily Macel- 
licephaloidinae Pettibone, 1976: Of partic- 
ular interest is the find of a single specimen 
from the Guaymas Basin that seems to fit 
into the genus Macellicephaloides, estab- 
lished by Uschakov (1955) for three 
ultra-abyssal species, M. grandicirrus, M. 
verrucosus, and M. vitiazi, from the Kurile- 
Kamchatka Trench in 7210-9950 m. Us- 
chakov placed the genus under Aphroditi- 



dae: Polynoinae. An additional species, M. 
uschakovi, was added to the genus from the 
same area in 8120 m by Levenstein (1971). 
In her catalogue, Hartman (1959:93) 
placed Macellicephaloides under Polynoi- 
dae: Lepidonotinae, and selected M. gran- 
dicirrus as the type species. Hartmann- 
Schroder (1974), in her revision of the 
subfamily Macellicephalinae Hartmann- 
Schroder, 1971, included Macellicepha- 
loides in the subfamily. As a part of my 
revision of the genus Macellicephala and the 
subfamily Macellicephalinae Hartmann- 
Schroder, I established the new subfamily 
Macellicephaloidinae for Macellicepha- 



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hides and the above four species, with sup- 
plementary descriptions, based on speci- 
mens sent in exchange by P. Uschakov and 
R. Levenstein. 

Levenstein (1975) added an additional 
species, M. sandvichensis, from the South 
Sandwich Trench in 7200-7934 m. Another 
species, M. villosus, was added by Leven- 
stein (1982) from the Japan Trench in 73 70- 
7380 m. Levenstein (1983) added a seventh 
species, M. improvisus, from the Kurile- 
Kamchatka Trench in 8035-8120 m and 
included a Key to the seven species of Ma- 
cellicephaloides. It probably should be not- 
ed that Uschakov (1982) did not recognize 
the subfamily Macellicephaloidinae and 
other subfamilies proposed by Pettibone but 
included the genus under Macellicephali- 
nae. 

In her review of the genera and species of 
the deep-sea polynoid subfamilies in the 
world oceans, Levenstein (1984:83, fig. 4) 
indicated that the distributional pattern of 
the Macellicephaloidinae showed a western 
Pacific origin, since six of the seven species 
of the genus Macellicephaloides were known 
to inhabit trenches of the western Pacific 
and only one species was known to inhabit 
the Atlantic sector of the Antarctic Ocean, 
the South Sandwich Trench. She suggested 
that representatives of this genus (or its 
ancestor) were probably widely distributed 
in the abyssal region of the Pacific Ocean 
and gave rise to species adapted to life in 
the oceanic trenches, producing the ultra- 
abyssal fauna of this subfamily. The pres- 
ence of M. sandvichensis in the South Sand- 
wich Trench could be explained on the basis 
of a close connection between the deep-sea 
faunas of the Pacific and Atlantic sectors of 
the Antarctic. The new species, M. alvini 
from 4000-5000 m shallower waters in the 
Gulf of California, might be considered to 
be close to the ancestor of the genus. It is 
considerably smaller than representatives 
from the other areas. 

All eight species have a relatively small 
number of segments ( 1 6-2 1 , 1 st achaetous) 
and eight pairs of elytrophores (elytra all 



missing) on segments 2, 4, 5, 7, 9, 11, 13, 
and 15. The prostomium is relatively small, 
bilobed, with the ceratophore of the median 
antenna in the middle of the prostomium 
and with palps but without lateral antennae 
or eyes. The first or tentacular segment en- 
closes the prostomium and is distinct dor- 
sally, with two pairs of tentacular cirri on 
tentaculophores, but without acicular lobes 
or setae; ventrally the tentacular segment 
forms the upper, lateral, and posterior lips 
of the ventral mouth. The second segment 
has the first pair of elytrophores, large par- 
apodia, and ventral cirri, with a deep mid- 
ventral depression covered by a large oval 
flap attached to segment 3 (as in M. alvini, 
Fig. 3B) or a rectangular flap (as in M. gran- 
dicirrus, Pettibone, 1976, fig. 25b). Para- 
podia are elongate, subbiramous, the no- 
topodium represented by a conical lobe with 
a more or less projecting, stout notoacicu- 
lum, without notosetae; the neuropodium 
is conical, with numerous, slender neuro- 
setae. Dorsal cirri on the non-elytragerous 
segments have moderately to very long, cy- 
lindrical cirrophores and short styles; dorsal 
tubercles are indistinct or rarely nodular, as 
in M. verrucosus. Ventral cirri are short. The 
pygidium is enclosed in the long parapodia 
of a variable number of posterior segments, 
and without anal cirri. The integument is 
smooth, rarely papillate dorsally, as in M. 
villosus, or with middorsal nodular tuber- 
cles, as in M. verrucosus. The large muscular 
pharynx has two pairs of hooked jaws and 
distal papillae, the lateral pair of papillae 
longer than the others. Lateral papillae 
sometimes very long, as in M. vitiazi (Pet- 
tibone, 1976, fig. 26c) and M. sandvichensis 
(Levenstein, 1975, fig. 3a), and similar to 
those of the pelagic polychaete family Al- 
ciopidae. 

Key to the Species of 
Macellicephaloides 

1. Segments 16 or 17 2 

- Segments 18 or 20-21 6 

2. Segments 16 3 

- Segments 17 4 



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165 




Fig. 5. Neoleanira racemosa, USNM 118714: A, Dorsal view of anterior end, ventral tentacular cirri not 
visible; left dorsal cirrus on segment 3 broken off; B, Right tentacular parapodium, outer view, aciculum dotted; 
C, Right elytragerous parapodium from segment 2, anterior view, acicula dotted; D, Right cirrigerous parapodium 
from segment 3, posterior view; E, Right elytragerous parapodium from segment 4, anterior view, acicula dotted. 
Scales = 1.0 mm for A; 0.5 mm for B-E. au, auricle; br, branchia; ct, ctenidium; dc, dorsal cirrus; dtc, dorsal 
tentacular cirrus; itl. inner tentacular lobe; Ian. lateral antenna; man, median antenna; pa. palp; st. stylode; vtc, 
ventral tentacular cirrus. 



With paired, nodular dorsal tuber- 
cles on cirrigerous segments 6-14 
and middorsal tubercles on seg- 
ments 3-15 (Uschakov 1955, fig. 3; 
1982, pi. 12; Pettibone 1976, fig. 27 

M. verrucosus Uschakov 

Without dorsal tubercles on cirri- 
gerous segments and middorsal tu- 
bercles (Uschakov 1955, fig. 4; 1982, 
pi. 13; Pettibone 1976, fig. 26) ... 

M. vitiazi Uschakov 

Notoacicula not extra stout or long. 
Cirrophores of dorsal cirri extend- 
ing to about tips of neuropodia. 
Pharynx with pair of very long lat- 
eral papillae (Levenstein 1975. fig. 

3) M. sandxichensis Levenstein 

Notoacicula stout, extending far be- 
yond neuropodia in posterior seg- 



ments. Pharynx with lateral papillae 
not extra long 5 

5. Cirrophores of dorsal cirri extend- 
ing far beyond tips of neuropodia 
(Uschakov 1955, fig. 2; 1982, pi. 11; 
Pettibone 1976, fig. 25) 

M. grandicirrus Uschakov 

- Cirrophores of dorsal cirri not as 
long, extending to tips of neuropo- 
dia or shorter (Figs. 3. 4) 

M. alvini, new species 

6. Segments 18. Cirrophores of dorsal 
cirri very short. Notopodia with 
flexible, thin acicula (Levenstein 
1983, fig. a-c) 

M. improvisus Levenstein 

- Segments 20-21. Notopodia other- 
wise 

7. Dorsal side of body with conical pa- 



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Fig. 6. Neoleanira racemosa, USNM 1 18714: A, Right parapodium from posterior region (about segment 
35), anterior view, acicula dotted; B, Same, posterior view; C, Upper, middle, and lower compound neurosetae, 
with detail of distal part of stems. Scales = 0.5 mm for A, B; 0.1 for C. br, branchia; ct, ctenidium; st, stylode. 



pillae (Levenstein 1982, fig. 2) ... 

M. villosus Levenstein 

- Integument smooth (Levenstein 
1971, fig. 6) . .M. uschakovi Levenstein 

Family Sigalionidae 

Genus Neoleanira Pettibone, 1970 

Neoleanira racemosa (Fauchald) 

Figs. 5, 6 

Sthenolepis racemosa Fauchald, 1972:34, 
pi. 2, figs. a-d. 

Neoleanira racemosa. — Fauchald & Han- 
cock, 1981:23. 

Remarks. —The species was originally de- 
scribed by Fauchald (1972) under Stheno- 
lepis, based on specimens collected in the 
Guaymas Basin in 1774-2259 m and later 
referred to Neoleanira by Fauchald and 
Hancock (1981), with additional specimens 
reported from off central Oregon in 2000- 
2900 m. The additions to the description of 
the species are based on additional speci- 
mens from the Guaymas Basin. All speci- 
mens are incomplete posteriorly and elytra 
are missing in both the previous and the 
present collections. 

Material. —Gulf of California, Alvin dives 



in Guaymas Basin, Jan 1972, 27°01'N, 
111°25'W, 2020 m: Dive 1170, 2 m from 
vestimentiferans, Box core 1, small speci- 
men (USNM 1 18713). Dive 1 173, on slope 
of mound, Box core 1 , one specimen (USNM 
118714), Box core 2b, subcore D, two spec- 
imens (USNM 118715). Dive 1174, about 
15 m from mound, Box core la, one spec- 
imen (USNM 118712), Box core lc, one 
small specimen (USNM 118710). Dive 
1177, about 3 m from mound with tube 
worms, Box core 2, one specimen (USNM 
118711). 

Supplementary description. —Incomplete 
specimen of 44 segments, 16+ mm long, 3 
mm wide with setae (USNM 1 18714). Ely- 
tra all missing; elytrophores on segments 2, 
4, 5, 7, continuing on alternate segments to 
23, then on all segments. Prostomium 
rounded, ceratophore of median antenna on 
anterior margin, with pair of lateral auricles 
and long tapering style; parapodia of first or 
tentacular segment partially fused to lateral 
and anterior sides of prostomium, with long 
lateral antennae attached on dorsal, inner 
side; long dorsal and short ventral tenta- 
cular cirri with short inner tentacular lobe; 
long tapering palps inserted ventral to ten- 



VOLUME 102, NUMBER 1 



167 



tacular parapodia; with aciculum and 2 ra- 
diating bundles of capillary setae (Fig. 5A, 

B). 

Parapodia of segments 2 and 3 directed 
anteriorly (Fig. 5 A). Segment 2 with first 
pair of elytrophores, biramous parapodia, 
and ventral buccal cirri longer than follow- 
ing ventral cirri (Fig. 5A, C). Parapodium 
of segment 3 with long dorsal cirrus on short 
cirrophore and small oval ventral ctenid- 
ium medial to ventral cirrus (Fig. 5A, D). 
Segment 4 with 3 oval ctenidia between dor- 
sal base of notopodium and elytrophore, 
with very small lateral branchia (Fig. 5E). 
Anterior neuropodia with numerous fim- 
briae or stylodes on tips of acicular lobes 
and bilobed posterior bracts (Fig. 5C-E). 

More posterior parapodia with well-de- 
veloped branchiae attached to lateral sides 
of elytrophores or dorsal tubercles and 3 
ciliated ctenidia in curved part; notopo- 
dium with stylodes and long capillary no- 
tosetae, finely to coarsely spinous; neuro- 
podium conical, with stylodes, fan-shaped 
bundle of slender neurosetae, ventral cirrus, 
and small bulbous ventral ctenidium (Fig. 
6A, B). Neurosetae compound spinigers, 
upper and lower ones with shorter blades, 
middle ones with longer blades, none can- 
aliculate; long slender shafts distally dentate 
(Fig. 6C). 

Distribution.— Gulf of California and off 
Central Oregon. In 1774-2900 m. 

Key to the Species of Neoleanira 

1. Blades of compound spinigerous 
neurosetae not canaliculate. Ventral 
ctenidia small, bulbous. Branchiae 
without prominent spurlike pro- 
cesses. Elytra missing (Figs. 5, 6; 
Fauchald 1972, figs, a-d) 

N. racemosa (Fauchald) 

- Blades of compound spinigerous 
neurosetae canaliculate 2 

2. Elytra with borders entire, without 
lateral fringes of papillae. [Ventral 
ctenidia small, clavate. Branchiae 
without prominent basal spurlike 
processes] (Pettibone 1970, fig. 7) 



TV. magellanica (Mcintosh) 

- Elytra with delicate lateral fringes of 
papillae 3 

3. Ventral ctenidia small, clavate. 
Branchiae without prominent basal 
spurlike processes (Pettibone 1970, 
figs. 1-4) N. tetragona (Oersted) 

- Ventral ctenidia large, boot-shaped. 
Branchiae with prominent basal 
spurlike processes (Pettibone 1970, 
figs. 5, 6) N. areolata (Mcintosh) 

Acknowledgments 

My thanks go to J. Fred Grassle, Susan 
Brown-Leger, and Rosemarie Petrecca of the 
Woods Hole Oceanographic Institution for 
the scaled polychaetes from the Guaymas 
Basin and for the additional polynoids com- 
mensal with the Galapagos mussels, sepa- 
rated by Vida Kenk. I also thank Barbara 
Hecker for the additional polynoids com- 
mensal with the Gulf of Mexico mussels, 
separated by Ruth Turner. Additional col- 
lections from the Galapagos and the East 
Pacific Rise at 21°N, received from WHOI 
through the Smithsonian Sorting Center, are 
appreciated. The manuscript benefited from 
the careful reviews of Thomas H. Perkins 
and Nancy J. Maciolek. 

Literature Cited 

Desbruyeres, D., F. Gaill, L. Laubier, & Y. Fouquet. 
1985. Polychaetous annelids from hydrother- 
mal vent ecosystems: An ecological overview. — 
Bulletin of the Biological Society of Washington 
6:103-116. 

Fauchald, K. 1972. Benthic polychaetous annelids 
from deep water off Western Mexico and ad- 
jacent areas in the Eastern Pacific Ocean.— Al- 
lan Hancock Monographs in Marine Biology 7: 
1-575. 

, & D. R. Hancock. 1981. Deep-water poly- 
chaetes from a transect off Central Oregon.— 
Allan Hancock Foundation Monograph 1 1 : 
1-73. 

Grassle, J. F. 1986. The ecology of deep-sea hydro- 
thermal vent communities. — Advances in Ma- 
rine Biology 23:301-362. 

Hartman, O. 1959. Catalogue of the polychaetous 
annelids of the world, Part I. — Allan Hancock 
Foundation Publications Occasional Paper 23: 
1-353. 



168 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Hartmann-Schroder, G. 1974. Die Unterfamilie Ma- 
cellicephalinae Hartmann-Schroder, 1971 (Po- 
lynoidae, Polychaeta). Mit Beschreibung einem 
neuen Art, Macellicephala jameensis n. sp., aus 
einem Hohlengewasser von Lanzarote (Kanar- 
ische Inseln).— Mitteilungen aus dem Hambur- 
gischen Zoologischen Museum und Institut 7 1 : 
75-85. 

Hecker, B. 1985. Fauna from a cold sulfur-seep in 
the Gulf of Mexico: Comparison with hydro- 
thermal vent communities and evolutionary 
implications. — Bulletin of the Biological Society 
of Washington 6:465-473. 

Kenk, V. C, & B. R. Wilson. 1985. A new mussel 
(Bivalvia, Mytilidae) from hydrothermal vents 
in the Galapagos rift zone. — Malacologia 26(1— 
2):253-271. 

Levenstein, R. J. 1971. [Polychaete worms of the 
genera Macellicephala and Macellicephaloid.es 
(Family Aphroditidae) from the Pacific Ocean. 
In Fauna of the Kurile-Kamchatka Trench.] — 
Trudy Institut Okeanologii P.P. Shirshov Aca- 
demiia Nauk SSSR 92:18-35 [in Russian, En- 
glish summary]. 

. 1975. [Polychaetous annelids of the deep-sea 

trenches of the Atlantic Sector of the Antarctic 
Ocean.]— Trudy Institut Okeanologii P.P. Shir- 
shov Academiia Nauk SSSR 103:119-142 [in 
Russian, English summary]. 

. 1982. [On the polychaete fauna of the family 

Polynoidae from the trench of Japan.]— Trudy 
Institut Okeanologii P.P. Shirshov Academiia 
Nauk SSSR 117:59-62 [in Russian, English 
summary]. 

. 1983. \M acellicephaloid.es improvisa sp. n. 

(Polychaeta, Polynoidae) from Kurile-Kam- 
chatka Trench.]— Zoologichesky Zhurnal 62(9): 
1419-1421 [in Russian, English summary]. 

. 1984. On the ways of formation of the deep- 
sea polychaete fauna of the family Polynoidae. 
Pp. 72-85 in P. A. Hutchings, ed., Proceedings 
of the First International Polychaete Confer- 
ence, Sydney, Australia, 1983. The Linnean So- 
ciety of New South Wales. 

Lonsdale, P. 1 984. Hot vents and hydrocarbon seeps 
in the Sea of Cortez.— Oceanus 27(3):21-24. 

Pettibone, M. H. 1970. Two new genera of Sigalio- 
nidae (Polychaeta).— Proceedings of the Biolog- 
ical Society of Washington 83(34):365-386. 

. 1976. Revision of the genus Macellicephala 

Mcintosh and the subfamily Macellicephalinae 
Hartmann-Schroder (Polychaeta: Polynoi- 
dae).— Smithsonian Contributions to Zoology 
229:1-71. 

. 1983. A new scale worm (Polychaeta: Poly- 
noidae) from the hydrothermal rift-area off 
Western Mexico at 2 1°N. — Proceedings of the 
Biological Society of Washington 96(3):392-399. 



. 1984a. A new scale-worm commensal with 

deep-sea mussels on the Galapagos hydrother- 
mal vent (Polychaeta: Polynoidae).— Proceed- 
ings of the Biological Society of Washington 
97(l):226-239. 

. 1984b. Two new species of Lepidonotopo- 

dium (Polychaeta: Polynoidae: Lepidonotopo- 
dinae) from hydrothermal vents of the Gala- 
pagos and the East Pacific Rise at 21°N.— 
Proceedings of the Biological Society of Wash- 
ington 97(4):847-863. 

. 1985a. An additional new scale worm (Po- 
lychaeta: Polynoidae) from the hydrothermal rift- 
area off Western Mexico at 2 1°N.— Proceedings 
of the Biological Society of Washington 98(1): 
150-157. 

. 1985b. Additional branchiate scale-worms 

(Polychaeta: Polynoidae) from Galapagos hy- 
drothermal vent and rift-area off Western Mex- 
ico at 2 1°N.— Proceedings of the Biological So- 
ciety of Washington 98(2):447^169. 

. 1985c. New genera and species of deep-sea 

Macellicephalinae and Harmothoinae (Poly- 
chaeta: Polynoidae) from the hydrothermal rift 
areas of the Galapagos and Western Mexico at 
21°N and from the Santa Catalina Channel.— 
Proceedings of the Biological Society of Wash- 
ington 98(3):740-757. 

. 1986. A new scale-worm commensal with 

deep-sea mussels in the seep-sites at the Florida 
Escarpment in the eastern Gulf of Mexico (Po- 
lychaeta: Polynoidae: Branchipolynoinae).— 
Proceedings of the Biological Society of Wash- 
ington 99(3):444-451. 

. 1988. New species and new records of scaled 

polychaetes (Polychaeta: Polynoidae) from hy- 
drothermal vents of the Northeast Pacific Ex- 
plorer and Juan de Fuca ridges. — Proceedings 
of the Biological Society of Washington 101(1): 
192-208. 

Uschakov, P. V. 1955. [Polychaete worms of the fam- 
ily Aphroditidae from the Kurile-Kamchatka 
Trench.]— Trudy Institut Okeanologii Akade- 
miia Nauk SSSR 12:311-321 [in Russian: 
translated in 1 969 by the Bureau of Commercial 
Fisheries, 10 pages]. 

. 1982. [Polychaetes of the Suborder Aphro- 

ditiformia of the Arctic Ocean and the north- 
western part of the Pacific, families Aphroditi- 
dae and Polynoidae.] In Fauna of the USSR: 
Polychaetes, Vol. 2, Part I, Academy of Sciences 
of the USSR. Zoological Institute, 272 pp. [in 
Russian]. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
102(1), 1989, pp. 169-175 

A NEW EULIMID GASTROPOD, 

TROCHOSTILIFER EUCIDARICOLA, PARASITIC 

ON THE PENCIL URCHIN EUCIDARIS TRIBULOIDES 

FROM THE SOUTHERN CARIBBEAN 

Anders Waren and Robert Moolenbeek 

Abstract. — Trochostilifer eucidaricola (Prosobranchia, Eulimidae) is de- 
scribed from Aruba, an island north of Venezuela. The parasite lives on the 
ventral side of the host, close to the peristome, permanently attached by its 
snout. This is the first Atlantic species of Trochostilifer, a genus previously 
known only from the Indo-Pacific area. 



The family Eulimidae contains a large 
number of species, all of which as far as 
known, parasitize echinoderms (Waren 
1984). Many species have been described 
from the western Atlantic including the Ca- 
ribbean, but at present the host is known 
for only a small number of them. The eu- 
limids represent all different levels of par- 
asitism, from species with normal gastro- 
pod anatomy that suck body fluids from the 
host for a comparatively short time and then 
leave the host to digest their meal, to species 
that are permanent endoparasites and can 
be recognized as gastropods only from the 
veliger larvae. 

The genus Trochostilifer Waren, 1980 
contains five species, all parasitic on cida- 
roid sea urchins. Four of these species live 
in galls in the spines of their hosts. The fifth 
species, Trochostilifer striatus (Hedley, 1 90 1) 
lives attached to the test of the host close 
to the peristome in a way similar to the 
present new species. 

Trochostilifer eucidaricola, new species 
Figs. 1,3-8, 14 

Holotype. —Zoological Museum, Am- 
sterdam (ZMA Moll. 3.88.004). Female 
from host number 9 (Fig. 4). Allotype.— 
Zoological Museum, Amsterdam (ZMA 
Moll. 3.88.005). Male attached with holo- 
type (Fig. 6). 



Paratypes. —Remaining specimens in 
Zoological Museum, Amsterdam, Swedish 
Museum of Natural History and U.S. Na- 
tional Museum of Natural History, see Ta- 
ble 1. 

Type locality. —Parasitic around peri- 
stome of Eucidaris tribuloides (Lamarck). 
Caribbean, Aruba, Malmok, Sandy Thalas- 
sia flat with Pontes, 2-7 m depth, under 
corals and diorite boulders (12°36'N, 
70°3.5'W). Leg. I. Peters, Dec 1986. 

Material examined. —Known only from 
the type locality and the material examined, 
listed in Table 1 . 

Description.— Female: Shell of medium 
size, conical, transparent, rather solid, with 
high, distinctly shouldered whorls and 
sharply set off larval shell. Larval shell (Figs. 
1, 3) of typical eulimid shape, consisting of 
about four slightly convex, perfectly smooth 
whorls, of which 650 jum are visible above 
the teleoconch (Fig. 3). Holotype with about 
five teleoconch whorls, of which the apical 
2.5 whorls are distinctly shouldered and the 
lower 1.5 almost perfectly flat. Between these 
there is one transitional whorl. Body whorl 
distinctly angulated at the periphery with a 
demarcated basal surface. Shell surface 
marked by very shallow, indistinct spiral 
furrows and more obvious, sharp incre- 
mental lines plus irregularly appearing, 
strong incremental scars. Aperture strongly 



170 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1.— Specimens of Eucidaris tribuloides para- 
sitized by culimids. Explanation: F— female; M— male; 
O— sex characteristics not developed. Numbers in pa- 
renthesis are register numbers in ZMA— Department 
of Malacology, Zoological Museum of Amsterdam; 
SMNH — Department of Invertebrate Zoology, Swed- 
ish Museum of Natural History; USNM— Division of 
Mollusks, U.S. National Museum of Natural History. 
Other specimens kept in ZMA and SMNH. 









Euli- 


Specimen 






mid 


number 


T. eucidaricola 


5. troglodytes 


n. sp. 


1 (Aug 


F 3.6 mm, 


— 


3 


1986) 


M 2.4 mm 
(SMNH 
3847, left on 
host) 






2 (Dec 


F 3.6 mm, 


— 


7 


1986) 


M 2.0 mm, 
eggs (SMNH 
3848) 






3 (May 


O 1.6 mm 


2 apical, emp- 


7 


1986) 


(ZMA Moll. 


ty galls; 1 






3.88.006 


basal gall 
(Fig. 13), 
F 4.6 mm, 
M 2.2 mm, 
M 2.3 mm, 
M 2.2 mm, 
O 0.9 mm 
(Figs. 9-11) 
(galls never 
closed) 




4 (??? 


— 


(1 open apical 


— 


1986) 




gall, F 2.8 
mm, M 2.2 
mm, eggs) 




5 (Dec 


1 pair lost; 


— 


— 


1986) 


F 4.0 mm, 
M 2.6 mm, 
eggs (ZMA 
Moll. 
3.88.007- 
008, left on 
host) 






6 (Dec 


O 2.0 mm, 


— 


— 


1986) 


O 2.0 mm 
(ZMA Moll. 
3.88.009- 
010) 






7 (Dec 


O 2.2 mm, 


— 


— 


1986) 


O 2.2 mm 
(ZMA Moll. 
3.88.011- 

012) 







Table 1. 


—Continued. 






Specimen 
number 


T. eucidaricola 


5. troglodytes 


Euli- 
mid 
n. sp. 


8 (Dec 


O 1.6 mm, 


— 


— 


1986) 


O 0.6 mm 
(ZMA Moll. 
3.88.013- 
014) 






9 (Dec 


F 3.6 mm, 


empty gall 


2 


1986) 


M 2.0 mm 
(USNM 
859313); 
F 4.8 mm 
(holotype), 
M 2.6 mm 
(allotype) 
(ZMA Moll. 
3.88.004 
and 005) 






10 (Dec 


— 


O 0.6 mm, on 


— 


1986) 




spine 




1 1 (Dec 


— 


1 gall with 


— 


1986) 




M 2.2 mm, 
M 2.2 mm, 
M 1.8 mm, 
M 1.8 mm 
(Fig. 15). 





prosocline. Outer lip thickened, with the 
most protruding part situated one-seventh 
of total height from suture. Inner lip thick- 
ened, reflected over columella. Height of 
holotype 4.68 mm (larval shell broken). 

Soft parts (based on rehydrated speci- 
mens): Head-foot can be almost completely 
retracted into pallial cavity. Animal pig- 
mented with bright red, especially along right 
side of visceral mass. Tentacles short, flat- 
tened, with subcutaneous eyes situated 
somewhat laterally to center of bases, meet- 
ing above snout to form a "V." Snout large, 
cylindrical, with distal attachment disc with 
anterior notch and central proboscis open- 
ing. Foot smaller than snout, shapeless but 
with distinct opercular lobes and large oper- 
culum. One large pedal fold on each side of 
foot. Body of holotype consisting of 2.5 
whorls, of which pallial cavity occupies 0.5. 
Holotype female with large closed pallial 
oviduct, no penis. Gill with about 25 low 



VOLUME 102, NUMBER 1 



171 




Figs. 1-6. 1 , Trochostilifer eucidaricola new species, recently settled larva, from host 8, 0.67 mm; 2. Sabinella 
troglodytes, recently settled larva from host 10. 0.67 mm; 3-6, Trochostilifer eucidaricola new species; 3, apex 
of allotype, 0.65 mm of larval shell visible: 4. holotype. 4.68 mm; 5, female paratype from host 2, 3.6 mm; 6. 
male paratype from host 2. 2.6 mm. 



172 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 7-8. Trochostilifer eucidaricola, operculum, 
female and male, maximum diameter 1 .4 and 0.79 mm 
respectively. 

leaflets, occupying almost all space between 
osphradium and oviduct. Hypobranchial 
gland anteriorly occupying small, short, tri- 
angular area between gill and oviduct, pos- 



teriorly forming a very narrow band. Os- 
phradium a simple low ridge in shallow 
furrow to the left of and along whole length 
of gill. Operculum (Figs. 7-8) yellowish, thin, 
paucispiral with thickened part inside the 
nucleus serving as a muscular attachment. 

Male: In fully grown pairs it is less than 
0.6 the size of the female. Shell consisting 
of up to 2.5 teleoconch whorls of shape 
identical to corresponding whorls of female. 
Soft parts similar to those of female, but 
with a simple tentacle-like penis curving 
forwards, to the right and then backwards 
along right side of pallial cavity. No trace 
of female reproductive organs seen. Height 
of allotype 2.72 mm. 

Remarks.— Specimens of T. eucidaricola 
and Sabinella troglodytes can easily be dis- 
tinguished by the apical teleoconch whorls 
which are strongly shouldered in T. euci- 
daricola and evenly convex in S. troglo- 




Figs. 9-11. 
respectively. 



Sabinella troglodytes, female (Fig. 9) and two males from host 3, height 4.6, 2.2, and 2.3 mm 



VOLUME 102, NUMBER 1 



173 




Figs. 12-15. 12, Epidermal fold formed by the host, normally covering distal part of snout of Sabinella 
troglodytes, detail of lower left scar in Fig. 13; 13, Spine of Eucidaris tribuloides with deformed (cf. Fig. 15) gall 
caused by S. troglodytes, apical view. The spine has formed 3 "secondary" spines at a right angle to the axis of 
the original spine. Four attachment scars of epidermal tissue visible. Maximum diameter of spine 5 mm; 14, 
Eucidaris tribuloides, host 1, with male and female of Trochostilifer eucidaricola. Primary spine just to the right 
and dorsal to the pair has been removed to show the pair better. Diameter of test 20 mm; 15, normally formed 
gall caused by Sabinella troglodytes, host 1 1, length 9.7 mm. 



dytes. Monogamus minibulla (Olsson & 
McGinty, 1958) bears some resemblance to 
T. eucidaricola in having the first part of the 
teloconch distinctly shouldered, but is a 



much smaller species, ca. 1.5 mm; it has a 
larval shell of about 1.0 whorl and is para- 
sitic on the sea urchin Echinometra lucunter 
(L.) (Waren, unpub. data). 



174 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Among the described species of Trochos- 
tilifer only T. striatus bears any resemblance 
to T. eucidaricola. The other species of the 
genus have a shell of typical "Trochus shape" 
and live in galls in the spines of their hosts. 
Trochostilifer striatus can, however, easily 
be distinguished by having a much more 
distinct spiral sculpture, a larval shell of 1.5 
whorls and by having evenly convex apical 
whorls. 

The newly settled larva (Fig. 1) is virtually 
indistinguishable from that of Sabinella 
troglodytes (Fig. 2), but it is possible that 
they can be distinguished by the preferred 
site on the host, because the specimen in 
Fig. 2 had already started to deform the 
spine where it was found. This is also the 
basis for our determination of these two 
specimens. These determinations are not 
contradicted by the shape and size of the 
larval shells of specimens with enough post- 
larval growth to allow identification by the 
shape of the teleoconch. 

Several dozen specimens of the host ur- 
chin were examined and of these 8 were 
parasitized by this new species. These spec- 
imens are enumerated in Table 1. During 
the examination two more species of eu- 
limids were found on Eucidaris, viz. Sabi- 
nella troglodytes (Thiele, 1925) (Figs. 9-13, 
1 5) and specimens of an undescribed, small, 
slender, straight- sided species. Because of 
problems with generic assignment of such 
species, and because they seemed juvenile, 
the latter are left undescribed. 

A few years ago the senior author ex- 
amined all material of Eucidaris tribuloides 
in USNM and found several specimens par- 
asitized by Sabinella troglodytes from the 
Caribbean area. No specimens of Trochos- 
tilifer eucidaricola were then noticed and we 
assume it to be a more rare and/or locally 
distributed species. 

Sabinella troglodytes lives in galls of the 
spines of the host and has been reported in 
the literature several times (Waren 1984). 
We figure the latter species for comparison, 



together with two galls. These specimens are 
also listed in Table 1 . 

Trochostilifer eucidaricola lives attached 
to the test of the host by the disc-shaped 
distal part of the snout, and it seems very 
unlikely that it is able to move. In the center 
of the scar on the test of the host there is 
a very narrow hole, probably caused by the 
proboscis, but in these dried specimens it 
proved impossible to trace the proboscis to 
find the precise way of parasitism. The spec- 
imens were almost always attached in cou- 
ples, between a row of primary spines and 
the peristome. In two cases also the egg cap- 
sules, small globular sacs, were present, at- 
tached to the test by a short stalk. Each 
capsule contained 100-300 eggs, embryos, 
or veliger larvae ready to hatch. The di- 
ameter of the shell of the veligers, consisting 
of about 0.5 whorls is about 90 iim. This 
clearly shows that the species has plankto- 
trophic development. 

In the five cases when the sex of the spec- 
imens in a pair could be determined, the 
couples consisted of a small male and a larg- 
er female. In one case the sex could not be 
determined, because the specimens had not 
begun to develop a penis or pallial oviduct. 
A single specimen was found only on one 
host, which clearly shows that the distri- 
bution is not random, but that larvae prefer 
to settle on a host already parasitized by a 
single specimen. 

Acknowledgments 

We thank Mrs. Ineke Peeters, Aruba, who 
collected the sea urchins and most kindly 
allowed us to describe their molluscan fau- 
na, and Frere Fredericus, who forwarded 
the material to ZMA. 



Literature Cited 

Hedley, C. 1900. Studies on Australian Mollusca. 

2. — Proceedings of the Linnean Society of New 

South Wales 26:16-25. 
Olsson, A. A. , & T. L. McGinty . 1958. Recent marine 



VOLUME 102, NUMBER 1 



175 



mollusks from the Caribbean coast of Panama 
with the description of some new genera and 
species.— Bulletins of American Paleontology 
39(177):l-58. 

Thiele, J. 1925. Gastropoda der deutschen Tiefsee- 
Expedition. II. Teil. — Wissenschaftliche Ergeb- 
nisse der deutschen Tiefsee-Expedition auf dem 
Dampfer "Valdivia" 1898-1899 7(2):36-382. 

Waren, A. 1980. Descriptions of new taxa of Eulim- 
idae (Mollusca, Prosobranchia) with notes on 
some previously described genera.— Zoologica 
Scripta 9:283-306. 



. 1 984. A generic revision of the family Eulim- 

idae (Gastropoda, Prosobranchia).— The Jour- 
nal of Molluscan Studies, Supplement 13. 96 
PP. 

(AW) Swedish Museum of Natural His- 
tory, Box 50007, S- 10405 Stockholm, Swe- 
den; (RM) Zoological Museum, Postbus 
20125, 1000 HC Amsterdam, The Neth- 
erlands. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 176-248 



SPRINGSNAILS (GASTROPODA: HYDROBIIDAE) 

OF OWENS AND AMARGOSA RIVER 

(EXCLUSIVE OF ASH MEADOWS) DRAINAGES, 

DEATH VALLEY SYSTEM, CALIFORNIA-NEVADA 

Robert Hershler 



Abstract. —Thirteen springsnail species (9 new) belonging to Pyrgulopsis Call 
& Pilsbry, 1886, and Tryonia Stimpson, 1865 are recorded from the region 
encompassing pluvial Owens and Amargosa River (exclusive of Ash Meadows) 
drainages in southeastern California and southwestern Nevada. Discriminant 
analyses utilizing shell morphometric data confirmed distinctiveness of the 
nine new species described herein, as: Pyrgulopsis aardahli, P. amargosae, P. 
owenensis, P. pertubata, P. wongi, Tyronia margae, T. robusta, T. rowlandsi, 
and T. salina. Of the 22 springsnails known from Death Valley System, 1 7 
have very localized distributions, with endemic fauna concentrated in Owens 
Valley, Death Valley, and Ash Meadows. A preliminary analysis showed only 
partial correlation between modern springsnail zoogeography and configuration 
of inter-connected Pleistocene lakes comprising the Death Valley System. 



This constitutes the second part of a sys- 
tematic treatment of springsnails from the 
Death Valley System, a large desert region 
in southeastern California and southwest- 
ern Nevada integrated by a series of lakes 
during Pleistocene times. An earlier paper 
(Hershler & Sada 1987) dealt with the Ash 
Meadows faunule, while this document pro- 
vides descriptions of fauna collected during 
1985-1987 survey of much of remaining 
portions of the System, including waters in 
Mono, Adobe, Long, Owens, Indian Wells, 
Panamint, and Death Valleys; Amargosa 
River drainage; and some areas adjacent to 
the above (Fig. 1). A brief discussion of 
springsnail zoogeography also is provided, 
although a more extensive treatment will be 
given following survey of remaining por- 
tions of Death Valley System (notably Mo- 
jave River drainage) and additional periph- 
eral areas. 



List of Recognized Taxa 

Pyrgulopsis aardahli, new species. 

P. amargosae, new species. 

P. micrococcus (Pilsbry, 1893). 

P. owensensis, new species. 

P. perturbata, new species. 

P. cf. stearnsiana (Pilsbry, 1899). 

P. wongi, new species. 

Tryonia margae, new species. 

T. protea (Gould, 1855). 

T. robusta, new species. 

T. rowlandsi, new species. 

T. salina, new species. 

T. variegata Hershler & Sada, 1987. 

Materials and Methods 

Localities visited, consisting of low- to 
mid-elevation (<2500 m) springs and pe- 
rennial streams, are shown in Figs. 2-7 and 
listed in the Appendix. Snails were found 



VOLUME 102, NUMBER 1 



177 



af c£) 



"V 



\ 



(-) 



\ 



\ 

Mono Valley "vS^, 






Adobe Valley 



ft 



r ; 



\ 






\ 



\) 



Long Valley ^\ 



36° 





N 



i^ 



\ 



s 




«* 


u 








\ **© Asn Meadows 






-a 




o 

s 


^ 




% 


\ 




< 

9 


\ 




•< 


\ 



f \ 

( \ 

V.' 

i. Searles Valley 



1. V 



10 10 30 50 



i ' i ' ' ■ i_ 



-i i Kilometers 



118 c 



117< 



Fig. 1 . Map showing desert basins of southeastern California and southwestern Nevada comprising study 
area. Stippled areas indicate mountain ranges. 



in springs of varying sizes and appearance 
as well as low- to moderate-energy (spring- 
fed) streams. Photographs of representative 
sites are in Figs. 8 and 9. 

Snails were relaxed in the field with men- 
thol crystals, fixed in 4% buffered formalin 
and preserved in 70% ETOH. Material was 
collected by author unless otherwise indi- 



cated. Water temperature and conductivity 
were measured with a temperature-com- 
pensated, HACH 16300 conductivity me- 
ter. 

Methods of anatomical study and pho- 
tography of shells and other morphologic 
features are routine (Hershler & Sada 1987). 
Generalized radular formulae are based on 



178 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1.— Selected shell parameters for Pyrgulopsis species. Shell height (SH) and width (SW) are given in 
mm. Data for each population include mean values (upper line) and standard deviations (lower). Number of 
populations and specimens for which data were obtained are given in parentheses. NW = number of whorls, T 
= translation rate, D = distance of generating curve from coiling axis, AS = aperture shape, W = whorl expansion 
rate. 











Parameter 








Species 


NW 


SH 


sw 


T 


D 


AS 


w 


P. aardahli 


3.98 


2.90 


2.20 


4.24 


0.56 


1.15 


2.11 


(1-15) 


0.15 


0.21 


0.20 


0.54 


0.04 


0.04 


0.19 


P. amargosae 


3.86 


1.97 


1.45 


4.43 


0.57 


1.14 


2.03 


(3, 45) 


0.20 


0.28 


0.25 


0.67 


0.05 


0.06 


0.21 


P. crystalis 


3.17 


2.02 


1.95 


3.44 


0.50 


1.18 


2.56 


(1,3) 


0.29 


0.24 


0.18 


0.32 


0.05 


0.06 


0.76 


P. erythropoma 


3.46 


2.00 


1.82 


3.52 


0.56 


1.15 


2.75 


(5,68) 


0.27 


0.31 


0.24 


0.64 


0.05 


0.06 


0.87 


P. fairbanksensis 


3.29 


2.67 


2.23 


4.23 


0.61 


1.09 


2.80 


(1, 14) 


0.34 


0.21 


0.15 


1.03 


0.09 


0.13 


0.07 


P. isolata 


4.00 


2.75 


2.18 


3.81 


0.58 


1.12 


2.09 


(1,13) 


0.14 


0.13 


0.11 


0.36 


0.03 


0.04 


0.20 


P. micrococcus 


4.16 


2.16 


1.42 


5.17 


0.62 


1.19 


1.89 


(27, 362) 


0.33 


0.31 


0.17 


1.05 


0.06 


0.08 


0.28 


P. nanus 


3.55 


1.74 


1.53 


3.52 


0.55 


UQ9 


2.44 


(3, 42) 


0.24 


1.74 


1.53 


3.52 


0.55 


1.09 


0.34 


P. owensensis 


3.61 


2.16 


1.62 


4.56 


0.59 


1.16 


2.08 


(6, 86) 


0.26 


0.28 


0.18 


0.98 


0.05 


0.05 


0.24 


P. perturbata 


4.37 


3.29 


2.22 


5.26 


0.62 


1.18 


2.01 


(3, 39) 


0.19 


0.39 


0.31 


0.85 


0.62 


1.18 


0.19 


P. pisteri 


3.97 


2.31 


1.99 


3.46 


0.54 


1.10 


2.37 


(2,31) 


0.27 


0.18 


0.13 


0.45 


0.04 


0.04 


0.31 


P. cf. stearnsiana 


4.14 


2.52 


1.68 


5.73 


0.61 


1.24 


1.97 


(2, 25) 


0.15 


0.18 


0.12 


1.21 


0.05 


0.06 


0.21 


P. wongi 


3.86 


2.11 


1.62 


4.10 


0.59 


1.19 


2.23 


(19,286) 


0.27 


0.39 


0.24 


0.70 


0.05 


0.05 


0.29 



examination of SEM micrographs from 
varying numbers of populations, and are 
presented in following order of tooth types: 
centrals, laterals, inner marginals, outer 
marginals. 

Whorl counts, standard measurements, 
and Raupian parameters were obtained from 
selected series of live-collected, adult spec- 
imens, recognizable by their complete and 
thickened inner shell lips. After whorls were 
counted (NW), shells were imbedded in clay 
with aperture facing up and coiling axis per- 
pendicular to the observer's line of sight. 
Shell outlines were traced using camera lu- 
cida (25 x or 50 x ) and points on these dig- 



itized for calculation of the following pa- 
rameters (largely following Kohn & Riggs 
1975): shell height (SH), shell width (SW), 
length of body whorl (LBW), width of body 
whorl (WBW), aperture length (AL), aper- 
ture width (AW), translation rate (T), whorl 
expansion rate (W), distance of generating 
curve from coiling axis (D), and aperture 
shape (SA). In order to facilitate compari- 
sons, these parameters were also obtained 
from the entire set of shells of Ash Meadows 
springsnails used in an earlier morphomet- 
ric study (which involved generation of a 
subset of the above parameters using a 
somewhat different methodology; Hershler 



VOLUME 102, NUMBER 1 



179 



Table 2.— Selected shell parameters for Tryonia species. Shell height (SH) and width (SW) are given in mm. 
Data for each population include mean values (upper line) and standard deviations (lower). Number of popu- 
lations and specimens for which data were obtained are given in parentheses. NW = number of whorls, T = 
translation rate, D = distance of generating curve from coiling axis, AS = aperture shape, W = whorl expansion 
rate. 











Parameter 








Species 


NW 


SH 


sw 


T 


D 


AS 


w 


T. angulata 


5.70 


3.24 


1.56 


7.99 


0.58 


1.50 


1.66 


(3, 39) 


0.52 


0.37 


0.19 


1.29 


0.07 


0.08 


0.22 


T. elata 


5.99 


1.96 


0.82 


8.96 


0.59 


1.34 


1.42 


(2, 30) 


0.53 


0.29 


0.07 


1.15 


0.05 


0.08 


0.11 


T. ericae 


4.71 


1.40 


0.76 


6.42 


0.57 


1.25 


1.65 


(2, 19) 


0.66 


0.19 


0.06 


1.06 


0.06 


0.06 


0.11 


T. margae 


6.21 


2.78 


0.98 


10.80 


0.59 


1.48 


1.37 


(1,6) 


0.10 


0.12 


0.02 


0.93 


0.59 


0.03 


0.19 


T. protea 


5.33 


3.53 


1.59 


9.45 


0.66 


1.38 


1.51 


(1,6) 


0.52 


0.45 


0.13 


0.86 


0.04 


0.05 


0.19 


T. robusta 


4.22 


1.63 


1.06 


5.03 


0.60 


1.25 


1.81 


(2, 28) 


0.34 


0.31 


0.18 


0.83 


0.04 


0.04 


0.17 


T. rowlandsi 


4.34 


1.95 


1.19 


6.21 


0.62 


1.21 


1.69 


(1,16) 


0.18 


0.16 


0.07 


1.05 


0.06 


0.08 


0.15 


T. salina 


4.83 


2.89 


1.70 


5.72 


0.53 


1.25 


1.64 


(1,15) 


0.20 


0.14 


0.08 


0.60 


0.04 


0.03 


0.09 


T. variegata 


6.81 


4.27 


1.69 


8.93 


0.56 


1.50 


1.44 


(17,226) 


0.91 


1.18 


0.32 


2.02 


0.07 


0.11 


0.15 



& Sada 1987). Digitizing was done using 
CONCH (methodology described in Chap- 
man et al., in prep) and a GTCO Micro- 
Digi Pad 1 2 x 1 2 linked to a KAYPRO 2000 
microcomputer. 

Descriptive statistics for all morpholog- 
ical variables were obtained for each pop- 
ulation and species, and a selection of these 
are summarized (by species) in Tables 1 and 
2. Discriminant analysis (DA) incorporated 
all of the shell morphometric parameters 
and were constructed using all specimens 
measured for each species (considered as 
separate groups). All computations were 
performed using SYSTAT (Wilkinson 1986) 
on an IBM-AT. 

Systematics 

Generic diagnoses are in Hershler & Sada 
(1987). Species accounts include common 
name, synonomy, material examined (lo- 
calities numbered [in parentheses] as in Ap- 



pendix), diagnosis, description, remarks, 
type locality, distribution and habitat, 
etymology, and remarks. Asterisked (*) lots 
were used for shell measurements. Descrip- 
tions emphasize distinctive features from 
shell, radula, body pigmentation, and re- 
productive systems of both sexes. 

Family Hydrobiidae Troschel, 1857 

Genus Pyrgulopsis Call & Pilsbry, 1886 

Pyrgulopsis aardahli, new species 

Benton Valley springsnail 

Figs. 10-14 

Material examined.— California: Mono 
Co.; Spring at Bramlette Ranch (87), USNM 
860406 (holotype), *857951 (paratypes), 21 
Apr 1987. 

Diagnosis.— A large-sized species with 
broadly conical shell. Penis with elongate 
filament, very small lobe, and ventral swell- 
ing. Penial glandular ridges, 1-3. 

Description.— Shell (Fig. 10) 2.6-3.4 mm 



180 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Sampling localities in southern Owens Val- 
ley and western Indian Wells Valley. Solid lines indi- 
cate selected elevation contours (light) and modern 
drainage (dark; stipple indicates lakes); dashed lines 
indicate historic drainage, including dry lake beds. 



high, height/width, 140-170%. Whorls, 
3.75-4.25, moderately convex, moderately 
shouldered. Body whorl inflated, height 76- 
82% of that of shell. Aperture ovate, aper- 
tural plane near-parallel to coiling axis (Fig. 
10c). Inner lip thickened, slightly reflected, 
adnate to small portion of or slightly sep- 
arated from body whorl. Outer lip straight, 
thin. Umbilicus moderately open. 

Dark, grey-black epithelial (melanic) pig- 
ment on most of snout (to just posterior of 
cephalic tentacles), along anterior edge of 
foot, on operculigerous lobe (Fig. 11). Cen- 
tral portions of sides of head/foot lightly 




Fig. 3. Sampling localities in northern Owens Val- 
ley. 



dusted or unpigmented. Tentacles unpig- 
mented except for dark ring along bases. 
Brown-black subepithelial pigment gran- 
ules sometimes forming dark band along 
posterior edge of "neck." 

Radular (Fig. 12) formula: 5-1-5/1-1, 
2(3)-l-3, 22-24, 28-32 (from paratypes). 
Central tooth broadly trapezoidal; basal 
process moderately excavated. Penis (Fig. 
1 3b-e) large (extending beyond mantle col- 
lar), thin, considerably longer than wide. 
Filament slender, moderate in length. Lobe 
reduced or absent, with blunt distal edge. 
Large, elongate glandular ridge borne on 
elongate swelling of ventral penial surface. 
Single, smaller ridges sometimes found on 



VOLUME 102, NUMBER 1 



181 



.-^ 



{>. 






J51 M 0no Valley /y %<&j 




i l I i 



Fig. 4. Sampling localities in Mono, Long, and 
Adobe Valleys. 



tip of lobe, and on dorsal penial surface 
proximal to base of filament. Filament 
darkly pigmented with subepithelial gran- 
ules (Fig. 1 3b). Capsule gland sub-equal to 
albumen gland (Fig. 13a). Seminal recep- 
tacle minute, positioned entirely anterior to 
bursa copulatrix. Bursa pear-shaped, mod- 
erate in size relative to pallial oviduct, po- 
sitioned partly posterior to albumen gland. 

Type locality. —Spring at Bramlette Ranch 
in Benton Valley, Mono County, California. 

Distribution and habitat.— Endemic to 
type locality (Fig. 14). Common in dense 
watercress in uppermost portion of outflow 
of small, highly degraded spring (20°C, 1 90 
micromhos/cm). 

Etymology. —Named after Jeffrey B. Aar- 
dahl, Bureau of Land Management (Cali- 
fornia Desert District), who provided di- 
verse forms of assistance throughout the 
tenure of this project. 

Remarks.— Distinguished from similar P. 
perturbata by shell appearance and highly 
reduced penial lobe. 




Fig. 5. Sampling localities in Saline, Eureka, and 
Deep Springs Valleys. 



Pyrgulopsis amargosae, new species 

Amargosa springsnail 

Figs. 15, 16, 17a, b, 18, 19 

Material Examined. —California: San 
Bernardino Co.; Saratoga Spring (204), 
USNM 860401 (holotype), *853515 (para- 
types), 7 Feb 1985; 857972, 10 Jul 1986.- 
Inyo Co.; Spring crossing path on south side 
of Amargosa Gorge (208), *853516, 13 Mar 
1986; 857973, 16 Mar 1987. -Spring feed- 
ing Amargosa River in Amargosa Gorge 
(208), 853517, 13 Mar 1985. -Spring in 
marsh east of Grimshaw Lake (209), 
*853518, 13 Mar 1985. 

Diagnosis.— A small-sized species with 
globose to low conical shell. Penis large rel- 
ative to head/foot; penial lobe reduced; fil- 
ament long, wide. Penial glandular ridges, 
0-2. 

Description.— Shell (Fig. 15) 1.5-2.7 mm 
high, height/width, 1 10-160%. Whorls, 3.5- 



182 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



f^rV 




Indian Wells 
Valley 



I I I I I Km 



Fig. 6. Sampling localities in Panamint and north- 
ern Indian Wells Valleys. 



4.25, convex, moderately shouldered. Body 
whorl 74-85% of shell height. Apex, parts 
of teleoconch whorls often appearing white 
due to erosion. Aperture broadly ovate, 
apertural plane near-parallel to coiling axis 
(Fig. 15b). Inner lip thickened, reflected 
above, adnate to small portion of or slightly 
separated from body whorl. Outer lip 
straight, thin. Umbilicus chink-like to 
broadly open. Spiral striations sometimes 
prominent on teleoconch. 

Dark, brown-black epithelial pigment on 
most of snout (except distal tip), along an- 
terior and posterior edges of "neck" (central 
area lighter), on part or all of operculigerous 
lobe. Tentacles lightly pigmented. 

Radular (Fig. 16) formula: 5(6)-l-5(6)/ 
1-1, 3-1-4, 27-28, 24-26 (from paratypes). 
Central tooth broadly trapezoidal; basal 
process moderately excavated. Penis (Figs. 



1 7a, b, 1 8b-e) thickened, longer than wide. 
Filament much longer than lobe, about equal 
to remaining penis length. Lobe narrow, 
variable in shape (Fig. 18b-e). Filament 
having highly muscular appearance; surface 
lined with weak striations. One or two small 
glandular ridges sometimes present at tip of 
lobe. Dark, grey-black subepithelial pig- 
ment filling filament length almost to distal 
tip as well as portions of remaining distal 
penis (Fig. 18b). Albumen and capsule 
glands about equal in length (Fig. 18a). Sem- 
inal receptacle minute, positioned well an- 
terior to bursa copulatrix. Bursa copulatrix 
club-shaped, small, positioned partly pos- 
terior to albumen gland. 

Type locality. — Saratoga Spring, southern 
Death Valley, San Bernardino Co., Califor- 
nia. 

Distribution and habitat. —Found in a few 
near-brackish springs along lower course of 
Amargosa River at Saratoga Spring, Amar- 
gosa Gorge, and Grimshaw Lake area (Fig. 
19). Type locality an extensive marsh area 
(with several large, open pools) fed by sev- 
eral small springs. Snails were common in 
westernmost spring, both on rocks and sub- 
mergent macrophytes in small springpool 
(28°C, 5000 micromhos/cm), and on emer- 
gent sedges and firm bottom in shallow 
stream outflow. At other localities, consist- 
ing of small rheocrenes (16-27°C, 2000-10, 
150 micromhos/cm), snails were common 
on all substrates. 

Etym ology. —Referring to distribution 
along Amargosa River. 

Remarks. —Distinguished from P. micro- 
coccus by globose shell; large penis; large 
penial filament; small, narrow penial lobe; 
and highly reduced glandular ridge. Ranges 
of these very similar snails (previously con- 
fused by Hershler and Sada 1987:788) are 
contiguous. 

Pyrgulopsis micrococcus (Pilsbry, 1893) 

Oasis Valley springsnail 

Figs. 17c, d, 20-25 

Amnicola micrococcus Pilsbry in Stearns, 
1893:277. 



VOLUME 102, NUMBER 1 



183 




Fig. 7. Sampling localities in Death Valley and Amargosa Basin. 



Material examined.— Nevada: Nye Co.; 
Spring in Oasis Valley (204), USNM 
*850297, 18 Nov 1985; 857961, 16 Mar 
1987.-Cane Spring (239), *857936, 10 July 
1986.— California: Inyo Co.; Northernmost 
of Tecopa Hot Springs (210), *853502, 12- 
13 Mar 1985; 857962, 16 Mar 1987.- 
Shoshone Spring (211), *853501, 12 Mar 
1985; 857962, 16 Mar 1987. -Grapevine 
Springs (241): northwest-most spring in 
complex, 853503, 1 Mar 1985; Spring 
emerging below limestone bench northwest 
of ranch, 853504, 26 Feb 1985; Spring on 
bench above ranch, 853505, 25 Feb 1985; 
Spring at ranch, *853506, 25 Feb 1985; 
857964, 14 Mar 1985. -Spring 1.6 km east 
of Scotty's Castle (242), *853507, 26 Feb 
1985; 857965, 14 Mar 1987,-Hanaupah 
Spring (222), *853508, 16 Feb 1985; 



857966, 21 Mar 1987. -Lower spring in 
Johnson Canyon (220), *853509, 18 Feb 
1985; 857967, 21 Mar 1987. -Stream in 
Jail Canyon (196), *853513, 14 Feb 1985; 
857970, 19 Mar 1987. -Stream in Surprise 
Canyon (194), *857937, 19 Mar 1987; 
Stream in Pleasant Canyon (190), *857940, 
19 Mar 198 7. -China Garden Spring (184), 
853511, 9 Feb 1985. -Stream below Dar- 
win Falls (184), 857969, 23 Mar 1987.- 
Spring above Darwin Falls (184), *853512, 
10 Feb 1985. — Stream in Snow Canyon 
(182), *857938, 16 Sep 1987, J. Aardahl!- 
Stream in Knight Canyon (180), *857939, 
14 Apr 1987. — Tennessee Spring (178), 
*853514, 4 Mar 1985; 857971, 13 May 
1987.-SalineMarsh(152),*853510, 10 Feb 
1985; 857968, 28 Mar 1987. 
Diagnosis.— A small-sized species, with 



184 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




I 



j* 






Fig. 8. Photographs of selected springsnail localities: a, stream in Cottonball Marsh, Death Valley, type 
locality of T. salina (3-10-87; photo by P. Rowlands): b. Grapevine Springs, Death Valley, type locality of T. 
margae and T. rowlandsi is on the large travertine mound (3-14-87); c, Saline Marsh, Saline Valley (3-29-87); 
d, Sand Canyon stream. Indian Wells Valley (3-26-87); e, Fish Slough. Owens Valley, spring sources (including 
type locality of P. perturbata) visible in lower portion of photograph (ca. 1975; aerial photo by P. Pister); f, 
"NE" Spring at Fish Slough (2-7-85). 



globose to ovate-conic shell. Penis with 
moderate- sized lobe; distal edge of lobe usu- 
ally ornamented with glandular ridge. 

Description. -Shell (Figs. 20, 21) 1.1-3.1 
mm high, height/width, 99-190%. Whorls. 
3.25-5.0, well-rounded, with slight shoul- 
ders. Spire often irregularly convex due to 



bulging of whorls. Body whorl 65-89% of 
shell height. Aperture ovate, apertural plane 
near-parallel to coiling axis (Fig. 21b). Inner 
lip moderately thickened and reflected, usu- 
ally slightly separated from body whorl. 
Outer lip straight, thin. Umbilicus chink- 
like to moderately open. 



VOLUME 102, NUMBER 1 



185 




Fig. 9. Photographs of springsnail localities: a, Warm Springs. Owens Valley (4-19-87); b. Small spring S of 
Warren Lake, Owens Valley (4-18-87); c. Spring in Owens Gorge. Owens Valley (5-7-87): d. Hot Creek. Long 
Valley (4-23-87). 



186 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 10. SEM micrographs of P. aardahli from spring at Bramlette Ranch: a, Holotype, USNM 860406 (shell 
height, 3.26 mm); b-d, Paratypes, USNM 857951 (scale of "b" and "c" as in "a"; bar = 150 /mi). 



VOLUME 102, NUMBER 1 



187 



Head/foot epithelial pigment ranging from 
absent to near-uniformly black (lighter on 
central portions of sides of "neck"). 

Radular (Figs. 22, 23) formula: (4-7)-l- 
(4_7)/l_l, (2-4)-l-(3-6), 17-26, 19-32 
(from numerous populations). Central tooth 
moderately to broadly trapezoidal; basal 
process moderately excavated. 

Penis (Figs. 1 7c, d, 24b-i) barely extend- 
ing beyond mantle collar, moderately thick- 
ened, longer than wide. Filament slender, 
usually extending well beyond edge of lobe. 
Lobe highly variable in size and shape, ta- 
pering slightly distally. Glandular ridge 
(rarely absent) small, positioned on or near 
distal edge of lobe. Capsule gland smaller 
than albumen gland (Fig. 24a). Seminal re- 
ceptacle small, positioned largely anterior 
to bursa copulatrix. Bursa pear-shaped, 
small to fairly large relative to pallial ovi- 
duct, most of length positioned on albumen 
gland. 

Type locality. — Small spring in Oasis Val- 
ley, Nye County, Nevada (precise location 
unknown). 

Distribution and habitat.— A widespread 
form (contrary to Taylor 1985:317) found 
in small springs and stream outflows (see 
Figs. 8b, c) of Amargosa River drainage as 
well as interior drainage in Death, Pana- 
mint, and Saline Valleys (5-3 2°C, 300-4000 
micromhos/cm) (Fig. 25). Typically com- 
mon on stone, bits of travertine, watercress, 
and plant debris. 

Pyrgulopsis owensensis, new species 

Owens Valley springsnail 

Figs. 26a-d, 27-32 

Material examined. —California: Inyo 
Co.; Spring ca. 1.6 km south of Mule Spring 
(92), USNM 853541, 5 Dec 1987, D. Giu- 
liani.-Mule Spring (93), *853540, 19 Nov 
1985, D. Wong; 857983, 20 Apr 1987.- 
Spring at Graham Ranch (94), *853543, 8 
Feb 1985; 857985, 19 Apr 1987. -Wilk- 
erson Springs (96), *853542, 8 Feb 1985; 
857984, 18 Apr 1987,-Spring at Toll House 




Fig. 1 1 . Right lateral view of head-foot of P. aar- 
dahli, USNM 85795 1 , spring at Bramlette Ranch. Stip- 
ples indicate pigmented areas. 



(97), *857950, 26 Apr 1987. -Warm Springs 
(98), Main spring, *853544, 7 Feb 1985, 
857988, 19 Apr 1987; Small spring north 
of above, 853545, 7 Feb 1985; 857989, 19 
Apr 1987. Mono Co.: Stream in canyon S 
of Piute Creek (104), 860404 (holotype), 
*857955 (paratypes), 8 May 1987,-Springs 
on bench south of Piute Creek (103), North 
spring, 857986, 21 Apr 1987; South spring, 
857987, 22 Apr 1987. 

Diagnosis.— A small- to moderate-sized 
species with globose to ovate-conic shell. 
Penis large relative to head/foot, filament 
short, lobe enlarged, ventral swelling pres- 
ent. Penial glandular ridges, 2-6. 

Description.— Shell (Figs. 26a-d, 27, 28) 
1.5-2.8 mm high, height/width, 1 10-160%. 
Whorls, 3.0-4.25, well-rounded, sutures 
slightly impressed. Protoconch sometimes 
slightly tilted. Body whorl inflated, height 
74-90% of that of the shell. Apical whorls 
sometimes eroded, shell often covered by 
deposits. Aperture ovate, angled above; 
moderate to large in size. Aperture plane 
near-parallel to somewhat tilted relative to 
coiling axis (Figs. 27b, 28b). Inner lip slight- 
ly to considerably thickened and reflected, 
adnate to or (more commonly) slightly sep- 
arated from body whorl. Outer lip thin, 
straight or slightly sinuate. Umbilicus chink- 
like to moderately open. 

Dark brown or grey-black epithelial pig- 
ment on snout, peripheral portions of 



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Fig. 12. Radula of P. aardahli, USNM 857951, spring at Bramlette Ranch: a, Centrals (bar = 12 /mi); b, 
Laterals (bar = 8.6 /mi); c, Inner marginal (bar = 6.7 /mi); d, Outer marginals (bar = 6.0 /mi). 



"neck," and dorsal surfaces of tentacles. Radular (Figs. 29, 30) formula: 5(6)— 1— 

Dark subepithelial granules prominent in 5(6)/ 1-1, 3-1-3(4), 18-23, 25-37 (from 

central areas of sides and sometimes along three populations). Central tooth broadly 

posterior edges of "neck." trapezoidal; basal process well-excavated. 



VOLUME 102. NUMBER 1 



189 



Penis (Fig. 3 lb— i) extending well beyond 
mantle collar, elongate. Narrow filament 
shorter than lobe. Lobe massive, tapering 
only slightly, distal edge blunt. Edge of lobe 
ornamented with single elongate or two 
shorter glandular ridges. Dorsal penis some- 
times having small glandular ridges on lobe 
and or near base of filament. Ventral swell- 
ing small, located filament base. Ridge or- 
namenting swelling variably-sized. Fila- 
ment darkly pigmented with sub-epithelial 
granules along most of length (Fig. 31b). 
Capsule gland equal to (Fig. 31a) or slightly 
smaller than albumen gland. Seminal re- 
ceptacle minute, positioned largely or en- 
tirely anterior to bursa. Bursa pear-shaped, 
small- to moderate-sized, with about half 
of length posterior to albumen gland. 

Type locality.— Stream in canyon S of 
Piute Creek. Mono Co.. California. 

Distribution and habitat. — Found along 
escarpments of White and Inyo Mountains 
east side of Owens Valley (Fig. 32). Habitat 
consisting of small springbrooks (10-29°C. 
300-780 micromhos cm) (Fig. 9a) where 
snails typically common in watercress and 
or on bits of travertine and stone. Syntopic 
with P. wongi in spring at Toll House. 

Etymology— -Referring to occurrence in 
Owens Valley. 

/femor&s.— Distinguished from similar P. 
pertwbata by shell appearance, more en- 
larged penial lobe, and (usually) more nu- 
merous penial glandular ridges. 

Pyrgulopsis perturbata, new species 

Fish Slough springsnail 

Figs. 26e-g. 33-36 

Material examined. —California: Mono 
Co.: Fish Slough: "Northwest Springs."" 
South spring (76). USNM 860407 (holo- 
type). *853546 (paratypes). 7 Feb 1985: 

857990. 24 Apr 198" 7 : "Northwest Springs." 
North spring (76). 853547. 7 Feb 1985: 

857991. 20 Apr 1987: "NE Spring" (77). 
*853548. 7 Feb 1985: 857992. 20 Apr 1987: 
"BLM Spring" (" 7 5). *853549. 7 Feb 1985. 




Fig. 13. Reproductive anatomy of P. aardahli. 
USXNi S5~951. spring at Bramlette Ranch: a. Left 
lateral view of pallial oviduct complex: b-e. Penes (b. 
c. dorsal: d. e. ventral). Stippled areas on penes indicate 
glandular ridges (enclosed stipples) and pigmented re- 
gions (open). Ag = albumen gland: Bu = bursa copu- 
latrix: Cg = capsule gland: Cga = capsule gland open- 
ing: Emc = posterior end of pallial cavity; Ov = oviduct: 
Pf =penial filament: Plo = penial lobe: Sr = seminal 
receptacle: Vc = ventral channel of capsule gland. 



Diagnosis.— A large-sized species with 
thickened, low-conical shell. Penis large, 
lobe and filament small, ventral swelling 
present. Penial glandular ridges. 3 or 4. 

Description. — Shell (Figs. 26e-g. 33) 2.7- 
4.0 mm high, height width. 13O-180°o. 
Whorls. 4.25-5.0. only slightly convex, with 
pronounced sub-sutural angulations. Body 
whorl inflated. 7 l-85°o of shell height. Ap- 
erture broadly ovate. Apertural plane tilted 
relative to coiling axis (Fig. 33b). Inner lip 
usually well-thickened and reflected, broad- 
ly adnate to body whorl. Outer lip straight. 
thin or moderately thickened. Umbilicus 
chink-like. 

Dark brown-black epithelial pigment 
prominent on snout, tentacles, and periph- 
eral portions of sides of head foot. Pigment 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 14. Map showing distribution of P. aardahli. 



either light or absent on central portions of 
sides of head/foot. 

Radular (Fig. 34) formula: 5-1-5/1-1, 3- 
1-4(5), 18-19, 29-31 (from paratypes). 
Central tooth broadly trapezoidal, basal 
process well-excavated. Penis (Fig. 35b-e) 
extending well beyond edge of mantle collar, 



thin, considerably longer than wide. Fila- 
ment both narrow and short, barely extend- 
ing beyond tip of lobe. Lobe tapering only 
slightly distally, bearing elongate glandular 
ridge along distal edge. Dorsal surface of 
penis with small ridge proximal to base of 
filament. Ventral swelling arising slightly 



VOLUME 102, NUMBER 1 



191 




Fig. 15. SEM micrographs of P. amargosae from Saratoga Spring: a. Holotype, USNM 860401 (2.28 mm); 
b, c (bar = 136 ptm), Paratypes, USNM 853515. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 16. Radula of P. amargosae, USNM 853515, Saratoga Spring: a, Centrals (bar =12 nm); b, Laterals 
(bar = 8.6 ^m); c, Inner marginals (bar = 8.6 Aim); d, Outer marginal (bar = 3.8 ^m). 



proximal to base of filament; ridge (rarely 
two) on swelling elongate. Filament dark- 
ened along much of length by sub-epithelial 
pigment granules. Capsule gland sub-equal 



to albumen gland (Fig. 35a). Bursa small, 
with ca. 50% of length posterior to albumen 
gland. Seminal receptacle minute, posi- 
tioned entirely anterior to bursa. 



VOLUME 102, NUMBER 1 



193 




Fig. 17. SEM micrographs of head-foot and dorsal penes of P. amargosae and P. micrococcus: a (bar = 0.43 
mm), b (bar = 0.38 mm). P. amargosae, USNM 853515. Saratoga Spring; c (bar = 0.3 mm), d (bar = 200 nm), 
P. micrococcus, USNM 857961, spring in Oasis Valley. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Type locality. —"Northwest Springs," 
south springpool, Fish Slough, Mono Co., 
California. 

Distribution and habitat. —Found in three 
of the four main springs in Fish Slough (ex- 
tinct in "BLM Spring") (Figs. 8e, f, 36). 
Springs issuing relatively cool (22-26°C), 
soft (200-360 micromhos/cm) water. Snails 
found only in small vestiges of rheocrene 
habitat at various small orifices in north- 
west springs (rare at both sites on vegetation 
and stones), and at beginning of outflow from 
"NE Spring" (common in Chara). 

Etymology. —Referring to various distur- 
bances to endemic locale, including severe 
reduction of original rheocrene habitat as 
unanticipated side effect of efforts to con- 
serve Owens pupfish (Miller and Pister 
1971). 

Pyrgulopsis cf. stearnsiana 

(Pilsbry, 1899) 

Figs. 37^0 

Paludestrina stearnsiana Pilsbry, 1899:1 24. 
Fontelicella stearnsiana.— Gregg & Taylor 

1965:108. 
Pyrgulopsis stearnsiana. — Hershler & 

Thompson 1987:30. 

Material examined. —California: Kern 
Co.; Stream in Sage Canyon (1), USNM 
*853520, 26 Mar 1987; 857975, 10 Nov 
1987. — Stream in Sand Canyon (6), 
*853519, 26 Mar 1987; 857974, 22 Dec 
1987, J. Aardahl. 

Diagnosis.— A small-sized species with 
low conical shell. Penis small relative to 
head/foot, penial lobe reduced, filament 
elongate, ventral swelling present. Penial 
glandular ridges, 1-3. 

Description.— Pertaining only to popula- 
tions in study area. Shell (Fig. 37) 2.0-2.9 
mm high, height/width, 130-170%. Whorls, 
4.0-4.5, convex. Body whorl ca. 75% of shell 
height. Shell surface encrusted with thick, 
black deposits. Aperture broadly ovate, 
somewhat angled above, about half as tall 




Fig. 18. Reproductive anatomy of P. amargosae: 
a, Left lateral view of pallial oviduct complex, USNM 
853515, Saratoga Spring; b, c, Dorsal penes, USNM 
853515, Saratoga Spring; d, e, Dorsal penes, USNM 
857973, spring in Amargosa Gorge, "b" through "e" 
drawn to same scale. 



as body whorl. Inner lip slightly thickened 
and reflected, adnate to small portion of or 
slightly separated from body whorl. Outer 
lip straight, thin; apertural plane slightly 
tilted relative to coiling axis (Fig. 37b). 

Dark, grey-black epithelial pigment on 
most of snout (except distal tip), proximal 
portion of cephalic tentacles, along anterior 
and posterior edges of "neck," part or all of 
operculigerous lobe. Pigment on central 
portions of sides of neck absent to dark (sub- 
epithelial pigment cluster dense in area). 

Radular (Fig. 38) formula: 5(6)-l-5(6)/ 
1-1, 2(3,4)-l-3(4,5), 23-24, 23-29 (from 
single population). Central tooth broadly 
trapezoidal; basal process moderately ex- 
cavated. Penis (Fig. 39b-e) rarely protrud- 
ing beyond edge of mantle collar, longer than 
wide. Filament slender, sub-equal to re- 
maining penis length. Reduced lobe scarcely 



VOLUME 102, NUMBER 1 



195 




Fig. 19. Map showing distribution of P. amargosae. 



projecting distal to base of filament. Tip of 
lobe often ornamented with small glandular 
ridge; similar, single ridges on dorsal penial 
surface about halfway from base of penis to 
base of filament, on ventral surface near 
(sometimes on) inner edge. The latter ridge 
enlarged, borne on low swelling (Fig. 39e). 



Filament with dark sub-epithelial pigment 
streak (Fig. 39b). Albumen gland sub-equal 
to capsule gland (Fig. 39a). Seminal recep- 
tacle small, positioned anterior to bursa 
copulatrix. Bursa copulatrix small relative 
to capsule gland, positioned partly posterior 
to gland. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 








Fig. 20. Camera lucida drawings of shells of P. micrococcus: a, USNM 857936, Cane Spring (2.24 mm); b, 
USNM 853501. Shoshone Spring (1.96 mm); c, USNM 853502, Tecopa Hot Springs (1.90 mm); d, USNM 
853506, Grapevine Springs (2.16 mm); e, USNM 853508, Hanaupah Spring (2.26 mm); f, USNM 853510, 
Saline Marsh (1.96 mm); g, USNM 853512, spring above Darwin Falls (1.94 mm); h, USNM 857937, Surprise 
Canyon stream (1.88 mm). 



Type locality. —Near Oakland, Alameda 
County, California. 

Distribution and habitat.— Distribution 
given by Taylor (1 98 1 : 1 52) as "Central Cal- 
ifornia, from Sonoma County to Monterey 
County along the coast, and inland in the 
foothills of the Sierra Nevada." Found in 
the study area in two moderate-sized, cool 
streams (16-20°C, 550 micromhos/cm) sep- 
arated by 25 km along eastern slope of Sier- 
ra Nevada in Indian Wells Valley (Fig. 40). 
Snails common in watercress and Char a. 

Remarks.— Shells of the Indian Wells 
Valley snails closely resemble P. stearnsiana 
from the San Francisco Bay area and their 
penes conform to a general description for 
the species provided by Gregg and Taylor 
(1965:107). Additional study, including ex- 
amination of anatomy of topotypes or near- 
topotypes, will be necessary to confirm 



identity of the snails from Indian Wells Val- 
ley. 

Pyrgulopsis wongi, new species 

Wong's springsnail 

Figs. A\~M 

Material examined. —California: Inyo. 
Co.; Spring at Little Lake, east of HW 395 
(10), USNM *853521, 9 Feb 1985. -Hog- 
back Creek (Monache Mtn. quad.) (20), 
*853522, 28 Jun 1985, D. Wong.-Summit 
Creek (21), 853523, 12 Apr 1987. -Spring 
ca. 3.0 km north of Summit Creek (21), 
*853524, 12 Apr 1987,-Spring at Cabin 
Bar Ranch (25), 853525, 12 Oct 1987, J. 
Goldberg and M. Shumway.— Lubkin Creek 
(34), 853526, 16 Apr 198 7. -Spring feeding 
Lubkin Creek (34), 857942, 25 Apr 1987,- 
Spring along east side of Tuttle Creek (35), 



VOLUME 102. NUMBER 1 



197 



a 





Fig. 21. SEM micrographs of P. micrococcus: a. USNM 853502. Tecopa Hot Springs (1.98 mm): b. USNM 
153512. spring above Darwin Falls: c. USNM 853512. Tecopa Hot Springs (bar = 120 ^m). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 22. Central radular teeth of P. micrococcus: a, USNM 857936, Cane Spring (bar = 7.5 /mi); b, USNM 
857963, Tecopa Hot Springs (bar = 7.5 /mi); c, USNM 857965, spring E of Scotty's Castle (bar = 7.5 /mi); d, 

USNM 857968, Saline Marsh (bar = 8.6 /an). 



857943, 30 Apr 1987. -Hogback Creek 
(Lone Pine quad.) (36), *853527, 8 Feb 1985; 
857976, 16 Apr 1987. -Boron Springs (40), 
*853528, 17 Apr 1987. -Stream in Charlie 
Canyon (43), *853529, 12 Jun 1987, D. 



Giuliani. — Spring on hill south of Warren 
Lake (57), *853530, 18 Apr 1987. -Spring 
in canyon south of Shannon Canyon (59), 
857944, 20 Apr 1987. -Spring along north 
side of upper Pine Creek (69), *853531, 24 



VOLUME 102, NUMBER 1 



199 




Fig. 23. Lateral and marginal radular teeth of P. micrococcus: a, Laterals, USNM 857936, Cane Spring (bar 
= 6.0 /mi); b, Laterals, USNM 857966, Hanaupah Spring (bar = 7.5 /mi); c, Inner marginals, USNM 857937, 
Surprise Canyon stream (bar = 5.0 /urn); d, Outer marginals, USNM 857963, Tecopa Hot Springs (bar = 4.3 
Mm). 



Apr 1987; 857977, 15 Sep 1987, D. Wong.- 
West spring in Birchim Canyon (70), 860403 
(holotype), *857941 (paratypes), 3 May 
1987.— East spring in Birchim Canyon (70), 



857945, 3 May 1987. -Spring in Owens 
Gorge (74), *853532, 14 May 1985, D. 
Wong; 857978, 8 May 1987. -French Spring 
(89), *857947, 4 May 1 987. -Barrel Springs 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 24. Reproductive anatomy of P. micrococcus: 
a, Left lateral view of pallial oviduct complex, USNM 
857961, spring in Oasis Valley; b-i, Penes (all dorsal 
except "i") (b, i, USNM 857936, Cane Spring; c, USNM 
853505, Grapevine Springs; d, USNM 857962, Shosh- 
one Spring; e, USNM 857963, Tecopa Hot Springs; f, 
USNM 857967, Johnson Canyon spring; g, USNM 
857971, Tennessee Spring; h, USNM 853512, spring 
above Darwin Falls). 



(91),*857948, 18 Aug 1987.-SpringatToll 
House (97), *857949, 26 Apr 1987. -Corral 
Springs (160), *853535, 8 Feb 1985; 857979, 
29 Mar 1987. -Antelope Spring (162), 
♦853536, 8 Feb 1985. Mono Co.; Spring 
along Marble Creek (82), *853533, 22 Apr 
1 987. — Springs at north end of Blind Spring 
Valley (86); North Spring, *857946, 21 Apr 
1987; South spring, 853534, 21 Apr 1987.- 
Layton Springs (122), *853537, 6 Feb 1985; 
857980, 23 Apr 1987.-River Springs (132), 
*853538, 6 Feb 1985; 857981, 21 Apr 1987. 
Diagnosis.— A small- to moderate-sized 
species with globose to low conical shell. 
Penis massive, with large filament and mod- 
erate-sized lobe. Ventral penis with two 
prominent swellings bearing glandular 
ridges. Glandular ridges, 7-12. 



Description.— Shell (Figs. 41, 42) 1.2-3.0 
mm high, height/width, 100-160%. Whorls, 
3.25-4.5, well rounded, with slightly an- 
gulated shoulders. Body whorl 76-87% of 
shell height. Aperture broadly ovate, angled 
above. Apertural plane slightly tilted rela- 
tive to coiling axis (Fig. 42c). Inner lip 
slightly thickened, usually only minimally 
reflected, either adnate to portion of or 
slightly separated from body whorl. Outer 
lip straight, thin. Umbilicus chink-like to 
moderately open. 

Dark grey-black epithelial pigment on 
snout, proximal tentacles, and peripheral 
portions of sides of head/foot. Entirety of 
head/foot often very dark. Central portions 
of sides of head/foot usually with densely 
clustered dark, subepithelial pigment gran- 
ules. 

Radular (Figs. 43, 44) formula: (4-7)- 1- 
(4_7)/l_l ? (2-5)-l-(3-6), 17-26, 20-34 
(from numerous populations). Central tooth 
relatively square-shaped; basal process well- 
excavated. Penis (Figs. 45b-l, 46) flattened, 
longer than wide. Filament fairly broad, 
short relative to length of remaining penis. 
Lobe slightly shorter than filament, broad, 
often widening distally. Glandular ridge ar- 
rangement typically as follows: 1) elongate 
ridge lining edge of lobe; 2) similar, stalked 
ridge on ventral surface just proximal to 
lobe; 3) similar ridge (rarely very reduced) 
on fleshy projection of ventral surface prox- 
imal to above; 4) elongate ridge covering 
proximal-most two-thirds of dorsal fila- 
ment; 5) very elongate ridge on dorsal sur- 
face curving from just proximal to filament 
to lobe; 6) shorter ridges on dorsal surface 
along right side of (5) and in area between 
base of filament and edge of lobe. Modifi- 
cations of above include addition of a few 
small ridges on central dorsal surface, ad- 
dition of a single small ridge (sometimes 
stalked) on ventral surface, apparent frag- 
mentation of ridges (particularly "1," "5"); 
apparent merging of adjacent ridges on dor- 
sal surface. Filament darkened with sub-ep- 
ithelial pigment along most of length. Cap- 
sule gland sub-equal to albumen gland (Fig. 



VOLUME 102, NUMBER 1 



201 



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Fig. 25. Map showing distribution of P. micrococcus. Snail occurs at 15 localities in Ash Meadows. 



45a). Seminal receptacle small, positioned 
lateral to anterior bursa copulatrix. Bursa 
pear-shaped, massive, with about one-third 
of length posterior to albumen gland. 

Type locality. — Spring in Birchim Can- 
yon north of Bishop, Inyo Co., California. 

Distribution and habitat. —Widespread in 
Owens Valley along eastern escarpment of 
Sierra Nevada from Pine Creek south to 



Little Lake, and along western side of val- 
ley from French Spring to Marble Creek. 
Also found in a few sites in Long, Adobe, 
and Deep Springs Valleys (Fig. 47). Habitat 
includes seeps and spring-fed streams of 
small-moderate size (9.5-22°C, 85-450 
micromhos/cm) (Fig. 9b, c). Snails typically 
common in watercress and/or on small bits 
of travertine and stone. 



202 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






g 




Fig. 26. Camera lucida drawings of shells of P. owensensis and P. perturbata: a-d, P. owensensis (a, USNM 
853540, Mule Spring [2.1 mm]; b, USNM 857950, spring at Toll House [2.3 mm]; c, USNM 857955, stream 
in canyon S of Piute Creek [2.08 mm]; d, USNM 853543. spring at Graham Ranch [2. 1 6 mm]); e-g, P. perturbata 
(e, USNM 853546, Fish Slough, "Northwest Springs," [3.93 mm]; f, USNM 853549, Fish Slough, "BLM Spring," 
[3.22 mm]; g, USNM 853548, Fish Slough, "NE Spring" [3.97 mm]). 



Etymology. —Named after Darrell Wong, 
California Fish and Game, for his assistance 
during the project and interest in conserving 
Owens Valley springsnails. 

Remarks. —Similar in penial morphology 
to P. californiensis (Gregg & Taylor) from 
western California (Taylor 1981:154). Dis- 
tinguished from above by globose shell, and 
more numerous stalked glandular ridges on 
ventral penial surface. 

Genus Tryonia Stimpson, 1865 

Tryonia margae, new species 

Grapevine Springs elongate tryonia 

Figs. 48-50, 51a, 52 

Material examined. —California: Inyo 
Co.; Grapevine Springs (241); (upper) Warm 



spring on limestone bench, USNM 860408 
(holotype), *857952 (paratypes), 14 Mar 
1987; Spring on limestone bench above 
ranch 853555, 25 Feb 1987; 857996, 14 
Mar 1987. 

Diagnosis.— A small-sized species with 
narrow, turriform shell. Penis small, with 3 
papillae on inner curvature, and a single 
papilla (sometimes absent) on outer cur- 
vature. Distal tip of penis with elongate 
swelling on inner side. 

Description.— Shell (Fig. 48) 1.5-2.9 mm 
high, height/width, 250-300%. Whorls, 4.5- 
6.25, rounded, with angulated shoulders. 
Sutures often deeply impressed. Apex blunt, 
protoconch slightly depressed (Fig. 48g). 



VOLUME 102, NUMBER 1 



203 




Fig. 27. SEM micrographs of P. owensensis from stream in canyon S of Piute Creek: a, Holotype, USNM 
860404 (2.0 mm): b-d, Paratypes. USNM 857955 (bars = 120 pm, 176 urn). 



Body whorl 45-55% of shell height. Marl 
frequently covering apical whorls, with near- 
entirety of shell sometimes blanketed. Ap- 
erture narrowly ovate. Inner lip well-reflect- 
ed, slightly thickened, adnate to or slightly 
separated from body whorl. Outer lip sin- 
uate, thin (Fig. 48d, f). Umbilicus chink- 
like. Protoconch near-smooth (Fig. 48g). 
Collabral growth lines well developed on 



teleoconch, often elevated at irregular in- 
tervals. Spiral sculpture consisting of weak 
striations. 

Dark brown-black epithelial pigment 
covering most of head foot, although some- 
times absent on tentacles, and lighter on 
snout and central portion of sides of "neck." 

Radular (Figs. 49. 50) formula: 4(5)- 1 - 
4(5)/l(2)-l(2), 2(3)-l-3(4). 20-21, 22-28. 



204 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



a 




b 





v 





Fig. 28. SEM micrographs of P. owensensis from Warm Springs: a (1.92 mm), b, d, USNM 853544, main 
spring; c, USNM 853545, spring N of above (bar = 136 jtrni). 



Central tooth broadly trapezoidal, basal 
cusps large, basal process well excavated. 
Central cusps of lateral tooth only slightly 
enlarged (Fig. 49c, d). Penis (Fig. 51a) ex- 
tending only slightly beyond edge of mantle 
collar. Enlarged papilla located on outer edge 
of penis at base. Two smaller papillae po- 
sitioned along distal half on inner edge; sim- 
ilar small papilla usually found near base 



on inner edge. Brown-black epithelial pig- 
ment usually prominent near distal tip, at 
bases of two distal papillae. Lighter pigment 
dusting much of dorsal surfaces of penis and 
2 proximal papillae. 

Type locality. —Warm spring on lime- 
stone bench in Grapevine Springs complex, 
northern Death Valley, Inyo Co., Califor- 
nia. 



VOLUME 102, NUMBER 1 



205 




Fig. 29. Radula of P. owensensis, USNM 857955, stream in canyon S of Piute Creek: a, Centrals (bar = 10 
nm); b, Laterals (bar = 6.7 /urn); c, Inner marginal (bar = 6.0 ^m); d, Outer marginals (bar = 6.0 juin). 



Distribution and habitat.— Found in two 
sites (separated by about 1 km) among 
Grapevine Springs in northern Death Val- 
ley (Fig. 52). Habitat consists of upper por- 
tions of shallow spring brooks flowing over 
travertine (Fig. 8b), on which snails were 
moderately common. Springs were highly 



mineralized (1050-1075 micromhos/cm), 
but differed considerably in temperature 
(west spring, 26-27°C; east, 36-37°C). Syn- 
topic with T. rowlandsi in the thermal spring. 
Etymology.— From Latin marga, mean- 
ing marl, and referring to occurrence of snail 
in travertine springs. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 30. Radula of P. owensensis, USNM 857988, Warm Springs (main spring): a, Centrals (bar = 6.7 urn); 
b, Laterals (bar = 6.0 fim); c, Inner marginal (bar = 6.0 urn); d, Outer marginals (bar = 3.8 /mi). 



VOLUME 102, NUMBER 1 



207 



Tryonia protea (Gould, 1855) 

Desert tryonia 

Figs. 52-54 

Amnicola protea Gould 1855:129. 
Melania exigua Conrad 1855:269. 
Pyrgulopsis blakeana Taylor 1950:30. 
Pyrgulopsis cahuillarum Taylor 1950:31. 

Material examined.— California: Mono 
Co.; Hot Creek, USNM *857954, 23 Apr 
1987. 

Diagnosis.— A moderate to large-sized 
species (shell height typically 3-7 mm) with 
elongate-conic to turriform shell. Shell 
smooth or with spiral and/or collabral 
sculpture varying from faint striations to 
elevated ridges. 

Description of Hot Creek population. — 
Shell (Fig. 53) 3.0-4.5 mm high, height/ 
width, 210-242%. Whorls, 4.75-6.25, 
rounded. Sutures moderately impressed. 
Body whorl 55-61% of shell height. Pro- 
toconch slightly depressed (Fig. 53b), apex 
of adults invariably highly eroded. Aperture 
ovate, apertural plane tilted relative to coil- 
ing axis. Inner lip only slightly thickened 
and reflected, broadly adnate to body whorl. 
Outer lip thin, slightly sinuate. Umbilicus 
chink-like, near-absent. Smooth surface of 
protoconch lined with well-spaced rows of 
numerous, shallow perforations (Fig. 53b, 
c). Teleoconch with strong growth lines; 10- 
20 spiral lines prominent on middle whorls, 
sometimes extending to aperture. 

Brown epithelial pigment dense on most 
of snout. Black patches of subepithelial pig- 
ment along edges of sides of head/foot. Pal- 
lial roof with paired, black pigment patches 
extending along length of ctenidium. 

Radular (Fig. 54) formula: 5(6)-l-5(6)/ 
2(3)-2(3), (3-5)-l-(6-8), 35-36, 28-31 
(from single population). Central tooth 
broadly trapezoidal; basal cusps large, basal 
process moderately excavated. No males 
found among the more than 1 00 specimens 
collected. 

Type locality. —Colorado Desert (proba- 




Fig. 31. Reproductive anatomy of P. owensensis: 
a, Left lateral view of pallial oviduct complex, USNM 
857955, stream in canyon S of Piute Creek; b-i, Penes 
(b [dorsal], i [ventral], USNM 857955, stream in can- 
yon S of Piute Creek; c [dorsal], f [ventral], USNM 
857983, Mule Spring; d [dorsal], g [ventral], h [ventral], 
USNM 857988, Warm Spring (main spring); e [ven- 
tral], USNM 857989. Warm Springs [spring N of 
above]), "b" through "i" drawn to same scale. 



bly in Riverside County, California [based 
on Blake 1857]). 

Distribution and habitat. — "Western Utah 
to southeastern California, adjacent Baja 
California, and southeastern Arizona" 
(Taylor 1981:155), with only two living 
populations known from California: that of 
Hot Creek (Fig. 52) and Dos Palmas Spring 
in Riverside County (Taylor 1981:154). At 
Hot Creek (Fig. 9d), snails were rare on var- 
ious submergent macrophytes lining sides 



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Fig. 32. Map showing distribution of P. owensensis. 



of the swiftly flowing thermal stream (32°C, 
480 micromhos/cm). 

Remarks. —The taxonomic concept of this 
form is vague (Taylor 1966:53-54) and sta- 
tus of the few living (and mostly widely sep- 
arated) populations (Taylor 1985:fig. 35) re- 
mains unclear. The Hot Creek population 
is referable to T. protea on basis of shell 
form, size, and sculpture, but confirmation 



by anatomical comparison with snails from 
Dos Palmas Spring (that represent near to- 
potypes) would be desirable. 

Tryonia robusta, new species 

Robust tryonia 

Figs. 5 Id, 52, 55-58 

Material examined. —California: Inyo 
Co.; Nevares Springs (230); Spring on trav- 



VOLUME 102, NUMBER 1 



209 



a 




Fig. 33. SEM micrographs of P. perturbata from Fish Slough. "Northwest Springs. 
Holotype. USNM 860407 (3.57 mm): c. Paratype. USNM 853456 (bar = 200 Mm). 



south spring: a. b. 



ertine mound. USNM 860411 (holotype). 
*853557 (paratypes). 2 Feb 1985: 857999. 
16 Nov 1985: 857958. 12 Mar 1987: Spring 
issuing from base of mound. 853558. 2 Feb 



1985 — Travertine Springs: South spring. 
*853559, 2 Feb 1985: 857959. 12 Mar 1987: 
North spring. 857960. 12 Mar 1987. 
Diagnosis.— A small-sized species with 



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Fig. 34. Radula of P. perturbata, USNM 857990, Fish Slough, "Northwest Springs," south spring: a, Centrals 
(bar = 13.6 /Ltm); b, Laterals (bar =10 Aim); c, Laterals and inner marginals (bar =12 /urn); d, Outer marginals 
(bar = 10 Mm). 



unusually broadly conical shell for genus, high, height/width, 140-170%. Whorls, 

Penis moderate- sized, with single papilla on 3.75-4.75, rounded, with pronounced 

outer curvature. shoulders. Sutures impressed. Body whorl 

Description.— Shell (Fig. 55) 1.1-2.1 mm 68-80% of shell height. Protoconch (Figs. 



VOLUME 102, NUMBER 1 



211 



56a-c) simple, sometimes eroded. Aperture 
near-ovate, sometimes sharply angled 
above; apertural plane slightly tilted relative 
to coiling axis. Inner lip thickened, mod- 
erately reflected, usually slightly separated 
from body whorl. Outer lip thin, near- 
straight. Umbilicus chink-like. Protoconch 
surface wrinkled (Figs. 56a-c). Teleoconch 
often with weak spiral striations, particu- 
larly on upper whorls. 

Operculum (Fig. 56d) thin, paucispiral, 
with 2.5-3.0 rapidly expanding whorls. 

Brown-black melanic pigment dark on 
snout and posterior edge of "neck," variable 
on central portions of sides of head/foot; 
sub-epithelial pigment granules prominent 
in latter. Mantle roof epithelial pigment typ- 
ically uniformly dark brown-black. 

Radular (Fig. 57) formula: (5-7)- 1 -(5-7)/ 
1_1 , 3(4)_i_5(6), 24, 22-27 (from two pop- 
ulations). Central tooth relatively square- 
shaped; central cusps elongate, basal cusps 
small, basal process highly excavated. Ce- 
phalic tentacles covered with sparse, irreg- 
ular patches of hypertrophied cilia, rarely 
forming short tracts. Penis (Figs. 5 Id, 58) 
considerably longer than wide, moderately 
thickened. Attachment area broad, with rest 
of narrow length coiled in tight, clockwise 
fashion. Papilla small, conical, positioned 
along outer curvature just proximal to mid- 
length. Distal tip blunt, with inner side 
slightly swollen; surface of tip densely cov- 
ered with elongate cilia. Distal tip of papilla 
simple, lightly ciliated (Fig. 58b). 

Type locality. — Nevares Springs, Death 
Valley, Inyo County, California. 

Distribution and habitat. —Found at Ne- 
vares and Travertine Springs along base of 
Funeral Mountains in mid-eastern Death 
Valley (Fig. 52). Snails found commonly on 
plant debris and travertine in shallow out- 
flows of four thermal springs (30-3 7°C, 900- 
975 micromhos/cm). Extinct at Texas 
Springs (located between above complexes). 

Etymology. —Referring to unusually 
broad shell for genus. 

Remarks.— Shell form and penial shape 




Fig. 35. Reproductive anatomy of P. perturbata 
from Fish Slough, " Northwest Springs": a, Left lateral 
view of pallial oviduct complex, USNM 857990, south 
spring; b-e, Penes (b [dorsal], c [dorsal], d [ventral], 
USNM 857990, south spring; e [ventral], USNM 
857991, north spring). 



and lobation pattern are highly distinctive, 
and relationship between this local endemic 
and other congeners in region is unclear. 
Species closely conforms to typical Tryonia 
in other features (including details of female 
reproductive morphology). 

Tryonia rowlandsi, new species 

Grapevine Springs squat tryonia 

Figs. 51b, 52, 59, 60 

Material examined. —California: Inyo 
Co.; Grapevine Springs (241); (upper) Warm 
spring on limestone bench, USNM 860409 
(holotype), *857953 (paratypes), 14 Mar 
1987; (lower) Warm spring on limestone 
bench, 857997, 14 Mar 1987. 

Diagnosis.— A small-sized species with 
elongate-conic shell. Penis large, with single 
papilla on outside curvature and 2 papillae 
on inside curvature. 

Description.— Shell (Fig. 59) 1.7-2.3 mm 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



"V 



\ 



(-; 



\ 



M* 






,F 



r^ 



\J 




36 c 



N 



& 



is 



f \ 

( v 



10 10 30 50 
1 1 1 1 1 1 i i i i i Kilometers 



118 c 



117' 



Fig. 36. Map showing distribution of .P. perturbata. 



high, height/width, 1 40-1 90%. Whorls, 4.0- 
4.75, rounded, somewhat shouldered. Su- 
tures impressed. Body whorl 62-74% of shell 
height. Protoconch tilted relative to teleo- 
conch. Aperture ovate, apertural plane 
slightly tilted relative to coiling axis. Inner 
lip thin, only very slightly reflected, adnate 
to or slightly separate from body whorl. 



Outer lip thin, near-straight. Umbilicus nar- 
row, chink-like. Protoconch mottled with 
series of short, predominantly spiral stria- 
tions (Fig. 59e). Teleoconch with weakly de- 
veloped spiral lines, particularly on upper 
whorls. 

Head/foot and visceral coil usually uni- 
formly coated with dark brown epithelial 



VOLUME 102, NUMBER 1 



213 





J 



Fig. 37. SEM micrographs of P. cf. stearnsiana, USNM 853519, Sand Canyon (a, 3.04 mm; bar = 136 Mm). 



pigment. Central portions of sides of head/ Central tooth broadly trapezoidal; basal 

foot consisting of either a thin band or broad cusps small, basal process only moderately 

patch, often lighter or unpigmented. excavated. Lateral tooth having unusual 

Radular formula (Fig. 60): 4(5)-l-4/ 1 (2)- "claw-like" appearance, with cusps few and 

1, 2(3)-l-2(3), 16, 20 (from paratypes). widely separated. Penis (Fig. 51b) elongate, 



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Fig. 38. Radula of P. cf. stearnsiana, USNM &S7974, Sand Canyon: a, Centrals (bar = 10 ^m); b, Laterals 
(bar = 6.0 |im); c, Inner marginal (bar = 8.6 nm); d, Outer marginals (bar = 6.0 nm). 



distal tip with slightly swollen inner edge. 
Attachment area broad, with remaining pe- 
nis near-totally uncoiled. Single, small pa- 
pillae located on outer curvature near mid- 
point, and on inner curvature near distal 



tip. Inner curvature also bearing enlarged, 
basal papilla. Penial pigmentation consist- 
ing of small, dark patch near distal tip and/ 
or light dusting on broad area near base. 
Type locality. —Grapevine Springs, 



VOLUME 102, NUMBER 1 



215 



Grapevine Mountains, Death Valley, Inyo 
Co., California. 

Distribution and habitat.— Found in two 
thermal, highly mineralized springs on 
limestone bench in northeast portion of 
Grapevine Spring complex (32-36°C, 1 100- 
1050 micromhos/cm) (Fig. 52). 

Etymology.— Named after Peter Row- 
lands, Resources Management Division, 
National Park Service, Death Valley, for his 
efforts in both assisting this study and con- 
serving local aquatic gastropods in the Mon- 
ument. 



Tryonia salina, new species 

Cottonball Marsh tryonia 

Figs. 51e, 52, 61, 62 

Tryonia sp.— Taylor in LaBounty and Dea- 
con 1972:775. 

Material examined. — California: Inyo 
Co.; Spring in Cottonball Marsh (231), 
USNM 860410 (holotype), *853556 (para- 
types), 22 Feb 1985; *857998, 1 1 Mar 1987. 

Diagnosis. —A moderate-sized species 
with elongate-conic shell bearing well-de- 
veloped spiral striae. Penis with three pa- 
pillae on inner curvature and single papilla 
on outer curvature. 

Description.— Shell (Fig. 61) 2.4-3.1 mm 
high, height/width, 150-190%. Whorls, 4.5- 
5.25, well-rounded, shouldered above. Su- 
tures impressed. Shell loosely coiled, with 
slight separation of upper whorls common. 
Body whorl 61-68% of shell height. Pro- 
toconch tilted relative to teleoconch. Ap- 
erture ovate, apertural plane slightly tilted 
relative to coiling axis. Inner lip thin, only 
slightly reflected, narrowly adnate to or 
slightly separated from body whorl. Outer 
lip thin, near-straight. Umbilicus chink-like 
to moderately open. Protoconch smooth 
(Fig. 61c). Teleoconch with strong, regular- 
ly-spaced spiral striations, usually continu- 
ing onto body whorl, interacting with oc- 
casionally highly pronounced growth lines 
to produce a weakly cancellate appearance. 

Brown epithelial pigment dark on snout, 




Fig. 39. Reproductive anatomy of P. cf. stearn- 
siana, USNM 857974, Sand Canyon: a, Left lateral 
view of pallial oviduct complex; b-e, Penes (b, c, dorsal 
aspects; d, e, ventral aspects). 



dorsal tentacles, and all but narrow central 
strip of head/foot. 

Radular (Fig. 62) formula: 5(6)-l-5(6)/ 
2(3)-2(3), 3(4)-l-4(5), 19-26, 26-28 (from 
paratypes). Central tooth broadly trapezoi- 
dal, basal cusps small, basal process weakly 
excavated. Penis (Fig. 5 le) longer than wide, 
moderately thickened. Distal tip with 
prominent swelling of inner side. Inner cur- 
vature with two small papillae positioned 
on inner curvature near tip, and larger basal 
papilla. Outer curvature with enlarged basal 
papilla. Dorsal penis variably pigmented by 
brown epithelial melanin, with especially 
dark cover on basal penis and papillae. 

Type locality.— Spring brook in Cotton- 
ball Marsh, near base of Panamint Moun- 
tains at west side of salt pan in Death Valley, 
Inyo Co., California. 

Distribution and habitat. — Found in a few 
small, cool, brackish (17°C, >5000 mi- 
cromhos/cm) spring brooks emerging in 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 40. Map showing distribution of P. cf. stearnsiana in the study area. 



Cottonball Marsh, ca. 1.6 km out onto salt 
pan east of Salt Springs. Snails common on 
large, actively growing tufa blocks. Highly 
remote area in near-pristine condition. 

Etymology.— Referring to highly miner- 
alized spring habitat. 

Remarks.— -Very similar to T. rowlandsi 
from northern Death Valley, but distin- 
guished by larger size, more highly devel- 



oped shell sculpture, penial lobation pat- 
tern, and morphology of lateral radular teeth. 

Tryonia variegata 

Hershler and Sada, 1987 

Amargosa tryonia 

Figs. 51c, 52, 63, 64 

Tryonia variegata Hershler & Sada 1987: 
817. 



VOLUME 102, NUMBER 1 



217 






Fig. 41. Camera lucida drawings of shells of P. wongi: a, USNM 853531, spring alongside upper Pine Creek 
(2.18 mm); b, USNM 853524, spring N of Summit Creek (2.24 mm); c, USNM, 853527, Hogback Creek (Lone 
Pine) (1.74 mm); d, USNM 853530, spring S of Warren Lake (2.08 mm); e, USNM 853521, spring at Little 
Lake (1.44 mm); f, USNM 857947, French Spring (2.32 mm); g, USNM 857949, spring at Toll House (1.66 
mm); h, USNM 857946, spring (north) at N end of Blind Spring Valley (2.34 mm); i, USNM 853536, Antelope 
Spring (1.90 mm); j, USNM 853535, Corral Springs (1.80 mm); k, USNM 853538, River Springs (2.66 mm); 
1, USNM 853537, Layton Spring (1.78 mm). 



Material examined. —California: San 
Bernardino Co.; Saratoga Spring (204), 
USNM *853554, 27 Feb 1985; 857995, 10 
Jul 1986. Inyo Co.; Spring crossing path on 
south side of Amargosa Gorge (208), 853553 
(empty shell). — Spring in marsh east of 
Grimshaw Lake (209), *853551, 13 Mar 



1985; 857994, 22 Mar 1987. -Shoshone 
Spring (211), *853550, 12 Mar 1985; 
857993, 16 Mar 1987. -Resting Spring 
(212), 853552, 13 Mar 1985. 

Diagnosis.— A variably- sized species with 
turriform-aciculate shell. Penis large, with 
3 or 4 papillae on inner curvature and single 



218 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



a 





Fig. 42. SEM micrographs of P. wongi from west spring in Birchim Canyon: a, Holotype, USNM 860403 
(2.14 mm); b, c, Paratypes, USNM 857941 (bar = 136 fim). 



papilla (sometimes absent) on outer cur- 4.75-9.75, slightly to moderately rounded, 

vature. sometimes slightly shouldered. Sutures im- 

Description. —Shell (Fig. 63) 1 .2-7.5 mm pressed. Body whorl 37-63% of shell height. 

high, height/width, 170-350%. Whorls, Aperture narrowly ovate, apertural plane 



VOLUME 102, NUMBER 1 



219 




Fig. 43. Central radular teeth of P. wongi: a, USNM 857941, west spring in Birchim Canyon (bar = 10 yum); 
b, USNM 853521, spring at Little Lake (bar = 7.5 ^m); c, USNM 853536, Antelope Spring (bar = 7.5 ^m); d, 
USNM 857981, River Springs (bar = 12 /mi). 



near-parallel with coiling axis. Inner lip Umbilicus chink-like to moderately open, 

slightly thickened and reflected, adnate to Protoconch smooth to irregularly wrinkled 

or slightly separated from body whorl. Out- (Fig. 63d). Growth lines often highly pro- 

er lip thin, slightly to moderately sinuate, nounced. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 44. Lateral and marginal teeth of P. wongi: a (laterals), c (laterals and inner marginals), d (outer marginals), 
USNM 857941, west spring in Birchim Canyon (bars = 6.0 Aim); b, Laterals, USNM 853536, Antelope Spring 
(bar = 6.0 tim). 



VOLUME 102, NUMBER 1 



221 




Fig. 45. Reproductive anatomy of P. wongi: a, Left lateral view of pallial oviduct complex, USNM 857941, 
west spring in Birchim Canyon; b-1, Dorsal penes (b, USNM 857941, west spring in Birchim Canyon; c, USNM 
853530, spring S of Warren Lake; d, USNM 857981, River Springs; e, USNM 857979, Corral Springs; f, USNM 
857980, Layton Spring; g, USNM 857947, French Spring; h, USNM 857949, spring at Toll House; i, USNM 
853534, spring (north) at northern end of Blind Spring Valley; j, USNM 857978, spring in Owens Gorge; k, 
USNM 857942, spring alongside Lubkin Creek; 1, USNM 853523, Summit Creek). " c "-"l" drawn to same 
scale. 



Radular (Fig. 64) formula: (4-7)- 1 -(4-7)/ 
2(3)-2(3), 3(4)-l-4(5), 17-30, 22-33 (from 
numerous populations). Central tooth typ- 
ically broadly trapezoidal; basal cusps mod- 
erate to large-sized, basal process moder- 
ately excavated. 

Epithelial pigment usually dark brown- 



black on snout and sides of head/foot, with 
lighter cover on central portions Of latter. 

Type locality. —Five Springs, Ash Mead- 
ows, Nye County, Nevada. 

Distribution and habitat. — Amargosa 
River drainage, from Ash Meadows south 
to Saratoga Spring. Snails typically common 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 46. Penes (ventral aspects) of P. wongi: a, USNM 857941, west spring in Birchim Canyon; b, USNM 
857947, French Spring; c, USNM 853536, Antelope Spring; d, USNM 857942, spring alongside Lublin Creek; 
e, USNM 853534, spring (north) at northern end of Blind Spring Valley; f, USNM 857981, River Springs; g, 
USNM 857949, spring at Toll House; h, USNM 857979, Corral Spring; i, USNM 853523, Summit Creek, "a"- 
"e" and "g"-"i" drawn to same scale. 



on various substrates in small springpools 
and upper portions of outflows (7-3 2°C, 
430-10,150 micromhos/cm). 

Remarks. —Populations from lower 
Amargosa River drainage are clearly as- 
signable to T. variegata on basis of shell, 
radular, and penial features. 

Morphometries 

Discriminant analyses were performed on 
each of three groups of congeners from giv- 
en portions of the study area: a) P. amar- 
gosae and P. micrococcus, very similar forms 
with contiguous ranges; b) Pyrgulopsis fau- 
na of Owens Valley (P. aardahli, P. owen- 
sensis, P. perturbata, P. cf. stearnsiana, P. 



wongi); and c) small-sized Tryonia spp., in- 
cluding those of northern Death Valley (T. 
salina, T. rowlandsi, T. margae) and Ash 
Meadows (T. ericae, T. elata). 

Results (summarized in Tables 3-5) sup- 
port taxonomy presented herein, insofar as 
high correct classifications (81-87% [over- 
all]) indicate that these snails are well-dif- 
ferentiated in shell features. In the analysis 
involving P. amargosae and P. micrococcus, 
correct classification was 89% (40/45) and 
80% (291/362) for each species, respective- 
ly. Correct classification for two populations 
of P. amargosae (Amargosa Gorge, Grim- 
shaw Lake) from Tecopa area (where the 
ranges abut) was high (83%), as was that 
(92%) for two nearby populations of P. mi- 



VOLUME 102, NUMBER 1 



223 




Fig. 47. Map showing distribution of P. wongi. 



crococcus (Tecopa Hot Springs, Shoshone 
Spring). Misclassified specimens of P. mi- 
crococcus were concentrated in a few pop- 
ulations in northern Death, Panamint, and 
Saline Valleys (well separated from the range 
of P. amargosae), and indicate local differ- 
entiation of this relatively widespread form. 



In this analysis, discrimination was based 
largely on height of shell and aperture. 

For the Owens Valley Pyrgulopsis, correct 
classification was 84% overall (382/451), 
and ranged from 79-87% by species. The 
sole species having <80% correct classifi- 
cation, P. owensensis, was most frequently 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 48. SEM micrographs of T. margae: a-d, Grapevine Springs, upper warm spring (a, holotype, USNM 
860408, [3.52 mm]; b-d, Paratypes, USNM 857952); e-g, USNM 853555, Grapevine Springs, spring above 
shack (bar = 86 nm). 



VOLUME 102, NUMBER 1 



225 




..« 






■^ 


L.wA 




K %L ' 




1 ^v 


ij 


^^B ^B |A 




lA ^ 




■■ jp 


^K J I i ^H 


^^J 


iiVP 


^^^^^ 





,. U.Jifc^ 




Fig. 49. Central and lateral radular teeth of T. margae: a, c, USNM 857953, Grapevine Springs, upper warm 
spring (a, centrals, bar = 6.7 ^m; c, laterals and inner marginals, bar = 6.7 /um); b, d, USNM 857996, Grapevine 
Springs, spring above shack (b, centrals, bar = 6.7 ^m; d, laterals, bar = 6.7 yum). 



(10/14) misclassified as P. wongi, which nant function were related to size and width 

cannot be confused with the former when of shell and aperture, 

penial morphology is considered. Parame- Classification of each of the three small- 

ters weighing heavily on the first discrimi- sized Tryonia spp. from northern Death 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 50. Marginal radular teeth of T. margae: a, c, USNM 857953, Grapevine Springs, upper warm spring 
(a, laterals and inner marginals, bar = 10.0 /mi; c, outer marginal, bar = 3.8 /mi); b, d, USNM 857996, Grapevine 
Springs, spring above shack (b, laterals and inner marginals, bar =7.5 /mi; d, outer marginal, bar = 3.8 /mi). 



Valley was 100%, while that of T. elata and 
T. ericae from Ash Meadows was 93% and 
95%, respectively. Despite their general 
similarity in form, T. margae and T. elata 



were well-discriminated, with only a single 
specimen (of the latter) misclassified (as the 
former). Parameters weighing heavily on the 
first discriminant function were shell height, 



VOLUME 102, NUMBER 1 



227 




Fig. 51. Penes (dorsal aspects) of Tryonia spp.: a, 
T. margae, USNM 857952, Grapevine Springs, upper 
warm spring; b, T. rowlandsi, USNM 857953, Grape- 
vine Springs, upper warm spring; c, T. variegata, USNM 
857994, spring at Grimshaw Lake; d, T. robusta, USNM 
857999, Nevares Springs, upper spring; e, T. salina, 
USNM 857998, Cottonball Marsh, "a," "b," "d," "e" 
drawn to same scale. 



width of body whorl, and aperture size and 
shape. 

Discussion 

Modern drainage in the Death Valley area 
includes Owens, Amargosa, and Mojave 
Rivers, the latter two of which are dry along 
most of their courses. Although now iso- 
lated from one another, these river systems 
were integrated during wetter, pluvial times 
during the Pleistocene as a series of lakes 
overflowed along their courses to terminate 
in Death Valley. This Death Valley System 
(named by Miller 1943:69) was initially 
proposed by Gale (1914) and Blackwelder 
(1933), and has been discussed in detail by 
many others (Hubbs & Miller 1948; Miller 
1946, 1948; Smith 1978; Soltz & Naiman 



Table 3.— Results of discriminant function analysis 
on Pyrgulopsis amargosae and P. micrococcus. Also 
given are the canonical correlation, number of speci- 
mens used (by species, ordered as above), and percent 
correct classification (overall). 



Variables 



Standardized 
coefficients 



Correlations 





WH 


0.365 


0.554 




SH 


1.194 


0.354 




SW 


0.398 


-0.094 


E 


LBW 


-0.717 


0.176 


E 


WBW 


0.749 


0.146 


o 


AL 


-1.728 


0.055 




AW 


-0.067 


-0.108 




W 


0.176 


-0.292 




D 


0.798 


0.536 




T 


-0.390 


0.425 




SA 


0.871 


0.351 




C. correlation 


0.475 






N 


45,362 






% correct 








classification 


81 





1978; Smith & Street-Perrott 1983; Minck- 
ley et al. 1986; Taylor 1986). Configuration 
of the System is portrayed in Fig. 65 as pre- 
lude to discussion of springsnail zoogeog- 
raphy. 

Springsnail distributions reflect pro- 
nounced local endemism of the modern fau- 
na. Twenty of 22 species known from Death 
Valley System area are restricted to its con- 
fines. Of these, 10 are endemic to single 
springs or spring complexes, and another 
seven occur in relatively small portions of 
single modern drainage basins. Locally en- 
demic Pyrgulopsis are concentrated in Ow- 
ens Valley and Ash Meadows, while endem- 
ic Tryonia occur in the latter and northern 
Death Valley. Three forms are relatively 
widespread in the system: P. wongi, of plu- 
vial Owens River drainage (Long. Adobe, 
Owens Valleys); P. micrococcus, widely dis- 
tributed from Amargosa Valley west to Pan- 
amint Valley; and T. variegata, of pluvial 
Amargosa River drainage (Ash Meadows, 
Tecopa Basin, Saratoga Spring [southern 
Death Valley]). 

Endemic Pyrgulopsis of Owens Valley and 



228 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



38° 



36 c 



T. margae □ 

T. protea a 

T. robusta ■ 

T. rowlandsi o 

T. salina a 

T. variegata* 





Ik 


Ash Meadows 


\ 


x / 


■ 
■ 


\ • 




\ 




\ *„ 




V 




\ 




*fe\ 



\ 



N 



lk^ 



c j' 



f \ 
( \ 



10 10 30 50 
1 1 1 in 1 1 1 1 1 Kilometers 



1 18 c 



1 17 c 



Fig. 52. Map showing distribution of Tryonia spp. in study area. Tyronia variegata occurs at 18 localities 
in Ash Meadows. 



Amargosa Basin are sharply differentiated 
on basis of penial morphology and represent 
separate local radiations. Owens Valley 
forms, characterized in part by possession 
of a ventral penial swelling bearing glan- 
dular ridge (absent in forms endemic to 



Amargosa drainage), include a group com- 
prising P. cf. stearnsiana and similar, en- 
demic forms (P. aardahli, P. owensensis, P. 
perturbata); and P. wongi, which has no close 
relatives in the study area. Ash Meadows 
(Amargosa Basin) endemics represent at 



VOLUME 102, NUMBER 1 



229 




Fig. 53. SEM micrographs of T. protea, USNM 857954, Hot Creek. Shell "a" is 3.64 mm tall, "b," "c" 
show sculpture on protoconch of embryonic shell (bars = 60, 17.6 jrni). 



least two lineages (both including Flumin- 
icola-Mke representatives): snails with re- 
duced or absent penial lobes and a glandular 
ridge on mid-ventral penial surface (P. 
erythropoma, P. crystalis, P. pisteri); and 
those having a ridge on terminal portion of 
moderate-large sized penial lobes (P. fair- 



banksensis, P. nanus, P. isolatus). The latter 
may be related to the probable sister species 
pair of widespread P. micrococcus and P. 
amargosae from southern Amargosa River 
drainage. 

Relationships among regional Tiyonia are 
more speculative. Endemic Tryonia in 



Table 4.— Results of discriminant function analysis on Owens Valley Pyrgulopsis (P. aardahli, P. owensensis, 
P. perturbata, P. cf. stearnsiana, P. wongi). Also given are canonical correlations, number of specimens used 
(by species, ordered as above), and percent correct classification (overall). 





Standardized coefficients 




Correlations 


Variable 


Function 1 




Function 2 


Function 1 


Function 2 


WH 


0.580 




-0.841 


-0.277 


-0.711 


SH 


-2.972 




0.489 


-0.708 


-0.534 


SW 


1.037 




2.483 


-0.549 


-0.466 


LBW 


0.513 




-2.301 


-0.641 


-0.527 


WBW 


1.367 




-2.253 


-0.558 


-0.555 


AL 


2.937 




-0.550 


-0.553 


-0.424 


AW 


-3.827 




1.822 


-0.617 


-0.387 


W 


0.075 




-0.317 


0.255 


-0.068 


D 


0.285 




0.453 


-0.105 


-0.120 


T 


-0.143 




0.904 


-0.397 


-0.042 


SA 


-0.810 




0.600 


0.085 


-0.164 


C. correlation 


0.771 




0.687 






N 


15.86.39. 


25, 


286 






% correct 












classification 


84 











230 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 







Fig. 54. Radula of T. protea, USNM 857954, Hot Creek: a, Centrals (bar = 12 /mi); b, c, Laterals and inner 
marginals (bars = 8.6 /mi); d, Outer marginals (bar = 7.5 /im). 



VOLUME 102, NUMBER 1 



231 




Fig. 55. SEM micrographs of T. robusta from Nevares Springs, upper spring: a, Holotype, USNM 86041 1 
(1.92 mm); b-d, Paratypes, USNM 853557. 



northern Death Valley and Ash Meadows, 
apart from enigmatic T. robusta, apparently 
comprise a single local radiation. T. protea 
from west of Death Valley (Long and [fossil; 
Taylor 1985:317] Panamint Valleys) is quite 
similar (in shell) to at least one represen- 



tative from the east, T. variegata (contrary 
to Taylor 1985:317) and may not represent 
a separate lineage (as implied by Taylor 
1985:317). 

Pluvial waters of the Death Valley System 
provided aquatic continuity (and thus the 



Table 5.— Results of discriminant function analysis on small-sized Tryonia (T. elata, T. ericae, T. margae, 
T. rowlandsi, T. salina). Also given are canonical correlations, number of specimens used (by species, ordered 
as above), and percent correct classification (overall). 





Standardized coefficients 




Correlations 


Variable 


Function 1 


Function 2 


Function 1 


Function 2 


WH 


-0.044 


0.672 


0.109 


0.487 


SH 


1.592 


0.600 


-0.178 


0.699 


SW 


0.393 


0.735 


-0.580 


0.432 


LBW 


0.423 


-0.227 


-0.402 


0.535 


WBW 


-1.357 


-0.908 


-0.521 


0.493 


AL 


-1.813 


-0.341 


-0.510 


0.580 


AW 


0.003 


0.634 


-0.622 


0.405 


w 


-0.031 


0.081 


-0.077 


-0.312 


D 


0.286 


0.191 


0.032 


-0.030 


T 


-0.071 


0.429 


0.131 


0.427 


SA 


0.736 


0.828 


0.073 


0.353 


C. correlation 


0.994 


0.975 






N 


30, 19,6, 16, 15 








% correct 










classification 


97 









232 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 56. SEM micrographs of T. robusta, Nevares Springs, upper spring, USNM 853557: a-c, Views of apex 

(bars = 75, 43, 100 ^m); d. Dorsal operculum (bar = 200 ^ni). 



VOLUME 102, NUMBER 1 



233 




Fig. 57. Radula of T. robusta, Nevares Springs, upper spring, USNM 857999: a, Centrals (bar = 6.0 jrni); 
b, c, Laterals and inner marginals (bars = 3.8, 5.0 /im); d, Outer marginals (bar = 3.8 nm). 



234 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 58. SEM micrographs of penes of T. robusta, USNM 857999, Nevares Springs, upper spring: a, Dorso- 
right lateral aspect, with attachment to head on lower left (bar = 136 nm); b, Distal tip of papilla (bar = 23.1 
/mi); c, d, Distal tips of penis, showing ciliation (bars = 30, 20 /mi). 



VOLUME 102, NUMBER 1 



235 




Fig. 59. SEM micrographs of T. rowlandsi from Grapevine Springs, upper warm spring: a, Holotype, USNM 
860409 (2.18 mm); b-e, Paratypes, USNM 857953 (bar = 100 urn). 



236 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 60. Radula of T. rowlandsi, USNM 857953, Grapevine Springs, upper warm spring: a, Centrals (bar 
15 nm); b, Laterals (bar = 13.6 nm); c, Inner marginal (bar = 6.0 ^m); d, Outer marginals (bar = 5.0 yum). 



means for potential exchange of aquatic fau- ern springsnail zoogeography indicates that 

na) across current drainage divides imposed the above did not facilitate much mixing 

by the region's predominant north-south between pluvial Owens and Amargosa Riv- 

trending basin and range topography. Mod- er faunas. These results support earlier con- 



VOLUME 102, NUMBER 1 



237 









Fig. 61. SEM micrographs of T. salina from Cottonball Marsh: a, Holotype, USNM 860410 (3.27 mm); b, 
c, Paratypes, USNM 853556 (bar = 120 fim). 



elusions ofTaylor(m Miller 1981:58, 1985: 
317-318), although his observation that 
"Spillover from Lake Panamint into Death 
Valley . . . had no recognizable effect on 
mollusc distribution" (Taylor 1985:318) is 
contradicted by distribution of P. micro- 
coccus (and possible close affinities between 
T. protea and congeners to the east). 

Taylor (1985:317) suggested that faunal 
exchange between Amargosa and Owens 
River drainages might have been precluded 
by a saline dispersal barrier consisting of an 
arm of Bouse embayment (from former Gulf 
of California) extending into Death Valley. 
Although this hypothesis appears plausible, 
it may not need to be invoked to explain 
patterns described above. Apart from ad- 
ditional influences on snail distribution im- 
posed by far older lacustrine episodes than 
considered above (Smith 1984) and other 
aspects of complex regional geologic/hydro- 
logic history (Minckley et al. 1986), fun- 
damental questions relating to types of hab- 
itat occupied by Pleistocene springsnails and 
plausibility of dispersal of such (frequently 
stenotopic) organisms along chains of plu- 



vial lakes have not been thoroughly inves- 
tigated (see Hershler and Minckley 1986). 
Details such as size, number, and salinity 
of inter-connected lakes; and length and gra- 
dient of lake outlets obviously would affect 
probability of snail dispersal. Further, other 
modes of dispersal undoubtedly occurred 
and could have produced distributions 
comparable to or conflicting with those pre- 
dicted by pluvial drainage models. The 
presence of fauna in isolated basins recently 
created or devastated by volcanic activity 
(Long, Adobe Valleys, respectively), or those 
that either lacked pluvial connections (Sa- 
line Valley, Frenchman Flat [Cane Spring]; 
Hubbs and Miller 1948) or connected with 
snailless areas (Deep Springs Valley, whose 
pluvial lake spilled into Eureka Valley; Mil- 
ler 1928) attests to importance of factors 
other than late Pleistocene pluvial drainage 
systems in effecting modern snail distribu- 
tions in the region. A detailed zoogeograph- 
ic analysis encompassing the above consid- 
erations will be provided following 
completion of springsnail survey of remain- 
ing portions of the Death Valley System. 



238 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 62. Radula of T. salina, USNM 857998, Cottonball Marsh: a, Centrals (bar = 10 urn); b, Laterals and 
arginal (bar = 7.5 urn); c, Inner and outer marginals (bar = 8.6 /*m); d, Outer marginal (bar = 5.0 /mi). 



inner m 



VOLUME 102, NUMBER 1 



239 




Fig. 63. SEM micrographs of T. variegata: a (2.8 mm), b, USNM 853554, Saratoga Spring; c (2.86 mm), d, 
USNM 853550, Shoshone Spring (bar = 86 urn). 



""^ItfJI 


kjf 


^B I ^k. 






i 


^^r fry 

w5 


L^| 


B^ 


^V^fli 









Fig. 64. Radula of T. variegata: a, b, USNM 857995, Saratoga Spring (a, centrals, bar = 10.0 /tm; b, laterals, 
bar = 7.5 jum); c, d, USNM 857993, Shoshone Spring (c, laterals and inner marginals, bar = 6.0 nm; d, outer 
marginal, bar = 4.3 /nm). 



Fig. 65. Map of Pleistocene Death Valley System showing drainage relations (not necessarily contempora- 
neous). Stippled areas encircled by dashed lines may not have contained lakes for significant portions of the 
pluvial period. In the northeast, Lake Russell (Mono Valley) spilled into Lake Adobe (Adobe Valley), which in 
turn overflowed (through a gap in Benton Range) to enter the (now mostly dry) north fork of ancestral Owens 
River (Putnam 1949; Hubbs and Miller 1948; Gilbert et al. 1968). Lake Long Valley filled shortly after Long 
Valley caldera was created by explosive eruption of Bishop tuff 700,000 BP (Bailey et al. 1976), and spilled into 
south fork of Owens River. Outflow from Owens Lake, possibly initiated as a result of increased flow due to 

240 




capture of upper San Joaquin River drainage in Long Valley (Huber 1981, Smith et al. 1983), passed between 
Sierra Nevada and Coso Range to enter and fill China and Searles Lakes. Searles Lake overflowed into Panamint 
Valley, which held a large lake in its southern portion that spilled into Death Valley during several periods 
between 120,000 and 20,000 BP (Gale 1914, Hooke 1972, Smith 1976, Hale 1984). To the west, the lush 
wetland of Ash Meadows (that did not have a large pluvial lake; Mifflin and Wheat 1979, Hay et al. 1986) 
drained at least intermittently to Amargosa River during Pliocene-Pleistocene. Further downflow the river was 
ponded in mid- to late-Pleistocene by alluvial fans above Amargosa Gorge to form Lake Teopa (Sheppard and 
Gude 1968, Starkey and Blackmon 1979), which spilled into southern Death Valley. Mojave River filled a series 
of three pluvial lakes, with Lake Mojave overflowing toward southern Death Valley during several intervals of 
late Pleistocene from > 1 4,500-9000 BP (Ore and Warren 1971). Overflow from Death Valley System to Colorado 
River during an extreme pluvial period is conjectural, and has often been hypothesized based on distribution 
of aquatic biota (Miller 1981, Taylor 1986). Hale (1984) described an old (pre-Pleistocene) fluvial channel at 
Ash Hill near Ludlow, CA, that could be a product of such outflow. The channel is at 594 m elevation, which 
would require a vast pluvial lake (>300 km long and >650 m maximum depth) extending to northern Death 
Valley: additional geological evidence is needed to confirm this remarkable possibility. 

241 



242 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Acknowledgments 

Fieldwork in the Death Valley area was 
supported by Smithsonian Institution Re- 
search Opportunity Fund (ROF), Wildlife 
Conservation International of the New York 
Zoological Society, and contracts awarded 
by U.S. Fish and Wildlife Service, Great 
Basin Complex (Order No. 14320-0182); 
Bureau of Land Management, California 
Desert District (CA-060-CT5); and Califor- 
nia Fish and Game (C-1922). Collecting 
permits were provided by National Park 
Service (Death Valley National Monument) 
and State of California, and access to sites 
on Nevada Test Site, China Lakes Naval 
Weapons Center, and Fort Irwin is also 
gratefully acknowledged. Bureau of Land 
Management (District offices in Ridgecrest 
and Bishop), California Fish and Game 
(Bishop office), and National Park Service 
(Death Valley National Monument) loaned 
field vehicles and provided other forms of 
logistic support. Field assistance was pro- 
vided by numerous individuals, especially 
J. Aardahl, R. Brown, W. Cassidy, T. Ford, 
D. Giuliani, D. Herbst, B. Kohfield, P. 
Rowlands, and D. Wong. Heidi Wolf sorted 
material and produced excellent SEM mi- 
crographs. Paul Greenhall digitized shell 
material. Molly Ryan drew shells and draft- 
ed maps (with assistance from C. Flamer). 

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Appendix 

Collection localities, numbered as in Figs. 2-7. Data 
include name of site, state, county, topographic sheet, 
township and range coordinates, site elevation, and 
date of visitation (for negative sites only). 

1 . Stream in Sage Canyon. CA: Kern; Horse Canyon, 
CA (7.5), 7.0 km SW of NE comer of quadrangle, 1342 
m. 2. Boulder Spring. CA: Kern; Horse Canyon, CA, 
5.6 km S-SW of NE corner of quadrangle, 1251 m, 
3-26-87. 3. Stream in Indian Wells Canyon. CA: Kern; 
Inyokern, CA, NW Va sec. 17, T 26S, R 38E, 1068 m, 
3-26-87. 4. Spring in SW corner of Short Canyon. CA: 
Kern; Inyokern, CA, NW Va sec. 5, T 26S, R 38E, 1 129 
m, 3-26-87. 5. Stream in Grapevine Canyon. CA: Kem; 
Inyokern, CA, center of sec. 29, T 25S, R 38E, 946 m, 
3-26-87. 6. Stream in Sand Canyon. CA: Kern; Little 
Lake, CA, center of sec. 7, T 25S, R 38E, 1068 m. 7. 
Stream in Noname Canyon. CA: Kern; Little Lake, 
CA, 10.0 km NE of SW corner of quadrangle, 976 m, 
3-26-87. 8. Stream in Ninemile Canyon. CA: Inyo; 
Little Lake, CA, 12.2 km NE of SW corner of quad- 
rangle, 976 m, 3-26-87. 9. Spring 0.8 km S of Little 
Lake, W of HW 395. CA: Inyo; Little Lake, CA, SE Va 
sec. 18, T 23S, R 38E, 946 m, 4-1-87. 10. Spring at 
Little Lake, E of HW 395. CA: Inyo; Little Lake, CA, 
NW 1/4 sec. 17, T 23S, R 38E, 946 m. 11. Stream in 
Little Lake Canyon. CA: Inyo; Little Lake, CA, NE Va 
sec. 12, T 23S, R 37E, 1 129 m, 4-1-87. 12. Springs ca. 
1 .0 km N of Little Lake Canyon. CA: Inyo; Little Lake, 
CA, SW Va sec. 1, T 23S, R 37E, 1159 m, 4-30-87. 13. 
Stream in canyon ca. 3.0 km N of Little Lake Canyon. 
CA: Inyo; Little Lake, CA, 4.4 km SE of NW corner 
of quadrangle, 1220 m, 4-30-87. 14. Stream in Por- 
tugese Canyon. CA: Inyo; Haiwee Reservoir, CA, 2. 19 
km NE of SW corner of quadrangle, 1342 m, 4-1-87. 
15. Springs on Portugese Bench. CA: Inyo; Haiwee 
Reservoir, SW corners of sees. 3, 10, T 22S, R 37E, 
1 160-1220 m, 4-30-87. 16. Lower spring in Tunawee 
Canyon. CA: Inyo; Haiwee Reservoir, CA, SW Va sec. 
33, T 2 IS, R 37E, 1373 m, 4-25-87. 17. Upper spring 
in Tunawee Canyon. CA: Inyo; Monache Mtn., CA, 
7.0 km N-NW of SE corner of quadrangle, 1525 m, 
4-30-87. 18. Haiwee Creek. CA: Inyo; Monache Mtn., 



CA, 12.6 km S-SW of NE corner of quadrangle, 1586 
m, 4-30-87. 19. Springs in (2) unnamed canyons N of 
Haiwee Canyon. CA: Inyo; Monache Mtn., CA, NW 
corners sees. 30, 31, T 20S, R 37E, 1556 m, 4-30-87. 
20. Hogback Creek. CA: Inyo; Monache Mtn., CA, 7.3 
km SW of NE corner of quadrangle. 1586 m. 21. Sum- 
mit Creek. CA: Inyo; Monache Mtn., CA, NE Va sec. 
7, T 20S, R 37E, 1373 m. 22. Spring ca. 3.0 km N of 
Summit Creek. CA: Inyo; Monache Mtn, CA, center 
of sec. 6, T 20S, R 37E, 1281 m. 23. Walker Creek. 
CA: Inyo; Monache Mtn., CA, NE Va sec. 34, T 19S, 
R 37E, 1769 m, 4-16-87. 24. Cartago Creek. CA: Inyo; 
Olancha, CA, NE % sec. 11, T 19S, R 37E, 1159 m, 
4-16-87. 25. Spring at Cabin Bar Ranch, ca. 2.5 km N 
of Olancha. CA: Inyo; Olancha, CA, SW Va sec. 6, T 
19S, R 37E, 1098 m. 26. Braley Creek and springs just 
to S. CA: Inyo; Olancha, CA, 10.0 km NW of SE corner 
of quadrangle, 1190 m, 4-16-87. 27. Springs on edge 
of Owens Lake at Permanente. CA: Inyo; Olancha, CA, 
13.8 km N of SE corner of quadrangle, 1068 m, 2-9- 
85. 28. Ash Creek. CA: Inyo; Olancha, CA, 15.6 km 
NW of SE corner of quadrangle, 1068 m, 4-16-87. 29. 
Cottonwood Creek. CA: Inyo; Olancha, CA, 9.6 km 
SW of NE corner of quadrangle, 1037 m, 4-16-87. 30. 
Lower Centennial Spring. CA: Inyo; Keeler, CA, 2.3 
km W of SE corner of quadrangle, 1769 m, 3-31-87. 
31. Dirty Socks (Hot Spring). CA: Inyo; Keeler, CA, 
NE Va sec. 34, T 18S, R 37E, 1098 m, 2-9-85. 32. 
Springs at S end of Owens Lake, ca. 3.5 km NW of 
Dirty Socks. CA: Inyo; Keeler, CA, NW Va sec. 17, T 
18S, R 38E, 1098 m, 2-9-85. 33. Springs on edge of 
Owens Lake, ca. 3.0 km S of Keeler. CA: Inyo; Keeler, 
CA, NW y 4 sec. 22, T 17S, R 38E, 1098 m, 4-25-87. 
34. Lubkin Creek and spring feeding creek from south. 
CA: Inyo; Lone Pine, CA, SE 1/4 sec. 16, T 16S, R 36E, 
1220 m. 35. Spring along E side of Tuttle Creek. CA: 
Inyo; Lone Pine, CA, NE 1/4 sec. 6, T 16S, R 36E, 128 1 
m. 36. Hogback Creek. CA: Inyo; Lone Pine, CA, NW 
1/4 sec. 2, T 15S, R 35E, 1 159 m. 37. Spring at NE end 
of Alabama Hills, ca. 4.2 km N-NW of Lone Pine. CA: 
Inyo; Lone Pine, CA, NE % sec. 31, T 14S, R 36E, 
1159 m, 4-25-87. 38. George Creek. CA: Inyo; Lone 
Pine, CA, NE Va sec. 27, T 14S, R 35E, 1251 m, 4-25- 
87. 39. Independence Creek. CA: Inyo; Independence, 
CA, SE 1/4 sec. 23, T 13S, R 34E, 1342 m, 4-17-87. 40. 
Boron Springs. CA: Inyo; Mt. Pinchot, CA, NW Va sec. 
22, T 13S, R 34E, 1556 m. 41. Oak Creek, south fork. 
CA: Inyo; Mt. Pinchot, CA, SW 1/4 sec. 10, T 13S, R 
34E, 1525 m, 4-17-87. 42. Springs ca. 1.0 km W of 
Mt. Whitney Fish Hatchery. CA: Inyo; Mt. Pinchot, 
CA, SE 1/4 sec. 3, T 13S, R 34E, 1342 m, 4-18-87. 43. 
Stream in Charlie Canyon. CA: Inyo; Mt. Pinchot, CA, 
SW 1/4 sec. 3, T 1 3S, R 34E, 1 6 1 7 m. 44. Springs feeding 
N fork Oak Creek. CA: Inyo; Mt. Pinchot, CA, SW Va 
sec. 3, T 13S, R 34E, 1586 m, 4-25-87. 45. Oak Creek, 
north fork. CA: Inyo; Mt. Pinchot, CA, center of sec. 
3, T 13S, R 34E, 1525 m, 4-17-87. 46. Grover Anton 
Spring. CA: Inyo; Mt. Pinchot, CA, SW Va sec. 20, T 



VOLUME 102. NUMBER 1 



245 



12S. R 34E. 1 586 m. 4- 1 8-87. 47. Ham Birch Springs. 
CA: Inyo: Mt. Pinchot. CA. NW Ha sec. 17. T 12S. R 
34E. 1464 m. 4-18-87. 48. Sawmill Creek. CA: Inyo: 
Mt. Pinchot. CA. SW Ha sec. 9. T 12S. R 34E. 1251 
m, 4-17-87. 49. Division Creek and Scotty Springs just 
to N. CA: Inyo: Mt. Pinchot. CA. SW Ha sec. 1. T 12S. 
R 33E. 1 739 m. 4- 1 7-87. 50. Goodale Creek. CA: Inyo: 
Mt. Pinchot. CA. SE Va sec. 17. T US. R 34E. 1251 
m. 4-18-87. 51. Taboose Creek. CA: Inyo: Big Pine. 
CA. SE >4sec. 8. T US. R 34E. 1281 m. 4-18-87. 52. 
Tinnemaha Creek. CA: Inyo: Big Pine. CA. NW Va sec. 
28. T 10s. R 34E. 1281 m. 4-18-87. 53. Springs N of 
Tinnemaha Creek and W of Poverty Hills. CA: Inyo: 
Big Pine. CA, NE Va sec. 30. T 10S. R 34E. 1403 m. 
4-18-87. 54. Big Pine Creek. CA: Inyo: Big Pine. CA. 
SE U sec. 24. T 9S. R 33E. 1373 m. 4-18-87. 55. Baker 
Creek. CA: Inyo: Big Pine. CA. NE Va sec. 13. T 9S. R 
33E. 1312 m. 4-18-87. 56. Big Pine Spring. CA: Inyo: 
Big Pine. CA. NE Va sec. 16. T 9S. R 33E. 1891 m. 
4-18-87. 57. Spring on hill S of Warren Lake. CA: Inyo: 
Big Pine. CA NE Va sec. 16. T 9S. R 33E. 1220 m. 58. 
Springs in canyon W of Warren Lake. CA: Inyo: Big 
Pine. CA. SW Va sec. 2. T 9S. R 33E. 1251 m. 4-18- 
87. 59. Springs in canyon S of Shannon Canyon. CA: 
Inyo: Big Pine. CA. SW Va sec. 33. T 8S. R 33E. 1373 
m. 60. Springs N of Klondike Lake. CA: Inyo: Big Pine. 
CA, SW Va sec. 23. T 8S. R 34E. 1208 m. 5-1-87. 61. 
Freeman Creek. CA: Inyo: Big Pine. CA. NE Ha sec. 
20. T 8S. R 33E. 1434 m. 4-18-87. 62. Spring S of 
Rawson Creek. CA: Inyo: Bishop. CA. center of sec. 
6. T 8S. R 33E. 1434 m. 4-18-87. 63. Spring in Chip- 
munk Canyon. CA: Inyo: Bishop. CA. SW Va sec. 28. 
T 7S. R 32E, 1708 m. 4-20-87. 64. Stream in Butter- 
milk Country. CA: Inyo: Mt. Tom. CA. NE Va sec. 3 1 . 
T 7S. R 3 IE. 2349 m. 5-1-87. 65. Stream in McGee 
Meadow. CA: Inyo: Mt. Tom. CA, NE Va sec. 22. T 
7S. R 3 IE. 1800 m. 4-24-87. 66. Spring in Deep Can- 
yon. Mt. Tom, CA. center of sec. 12. T 7S. R 3 IE. 
1555 m. 4-29-87. 67. Horton Creek. CA: Inyo: Mt. 
Tom. CA. center of sec. 33. T 6S. R 31E. 1495 m. 
4-28-87. 68. Springs N of Horton Creek. CA: Inyo: Mt. 
Tom. CA. SW '4 sec. 32. T 6S. R 3 IE. 1586 m. 4-28- 
87. 69. Spring along N side of upper Pine Creek. CA: 
Inyo: Mt. Tom. CA. NE Ha sec. 26. T 6S. R 30E. 1830 
m. 70. Springs entering Pine Creek in Birchim Canyon. 
CA: Inyo: Mt. Tom. CA. SE Ha sec. 9. T 6S. R 3 IE. 
1373 m. 71. Spring in Wells Meadow. CA: Inyo: Mt. 
Tom. CA. NW Va sec. 2. T6S. R 3 IE. 1617 m. 4-24- 
87. 72. Spring W of Sierra Paradise. CA: Mono: Mt. 
Tom. CA, NW Va sec. 25. T 5S. R 30E. 1769 m. 4-24- 
87. 73. Stream in Swall Meadow. CA: Mono: Casa 
Diablo Mtn.. CA. NW Ha sec. 14. T 5S. R 30E. 2196 
m. 4-24-87. 74. Spring in Owens Gorge. CA: Inyo: Casa 
Diablo Mtn.. CA. SW Va sec. 31. T 4S. R 3 IE. 1830 
m. 75. Fish Slough. "BLM Spring." CA: Mono: Bishop. 
CA. SW Ha sec. 30. T 5S. R 33E. 1251 m. 76. Fish 
Slough. "Northwest Springs." CA: Mono: White Mtn. 
Peak. CA. SE Ha sec. 13. T 5S. R 32E. 1281 m. 77. Fish 



Slough. "NE Spring.'* CA: Mono: White Mtn. Peak. 
CA. NW 7 Ha sec. 18. T 5S. R 33E. 1281 m. 78. .Antelope 
Spring. CA: Mono: Casa Diablo Mtn.. CA. SW Va sec. 
9. T 4S. R 3 IE. 1830 m. 5-1-87. 79. Moran Spring. 
CA: Mono: Casa Diablo Mtn.. CA. SW Ha sec. 29. T 
3S. R 31E. 2104 m. 5-1-87. 80. Banner Springs. CA: 
Mono: Casa Diablo Mtn.. CA. SE Va sec. 18. T 3S. R 
3 IE. 2196 m. 5-1-87. 81. Spring by Tower Mine. CA: 
Mono: Casa Diablo Mtn.. CA. SE Ha sec. 3. T 3S. R 
3 IE. 1952 m. 4-21-87. 82. Springs entering Marble 
Creek. CA: Mono: White Mtn. Peak. CA. SE ] 4 sec 28. 
T 2S. R 32E. 1525 m. 83. Marble Creek at HW 6. CA: 
Mono: White Mtn. Peak. CA. NE Ha sec. 28. T 2S. R 
32E. 1525 m. 4-21-87. 84. Springs ca. 3.0 km S of 
Benton. W of HW 6. CA: Mono: Benton. NV-CA. SE 
Ha sec. 8. T 2S. R 32E. 1617 m. 4-21-87. 85. Benton 
Hot Springs. CA: Mono: Glass Mountain. CA-NV. SW 
Ha sec. 2. T 2S. R 3 IE. 1 7 08 m. 4-21-87. 86. Springs 
at N end of Blind Spring Valley. CA: Mono: Glass 
Mountain. CA-NV. SW Ha sec. 36. T IS. R 3 IE. 1708 
m. 87. Springs at Bramlette Ranch. CA: Mono: Benton. 
NV-CA. SW Va sec. 6 (springsnail positive). SE Ha sec. 
6 (negative). T IS. R 32E. 1678 m. 4-21-87. 88. Stream 
in Long John Canyon. CA: Inyo: New York Butte. CA, 
14.4 km N-NE of SW corner of quadrangle. 1 " 7 69 m. 
5-4-87. 89. French Spring. CA: Inyo. New York Butte. 
CA. 8.8 km S of NW corner of quadrangle. 1617 m. 
90. Spring in Willow Springs Canyon. CA: Inyo: In- 
dependence. CA. 6.5 km NW of SE corner of quad- 
rangle. 1861 m. 8-18-87. 91. Barrel Springs. CA: Inyo. 
Independence. CA. 13.6 km SW of NE corner of quad- 
rangle. 1952 m. 92. Spring ca. 1.6 km SE of Mule 
Spring. CA: Inyo: Waucoba Mtn.. CA. 11.7 km N-NE 
of SW corner of quadrangle. 1586 m. 93. Mule Spring. 
CA: Inyo. Waucoba Mtn.. CA. SE Ha sec. 1. T 10S, R 
34E. 1312 m. 94. Spring at Graham Ranch. CA: Inyo. 
Waucoba Mtn.. CA. NE Ha sec. 12. T9S. R 34E. 1373 
m. 95. Llymeyer Spring. CA: Inyo: Waucoba Mtn., 
CA. NE »4 sec. 10. T 9S. R 34E. 1235 m. 4-19-87. 96. 
Wilkerson Springs. CA: Inyo. Waucoba Mtn.. CA. NW 
'4 sec. 10. T 9S. R 34E. 1220 m. 97. Spring at Toll 
House. CA: Inyo. Waucoba Mtn.. CA. NE '4 sec. 24. 
T 8S. R 34E. 1861 m. 98. Warm Springs. CA: Inyo. 
Bishop. CA. SW Va sec. 8. T 8S. R 34E. 1220 m. 99. 
Spring S of Poleta Canyon. CA: Inyo: Bishop. CA. SE 
Va sec. 18. T 7S. R 34E. 1373 m. 5-4-87. 100. Spring 
S of Silver Canyon. CA: Inyo: Bishop. CA. SE '4 sec. 
26. T6S. R 33E. 1281m. 4-26-87. 101. Stream in Silver 
Canyon. CA: Inyo: Bishop. CA. SE Ha sec. 24. T 6S. R 
33E. 1434 m. 4-26-87. 102. Stream in Coldwater Can- 
yon. CA: Mono: Bishop. CA. SE Ha sec. 26. T 5S. R 
33E. 1494 m. 5-3-87. 103. Springs on bench S of Piute 
Creek. CA: Mono. Bishop. CA. SE Va sec. 22. T 5S. R 
33E. 1342 m. 104. Stream in canyon S of Piute Creek. 
CA: Mono. Bishop. CA. NE Ha sec. 23. T 5S. R 33E. 
1617 m. 105. Spring S of Millner Creek at Copper 
Queen Mine. CA: Mono: White Mtn. Peak. CA. NE 
Va sec. 21. T 4S. R 33E. 1586 m. 4-28-87. 106. Spring 



246 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



S of Lone Tree Creek at Hill Ranch. CA: Mono; White 
Mtn. Peak, CA. NE Va sec. 5, T 4S, R 33E. 1617 m. 
4-22-87. 107. Birch Creek. CA: Mono; White Mtn. 
Peak, CA, SE Va sec. 7, T 3S, R 33E, 1617 m, 4-22- 
87. 108. Orchard Spring. NV: Mineral; Benton, NV- 
CA, SW Va sec. 35, T IN, R 32E, 2135 m. 5-10-87. 
109. Spring in Queen Canyon. NV: Mineral; Benton, 
NV-CA, SE Va sec. 25, T IN, R 32E, 2166 m, 5-10- 
87. 1 10. Stream in Queen Canyon (above spring). NV: 
Mineral; Benton, NV-CA, NE Va sec. 36, T IN, R 32E, 
2288 m, 5-10-87. 111. Springs just downslope from 
Long Valley Dam. CA: Mono; Casa Diablo Mtn., CA, 
NW Va sec. 20, T 4S, R 30E, 1922 m, 5-3-87. 112. 
Stream on W side of Little Round Valley. CA: Mono; 
Casa Diablo Mtn., CA, SE Va sec. 36, T 4S, R 29E, 
2105 m, 4-23-87. 113. McGee Creek. CA: Mono; Mt. 
Morrison, CA, 7.9 km NW of SE corner of quadrangle, 
2501 m, 4-23-87. 114. Stream (W of Convict Creek) 
feeding Convict Lake. CA: Mono; Mt. Morrison, CA, 
SW Va sec. 22, T 4S, R 28E, 2562 m, 4-27-87. 115. 
Springs on S side of HW 395 W of Crowley Lake. CA: 
Mono; Mt. Morrison, CA, SW Va sees. 1 (4-23-87) and 
2 (4-29-87), T 4S, R 28E, 2172 m. 116. Spring E of 
Laurel Creek. CA: Mono; Mt. Morrison, CA, NE Va 
sec. 9, T 4S, R 28E, 2196 m, 4-29-87. 117. Whitmore 
Hot Springs. CA: Mono; Mt. Morrison, CA, NE Va sec. 
6, T4S, R29E, 2105 m, 4-29-87. 118. Springs at Hot 
Creek Hatchery. CA: Mono; Mt. Morrison, CA, SW Va 
sec. 35, T 3S, R 28E, 2166 m, 4-23-87. 119. Springs 
at Hot Creek Ranch. CA: Mono; Mt. Morrison, CA, 
SE Va sec. 35, T 3S, R 28E, 2166 m, 4-23-87. 120. Casa 
Diablo Hot Springs (cool seeps). CA: Mono; Mt. Mor- 
rison, CA, NW Va sec. 32, T 3S, R 28E, 2257 m, 4-23- 
87. 121. Hot Creek. CA: Mono, Mt. Morrison, CA, 
NE Va sec. 25, T 3S, R 28E, 2196 m. 122. Layton 
Springs (unnamed on topographic sheet). CA: Mono, 
Casa Diablo Mtn., CA, SE Va sec. 36, T 3S, R 29E, 
2074 m. 123. Springs feeding Little Alkali Lake. CA: 
Mono; Mt. Morrison, CA, SW Va sec. 20, T 2S, R 29E, 
2104 m, 4-29-87. 124. Little Hot Creek (source). CA: 
Mono; Mt. Morrison, CA, NW Va sec. 13, T 3S, R 28E, 
2074 m, 5-6-87. 125. Deadman Creek. CA: Mono; Mt. 
Morrison, CA, NE Va sec. 27, T2S, R 27E, 2257 m, 
4-29-87. 126. Big Springs. CA: Mono; Cowtrack Mtn., 
CA, NE Va sec. 25, T 2S, R 27E, 2166 m, 4-23-87. 127. 
Spring ca. 0.5 km E-NE of Arcularius Ranch. CA: Mono; 
Cowtrack Mtn., CA, SE Va sec. 20, T 2S, R 28E, 2135 
m, 5-6-87. 128. Springs along lower section of Mc- 
Laughlin Creek. CA: Mono; Cowtrack Mtn., CA, NE 
Va sec. 15, T 2S, R 28E, 2318 m, 5-6-87. 129. Mc- 
Laughlin Spring. CA: Mono; Cowtrack Mtn., CA, NE 
Va sec. 12, T 2S, R 28E, 2654 m, 5-6-87. 130. Springs 
at S end of Black Lake. CA: Mono; Glass Mountain, 
CA-NV, SW Va sec. 4, T 2S, R 3 IE, 1922 m, 4-21-87. 
131. Springs at N end of Black Lake. CA: Mono; Glass 
Mountain, CA-NV, SE Va sec. 29, T IS, R 3 IE, 1952 
m, 4-21-87. 132. River Spring. CA: Mono, Glass 
Mountain, CA-NV, NE Va sec. 24, T IN, R 30E, 1983 



m. 133. Stream in Crooked Meadows. CA: Mono; 
Cowtrack Mtn., CA, NW Va sec. 36, T IS, R 28E, 2654 
m. 5-6-87. 134. Pilot Spring. CA: Mono; Cowtrack 
Mtn., CA, SE Va sec. 27, T IS, R 28E, 2623 m, 5-6- 
87. 135. Baxter Spring. CA: Mono; Cowtrack Mtn., 
CA, SE Va sec. 1, T IS, R 28E, 2623 m, 5-6-87. 136. 
Gaspipe Spring. CA: Mono; Cowtrack Mtn., CA, NW 
Va sec. 1, T IS, R 28E, 2440 m, 5-6-87. 137. Indian 
Spring. CA: Mono; Cowtrack Mtn., CA, 13.4 km SE 
of NW corner of quadrangle. 2288 m, 5-6-87. 138. 
Simons Spring (and others in large marshy area). CA: 
Mono; Cowtrack Mtn., CA, 6.9 km SE of NW corner 
of quadrangle, 1968 m, 5-5-87. 139. Willow Spring 
(not shown on topographic sheet). CA: Mono; Cow- 
track Mtn., CA, NE Va sec. 15, T IN, R 27E, 1952 m, 
5-5-87. 140. Spring along Lee Vining Creek. CA: Mono; 
Mono Craters, CA, NW Va sec. 19, T IN, R 26E, 2227 
m, 5-5-87. 141. Springs feeding lower section of Lee 
Vining Creek. CA: Mono; Mono Craters, CA, NW Va 
sec. 9, T IN, R 26E, 2000 m, 5-5-87. 142. Springs 
along HW 395 ca. 5.0 km NW of Lee Vining. CA: 
Mono; Mono Craters, CA, NE Va sec. 3 1 , T 2N, R 26E, 
1950-2075 m, 5-6-87. 143. Lower Dechambeau Creek 
and associated springs. CA: Mono; Bodie, CA, SE Va 
sec. 19, T 2N, R 26E, 1922 m, 5-6-87. 144. Spring ca. 
2.0 km S of Dechambeau Creek along HW 395. CA: 
Mono; Bodie, CA, SE Va sec. 30, T 2N, R 26E, 1922 
m, 5-6-87. 145. Mill Creek. CA: Mono; Bodie, CA, 
NW Va sec. 15, T 2N, R 25E, 2379 m, 5-6-87. 146. 
Stream N of Mill Creek. CA: Mono; Bodie, CA, SE Va 
sec. 1, T 2N, R 25E, 2105 m, 5-6-87. 147. Stream in 
Rattlesnake Canyon. CA: Mono; Bodie, CA, NW Va 
sec. 30, T 3N, R 26E, 2288 m, 5-6-87. 148. Murphy 
Spring. CA: Mono; Bodie, CA, SE Va sec. 24, T 4N, R 
26E, 2501 m, 5-6-87. 149. Stream S of Murphy Spring. 
CA: Mono; Bodie, CA, NE Va sec. 25, T 4N, R 26E, 
2410 m, 5-6-87. 150. Stream in Cottonwood Canyon. 
CA: Mono; Bodie, CA, SE Va sec. 32, T 4N, R 27E, 
2227 m, 5-6-87. 151. Sulphur Pond and pool to E. CA: 
Mono; Trench Canyon, CA-NV, SE Va sec. 26, T 3N, 
R 27E, 1952 m, 5-6-87. 152. Saline Marsh. CA: Inyo; 
New York Butte, CA, center of sec. 27, T 14S, R 28E, 
335 m. 153. Stream in Hunter Canyon. CA: Inyo; New 
York Butte, CA, 14.5 km SE of NW corner of quad- 
rangle, 549 m, 3-28-87. 154. Stream in Beveridge Can- 
yon. CA: Inyo; New York Butte, CA, 12.3 km E-SE of 
NW corner of quadrangle, 580 m, 3-28-87. 155. Bad- 
water Spring. CA: Inyo; Waucoba Wash, CA, 10.0 km 
NE of SW corner of quadrangle, 488 m, 3-28-87. 156. 
Springs S of Lower Warm Springs. CA: Inyo; Waucoba 
Wash, CA, NE Va sec. 30, T 13S, R 39E, 381 m, 3-29- 
87. 157. Springs NW of Upper Warm Springs. CA: 
Inyo; Waucoba Wash, CA, SE Va sec. 5, T 13S, R 39E, 
549 m, 3-29-87. 158. Upper Warm Spring. CA: Inyo; 
Dry Mountain, CA, NE Va sec. 9, T 13S, R 39E, 488 
m, 2-11-85. 159. Willow Creek. CA: Inyo; Waucoba 
Wash, CA, 12.3 km N-NE of SW corner of quadrangle, 
915 m, 3-28-87. 160. Corral Springs. CA: Inyo; Blanco 



VOLUME 102, NUMBER 1 



247 



Mtn., CA, SW Va sec. 3, T 8S, R 36E, 1525 m. 161. 
Bog Mounds Springs. CA: Inyo; Blanco Mtn., CA, SW 
Va sec. 32, T 7S, R 36E, 1 508 m, 4-26-87. 1 62. Antelope 
Spring. CA: Inyo; Blanco Mountain, CA, NW Va sec. 
24, T 7S, R 35E, 1708 m. 163. Samms Spring. CA: 
Inyo; Blanco Mountain, CA, SW Va sec. 13, T 7S, R 
35E, 1739 m, 3-29-87. 164. Willow Spring. CA: Inyo; 
Magruder Mtn., NV-CA, 10.0 km N-NE of SW comer 
of quadrangle, 1739 m, 3-30-87. 165. Wheelbarrow 
Spring (not shown on topographic sheet). CA: Inyo; 
Soldier Pass, CA-NV, NE Va sec. 5, T 7S, R 37E, 1708 
m, 3-30-87. 166. Wyman Creek. CA: Inyo; Soldier 
Pass, CA-NV, SW Va sec. 23, T 6S, R 36E, 1769 m, 
3-29-87. 167. Stream in Indian Joe Canyon. CA: Inyo; 
Trona, CA, NW Va sec. 24, T 24S, R 42E, 732 m, 4- 1 3- 
87. 168. Allen Spring (not shown on topographic 
sheet). CA: Inyo; Trona, CA, NE Va sec. 24, T 24S, R 
42E, 854 m, 4-13-87. 169. Stream in Great Falls Basin. 
CA: Inyo; Trona, CA, SW Va sec. 12, T 24S, R 42E, 
854 m, 4-13-87. 170. Stream in Canyon N of Great 
Falls Basin. CA: Inyo; Trona, CA, NW Va sec 7, T 24S, 
R 43E, 915 m, 3-25-87. 171. Stream in S fork Home- 
wood Canyon. CA: Inyo; Trona, CA, NW Va sec 2, T 
24S, R 42E, 1 159 m, 3-25-87. 172. Benko Spring. CA: 
Inyo; Trona, CA, NE Va sec. 34, T 23S, R 42E, 1037 
m, 3-25-87. 173. Stream in N fork Homewood Can- 
yon. CA: Inyo; Trona, CA, SW Va sec. 25, T 23S, R 
42E, 1098 m, 3-25-87. 174. Bircham Springs. CA: Inyo; 
Trona, CA, NE Va sec. 20, T 23S, R 42E, 1708 m, 3- 
4-85. 175. Wild Rose Spring. CA: Inyo; Mountain 
Springs Canyon, CA, SW Va sec. 11, T 23S, R 4 IE, 
1525 m, 3-4-85. 176. LaMotte Spring. CA: Inyo; Trona, 
CA.NE^sec. 31, T 22S, R42E, 1525 m, 3-4-85. 177. 
Stream in N fork Water Canyon. CA: Inyo; Trona, CA, 
NE Va sec. 36, T 22S, R 42E, 1098 m, 4-11-87. 178. 
Tennessee Spring. CA: Inyo; Coso Peak, CA, SW Va 
sec. 13, T 2 IS, R 4 IE, 1830 m. 179. Haiwee Spring. 
CA: Inyo; Haiwee Reservoir, CA, SW Va sec. 10, T 2 IS, 
R39E, 1403 m, 3-5-85. 1 80. Stream in Knight Canyon. 
CA: Inyo; Maturango Peak, CA, SE Va sec. 14, T 2 IS, 
R 42E, 1037 m. 181. Stream in Revenue Canyon. CA: 
Inyo; Maturango Peak, CA, NW Va sec. 3, T 2 IS, R 
42E, 976 m, 4-14-87. 182. Stream in Snow Canyon. 
CA: Inyo; Maturango Peak, CA, 5.1 km S-SE of NW 
corner of quadrangle, 1342 m. 183. Black Spring. CA: 
Inyo; Coso Peak, CA, 1 .6 km E of NW corner of quad- 
rangle, 1769 m, 3-31-87. 184. Darwin Wash. CA: Inyo; 
Darwin, CA, China Garden Spring, NE Va sec. 4, T 
19S, R 4 IE, 1037 m; spring above Darwin Falls, NW 
Va sec. 3, T 19S, R 4 IE, 854 m; stream below falls, SW 
■A sec. 34, T 18S, R41E, 732 m. 185. Stream in Grape- 
vine Canyon. CA: Inyo; Ubehebe Peak, CA, 8.4 km 
NW of SE corner of quadrangle, 1525 m, 3-28-87. 186. 
Big Dodd Spring. CA: Inyo; Ubehebe Peak, CA, 1 1.5 
km NW of SE corner of quadrangle, 1 159 m, 3-12-87. 
187. Stream in Goler Canyon. CA: Inyo; Manley Peak, 
CA, 15.3 km NE of SW corner of quadrangle, 549 m, 
3-31-87. 188. Anvil Spring. CA: Inyo; Manley Peak, 



CA, 1 1.3 km SW of NE corner of quadrangle, 1373 m, 
2-18-85. 189. Seepage on SW side of Panamint Lake 
bed. CA: Inyo; Manley Peak, CA, E sides of sees. 28 
and 34, T 22S, R 44E, 317 m, 3-31-87. 190. Stream 
in Pleasant Canyon. CA: Inyo; Telescope Peak, CA, 
6.9 km NE of SW corner of quadrangle, 793 m. 191. 
Post Office Spring. CA: Inyo; Telescope Peak, CA, NE 
Va sec. 9, T 22S, R 44E, 317 m, 2-14-85. 192. Stream 
in Happy Canyon. CA: Inyo; Telescope Peak, CA, 11.3 
km NE of SW corner of quadrangle, 1098 m, 4-1 1-87. 
193. Spring mounds on Panamint Lake bed NW of 
Ballarat. CA: Inyo; Maturango Peak, CA, NW Va sec. 
20, T 21S, R 44E, 323 m, 4-14-87. 194. Stream in 
Surprise Canyon. CA: Inyo; Telescope Peak, CA, 15.3 
km NE of SW corner of quadrangle, 1098 m. 195. 
Warm Sulphur Springs. CA: Inyo; Telescope Peak, CA, 
SE Va sec. 10, T 21S, R 44E, 305 m, 2-13-85. 196. 
Stream in Jail Canyon. CA: Inyo; Telescope Peak, CA, 
10.0 km SEofNW corner of quadrangle, 1220 m. 197. 
Stream in Tuber Canyon. CA: Inyo; Telescope Peak, 
CA, 5.6 km SE of NW corner of quadrangle, 1098 m, 
4-11-87. 198. Wildrose Spring. CA: Inyo; Emigrant 
Canyon, CA, 3.8 km NE of SW corner of quadrangle, 
1080 m, 2-15-85. 199. Springs in upper Wildrose Can- 
yon. CA: Inyo; Emigrant Canyon, CA, 5.3 km NE of 
SW corner of quadrangle, 1220 m, 2-15-85. 200. Cave 
Spring. CA: San Bernardino; Avawatz Pass, CA, 7.7 
km NE of SW corner of quadrangle, 1 129 m, 4-9-87. 
201. Sheep Creek Spring. CA: San Bernardino; Ava- 
watz Pass, CA, 3.8 km NW of SE corner of quadrangle, 
427 m, 4-10-87. 202. Salt Creek E of HW 127. CA: 
San Bernardino; Avawatz Pass, CA, 14.1 km S-SW of 
NE corner of quadrangle, 92 m, 3-15-87. 203. Owl 
Hole Springs. CA: San Bernardino; Leach Lake, CA, 
NW Va sec. 22, T 19N, R 3E, 641 m, 3-16-85. 204. 
Saratoga Spring. CA: San Bernardino; Avawatz Pass, 
CA, NW Va sec. 2, T 18N, R 5E, 61 m. 205. Ibex Spring. 
CA: San Bernardino; Shoshone, CA, 8.5 km E-NE of 
SW corner of quadrangle, 366 m, 3-22-87. 206. Horse 
Thief Springs. CA: San Bernardino; Horse Thief Springs, 
CA-NV, 10.4 km E-NE of SW corner of quadrangle. 
1403 m, 4-10-87. 207. Willow Spring. CA: Inyo; Te- 
copa, CA, SW Va sec. 25, T 20N, R 7E, 427 m, 3-22- 
87. 208. Springs in Amargosa Gorge, S of Tecopa. CA: 
Inyo; Tecopa, CA, NW Va sec. 15, T 20N, R 7E, 397 
m. 209. Spring in marsh E of Grimshaw Lake. CA: 
Inyo; Tecopa, CA, NE Va sec. 9, T 20N, R 7E, 427 m. 
210. Northernmost of Tecopa Hot Springs. CA: Inyo; 
Tecopa, CA, NW Va sec. 33, T 2 IN, R 7E, 397 m. 21 1. 
Shoshone Spring. CA: Inyo; Shoshone, CA, NW Va sec. 
30, T 22 N, R 7E, 519 m. 212. Resting Spring. CA: 
Inyo; Tecopa, CA, SE Va sec. 31. T 2 IN, R 8E, 549 m. 
213. Lost Spring (unnamed on quadrangle). CA: Inyo; 
Wingate Wash, CA. 1 2.8 km SE of NW corner of quad- 
rangle, 702 m, 3-10-87. 214. Salsberry Spring. CA: 
Inyo; Shoshone, CA. 10.6 km SW of NE corner of 
quadrangle, 366 m. 3-14-85. 215. Warm Spring. CA: 
Inyo; Wingate Wash, CA. 7.2 km SE of NW corner of 



248 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



quadrangle. 732 m, 2-18-85. 216. Lower Talc Mine 
Spring. CA: Inyo; Bennetts Well, CA, 6.4 km E-NE of 
SW corner of quadrangle. 9 1 5 m, 3- 1 0-87. 2 1 7. Willow 
Spring. CA: Inyo; Funeral Peak, CA, 7.8 km NE of SW 
corner of quadrangle. 732 m, 3-20-87. 218. Hidden 
Spring. CA: Inyo; Funeral Peak, CA, 15.3 km NE of 
SW corner of quadrangle, 1 342 m, 3- 1 7-87. 219. Springs 
at Mormon Point. CA: Inyo; Bennetts Well, CA, 6.7 
km N-NW of SE corner of quadrangle, —64 m, 3-9- 
85. 220. Spring (lower) in Johnson Canyon. CA: Inyo; 
Telescope Peak, CA, SW Va sec. 22, T 2 1 S, R 46E, 1 1 90 
m. 221. Eagle Borax Spring. CA: Inyo; Bennetts Well, 
CA, 13.3 km SE of NW corner of quadrangle, — 79 m, 
2-16-85. 222. Hanaupah Spring. CA: Inyo; Telescope 
Peak, CA, 7.6 km S-SW of NE corner of quadrangle, 
1281 m. 223. Badwater Spring. CA: Inyo; Bennetts 
Well, CA, 2.7 km SW on NE corner of quadrangle, 
-79 m, 2-16-85. 224. Tule Spring. CA: Inyo; Bennetts 
Well, CA, 10.8 km E-SE of NW corner of quadrangle, 
— 79 m, 2-28-85. 225. Grapevine Springs. NV: Nye; 
Ash Meadows, NV-CA, NE Va sec. 2, T 19S, R 50E, 
732 m, 3-14-85. 226. Blackwater Spring. CA: Inyo; 
Emigrant Canyon, CA, 12.9 km S-SW of NE corner of 
quadrangle, 946 m, 3-10-87. 227. Navel Spring. CA: 
Inyo; Ryan, CA-NV, NW Va sec. 13, T 26N, R 2E, 640 
m, 3-17-87. 228. Travertine Springs. CA: Inyo; Fur- 
nace Creek, CA, NW Va sec. 25, T 27N, R IE, 122 m. 
229. Salt Springs, NW of Cow Creek. CA: Inyo; Chlo- 
ride Cliff, CA-NV, SW Va sec. 21, T 28N, R IE, -79 
m, 2-3-85. 230. Nevares Springs. CA: Inyo; Chloride 
Cliff, CA-NV, NE Va sec. 36, T 28N, R IE, 275 m. 231. 
Cottonball Marsh. CA: Inyo; Chloride Cliff, CA-NV, 



2.8 km NE of SW corner of quadrangle, -79 m. 232. 
Salt Springs, W of Cottonball Marsh. CA: Inyo; Chlo- 
ride Cliff, CA-NV, 1.0 km NE of SW corner of quad- 
rangle, -73 m, 2-22-85. 233. Salt Springs S of Cow 
Creek. CA: Inyo; Furnace Creek, CA, SE Va sec. 3, T 
27N, R IE, 31 m, 2-5-85. 234. Stream in Cottonwood 
Canyon. CA: Inyo; Marble Canyon, CA, stream 10.0 
km NW of SE corner of quadrangle, 732 m, 2-15-85, 
stream 11.2 km E-NE of SW corner of quadrangle, 
1037 m, 3-12-87. 235. McLean Spring (Salt Creek). 
CA: Inyo; Stovepipe Wells, CA, SW Va sec. 31, T 15S, 
R 46E, 37 m, 3-8-85. 236. Springs NW of Keane Won- 
der Mine. CA: Inyo; Chloride Cliff, CA-NV, SE Va sec. 
1,T15S,R46E, 366 m, 2- 13-85. 237. Monarch Spring. 
CA: Inyo; Chloride Cliff, CA-NV, SE Va sec. 24, T 14S, 
R 46E, 915 m, 2-4-85. 238. Keane Spring. CA: Inyo; 
Chloride Cliff, CA-NV, NW Va sec. 8, T 30W, R IE, 
1159 m, 2-4-85. 239. Cane Spring. NV: Nye; Cane 
Spring, NV, 6.0 km N-NE of SW corner of quadrangle, 
1238 m. 240. Spring in Oasis Valley, 13.2 km N of 
Beatty. NV: Nye County; Thirsty Canyon, NV, SE Va 
sec. 32, T 1 OS, R 47E, 1 1 28 m. 24 1 . Grapevine Springs. 
CA: Inyo; Ubehebe Crater, CA-NV, NE Va sec. 3, T 
US, R 42E, 824 m. 242. Spring 1.6 km E of Scotty's 
Castle. CA: Inyo; Ubehebe Crater, CA-NV, NW Va sec 
5, T 1 IS, R 42E, 946 m. 243. Little Sand Spring. CA: 
Inyo; Last Chance Range, CA-NV, SE Va sec. 17, T 9S, 
R41E, 915 m, 387. 244. Sand Spring. CA: Inyo; Last 
Chance Range, CA-NV, SE Va sec. 7, T 9S, R 4 IE, 946 
m, 3-30-87. 245. Last Chance Spring. CA: Inyo; Ma- 
gruder Mtn., NV-CA, SE Va sec. 2, T 8S, R 39E, 1739 
m, 3-30-87. 



PROC. BIOL. SOC. WASH. 
102(1). 1989. pp. 249-263 

FOUR NEW SPECIES OF FREE-LIVING MARINE 

NEMATODES IN THE GENUS PAREURYSTOMINA 

(NEMATODA: ENOPLIDA) WITH OBSERVATIONS ON 

OTHER MEMBERS OF THE GENUS 

Edwin J. Keppner 

Abstract. — Four new species of Pareurystomina and the male of P. flagelli- 
caudata are described for the first time from sediments in St. Andrew Bay and 
the Gulf of Mexico. Bay County. Florida. U.S.A. Additional information is 
provided for P. bissonettei, P. floridensis. and P. acuminata. P. americana n. 
sp. and P. xaughtae n. sp. differ from one another in the shape of the spicules 
and gubernaculum. and from P. atypica in the location of the excretory pore 
and shape of the spicules and gubernaculum. P. paraflohdensis n. sp. differs 
from P. floridensis in the number of circles of denticles in the stoma and in 
the size and arrangement of the denticles. P. alima n. sp. differs from P. 
parafloridensis n. sp. in the number of circles of denticles in the stoma, in the 
absence of a gubernaculum. and in the shape of the spicules. It differs from P. 
agubernacula in the number of circles of denticles in the stoma and the length 
and shape of the spicules. The genus Megewystomina is placed in synonymy 
with Parewystomina, and M. combesi becomes P. combesi n. comb. A key to 
the species of the genus Parewystomina is given. 



The genus Parewystomina Micoletzky & 
Kreis, 1930. was erected for those species 
of free-living marine nematodes of the 
subfamily Eurystomininae Filipjev. 1921, 
in which the spinneret and caudal glands 
are absent, and the terminus of the tail is 
acute. Wieser (1959) described Parewys- 
tomina pugetensis, described the cervical 
setae as flattened in this species, and re- 
viewed the known species in the genus. He 
recognized as valid only those species for 
which males had been described. He ex- 
pressed doubt as to the inclusion of Pareu- 
rystomina tenuicauda Stekhoven. 1950. in 
the genus because circles of denticles are 
absent from the stoma in this species where- 
as they are present in all other species in the 
genus. Luc & De Coninck (1959) described 
Pareurystomina armorica and provided a 
key to six species of the genus. 

Chitwood ( 1960) described Pareurystom- 
ina atypica and provided a key to the nine 



species then in the genus and included P. 
tenuicauda. The key used a number of char- 
acters to differentiate the species including 
the number of circles of denticles in the sto- 
ma, tail length, length of spicules, and pres- 
ence or absence of flattened cervical setae. 
He also recognized the problem of the ab- 
sence of male descriptions for some species. 
Hopper (1963) discussed the presence of 
large hypodermal gland cells in Parewys- 
tomina. Hopper (1970) described Parewys- 
tomina bissonettei. discussed the presence 
of flattened cervical setae, the arrangement 
of the denticles in the stoma in Parewys- 
tomina, and emended the description of P. 
atypica to include flattened cervical setae. 
Lambshead &. Piatt (1979) described a new 
genus Bathewystomina. provided a tabular 
key to the genera of the Eury stomininae. 
and discussed each genus in the subfamily. 
They considered male specimens to be crit- 
ical to an understanding of the Eurystomi- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ninae. They considered Eurystomina re- 
panda Wieser, 1959, to be a synonym of P. 
pugetensis based on the presence of flat- 
tened cervical setae and similarity of the 
male genital apparatus. They considered all 
species of Pareurystomina for which males 
are unknown as species dubiae, and P. ten- 
uicauda was considered species dubia due 
to the absence of information pertaining to 
the stoma. 

Collections of free-living marine nema- 
todes from various habitats in St. Andrew 
Bay, and the Gulf of Mexico, Bay County, 
Florida, U.S.A. from 1981-1987 yielded 1 74 
specimens of nine species in the genus Par- 
eurystomina. The purpose of this paper is 
to describe the species collected, provide 
some additional information for known 
species and present a key to the species of 
the genus Pareurystomina. 

Specimens were recovered from sediment 
from a number of subtidal sites in the above- 
mentioned waterbodies with core samplers 
of variable diameters. Nematodes were ex- 
tracted from the samples by repeated wash- 
ing of the sediment in water from the site. 
Nematodes were removed alive from the 
settled material of the final washing, fixed 
in hot alcohol-formalin-acetic acid or hot 
4% formalin in sea water for 24 h and de- 
hydrated and mounted in anhydrous glyc- 
erine on Cobb slides. 

All measurements are given in ^m unless 
otherwise stated, and the mean is followed 
by the range in parentheses. The spicular 
lengths are given as the chord of the arc 
unless otherwise indicated. The taxonomic 
hierarchy is from Gerlach & Riemann 
(1974). Type specimens and specimens of 
known species have been deposited in the 
U.S. National Museum of Natural History 
(USNM) and the Florida Nematode Col- 
lection, University of Florida, Gainesville 
(FNC). 

Taxonomic Account 

The specimens of Pareurystomina col- 
lected during this study presented a variety 



of spicular and gubernacular shapes and 
sizes, a variable number of circles of den- 
ticles in the stoma, and a variety of denticu- 
lar shapes and arrangements. Particular em- 
phasis was given to placing males collected 
during this study with previously described 
species in which the males are unknown. 
The characters used to place males with fe- 
males were the number of circles of stomatal 
denticles, shape and size of the denticles in 
each circle, tail length, size of amphid, and 
location of the excretory pore. Comparison 
of males with males of recognized species 
included the above characters as well as the 
shape of the male genital apparatus. 

All specimens examined during this study 
have the following features in common (Figs. 
1-3). Body size and shape is variable within 
a species as evidenced by the wide range of 
lengths and wide range of the demanian "a" 
values in some species. The cuticle is thin 
and smooth, and the terminus of the tail is 
acute. Caudal glands and spinneret were not 
observed. Hypodermal gland cells are pres- 
ent and arranged as described by Hopper 
(1963, 1970). The gland cells are more 
prominent and abundant in some species 
than in others. The head is retracted in some 
specimens which gives the appearance of an 
anterior displacement of the amphid and 
cephalic setae. The head has six small lips 
each with a papilla, and a single circle of 10 
(6 + 4) cephalic setae. Four of the subme- 
dian cephalic setae are shorter than the oth- 
er six cephalic setae. The lip region is de- 
lineated from the remainder of the head by 
a cephalic groove. The amphid is large, wid- 
er than long, and located over the stoma. 
Stoma is large with heavily cuticularized 
walls and is divided into an anterior and 
posterior chamber by a stomatal suture at 
about midlevel. The stoma has a single, 
large, right subventral tooth and a variable 
number of circles of denticles. The circles 
of denticles can be complete or interrupted 
on the right subventral wall of the stoma as 
described by Hopper (1970). The excretory 
pore is anterior to the level of the cephalic 



VOLUME 102, NUMBER 1 



251 



setae. Cervical setae are compressed and ap- 
pear flattened with a broad base and ter- 
minus. The width and length of these setae 
vary somewhat among species. The female 
reproductive system is amphidelphic and 
the ovaries are antidromous. Males have 
two spicules, and two cuticularized, cup- 
shaped, pre-anal supplements. Two excep- 
tions to these statements were observed 
during this study; a single male of P. flori- 
densis with only one pre-anal supplement 
and a male of P. floridensis in which the 
gubernaculum was absent. Gerlach (1952) 
described a male of Pareurystomina acu- 
minata (De Man, 1889) with a single pre- 
anal supplement. A gubernaculum is absent 
or reduced in some species of Pareurysto- 
mina. 

The characters of most value in differ- 
entiating the species of Pareurystomina are 
the shape and size of the spicules and the 
presence or absence and shape of the gu- 
bernaculum. The number of circles of den- 
ticles in the stoma, the shape of the denticles 
in each circle, tail length and position of the 
excretory pore were also of value in differ- 
entiating both males and females of the var- 
ious species. The use of the number of cir- 
cles of denticles in the stoma can be a 
problem because the posterior circles are 
often incomplete. Hopper (1970), in the de- 
scription of P. bissonettei, noted that the 
fewest rows of denticles occur on the wall 
of the stoma from which the large subven- 
tral tooth originates, and the largest number 
of rows of denticles occurs on the stomatal 
wall opposite the origin of the subventral 
tooth. The presence of partial circles of den- 
ticles and the location of the interruption 
requires examination from lateral aspects in 
order to determine accurately the maxi- 
mum number of denticular circles in the 
stoma. Some descriptions of Pareurysto- 
mina species are not clear as to whether the 
maximum or minimum number of denticu- 
lar circles is stated. It is assumed, for pur- 
poses of this paper, that the stated or figured 
number of circles of denticles for each pre- 




Figs. 1-3. Pareurystomina floridensis: 1, Male, head, 
lateral view; 2, Entire male, lateral view; 3, Male, an- 
terior pre-anal supplement, lateral view. Scale bars in 
nm. Legend: a = amphid. aa = anterior apophysis, eg 
= cervical groove, cs = cephalic setae, d = denticles, 
ep = excretory pore, es = esophagus, fs = flat cervical 
setae, ga = gubernacular apophysis, hg = hypodermal 
gland cell, in = intestine, lp = labial papilla, nr = nerve 
ring, p = papilla, pa = posterior apophysis, ps = pre- 
anal supplement, r = rim of cup. s = spicule, ss = 
stomatal suture, st = subventral tooth, t = testis. 



viously described species is the maximum 
number. 

Wieser (1959) recognized that the size and 
shape of the denticles can vary between cir- 
cles in Pareurystomina and used these dif- 
ferences with other characters to distinguish 
the species. In the specimens examined dur- 
ing this study, the size and shape of the 
denticles varied between circles in some 
species but were of the same size and shape 
within a given circle in the same species. 

Pareurystomina americana, new species 
Figs. 4-12 

Diagnosis. —(Ten males, 3 females and 5 
juveniles from St. Andrew Bay.) Body long, 
slender. Four submedian cephalic setae 



252 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 4-12. Pareurystomina americana: 4, Male 
paratype, head, lateral view; 5, Male holotype, head, 
lateral view; 6, Male paratype, head, lateral view; 7, 
Male holotype, posterior end, lateral view; 8, Male 
holotype, pre-anal supplements, lateral view (anterior, 
left; posterior, right); 9, Male holotype, right spicule 
and gubernaculum, lateral view; 10, Male paratype, 
right spicule and gubernaculum, lateral view; 1 1 , Male 
paratype, pre-anal supplements, lateral view (anterior, 
left; posterior, right); 12, Female, posterior end, lateral 
view. Scale bars in /xm. 



about l h length of other 6. Flat cervical setae 
with narrow base present from base of sto- 
ma to about Vi distance to nerve ring; one 
male with additional setae for about l h dis- 
tance from nerve ring to base of esophagus. 
Somatic setae not observed; caudal setae 
sparse in male, not observed in female. Am- 
phids over midlevel of stoma. Single com- 
plete circle triangular denticles and one in- 
complete circle small, round denticles 
anterior to stomatal suture; single incom- 
plete circle large, triangular denticles pos- 
terior to suture. Excretory pore anterior to 
cephalic setae. Tail conical with acute ter- 
minus. 

Males (n = 6): Length 4.52 mm (4.24- 



4.94), width at midbody 35.8 (30-43). Head 
diameter 27 (25-29) at level of cephalic se- 
tae; cephalic setae 15 (14-17) and 5.3 (5-6) 
long. Amphid 13.7 (13-16) wide. Stoma 24.2 
(23-26) long, 17.5 (16-19) wide. Esophagus 
0.964 mm (0.730-1.10) long; nerve ring 
188.2 (179-195) from anterior end. Tail 

140.2 (138-163) long, 34.2 (32-37) wide at 
anus. Two midventral, cuticularized, cup- 
shaped, pre-anal supplements present 119 
(107-136) and 228 (208-248) anterior to 
anus. Posterior apophysis of each supple- 
ment larger than anterior; rim of cup striat- 
ed. Two pairs subventral papillae between 
anus and first pre-anal supplement; two pairs 
subventral papillae between first and second 
pre-anal supplements. Two pairs subventral 
setae immediately anterior to anus; subven- 
tral, lateral, subdorsal setae present poste- 
rior to anus. Spicules gently curved 41.8 
(37-45) long; tips blunt, barb absent. Gu- 
bernaculum with narrow corpus surround- 
ing spicules; gubernacular apophysis 14.7 
(14-16) long, bent dorsally at midpoint, a 
= 125.9 (1 14.9-147.8); b = 4.77 (3.96-6.03); 
c = 30.8 (27.0-33.6). 

Females (n = 3): Length 4.19 mm (3.36- 
4.65), width at midbody 42.3 (42-45). Head 
diameter 28.7 (28-30) at level of cephalic 
setae; cephalic setae 15 (14-17) and 5.3 (5- 
6) long. Amphid 12.3 (11-13) wide. Stoma 
24 (22-26) long and 18.7 (18-19) wide. 
Esophagus 848 (789-896) long, nerve ring 

182.3 (171-200) from anterior end. Tail 
186.7 (165-203) long, 33.7 (32-35) wide at 
anus. Vulva 2.76 mm (2.15-3.1 1) from an- 
terior end. a = 98.7 (84.0-1 10.7); b = 4.92 
(4.26-5.31); c = 23.7 (17.5-31.2); V = 65.7% 
(64-68). 

Type specimens. —Holotype male, USNM 
77137; paratype males, USNM 77138- 
77140; allotype female, USNM 77141; 
paratype female, USNM 77142. Paratype 
males, FNC A- 144 and A- 146. 

Type locality. —Holotype male from non- 
vegetated sediment from the north shore of 
St. Andrew Bay, Bay County, Florida 0.3 
km west of the inlet to Lake Caroline 



VOLUME 102, NUMBER 1 



253 



(85°41'10"W, 30°09'45"N). Also in vegetat- 
ed and nonvegetated sediment from Fresh- 
water Bayou, St. Andrew Bay, Bay County, 
Florida, (85°39'00"W, 30°07'30"N). 

Etymology. —Named for the place of col- 
lection, America. 

Remarks. —Pareurystomina americana is 
most similar to Pareurystomina atypica 
Chitwood, 1 960, from which it differs in the 
location of the excretory pore (anterior to 
cephalic setae vs. 104 from anterior end), 
in the width of the amphid (11-16 vs. 7), 
in the shape of the spicules (blunt tip vs. 
narrow, pointed tip) and in the shape of the 
gubernacular apophysis (bent dorsally vs. 
bent ventrally). P. americana is also similar 
to P. vaughtae described next but differs 
from P. vaughtae in the shape of the spicules 
(absent of barb at tip), the corpus of the 
gubernaculum is not conical and the gu- 
bernacular apophysis is bent dorsally rather 
than straight. 

Pareurystomina vaughtae, new species 
Figs. 13-20 

Diagnosis.— (Five males, 3 females, and 
3 juveniles from mouth of Freshwater Bay- 
ou and Grand Lagoon, St. Andrew Bay.) 
Body long, slender. Four submedian ce- 
phalic setae about 28% length of other 6. 
Flat cervical setae narrow; present from base 
of stoma almost to nerve ring. Somatic setae 
sparse; caudal setae abundant in males, 
sparse in females. Amphids over midlevel 
of stoma. Single complete circle large tri- 
angular denticles anterior to stomatal su- 
ture, single incomplete circle large, trian- 
gular denticles posterior to suture. Excretory 
pore anterior to cephalic setae. Tail conical 
with acute terminus. 

Males (n = 5): Length 3.24 mm (2.63- 
4.70), width at midbody 33.8 (30-37). Head 
diameter 26.6 (26-29) at level of cephalic 
setae; cephalic setae 18.1 (16-20) and 4.8 
(4-5) long. Amphid 1 1.4 (1 1-12) wide. Sto- 
ma 24.6 (24-27) long, 17.2 (16-18) wide. 
Esophagus 713.2 (606-936) long; nerve ring 
179 (168-200) from anterior end. Tail 138 




Figs. 13-20. Pareurystomina vaughtae: 13, Male 
holotype, head, ventral view; 14, Male paratype, head, 
lateral view; 15, Male holotype, posterior end, lateral 
view; 16, Male, anterior pre-anal supplement, lateral 
view; 17, Male holotype, left spicule and gubernacu- 
lum, lateral view; 18, Male holotype, anterior pre-anal 
supplement, lateral view; 19, Male paratype, left spic- 
ule and gubernaculum, lateral view; 20, Female, tail, 
lateral view. Scale bars in ^m. 



(126-144) long, 31.6 (29-35) wide at anus. 
Two cuticularized, cup-shaped, midventral, 
pre-anal supplements present 118.8 (114- 
136) and 200.4 (184-216) anterior to anus. 
Posterior apophysis of each supplement 
larger than the anterior; rim of cup not 
striated. Two pairs subventral papillae be- 
tween anus and first pre-anal supplement. 
Two pairs subventral setae immediately an- 
terior to anus. Spicules 41.8 (38-45) long, 
gently curved, tip with small barb. Corpus 
of gubernaculum 7.2 (6-8) long, conical, 
surrounds distal end of spicules; gubernacu- 
lar apophysis straight, 11 (8-13) long, di- 
rected dorsally. Post-anal setae abundant, a 
= 95.0 (80.0-127.0); b = 4.50 (3.95-5.02); 
c = 23.4(18.7-32.6). 



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Females (n = 3): Length 3.32 mm (3.07- 
3.54), width at midbody 45.3 (43-48). Head 
diameter 28.3 (27-29) at level of cephalic 
setae; cephalic setae 19.3 (18-21), 4.7 (4-5) 
long. Amphid 11.7(11-13) wide. Stoma 23.3 
(22-24) long, 19 (18-20) wide. Tail 166.3 
(163-168) long, 34.7 (32-37) wide at anus. 
Vulva 2.05 mm (1.91-2.16) from anterior 
end. a = 73.4 (63.9-78.7); b = 3.83 (3.69- 
4.02); c = 19.9 (18.8-21.1); V = 61.7% (61- 
62). 

Type specimens. — Holotype male, USNM 
77143; paratype males, USNM 77144- 
77145; allotype female, USNM 77146. 
Paratype male, FNC A- 155; paratype fe- 
male, FNC A-156. 

Locality. —Holotype from sediments in a 
bed of Shoalgrass (Halodule wrightii) at the 
mouth of Freshwater Bayou, St. Andrew 
Bay, Bay County, Florida (85°39'00 ,, W, 
30°07'30"N). Other site, Grand Lagoon off 
St. Andrew Bay, Bay County, Florida 
(85°43'15' , W, 30°07'50' , N). 

Etymology. —Named for Ms. Rosalie 
Vaught, librarian. This and other studies 
could not have been completed without her 
voluntary dedication to obtaining the re- 
quired literature. 

Remarks.— Par eurystomina vaughtae is 
most similar to P. americana but differs from 
it in the presence of the large cone-shaped 
corpus of the gubernaculum, straight rather 
than bent gubernacular apophysis, and the 
presence of two rather than three circles of 
denticles in the stoma. P. vaughtae is also 
similar to P. armorica in the presence of 
two circles of denticles in the stoma. It dif- 
fers from that species in that the excretory 
pore is anterior to the cephalic setae rather 
than at the cephalic groove, the amphid is 
larger (50% of head diameter vs. 31% of 
head diameter), and the tail is shorter (4.8 
anal diameters vs. 6 anal diameters). 

Pareurystomina parafloridensis, new species 
Figs. 21-28 

Diagnosis. —(Eleven males, 4 females and 
6 juveniles from various sites in St. Andrew 



Bay). Body long, broad. Four submedian 
cephalic setae about l h length of other 6. 
Flat cervical setae broad; present from base 
of stoma almost to nerve ring. Somatic setae 
and caudal setae sparse. Amphids over mid- 
level of stoma. Stoma with 4 circles of den- 
ticles anterior to stomatal suture, one circle 
posterior to suture. Last circle anterior to 
suture and circle posterior to suture incom- 
plete. Small triangular denticles in anterior- 
most 4 circles uniform in size within each 
circle and between circles. Posterior-most 
circle with larger triangular denticles. Ex- 
cretory pore anterior to cephalic setae. Male 
tail, short, conical with acute terminus. Fe- 
male tail longer, conical then narrow to acute 
terminus. 

Males (n = 6): Length 4.59 mm (3.95- 
5.10), width at midbody 70 (61-75). Head 
diameter 37.3 (37-38) at level of cephalic 
setae; cephalic setae 16.3 (15-18) and 5 (4- 
6) long. Amphid 20.3 (19-21) wide. Stoma 
30.2 (29-32) long, 24.6 (24-26) wide. 
Esophagus 1.02 mm (0.971-1.08) long; 
nerve ring 230.7 (224-242) from anterior 
end. Tail 188.8 (157-222) long, 57.5 (56- 
61) wide at anus. Two cuticularized, cup- 
shaped, midventral, pre-anal supplements 
present 179.5 (152-208) and 295 (240-360) 
anterior to anus. Posterior apophysis of each 
supplement about twice size of anterior 
apophysis; rim of cup striated. Two pairs 
small subventral papillae immediately an- 
terior to anus and another more lateral pair 
posterior to anus. Spicules 78.3 (75-82) long, 
gently curved then recurved at tip; tip broad 
with large barb. Gubernaculum with small 
corpus and narrow apophysis 34.3 (32-38) 
long; lateral parts of apophysis more heavily 
cuticularized; apophysis directed almost 
dorsally. a = 65.3 (60.3-70.8); b = 4.51 
(3.91-4.95); c = 24.5 (20.6-27.7). 

Females (n = 4): Length 4.39 mm (3.63- 
4.96), width at midbody 104.8 (99-109). 
Head diameter 37.5 (34-40) at level of ce- 
phalic setae; cephalic setae 17 (16-18) and 
5.5 (5-6) long. Amphid 18.8 (18-19) wide. 
Stoma 32 (30-34) long, 27.3 (26-29) wide. 



VOLUME 102, NUMBER 1 



255 



Esophagus 1.06 mm (0.846-1.12) long; 
nerve ring 240.8 (213-254) from anterior 
end. Tail 283.5 (258-323) long, 52 (48-54) 
wide at anus. Caudal setae restricted to anal 
region. Vulva 2.68 mm (2.03-3.12) from 
anterior end. a = 43.7 (36.7-51.7); b = 4.15 
(3.94-^4.31); c = 15.5 (14.0-17.1); V = 62% 
(60-64). 

Type- specimens.— Male holotype, USNM 
77147; male paratypes, USNM 77148- 
77149; female allotype, USNM 77150; fe- 
male paratype, USNM 77151. Paratype 
males, FNC A- 152, A- 153; female para- 
type, FNC A-154. 

Locality. —Holotype male from sedi- 
ments in a bed of shoalgrass {Halodule 
wrightii) in Grand Lagoon, St. Andrew Bay, 
Bay County, Florida (85°43'15"W, 
30°07'50"N). Other locality, sediments in a 
bed of manatee grass (Syringodium fili- 
forme) at mouth of Freshwater Bayou, Bay 
County, Florida (85°39'00' , W, 30°07'30"N). 

Etymology. —From the Greek para 
meaning beside and floridensis referring to 
Pareurystomina floridensis. 

Remarks. —Pareurystomina paraflori- 
densis is similar to P. floridensis in the length 
and shape of the spicules and gubernaculum 
in males and most characters in females. P. 
parafloridensis differs from P. floridensis 
primarily in the number of circles of den- 
ticles in the stoma and the shape and size 
of the denticles in each circle. P. paraflori- 
densis has five circles of stomatal denticles 
and the anterior four circles are of equal size 
and shape; each denticle is a small triangular 
structure. The posterior-most circle has 
larger triangular denticles. In P. floridensis 
there are three to four circles of denticles of 
variable size and shape; the denticles of the 
anterior-most circle are large elongate trian- 
gules, the following two circles are small and 
rounded and the posterior circle is large and 
rounded. P. parafloridensis is also similar 
to Pareurystomina combesi (Luc & De Con- 
inck, 1959) n. comb, in the number of rows 
of denticles in the stoma. It differs from P. 
combesi in that the excretory pore is ante- 




Figs. 21-28. Pareurystomina parafloridensis: 21, 
Male holotype, head, lateral view; 22, Male holotype, 
head, lateral view; 23, Male holotype, posterior end, 
lateral view; 24, Male holotype, left spicule and gu- 
bernaculum, lateral view; 25, Male holotype, pre-anal 
supplements, lateral view (posterior, top right; anterior, 
bottom left); 26, Female, posterior end, lateral view; 
27, Male paratype, left spicule and gubernaculum, lat- 
eral view; 28, Female, head, lateral view. Scale bars in 
urn. 



rior to the cephalic setae rather than pos- 
terior to the cephalic groove and the female 
tail is longer (c = 14-17.1 vs. c = 47.8). 

Pareurystomina alima, new species 
Figs. 29-36 

Diagnosis. — (Four males, 3 females, and 
5 juveniles from mouth of Freshwater Bay- 
ou and Grand Lagoon, St. Andrew Bay). 
Body short, broad. Four submedian ce- 
phalic setae about Va length of other 6. Flat 
cervical setae broad; extend from base of 
stoma to nerve ring; somatic setae not ob- 
served; caudal setae present. Amphids over 
level of stomatal suture when relaxed, an- 
terior to stomatal suture when contracted. 
Denticles in 2 broad bands of 3 circles each; 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 29-36. Pareurystomina alima: 29, Male ho- 
lotype, head, right lateral view (contracted); 30, Male 
holotype, posterior end, lateral view; 3 1 , Male holotype 
pre-anal supplements, lateral view (anterior, top; pos- 
terior, bottom); 32, Male paratype, right spicule, lateral 
view; 33, Female, tail, lateral view; 34, Male holotype, 
right spicule, lateral view; 35, Female, head, lateral 
view; 36, Male paratype, head, lateral view. Scale bars 
in mm. 



first 4 circles anterior to suture complete 
with small, triangular denticles; next 2 cir- 
cles posterior to suture incomplete; first with 
small, round denticles; second with large, 
triangular denticles. Excretory pore anterior 
to cephalic setae. Tail conical, terminus 
acute. 

Males (n = 4): Length 3.22 mm (3.05- 
3.57), width at midbody 75.5 (53-86). Head 
diameter 38 (32-40) at level of cephalic se- 
tae; cephalic setae 13.5 (13-14), 3.7 (3-4) 
long. Amphid 16.5 (14-19) wide. Stoma 26 
(24-27) long, 23.8 (21-25) wide. Esophagus 
709.8 (598-792) long; nerve ring 210.5 (194- 



219) from anterior end. Tail 168.5 (128- 
184) long, 46 (38-50) wide at anus. Two 
midventral, cuticularized, cup-shaped, pre- 
anal supplements present 130.5 (70-160) 
and 21 1.8 (101-272) anterior to anus. Pos- 
terior apophysis of each supplement about 
twice size of anterior apophysis; rim of cup 
not striated. Two pairs circular, subventral, 
plaque-like papillae between anus and first 
pre-anal supplement. Two pairs small, sub- 
ventral setae immediately anterior to anus. 
Spicules straight, broad, 50 (45-54) long. 
Gubernaculum absent, a = 44.0 (36.6-57.4); 
b = 4.57 (4.36-5.10); c = 19.5 (17.1-23.8). 

Females (n = 3): Length 4.19 mm (3.24- 
4.90), width at midbody 90 (80-104). Head 
diameter 42.3 (40-45) at level of cephalic 
setae; cephalic setae 14.7 (14-16), 3.7 (3-4) 
long. Amphid 17.3(16-1 9) wide. Stoma 30.3 
(27-32), 27.7 (26-30) wide. Esophagus 1.03 
mm (0.843-1.15) long; nerve ring 236.7 
(213-259) from anterior end. Tail 282.6 
(248-328) long, 50 (48-51) wide at anus. 
Caudal setae sparse. Vulva 2.46 mm (1.83- 
2.94) from anterior end. a = 47.8 (31.2- 
57.0); b = 4.04 (3.84-4.26); c = 14.8 (13.1- 
16.3); V = 58.3% (56-60). 

Type-specimens.— Holotype male, USNM 
77152; paratype males, USNM 77153- 
77154; allotype female, USNM 77155; 
paratype female, USNM 77156. Paratype 
male, FNC A- 143; paratype female, FNC 
A-145. 

Locality.— Holotype male nonvegetated 
sediments from the mouth of Freshwater 
Bayou, St. Andrew Bay, Bay County, Flor- 
ida (85°39'00"W, 30°07'30"N). Other site, 
nonvegetated sediments in Grand Lagoon 
off St. Andrew Bay, Bay County, Florida 
(85°43'15"W, 30°07'50"N). 

Etymology. — From the Greek halimos 
meaning pertaining to the sea. 

Remarks. —Pareurystomina alima is most 
similar to P. parafloridensis in the stomatal 
characters. It differs in the greater number 
of circles of denticles (6 vs. 5), in the shape 
and size of the denticles in the posterior- 
most two circles, and the denticles in each 



VOLUME 102, NUMBER 1 



257 



circle are smaller. P. alima males differ from 
P. parafloridensis in the absence of a gu- 
bernaculum and in the short straight spic- 
ule. P. alima is similar to P. agubernacula 
Keppner, 1986, in the absence of a guber- 
naculum and the presence of a short straight 
spicule. It differs from P. agubernacula in 
the greater number of circles of denticles (6 
vs. 2) and shape and length of the spicules 
(45-54 vs. 25-27). 

Pareurystomina flagellicaudata 

Stekhoven, 1946 

Figs. 37-41 

Diagnosis. — (One male from site in Gulf 
of Mexico, Bay County, Florida.) Body very 
long, slender. Four submedian cephalic se- 
tae about l h length of other 6. Rat cervical 
setae short, broad; present from base of sto- 
ma to about 2 h distance to nerve ring. So- 
matic setae and caudal setae sparse. Am- 
phid just posterior to midlevel of stoma. 
Denticles in 2 circles, one complete circle 
large, triangular denticles anterior to suture, 
one incomplete circle large, triangular den- 
ticles posterior to suture. Excretory pore an- 
terior to cephalic setae. Tail conical then 
flagellate with acute terminus. 

Male (n = 1): Length 7.83 mm, width at 
midbody 48. Head diameter 30 at level of 
cephalic setae; cephalic setae 22 and 7 long. 
Amphid 14 wide. Stoma 29 long, 18 wide. 
Esophagus 782 long; nerve ring 248 from 
anterior end. Tail 600 long and 45 wide at 
anus. Two cuticularized, cup- shaped, mid- 
ventral supplements present 139 and 304 
anterior to anus. Anterior and posterior 
apophyses of each supplement about equal; 
rim of cup not striated. Two pairs subven- 
tral setae immediately anterior to anus; sub- 
ventral and dorsolateral setae present from 
anus to just anterior to anterior supplement. 
Conical portion of tail with numerous small 
setae. Spicules almost straight, 48 long with 
blunt tip. Gubernaculum with short, weakly 
cuticularized apophysis 6 long, corpus a thin 
distal extension, a = 163: b = 10.0: c = 13.5 




Figs. 37-41. Pareurystomina flagellicaudata: 37, 
Male, anterior end, lateral view; 38, Male, posterior 
end, lateral view; 39, Male, pre-anal supplements, lat- 
eral view (posterior, top; anterior, bottom); 40, Male, 
head, lateral view; 4 1 , Male, right spicule and guber- 
naculum, lateral view. Scale bars in yum. 

Specimen.— One male, USNM 77157. 

Locality.— Coarse sand and shell near 
ledge in water 24 m deep in the Gulf of 
Mexico, Bay County, Florida (85°49'30"W, 
30°01'30"N). 

Remarks. —Pareurystomina flagellicau- 
data was described from a single female 
specimen. Hopper (1963) observed a spec- 
imen of P. flagellicaudata from Woods Hole, 
Massachusetts U.S.A., but did not mention 
the sex or provide figures of the specimen. 
To the writer's knowledge, the male of this 
species has not been described. The speci- 
men described above is considered to be P. 
flagellicaudata based upon the number of 
rows of denticles in the stoma and the length 
of the tail. It differs from the original de- 
scription in the length of the cephalic setae 
and in having distinctly flattened cervical 
setae. However, the original description did 
not figure or state that the cervical setae 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 42-46. Pareurystomina acuminata: 42, Male, 
head, right lateral view; 43, Male, head, left lateral 
view; 44, Male, posterior end, lateral view; 45, Male, 
pre-anal supplements, lateral view (posterior, left; an- 
terior, right); 46, Male, right spicule and gubernaculum, 
lateral view. Scale bars in nm. 



were flat or not, and the position of the ex- 
cretory pore was not mentioned. 

Pareurystomina acuminata 

(DeMan, 1889) 

Figs. 42-46 

Diagnosis, —(one male from sediment in 
St. Andrew Bay.) Body long, slender. Four 
submedian cephalic setae about l h length of 
other 6. Fiat cervical setae broad, restricted 
to region just posterior to stoma. Somatic 
and caudal setae sparse. Amphid wide, an- 
terior to stomatal suture. Two circles tri- 
angular denticles present anterior to sto- 
matal suture, posterior circle incomplete 
with slightly smaller denticles; denticles ab- 
sent posterior to suture. Excretory pore an- 
terior to cephalic setae. Tail conical, tapers 
rapidly to acute terminus. 



Male (n = 1): Length 3.50 mm, width at 
midbody 37. Head diameter at level of ce- 
phalic setae 24; cephalic setae 1 5 and 5 long. 
Amphid 13 wide. Stoma 28 long, 14 wide. 
Esophagus 651 long; nerve ring 194 from 
anterior end. Tail 235 long, 37 wide at anus. 
Two midventral, cuticularized, cup-shaped, 
pre-anal supplements present 165 and 309 
anterior to anus. Anterior and posterior 
apophyses of each supplement about equal. 
Two pairs subventral papillae between anus 
and first pre-anal supplement. Single pair 
subventral setae immediately anterior to 
anus. Spicules 55 (chord), 72 (arc) long, 
strongly arcuate, tip with narrow velum and 
small lateral barb. Corpus of gubernaculum 
14 long, does not surround spicules; gu- 
bernacular apophysis 16 long, straight, 
broad, heavily cuticularized, directed pos- 
tero-dorsally. a = 94.4; b = 5.38; c = 14.9. 

Specimen. -Male, USNM 77158. 

Locality.— Coarse sand and shell from 
water about 9 m deep in St. Andrew Bay off 
the mouth of Freshwater Bayou, Bay Coun- 
ty, Florida (85°39'17"W, 30°07'45"N). 

Remarks. — Gerlach (1952) described a 
male of P. acuminata and figured the am- 
phid as spiral in shape. Blome (1974) de- 
scribed two males of P. acuminata and fig- 
ured the amphid as a loose spiral. Neither 
author mentioned or figured the position of 
the excretory pore, and both figured the cer- 
vical setae as not flattened. The specimen 
described herein as P. acuminata is similar 
in all important characters to those de- 
scribed above with the exception of the shape 
of the amphid and the presence of flat cer- 
vical setae. 

Hopper (1963) described a female nema- 
tode collected at Gulf Shores, Alabama, 
U.S.A. as P. acuminata. However, this 
specimen has three circles of denticles in the 
stoma, a long tail, an oblong amphid, and 
the cephalic setae are shorter than those de- 
scribed for P. acuminata. The description 
and figures given by Hopper (1963) appear 
to be that of P. americana n. sp. with the 
exception of the absence of flat cervical se- 



VOLUME 102, NUMBER 1 



259 



tae. The cervical setae in P. americana are 
narrow and could be overlooked. Therefore, 
P. acuminata sensu Hopper (1963) is con- 
sidered to be a synonym of P. americana. 

Pareurystomina bissonettei Hopper, 1970 
Figs. 47-50 

This species was the second most abun- 
dant Pareurystomina collected (20 males, 
14 females, and 9 juveniles). It was present 
in nonvegetated and vegetated, shallow 
water, estuarine sediments and one male was 
obtained from nonvegetated sediments in 
the Gulf of Mexico. Specimens agree with 
the description given by Hopper (1 970). The 
notable exception is the male collected in 
the Gulf of Mexico. This male is appreciably 
larger but morphologically similar to the 
males from shallow, estuarine sediments. 
The body measurements are about equally 
proportional to the smaller estuarine males. 
A description of this male follows. 

Diagnosis. — Body long, slender. Large, 
vacuolated, subcuticular cells present in ad- 
dition to the hypodermal gland cells. Four 
submedian cephalic setae about l A length of 
other 6. Flat cervical setae short, narrow, 
extend about Vi distance to nerve ring. So- 
matic setae short, sparse; caudal setae nar- 
row, flattened in males. Amphid wide, at 
midlevel of stoma. Denticles in 5 circles; 
those in 2 posterior circles slightly larger 
than those in anterior circles. Tail short, 
with narrow terminal spike with minute 
spines. 

Male (n = 1): Length 7.56 mm, width at 
midbody 107. Head diameter 85 at level of 
cephalic setae; cephalic setae 40 and 1 1 long. 
Stoma 66 long, 56 wide. Amphid 35 wide. 
Esophagus 1.70 mm long; nerve ring 320 
from anterior end. Tail 145 long, 88 wide 
at anus; caudal setae numerous. Two cuti- 
cularized, cup-shaped, midventral, pre-anal 
supplements present 221 and 422 anterior 
to anus. Cup of each supplement with 4 
arms each with striated distal terminus; arms 
directed anteriorly, posteriorly, and later- 
ally. Posterior apophysis of each supple- 




Figs. 47-54. Pareurystomina bissonettei: 47, Male, 
head, sublateral view; 48. Male, pre-anal supplements, 
lateral view (anterior, left; posterior, right); 49, Male, 
posterior end, lateral view; 50, Female, tail, lateral view. 
Pareurystomina floridensis: 5 1 , Male, head, lateral view; 
52, Male, left spicule and gubernaculum. lateral view. 
Pareurystomina agubernacula: 53, Male, head, lateral 
view; 54, Male, left spicule, lateral view. Scale bars in 
/urn for all figs. 

ment very large, anterior apophysis small. 
Two pairs subventral papillae between anus 
and first pre-anal supplement. Single pair 
subventral setae immediately anterior to 
anus. Spicules 128 long, arcuate with small 
barb at tip. Corpus of gubernaculum does 
not surround spicules; gubernacular apoph- 
ysis 26 long, directed dorsally. a = 70.7; b 
= 4.45; c = 52.1 

Specimen.— One male, USNM 77165; 
other males. 77159-77163: females. 77164 
and 77 166. Two males. FNC A- 147. A- 148, 
A- 149; female, A- 150. 

Locality.— Coarse sand and shell near 
ledge in water about 24 m deep. Gulf of 
Mexico, Bay County. Florida (85°49'30"W, 
30°0 1 '30"N). Other specimens from various 
sites in St. Andrew Bay. Florida. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Pareurystomina floridensis Keppner, 1986 
Figs. 51-52 

This species was the most abundant en- 
countered during this study (33 males, 16 
females, and 6 juveniles). It was recovered 
only from shallow water, nonvegetated sed- 
iments at a variety of sites in St. Andrew 
Bay and Lake Powell, Bay County, Florida. 
Examination of specimens recovered after 
the original description reveals that in some 
of the specimens there is an additional, par- 
tial circle of very small, round denticles 
present just anterior to the stomatal suture 
in addition to the three circles originally 
described. The apophysis of the gubernac- 
ulum in the specimens examine curves 
slightly ventrally at the tip. Figures are in- 
cluded for comparative purposes. 

Pareurystomina agubernacula 

Keppner, 1986 

Figs. 53-54 

This species has not been recovered since 

the original description, and the figures are 

included for comparative purposes. 

Megeurystomina combesi 
Luc & De Coninck, 1959 

Luc & De Coninck (1959) described a 
new genus and species of Eurystomininae, 
Megeurystomina combesi. They differen- 
tiated the genus Megeurystomina from Par- 
eurystomina on the basis of the short length 
of the tail, the structure of the head partic- 
ularly its ability to retract and protract, and 
body size (10 mm and greater) in Megeu- 
rystomina. Lambshead & Piatt (1979) con- 
sidered Megeurystomina to be a dubious 
genus due to the absence of a description of 
the male. The tail length (2 anal diameters), 
as a generic character for Megeurystomina, 
is contradicted by P. bissonettei that also 
has a short tail (1 .5-2.5 anal diameters). The 
large male of P. bissonettei collected during 
this study from the Gulf of Mexico is about 
1.6 times larger than the average P. bisso- 
nettei male collected from inshore waters. 



This reduces the importance of body size as 
a generic character for Megeurystomina. The 
retracted head observed in some of the spec- 
imens examined during this study reduces 
the importance of this as a generic character. 
M. combesi, therefore, may be a large form 
of a species of Pareurystomina. The struc- 
ture of the head is similar to that of the 
species of Pareurystomina with five or six 
circles of stomatal denticles. For these rea- 
sons, Megeurystomina is considered to be 
a synonym of Pareurystomina and the 
species Pareurystomina combesi (Luc & De 
Coninck, 1959) n. comb, is included in the 
key. It can be differentiated from the other 
species on the basis of the number of circles 
of denticles in the stoma, the position of the 
excretory pore, and the tail length. 

Key to Species of 
Adult Pareurystomina 

The taxonomic history of the genus Par- 
eurystomina is similar to that of the genus 
Eurystomina Filipjev, 1921, with regard to 
the characters of importance in differen- 
tiating species. Wieser (1953) stressed the 
importance of the characters of the head in 
distinguishing the species of Eurystomina. 
Inglis (1962) stressed the importance of the 
male genital apparatus, particularly the 
shape of the gubernaculum, in distinguish- 
ing the species of Eurystomina, and Yeates 
(1967) agreed with this approach. Wieser & 
Hopper (1 967) recognized the value of using 
a combination of characters involving the 
head and male genital apparatus. As in the 
Eurystomina, the identification of the 
species of Pareurystomina is best achieved 
when males are available for study. 

In the following key, the maximum num- 
ber of circles of denticles in the stoma, po- 
sition of the excretory pore, and tail length 
are used to differentiate those species for 
which only females are known. The use of 
these characters allowed all species previ- 
ously described to be placed in the key with 
the exception of P. parapugetensis Vitiello, 
1970, described from one juvenile and P. 



VOLUME 102, NUMBER 1 



261 



tenuissima (Filipjev, 1927) Filipjev 1946 
described from one female. Both species 
could not be placed in the key because the 
position of the excretory pore is unknown. 
Therefore, these two species remain species 
dubiae in accordance with Lambshead & 
Piatt (1979). The presence or absence of flat 
cevical setae was not used in the key because 
their presence on the species described prior 
to Hopper (1970) is uncertain. 

1 . Circles of denticles present in 
stoma 2 

- Circles of denticles absent from 
stoma 

. . .P. tenuicauda Stekhoven, 1950 
2(1). One to two circles of denticles 

in stoma 3 

- Three or more circles of den- 
ticles in stoma 11 

3(2). One circle of denticles in sto- 
ma 4 

- Two circles or denticles in sto- 
ma 5 

4(3). Tail 1 8 anal diameters long . 

P. filicaudata Allgen, 1934 

- Tail 4.5 anal diameters long 

... P. typica Micoletzky & Kreis, 

1930 
5(3). Excretory pore posterior to 

base of stoma 6 

- Excretory pore anterior to base 

of stoma 7 

6(5). Stoma twice as long as wide; 
tail 10-11 anal diameters long 

(male unknown) 

. . . P. biserialis Stekhoven, 1946 

- Stoma about 1.5 times as long 
as wide; tail less than 10 anal 
diameters long; spicules slen- 
der with recurved tip 

P. atypica Chitwood, 1960 

7(5). Excretory pore at cephalic 
groove; amphid 31% of head 
diameter; tail 6 anal diameters 

long (male unknown) 

. .P. armorica Luc & De Coninck, 

1959 



- Excretory pore anterior to ce- 
phalic groove and setae; am- 
phid greater than 40% of head 
diameter; tail less than 6 anal 
diameters long 8 

8(7). Gubernaculum absent; spic- 
ules straight 

. . P. agubernacula Keppner, 1986 

- Gubernaculum present; spic- 
ules straight to curved 9 

9(8). Gubernacular apophysis re- 
duced; spicules straight; tail 1 5 

anal diameters long 

. . . P. flagellicaudata Stekhoven, 

1946 

- Gubernacular apophysis dis- 
tinct, elongate; spicules curved; 
tail less than 6 anal diameters 
long 10 

1 0(9). Spicules thin, strongly arcuate, 
tip not recurved, with small 
lateral barb; corpus of guber- 
naculum elongate, does not 
surround spicules; gubernac- 
ular apophysis straight, direct- 
ed postero-dorsally 

. . P. acuminata (De Man, 1889) 

- Spicules broad, gently curved, 
tip recurved, small lateral barb 
absent; corpus of gubernacu- 
lum cone-shaped, surrounds 
spicules; gubernacular apoph- 
ysis straight, directed dorsally 

P. vaughtae, n. sp. 

1 1(2). Tail short, cylindrical, abrupt- 
ly narrows to terminal spike 
with minute spines in male, 
terminal spines absent in fe- 
male P. bissonettei Hopper, 1970 

- Tail conical, tip without mi- 
nute spines in males and fe- 
males 12 

12(11). Stoma with 5-6 circles of den- 
ticles, denticles in circles an- 
terior to stomatal suture small 
and of equal size 13 

- Stoma with 3-4 circles of den- 
ticles, denticles in circles an- 



262 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tenor to stomatal suture equal 
or unequal in size 15 

13(12). Excretory pore posterior to ce- 
phalic groove; tail short (2 anal 
diameters) . . P. combesi, n. comb. 
- Excretory pore anterior to ce- 
phalic groove and setae; tail 
longer (4 anal diameters or 
more) 14 

14(13). Stoma with 5 circles of denti- 
cles; spicules curved, tip re- 
curved with terminal barb; gu- 
bernaculum present; female 
cephalic setae 45% and amphid 

50% of head diameter 

P. parafloridensis, n. sp. 

- Stoma with 6 circles of denti- 
cles; spicules straight, tip with- 
out terminal barb; gubernac- 
ulum absent; female cephalic 
setae 35% and amphid 40% of 
head diameter . . . P. alima, n. sp. 

15(12). Tail 14 anal diameters long . . 

.... P. micoletzkyii Filipjev, 1946 

- Tail less than 10 anal diame- 
ters long 16 

16(15). Spicules without terminal 

barb; gubernacular apophysis 

bent dorsally at midpoint; male 

post-anal papillae absent . . . 

P. americana, n. sp. 

- Spicules with terminal barb; 
gubernacular apophysis not 
bent dorsally at midpoint; male 
post-anal papillae present or 
absent 17 

17(16). Male post-anal papillae ab- 
sent; spicules with broad, cup- 
shaped tip with large barb . . 
.... P. floridensis Keppner, 1986 

- Male post-anal papillae pres- 
ent; spicules with narrow tip 
and small barb 18 

18(17). Male post-anal papillae about 
l h distance from anus to tail 
tip; gubernacular apophysis 

short, directed dorsally 

. . P. scillionensis Warwick, 1977 



- Male post-anal papillae im- 
mediately post-anal; guber- 
nacular apophysis long, di- 
rected postero-dorsally 

P. pugetensis Wieser, 1959 

Acknowledgments 

Sincere appreciation is expressed to Dr. 
W. Duane Hope, National Museum of Nat- 
ural History, for his review of the manu- 
script, examination of the specimens, and 
continued encouragement. Appreciation is 
also expressed to Dr. Armen C. Tarjan, 
University of Florida, for his review of the 
manuscript and his encouragement. Mr. and 
Mrs. William Fable, National Marine Fish- 
eries Service, collected the sediment sam- 
ples from the Gulf of Mexico. 

Literature Cited 

Blome, D. 1974. Zur Systematik von Nematoden aus 
dem Sandstrand der Nordseeinsel Sylt.— Mik- 
rofauna Meeresboden 33:1-25. 

Chitwood, B. G. 1960. A preliminary contribution 
on the marine nemas (Adenophorea) of North- 
ern California.— Transactions of the American 
Microscopical Society 79:347-384. 

Gerlach, S. A. 1952. Nematoden aus dem Kiisten- 
grundwasser. — Abhandlungen der Akademie der 
Wissenschaften und der Literature, Mainz 6:315- 
372. 

Hopper, B. E. 1 963. Marine nematodes from the coast 
line of the Gulf of Mexico III. Additional species 
from Gulf Shores, Alabama.— Canadian Jour- 
nal of Zoology 4:841-863. 

. 1970. Free-living marine nematodes from 

Biscayne Bay, Florida, III. Eurystominidae: 
Pareurystomina bissonettei sp. n. from Biscayne 
Bay and other locations.— Proceedings of the 
Helminthological Society of Washington 37:175- 
178. 

Inglis, W. G. 1962. Marine nematodes from Banyuls- 
sur-Mer: With a review of the genus Eurysto- 
mina. — Bulletin of the British Museum (Natural 
History) Zoology 10:531-552. 

Lambshead, P. J. D., & H. M. Piatt. 1979. Batheu- 
rystomina, a new genus of free-living marine 
nematodes (Enchelidiidae) from the Rockall 
Trough.— Cahiers de Biologie Marine 20:371- 
380. 

Luc, M., & L. A. P. De Coninck. 1959. Nematodes 
libres marins de la region de Roscoff. —Archives 



VOLUME 102, NUMBER 1 



263 



de Zoologie Experimentale et Generate 98: 103- 
165. 

Wieser. W. 1953. Free-living marine nematodes. I. 
Enoploidea. — Acta Universitatis Lundensis. 
Medica, Mathematica. Scientiae Rerum Natur- 
alium(N. F. 2)49:1-155. 

. 1959. Free-living nematodes and other small 

invertebrates of Puget Sound beaches. Univer- 
sity of Washington Press, Seattle. Wash. 1 79 pp. 

. & B. E. Hopper. 1967. Marine nematodes of 

the east coast of North America. I. Florida.— 



Bulletin of the Museum of Comparative Zool- 
ogy, Harvard University 135:239-344. 
Yeates, G. W. 1967. Studies on nematodes from dune 
sands 3. Oncholaimidae. Ironidae. Alaimi- 
dae.— New Zealand Journal of Science 10:299- 
321. 

306 Hibiscus Avenue, Panama City 
Beach, Honda 32407. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 264-271 

NEOTROPICAL MONOGENEA. 14. REVISION OF 

CALLORHYNCHOCOTYLE SURIANO AND INCORVAIA, 

1982 (HEXABOTHRIIDAE) WITH THE 

DESCRIPTION OF C. AMATOI 

Walter A. Boeger, Delane C. Kritsky, and Joaber Pereira, Jr. 

Abstract. —The generic diagnosis of Callorhynchocotyle Suriano & Incorvaia, 
1982, is emended based on new information obtained from specimens of the 
type species, C. marplatensis, collected from Callorhinchus callorynchus from 
the coasts of Uruguay and Argentina. Callorhynchocotyle marplatensis is re- 
described; Squalonchocotyle callorhynchi is redescribed and transferred to Cal- 
lorhynchocotyle; and C amatoi is proposed for hexabothriids reported previ- 
ously from Callorhinchus milii by Manter (1955) and Dillon & Hargis (1968). 



All known Hexabothriidae are parasitic 
on gills of elasmobranch or holocephalan 
fishes (Class Chondrichthyes). Approxi- 
mately 60 species are described from elas- 
mobranchs; two are known from holo- 
cephalans. Manter (1955) described 
Squalonchocotyle callorhynchi from the 
holocephalans, Callorhinchus capensis Du- 
meril and C milii Bory, from marine waters 
off South Africa and New Zealand, respec- 
tively. Dillon & Hargis (1968) redescribed 
this species as Erpocotyle callorhynchi 
(Manter, 1955) Yamaguti, 1963, based on 
specimens from C milii in New Zealand 
waters. Lebedev & Parukhin (1969) report- 
ed the parasite under the latter name from 
C. capensis in Wallfish Bay (southwestern 
Africa); and in 1970, Kuznetsova recorded 
S. callorhynchi from Callorhinchus antarc- 
ticus Fleming (=C callorynchus) from the 
Patagonian Shelf off South America. Suri- 
ano and Incorvaia (1982) proposed Callo- 
rhynchocotyle (Hexabothriidae) for their new 
species, C marplatensis, from Callorhin- 
chus callorynchus (Linnaeus) off Argentina 
(38°S, 57°W). In the present study, Callo- 
rhynchocotyle Suriano & Incorvaia, 1982, 
is revised; C marplatensis is redescribed; 
S. callorhynchi is redescribed and trans- 
ferred to Callorhynchocotyle; and C. ama- 
toi, n. sp., is proposed for hexabothriids re- 



ported from Callorhinchus milii by Manter 
(1955) and Dillon & Hargis (1968). 

Materials and Methods 

Hosts {Callorhinchus callorynchus) were 
collected by trawl off the coasts of Uruguay 
and Argentina during May 1985 and June 
1986. Methods of parasite collection, prep- 
aration, measurement, and illustration are 
as described by Kritsky et al. (1986). Mea- 
surements are in micrometers; averages are 
followed by ranges in parentheses. Mea- 
surements of curved structures represent 
straight-line distances between extreme 
points; see Fig. 4 for those obtained from 
each sclerite. Vouchers (specimens collected 
during the present study) are deposited in 
helminthological collections of the National 
Museum of Natural History (USNM), which 
is housed at the USDA in Beltsville, Mary- 
land; the University of Nebraska State Mu- 
seum (HWML), Lincoln, Nebraska; the 
Fundacao Instituto Oswaldo Cruz (FIOC), 
Rio de Janeiro, Brazil; and the Museo de 
La Plata (MP), La Plata, Argentina. 

Callorhynchocotyle 
Suriano & Incorvaia, 1982 

Emended diagnosis. —Hexabothriidae. 
Body elongate. Tegument thin, smooth. 



VOLUME 102, NUMBER 1 



265 



Caeca confluent in peduncle posterior to 
testes. Testes numerous, irregular; vas def- 
erens sinuous, surrounded by small gland 
cells along most of its length, with small 
loop proximal to entrance into cirrus. Cirrus 
comprising two parts: distal part ovate, bul- 
bous; proximal part a thick-walled nonex- 
panded tube. Genital pore at level of gut 
bifurcation. Ovary lobate anteriorly, coiled 
posteriorly; oviduct originating from pos- 
terior end of ovary; seminal receptacle sac- 
like, reduced; ootype smooth; uterus dorsal 
to ovary, ventral to vas deferens. Vagina 
parallel, comprising two portions; distal 
portion glandular, proximal portion deli- 
cate. Vaginal pores on ventral surface about 
midway between body midline and lateral 
margin on each side of cirrus. Vitellaria 
comprising 2 bilateral bands extending from 
level of vaginal pores into peduncle; vitel- 
line commissure, genitointestinal canal dor- 
sal to ovary. Eggs with two polar filaments, 
forming continuous chain by united fila- 
ments. Excretory pores marginal at level of 
genital pore. Haptor asymmetrical; with 2 
small, 4 large sucker-sclerite complexes; 
small complexes lying on same side of lon- 
gitudinal axis of haptor; appendix originat- 
ing from dorsal haptoral surface lateral to 
body midline, armed with 2 anchors, 2 ter- 
minal suckers. Parasites of Callorhinchidae. 

Type species. — Callorhynchocotyle mar- 
plat ensis Suriano and Incorvaia, 1982, from 
Callorhinchus callorynchus. 

Other species. — Callorhynchocotyle cal- 
lorhynchi (Manter, 1955), n. comb., from 
Callorhinchus capensis; Callorhynchocotyle 
amatoi, n. sp., from Callorhinchus milii. 

Callorhynchocotyle marplatensis 

Suriano & Incorvaia, 1982 

Figs. 1-9, 10a, 11a, 12a 

Synonyms. — Callorhynchocotyle callo- 
rhynchi Suriano & Incorvaia, 1982; C. cal- 
lorhynchy Suriano & Incorvaia, 1982. 

Host. — Callorhinchus callorynchus (Lin- 
naeus). 



Distributions.— Coasts of Uruguay, Ar- 
gentina (holotype). 

Specimens studied. —Holotype, MP 12; 
17 vouchers, USNM 80279, HWML 20705, 
20706, FIOC 32.442, MP 1613 D. 

Description (measurements in Table 1).— 
Weak genital sucker, 94-1 10 testes. Papillae 
lacking on rim and inner wall of oral, hap- 
toral suckers. Sucker sclerite 2, 3 with elon- 
gate point, evenly curved shaft and point. 
Anchor with wide base, short roots, medi- 
ally curved shaft, short point. 

Remarks.— Suriano & Incorvaia (1982) 
treat this species under three names: C 
marplatensis, C callorhynchi, and C cal- 
lorhynchy. The latter two apparently rep- 
resent typographic and/or editing errors. It 
is apparent that these authors were not re- 
ferring to other species of Callorhynchoco- 
tyle since they consider the genus monotyp- 
ic. Clearly, the authors intended the name 
of their new species to be C marplatensis 
which is considered herein as the valid name 
of the taxon. 

Based on the holotype and voucher spec- 
imens, it is evident that Suriano & Incorvaia 
(1 982) confused polarity of the dorso ventral 
axis in their specimens. This is supported 
by their illustration of the holotype pre- 
sented as a dorsal rather than a ventral view 
as the specimen is mounted. Consequently, 
the sinistrodextral orientations of features 
of the parasite are reversed in the original 
description. Also contrary to the original 
description, the haptoral appendix origi- 
nates from the dorsal surface of the haptor, 
and the genitointestinal canal, uterus and 
vitelline commissure are dorsal to the ovary. 

The holotype is a damaged, excessively 
flattened, and contracted specimen. One of 
the haptoral suckers and its sclerite are torn 
away (not shown in fig. 1 of Suriano & In- 
corvaia 1982). Apparently, these original 
authors added the missing sucker to their 
drawing of the holotype based on the pres- 
ence of six suckers in other specimens of 
their collection. This, however, resulted in 
their apparent confusion of a damaged mus- 



266 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1 . — Measurements (in micrometers) of Callorhynchocotyle species. 





C. callorhynchi 
(n = 2) 


C. marplatensis 
(n= 17) 


C. amatoi 
(n = 8) 


Body length 
Body width 


6052(4995-7109) 
845 (669-1021) 


9989(7200-12,430) 
939(649-1217) 


7091 (5329-8474) 
989(706-1316) 


Haptor 








Length 
Width 


2007 


2719(2051-3324) 
1596(1331-1918) 


1959(1451-2482) 
1201 (755-1639) 


Oral sucker 








Width 


284 (248-322) 


302(231-362) 


252(191-356) 


Pharynx 








Length 
Width 


72 

64 (56-72) 


71 (58-83) 
71 (60-78) 


79(69-91) 
70 (62-89) 


Genital sucker 








Diameter 


— 


205(153-262) 


— 


Egg 








Length 
Width 


152(136-168) 
58 (53-62) 


145(125-193) 
59 (52-66) 


156(144-169) 
54 (48-60) 


Ovary 








Length 


639(459-818) 


1147(816-1505) 


859(651-971) 


Distal vagina 


276 


180(136-203) 


350 (269^132) 


Appendix 








Length 
Width 


2104(1844-2363) 
997(972-1021) 


1397(1275-1637) 
256 (149-350) 


1108(674-1429) 
850(541-1206) 


Appendix sucker 








Length 
Width 


289 
109 


181 (164-204) 
195(154-190) 


272 (236-320) 
144(102-185) 


Anchor 








Length 
Base width 


194(186-203) 


190(144-210) 
87 (72-98) 


182(172-195) 
85 (83-87) 


Diameter of haptoral sucker 








Pair 1 
Pair 2 
Pair 3 


274 (258-290) 

362 

319 


305 (243-346) 
365 (270-419) 
345 (267-393) 


267(188-362) 
332 (232^35) 
310(217^115) 


Sclerite #1 








Total 
Shaft 
Point 


283(281-285) 
31-32 
42 (39^5) 


353 (289-385) 
33 (27^14) 
50 (42-57) 


297 (239-355) 
20(18-22) 
37 (36-39) 


Sclerite #2 








Total 
Shaft 
Point 


362 

88 

106 


460 (362^99) 

76 (59-92) 

117(95-141) 


412(347^152) 
54 (46-70) 
83 (70-98) 


Sclerite #3 








Total 

Shaft 
Point 


351 
84 (83-86) 
123-124 


454(382-512) 

80 (65-92) 

129(99-145) 


408 (332^169) 
56 (45-66) 
92 (86-98) 



VOLUME 102, NUMBER 1 



267 




Figs. 1-6. Callorhynchocotyle marplatensis: 1, Whole mount (ventral); 2, Sucker-sclerite complex; 3, Sclerite 
of complex 1; 4, Sclerite of complex 2; 5, Sclerite of complex 3; 6, Anchor. All figures are reproduced to respective 
scales except Figs. 3, 4, 5 are drawn to the 250-micrometer scale. Circled numbers identify sucker-sclerite pairs 
in the haptor. Fig. 4 shows measurements taken from each sclerite (Table 1): a = total length; b = shaft length; 
c = point length. 



cle as a haptoral extension of the gut. In the 
holotype the gut is not visible in the haptor, 
although the structure depicted as gut (the 
muscle which apparently served the lost 
sucker) is evident. Our specimens show that 



the esophagus has lateral diverticulae, the 
caeca are confluent posterior to the testes, 
and the intestine extends into the haptor 
with branches, one of which occurs in the 
haptoral appendix. 



268 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Figs. 7-9. Callorhynchocotyle marplatensis: 7, Ventral view of anterior end; 8, Egg chain from uterus; 9, 
Female reproductive system (ventral). Figures are to respective 200-micrometer scales. 



Callorhynchocotyle amatoi, new species 
Figs. 10c, lie, 12c 

Synonyms.— Squalonchocotyle callo- 
rhynchi Manter, 1955 (part); Erpocotyle 
cat 'lor hy nchi (Manter, 1955)Yamaguti, 1963 
(part). 

Host. — Callorhinchus milii Bory. 

Type locality. — Coast of New Zealand. 

Specimens studied. — Holotype, USNM 
37448; seven paratypes, USNM 71197, 
HWML 1442. 

Description (measurements in Table 1). — 
Genital sucker absent; 94-120 testes. Pa- 
pillae present on rim and inner wall of oral, 
haptoral suckers. Sucker sclerite 2, 3 with 
short point; indistinct angle formed at union 
of shaft and point of sclerite 3; sclerite 1 
with delicate point, short shaft. Anchor with 
wide base, short roots, medially curved shaft, 
short point. 

Etymology. — This species is named for 
Dr. Jose Felipe R. Amato, Universidade 
Federal Rural do Rio de Janeiro, in recog- 
nition of his contributions in marine par- 
asitology. 

Remarks.— Our study of the type speci- 
mens of Squalonchocotyle callorhynchi 



Manter, 1955, and specimens identified as 
Erpocotyle callorhynchi (Manter, 1955) by 
Dillon & Hargis (1968) revealed that those 
from Callorhinchus milii from New Zea- 
land represent a species distinct from that 
occurring on C. capensis. Although Dillon 
& Hargis (1968) recognized the morpholog- 
ic differences between these specimens, they 
did not question their conspecificity. 

Since the holotype of Squalonchocotyle 
callorhynchi is a specimen collected from 
the gills of Callorhinchus capensis captured 
off South Africa, Callorhynchocotyle amatoi 
is proposed for those specimens from Cal- 
lorhinchus milii from New Zealand. Cal- 
lorhynchocotyle amatoi differs from C. cal- 
lorhynchi and C. marplatensis by having 
shorter shafts and points of all sucker scler- 
ites (Fig. 1 3) and an indistinct angle between 
the point and shaft of sclerite 3 (Fig. 12c). 



Callorhynchocotyle callorhynchi 

(Manter, 1955), new combination 

Figs. 10b, lib, 12b 

Synonyms.— Squalonchocotyle callo- 
rhynchi Manter, 1955 (part); Erpocotyle cal- 



VOLUME 102, NUMBER 1 



269 




Figs. 10-12. Sclerites of Callorhynchocotyle spp.: 10, Sclerite of complex 1; 11, Sclerite of complex 2; 12, 
Sclerite of complex 3. a = C. marplatensis; b = C. callorhynchi; c = C. amatoi. All figures are drawn to the 
same scale. 



lorhynchi (Manter, 1955) Yamaguti, 1963 
(part). 

Host. — Callorhinchus capensis Dumeril. 

Type locality.— Coast of South Africa. 

Specimens studied. — Holotype, paratype, 
USNM 37447. 

Description (measurements in Table 1).— 
Genital sucker absent, 96-98 testes. Pa- 
pillae present on rim and inner wall of oral, 
haptoral suckers. Sucker sclerite 1,2,3 with 
short point, moderate shaft, evenly curved 
shaft and point. Anchor with wide base, 
short roots, medially curved shaft, short 
point. 

Remarks. — Callorhynchocotyle callo- 
rhynchi is most similar to C. marplatensis, 
from which it differs by possessing papillae 
on the rim and inner walls of the oral and 
haptoral suckers, a comparatively long dis- 
tal portion of the vaginae, absence of a gen- 
ital sucker, and by the comparative mor- 
phology of the shaft and point of the sucker 
sclerites (Figs. 10-12). 

Discussion 

Callorhynchocotyle was proposed by 
Suriano & Incorvaia (1982) to accommo- 
date C. marplatensis. Among characters 
used to define the genus was the presence 
of marginal vaginal pores. Examination of 
the holotype (MP 1 2) of C. marplatensis and 
vouchers collected during the present study 



shows that the vaginae actually open sub- 
marginally as they do in members of all 
other hexabothriid genera. Apparently Suri- 
ano and Incorvaia (1982) confused the lat- 
eral excretory pores as the vaginal apertures. 
Other characters used by Suriano and In- 
corvaia (1982) for diagnosis of the genus 
include: 1) an asymmetric haptor with 6 
suckers armed with sclerites of different 
sizes, 2) a sac-like seminal receptacle, 3) par- 
allel bilateral vaginae, 4) an unarmed cirrus, 
5) a smooth ootype, and 6) the absence of 
a glandular region of the cirrus. Of these, 
only the absence of a glandular region of the 
cirrus is unique for the genus. In Callo- 



H 160 

E 

3. 140 
+* 120- 

O 100-1 
<4-h 80 

o 

5 60 1 

c 

CD 
— 



20 + 






rt» oo • • 



o o *0 



>e°o 



g 



° • C marplatensis 

a * C amatoi 

♦ o C callorhynchi 



15 30 45 60 75 90 105 

Length of shaft (yum) 13 

Fig. 13. Scatter diagram of length of point versus 
length of shaft of the sclerite for three species of Callo- 
rhynchocotyle. The left column of symbols of the leg- 
end refers to sclerite 1 of respective species; the second 
column refers to sclerites 2, 3 of the respective species. 



270 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



rhynchocotyle, the glandular region of the 
cirrus is apparently replaced by an extensive 
glandular segment of the vas deferens. These 
characters may represent an example of 
compensatory change (evolution) as defined 
by Brooks and Wiley (1986). 

Other hexabothriid genera with asym- 
metric haptors include Heteronchocotyle 
Brooks, 1934, Paraheteronchocotyle Mayes, 
Brooks and Thorson, 1981, Epicotyle Euzet 
& Maillard, 1974, Rhinobatonchocotyle 
Doran, 1953, and Neonchocotyle Ktari & 
Maillard, 1972. The haptor of Callorhyn- 
chocotyle most closely resembles that of 
Neonchocotyle in that the asymmetry re- 
sults from the apparent migration of one 
sucker-sclerite complex to the opposite side 
of the haptor. Further, the appendix does 
not originate from the midline which adds 
to haptoral asymmetry. 

In addition to the above, Callorhyncho- 
cotyle is distinguished from other hexa- 
bothriid genera by the combined presence 
of the following characters: presence of a 
bulbous distal cirrus, a dorsal instead of 
terminal haptoral appendix, and a terminal 
glandulomuscular portion of the vagina. 

Hargis (1955) reported that the genitoin- 
testinal canal of Heteronchocotyle leucas 
Hargis, 1955 (Hexabothriidae), apparently 
joined the left intestinal caecum and sug- 
gested that this condition was rare among 
the higher Monogenea. In individual spec- 
imens of Callorhynchocotyle, the genitoin- 
testinal canal was found to be united to the 
intestinal caecum which corresponded to the 
respective position of the ovary, i.e., if the 
ovary occurred to the right of the body mid- 
line (Fig. 9) the genitointestinal canal united 
with the right caecum and conversely with 
the left caecum if the ovary occurred to the 
left of the body midline (Fig. 1). Similarly, 
the haptoral appendix and the migrated 
sucker-sclerite complex occur either to the 
left or right of the longitudinal haptoral axis. 
However, there was no correlation between 
the asymmetry of the haptor and the loca- 
tion of the ovary. The sinistral or dextral 



position of the ovary and genitointestinal 
canal and the position of the appendix and 
sucker-sclerite in the haptor are examples 
of intraspecific variation and do not con- 
stitute generic characteristics for the Hexa- 
bothriidae. 

Acknowledgments 

The authors wish to thank Drs. O. M. 
Blanco (MP), J. R. Lichtenfels (USDA), and 
M. Pritchard (HWML) for allowing us to 
examine specimens in their care. The Con- 
selho Nacional de Desenvolvimento Cien- 
tifico e Tecnologico provided a study grant 
(20.01 15/84) to WAB. 

Literature Cited 

Brooks, D. R., & E. O. Wiley. 1986. Evolution as 
entropy. Toward a unified theory of biology. 
University of Chicago Press, Chicago, Illinois. 
335 pp. 

Dillon, W. A., & W. J. Hargis. 1968. Monogenetic 
trematodes from the southern Pacific Ocean. Part 
IV. Polyopisthocotyleids from New Zealand 
fishes: The families Mazocraeidae, Diclidopho- 
ridae and Hexabothriidae.— Proceedings of the 
Biological Society of Washington 81:351-366. 

Hargis, W. J., Jr. 1955. Monogenetic trematodes of 
Gulf of Mexico fishes. Part VI. The superfam- 
ilies Polystomatoidea Price, 1936 and Dicli- 
dophoroidea Price, 1936.— Transactions of the 
American Microscopical Society 74:361-377. 

Kritsky, D. C, V. E. Thatcher & W. A. Boeger. 1986. 
Neotropical Monogenea. 8. Revision of Uro- 
cleidoides (Dactylogyridae, Ancyrocephali- 
nae).— Proceedings of the Helminthological So- 
ciety of Washington 53:1-37. 

Kuznetsova, I. G. 1970. Parasitofauna of cartilagi- 
nous fishes of the Patagonian Shelf. Pp. 52-54 
in Issues on marine parasitology, materials from 
the 1st annual All-Union Symposium on Par- 
asites and Diseases of Marine Life, Sevastopol. 
Publ. Naukova Dumka, Kiev. 

Lebedev, B. I., & A. M. Parukhin. 1969. Monogenea 
of some fish from Wallfish Bay (south-western 
Africa). — Gidrobiologicheskii Zhurnal 5:70-8 1 . 

Manter, H. W. 1955. Two new monogenetic tre- 
matodes from elephant fishes (Callorhynchus) 
from South Africa and New Zealand. Pp. 21 1- 
220 in Essays in the natural sciences in honor 
of Captain Allan Hancock. University of South- 
ern California Press, Los Angeles. 

Suriano, D. M., & I. S. Incorvaia. 1982. Sistematica 



VOLUME 102, NUMBER 1 



271 



y biologia de Callorhynchocotyle marplatensis 
gen. et sp. n. (Monogenea: Polyopisthocotylea) 
parasita de las branquias de Callorhynchus cal- 
lorhynchus (Linne, 1758)Garman. 1904 (Pisces: 
Holocephali) de la region costera de Mar del 
Plata. — Comunicaciones del Museo Argentino 
de Ciencias Naturales Bernardino Rivadavia e 
Instituto Nacional de Investigacion de las Cien- 
cias Naturales 2:19-32. 

(WAB) Department of Biological Sci- 
ences, Idaho State University, Pocatello, 



Idaho 83209, and Instituto Nacional de 
Pesquisas da Amazonia, Manaus, Amazon- 
as, Brazil; (DCK) Department of Allied 
Health Professions and Idaho Museum of 
Natural History, Idaho State University, 
Pocatello, Idaho 83209; (JP) Departamento 
de Ciencias Morfobiologicas, Fundagao 
Universidade do Rio Grande, Rio Grande, 
Brazil. 



PROC. BIOL. SOC. WASH. 

102(1), 1989, pp. 272-279 

PHYTOPLANKTON COMPOSITION IN A 
BORROW PIT LAKE IN VIRGINIA 

Seba B. Sheavly and Harold G. Marshall 

Abstract. — The phytoplankton assemblages in Lake Trashmore, Virginia, a 
borrow pit, were dominated by centric diatoms and cyanobacteria, with sea- 
sonal pulses of cryptomonads, euglenoids, and chlorophyceans. Ninety species 
were identified and their abundance levels noted for a 12 -month period. 



Borrow pit lakes are generally associated 
with highway construction and residential 
landscaping. In contrast, this borrow pit lake 
was formed as part of an above-ground 
landfill operation. Alternate layers of soil 
and refuse were laid down to eventually form 
a mound that was capped with soil, then 
landscaped. The pit formed in this process 
was gradually filled with water by 1 97 1 . The 
lake is located in Virginia Beach, Virginia, 
where it is now a part of a recreational park 
complex. The lake has a mean depth of 4 
m, a maximum depth of 7.3 m, a surface 
area of 21 ha, and a volume of 8.4 x 10 5 
m 3 (Virginia State Water Control Board 
1982). The surrounding area is highly de- 
veloped. The lake is flanked to the north 
and south by major highways and is adja- 
cent to the disposal mound (Mount Trash- 
more) on the west, with residential devel- 
opment along the eastern section. 

The purpose of this study was to identify 
the seasonal phytoplankton composition and 
concentrations of this lake and to make 
comparisons with a phytoplankton survey 
of the lake made over a decade ago (Cocke 
1973). 

Other phytoplankton studies in Virginia 
have concentrated on the two natural lakes 
in the state, Lake Drummond and Moun- 
tain Lake (Simmons & Neff 1974, Marshall 
1979). In addition, the National Eutrophi- 
cation Survey (Anonymous 1972) included 
eight other Virginia lakes and reservoirs that 



were of greater size than Lake Trashmore, 
and all were found to be eutrophic. Phyto- 
plankton were included in a limited water 
quality study during the early formation of 
Lake Trashmore by Beck (1973). He indi- 
cated an abundance of diatoms, chlorophy- 
ceans, cyanobacteria and phytoflagellates. 
Later Cocke (1973) completed a one year 
study of the lake and reported 45 taxa, with 
chlorophyceans and diatoms dominant. 

In general, borrow pit lake studies are rare. 
In Nebraska, McCarraher et al. (1974) sur- 
veyed 4 1 borrow pit lakes in the Platte River 
Valley, with Adrian et al. (1 970) conducting 
a primary productivity study on one lake. 
Seven Illinois borrow pit ponds were stud- 
ied by Lipsey (1980). In all of these studies, 
the formation of borrow pits was associated 
with a major highway development. Dia- 
toms, cyanobacteria and chlorophyceans 
were generally the dominant forms in these 
ponds or lakes. 

Methods. —One collection station was es- 
tablished at the center and deepest (7.3 m) 
portion of the lake. Samples were collected 
monthly from the upper and lower 0.5 m 
of the euphotic zone from Mar 1985 to Feb 
1986. The lower sampling depth was deter- 
mined from transparency measurements 
with a Secchi disk (15 cm diam), according 
to Holmes (1970). A Kemmerer water bot- 
tle (2 liter) was used to obtain 500 ml water 
samples from both depths. These were pre- 
served immediately with Lugol's solution 



VOLUME 102, NUMBER 1 



273 



and returned to the laboratory, where a set- 
tling and siphoning procedure was followed 
to obtain a 40 ml concentrate. Aliquots were 
taken from this concentrate and placed in 
settling chambers for examination with a 
Zeiss inverted plankton microscope. A ran- 
dom field and minimum count procedure 
was followed at magnifications of 3 1 2 x and 
500 x for micro-, nano-, and picoplankton 
for a precision estimate of 85%. In addition, 
the net phytoplankton were counted by 
scanning the entire chamber at 125 x . 

Results. —Mean water temperatures 
ranged from 4.0°C in January to 27.3°C in 
August, with only small differences gener- 
ally found with depth in the euphotic zone. 
The mean monthly pH value was 7.52, with 
a range from 6.5 (Dec) to 8.6 (Sep). Secchi 
disk readings ranged from 0.47 m (Mar) to 
1.18 m (Aug), averaging 0.84 m. 

Ninety phytoplankton species were iden- 
tified in this study (Table 1), including 32 
Chlorophyceae, 28 Bacillariophyceae, 16 
Cyanobacteria, 8 Euglenophyceae, 5 Di- 
nophyceae, and 1 Cryptophyceae. Uniden- 
tified picoplankton and nanoplankton cells 
were counted by size groups <3, 3-5 and 
5-10 jLtm. They were present throughout the 
year, with lowest concentrations in May (3.7 
x 10 6 cells/liter), gradually increasing to a 
December high (61.5 x 10 6 cells/liter). The 
majority of these cells were <3 ^m in size 
and under epifluorescence microscopy 
proved to be cyanobacteria. In contrast, 
larger and identifiable cyanobacteria had one 
major pulse in summer and early fall, reach- 
ing 37.0 x 10 6 cells/liter in September. Eu- 
glenoids and dinoflagellates also had high 
summer concentrations. In contrast, the 
cryptomonads were in lowest concentrations 
in summer and early fall and became more 
abundant during winter and spring. The 
diatoms had low concentrations in spring 
and summer, gradually increased into fall 
and early winter and peaked in December 
(9.4 x 10 6 cells/liter). The dominant species 
were Cyclotella spp. and Melosira spp., with 
representatives from these genera most 



common in the unidentified category of cen- 
tric diatoms <20 nm size. 

Discussion. — In comparison to early data 
on phytoplankton populations in Lake 
Trashmore (Beck 1973, Cocke 1973), there 
has been a shift in composition and domi- 
nant species. Cocke identified 45 species in 
a seven-month study (Aug-Feb), with the 
two dominant groups: pennate diatoms (e.g., 
Pleurosigma normanii, P. strigosum) and 
chlorophyceans {Pediastrum simplex) the 
major species. He found cyanobacteria 
(blue-green algae) to be common, but not 
abundant. Other dominant species included 
the desmids Closterium lunuae and Cos- 
marium circulare, the diatoms Fragilaria 
crotonensis, Melosira spp., Navicula spp., 
and the dinoflagellate Gymnodinium sim- 
plex. Secchi disk readings at this time ranged 
between 24.0 and 54.0 cm, averaging 40.9 
cm. The range of the surface pH values was 
6.6 to 7.1, averaging 6.8. The present study 
indicated a more basic pH mean of 7.52, 
ranging between 6.5 and 8.6. However, the 
Secchi disk readings were higher, averaging 
84 cm, with readings between 47 cm (Mar) 
and 1 1 8 cm (Aug). 

In addition to an increased diversity of 
species, there has been a change over the 
past decade in the phytoplankton popula- 
tions in Lake Trashmore. The transition has 
been from a dominance of pennate diatoms, 
chlorophyceans, and a filamentous-coccoid 
assemblage of cyanobacteria to the current 
status, where cyanobacteria, centric dia- 
toms (e.g., Cyclotella spp., Melosira spp.), 
plus a seasonal abundance of cryptomo- 
nads, euglenoids, and chlorophyceans are 
dominant. It is impossible to evaluate the 
significance of the high picoplankton and 
nanoplankton (< 10 /xm) concentrations be- 
cause Cocke's collection procedure by tow 
net would not have collected many of these 
cells. However, these changes in phyto- 
plankton composition and abundance are 
assumed to be in association with the chang- 
ing and advancing eutrophic state of the lake 
over the past decade. 



274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Table 1.— Mean monthly abundance for each identified taxon (cells/liter). 





Mar 


Apr 


May 


Jun 


Bacillariophyceae 










Achnanthes sp. 


47,268 


639 


5373 


757 


Amphora costata W. Smith 











213 


Amphora sp. 





21 








Biddulphia alternans (Bailey) Van Heurck 








5 





Cocconeis distans Gregory 


21 











Cyclotella meneghiniana Kiitzing 





77,867 


29,373 


18,131 


Cy clot el I a sp. 


273,843 


27,084 


28,213 


4578 


C. striata (Kiitzing) Grunow 








18,056 





Cylindrotheca closterium (Ehrenberg) 










Reimann & Lewin 


277 








53 


Cymbella sp. 


9515 


602 


123 


85 


Diploneis sp. 








16 


12 


Fragilaria sp. 











10,579 


Gomphonema sp. 








16 


53 


Gyrosigma sp. 











21 


Melosira distans (Ehrenberg) Kiitzing 


33,824 











M. granulata (Ehrenberg) Ralfs 


221,898 


9649 


299 


1664 


M. islandica Muller 


5195 





4557 


85 


M. moniliformis (Muller) Agardh 








32 





Melosira sp. 





37 


107 


427 


Navicula spp. 


9813 


37 


32 


117 


Nitzschia clausii Hantzsch 











64 


N. pungens Grunow 


43 








75 


N. seriata Cleve 


43 











Nitzschia sp. 


14,411 


16 





32 


Rhizosolenia eriensis H. L. Smith 














Synedra acus Kaiitzing 














Synedra spp. 





21 


85 


587 


Thalassionema nitzschioides (Grunow) 










Grunow & Hustedt 





16 





384 


centrics (unid.) < 20 jiim diam. 


285,376 


293,413 


178,303 


81,607 


centrics (unid.) 20-100 Aim diam. 


109,243 








14,374 


pennates (unid.) >20 /urn in length 


51,857 


56,426 


28,341 


22,072 


pennates (unid.) <20 nm in length 


26,076 


112 


101 


21 


subtotal 


1,088,703 


465,940 


294,032 


156,000 


Dinophyceae 










Ceratium hirundinella (O. F. Muller) 










Dujardin 








368 


19,040 


Glenodinium gymnodinium Penard 











43 


Glenodinium sp. 





3 








Gymnodinium sp. 











36,113 


Protoperidinium sp. 











96 


subtotal 





3 


368 


55,292 


Cyanobacteria 










Anabaena sp. 


725 


23,138 





576 


Chroococcus limnecticus Lemmerman 











477,533 


Dactylococcopsis raphidiodes Hansgirg 


2,607,829 


920,865 


478,484 


101,230 


Gomphosphaeria aponina Kutzing 


120,610 


152 


2294 


9028 


Johannesbaptistia pellucida (Dickie) 










Taylor & Drouet 














Lyngbya controta Lemmermann 















VOLUME 102, NUMBER 1 



275 



Table 1.— Continued. 



Jul 


Aug 


Sep 


Oct 


Nov 


Dec 


Jan 


Feb 


1984 














256 








469 


64 


107 














64 

















247 
























































18,057 


18,078 


36,113 





36,455 


180,907 


55,297 


483,016 


9028 


135,532 


270,850 


81,255 


81,255 


424,328 


40,626 


731,316 








22,571 














40,628 





32,154 


46,187 





128 


213 


208 


725 


43 


64 


277 


128 


128 


640 


32 


139 


43 























533 


853 


36,849 





341 


768 











64 





















































36,415 


1,363,278 


3,403,654 


137,677 


153,780 


28,557 


31,864 


248,919 


768,108 


806,756 


3,710,612 


343,064 


690,668 


320 


1472 


427 


198,623 






































9220 








40 














768 


939 


2965 


1493 


256 


1963 


64 


18,099 


9114 


4557 











256 
























































128 





555 




















117,432 


63,198 

















43 


875 


53,675 


128 


469 


700 


517 


789 


597 


51,789 


124,323 


49,878 


3029 


3212 


171 


299 


299 


18,505 


272,215 








180,799 








117,368 


153,482 


329,534 


135,425 


261,864 


1,426,466 


279,190 


1,923,035 


21 


21 


18,227 





213 


640 


38,369 


31,599 


22,571 


9028 


99,312 


54,170 


939 


232,435 


58,682 


90,282 


85 


40,734 


341 


85 


768 


1244 


8028 


533 


219,248 


617,507 


1,626,645 


1,325,748 


2,555,879 


9,569,435 


961,930 


4,164,983 


5589 


3392 


23,467 


13,227 


1408 


537 


16 





85 


1835 


1771 


981 














213 





1600 


18,185 














64 

















22,570 





43 























5994 


5227 


26,838 


32,393 


1408 


537 


22,586 





1,173,683 


9,619,689 


6,238,578 


1,760,525 


2987 


356,250 








537,186 


64,009 


643,401 


451,417 


504,161 


2,644,782 


205,091 


175,549 


94,947 





225,708 


370,162 


353,897 


839,628 


815,906 


1,047,287 





128 


100,784 


397,247 


91,862 


58,479 


21,479 


32,815 


341 


























794,493 


174,098 


64,350 















276 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1.— Continued. 



Mar 



Apr 



May 



Jun 



Merismopedia glauca (Ehrenberg) 

Naegeli 
Merismopedia punctata Meyen 
Merismopedia sp. 

Merismopedia tenuissima Lemmermann 
Microcystis aeruginosa Kutzing 
Nostoc commune Vaucher 
Oscillatoria limnetica Lemmermann 
Oscillatoria sp. 
Spirulina laxa G. M. Smith 
Spirulina subsalsa Oersted 
blue green spheres (unid) 
blue green trichomes (unid) 
subtotal 

Euglenophyceae 
Euglena acus Ehrenberg 
Euglena sp. 

Eutreptia lanowii Steuer 
Eutreptia viridis Perty 
Phacus longicaudus (Ehrenberg) Dujardin 
Phacus curvicaudus Swirenko 
Trachelomonas hispida (Perty) Stein 
Trachelomonas volvocina Ehrenberg 
subtotal 

Chlorophyceae 

A nkistrodesmus falcatus Beijerinck 
Ankistrodesmus fr actus (West & West) 

Brunnthaler 
Chlorella sp. 

Chlorella vulgaris Beijerinck 
Cosmarium botrytis Meneghini 
Crucigenia apiculata (Lemmermann) 

Schmidle 
Crucigenia fenestrata Schmidle 
Crucigenia quadrata Morren 
Crucigenia sp. 
Crucigenia tetrapedia (Kirchner) West 

& West 
Dictyosphaerium pulcellum Wood 
Euastrum denticulatum (Kirchner) Gay 
Franceia droescheri (Lemmermann) 
Kirchneriella contorta (Schmidle) Bohlin 
Lagerheimia ciliata (Lagerheim) Chodat 
Lagerheimia quadriseta Lemmermann 

(G. M. Smith) 
Micractinium pusillum Frensenius 
Oedogonium sp. 
Oocystis borgei Snow 
Pediastrum duplex Meyen 
Pediastrum simplex (Meyen) 

Lemmermann 
Scenedesmus armatus (Chodat) G. M. Smith 





















8038 


18,526 


























408,516 


2,471,603 


2,059,101 


2,719,878 


277 


15,738 


432 


779 

















32 


208 


4274 











53 











32 








54,168 





64 


1129 


6782 


23,010 


3,138,021 


3,432,657 


2,609,507 


3,354,919 























1821 


2257 


52,529 


22,591 


87,704 


85 





5 


43 















































76,608 


2342 


52,529 


22,591 


182,572 



156,672 



27,655 



31,316 



31,445 















9028 




















4104 


























92,219 








18,057 








9113 


744,539 








4514 





,831,853 


143,152 





207,908 


1365 


309,316 


143,839 


2,013,046 











21 














191,125 


293,977 


302,438 


283,177 












































421 





114,091 


129,214 


105,047 


250,682 














36,147 


421 


1259 


5845 


363 












VOLUME 102, NUMBER 1 277 

Table 1.— Continued. 



Jul 


Aug 


Sep 


Oct 


Nov 


Dec 


Jan 


Feb 





115,257 





341 

















171 


2123 


1024 


308,329 


704 




















288,907 

















15,924,935 


5,673,395 


870,724 


928 


18,056 





10,251,540 


7,031,505 


8,564,855 


9,331,530 


6,593,405 


4,709,592 


82,145 


251,908 


2,902,609 


6,414,631 


4,441,940 


5,254,490 


3,891,212 


2,146,677 


928,350 


853 





993,117 


672,611 


361,133 














22,912 


13,756 





18,057 





427 


32 








18,675 


85 





171 














9071 












































4514 


9028 


90,283 


18,057 


43 





48 


213 


14,987,732 


25,083,530 


37,079,401 


23,701,728 


1,290,568 


10,757,467 


2,071,107 


150,862 





192 


299 

















39,953 


12,634 


186,958 


55,493 


768 


341 








126,397 


54,234 


288,907 


180,567 


63,284 


208,333 


80 


4099 





21 


























1664 





128 














256 


77,091 


95,446 


1323 





32 





448 


832 


832 


46,282 


118,307 


2603 


16,992 


49,763 


32,218 








27,085 


72,227 


54,170 


16,928 


63,433 


198,568 


67,785 


555,751 


404,873 


256,037 


265,447 


34,032 


154,186 



66,646 9600 46,780 317,562 72,133 8021 437 405 


















112,487 


6485 


2112 
























































36,113 





384 























583,616 


1,529,376 


252,869 


597 














63,198 





341 


77,082 


72,376 


162,851 


110,900 


580,928 


361,133 


1,011,507 


27,575 


1280 


18,057 




















235 


341 





144,453 


306,963 


184,151 


1,594,264 


188,460 


277,399 


























128 


9178 


135,425 


597 


299 


1024 








18,057 











384 











18,057 








4949 


383,115 


1,119,844 


125,264 


112,876 





22,571 




















63,198 


58,684 


18,057 


171 





341 

















4779 

















171 




















176,053 


361,389 


487,701 


397,247 


297,935 


2,058,442 


164,761 


171,773 


























10,197 


10,603 


63,957 


88,832 


153,583 


55,808 


599 


1792 



























278 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1. — Continued. 



Mar 



Apr 



May 



Jun 



Scenedesrnus bijugus Turpin Lager. 
Scenedesmus dimorphus (Turpin) Kutzing 
Scenedesrnus quadricaudus (Turpin) 

Brebisson 
Scenedesmus sp. 
Selenastrum gracile Reinsch 
Staurastrum americanum (West & West) G. M. 

Smith 
Staurastrum leptocladum var. insigne West 

& West 
Staurastrum paradoxum Meyen 
Staurastrum sp. 

Tetraedron minimum (Braun) Hansgirg 
Chlorophyceans (unid.) 
subtotal 

Cryptophyceae 
Cryptomonas sp. 
subtotal 

Other taxa 

micro-phytoflagellates <10 nm 
micro-phytoflagellates >10 /j.m 
small green spheres ( < 3 pm) 
small green spheres (3-5 iim) 
small green spheres (5-10 ^ni) 
subtotal 

Total 



20,096 


9625 

85 






2343 



40,405 

11,285 










27,276 
4530 



11 



37,685 
85 

196,406 
18,398 













80 


3008 








11 


341 





37 


43 


117 


64 


580 


21 


102,122 


171 


16,928 


136,549 


650,038 


2,462,904 


975,313 


766,468 


4,567,024 


825,700 


1,816,905 


1,271,836 


554,242 


825,700 


1,816,905 


1,271,836 


554,242 


47,637 








175,407 








8215 


43,810 


18,320,895 


3,792,191 


3,348,778 


9,835,356 


4,786,425 


470,241 


262,864 


675,405 


997,156 


131,192 


84,883 


115,001 


24,152,113 


4,393,624 


3,704,740 


10,844,979 


31,669,783 


11,136,971 


869,542 


19,715,028 



Literature Cited 

Adrian, G. L., C. Throckmorton, & R. McDonald. 
1970. A study of primary production in a Ne- 
braska Interstate 80 lake.— Transactions of the 
Kansas Academy of Science 73(2):227-236. 

Anonymous. 1972. Water quality criteria. A report 
of the Committee on Water Quality Criteria.— 
National Academy of Science and National 
Academy of Engineering. Washington, D.C., 219 
pp. 

Beck, W. M. , Jr. 1973. Building an amphitheater and 
costing ramp of municipal solid waste. U.S. En- 
vironmental Protection Agency. SW-52d.of. Of- 
fice of Solid Waste Programs Vol. I and II. 
Washington, D.C., 265 pp. 

Cocke, G. R. 1973. A comparative limnological sur- 
vey of a brackish water lake and freshwater lake 
at Mount Trashmore, Virginia Beach, Virginia. 
Masters Thesis. Old Dominion University, 
Norfolk, Virginia, 120 pp. 



Holmes, R. W. 1970. The secchi disk in turbid coastal 
waters.— Limnology and Oceanography 15:688- 
694. 

Lipsey, L. L. 1980. Phytoplankton of selected borrow 
pit ponds in northern Illinois.— Ohio Journal of 
Science 80(3): 108-1 13. 

Marshall, H. G. 1979. Lake Drummond: With a dis- 
cussion regarding its plankton composition. Pp. 
169-182 in P. W. Kirk, ed., The Great Dismal 
Swamp. University Press, Charlottesville, Vir- 
ginia. 

McCarraher, D. B., R. McDonald, & G. Adrian. 1974. 
Some hydrobiological characteristics of Inter- 
state-80 highway lakes in Nebraska.— Trans- 
actions of the Kansas Academy of Science 77(2): 
93-102. 

Simmons, G. M., & S. E. Neff. 1974. Observations 
on limnetic carbon assimilation rates in Moun- 
tain Lake, Virginia during its thermal stratifi- 
cation periods.— Virginia Journal of Science 
24(4):206-211. 



VOLUME 102, NUMBER 1 



279 



Table 1.— Continued. 



Jul 


Aug 


Sep 


Oct 


Nov 


Dec 


Jan 


Feb 


388,218 
128 


173,006 
341 


439,333 

72,483 


507,080 
683 


257,401 



15,104 

1024 


299 



1365 




193,623 

49,656 




55,109 




218,088 



21,662 


361,133 

153,482 

128 


3,250,202 

35,455 




326,042 

270,847 




84,234 




198,623 





981 



64 



555 


2688 


1536 


1621 


1579 


512 


16 





128 


1152 


1314 


724 


384 


256 





64 


13,628 


85 














32 





72,227 


148,968 


230,223 


252,793 


99,696 


622,950 


27,153 


27,192 


1,060,829 


871,234 


3,954,410 


21,397,150 


1,932,063 


3,186,975 


354,349 


442,388 


2,227,102 


1,887,651 


5,982,435 


24,376,822 


4,752,529 


11,915,805 


1,232,874 


1,315,247 


306,963 


446,903 


979,574 


451,417 


486,558 


1,733,426 


822,677 


1,584,473 


306,963 


446,903 


979,574 


451,417 


486,558 


1,733,426 


822,677 


1,584,473 


81,255 


279,878 


419,818 


343,077 


379,190 


442,385 


144,448 


18,057 


15,335 


153,353 








591,435 


219,050 


240,959 


503,816 


14,763,970 


14,238,250 


18,641,155 


13,708,148 


16,143,985 


56,011,283 


5,542,049 


12,748,730 


1,171,918 


104,048 


3,066,790 


1,533,350 


1,555,255 


3,986,723 


353,223 


657,151 


328,575 


394,290 


6,900,008 


635,245 


240,955 


876,203 


35,596 


120,478 


16,361,053 


16,106,241 


22,817,681 


16,219,820 


18,910,820 


61,535,644 


6,316,275 


14,048,232 


34,306,660 


44,214,844 


69,068,298 


66,512,801 


39,870,929 


95,777,761 


11,461,481 


22,775,746 



Virginia State Water Control Board. 1982. Classifi- 
cation and Priority Listing of Virginia Lakes. 
U.S. Environmental Protection Agency. S- 
003219-010. Richmond, Virginia, 727 pp. 



Department of Biological Sciences, Old 
Dominion University, Norfolk, Virginia 
23529. 



280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



INTERNATIONAL COMMISSION ON ZOOLOGICAL 

NOMENCLATURE 

% British Museum (Natural History) 
Cromwell Road 
London, SW7 5BD 

23 September 1988 

The following applications were published on 23 September 1988 in Vol. 45, Part 
3 of the Bulletin of Zoological Nomenclature. Comment or advice on these appli- 
cations is invited for publication in the Bulletin and should be sent to the Executive 
Secretary, ICZN, British Museum (Natural History), Cromwell Road, London SW7 
5BD, U.K. 

Case No. 

2449 Septotrochammina Zheng, 1979 (Foraminiferida): proposed designation of 

Remaneica gonzalezi Seiglie, 1964 as the type species. 
2622 Pleuromma princeps Scott, 1894 (currently Gaussia princeps; Crustacea, Co- 

pepoda): proposed conservation of the specific name. 
2568 acrididae Karny, 1907, oedipodidae Walker, 1870 and locustidae La- 

treille, 1 802 (Insecta, Orthoptera): proposed order of precedence. 
2618 Bruchus Linnaeus, 1767, Ptinus Linnaeus, 1767 and Mylabris Fabricius, 1775 

(Insecta, Coleoptera): proposed conservation. 

2627 Coryphium angusticolle Stephens, 1834 (Insecta, Coleoptera): proposed con- 

servation of both the generic and specific names. 
2632 Tachina orbata Wiedemann, 1830 (currently Peribaea orbata; Insecta, Dip- 
tera): proposed confirmation of neotype designation. 

2628 Tenthredo zonula Klug, 1817 (Insecta, Hymenoptera): proposed conservation 

of the specific name. 

2625 Saccopharynx Mitchill, 1 824 (Osteichthyes, Saccopharyngiformes): proposed 
conservation. 

2616 ichthyophiidae Taylor, 1968 (Amphibia, Gymnophiona): proposed con- 
servation. 

2650 Thorius pennatulus Cope, 1 869 (Amphibia, Caudata): proposed conservation 
of the specific name. 

2441 Semioptera wallacii Gray, 1859 (Aves, Paradisaeidae): proposed confirma- 
tion as the correct spelling. 

2640 Mus musculus domesticus Schwarz & Schwarz, 1943 (Mammalia, Rodentia): 
proposed conservation. 

J. D. D. SMITH 
Scientific Administrator 




VOLUME 102, NUMBER 1 281 



INTERNATIONAL COMMISSION OF ZOOLOGICAL 

NOMENCLATURE 

% British Museum (Natural History) 
Cromwell Road 
London, SW7 5BD 

23 September 1988 

The following Opinions were published on 23 September 1988 in Vol. 45, Part 
3 of the Bulletin of Zoological Nomenclature. 

Opinion No. 

1501 Alveolina d'Orbigny, 1826 (Foraminiferida): Oryzaria boscii Defrance in 

Bronn, 1825 designated as the type species. 

1502 Conus fergusoni G. B. Sowerby III, 1873 (Mollusca, Gastropoda): specific 

name conserved. 

1503 pseudocalanidae Sars, 1901 (Crustacea, Copepoda): not to be given pre- 

cedence over clausocalanidae Giesbrecht, 1892. 

1504 berytidae Fieber, [1851] and Berytinus Kirkaldy, 1900 (Insecta, Heterop- 

tera): conserved. 

1505 Sigara scholtzi Fieber, [ 1 860] (currently Micronecta (Dichaetonecta) scholtzi; 

Insecta, Heteroptera): specific name conserved. 

1506 Oncomera Stephens, 1829 (Insecta, Coleoptera): Dryops femorata Fabricius, 

1792 designated as the type species. 

1507 Musca marginalis Wiedemann, 1830 (currently Chrysomya marginalis; In- 

secta, Diptera): specific name conserved. 

1508 Simulium austeni Edwards, 1915 (Insecta, Diptera): not to be given prece- 

dence over Simulium posticatum Meigen, 1838. 

1509 Paraphytomyza Enderlein, 1936 (Insecta, Diptera): Phytagromyza luteo- 

scutellata de Meijere, 1924 designated as the type species. 

1510 Microgaster Latreille, 1804 (Insects, Hymenoptera): Microgaster australis 

Thomson, 1895 deisgnated at the type species. 

1511 Halictus costulatus Kriechbaumer, 1873 (currently Lasioglossum costulatum; 

Insecta, Hymenoptera): specific name conserved. 

1512 Desorella Cotteau, 1855 (Echinodermata, Echinoidea): confirmation of Hy- 

boclypus elatus Desor, 1 847 as the type species. 

1513 Synapturanus Carvalho, 1954 (Amphibia, Anura): Synapturanus miranda- 

riberoi Nelson & Lescure, 1975 designated as the type species. 

1514 Liasis Gray, 1842 (Reptilia, Serpentes): Liasis mackloti Dumeril & Bibron, 

1844 designated as the type species. 

1515 laridae Rafinesque Schmaltz, 1815 (Aves) and larini LeConte, 1861 (In- 

secta, Coleoptera): homonymy removed. 

1516 Taeniolabis Cope, 1882 (Mammalia, Multituberculata): Poly mastodon taoensis 

Cope, 1882 designated as the type species. 

1517 Viverravus gracilis Marsh, 1872 (Mammalia, Carnivora): generic and specific 

names conserved. 



282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

REVIEWERS 

The following people reviewed manuscripts for the Proceedings in 1988. A. Alvarino, M. V. Angel, 
F. M. Bayer, B. Bieri, R. Birdsong, J. A. Blake, T. Bongers, J. Bouillon, T. E. Bowman, R. O. 
Brinkhurst, B. Brown, S. D. Cairns, D. R. Calder, R. Castro, F. L. Chace, Jr., L. L. Cole, B. B. Collette, 
P. F. S. Cornelius, R. F. Cressey, A. Crosnier, D. Davis, G. M. Davis, H. DeWitt, M. Dojiri, S. 
Edmonds, D. J. Eernisse, C. Ehrlich, K. Fauchald, K. Fitzhugh, V. A. Funk, A. L. Gardner, S. Gelder, 
I. M. Goodbody, T. M. Gosliner, G. R. Graves, K. Green, M. Harasewych, E. Harrison-Nelson, C. 
W. Hart, Jr., W. R. Heyer, H. H. Hobbs, Jr., W. D. Hope, J. Iverson, S. C. Jameson, B. F. Kensley, 
L. S. Kornicker, J. D. Kudenov, K. Kuroda, D. B. Lellinger, R. J. L. Lester, M. S. Loden, J. Lynch, 
W. G. Lyons, N. J. Maciolek, R. B. Manning, J. W. Martin, R. W. McDiarmid, J. H. McLean, M. 
Milligan, G. C. Mitchell, R. F. Modlin, R. J. Mooi, T. A. Munroe, A. T. Newberry, R. T. O'Grady, 
S. L. Olson, L. Parenti, K. Parks, D. L. Pawson, T. H. Perkins, M. H. Pettibone, W. L. Pratt, J. W. 
Reid, M. E. Rice, M. Robbins, C. R. Robins, W. Rudman, D. E. Russell, J. M. Savage, V. G. Springer, 
W. C. Starnes, M. Stiassny, F. G. Thompson, D. A. Thoney, R. P. Vari, M. Vecchione, T. C. Walter, 
L. Watling, S. H. Weitzman, M. J. Wetzel, A. B. Williams, S. Williams, R. Winterbottom, J. J. 
Wurdack, R. Zottoli, G. R. Zug, R. L. Zusi. 



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Eigenmann, C. H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South 

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CONTENTS 

Paleontological type specimens in the Santa Barbara Museum of Natural History 

Scott E. Miller 1 

A new generic name for Tate's (1933) Microtarsus group of South American mouse opossums 

(Marsupialia: Didelphidae) Alfred L. Gardner and G. Ken Creighton 3 

Description of a rock-dwelling cichlid (Teleostei: Cichlidae) from Lake Malawi, Africa 

J. R. Stauffer, Jr. and J. M. Boltz 8 

Ascidians collected around the Galapogos Islands using the Johnson-Sea-Link research sub- 
mersible Claude Monniot and Francoise Monniot 14 
Eukrohnia calliops, a new species of Chaetognatha from the northern Gulf of Mexico with 

notes on related species Jerry A. McLelland 33 

Smalleyus tricristatus, new genus, new species, and Pseudothelphusa parabelliana, new species 

(Brachyura: Pseudothelphusidae) from Los Tuxtlas, Veracruz, Mexico Fernando Alvarez 45 
Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae) 

Raymond B. Manning and L. B. Holthuis 50 
Synalpheus arostris and Philocheras lapillus, two new species of caridean shrimp (Crustacea) 

from the tropical eastern Pacific Mary K. Wicksten 78 

The holotype of Heterocarpus alexandh A. Milne-Edwards (Crustacea: Decapoda: Pandalidae) 

Fenner A. Chace, Jr. 84 
Gammaropsis arawakia, a new species of marine Amphipoda (Crustacea) from Jamaica i 

James Darwin Thomas and J. L. Barnard 89 4 
Natatolana pastorei (Giambiagi, 1925) (Crustacea, Isopoda, Cirolanidae) from the Straits of 

Magellan, South America: Redescription and notes on functional morphology 

Johann Wolfgang Wagele and Niel L. Bruce 95 
Neobrachiella anisotremi (Copepoda: Lernaeopodidae), a new species parasitic on an inshore 

fish, Anisotremus scapularis, off the Chilean coast 

Raul Castro Romero and Hernan Baeza Kuroki 106 
The First Annual Riser Lecture: Meiofauna and Microbes— the interactive relations of annelid 

hosts with their symbiotic bacteria Olav Giere 109 

Revision of the family Lysaretidae, and recognition of the family Oenonidae Kinberg, 1865 

(Eunicida: Polychaeta) G. Kent Colbath 1 1 6 

Redescription of Nephtys squamosa Ehlers (Polychaeta: Nephtyidae) Takashi Ohwada 124 

Phallodrilus hessleri, new species (Oligochaeta: Tubificidae) from abyssal depths in the western 

Pacific Ocean Christer Erseus 1 3 1 

Phallodrilus vescus, new species (Oligochaeta, Tubificidae) from the Gulf of Mexico 

Christer Erseus 1 34 
New species of scale-worms (Polychaeta: Polynoidae) from the hydrothermal rift-area of the 

Mariana Back- Arc Basin in the western central Pacific Marian H. Pettibone 137 

Polynoidae and Sigalionidae (Polychaeta) from the Guaymas Basin, with descriptions of two 

new species, and additional records from hydrothermal vents of the Galapagos Rift, 21°N, 

and seep-sites in the Gulf of Mexico (Florida and Louisiana) Marian H. Pettibone 1 54 

A new eulimid gastropod. Trochostilifer eucidaricola, parasitic on the pencil urchin Eucidaris 

tribuloides from the southern Caribbean Anders Waren and Robert Moolenbeek 169 

Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa River (exclusive of Ash 

Meadows) drainages. Death Valley System, California-Nevada Robert Hershler 176 

Four new species of free-living marine nematodes in the genus Pareurystomina (Nematoda: 

Enoplida) with observations on other members of the genus Edwin J. Keppner 249 

Neotropical Monogenea. 14. Revision of Callorhynchocotyle Suriano and Incorvaia, 1982 

(Hexabothriidae) with the description of C. amatoi 

Walter A. Boeger, Delane C. Kritsky, and Joaber Pereira, Jr. 264 
Phytoplankton composition in a borrow pit lake in Virginia / 

Seba B. Sheavly and Harold G. Marshall 272 / 
International Commission on Zoological Nomenclature: Applications and Opinions 280 

Reviewers- 1988 282 






PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY 

OF 

WASHINGTON 



VOLUME 102 
28 JUNE 1989 



NUMBER 2 



ISSN 0006-324X 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

1988-1989 
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Elected Council 

Gary R. Graves Meredith L. Jones 

W. Ronald Heyer Raymond B. Manning 

W. Duane Hope Wayne N. Mathis 

Custodian of Publications: David L. Pawson 



PROCEEDINGS 

Editor: C. Brian Robbins 

Associate Editors 

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Plants: David B. Lellinger Raymond B. Manning 

Insects: Wayne N. Mathis Vertebrates: G. David Johnson 

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PROC. BIOL. SOC. WASH. 
102(2), 1989, pp. 283-299 

A REVISION OF ARABELLA MUTANS 

(CHAMBERLIN, 1919) AND RELATED SPECIES 

(POLYCHAETA: ARABELLIDAE) 

G. Kent Colbath 

Abstract. —Arabella mutans (Chamberlin, 1919), previously considered cir- 
cumtropical in distribution, is revised using a combination of soft-part, setal, 
and maxillary characters. Emphasis is placed on adopting a population ap- 
proach to evaluating character states. Poorly preserved, isolated, or juvenile 
individuals may not be identifiable to species based on the criteria advocated 
here. Seven species with non-overlapping geographic ranges are recognized. 
Arabella mutans s. s. is restricted to tropical Polynesia. Arabella novecrinita 
Crossland, 1924 is recorded from the western Indian Ocean. Arabella logani 
Crossland, 1924 and A. atlantica Crossland, 1924 are recognized as distinct 
species, and recorded from the Red Sea and Cape Verde Islands, respectively. 
Arabella multidentata (Ehlers, 1887) is recognized to encompass western At- 
lantic and Carribean forms. Two new species are erected: Arabella panamensis 
for specimens from Pacific Panama, and A. monroi from the Galapagos. 



The genus Arabella is distributed 
throughout the world ocean, and includes 
20 named species (Fauchald 1977b). Many 
of these species are relatively rare, and have 
not been satisfactorily circumscribed in 
terms of either morphology or distribution. 

Arabella mutans (Chamberlin, 1919) is 
one such species. Although A. mutans is re- 
ported to have a circumtropical distribution 
(Monro 1928, 1933a, b; Day 1967; Reish 
1968; Gardiner 1976), this assertion has not 
been supported by detailed morphological 
comparisons of specimens (Fauchald 1970). 
The present study attempts to partially re- 
dress this situation by presenting informa- 
tion compiled from the detailed examina- 
tion of numerous specimens, including the 
holotypes of both A. mutans and the similar 
A. novecrinita Crossland, 1924. 

From a paleontological perspective, Ar- 
abella mutans is of interest because of the 
wide range of jaw morphologies which have 
been recorded for the species. Establishing 
constraints on the amount of variation in 
jaw morphology that can be expected within 



modern species should aid in the interpre- 
tation of fossil jaw assemblages. 

History of Study 

Chamberlin (1 9 1 9:329-332) described the 
new genus and species Cenothrix mutans 
based on a single specimen collected from 
Easter Island by the Albatross Expedition. 
He distinguished the genus Cenothrix from 
Arabella Grube, 1850 on the presence of a 
ventral, hooded seta in medial and posterior 
parapodia in the former, and the supposed 
absence of such setae in the latter. The new 
species was distinguished on the same taxo- 
nomic character. 

Crossland (1924:70) considered the pres- 
ence of a single hooded seta per parapodium 
insufficient grounds for recognition of a new 
genus, and transferred C mutans into Ar- 
abella. He described the new species Ara- 
bella novecrinita based on 16 specimens 
from the Atlantic and Indian Oceans, and 
noted that A. novecrinita "... is like, and 
probably identical with A. {Cenothrix) mu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tans," (Crossland 1924:71). Crossland ap- 
parently considered the jaws of the type 
specimen of A. mutans atypical, and justi- 
fied the erection of a new species on that 
basis. He recognized four varieties of A. no- 
vecrinita: the type material from Zanzibar 
and the Maldive Islands, A. novecrinita var. 
logani from the Red Sea, and A. n. var. 
atlantica and A. n. var. asymmetrica from 
the Cape Verde Islands. 

Monro (1928:91; 1933b:88) considered 
A. novecrinita a junior synonym of A. mu- 
tans, and considered the species circum- 
tropical in distribution. Monro essentially 
used Chamberlin's specific name for spec- 
imens included within the range of variation 
described by Crossland. Subsequent work- 
ers prior to Orensanz (1974) followed Mon- 
ro, although Fauchald (1970:129) noted that 
Monro's revision did not involve exami- 
nation of the type material of either species, 
and suggested that A mutans, as considered 
by Monro, might actually include several 
species with more restricted distributions. 

Orensanz (1974:388) used Cenothrix as a 
subgenus of Arabella, and recognized three 
species within the subgenus, which he dis- 
tinguished based on their maxillary mor- 
phology. A. mutans was recognized for spec- 
imens with asymmetrical maxillae with 
falcate MI elements, A. asymmetrica for 
those with asymmetrical maxillae with a bi- 
fid left MI, and A. novecrinita for those with 
symmetrical maxillae. 

Perkins (1979:445-450) followed this ar- 
rangement and placed three species in Ar- 
abella {Cenothrix). He made the important 
observation that the type specimens of both 
Arabella multidentata (Ehlers, 1887) and A 
maculosa Verrill, 1 900 have "hooded" ven- 
tral setae, and therefore should not be con- 
sidered junior synonyms of Arabella iricolor 
(Montagu, 1804) as proposed by Hartman 
(1944:173). Perkins distinguished A. mul- 
tidentata from A. mutans based on jaw sym- 
metry, and considered the two species sym- 
patric in the Florida region. He recognized 



A. maculosa based on the extreme asym- 
metry of the "hoods" on the ventral setae 
on the type specimen from Bermuda. 

Materials and Methods 

The extensive collection of specimens 
catalogued as Arabella mutans in the U.S. 
National Museum of Natural History 
(USNM) was examined in its entirety, as 
were samples identified as Arabella novecri- 
nita. Additional material was obtained from 
the British Museum (Natural History) 
(=ZK), including Crossland's type material 
for A. novecrinita. Selected specimens of Ar- 
abella iricolor from the National Museum 
collection and the Florida Department of 
Natural Resources (FSBC, EJ) were also ex- 
amined for comparative purposes. 

On each dissected specimen the lengths 
of the first maxillae (Mis), length and width 
of the prostomium, and length of the peri- 
stomium were measured. Measurements 
were made using an ocular micrometer on 
a Wild M5 wide-field microscope. Denticles 
were counted (where possible) for all max- 
illary jaws. 

Setae were prepared for scanning electron 
microscopy by allowing parapodia suspend- 
ed in alcohol to dry onto glass cover slips, 
which were in turn glued onto stubs. Jaws 
were mounted by leaving a flange of muscle 
tissue around the jaw margins which, when 
dried, fixed the jaws firmly onto glass cover 
slips. All specimens were coated with a thin 
film of carbon, then sputter coated with gold 
palladium. Observations were made on a 
Cambridge scanning electron microscope at 
the National Museum of Natural History, 
Smithsonian Institution. 

Taxonomic Characters 

Arabella mutans has been recognized by 
most authors based on the presence of a 
single morphologic character (the "hooded" 
ventral seta). Consideration of additional 
characters coupled with a closer examina- 



VOLUME 102, NUMBER 2 



285 



Table 1 . — Distribution of taxonomic characters in species of Arabella under consideration. 





Nuchal 


Anal 


Postsetal 








Species 


papillae 


cirri 


lobe 


Ventral setae 


Maxillary symmetry 


Robust MI 


iricolor 


no 


2 


short 


ridged, gradual 


invariant 


bifid-falcate 


atlantica 


no 


? 


short 


ridged, abrupt 


invariant 


falcate 


logani 


no 





short 


ridged, gradual 


? 


absent 


monroi 


no 





short 


ridged, abrupt 


invariant 


falcate 


multidentata 


no 


4 


short 


ridged, abrupt 


polymorphic 


bifid 


mutans 


no 


4 


short 


smooth, abrupt 


polymorphic 


dentate 


novecrinita 


no 


4 


short 


smooth, abrupt 


invariant 


absent 


panamensis 


no 


2 


long 


ridged, abrupt 


polymorphic 


falcate 


sp. A 


yes 


? 


short 


ridged, abrupt 


polymorphic 


bifid 



tion of the setae has allowed separation of 
discrete morphological subgroups, here rec- 
ognized as distinct species with discrete geo- 
graphic ranges. The taxonomic characters 
are discussed below, and the character states 
exhibited by each of the species are listed 
in Table 1 . 

Soft-part morphology. —Soft-part mor- 
phology within the genus Arabella is noto- 
riously uniform and relatively simple, which 
accounts in part for the difficulty encoun- 
tered in treating these worms taxonomical- 
ly. The relative proportions of the prosto- 
mium are fairly uniform throughout the A. 
mutans species complex, and the ventral 
furrowing of the peristomium, although 
variable, does not seem to vary in a system- 
atic fashion. 

There are three features which are useful 
in differentiating these species, however. The 
postsetal lobe in parapodia from A. pana- 
mensis, n. sp. is markedly longer than that 
in the other species. 

Nuchal papillae are present in some large 
individuals from Guayanilla Bay, Puerto 
Rico, a unique feature in the arabellids. The 
specimens are from a population of other- 
wise identical worms which lack papillae, 
however, and the taxonomic significance of 
this character is unclear. 

The morphology of the pygidium is an 
extremely useful character, limited only by 
the difficulty of obtaining complete speci- 



mens. Species are characterized by having 
two anal cirri (Fig. 3b, f), four anal cirri (Fig. 
3c, d), or two swollen pads lacking cirri (Fig. 
3a, e). 

Setae. — The most important point to 
emerge from SEM examination of the setae 
from the specimens studied here is that the 
"hooded" ventral seta in medial and pos- 
terior parapodia used to define A. mutans 
is not hooded (in the sense of having a three 
dimensional structure enclosing the distal 
end of the seta). In all specimens examined, 
the distal end of the ventral seta consists of 
a relatively flat extension of the setal shaft. 

Although the diameter of the ventral seta 
is generally twice that of the other setae in 
these forms, it originates in the same mus- 
cular bundle, and is not an acicula as re- 
corded by some authors. The modification 
of the distal end of the ventral seta is dif- 
ferent only in degree, not in structure, from 
that observed in the ventral setae of Ara- 
bella iricolor. This observation strongly sup- 
ports Crossland's (1924) contention that 
Cenothrix Chamberlin should be regarded 
as a junior synonym of Arabella. 

The terminology adopted here to describe 
the ventral setae is illustrated in Fig. 1 . Three 
distinct morphologies are recognized. The 
shaft of the seta may be smooth with an 
abrupt taper from the shaft to the guard, 
ridged with an abrupt taper from the shaft 
to the guard, or ridged with a gradual taper 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



shaft 




guard 

Fig. 1 . Terminology used to describe morphology of the ventral seta in medial and posterior parapodia. 



from the shaft to the guard. Arabella pan- 
amensis, n. sp. has ventral setae which are 
somewhat modified from the second type 
above in having an additional ridge near the 
tip of the shaft (Fig. 3h). 

SEM examination of the other setae re- 
veals that the distinction between smooth 
and "toothed" geniculate setae made by both 
Chamberlin (1919) and Crossland (1924) is 
indeed significant. The "toothed" setae are 
not actually toothed, however, but are in- 
stead ringed with loose rows of large fibrils, 
in contrast to the tight, irregular arrange- 
ment of fibrils in the smooth setae (Fig. 4k, 

i). 

Jaws.— Maxillary jaws in eunicoid poly- 
chaetes are numbered, by convention, from 
posterior to anterior with roman numerals. 
Left and right jaws are indicated by a capital 
L or R respectively. Thus MIR refers to the 
posterior maxilla on the right side of the 
apparatus. Denticle formulas are indicated 
in sequence, with the left jaw in a given pair 
listed first. For populations, the low end of 
the range of denticles for a given jaw is listed 
first, with the high end of the range in pa- 
rentheses. 

Maxillae within some of the species are 
here considered polymorphic, but variation 
within single populations is limited to two 
or three morphs. The first and second max- 
illae (MI and Mil) were found to be the most 
significant in differentiating species. The 
morphs recognized here are illustrated in 
Fig. 2. The Mil jaws may be either "long," 



extending posteriorly to the base of the MI, 
or "short," extending posteriorly only half 
of the distance to the base of the MI (Fig. 
2a, b). A long Mil is invariably associated 
with a relatively slender MI (here termed 
"gracile") with a single falx (or hook) and a 
pronounced gap between the falx and the 
basal denticle row (Fig. 2e). Mis adjacent 
to short Mils are wider and generally slight- 
ly longer than the gracile Mis, and are here 
termed "robust." They may take one of three 
forms in adult worms. These jaws may have 
a single falx with pronounced gap (= "fal- 
cate," Fig. 2c), a bifid falx with pronounced 
gap (="bifid," Fig. 2d), or no distinct falx 
and no gap (="dentate," Fig. 2f). Dentate 
Mis are found in juvenile specimens of a 
number of different species, but in the ju- 
veniles with dentate Mis the anterior den- 
ticles project laterally rather than dorsally 
as in the adults (compare Fig. 4c, d). 

The statistical frequency of these jaw 
morphologies varies systematically among 
different populations of worms, and in most 
cases only one of the three morphologies of 
robust Mis is represented (if any). The pres- 
ence or absence of jaw symmetry polymor- 
phism within a population is also a useful 
taxonomic character. 

Within the species considered here, the 
lengths of Mis are highly correlated with the 
sizes of individual worms as measured by 
prostomial width (Fig. 7). A plot of MI length 
against prostomial width is similar for most 
of these species, with the exception of Ar- 



VOLUME 102, NUMBER 2 



287 





Fig. 2. Schematic representation of morphologies observed in first and second maxillae, a, short Mil; b, long 
Mil; c, falcate robust MI; d, bifid robust MI; e, gracile falcate MI; f, dentate robust MI. 



abella panamensis. In A. panamensis indi- 
viduals have relatively large jaws for their 
size. 

The number of denticles on a given max- 
illa is variable, and has generally been dis- 
counted as a useful taxonomic character in 
eunicoid polychaetes (Kielan-Jaworowska 
1966, Wolf 1980). With sufficient speci- 
mens, however, it may be possible to use 
the number of denticles on a statistical basis 
(see below under Arabella multidentata). 

The shape of the mandibles exhibits with- 
in-sample variation in these forms (Cross- 
land 1924), but proved of no value in dis- 
tinguishing species. A typical mandible is 
illustrated in Fig. 5b. 

Order Eunicida Fauchald, 1977 

Family Arabellidae Hartman, 1944 

Genus Arabella Grube, 1850 

Arabella Grube, 1850:17. 
Notopsilus Ehlers, 1868:406. 
Maclovia Grube, 1871:58. 
Cenothrix Chamberlin, 1919:329. 

Type species. —Arabella tricolor (Monta- 
gu, 1804). 

Remarks. -Fauchald (1977b:l 10-1 1) 
noted that generic diagnoses within the Ar- 



abellidae are inconsistent. In forms with 
projecting acicular spines, the shape of the 
MI is used to separate the genus Drilonereis 
Claparede (falcate Mis) from Notocirrus 
Schmarda (dentate Mis), while the genus 
Arabella (which lacks emergent acicular 
spines) includes forms with both maxillary 
morphologies. Fauchald suggested that 
either the genus Notopsilus Ehlers be rec- 
ognized to include species with dentate Mis 
which are presently placed in Arabella, or 
that Drilonereis and Notocirrus be consid- 
ered synonymous. 

The report here of species in which the 
jaws are polymorphic (including both den- 
tate and falcate Mis) strongly suggests that 
Notopsilus should be considered a junior 
synonym of Arabella. This does not imply, 
however, that Drilonereis and Notocirrus 
should also be considered synonymous. The 
basic arrangement of jaws is quite different 
in Drilonereis than it is in Notocirrus, and 
intermediate or hybrid maxillae have not 
been recorded. 

Arabella iricolor (Montagu, 1804) 
Figs. 3b, 4j, k, 5a, 6a 

Nereis iricolor Montagu, 1 804. 

Arabella iricolor. — Pettibone, 1963:269, fig. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. a, Arabella logani, holotype, ZK 1924.3.1.15, pygidium, left-lateral view; b, Arabella tricolor, USNM 
61732, pygidium, right-lateral view; c, Arabella novecrinita, lectotype, ZK 1924.3.1.15, pygidium, dorsal view; 
d, Arabella mutans, USNM 57434, pygidium, oblique right-lateral view; e, Arabella monroi, holotype, ZK 
1932.12.24.595, pygidium, right-lateral view; f-h, Arabella panamensis, USNM 98237, holotype; f, pygidium, 
dorsal oblique right-lateral view; g, posterior parapodium, anterior view, note elongate post-setal lobe; h, ventral 
seta of parapodium in g, note 2 ridges, abrupt taper of shaft; i, Arabella sp. A, USNM 54915, anterior end with 
pharynx extended, lateral view, note nuchal papillae. Scale bars on a-f, i = 0.5 mm; g = 0.2 mm; h = 20 jwn. 



71a-e; Gardiner, 1976:206-207, figs. 

27r-u, 28a; Uebelacker, 1984:42-5, fig. 

42-2. 

Materials examined. — USNM 30058, 
FSBC I 24159, Seashore Key, Florida (6); 
FSBCI 13436, Terra Ceia Bay, Florida(l).- 
FSBC I 13102, Lower Tampa Bay, Florida 
(l).-EJ-72-66, EJ-73-414, Hutchinson Is- 
land, Florida (3). -USNM 61732, USNM 
53026, Cape Lookout, North Carolina 
(ll).-USNM 4676, Woods Hole, Massa- 
chusetts (7). -USNM 33787, Province- 
town, Massachusetts (8). 



Remarks.— The specimens of Arabella 
iricolor examined here are characterized by 
having ventral setae in medial and posterior 
parapodia which are ridged, with a gradu- 
ally tapering shaft (Fig. 4j). The pygidium 
has two cirri which arise dorsally (Fig. 3b). 
No variation in jaw symmetry was observed 
among adult specimens. In each individual 
the MIL is robust with a short MIIL, the 
MIR gracile with a long MIIR. 

The Florida specimens differ from those 
from North Carolina and Massachusetts in 
that all adult worms have falcate robust Mis 



VOLUME 102, NUMBER 2 



289 




Fig. 4. Setae, all in anterior view, a, b, Arabella monroi, ZK 1932.12.24.337-340, paratype, large anterior 
fragment, ventral seta, setiger 143, note ridge, abrupt taper of shaft, c, Arabella novechnita, ZK 1924.3.1.15. 
lectotype, ventral seta, setiger 200, note smooth, abruptly tapered shaft, d, Arabella mutans, USNM 23933. 
ventral seta, posterior setiger, note smooth, abruptly tapered shaft, e, Arabella mutans, USNM 57434. ventral 
seta, posterior setiger, dorsal to bottom of page, note smooth, abruptly tapered shaft, f, Arabella atlantica, ZK 
1923.11.5.1-2, large syntype, ventral seta, setiger 122, note ridge, abrupt taper, g, h, Arabella logani, ZK 
1924.3.1.15, holotype, ventral seta, setiger 125, note ridge, gradually tapered shaft, i, Arabella multidentata, 
USNM 98154, ventral seta, posterior setiger, note ridge, abruptly tapered shaft, j, k, Arabella iricolor, USNM 
30058 (Florida); j, ventral seta, posterior setiger, note ridge, gradually tapered shaft; k, "smooth" seta, posterior 
setiger. 1, Arabella sp. A, USNM 54915, posterior setiger, smooth seta above, toothed seta below. All scale bars 
= 5 iiva. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



(n = 8), in contrast to the bifid robust mor- 
phology (Fig. 5 a) observed in the northern 
populations (25 of 26 adult specimens). 

Jaw morphology in these populations is 
quite stable, as the jaws are essentially iden- 
tical in specimens collected from Woods 
Hole in 1883 (USNM 4676), Provincetown 
in 1954 (USNM 33787), and North Caro- 
lina in 1974 (USNM 53026) and 1975 
(USNM 61732). Thus although the differ- 
ence between the Florida and northern 
populations is small, the stability of the 
character suggests that the difference is he- 
reditary, and may be used to subdivide east 
coast populations of A. iricolor. As presently 
recognized, A. iricolor is probably a species 
complex. 

Two juvenile specimens of A. iricolor from 
Florida have dentate MILs which resemble 
those in several other species of Arabella, 
including A. mutans (Fig. 5d). 

Arabella atlantica Crossland, 1924, 

new rank 

Fig. 4f 

Arabella novecrinita var. atlantica Cross- 
land, 1924:78-80, text-figs. 99-101, 105. 

Arabella novecrinita var. asymmetrica 
Crossland, 1924:80-83, text-figs. 102- 
104. 

Syntypes. -ZK 1923.11.5.1-2, Cape 
Verde Islands, two anterior fragments. 

Materials examined.— ZK 1923.11.5.1- 
2, Cape Verde Islands (two anterior frag- 
ments).— jaw mounts from six specimens 
from the Cape Verde Islands as follows: ZK 
1923.11.8.12-14, three unregistered slides 
labelled CV Isl. W9; three unregistered jaw 
mounts of Arabella novecrinita var. asym- 
metrica, Cape Verde Islands. 

Remarks.— Crossland (1924:78) record- 
ed 1 1 specimens from the Cape Verde Is- 
lands. Three small specimens which exhibit 
maxillae typical of juveniles in the genus 
Arabella were assigned to Arabella nove- 
crinita var. asymmetrica (Crossland, 1924, 



text-fig. 103). These specimens are here re- 
garded as juveniles of Arabella atlantica. 

Of the remaining eight specimens, two 
anterior fragments and six sets of jaws are 
represented in the collection of the British 
Museum (Natural History). At present it is 
impossible to determine which sets of jaws 
belong to the two preserved specimens, and 
thus no lectotype is designated. 

The soft-part morphology of this species 
is adequately described by Crossland (1924: 
78-80, text-fig. 99). The ventral setae are 
ridged, with an abrupt taper from the guard 
to the shaft (Fig. 4f). In the specimens avail- 
able for examination, the MIL is robust fal- 
cate, the MIIL short, and the MIR gracile 
falcate with a long MIIR. No symmetry 
polymorphism observed (n = 6). 

This species is distinguished from Ara- 
bella novecrinita by its ridged ventral setae 
and robust MIL. It differs from A. multi- 
dentata in having falcate rather than bifid 
robust Mis, and in not exhibiting symmetry 
polymorphism. A. atlantica most closely re- 
sembles A. monroi n. sp. from the Gala- 
pagos, but because neither preserved spec- 
imen of A. atlantica has an intact pygidium, 
it cannot be compared based on this im- 
portant character. 

Arabella logani Crossland, 1924, new rank 
Figs. 3a, 4g, h 

Arabella novecrinita var. logani Crossland, 
1924:75-78, text-figs. 96-98. 

Holotype. -ZK 1924.3.1.138, Red Sea. 

Materials examined. — ZK 1924.3.1.138, 
Red Sea (1). 

Remarks. —The soft-part morphology of 
the type and only specimen is accurately 
described by Crossland (1924:75-76). 
Noteworthy is the presence of two inflated 
pads on the pygidium (Fig. 3a) which lack 
anal cirri. No evidence for incomplete re- 
generation is apparent, and this may rep- 
resent the fully developed character state. 

The ventral setae in posterior parapodia 



VOLUME 102, NUMBER 2 



291 




Fig. 5. a, Arabella tricolor, USNM 33787, maxillae, dorsal view, note bifid MIL. b-d. Arabella mutatis, 
USNM 23933; b, mandibles, ventral view; c, maxillae of adult specimen, dorsal view, note dentate MIR. falcate 
tip of MIL broken during mounting; d, maxillae of juvenile specimen, note dentate MIL with anterior denticles 
which project laterally, e, Arabella sp. A, USNM 549 1 5, maxillae, dorsal view, example of symmetrical apparatus 
with 2 long Mils, 2 gracile falcate Mis. f, Arabella panamensis, USNM 66043. paratype. maxillae, dorsal view, 
note falcate MIL, falcate tip of MIR broken during mounting. Scale bars on a-c. e. f = 100 Mm: d = 50 nm. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



are ridged and gradually tapering (Fig. 4g, 
h), and are thus unlike those in other species 
within the Arabella mutans complex. 

The MIL is robust bifid, the MIIL short, 
and the MIR gracile falcate with a long MIIR 
(note that Crossland reversed the image in 
text-fig. 96, and incorrectly labeled the spec- 
imen as a "Cape Verde specimen"). The 
bifid MIL serves to distinguish this species 
from Arabella tricolor, which in the eastern 
Mediterranean and Red Sea has a falcate 
robust MIL (Ben-Eliahu 1976:172). No 
statement concerning jaw variability can be 
made based on a single specimen, and full 
characterization of the species must await 
further collecting. 

Arabella monroi, new species 
Figs. 3e, 4a, b 

Arabella mutans. —Monro 1933b:88-89 (in 
part). 

Holotype. -ZK 1932.12.24.595, washed 
from peat, Isla Santa Cruz (Indefatigable), 
Galapagos Islands. 

Etymology. —In honor of C. C. A. Monro. 

Materials examined. —Holotype; ZK 
1932.12.337-340 (paratypes), washed from 
peat, Isla Santa Cruz (Indefatigable), Ga- 
lapagos Islands (26). -ZK 1932.12.336, fine 
gravel in pools, Isla Isabela (Albermarle), 
Galapagos Islands (l).-ZK 1932.12.341, 
Post Office Bay, Isla Santa Maria (Charles), 
Galapagos Islands (3). 

Description. — Prostomium bullet-shaped, 
slightly longer than wide to slightly wider 
than long, V-shaped furrow on ventral sur- 
face, four eyes embedded in posterior dorsal 
margin. Peristomium divided into two rings, 
a shallow ventral median furrow in anterior 
ring only. 

Setigers numerous, exceeding 200 in large, 
adult specimens. Notopodia represented by 
small tubercle, neuropodia with blunt, sub- 
triangular postsetal lobe. Pygidium with two 
swollen pads (Fig. 3e), no anal cirri ob- 
served. 



Notoacicula small, 2-3 per setiger. Neu- 
roacicula large, amber colored, 3-4 per se- 
tiger with tips protruding slightly through 
parapodial wall. Anterior setigers with 5-6 
setae, 3 being smooth, 2-3 toothed. Medial 
and posterior setigers with 2 smooth, 1 
toothed, and 1 ventral seta. Ventral setae 
ridged, shaft tapering abruptly to distal guard 
(Fig. 4a, b). 

Mandibles H-shaped, typical of Arabella. 
Maxillae with 2 long carriers and unpaired 
ventral ligament (="third carrier" of some 
authors). Maxillary apparatus asymmetri- 
cal, no symmetry polymorphism observed. 
MIL robust falcate, MIIL short, MIR gracile 
falcate, MIIR long. Denticle formula: 6(9)- 
6(8); 7(8)-12(16); 5(6)-5(6); 4(5)-4; 1-1. 

Holotype large, complete individual with 
208 setigers, prostomium 0.68 mm long, 
0.55 mm wide, peristomium 0.54 mm long, 
MIL 0.36 mm long, MIR 0.35 mm long, 
denticle formula 9-8; 8-16; 6-6; 4-4; 1-1. 

Remarks.— The collection from the Ga- 
lapagos includes 13 complete specimens. 
None have anal cirri, and it seems likely 
that the presence of two pygidial lobes with 
no cirri represents the fully developed adult 
character state. 

Arabella monroi is distinguished from A. 
mutans based on its asymmetrical maxillae 
which are not polymorphic, falcate rather 
than dentate robust Mis, ridged rather than 
smooth ventral setae, and by the absence of 
anal cirri. It differs from A. multidentata in 
having falcate rather than bifid robust Mis, 
in not exhibiting symmetry polymorphism 
of the maxillae, and in lacking anal cirri. 

The maxillary apparatus in A. monroi is 
identical to that in A. atlantica. The pygid- 
ium in the latter species is unknown, how- 
ever, and I consider it prudent to describe 
the Galapagos specimens as a separate 
species pending collection of additional ma- 
terial from the type locality of A. atlantica. 

The geographic range of A. monroi is pres- 
ently restricted to the islands of the Gala- 
pagos. 



VOLUME 102, NUMBER 2 



293 



Arabella multidentata (Ehlers, 1887) 
Figs. 4i, 6b, 7 

Aracoda multidentata Ehlers, 1887:1 12, pi. 
34, figs. 8-10, pi. 35, figs. 1-4. 

Arabella multidentata. — Hartman, 1938:12; 
Perkins, 1979:447-449, fig. 13; Uebe- 
lacker, 1984:42-9, figs. 42-5, 6a-g. 

Arabella maculosa Verrill, 1900:651.— 
Treadwell, 1921:114, pi. 9, figs. 12-13, 
text-figs. 424-428; Perkins, 1979:450, fig. 
14. 

Arabella mutans.— Gardiner, 1976:206, figs. 
27o-q; Fauchald, 1977a:45 (in part); Per- 
kins, 1979:445-447, fig. 12; ?Duenas, 
1981:193. 

Arabella novecrinita. — Monro, 1933a:260- 
261. 

1 Arabella asymmetrica.-Orensa.nz, 1974: 
368-388, fig. III-6. 

Holotype.—MCZ 825, Florida. 

Materials examined. — USNM 55882, 
Gulf of Mexico (l).-USNM 54694-54696, 
Hutchinson Island, Florida (3).— USNM 
59526, N Atlantic off Florida (2); USNM 
59528, N Atlantic off N Carolina (1).- 
USNM 56844, N Atlantic off Virginia (1). - 
USNM 98142-98156, Bermuda (22).- 
USNM 53297, 54215, 54220, 54221, 54223, 
54818, 54820, 98238, Guayanilla Bay, 
Puerto Rico (22). -USNM 54217, 54218, 
54222, 54837-54840, Jobos Bay, Puerto 
Rico (8).-USNM 66044-66046, 73653, 
Galeta Reef, Panama (5).-?USNM 58273, 
Bay of Cartagena, Colombia (1). 

Remarks. -Perkins (1979:448-449) ac- 
curately described the soft-part morphology 
of A. multidentata. To his description of the 
setae I add that the ventral setae are ridged 
and abruptly tapered (Fig. 4i). 

Perkins (1979:450) distinguished Arabel- 
la maculosa Verrill, 1900 from A. multi- 
dentata based on the "more obviously 
asymmetrical" ventral setae in the syntype 
of the former. I found the degree of asym- 
metry variable on a single specimen, and 
was unable to separate the species on this 



Table 2. — Distribution of specimens of Arabella with 
particular MI morphologies in polymorphic species. 



Species 


2 robust 


1 gracile, 
1 robust 


2 gracile 


multidentata 








Bermuda 


1 


9 


8 


Others 


3 


15 


12 


mutans 


4 


4 


2 


panamensis 


3 


5 


— 


sp. A 


— 


4 


2 



basis. The syntype of A. maculosa has a 
regenerated pygidium with two poorly de- 
veloped lobes and no cirri (Perkins 1979, 
fig. 14a). Eight complete specimens from 
Bermuda, with no indication of posterior 
regeneration, have four anal cirri as is typ- 
ical of A. multidentata. The jaws of the Ber- 
muda specimens are indistinguishable from 
those of other west Atlantic and Caribbean 
populations of A. multidentata (Table 2), 
and I therefore consider A. maculosa a ju- 
nior synonym of A. multidentata. 

Perkins (1979:447) distinguished sym- 
patric Florida populations of A multiden- 
tata from A. mutans based on jaw sym- 
metry. Instead, I recognize a single species 
{A. multidentata) with polymorphic jaw 
symmetry which is distinct from A. mutans. 
A. multidentata has bifid (25 of 27 adult 
specimens with robust elements, remaining 
2 falcate) rather than dentate robust MI ele- 
ments (where present), and further differs 
from A. mutans in having ridged rather than 
smooth ventral setae. 

A comparison between A. multidentata 
and A. iricolor sensu Pettibone ( 1 963) is par- 
ticularly instructive, as the geographic ranges 
of the two species overlap along the east 
coast of the United States. A. multidentata 
has abruptly tapering shafts on the ventral 
setae (Fig. 4i), rather than gradually tapering 
shafts as in A. iricolor (Fig. 4j), is further 
characterized by having four rather than two 
anal cirri, and by having polymorphic max- 
illary symmetry. The number of denticles 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



30 - 



25 - 



to 

O 20- 



O 

w 15 - 

_Q 

E 

=> 10 - 

Z 



5 - 



- 



X = 12 2 



a 



X =13 8 



6 7 8 9 10 11 12 13 14 15 16 6 7 8 9 10 11 12 13 14 15 16 

Number of Denticles 

Fig. 6. Histograms of denticle number in Mil jaws in Arabella tricolor (a) and Arabella multidentata (b). 
Note that the distinction between short and long Mil elements is evident from the diagram. Means are indicated 
for long Mils. 



in long Mil elements overlaps between the 
two species (Fig. 6a, b), but the difference 
in means is highly significant (P < 0.001, t 
= 6.14). 

Dentate jaws were observed in four ju- 
venile specimens of A. multidentata. The 
juvenile dentition is indistinguishable from 
that in other species of Arabella. 

As defined here, Arabella multidentata is 
a west Atlantic species recorded from the 
east coast of the United States as far north 
as Virginia, from Bermuda, Puerto Rico, the 
Gulf of Mexico, Atlantic Panama, and ques- 
tionably from the Bay of Cartagena, Colom- 
bia (single specimen with falcate rather than 
bifid robust MIL). The specimens described 
by Orensanz (1974) as A. asymmetrica have 
maxillae identical to the present material, 
and ventral setae with abruptly tapered 
shafts. It's unclear whether or not the ven- 
tral setae are ridged, however, and the py- 
gidium was undescribed. This attribution 
must therefore be considered tentative. 



Arabella mutans (Chamberlin, 1919) 
Figs. 3d, 4d, e, 5b-d 

Cenothrix mutans Chamberlin, 1919:330- 
332, pi. 61, figs. 2-9, pi. 62, fig. 1. 

Arabella mutans. — Crossland, 1924:71; 
Hartman, 1954:624; Reish, 1968:220; 
Kohn and Lloyd, 1973:699. 

Arabella novecrinita. — Hartman, 1954:625. 

Holotype. -USNM 19740, Easter Island. 

Materials examined. —USNM 19740, 
57434, Easter Island (2). -USNM 23933- 
23935, Bikini Island (14). 

Remarks.— Chnmberlitfs (1919:330, pi. 
61, fig. 4) description of the anterior end of 
the holotype is accurate. The pygidium is 
poorly developed, and represents incom- 
plete regeneration. A second specimen from 
Easter Island (USNM 57434) has four anal 
cirri (Fig. 3d), as do two specimens from 
Bikini, and this is considered the fully de- 
veloped character state. 

As noted above, the hooded acicula (or 



VOLUME 102, NUMBER 2 



295 



"crochets") which Chamberlin (1919:331) 
originally described for this species are nei- 
ther hooded nor acicula, but rather are setae 
typical of the genus Arabella which have an 
abruptly tapering shaft. The shaft is smooth 
on ventral setae in the holotype, and on the 
other specimens examined (Fig. 4d, e). 

Chamberlin ( 1 9 1 9:6 1 , fig. 2) inaccurately 
figured the Mis in the holotype. The MIL 
is robust dentate, with two upturned ante- 
rior denticles (as in MIR in Fig. 5c) which 
were not depicted by Chamberlin. The MIR 
is gracile falcate with the anterior tip broken 
off. Examination of additional specimens 
reveals that maxillary symmetry is poly- 
morphic (Table 2), and that if robust Mis 
are present they are dentate (Fig. 5 c). Two 
juvenile specimens were observed with den- 
tate Mis of the type illustrated in Fig. 5d. 
Mandibles (Fig. 5b) are as described by 
Chamberlin (1919:332, pi. 61, fig. 3). 

Arabella mutans most closely resembles 
A. novecrinita as restricted here. In contrast 
to the symmetry polymorphism exhibited 
by A mutans, A. novecrinita is characterized 
by symmetrical maxillae with gracile Mis 
and long Mils. In view of the small number 
of A. novecrinita included here (n = 4), this 
distinction must be considered tentative. A. 
mutans has more denticles on long Mils on 
average (13.3, n = 12) than does A. nove- 
crinita (10.6, n = 5), but again the small 
sample size makes use of this distinction 
tentative. 

As restricted here, A. mutans has been 
recorded from Easter Island and the Mar- 
shall Islands in the tropical Pacific. A spec- 
imen from Guam (USNM 54426), which 
had been identified as A. mutans, is now 
missing its jaws, and the identification could 
not be confirmed. 

Day's (1975:196-197) report of A mu- 
tans from Western Australia was based on 
a single juvenile specimen, and must be con- 
sidered questionable on that basis. The jaws 
illustrated by Fauvel (1953:275, fig. 1401) 
from a small specimen collected from Cey- 



lon are also those of a juvenile, and the 
record cannot be accepted. Tampi & Ran- 
garajan (1964:11, figs. 29-31) did not de- 
scribe the jaws in the anterior fragment they 
identified as A. mutans from Car Nicobar, 
Indian Ocean, and their record must also 
be considered questionable. Silva (1961:182) 
described but did not illustrate a specimen 
from Ceylon as A. mutans which has two 
"hooded acicula" per parapodium, and 
asymmetrical maxillae with MIL robust fal- 
cate, MIIL short, MIR gracile falcate, and 
MIIR long. The presence of two "hooded" 
setae per parapodium is unusual, and the 
maxillae are unlike those in either A. mu- 
tans or A. novecrinita. 

Arabella novecrinita Crossland, 1924 
Figs. 3c, 4c 

Arabella novecrinita Crossland, 1924:71-75, 
text-figs. 89-95. 

Lectotype.—ZK 1924.3.1.15, Zanzibar; 
designated herein. 

Materials examined. — ZK 1924.3.1.15, 
Zanzibar (l).-ZK 1924.3.1.292, Minikoi 
Island, Maldives, mounted jaws and para- 
podia (1?). — uncatalogued anterior frag- 
ment, Aldabra. 

Remarks.— Crossland (1924:71) record- 
ed two specimens (with maxillary formulas) 
from the Maldives. The British Museum 
(Natural History) collection includes only 
mounted jaws and parapodia from this lo- 
cation which may be from a single speci- 
men. In light of the fragmentary nature of 
the Maldive material, and the wide geo- 
graphic separation between Crossland's 
original type localities, the Zanzibar speci- 
men is herein designated the lectotype. 

Crossland's (1924:71-75, text-fig. 89) de- 
scription of the soft-part morphology in this 
species is accurate. Noteworthy is the pres- 
ence of four anal cirri (Fig. 3c) on the lec- 
totype (Crossland 1924:75). The ventral se- 
tae in medial and posterior parapodia have 



296 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



a smooth shaft which tapers abruptly to the 
guard (Fig. 4c). The maxillae on the lecto- 
type are symmetrical, with gracile Mis and 
long Mils (Crossland 1924:72), as are the 
jaws on the two specimens Crossland (1924: 
73) recorded from the Maldives, and the 
anterior fragment recorded here from Al- 
dabra. The jaw formula for the lectotype is 
7-7; 7 + -10; 5-5; 5-4; 1-1. For the species it 
is 6(7)-7; 10(1 1)-10(1 1); 4(6)-4(5); 4(5)-3(4); 
1-1. The mandibles are typical of Arabella, 
as illustrated by Crossland (1924, text-fig. 

91). 

As noted above, the setae and soft-part 
characteristics of this species are similar to 
those in Arabella mutans, and the species 
are distinguished based on the maxillae. 

As restricted here, A. novecrinita is known 
only from islands in the Indian Ocean. Day 
(1967:446, fig. 17.18f-h) recorded speci- 
mens (as A. mutans) from Mozambique and 
South Africa which may belong in A. no- 
vecrinita based on the illustrated maxillary 
apparatus. It's not clear whether the ventral 
setae are ridged based on Day's (1967, fig. 
1 7. 1 8f) illustration, and thus the report can- 
not be securely credited to A. novecrinita. 
A South African specimen identified as A. 
mutans in the British Museum collection 
(ZK 1932.11.24.38) proved to be a mis- 
identified specimen of Arabella iricolor sen- 
su lato, and thus this question remains un- 
resolved. 

Arabella panamensis, new species 
Figs. 3f-g, 5f, 7 

Arabella mutans. — Fauchald, 1977a:45 (in 
part). 

Holotype. -USNM 98237, Paitilla Beach, 
Panama. 

Etymology. —From Panama, the type lo- 
cality. 

Materials examined. —Holotype; USNM 
66042, 66043, 66365 (paratypes), Paitilla 
Beach, Panama (9).-?USNM 80225, Golfo 
de Nicoya, Costa Rica (1). 

Description. — Prostomium bullet-shaped, 



slightly wider than long, V-shaped mark on 
ventral surface, four eyes embedded in pos- 
terior dorsal margin. Peristomium divided 
into two rings, ventral median furrow in 
anterior ring only. 

Setigers numerous, up to 177 in complete 
adults. Notopodia not expressed on surface, 
neuropodia with elongate postsetal lobe (Fig. 
3g) which is commonly pigmented. Pygid- 
ium with two ventral anal cirri (Fig. 3f). 

Notoacicula fine, from two to three per 
setiger. Neuroacicula large, amber colored, 
three per setiger, tips commonly protruding 
slightly through parapodial wall. Anterior 
setigers with three smooth and from two to 
three toothed setae, medial and posterior 
setigers with two smooth, from one to two 
toothed, and one ventral seta. Shafts on 
ventral setae taper abruptly to guard, two 
ridges present (Fig. 3h). 

Mandibles H-shaped. Maxillae with two 
long carriers and unpaired ventral ligament. 
Maxillae exhibit symmetry polymorphism. 
One or both MI elements robust falcate (Fig. 
51). Denticle formula: 5(8)-6(9); 6(7) or 
9-5(7) or 11(12); 4(6)-4(5); 3(4)-3(4); 1-1. 

Holotype large, complete specimen with 
177 setigers, prostomium 0.39 mm long, 
0.70 mm wide, peristomium 0.39 mm long, 
MIL 0.41 mm long, MIR 0.41 mm long, 
denticle formula 7-9; 6-11; 6-5; 4-4; 1-1. 

Remarks. —This species differs from oth- 
ers in the A. mutans complex in having long- 
er postsetal lobes on medial and posterior 
parapodia, two anal cirri which arise ven- 
trally, and in having two ridges on the ven- 
tral setae. It further differs in having some- 
what longer Mis in individuals of a given 
size (Fig. 7). The post-setal lobe in Arabella 
semimaculata (Moore, 1911) resembles that 
in A. panamensis (compare Fig. 3g; Fau- 
chald 1970, pi. 20, fig. g), but the former 
species has smoothly bilimbate ventral se- 
tae (Fauchald 1970:133), unlike the ridged, 
abruptly tapering ventral setae in A. pana- 
mensis. 

A juvenile specimen from the Pacific side 
of Costa Rica (USNM 80225) with dentate 



VOLUME 102, NUMBER 2 



297 



0-4- 










X 


X 


T 

E 

0-3- 










*" J>i' 


/^ 


•— • 








X 


^r* ' 


* 


of Robust 

o 

1 




X 


X y/' 








Length 

o 9 
i i 


1 1 


1 


1 


1 


1 


1 1 



0-2 03 0-4 0-5 

Width of Prostomium (mm) 



0-6 



07 



0-8 



Fig. 7. Plot of robust MI length against width of prostomium for Arabella multidentata (•), Arabella pana- 
mensis (x), and Arabella sp. A (*). Prostomial width is less subject to distortion by fixation than is total length, 
and is used here as a more reliable measure of specimen size (and, by inference, maturity). The spread of points 
for A. multidentata is typical of that observed for the other species considered here with the exception of A. 
panamensis. The regression line for A. multidentata is of the form y = 0.55x — 0.44 (r = 0.94). Note that all 
of the points for A panamensis plot above the regression line, suggesting that individuals of A panamensis have 
larger jaws for their size than do individuals of A multidentata. The slope of a linear regression for A panamensis 
(not shown) is significantly different from the slope for A multidentata (P = 0.005, t = 3.04, d.f. = 35). Note 
also that plots of points for papillate individuals (Arabella sp. A) fall within the large end of the range of variation 
for A. multidentata. 



MIR has setae comparable to A. panamen- 
sis, but lacks the elongate postsetal lobe in 
posterior parapodia. No data is available on 
ontogenetic changes to be expected in this 
character, and the specimen is only tenta- 
tively attributed to A. panamensis. 

The species is presently known only from 
the Pacific coast of Panama. 



fold (Fig. 3i). The jaws (Figs. 5e, 7) and setae 
are comparable to those in A. multidentata. 
No papillate specimens were recorded from 
any other locality, and the significance of 
the papillae is presently unclear. The char- 
acter is sufficiently distinctive to warrant 
segregation of these forms (informally as sp. 
A) pending further investigation. 



Arabella sp. A 
Figs. 3i, 41, 5e 

Materials examined. — USNM 54216, 
549 1 5, 98239, Guayanilla Bay, Puerto Rico 
(8). 

Remarks. — Three collections of Arabella 
multidentata from Guayanilla Bay included 
several relatively large individuals with two 
distinctive papillae arising from the nuchal 



Discussion 

The systematic treatment here afforded 
members of the Arabella mutans species 
complex should be regarded as a working 
hypothesis. The stability of the taxonomic 
characters employed can be evaluated by 
the collection of additional specimens, and 
data on additional characters (using electro- 
phoresis, for example) can be collected for 



298 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



comparative purposes. Studies on the 
breeding interactions in living populations 
would, of course, provide the most powerful 
test of the systematics, but at the same time 
may be the most challenging to implement. 

Particular emphasis is placed here on 
characterizing species based on populations 
rather than single individuals. Thus, poly- 
morphism of jaw symmetry is considered a 
useful taxonomic character, even though it 
is meaningless in regard to a single individ- 
ual. A single individual collected from a new 
location may thus not be identifiable to the 
species level. 

This restriction especially applies to ju- 
venile individuals. Similar maxillae have 
been recorded from juveniles in four (ques- 
tionably five) of the species of Arabella con- 
sidered here, although the adult maxillae in 
these forms exhibit significant differences. 

Studies of other species of Arabella should 
consider the maturity of specimens before 
maxillary form is included in the species 
diagnosis. In particular, the distinction 
commonly made between Arabella iricolor 
and A. genicnlata (Claparede, 1868) might 
be worth re-evaluation in this light (Fauvel 
1923:439^40, fig. 1751-1; Fauchald 1970: 
226; Ben-Eliahu 1976:172). 

As restricted here, two of the species con- 
sidered are incompletely characterized (Ar- 
abella atlantica, A. logani). The remaining 
five species are separated geographically as 
well as morphologically. The concept of A. 
mutans as a single circumtropical species is 
clearly incorrect. This observation serves to 
underline the futility inherent in attempting 
to use the distribution of rare, poorly under- 
stood species for the study of biogeography. 

The chief lessons for paleontologists 
studying fossil polychaete jaws are as fol- 
lows: 1) jaw symmetry may vary within a 
single population, and should be used as a 
taxonomic character with caution, 2) the 
morphologies of the MI and Mil maxillary 
elements may be useful for species level di- 
agnoses, 3) the denticles on jaws may be 
worth counting after all, but denticle num- 



ber should only be used as a taxonomic 
character on a population basis. The second 
point has already been deduced from the 
study of fossil material (Kielan-Jaworowska 
1966; Edgar 1984), suggesting that classifi- 
cation of recent and fossil material may be 
roughly comparable. The first point has been 
less fully appreciated, and some revision of 
generic and family level fossil taxa may be 
necessary on that basis. 

Acknowledgments 

I thank Kristian Fauchald and Marian H. 
Pettibone for informative discussions con- 
cerning polychaete systematics, and for crit- 
ically reading the manuscript. Alex I. Muir, 
British Museum (Natural History), and 
Thomas H. Perkins, Florida Department of 
Natural Resources, generously arranged the 
loan of type and comparative material. I 
also thank numerous members of the staff 
of the U.S. National Museum of Natural 
History, Smithsonian Institution, for their 
cooperation in the completion of this study. 
Joanne Simpson drafted Figs. 1, 2, 6, and 
7. This research was supported by a post- 
doctoral fellowship, Smithsonian Institu- 
tion. 

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2195 Deborah Way, Upland, California 
91786. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 300-304 

A NEW SPECIES OF BENHAMIPOLYNOE 

(POLYCHAETA: POLYNOIDAE: LEPIDASTHENIINAE) 

FROM AUSTRALIA, ASSOCIATED WITH THE 

UNATTACHED STYLASTERID CORAL 

CONOPORA ADETA 

Marian H. Pettibone 

Abstract.— Benhamipolynoe cairnsi, n. sp. from deep-water (398 m) off 
Queensland, Australia, was found in close symbiotic relationship with the 
unusual, unattached stylasterid coral Conopora adeta Cairns. The coral grows 
around the polynoid, enclosing it in a coiled tube. The species is compared 
with the type species of the genus, B. antipathicola (Benham), from off New 
Zealand, found in latticed tunnels formed by the antipatharian coral Paranti- 
pathes tenuispina (Silberfeld). Both polynoid species are referred to the new 
subfamily Lepidastheninae. 



While working on his description of the 
new species of stylasterid coral from Aus- 
tralia, the unique unattached Conopora ad- 
eta, Cairns (1987) noted the presence of a 
polychaete enclosed in each of his ten coral 
specimens. He brought them to my atten- 
tion and allowed me to extract the polynoids 
from two of the corals, which was no small 
task, since the coral paratypes had to be cut 
with the minimum of damage. On exami- 
nation, the commensal polynoids appeared 
to belong to an undescribed genus and 
species (Cairns 1987:143). After further 
study, they are described below as a new 
species of Benhamipolynoe Pettibone, 1970, 
with type species B. antipathicola, described 
by Benham (1927, under Lepidasthenia) and 
Pettibone (1 970, under the new genus), from 
off New Zealand, associated with the anti- 
patharian coral Parantipathes tenuispina. 

The types are deposited in the Northern 
Territory Museum, Darwin, Australia 
(NTM) and the National Museum of Nat- 
ural History, Smithsonian Institution 
(USNM). 



Family Polynoidae 
Lepidastheniinae, new subfamily 

Types genus: Lepidasthenia Malmgren, 
1867, with type species L. elegans (Grube, 
1840), by monotypy. 

Diagnosis. — Prostomium bilobed, with 3 
antennae and 2 palps; ceratophore of me- 
dian antenna in anterior notch, lateral an- 
tennae inserted terminally on anterior ex- 
tensions of prostomium, on same level as 
median antenna (or slightly subterminally); 
two pairs of eyes on posterior half of pro- 
stomium. First or tentacular segment not 
distinct dorsally; tentaculophores lateral to 
prostomium, each with aciculum and pair 
of dorsal and ventral tentacular cirri, with- 
out setae. Second or buccal segment with 
first pair of elytrophores and elytra, setig- 
erous parapodia and ventral or buccal cirri 
lateral to ventral mouth. Elytra smooth, 
without fringes of papillae, without tuber- 
cles, with or without scattered micropapil- 
lae. Parapodia subbiramous, with small no- 
topodium, notoaciculum, and without, or 



VOLUME 102, NUMBER 2 



301 



with very few, notosetae. Neuropodia with 
anterior and posterior lobes deeply cut dor- 
sally and ventrally, without projecting acic- 
ular lobes. Dorsal cirri on non-elytrigerous 
segments with basal cirrophores and distal 
styles; dorsal tubercles indistinct. Pharnyx 
with 9-13 pairs of border papillae and 2 
pairs of chitinous jaws. 

Remarks.— Lepidasthenia has been in- 
cluded in the Subfamily Lepidonotinae Wil- 
ley, 1902, chiefly on the basis of the simi- 
larity of the prostomium to Lepidonotus 
Leach, 1 8 1 6. It is separated herein based on 
the different types of parapodia and elytra. 
Also included in Lepidastheniinae are the 
following polynoid genera: Perolepis Ehlers, 
1908; Parahalosydna Horst, 1915; Hyper- 
halosydna Augener, 1922; Alentiana Hart- 
man, 1942; Telolepidasthenia Augener & 
Pettibone, 1970; Benhamipolynoe Petti- 
bone, 1970. A new species is described be- 
low under the last named genus. 

Benhamipolynoe Pettibone, 1970 

Type species: Lepidasthenia antipathicola 
Benham, 1927, by original designation. 

Diagnosis. — Body elongate, slender, flat- 
tened, segments numerous (more than 50, 
up to 200 or more). Elytra 10 or more pairs, 
on segments 2, 4, 5, 7, alternate segments 
to 15, then variable in arrangement, with 
either long posterior region without elytra 
or with elytra continuing posteriorly. Elytra 
without fringes of papillae, smooth except 
for scattered micropapillae. Prostomium 
lepidasthenoid, bilobed, with two palps and 
three antennae; ceratophore of median an- 
tenna in anterior notch of prostomium, lat- 
eral antennae inserted terminally on ante- 
rior extensions of prostomium, lateral to 
median antenna, with two pairs of eyes on 
posterior half of prostomium. Tentaculo- 
phores of tentacular segment lateral to pro- 
stomium, without setae, with two pairs of 
dorsal and ventral tentacular cirri. Second 
or buccal segment with first pair of elytra 



and long ventral buccal cirri; without nuchal 
lobe. Parapodia subbiramous, with small 
conical notopodia on anterodorsal faces of 
neuropodia, without notosetae. Neuropo- 
dia deeply cut dorsally and ventrally, with 
anterior and posterior subequal rounded 
lobes, without projecting acicular lobes. 
Neurosetae relatively few in number (5-12), 
of single type, rather stout, smooth or with 
slight indication of spinous rows, with fal- 
cate entire tips. Dorsal cirri with short cy- 
lindrical cirrophores and long, smooth, ta- 
pering styles; dorsal tubercles indistinct. 
Ventral cirri short, subulate. Pygidium with 
paired long anal cirri. Nephiridial papillae 
short, cylindrical, beginning on segment 8. 

Remarks. —Benhamipolynoe differs from 
Lepidasthenia Malmgren in regard to the 
number and arrangement of the elytra and 
the types of neurosetae. In Lepidasthenia, 
the neurosetae have well-developed short 
and wide spinous regions and bifurcate tips; 
in Benhamipolynoe, the neurosetae have 
spinous regions scarcely developed or lack- 
ing and entire falcate tips. 

Benhamipolynoe antipathicola (Benham, 
1927), originally described as Lepidasthen- 
ia, from off New Zealand in 1 28 meters, was 
associated with Parantipathes tenuispina 
(Silberfeld), where it was found in latticed 
tunnels formed by the serrated branchlets 
of the antipatharian. Additional specimens 
referred to B. antipathicola by Pettibone 
(1 970) include one from the Gulf Stream off 
Key West, Florida, in 237 meters, associ- 
ated with Antipathes columnaris Duchas- 
saing, and one from Indonesia, in 520 me- 
ters, found in an empty tube of a eunicid 
polychaete. 

Benhamipolynoe cairnsi, new species 
Figs. 1, 2 

Material.— Queensland, Australia, Mar- 
ion Plateau off MacKay, 20°46.2'S, 
152°51.8'E, 398-399 m, F.R.V. Soela sta 
25, 22 Nov 1985, A. Bruce, collector, re- 
moved from coiled galleries in stylasterid 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Benhamipolynoe cairnsi, A, C-H, Holotype, NTM W4900; B, Paratype, USNM 1 18373: A, Dorsal 
view of anterior end, styles of median and right lateral antennae, left dorsal and ventral tentacular cirri all 
missing; right parapodium and dorsal cirrus of segment 3 not shown; B, Ventral view of anterior end, styles of 
median and left lateral antennae, left palp, and all tentacular cirri missing; C, Right elytrigerous parapodium 
of segment 2, posterior view, acicula and small notopodium dotted, elytrophore not shown; D, Right cirrigerous 
parapodium of segment 3, anterior view, acicula dotted; E, Neurosetae from same; F, Right 1st elytron from 
segment 2, with few micropapillae (not to scale); G, Right 2nd elytron from segment 4; H, Right 9th elytron 
from segment 19. Scales = 1.0 mm for A. B; 0.5 mm for C, D, F-H; 0.1 mm for E. 



coral Conopora adeta Cairns, holotype 
(NTM W4900) and paratype (USNM 
118373). 

Description.— Holotype (in two pieces, 
incomplete posteriorly) with 38 segments, 
16 mm long, 2 mm wide; paratype (in three 
pieces, complete) with 63 segments, 33 mm 
long, 3 mm wide; both specimens females, 
full of large yolky eggs. 



Body elongate, flattened ventrally, arched 
dorsally, widest in anterior half, tapering 
gradually posteriorly; without color except 
for black eyes. Prostomium oval, bilobed; 
ceratophore of median antenna large, oval, 
in anterior notch, style missing; lateral an- 
tennae inserted terminally on anterior con- 
tinuations of prostomium, lateral to median 
antenna, with long styles about equal to stout 



VOLUME 102, NUMBER 2 



303 




Fig. 2. Benhamipolynoe cairnsi, Holotype, NTM W4900: A, Right cirrigerous parapodium from segment 
1 2, posterior view, extension on anterior side of cirrophore dotted; B, Right cirrigerous parapodium from segment 
21, posterior view; C, Upper and middle neurosetae from same, latter with tip worn; D, Right cirrigerous 
parapodium from segment 35, anterior view, acicula dotted; E, Upper and middle neurosetae from same. Scales 
= 0.5 mm for A, B, D; 0.1 mm for C, E. 



tapering palps; two pairs of subequal black 
eyes on posterior half of prostomium (Fig. 
1 A, B). First or tentacular segment not vis- 
ible dorsally; tentaculophores lateral to pro- 
stomium, without setae, each with pair of 
tentacular cirri, ventral one about as long 
as lateral antennae, dorsal one longer than 
palps; ventrally with bulbous facial tubercle 
(Fig. 1 A, B). Second or buccal segment with 
slightly developed nuchal lobe, first pair of 
elytrophores, subbiramous parapodia with 
few neurosetae, and ventrally with long buc- 
cal cirri lateral to ventral mouth (Fig. 1A- 
C). Segment 3 with first pair of dorsal cirri; 
cirrophores longer than neuropodia, cylin- 
drical with distal bulbous extension and long 
style; ventral cirri almost as long as buccal 
cirri of segment 2 (Fig. 1A, B, D). 

Elytra and elytrophores 10 pairs on seg- 
ments 2-4, 5-7-9-1 1-13-15- 19-23, with 
dorsal cirri on all segments from 24 on. Ely- 
tra oval, attached eccentrically near lateral 
side, delicate, with "veins" and scattered 
round micropapillae on surface (Fig. 1 F-H). 

Parapodia subbiramous; notopodia in 
form of small conical acicular lobes on 



anterodorsal faces of larger neuropodia, 
without notosetae; neuropodia rather deep- 
ly cut dorsally and ventrally, forming 
rounded presetal and postsetal lobes, with 
row of 5-10 short neurosetae (Figs. 1C, 2 A, 
B, D). Neurosetae stout, with faint spinous 
rows and slightly hooked entire tips, some- 
times slightly worn (Figs. IE, 2C, E). Cir- 
rophores of dorsal cirri cylindrical, extend- 
ing beyond neuropodia, with triangular 
extension on lower side, with long, smooth 
styles tapering to slender tips and extending 
far beyond parapodia; ventral cirri wider 
basally, with long slender tips, extending 
slightly beyond tips of neuropodia (Fig. 2 A, 
B, D). 

Nephridial papillae beginning on segment 
8, small at first, becoming longer about seg- 
ment 15, cylindrical, directed dorsally be- 
tween parapodia. Pharynx not extended and 
not examined. Pygidium forming small lobe 
with dorsal anus between last pair of small 
parapodia and pair of long anal cirri, similar 
to dorsal cirri. 

Biology.— The two symbiont polynoids 
were removed from two paratypes of the 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



colonial stylasterid coral Conopora adeta, 
well figured by Cairns (1987. figs. 1-6). The 
coral is unusual in that it is unattached, 
formed of a massive globose solid base with 
several (3-6) stout, radiating, stabilizing 
branches and two main ascending porous 
branches. The polynoid occupied a coiled 
runway in the globose base connected to a 
U-shaped tube in the vertical branches. The 
close symbiotic relationship of the polynoid 
with the coral begins at an early stage in 
development and determines the morphol- 
ogy of the colony. One of the openings near 
the base of the colony was large enough for 
the polynoid to be able to protrude from the 
tube. All of the ten coral colonies examined 
by Cairns contained a polynoid symbiont. 
Both specimens of Benhamipolynoe cairnsi 
were females, full of large yolky eggs. 

Remarks.— Benhamipolynoe cairnsi dif- 
fers from B. antipathicola in having only 10 
pairs of elytra and only dorsal cirri from 
segment 24 on. compared to numerous pairs 
of elytra continuing to the posterior end. In 
B. cairnsi, the cirrophores of the dorsal cirri 
have unique extra extensions: bulbous on 
segment 3 and triangular on the following 
cirrophores; this condition was not found 
on B. antipathicola and has not been noted 
previously on other polynoids. The two 
species differ in their positions with their 
respective coral hosts. As indicated above, 
the coiled runway of B. cairnsi is enclosed 
in the stylasterid coral Conopora adeta, al- 
lowing only limited movement to the ex- 
terior. For B. antipathicola, the antipathar- 
ian coral host. Parantipathes tenuispina, is 
a bottle-brush shaped colony where the in- 
terlacing branchlets form a complicated lat- 
tice-work tube that serves as an elongated 



"worm-run," described by Totton (1923: 
106, fig. 9), thus allowing much greater ex- 
ternal movement for the polynoid com- 
mensal. 

Etymology'. — The species is named for 
Stephen D. Cairns, who described and 
named the stylasterid coral host of the po- 
lynoid symbiont. 

Acknowledgments 

My thanks go to Stephen Cairns for bring- 
ing the polynoid commensal to my atten- 
tion and for allowing me to extract them 
from the stylasterid coral. The manuscript 
benefited from the reviews of Russel Hanley 
and Stephen Cairns. 

Literature Cited 

Benham. W. B. 1972. Polychaeta. — British Antarctic 
"Terra Nova" Expedition. 1910. Natural His- 
tory Reports. Zoology 7(2):47-182. 

Cairns, S. D. 1987. Conopora adeta, new species (Hy- 
drozoa: Stylasteridae) from Australia, the first 
known unattached stylasterid.— Proceedings of 
the Biological Society of Washington 1 00( 1): 141— 
146. 

Pettibone, M. H. 1970. Polychaeta errantia of the 
Siboga-Expedition. Part 4: Some additional 
polychaetes of the Polynoidae, Hesionidae, Ne- 
reidae. Goniadidae. Eunicidae. and Onuphidae. 
selected as new species by the late Dr. Hermann 
Augener with remarks on other related species. — 
Siboga-Expedite 24, Id: 199-270. 

Totton, A. K. 1923. Coelenterata. Part III. Antipa- 
tharia (and their Cirripede Commensals). —Brit- 
ish Antarctic ("Terra Nova") Expedition. 1910. 
Natural History Report. Zoology 5(3):97-120. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution. Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
102(2), 1989, pp. 305-310 

TWO NEW SPECIES OF HARMOTHOINAE 

(POLYCHAETA: POLYNOIDAE) FROM 

THE EAST PACIFIC RISE, COLLECTED BY 

ALVIN DIVES 2000 AND 2003 

Marian H. Pettibone 

Abstract.— Alvin dives in the East Pacific Rise in 2700 meters near 11°N 
collected two polynoid species of Harmothoinae: Harmothoe hollisi, n. sp. from 
dive 2000, associated with a "wood island," and Eunoe alvinella n. sp. from 
dive 2003, collected from an inactive sulfide chimney. 



As a part of the dive series of the DSRV 
Alvin on the East Pacific Rise, during Dive 
2000 in March 1988, near 1 1°51'N in 2750 
meters, the pilot picked up a piece of wood 
that was densely covered with biota, in- 
cluding serpulid worms, gastropods, gala- 
theids, ophiuroids, and additional fauna 
(Van Dover 1988). Among the latter were 
two polynoid polychaetes sent to me for 
study by Cindy Lee Van Dover of the Woods 
Hole Oceanographic Institution. The col- 
lection site was in a field of lightly sedi- 
mented pillow basalt, within a few kilo- 
meters of known hydrothermal vent 
communities. The fauna of this "wood is- 
land" is of particular interest in comparing 
it with the fauna associated with the hydro- 
thermal vents. 

The polynoid is described below as a new 
species of Harmothoe, H. hollisi, a widely 
distributed genus in Harmothoinae. A large 
polynoid, collected by Dive 2003 on an in- 
active sufide chimney, is described as a new 
species of Eunoe, E. alvinella, also a widely 
distributed genus is the same subfamily. 
Previously reported species of Harmothoe, 
associated with the "wood island" and panel 
experiments by Turner (1973, 1981) in the 
deep water of the North Atlantic, included 
two species described by Pettibone (1985a): 
H. ingolfiana Ditlevsen and H. vagabunda 
Pettibone. Harmothoe macnabi Pettibone 
was described by Pettibone (1985b) from 



Alvin dive 895 in the Galapagos Rift. The 
above three species of Harmothoe, as well 
as the new species, lack eyes, as character- 
istic of most deep sea polynoids. The new 
species of Eunoe differs from other species 
of Eunoe in having very large eyes, occu- 
pying the lateral borders of the prostomium. 

Subfamily Harmothoinae Horst, 1917 

Genus Harmothoe Kinberg, 1856 

Harmothoe hollisi, new species 

Figs. 1, 2 

Material— East Pacific Rise, Alvin dive 
2000, 22 Mar 1988, 1 1°52'N, 103°51'W, 
2750 m, "wood island," holotype (USNM 
1 18867) and paratype (USNM 1 18868). 

Description.— Length of complete holo- 
type 24 mm, width with setae 1 5 mm, seg- 
ments 37, last one minute. Length of in- 
complete paratype 15 mm, width 7 mm, 
segments 2 1 . Without color except for gold- 
en setae. 

Body flattened, subrectangular. tapered 
slightly anteriorly and posteriorly. Elytra 1 5 
pairs, on large prominent elytrophores on 
segments 2, 4, 5, 7, alternate segments to 
23, 26, 29, and 32 (Figs. 1A, 2A). Elytra 
large, overlapping, covering dorsum, oval 
to subreniform in shape, delicate (tear eas- 
ily), densely covered with conical micro- 
tubercles, smaller on anterior part, becom- 
ing gradually larger on posterior and lateral 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Harmothoe hollisi, A, B. D, holotype (USNM 118867); C, paratype (USNM 118868): A, Anterior 
end. dorsal view, palps missing: B. Same, ventral view: C. Anterior end. dorsal view, pharynx partially extended, 
right palp and right ventral tentacular cirrus shorter, regenerating, right dorsal tentacular cirrus missing; D. Right 
5th elytron from segment 9, with detail of microtubercles and papillae (not to scale). Scales =1.0 mm for A- 
C: 2.0 mm for D. 



borders; larger tubercles cylindrical or ovoid, 
with tips entire or with 2-8 nodular exten- 
sions; scattered filiform papillae on surface 
and lateral and posterior borders (Fig. ID). 
Non-elytra-bearing segments with dorsal 
cirri and nodular to digitiform dorsal tu- 
bercles (Figs. 1A, 2B). 

Prostomium deeply bilobed. with distinct 
cephalic peaks, tending to curl inward, with- 
out eyes; median antenna with large bul- 
bous ceratophore in anterior notch and long 
papillate style; lateral antennae with short 
bulbous ceratophores inserted ventrally, 
with styles short, bulbous and papillate ba- 
sally. and long filamentous tips; palps (miss- 
ing on holotype) stout, tapered, smooth (Fig. 



1A-C). Tentaculophores lateral to prosto- 
mium, each with small acicular process and 
2-3 short notosetae on inner side, and dor- 
sal and ventral tentacular cirri similar to 
median antenna (Fig. 1A-C). Second or 
buccal segment with first pair of large elytro- 
phores. biramous parapodia. and long ven- 
tral buccal cirri inserted basally on neuro- 
podia lateral to ventral mouth, similar to 
tentacular cirri (Fig. 1A, B). 

Parapodia biramous. with golden setae. 
Notopodia rounded basally, with projecting 
acicular lobe on lower side, almost as long 
as neuropodia (Fig. 2A, B). Notosetae nu- 
merous, forming radiating bundle, slightly 
curved, about four different lengths, shorter 



VOLUME 102, NUMBER 2 



307 




Fig. 2. Harmothoe hollisi, holotype (USNM 118867): A, Right elytrigerous parapodium from segment 1 1, 
anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 12, posterior view; C Long and 
short notosetae; D, Subacicular neurosetae, with detail of tips; E, Supraacicular neuroseta, with detail of tip. 
Scales =1.0 mm for A, B; 0.1 mm for C-E. 



than and much stouter than neurosetae; no- 
tosetae with numerous spinous rows and 
rather long, tapered bare tips (Fig. 2C). Cy- 
lindrical cirrophores of dorsal cirri on dor- 
soposterior sides of notopodia, with slender 
papillate styles extending beyond neurose- 
tae (Fig. 2B). Neuropodia with conical pre- 
setal lobe and projecting acicular process 
with small supraacicular extension, and 
shorter rounded postsetal lobe (Fig. 2A, B). 
Neurosetae numerous, forming fan-shaped 
bundle, with numerous spinous rows. Su- 



praacicular neurosetae more slender, with 
long spinous regions and bare split tips (Fig. 
2E). Subacicular neurosetae stouter, with 
shorter spinous regions, and slightly hooked 
bare tips, all with rather long secondary tooth 
(Fig. 2D). Ventral cirri short, tapered, with 
abruptly narrowed distal tip (Fig. 2A, B). 

Pharynx not extended and not examined. 
Inflated ventral nephridial areas with small 
papilla on posterior side, beginning on seg- 
ment 6 (Fig. 2B). Pygidium forming small 
lobe between small parapodia of 2 posterior 



Table 1. — Comparison of Harmothoe ingolfiana and Harmothoe hollisi. 



H. ingolfiana 



H. hollisi 



Cephalic peaks of prostomium 
Elytral macrotubercles 
Neurosetae 



small 
present 

upper ones with bifid tips, middle 
and lower ones with entire tips 



prominent, tending to curl inward 

absent 

all with bifid tips 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Eunoe alvinella, holotype (USNM 1 1 8866): A, Anterior end, dorsal view, pharynx partially extended, 
styles of median antenna, left lateral antenna, right and left dorsal and ventral tentacular cirri missing; B, Left 
2nd elytron from segment 4, with detail of microtubercles and papillae (not to scale); C, Left middle elytron, 
with detail of microtubercles and papillae (not to scale). Scale = 1.0 mm for A-C. 



segments, with pair of anal cirri (styles miss- 
ing). 

Etymology.— The species is named for 
Ralph Hollis, the pilot of Alvin dive 2000, 
who collected the interesting "wood island" 
with its associated biota. 

Comparison.— Harmothoe hollisi agrees 
in most respects with H. ingolflana Ditlev- 
sen, reported from the North Atlantic south 
of Iceland to off New England in 1 830-3506 
meters and found in burrows of wood-bor- 
ing bivalves (Ditlevsen 1917:32, Pettibone 
1985a: 143). The two species are compared 
in Table 1 . 



Genus Eunoe Malmgren, 1865 

Eunoe alvinella, new species 

Figs. 3, 4 

Material— East Pacific Rise, Alvin dive 
2003, 25 Mar 1988, 11°46'N, 104°47'W, 
2725 m, inactive sulfide chimney, sample 
no. 9, holotype (USNM 118866). 

Description. —Length of complete holo- 
type 41 mm, width with setae 14 mm, seg- 



ments 34, last one minute. Dorsum light 
brown, with colorless transverse bands. 

Body elongate-oval, slightly tapered an- 
teriorly and more so posteriorly, flattened 
ventrally, arched dorsally. Elytra 15 pairs, 
on large bulbous elytrophores on segments 
2, 4, 5, 7, alternate segments to 23, 26, 29, 
and 32 (Figs. 3 A, 4 A, C). Elytra large, over- 
lapping, covering dorsum, oval to subreni- 
form in shape, thickly covered with rusty 
yellow and brown foreign material; elytral 
surface with microtubercles on most of sur- 
faces large globular macronodules on pos- 
terior third of elytra, variable in number 
and size, and filiform papillae on lateral bor- 
der and scattered on surface (Fig. 3B, C). 
Non-elytra-bearing segments with dorsal 
cirri and nodular dorsal tubercles (Fig. 4B, 
D). 

Prostomium deeply bilobed, wider than 
long, without distinct cephalic peaks, with 
two pairs of very large eyes occupying lateral 
sides of prostomium; median antenna with 
large bulbous ceratophore in anterior notch, 
style missing (probably long); lateral anten- 



VOLUME 102, NUMBER 2 



309 




Fig. 4. Eunoe alvinella, holotype (USNM 118868): A, Right elytrigerous parapodium from segment 2, 
anterior view, acicula dotted, style of ventral buccal cirrus missing; B, Right cirrigerous parapodium from 
segment 3, posterior view, style of dorsal cirrus missing; C, Right elytrigerous parapodium from segment 13, 
anterior view, acicula dotted, style of ventral cirrus hidden from view; D, Right cirrigerous parapodium from 
segment 14, posterior view; E, Short and long notosetae; F, Supraacicular neuroseta; G, Middle and lower 
subacicular neurosetae. Scales = 1.0 mm for A-D; 0.1 mm for E-G. 



nae with short bulbous ceratophores in- 
serted ventrally, styles short, tapered; palps 
stout, tapered, about one and a half times 
length of prostomium (Fig. 3A). Tentacu- 
lophores lateral to prostomium, each with 
projecting aciculum and one or two noto- 
setae on inner side, and dorsal and ventral 
tentacular cirri, styles missing (probably 
long); bulbous facial tubercle medial to cer- 



atophores of lateral antennae (Fig. 3 A). Sec- 
ond or buccal segment with first pair of large 
elytrophores, biramous parapodia; cerato- 
phores of ventral buccal cirri inserted ba- 
sally on neuropodia lateral to ventral mouth, 
styles missing (probably longer than follow- 
ing ventral cirri) (Figs. 3A, 4A). 

Parapodia biramous, with golden setae. 
Notopodia smaller and shorter than neu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ropodia (Fig. 4C, D). Notopodium wider 
basally, tapering to digitiform acicular lobe 
on lower side, tip of notoaciculum project- 
ing (Fig. 4C). Notosetae moderate in num- 
ber, much stouter than neurosetae, upper 
ones shorter, curved, lower ones longer, 
nearly straight, both with numerous spinous 
rows and rather short bare blunt tips (Fig. 
4E). Cylindrical cirrophores of dorsal cirri 
on dorsoposterior sides of notopodia, with 
long papillate styles extending far beyond 
neurosetae (Fig. 4D). Neuropodium with 
conical presetal acicular lobe and projecting 
acicular process, tip of neuroaciculum pro- 
jecting; postsetal lobe shorter, rounded (Fig. 
4A-D). Neurosetae numerous, with spinous 
rows. Supraacicular neurosetae slender, with 
long spinous regions, tapering to slender 
straight bare tips (Fig. 4F); middle and low- 
er subacicular neurosetae stouter, with 
shorter spinous regions and slightly hooked 
bare tips (Fig. 4G). Ventral cirri short, ta- 
pered (Fig. 4D). 

Pharynx not extended and not examined. 
Nephridial papillae small, beginning on seg- 
ment 4. Pygidium rectangular lobe between 
small parapodia of last segment, with pair 
of anal cirri (styles missing). 

Etymology. —The species is named for the 
collecting vessel, the DSRV Alvin. 

Comparisons.— E. alvinella differs from 
other species of Eunoe in having very large 
eyes occupying the lateral sides of the pro- 
stomium. The elytra, with their unique type 
of numerous globular macronodules, also 
differ from other species of the genus. 



as well as the pilot and observers of the 
submersible Alvin, on their celebration of 
dives 2000 and 2003, for the material on 
which this study is based. The manuscript 
benefited from the reviews of James A. 
Blake, J. Fred Grassle, and Cindy Lee Van 
Dover. 

Literature Cited 

Ditlevsen, Hj. 1917. Annelides I.— The Danish In- 
golf-Expedition, Copenhagen 4(4): 1-71. 

Pettibone, M. H. 1985a. Polychaete worms from a 
cave in the Bahamas and from experimental 
wood panels in deep water of the North Atlantic 
(Polynoidae: Macellicephalinae, Harmothoi- 
nae). — Proceedings of the Biological Society of 
Washington 98(1): 127-149. 

. 1985b. New genera and species of deep-sea 

Macellicephalinae and Harmothoinae (Poly- 
chaeta: Polynoidae) from the hydrothermal rift 
areas off the Galapagos and western Mexico at 
21°N and from the Santa Catalina Channel.— 
Proceedings of the Biological Society of Wash- 
ington 98(3):740-757. 

Turner, R. D. 1973. Wood-boring bivalves, oppor- 
tunistic species in the deep sea.— Science 180: 
1377-1379. 

. 1981. "Wood islands" and "thermal vents" 

as centers of diverse communities in the deep 
sea.— The Soviet Journal of Marine Biology 7(1): 
1-9. 

Van Dover, C. L. 1988. Alvin Dive 2000. -Sea Fron- 
tiers, 34(6):326-331. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 



Acknowledgments 

I wish to thank Cindy Lee Van Dover of 
the Woods Hole Oceanographic Institution, 



PROC. BIOL. SOC. WASH. 
102(2), 1989, pp. 311-312 

VARICHAETADRILUS ANGUSTIPENIS 

(BRINKHURST AND COOK, 1966), 

NEW COMBINATION FOR LIMNODRILUS 

ANGUSTIPENIS (OLIGOCHAETA: TUBIFICIDAE) 

Ralph O. Brinkhurst 

Abstract. —Limnodrilus angustipenis Brinkhurst & Cook, 1966 is recognized 
as a member of the genus Varichaetadrilus Brinkhurst & Kathman, 1983. 



Re-examination of some of the type spec- 
imens of Limnodrilus angustipenis Brink- 
hurst & Cook, 1966 reveals that this taxon 
should be attributed to the genus Varichae- 
tadrilus Brinkhurst & Kathman, 1983. Five 
specimens of the original set collected from 
Saginaw Bay, Lake Huron in 1956 by the 
former United States Bureau of Commer- 
cial Fisheries Laboratory of Ann Arbor, 
Michigan, were examined. These were orig- 
inally donated to the author by J. K. Hil- 
tunen, now of Sugar Island, Michigan. 

As the species is usually identified by ref- 
erence to the shape of the somewhat elon- 
gate penis sheaths, very few if any dissec- 
tions or sections appear to have been 
prepared from this species. Furthermore, the 
male ducts have never been fully illustrated. 
The species was originally described in the 
third part of a series of publications on the 
North American aquatic Oligochaeta 
(Brinkhurst & Cook 1966) in which a few 
tubificid descriptions were added to a paper 
largely directed to the Family Lumbriculi- 
dae. No follow-up description has been 
published. 

The male ducts of one of the five speci- 
mens at hand are visible as a series of frag- 
ments from a dissection. 

The union of the vas deferens and atrium 
and the origin of the prostate gland close to 
this point is evident on both male ducts. 
The form of these could well be mistaken 
for those of Limnodrilus. Beyond this point, 
however, the atrium does not narrow to form 



an ejaculatory duct, but remains wide all 
the way to the base of the penis sheath on 
each side (Fig. 1). This unique construction 
of the male ducts is typical of Varichaeta- 
drilus. Neither of these male ducts is com- 
plete, and so the total length cannot be es- 
timated, but they would appear to be more 
like those of V. psammophilus (Loden, 1977) 
and V. minutus (Brinkhurst, 1965) than 
those of V. pacificus (Brinkhurst, 1981) or 




Fig. 1 . Fragmented male ducts and spermatheca of 
Varichaetadrilus angustipenis drawn with the aid of a 
drawing tube from a paratype in the author's collection. 
Scale in mm. a— atrium, p— prostate stalk, s— penis 
sheath, sp— spermatheca, sz— spermatozeugmata, v— 
vas deferens. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




o 01 



Fig. 2. Chaetae from segments II and VI of the 
dissected specimen, posterior chaetae from another 
whole mount. Scale in mm. dll— dorsal chaetae of II, 
dVI— dorsal chaetae of VI, vVI— ventral chaetae of VI, 
pv— posterior ventral chaetae. 



V. fulleri Brinkhurst & Kathman, 1983, as 
illustrated by Brinkhurst (1986). 

In V. angustipenis the penis sac enclosing 
the long cuticular penis sheath lacks the spi- 
ral muscles diagnostic of the genus Lim- 
nodrilus s.s. (the positions of Limnodrilus 
grandisetosus Nomura, 1932 and Limnod- 
rilus silvani Eisen, 1879, might be chal- 
lenged in regard to this feature). The usual 
erectile penes of Varichaetadrilus appear to 
be reduced to the short conical structures in 
the bases of the penis sheaths in this species. 



In the original account, no mention was 
made of penial chaetae, and these do not 
seem to be modified on the specimens ex- 
amined here. The penial chaetae are mod- 
ified in V. pacificus and V. minutus (=V. 
nevadanus) but not in V. fulleri or V. psam- 
mophilus. Varichaetadrilus Israelis Brink- 
hurst, 1971 appears to lack them, too, but 
this requires confirmation. 

The somatic chaetae have reduced upper 
teeth in anterior bundles (Fig. 2) but the 
upper teeth are longer than the lower in most 
bundles, though they may vary within a 
bundle. There are mostly five chaetae in an- 
terior bundles, but only one or two poste- 
riorly. The upper teeth are especially short 
in chaetae from II to V. 

The form of the penis sheath, and in par- 
ticular its length, is diagnostic for this 
species. 

Literature Cited 

Brinkhurst, R. O. 1986. Guide to the freshwater 
aquatic microdrile oligochaetes of North Amer- 
ica.— Canadian Special Publication of Fisheries 
and Aquatic Sciences 84:1-259. 

, & D. G. Cook. 1966. Studies on the North 

American aquatic Oligochaeta. Ill: Lumbricu- 
lidae and additional notes and records of other 
families. — Proceedings of the Academy of Nat- 
ural Sciences of Philadelphia 118:1-33. 

Institute of Ocean Sciences, P.O. Box 
6000, Sidney, British Columbia V8L 4B2, 
Canada. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 313-323 

THE ADULT MALE OF THE TROGLOBITIC OSTRACODE 

SPELAEOECIA BERMUDENSIS ANGEL AND ILIFFE, 1987, 

FROM AN ANCHIALINE CAVE IN BERMUDA 

(CRUSTACEA: OSTRACODA: HALOCYPRIDOIDEA) 

Louis S. Kornicker 

Abstract.— The previously unknown adult male of Spelaeoecia bermudensis 
Angel & Iliffe, 1987, the type species of the monotypic genus Spelaeoecia Angel 
& Iliffe, 1987, is described and illustrated from a specimen collected in Won- 
derland Cave, Bermuda. The fifth limb bears a large sensory organ that is 
absent on the limbs of the adult female as well as the juvenile male, and has 
not been reported previously in the suborder Halocypridina. The rod-shaped 
posterior branch of the copulatory organ is not inside the canal of the broad 
anterior branch as in other genera of the superfamily Halocypridoidea. The 
separated anterior and posterior branches resemble those of the copulatory 
organ of adult males in the superfamily Thaumatocypridoidea, which together 
with the Halocypridoidea form the suborder Halocypridina. A brief supple- 
mentary description is given of the adult female. 



The species Spelaeoecia bermudensis was 
described by Angel & Iliffe (1987:545) from 
56 specimens (22 adults, but 2 of the adults 
represented by carapaces only) collected in 
8 anchialine caves in Bermuda. Angel & 
Iliffe (p. 552) stated, "The absence of males 
in the collections could indicate that the 
species breeds by parthenogenesis, but it 
seems more likely that breeding occurs in 
parts of the cave inaccessible to divers." 
Kornicker & Iliffe (1989) reported from 2 
anchialine caves in Bermuda 1 additional 
specimens including 2 A- 1 males, which they 
described briefly. They mentioned an adult 
male collected in Wonderland Cave, Ber- 
muda, which was encountered too late to 
conveniently include in their paper. That 
male is described herein. It was collected 
along with one juvenile and nine adult fe- 
males in a baited trap by Jan H. Stock in 
1984. Angel & Iliffe (1987:tab. 2) reported 
three adults (two represented by carapaces 
only) from the same cave. Unless the ratio 
of adult females to males in the species is 
unusually high it seems likely, as suggested 



by Angel & Iliffe (1987), that adult females 
are more widespread than adult males. Per- 
haps the adult male described herein was 
attracted from some distance by the bait in 
the trap, and trapping rather than netting 
may be a more efficient method for collect- 
ing males. A brief supplementary descrip- 
tion is presented based on a study of the 
holotype. 

Superorder Myodocopa Sars, 1866 

Composition. —This superorder compris- 
es the orders Myodocopida Sars, 1866, and 
Halocyprida Dana, 1853. 

Order Halocyprida Dana, 1853 

Composition. — This order comprises the 
suborders Cladocopina Sars, 1866, and Hal- 
ocypridina Dana, 1853. 

Suborder Halocypridina Dana, 1853 

Composition. —This suborder comprises 
the superfamilies Halocypridoidea Dana, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



1853, and Thaumatocypridoidea Muller, 
1906. 

Superfamily Halocypridoidea 
Dana, 1853 

Composition. —This superfamily con- 
tains the family Halocyprididae Dana, 1853. 

Family Halocyprididae Dana, 1853 

Composition.— This family comprises the 
subfamilies Halocypridinae Dana, 1853, 
Conchoecinae Claus, 1891, Archiconchoe- 
cinae Poulsen, 1969, Euconchoecinae Poul- 
sen, 1969, and Deeveyinae Kornicker & II- 
iffe, 1985. 

Subfamily Deeveyinae 
Kornicker & Iliffe, 1985 

Composition. —The subfamily comprises 
the genera Deeveya Kornicker & Iliffe, 1985, 
and Spelaeoecia Angel & Iliffe, 1987. 

Spelaeoecia Angel & Iliffe, 1987 

Type species.— Spelaeoecia bermudensis 
Angel & Iliffe, 1987. 

Composition. —The genus is monotypic. 

Terminology. —The lettering of bristles on 
the first and second antennae is the same as 
for the genus Deeveya explained in Kor- 
nicker and Palmer (1987:6 10). On the eighth 
joint of the first antenna of Deeveya the base 
of the d-bristle is slightly more lateral than 
on Spelaeoecia. Therefore, it is proposed 
that the anterior bristle regardless of wheth- 
er it is lateral or medial be designated the 
d-bristle; the definitely lateral bristle near 



midwidth of the joint and posterior to the 
d-bristle and stouter than other bristles on 
the joint be designated the e-bristle; the def- 
initely medial bristle near midwidth of the 
joint and with proximal part slightly angled 
ventrally be designated the f- bristle, and the 
posterior bristle that is lateral to the f-bristle 
and either slightly medial to, or at same 
level, as the e-bristle be designated the 
g-bristle (Fig. 2a). The e-bristle is the "prin- 
cipal bristle." The terminology of the sixth 
limb is that suggested by Angel & Iliffe (1 987: 
551) in which the joint previously inter- 
preted to be the first exopodial joint is now 
designated the basale, and a terminal dorsal 
process on that joint is now designated the 
endopodite. 

Spelaeoecia bermudensis 

Angel & Iliffe, 1987 

Fig. 1-5 

Spelaeoecia bermudensis Angel & Iliffe, 
1987:545, figs. 2-6. -Kornicker & Iliffe, 
1989, fig. 29. 

Holotype. -USNM 228468, adult female 
on five slides. 

Type locality.— Green Bay Cave, Ber- 
muda. 

Material— Wonderland Cave, Bermuda: 
USNM 193483, one adult male on slide and 
in alcohol; USNM 193484A, one adult fe- 
male in alcohol; USNM 193484B, one ju- 
venile (length 1.27 mm, height 0.74 mm) 
plus nine adult females, all in alcohol; all 
collected 4 Oct 1984 in baited trap at 0-1 
m depth by Dr. Jan H. Stock. Also exam- 
ined: holotype, and 2 A-l males (USNM 
1 93404A, 1 93405A) from Jane's Cave, Ber- 



Fig. 1. Spelaeoecia bermudensis, adult male, USNM 193483: a, Lateral view of specimen from left side 
showing representative surface lineations and some internal characters, length 1.37 mm; b, Dorsal view of 
carapace with valves partly open (dashed lines indicate ventral edges of valves); c, Detail of posterodorsal corner 
of carapace, from "a"; d, Detail of posterior of right valve; e, Specimen showing relationship of appendages 
(Bellonci Organ, furca, copulatory organ, and appendages of left side; not all bristles of appendages shown); f, 
Bellonci Organ and joints 1 and 2 of left 2nd antenna; g, Anterior of body and upper lip from right side, anterior 
to right; h, Upper lip from left side, anterior to left; i, j, Upper lip, anterior and posterior views, respectively; 



VOLUME 102, NUMBER 2 



315 



gland 




k, Ventral view of lower lip, anterior towards bottom. Abbreviations as follows: a.m. = adductor muscle 
attachment area of shell; B.O. = Bellonci Organ; c.o. = copulatory organ; endop. = endopodite; epip. = epipodite; 
exop. = exopodite; i.m. = inner margin of infold; mnd. = mandible; mx. = maxilla; r.v. = right valve; v.e. = 
valve edge. 1st = First antenna, 2nd = second antenna, 5th = fifth limb, 6th = sixth limb, 7th = seventh limb. 
Arabic numerals without a prefix refer to joints of endopodite or exopodite with 1 indicating proximal joint. 
Roman numerals identify endites with I indicating proximal endite. 



316 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Spelaeoecia bermudensis, adult male, USNM 1 93483: a, Left 1 st antenna, lateral view; b, c, Endopodite 
of right 2nd antenna, medial and lateral views, respectively; d, Endopodite of left 2nd antenna, lateral view; e, 
Exopodite of 2nd antenna; f, Right 6th limb (only proximal part of all except dorsal epipodial bristles shown), 
lateral view; g, Left 7th limb, lateral view, ventral to left; h, Left lamella of furca, apron, and bifurcate unpaired 
bristle; i, Copulatory organ, anterior to left. For abbreviations, see Fig. 1 . 



Fig. 3. Spelaeoecia bermudensis, adult male, USNM 193483: a-d, Left mandible: a, Dorsal view of coxale 
endite, anterior to right; b, Lateral view of mandible (not all bristles of basale and none of 3rd endopodial joint 



VOLUME 102, NUMBER 2 



317 




shown); c, Detail of proximal set of teeth of coxale endite, from "a"; d, Joints 2 and 3 of endopodite, medial view, 
e-g, Right mandible: e, Ventral view of outer distal set of teeth and part of proximal teeth of coxale endite (inner 
distal set not shown), anterior to right; f, Ventral view of proximal set of teeth, anterior to right; g. Lateral view 
of basale and joint 1 of endopodite, lateral view. Adult female, holotype, USNM 228468: h, i, bristles between 
proximal and distal sets of teeth of coxale endite: h, Ventral view of endite of right limb, anterior to right; i, 
Dorsal view of endite of left limb, anterior to right. For abbreviations, see Fig. 1 . 



318 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Spelaeoecia bermudensis, adult male, USNM 193483, maxilla: a, Right limb (bristles of endites not 
shown); b, Left limb (not all endite bristles shown); c-e, Endites I— III of left limb (only proximal part of dashed 
bristle of endite II shown). For abbreviations, see Fig. 1 . 



Fig. 5. Spelaeoecia bermudensis, adult male, USNM 193483, 5th limb: a, Lateral view of right limb (only 
proximal part of epipodial bristles shown); b, Lateral view of sensory organ of protopodite of left limb, anterior 
to left (epipodial bristles not shown); c, Medial view of endopodite of right limb, anterior to left; d, Ventral 



VOLUME 102, NUMBER 2 



319 




oblique view of sensory organ of protopodite of right limb, distal end of limb to lower right. For abbreviations, 
see Fig. 1. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



muda, both discussed in Kornicker & Iliffe 
(1989). 

Description of adult male (Figs. l-3g, 4, 
5). — Carapace uncalcified, elongate; dorsal 
margin straight, ventral margin broadly 
convex; anterior incisur just dorsal to mid- 
height (Fig. la); in dorsal view carapace 
broadest near middle and narrowing pos- 
teriorly (Fig. lb). Anterior edge of valve sin- 
uate and posterior to anterior end of ros- 
trum. 

Ornamentation: Surface with thin linea- 
tions (representative lineations near poste- 
rior of left valve illustrated in Fig. la), and 
few single hairs. 

Infold: Broad infold except along hinge, 
narrower just posterior to inner end of in- 
cisur (Fig. la). Narrow list distal to mid- 
width of infold extending from anteroven- 
tral to postero ventral corners of valve (Fig. 
la); a second list forming narrow bar pos- 
terior to posterior juncture of hinge, then 
extending ventrally as narrow list intersect- 
ing posterior edge of valve at midheight (Fig. 
lc, d); at intersection second list joins la- 
mellar prolongation of selvage suggesting 
that second list represents edge of valve and 
that segment of shell posterior to list is outer 
valve surface and similar in nature to outer 
surface of valve forming inner side of ros- 
trum. 

Glands (Fig. la-d): Postero ventral corner 
of right valve with about 10 minute glan- 
dular pores along dorsal edge. In lateral view 
posterodorsal corner (part bearing glands) 
of right valve extending dorsally past more 
rounded corner of left valve (Fig. lc). Outer 
edge of that part of infold extending from 
incisur to midheight of posterior end of valve 
with about 1 8 minute broad tube-like glan- 
dular openings, mostly along anteroventral 
margin and anterior three-fourths of ventral 
margin (last opening just ventral to inter- 
section of valve margin and second list); 
minute bristles along valve edge, generally 
one or two between adjacent tube-like glan- 
dular openings. 

Selvage: With narrow unfringed prolon- 



gation extending from inner corner of in- 
cisur to midheight of posterior end of each 
valve. Posterior end of prolongation ends 
at intersection between valve edge and ven- 
tral end of second list. 

Size: USNM 193483, length 1.37 mm, 
height 0.78 mm. 

First antenna (Figs, le, f, 2a): With eight 
distinct joints (Fig. 2a). First joint with ter- 
minal ventral extension with long spines. 
Second joint with distinct dorsal bristle with 
indistinct rings; medial surface of joint with 
distal spines. Third joint with ventral bristle 
with base on lateral surface. Fourth joint 
about half length of third, with dorsal bristle 
reaching past eighth joint, and very long 
ventral bristle. Fifth joint shorter than 
fourth, with long ventral bristle about three- 
times length of a-bristle of seventh joint. 
Sixth joint about same length as fifth joint, 
bare. Seventh joint longer than sixth, with 
short spinous a-bristle, long ventral b-bris- 
tle, and longer ventral c-bristle. Eighth joint 
with d-bristle anterior and slightly medial 
to e-bristle and about l h longer than a-bristle 
of seventh joint; e-bristle about twice length 
of d-bristle with indistinct proximal rings, 
f-bristle medial, bent ventrally, slightly 
shorter and narrower than e-bristle, g-bris- 
tle about same length as f-bristle. Long bris- 
tles of fourth, fifth, seventh, and eighth joints 
with indistinct minute marginal spines and 
longer terminal spine. 

Second antenna (Figs, le, 2b-e): Proto- 
podite bare (Fig. le). Endopodite three- 
jointed (Fig. 2b-d): First joint elongate with 
two slender spinous a- and b-bristles (a- 
bristle about two-thirds length of b-bristle); 
second joint with short c- and d-bristles with 
distal spines, one short lateral e-bristle, one 
long f-bristle, and one long g-bristle (g-bris- 
tle longer, stouter, and medial to f-bristle); 
third joint with long equilength h-, i-, and 
j -bristles, and terminal club-like process 
(process of right limb longer and stouter than 
that of left limb); long bristles of second and 
third joints with terminal papilla. Exopod- 
ite with nine joints (Figs, le, 2e): First joint 



VOLUME 102, NUMBER 2 



321 



divided into long proximal and short distal 
segments (well developed separation of seg- 
ments in sclerotized dorsal (concave) edge 
but not in ventral (convex) edge), and with 
long terminal bristle (with minute ventral 
spines) reaching well past ninth joint; bris- 
tles of joints two to eight stouter than bristle 
of first joint and with natatory hairs; ninth 
joint with four bristles (1 minute bare, one 
short with slender ventral spines, two long 
with natatory hairs). 

Mandible (Figs, le, 3a-g): Coxale endite 
with proximal and distal sets of teeth sep- 
arated by space (Fig. 3a-c, e, f ): proximal 
set comprising four stout cusps and with 
indistinct spinous bristle at each end; sur- 
face between cusps and proximal to cusps 
with slender spines; stout tooth and four 
spinous bristles between proximal and dis- 
tal sets of teeth. Distal set of teeth com- 
prising two flat teeth, each with seven cusps; 
one stout curved process and one small bris- 
tle proximal to flat teeth. Basale (Fig. 3b, 
g): distal edge with five terminal triangular 
cusps, one sharper triangular anterior cusp, 
and a smaller posterior cusp; lateral surface 
near distal edge with sharp tooth near mid- 
width; lateral surface at or distal to mid- 
length with one small bristle and five longer 
bristles; anterior margin with one long bris- 
tle distal to midlength; posterior margin hir- 
sute, with two distal ringed bristles (proxi- 
mal with slender tip, distal tubeformed); 
proximal medial surface with transparent 
plumose bristle on protuberance, and one 
short bristle near endopodite (Fig. 3g); two 
transparent plumose bristles present on or 
close to dorsal margin; lateral surface near 
insertion of endopodite with one long bare 
bristle. Endopodite (Figs, le, 3b, d, g): First 
joint with three bristles (one short, ventral, 
one long, medial, and one long, terminal, 
dorsal); second joint widening distally, with 
three terminal dorsal bristles (one claw-like, 
unringed, two ringed), one long unringed 
subterminal ventral bristle, and minute 
spines on dorsal margin and lateral surface 
near dorsal margin; third joint with two long 



stout claw-like spinous terminal bristles, four 
short ringed bristles forming medial row 
along terminal edge, and one longer ringed 
bristle on terminal lateral edge; anterior 
margin and medial surface of third joint hir- 
sute. 

Maxilla (Figs, le, 4): Endite I with two 
proximal and nine terminal bristles (three 
tubeformed) (Fig. 4c); endite II with two 
proximal and eight terminal bristles (two 
tubeformed) (Fig. 4d); endite III with one 
long proximal bristle and six terminal bris- 
tles (two tubeformed) (Fig. 4e); some bris- 
tles on each endite stout pectinate. Basale 
with long stout dorsal bristle with long mar- 
ginal spines and one long ventral bristle with 
long marginal spines (Fig. 4a, b); division 
of basale and first endopodial joint indicat- 
ed only by separation at dorsal edge, else- 
where joints appearing fused. Endopodite 
(Fig. 4a, b): First joint with five anterior 
bristles (four proximal, one distal), two 
proximal bristles near posterior margin, and 
six distal bristles on or near posterior mar- 
gin (because of orientation of limb on slide, 
the six distal bristles of the illustrated left 
maxilla are not close to the posterior mar- 
gin); end joint with two stout claw-like bris- 
tles and five slender ringed bristles, and hairs 
along surface anterior to claws and bristles. 

Fifth limb (Figs, le, 5): Epipodite with 
bristles forming three groups, each with four 
(dorsal group), six, and five plumose bristles 
(Fig. 5a). Protopodite with long striate sen- 
sory organ with many minute papillae at tip, 
and two endites (Fig. 5a, b, d): endite I with 
three bristles with long spines; endite II with 
four bristles (two with spines, two tube- 
formed). Basale with medial hairs, one long 
anterior bristle with long spines, two prox- 
imal medial bristles with short spines, and 
six ventral bristles (two pectinate claw-like, 
three tubeformed and either bare or with 
short spines, one with long spines) (Fig. 5a). 
Endopodite with medial hairs, one proxi- 
mal medial bristle (with short spines) and 
eight additional bristles (one short tooth- 
like medial, two tubeformed, two claw-like, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



two plumose, one long bare lateral) (Fig. 5a, 
c). First exopodial joint weakly divided into 
broad proximal and narrower distal parts 
(Fig. 5a): proximal part with eight bristles 
(three slender bare ventral bristles, one long 
plumose lateral bristle near ventral margin, 
one spinous medial distal bristle near ven- 
tral margin, one very long lateral plumose 
distal bristle at midheight of joint, one spi- 
nous lateral bristle near dorsal margin, and 
one spinous distal dorsal bristle); distal part 
with four bristles (one long subterminal bare 
dorsal bristle and three shorter bristles on 
or near ventral margin). Second exopodial 
joint: dorsal margin with one bare distal 
bristle; ventral margin with four slender 
bristles (bare or with short marginal spines) 
at midlength. Third exopodial joint with two 
stout claw-like unringed bristles (longer with 
indistinct minute ventral spines, other bare) 
and one slender bare ringed ventral bristle. 

Sixth limb (Figs, le, 2f): Epipodite with 
bristles forming three groups, each with six, 
six, and five plumose bristles (Fig. 2f ). Pro- 
topodite separated from basale by distinct 
suture and separated into two shallow lobes 
by ventral indentation and suture not reach- 
ing dorsal margin: proximal part with four 
ventral bristles, all with long spines; distal 
part with five bristles (four with long spines, 
one with short spines). Basale with six plu- 
mose bristles on or near ventral margin, and 
one distal lateral bristle at about midwidth 
of joint. Endopodite single-jointed with four 
long bristles (three with long spines, one 
bare). Exopodite three-jointed: first joint 
with four bare ventral bristles; second joint 
with three bare bristles (two ventral, one 
dorsal); third joint with three bristles (mid- 
dle bristle claw-like with short ventral spines, 
dorsal bristle bare, tending to be claw-like, 
with oblique rings, ventral bristle bare). 
Protopodite and basale with long medial 
hairs. 

Seventh limb (Figs, le, 2g): Elongate with 
three long terminal bristles. 

Furca (Figs, le, 2h): Each lamella with 



eight claws; claw 1 with two weak transverse 
sutures, claw 3 with three, claw 4 with two, 
claw 5 with one; claw 2 broken off near base; 
left lamella slightly anterior to right. Un- 
paired bristle bifurcate. 

Bellonci organ (Fig. le, f): Elongate with 
suture at proximal third, bifurcating at about 
two-thirds length; each branch with round- 
ed tip just reaching third joint of first an- 
tenna. 

Lips (Fig. lg-k): Anterior face with two 
small triangular processes (one on each side) 
and small glandular processes forming row 
on each side (four-six processes in each row) 
(Fig. lg-i). Terminal posterior edge with a 
minute spine-like process and slender spines 
at each side of shallow median concavity 
(Fig. 1 h, i). Lower lip with triangular process 
on each side of mouth (Fig. Ik). 

Copulatory organ (Figs, le, 2i): On left 
side of body and consisting of two parts: 
posterior rod-shaped organ with very long 
styliform process with hirsute tip; anterior 
part broad with slight knee-like bend at 
about distal two-thirds and terminating in 
foot- shaped process with rows of minute 
teeth in "heel" area. 

Apron (Fig. 2h): Reaching about three- 
fourths length of anterior margin of furcal 
lamellae. 

Supplementary description of adult fe- 
male (Fig. 3h, i). — Size (mm): USNM 
193484A, length 1.52, height 0.86; USNM 
193484B, six specimens (length:height): 
1.54:0.91, 1.49:0.85, 1.49:0.82, 1.43:0.83, 
1.53:0.85, 1.46:0.83. Range: length 1.43- 
1.52 mm, height 0.82-0.91 mm. Because 
some specimens were measured with valves 
slightly open, height measurement is less 
accurate than that of length. (Angel & Iliffe 
(1987:545) gave the dimensions of three fe- 
males: range of lengths 1 .58-1 .64 mm. They 
also presented (Fig. 7) a shell length-width 
graph with dimensions of 2 1 females having 
a length range of about 1 .4 1-1 .64 mm. Kor- 
nicker & Iliffe (1989) gave the dimensions 
of two females with lengths of 1 .54 mm and 



VOLUME 102, NUMBER 2 



323 



1.57 mm. The lengths of the females in the 
present collection fall within the range of 
previous collections.) 

First antenna: Ventral bristle of fourth 
joint about same length as a-bristle of sev- 
enth joint; limb otherwise similar to that of 
adult male. 

Mandible (Fig. 3h, i): Similar to that of 
male. The exact number of bristles adjacent 
to the stout tooth between proximal and 
distal sets of teeth of the coxale endite is 
difficult to resolve; it is interpreted to be 
four herein (Fig. 3h, i), but could be only 
three. Angel & Iliffe (1987:547, fig. 4B) in 
describing the coxale stated, "Between outer 
and middle list single long curved spine." 
The spine is present on one limb of the ho- 
lotype but not on the other, and examina- 
tion of the "spine" under oil immersion re- 
vealed it to be debris. 

Sexual dimorphism.— The carapace of the 
single adult male (length 1.37 mm) is short- 
er than known females (range of length 1.41- 
1.64 mm). The ventral bristle of the fourth 
joint of the first antenna is about the same 
length as the a-bristle of the seventh joint 
on the adult female and almost three-times 
the length on the adult male. The fifth limb 
of the adult male bears a well developed 
sensory organ not previously reported in the 
halocyprids. The fifth limbs of two adult 
females (Holotype and USNM 193484A) 
and 2 A-l males (USNM 193404A, 
193405 A) were examined and found not to 
have a sensory organ; the A-l males are 
from Jane's Cave, Bermuda, and were dis- 
cussed in Kornicker & Iliffe (1989). 

Discussion. —Although classed in the 
Halocypridoidea, the Deeveyinae in many 
nonsexual characters appear to be a link be- 
tween the Thaumatocypridoidea and Hal- 
ocypridoidea. This relationship is also ev- 
ident when comparing the copulatory organs 
of adult males in the two superfamilies. The 
copulatory organ of the adult male S. ber- 
mudensis has the rod-shaped process sep- 



arate from the broader anterior lobe, similar 
to the copulatory organ of thaumatocyprids. 
In halocyprids other than S. bermudensis 
the rod-shaped process of the adult male lies 
inside a canal within the anterior lobe. Ju- 
venile males of both thaumatocyprids and 
halocyprids have the rod-shaped process 
separate from the anterior lobe suggesting 
that having the rod-shaped process inside a 
canal within the anterior lobe is an apo- 
morphic character state. The endopodite of 
the adult male second antenna of S. 
bermudensis differs from those generally 
present in both thaumatocyprids and hal- 
ocyprids in having a straight rather than 
hook-shaped clasper. Not all halocyprids 
have a clasper. 

Acknowledgments 

I thank Dr. Jan H. Stock, Institut voor 
Taxonomische Zoology, Zoologisch Mu- 
seum, Amsterdam, Nederland, for the os- 
tracodes from Wonderland Cave, Mr. Jack 
Schroeder for inking my camera lucida 
drawings, and Ms. Elizabeth Harrison-Nel- 
son for general assistance. 

Literature Cited 

Angel, M. V., & T. M. Iliffe. 1987. Spelaeoecia ber- 
mudensis, new genus, new species, a halocyprid 
ostracod from marine caves in Bermuda.— 
Journal of Crustacean Biology, 7(3):54 1-553. 

Kornicker, L. S., & T. M. Iliffe. 1989. Ostracoda 
(Myodocopina, Cladocopina. Halocypridina) 
mainly from Anchialine Caves in Bermuda.— 
Smithsonian Contributions to Zoology 470: 
1-88. 

, & R. J. Palmer. 1987. Deeveya bransoni, a 

new species of troglobitic halocyprid ostracode 
from anchialine caves on South Andros Island. 
Bahamas (Crustacea: Ostracoda). — Proceedings 
of the Biological Society of Washington. 100(3): 
610-623. 

Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 
102(2), 1989, pp. 324-330 

NEW RECORDS OF ENTOCYTHERID OSTRACODS 

INFESTING BURROWING CRAYFISHES, WITH 

THE DESCRIPTION OF A NEW SPECIES, 

ASCETOCYTHERE STOCKERI 

Horton H. Hobbs, Jr., and Daniel J. Peters 

Abstract.— New records of the occurrence of entocytherids on burrowing 
crayfishes are reported from Kentucky, North Carolina, Virginia, and West 
Virginia. These include members of the genera Ascetocythere (3), Dactylocy there 
(3), Donnaldsoncy there (1), Entocythere (2), and Uncinocy there (2). Ascetocyth- 
ere stockeri, a close relative of A. cosmeta, is described from four localities in 
the Ohio Basin in Mason and Cabell counties, West Virginia, and a single 
locality in Carter County, Kentucky. 



In their study of burrowing crayfishes, 
Raymond F. Jezerinac and G. Whitney 
Stocker, of Ohio State University, Newark, 
saved the detritus from containers in which 
the crayfishes were preserved and have 
kindly permitted us to study the ostracods 
that were entrapped in it. Of the 20 samples 
received, 1 5 from the Ohio, Potomac, and 
Tennessee watersheds contained entocyth- 
erids among which were representatives of 
1 1 species belonging to 5 genera. These were 
infesting six species of crayfishes (the iden- 
tities of which were furnished to us by 
Messrs. Jezerinac and Stocker), five of which 
are largely restricted to burrows. The as- 
sociations of the ostracods and their crayfish 
hosts in the 1 5 localities are recorded below. 

For each species of ostracod, the original 
description and the first use of the currently 
accepted orthography are cited. These are 
followed by a list of the new localities, cray- 
fish hosts, and remarks on the ecological 
and geographic distribution of the species. 

Except as noted otherwise, the ostracods 
are deposited in the National Museum of 
Natural History, Smithsonian Institution. 

Ascetocythere cosmeta Hobbs & Hart 

Ascetocythere cosmeta Hobbs & Hart, 1966: 
46, figs. 24, 25. [Type locality: 4 mi N of 



jet of St. Rte. 89 and Blue Ridge Parkway, 
Grayson Co., Virginia. Host: Cambarus 
dubius Faxon.] 

New record. —North Carolina: Ashe 
County [New Basin], burrows near un- 
named trib of North Fork of New River on 
county road, 0.1 mi E of Crumpler, 8.8 mi 
NE of West Jefferson, 1 1 Aug 1984, G. W. 
Stocker, R. F. Jezerinac, and R. F. Thoma, 
host: Cambarus (Jugicambarus) dubius 
Faxon, associates: Dactylocy there daph- 
nioides (Hobbs, 1955), Entocythere harrisi 
Peters, 1975, and Uncinocythere simondsi 
(Hobbs and Walton, 1960). 

Habitat.— Crayfish burrows. 

Distribution. —The Catawba and New ba- 
sins in Alleghany, Ashe, and Avery coun- 
ties, North Carolina, and Grayson County, 
Virginia. 

Ascetocythere hyperoche Hobbs & Hart 

Ascetocythere hyperoche Hobbs & Hart, 
1966:41, figs. 12, 13. [Type locality: trib. 
of Cherokee Reservoir, 0.5 mi W of jet 
of U.S. Hwys. 1 IE and 25E, Grainger Co., 
Tennessee. Host: Orconectes spinosus 
(Bundy), Cambarus (Hiaticambarus) lon- 
girostris Faxon, and Cambarus (C.) bar- 
tonii cavatus Hay.] 



VOLUME 102, NUMBER 2 



325 




.05 mm 




Fig. 1. Ascetocythere stockeri, new sp.: a, Copulatory complex of holotypic male; b, Right valve of shell of 
holotype; c, Same of paratypic female (two irregular, overlapping masses in interior ova). 



New record.— Virginia: Russell County 
[Holston Basin], floodplain of tributary to 
Big Moccasin Creek on County Rd 657, 0.9 
air mi W of Hansonville, 15.9 air mi N of 
Bristol, 9 Aug 1986, G.W.S. andR.FJ., host: 
Cambarus (Jugicambarus) dubius Faxon, 
associate, Donnaldsoncy there donnaldson- 
ensis{¥A\e 1931). 

Habitat.— Probably largely confined to 
crayfish burrows. 

Distribution. —This ostracod was previ- 
ously known only from the type locality in 
the Holston River basin in Grainger Coun- 
ty, Tennessee, where Hobbs and Hart re- 
ported the hosts to be three stream-dwelling 
crayfishes. 

Ascetocythere stockeri, new species 
Fig. 1 

Diagnosis.— Eye pigmented and located 
about one-sixth shell length from anterior 
margin. Shell (Fig. lb) ovate with greatest 



height about one-third shell length from 
posterior end where about 1.5 times height 
at level of eye; margin entire, lacking emar- 
ginations and prominences. Submarginal 
setae present except dorsally. 

Copulatory complex (Fig. la) with peni- 
ferum expanded ventrally in bubiform en- 
largement following conspicuously slender 
constricted section about one-fourth of 
length of peniferum from distal extremity; 
anterior process lacking; ventral process 
digitiform extending ventrally much be- 
yond other processes; posterior process rep- 
resented by two acute teeth. Penis very 
prominent, spermatic and prostatic ele- 
ments not contiguous throughout their 
lengths, both C-shaped and emerging ven- 
trally at base of ventral process between it 
and small cephaloventrally directed lobe. 
Clasping apparatus broadly arched, obliter- 
ating junction of vertical and horizontal 
rami; postaxial border entire; preaxial bor- 



326 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1— Measurements (in mm) of Ascetocyt here 
stockeri, n. sp. 





Holotype 


Males (14) 


Females (5) 


Length (range) 


0.39 


0.35-0.43 


0.39-0.42 


Average 




0.38 


0.41 


Height (range) 


0.23 


0.21-0.25 


0.22-0.25 


Average 




0.23 


0.24 



der with three teeth grouped on thickened 
distal fifth. Dorsal finger moderately heavy 
and about one-half as long as curved, more 
slender ventral finger. 

Triunguis female.— Eye located about 
one-sixth shell length from anterior end. 
Shell (Fig. lc) subovate, weakly concave in 
anterior three-fifths. Greatest height dis- 
tinctly posterior to midlength where about 
1 .4 times that at level of eye. Submarginal 
setae present except dorsally. Genital ap- 
paratus supported by pair of ribs extending 
dorsomesially from complex of supports at 
base of third leg. Apices of ribs in contact 
with small, subspherical, homogeneous, 
hyaline body embraced dorsally and pos- 
teriorly by larger, heterogeneous mass. For 
measurements, see Table 1. 

Type locality: Chief Cornstalk Hunting 
Grounds Campground, 2.1 miles southwest 
of Beech Hill, 10.1 miles southeast of 
Gallipolis, Mason County, West Virginia 
[Kanawha Basin], where it was found in a 
collection of crayfishes containing represen- 
tatives of three species: Cambarus (Jugi- 
cambarus) dubius Faxon, Cambarus (La- 
cunicambarus) diogenes Girard, and 
Cambarus (C) bartonii cavatus Hay. These 
specimens were collected on 20 Apr 1986 
by G. Whitney Stocker, Dave Chrisman, 
and R. F. Jezerinac. 

Disposition of types: The holotypic male 
is deposited in the National Museum of 
Natural History (Smithsonian Institution), 
USNM 240114. Paratypes are in the col- 
lections of H. H. Hobbs III and the Smith- 
sonian Institution. 

Hosts: Cambarus (C.) bartonii cavatus 



Hay, C (Jugicambarus) dubius Faxon, and 
C (Lacunicambarus) diogenes Girard. C 
(J. ) dubius was present in all of the collec- 
tions containing specimens of Ascetocyt here 
stockeri, and we think it unlikely that this 
ostracod occurs on the first and last named 
crayfishes. 

Entocytherid associates: Dactylocythere 
macroholca Hobbs & Hobbs, 1970; Don- 
naldsoncy there donnaldsonensis (Klie, 
1931); Entocythere elliptica Hoff, 1 944; and 
Uncinocythere zancla Hobbs & Walton, 
1963. 

Range and specimens examined: Nine- 
teen specimens from the following locali- 
ties. Kentucky: Carter County [Little Sandy 
Basin], (1) roadside ditch along Dry Fork 
1.5 air mi S of Willard, 1 Sep 1986, Dave 
Hile, G.W.S., and R.F.J., host: C. (/.) du- 
bius, associates: Dactylocythere macroholca 
Hobbs & Hobbs, Entocythere elliptica Hoff, 
and Uncinocythere zancla Hobbs & Walton. 
West Virginia: Mason County [Kanawha 
Basin], (2) Type locality; (3) 3 mi S of type 
locality and 2.7 mi SE of Arlee, 2 May 1987, 
G. Lombardo, R. French, and G.W.S., host: 

C. (/.) dubius, no ostracod associates. Cabell 
County [Ohio Basin], (4) Spurlock Creek on 
Union Ridge Rd, 1 mi S of St. Rte. 2, 1.8 
mi N of Union Ridge, 3 May 1987, R.F., 

D. Chrisman, G.L., R.F.J., andG.W.S., host: 
Cambarus (J.) dubius, associate, Dn. don- 
naldsonensis', (5) along Brian Creek Rd, S 
of Union Ridge Rd, 4.4 mi S of Glenwood, 
3 May 1987, G.L., R.F., D.C., R.F.J., and 
G.W.S., host: C (J.) dubius, associate: Dn. 
donnaldsonensis. 

Relationships: Ascetocythere stockeri is a 
member of the Coryphodes Group (Hobbs 
& Hart 1966:44), characterized by the teeth 
on the preaxial margin of the clasping ap- 
paratus being grouped distally. Its closest 
relative is probably A. cosmeta Hobbs & 
Hart, 1966, from which it differs most con- 
spicuously in lacking an anterior process on 
the peniferum. It differs from the other 
members of the group, A. coryphodes Hobbs 
& Hart, 1966, and A. myxoides Hobbs & 



VOLUME 102, NUMBER 2 



327 



Hart, 1966, in lacking a snoutlike promi- 
nence on the anterior surface of the bulbous 
part of the peniferum. 

Etymology: This ostracod is named in 
honor of G. Whitney Stocker, who collected 
or participated in collecting most of the ma- 
terials on which this report is based. 

Dactylocy there crawfordi Hart 

Dactylocy there crawfordi Hart, 1965:255, 
figs. 1,2. [Type locality: Macacheek Creek 
on St. Rte. 287, Logan Co., Ohio. Host: 
Cambarus (Lacunicambarus) diogenes 
Girard.] 

New record. —West Virginia: Mason 
County [Ohio Basin], wet field along St. Rte. 
2, 2.0 mi S of U.S. Hwy. 35, 1.5 air mi E 
of Gallipolis, 19 Apr 1986, D.C., R.F.J., 
G.W.S., host: Fallicambarus (Creaserinus) 
fodiens (Cottle). 

Habitat. —Crayfish burrows. 

Distribution. —Ohio River Basin: White 
and Wabash drainages in Indiana, Little 
Miami, Scioto, and Symmes Creek in Ohio, 
and in Mason County, West Virginia. 

Dactylocythere daphnioides (Hobbs) 

Entocythere daphnioides Hobbs 1955:325, 

figs. 1, 2, 5, 6-9. 
Dactylocythere daphnioides.— Hart, 1962: 

130. 

New record. —North Carolina: Ashe 
County [New Basin], burrows near trib to 
North Fork of New River, 0.1 mi E of 
Crumpler; 8.8 mi NE of West Jefferson, 1 1 
Aug 1984, R.F.J., G.W.D., R.F.T., host: C. 
(J.) dubius, associates: As. cosmeta, E. har- 
risi, and U. simondsi. 

Habitat.— Except for the record cited 
above, all known specimens have been found 
on crayfishes living in lotic habitats. 

Distribution. —"From headwaters of the 
Little Tennessee and Pee Dee drainage sys- 
tems in North Carolina northward to the 
New and Big Sandy systems in Virginia and 
West Virginia. The Kentucky and Missouri 



localities cited by Hart and Hart (1974:56) 
should be confirmed" (Hobbs & Peters, 
1977:28). 

Dactylocythere macroholca 
Hobbs & Hobbs 

Dactylocythere macroholca Hobbs & Hobbs 
1970:6, fig. 3a-d. 

New record.— Kentucky: Carter County 
[Little Sandy Basin], roadside ditch 1.5 air 
mi S of Willard, near Lawrence Co line, 1 
Sep 1986, D.H., R.F.J., G.W.S., host: C. (J.) 
dubius, associates, As. stocked, E. elliptica, 
and U. zancla. 

Habitat.— Burrows and streams. 

Distribution. —Ohio River Basin in Ken- 
tucky: Barren, Licking, and Little Sandy wa- 
tersheds. 

Donnaldsoncythere donnaldsonensis (Klie) 

Entocythere donnaldsonensis Klie, 1931: 
334-341, figs. 1-9. [Type locality: Don- 
ald's Cave, Lawrence County, Indiana. 
Host/s: uncertain; however, probably Or- 
conectes inermis inermis Cope and/or 
Cambarus (Erebicambarus) tenebrosus 
Hay. 

Donnaldsoncythere donnaldsonensis. — Hart, 
1962:131. 

New record. —Kentucky: Lawrence Coun- 
ty [Big Sandy Basin], (1) roadside ditch 1.3 
air mi NW of Webbyville, 1.4 air mi S of 
Willard, 1 Sep 1986, D.H., G.W.S., and 
R.F.J. , host: C. (J.) dubius, associates: none. 
Lewis County [Licking Basin], (2) seep on 
St. Rte. 24, 3.8 mi E of county line, 2 air 
mi E of Foxport, 9 Jun 1984, G.W.S., Diane 
M. Williams, R.F.J. , host: C. (J.) dubius, no 
associates. Virginia: Russell County [Hol- 
ston Basin], (3) floodplain along trib of Big 
Moccasin Creek on Co Rd 657, 0.9 air mi 
W of Hanson ville, 15.8 air mi N of Bristol, 
9 Aug 1986, G.W.S. and R.F.J., host: C. (J.) 
dubius, associate: As. hyperoche. West Vir- 
ginia: Barbour County [Tygart Basin], (4) 
roadside ditch 0.8 mi E of U.S. Hwy. 119 



328 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



on Arden Rd, 1.3 air mi NE of Hopewell, 
R.FJ. and G.W.S., hosts: C (/.) dubius, C. 
(J.) diogenes, no associates. Cabell County 
[Ohio Basin]. (5) burrows along Brian Creek 
Rd. S of Union Ridge Rd, 4.4 mi S of Glen- 
wood, 3 May 1987, Greg Lombardo, Randy 
French, Dave Chrisman, R.F.J. , and G.W.S., 
host: C. (J.) dubius, associate: As. stockeri; 
(6) burrows on Union Ridge Rd, 1 mi S of 
St. Rte. 2, 1.75 mi N of Union Ridge, G.L., 
R.F., D.C., R.F.J., and G.W.S., host: C. (J.) 
dubius, associate: As. stockeri. Grant Coun- 
ty [Potomac Basin], (7) roadside ditch on 
Forest Rd 75, 10.2 air mi NW of Peters- 
burg, 28 Jun 1986, D.C. and G.W.S., host: 
C. (J.) monongalensis, no associates. Ma- 
son County [Kanawha Basin], (8) see type 
locality of As. stockeri, (9) burrows along 
dirt road 3 mi S of Chief Cornstalk Hunting 
Ground Campground, 2.7 mi SE of Arlee, 
2 May 1987, G.L., R.F., and G.W.S. (8) See 
Locality "2" for As. stockeri. Pocahontas 
County [Greenbrier Basin], (10) seep in Falls 
of Hills Creek Scenic Area off St. Rte. 39, 
1 .6 mi E of Greenbrier Co line, 11.1 air mi 
ESE of Richwood, 7 Jul 1985, V.L. Stocker 
and G.W.S., host: C. (J.) monongalensis, no 
associates. 

Habitat.— AW. of the records cited above 
are based on specimens collected from bur- 
rowing crayfishes. This ostracod, however, 
inhabits a wide range of epigean and hy- 
pogean habitats. 

Distribution.— "From northern Georgia 
to Indiana and Maine" (Hobbs & Peters 
1 977:44). All of the localities cited since this 
summary statement appeared (Hobbs & Pe- 
ters, 1982:312; and Hobbs &McClure 1983: 
772-778) lie within these limits. 

Entocythere elliptica Hoff 

Entocythere elliptica Hoff, 1944:328, figs. 
15-21. [Type locality: north of Fargo, 
Clinch County, Georgia. Host: Procam- 
barus (Ortmannicus) seminolae Hobbs.] 

New record. — Kentucky: Carter County 



[Little Sandy Basin], roadside ditch 1.5 air 
mi S of Willard near Lawrence County line, 
1 Sep 1986, D.H., G.W.S., R.F.J., host: C. 
(J.) dubius, associates: As. stockeri, Dt. ma- 
croholca, and U. zancla. 

Habitat.— Burrows and lentic and lotic 
habitats. 

Distribution. -"Hart & Hart (1974:[87- 
88], 90) recorded the presence of E. elliptica 
and/or E. internotalus from Texas to North 
Carolina and from Kentucky to Florida, and 
Peters (1975:34) reported it from a number 
of localities in the James River watershed 
in Virginia" (Andolshek & Hobbs, 1986: 
31). The locality cited above extends the 
range diagonally across the entire state of 
Kentucky. 

Entocythere harrisi Peters 

Entocythere harrisi Peters, 1975:32, figs. 
5-7. 

New record. —North Carolina: Ashe 
County [New Basin], burrows near trib of 
North Fork of New River, 0.1 mi E of 
Crumpler; 8.8 mi NE of West Jefferson, 1 1 
Aug 1984, R.F.J., G.W.S., R.F.T., host: C 
(J.) dubius, associates: As. cosmeta, Dt. 
daphnioides, and U. simondsi. 

Habitat.— Burrows and lotic habitats. 

Distribution. —On the Atlantic versant 
from the York River Basin in Virginia 
southward through the Catawba Basin in 
North Carolina, and the New River Basin 
in Ashe County, North Carolina. 

Uncinocythere simondsi (Hobbs & Walton) 

Entocythere simondsi Hobbs & Walton, 

1960:17, figs. 1-10. 
Uncinocythere simondsi. —Hart, 1 962: 138. 

New record. —North Carolina: Ashe 
County [New Basin], burrows near trib of 
North Fork of New River, 0.1 mi E of 
Crumpler; 8.8 mi NE of West Jefferson, 1 1 
Aug 1984, R.F.J., G.W.S., R.F.T., host: C 



VOLUME 102, NUMBER 2 



329 



(/.) dubius, associates: As. cosmeta, Dt. 
daphnioides, and E. harrisi. 

Habitat.— Burrows and a broad range of 
epigean lotic and lentic habitats. 

Distribution.— ¥rom Illinois, Kentucky, 
and North Carolina southward to Missis- 
sippi and northern Florida (Hart & Hart 
1974; Hobbs & Peters 1977, 1982; Andol- 
shek&Hobbs 1986). 

Uncinocy there zancla Hobbs & Walton 

Uncinocy there zancla Hobbs & Walton 
1963:456, figs. 1-3. 

New record.— Kentucky: Adair County 
[Green Basin], burrows along St. Rte. 205, 
0.8 mi NE of Neatsville, 10 mi N of Russell 
Springs, 15 Mar 1987, G.W.S. and R.F.J., 
host: Cambarus (Jugicambarus) sp.?; no as- 
sociates; Carter County [Little Sandy Ba- 
sin], roadside ditch 1.5 air mi S of Willard 
near Lawrence County line, 1 Sep 1986, 
D.H., G.W.S., R.F.J., host: C. (J.) dubius, 
associates: As. stocked, Dt. macroholca, and 
E. elliptica. 

Habitat.— This ostracod has been found 
most frequently infesting stream-dwelling 
crayfishes, but has also been found associ- 
ated with a troglobitic species in Hart Coun- 
ty, Kentucky, and with the burrower cited 
above. 

Distribution. —Known only from the 
Kentucky, Green, and Little Sandy wa- 
tersheds in Kentucky, and the Cumberland 
and Tennessee basins in Tennessee. 



Acknowledgments 

We are grateful to Raymond F. Jezerinac 
and G. Whitney Stocker for making possible 
our study of the material gleaned from their 
collections of burrowing crayfishes. Our 
thanks are also extended to C. W. Hart, Jr., 
of the Smithsonian Institution, and H. H. 
Hobbs III, of Wittenburg University for their 
criticisms of the manuscript. 



Literature Cited 

Andolshek, Margaret D., & Horton H. Hobbs, Jr. 1 986. 
The entocytherid ostracod faunas of southeast- 
ern Georgia.— Smithsonian Contributions to 
Zoology 424:1-43. 

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. 

. 1965. New entocytherid ostracods and dis- 
tribution records for five midwestern states.— 
Transactions of the American Microscopical 
Society 84(2):255-259. 

Hart, Dabney G., & C. W. Hart, Jr. 1974. The os- 
tracod family Entocytheridae.— Academy of 
Natural Sciences of Philadelphia Monograph 18: 
ix + 239 pages. 

Hobbs, Horton H., Jr. 1955. Ostracods of the genus 
Entocythere from the New River system in North 
Carolina, Virginia, and West Virginia.— Trans- 
actions of the American Microscopical Society 
74(4):325-333. 

, & C. W. Hart, Jr. 1966. On the entocytherid 

ostracod genera Ascetocythere, Plectocy there, 
Phymocythere (gen. nov.), and Cymocythere with 
descriptions of new species. —Proceedings of the 
Academy of Natural Sciences of Philadelphia 
11 8(2): 3 5-61. 

, & H. H. Hobbs, III. 1970. New entocytherid 

ostracods with a key to the genera of the subfam- 
ily Entocytherinae. — Smithsonian Contribu- 
tions to Zoology 47:1-19. 

, & Auden C. McClure. 1983. On a small col- 
lection of entocytherid ostracods with the de- 
scriptions of three new species.— Proceedings of 
the Biological Society of Washington 96(4):770- 
779. 

, & Daniel J. Peters. 1977. The entocytherid 

ostracods of North Carolina. — Smithsonian 
Contributions to Zoology 147:1-73. 

, and . 1982. The entocytherid ostracod 

fauna of northern Georgia. — Proceedings of the 
Biological Society of Washington 95(2):297-318. 

, & Margaret Walton. 1960. Three new ostra- 
cods of the genus Entocythere from the Hiwas- 
see drainage system in Georgia and Tennes- 
see.— Journal of the Tennessee Academy of 
Science 35(1): 17-23. 

. 1963. Three new ostracods (Ostracoda, En- 
tocytheridae) from the Duck River drainage in 
Tennessee. — American Midland Naturalist 
69(2):456-461. 

Hoff, C. Clayton. 1 944. New American species of the 
ostracod genus Entocythere.— American Mid- 
land Naturalist 32(2):327-357. 



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Klie. W. 1931. Campagne speologique de C. Bolivar 
et R. Jeannel dans l'Amerique du Nord (1928) 
3: Crustaces Ostracodes. — Biospeologica: Ar- 
chives de Zoologie Experimental et Generate 
71(3):333-344. 

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. 



(HHH) Department of Invertebrate Zo- 
ology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. 20560; (DJP) York High School, 9300 
George Washington Highway, Yorktown, 
Virginia 23692. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 331-338 

PSEUDOCYCLOPS LEPIDOTUS, A NEW SPECIES 

OF DEMERSAL COPEPOD 

(CALANOIDA: PSEUDOCYCLOPIDAE) FROM 

THE NORTHWESTERN PACIFIC 

Douglas J. Barr and Susumu Ohtsuka 

Abstract. —Pseudocy clops lepidotus, is described from Kuchinoerabu Island, 
off Kyushu, Japan. The main diagnostic characters are the lateral clefts in 
pediger 5, foliaceous urosomal scales, and structure of the fifth legs in both 
sexes. 



Although most oceanic calanoid cope- 
pods are planktonic, many coastal species 
are demersal and occupy restricted habitats 
on or near the substrate. Such species may 
be overlooked or misidentified owing to their 
small size and resemblance to cyclopoid co- 
pepods. Several families of calanoids are 
comprised entirely of demersal species, the 
largest and most widely distributed of these 
being the family Pseudocyclopidae Gies- 
brecht, 1893. This family is monogeneric, 
containing only the genus Pseudocyclops 
Brady, 1872. 

Pseudocyclops at present contains 26 de- 
scribed species, 15 of which have been col- 
lected only from the North Atlantic. Only 
three species have been described from the 
Pacific: P. australis Nicholls, 1944, from 
southern Australia; P. pacificus Vervoort, 
1964, from the Caroline Islands; and P. bi- 
lobatus Dawson, 1977, from southern Cal- 
ifornia. P. australis was subsequently re- 
ported by Tanaka (1966) from Kyushu, 
Japan, close to the type locality of the new 
species described below. Our specimens were 
obtained with a hand-held plankton net 
while using SCUBA over a coral sandy bot- 
tom at a depth of 9 m off Kuchinoerabu 
Island, Kyushu, Japan. 

Pseudocyclops lepidotus, new species 
Figs. 1-5 

Material. —Dissected female holotype 
(USNM 229956), and dissected male allo- 



type (USNM 229957) collected 10 Sep 1986 
from Nishino-Hama, Kuchinoerabu Island, 
Kagoshima Prefecture, Japan. Two female 
paratypes (USNM 229958), one dissected 
on slides and one whole, collected on same 
date from same locality. 

Description. —Female (Fig. 1 A, B): length 
0.90 mm, body compact, prosome oval in 
dorsal view; cephalosome partly fused with 
pediger 1, produced anteroventrally into 
acute rostrum (Fig. 1C); pediger 5 separate 
from pediger 4, with dorsolateral cleft bear- 
ing small seta (Figs. IB, D; 5 A). Urosome 
(Fig. IE) 4-segmented, segments bearing 
transverse rows of subtriangular foliaceous 
scales, many of which have fallen off during 
dissection (Fig. 5B); genital segment longer 
than following segments combined, open- 
ings widely separate; segment 3 (Fig. 5C) 
with two dorsal processes reaching mid- 
length of caudal rami; anal segment re- 
duced, telescoped into segment 3; caudal 
ramus serrate posterodorsally, bearing one 
dorsal, one lateral and four apical setae. 

First antenna (Fig. 1 F, G) 20-segmented, 
reaching posterior end of first pediger; seg- 
ment 1 with rows of anterior surface spi- 
nules, three large esthetascs; suture between 
segments 17 and 18 incomplete. 

Second antenna (Fig. 2A) with basipodal 
segments each bearing one seta on disto- 
medial margin; endopod three-segmented, 
segment 1 with two setae on medial margin 
and small spinules on outer distal margin, 



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Fig. 1. Pseudocyclops lepidotus, female: A, Habitus, dorsal; B, Habitus, lateral; C, Rostrum, ventral; D, 
Pedigers 4-5, lateral; E, Urosome, dorsal; F, First antenna; G, First antenna, segment 1. 



VOLUME 102, NUMBER 2 



333 




Fig. 2. Pseudocyclops lepidotus, female: A, Second antenna; B, Mandible; C, First maxilla; D, Second maxilla; 
E, Maxilliped, posterior; F, First leg, anterior. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Pseudocyclops lepidotus, female: A, Second leg, anterior; B, Third leg, anterior; C, Fourth leg, anterior; 
D, Fifth leg, anterior. 



VOLUME 102, NUMBER 2 



335 




\ B,D,E 



Fig. 4. Pseudocyclops lepidotus, male: A, Habitus, dorsal; B, Urosome, dorsal; C, Right first antenna; D, Left 
fifth leg, posterior; E, Right fifth leg, posterior. 



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Fig. 5. Pseudocyclops lepidotus, female: A, Pediger 5, lateral; B, Scales on urosome segment 2, dorsal; C, 
Urosome, posterodorsal. Male: D, Right first antenna, segments 14-18; E, Fifth legs, posterior; F, Right endopod, 
fifth leg, posterior. Scale bars = 10 microns. 



VOLUME 102, NUMBER 2 



337 



segment 2 with five medial and three ter- 
minal setae, segment 3 with seven terminal 
setae; exopod seven-segmented, segments 
1-6 each with one terminal seta, segment 7 
with one medial and four terminal setae. 

Mandible (Fig. 2B) with 8 heavily scler- 
otized teeth on gnathobase; basis with 3 
medial setae; endopod with 2 segments, 
bearing 3 and 1 setae, respectively; exopod 
4-segmented, bearing 1, 1, 1, and 3 setae, 
respectively. 

First maxilla (fig. 2C) with first inner lobe 
bearing 9 strong terminal spines, 4 setae on 
posterior surface, 1 seta on anterior surface; 
second and third inner lobes with 3 and 4 
setae, respectively; outer lobes 1 and 2 with 
8 and 1 setae, respectively; basis 2 bearing 
4 distomedial setae; endopod 2 segmented, 
bearing 8 medial and 7 terminal setae; ex- 
opod one-segmented, with 10 setae. 

Second maxilla (Fig. 2D) with five inner 
lobes bearing five, three, three, three, and 
seven setae, respectively; endopod reduced, 
indistinctly segmented, bearing seven setae. 

Maxilliped (Fig. 2E) with coxa carrying 
four lobes bearing one, two, three, and 
three setae, respectively, lobes three and four 
with surface spinules; basis with two lobes 
bearing three and two setae, respectively; 
endopod five-segmented, bearing four, four, 
three, four, and four setae, respectively. 

Legs 1-4 (Figs. 2F; 3A-C) with three-seg- 
mented rami, all bearing anterior and pos- 
terior surface spinules; leg 1 (Fig. 2F) with 
acute medial process and inner seta on ba- 
sis, distolateral margins of endopod seg- 
ments 2 and 3 acutely produced, distolateral 
margins of exopod segments 1 and 2 pro- 
duced into blunt serrate processes; leg 2 (Fig. 
3A) with serrate lateral margins of exopod 
segments 2 and 3; leg 4 (Fig. 3C) with short 
external seta on basis, endopod segment 3 
bearing 7 setae. 

Fifth leg (Fig. 3D) smaller than fourth, 
basis with short outer posterior seta; en- 
dopod three-segmented, segments 1 and 2 
produced distolaterally into bifurcate pro- 



cess, segment 3 bearing six setae; exopod 
segment 3 with four inner setae and four 
flanged spines, innermost spine narrow, only 
flanged at base. 

Male (Fig. 4 A): length 0.73 mm, prosome 
as in female. Urosome (Fig. 4B) 5 -seg- 
mented, adorned with rows of foliaceous 
"scales," anal segment and caudal rami as 
in female. 

Left first antenna 20-segmented, seg- 
ments 4 and 5 partly fused, otherwise as in 
female. Other cephalic appendages, rostrum 
and legs 1-4 as in female. 

Right first antenna (Fig. 4C) 19-segment- 
ed, geniculate between segments 1 5 and 1 6 
(Fig. 5D); segments 14 and 15 partly fused, 
bearing a sinuous process on anterior mar- 
gin; segment 16 with acute lateral process; 
segment 1 7 produced distally into sharp tri- 
angular process curving medially, almost 
reaching end of segment 18. 

Fifth legs (Figs. 4D-E, 5E) asymmetrical, 
biramous, highly modified. Basipodal seg- 
ments of left leg (Fig. 4D) fused, those of 
right leg (Fig. 4E) separate, all with posterior 
surface spinules. Left endopod one-seg- 
mented, reaching tip of left exopod, bearing 
five plumose setae; left endopodal process 
slender, narrowing distally, confluent with 
basis, reaching end of endopod. Left exopod 
two-segmented, segment 1 with terminal 
spine flanged laterally, segment 2 complex, 
membranous, bearing four lobes: innermost 
lobe spatulate, medial margin serrate; sec- 
ond innermost lobe elongate, narrowed at 
tip; second outermost lobe half as wide as 
long, bearing proximal seta, distal margin 
with v-shaped cleft; outermost lobe longest, 
three times as long as wide, tip dissected 
into three acute processes, bearing small seta 
at base. Right endopod (Fig. 5F) one-seg- 
mented, narrow; small posterior sensory pit 
on lateral margin; distal margin blunt, with 
patch of small setae. Right exopod one-seg- 
mented, with flanged outer spine and two 
long equal spines curving medially, inner- 
most spine narrowed to acute apex, tip 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



curved laterally, with two setae and naked 
process at base. 

Etymology.— The specific name, lepido- 
tus (G.), meaning "with scales," refers to 
the foliaceous scales on the urosome of both 
sexes. 

Discussion.— Known species of Pseudo- 
cyclops are demersal, spending the greater 
part of the diel cycle on or near the sub- 
strate. The nearly spherical shape and short 
first antennae limit the ability of these co- 
pepods to remain in the water column. P. 
lepidotus, however, is unique among mem- 
bers of the genus in possessing relatively 
long first antennae. In previously known 
species, the first antennae have from 1 5 to 
1 8 segments and scarcely extend beyond the 
cephalosome. In P. lepidotus the first an- 
tennae have 20 segments and extend be- 
yond the end of the first pediger. This fea- 
ture, combined with its comparatively large 
size and light purple coloration often as- 
sociated with planktonic species, suggests 
that P. lepidotus may spend more time in 
the water column than other species in the 
genus. 

The unique characteristics of urosomal 
scales, lateral clefts in pediger 5 and elongate 
first antennae, separate this species from all 
known species of the genus. At least one 
undescribed species of Pseudocyclops oc- 
curring in the western Pacific is closely re- 
lated to this new species (Barr, unpub- 
lished). This undescribed species, collected 
from coastal waters off Guam and the Phil- 
ippines, possesses all of the above-men- 
tioned characters. It differs from P. lepidotus 
in details of the fifth legs in both sexes and 
will be described in a forthcoming paper. 

Acknowledgments 

We thank Drs. Thomas E. Bowman 
(USNM), Taisoo Park (Texas A&M Uni- 



versity) and Merrill H. Sweet II (Texas A&M 
University) for their critical reviews of the 
manuscript. We also extend appreciation to 
Dr. Robert C. Burghardt (Texas A&M Uni- 
versity) for allowing access to the Electron 
Microscopy Center, and Dr. K. Gushima 
and Mr. M. Noda (Hiroshima University) 
for cooperation at sea. 

Literature Cited 

Brady, G. S. 1872. Contributions to the study of the 
Entomostraca, VII. A list of the non-parasitic 
marine Copepoda of the northeast coast of En- 
gland. —Natural History Transactions of North- 
umberland and Durham 4:423-445, pi. XVII- 
XX. 

Dawson, J. K. 1977. A new species of Pseudocyclops 
(Copepoda: Calanoida) from the southern Cal- 
ifornia coast.— Transactions of the American 
Microscopical Society 96:247-253. 

Giesbrecht, W. 1893. Mitteilungen iiber Copepoden. 
1-6. — Mitteilungen Zoologischen Station zu 
Neapel 11:56-106, pi. 5-7. 

Nicholls, A. G. 1 944. Littoral Copepoda from South 
Australia (II). Calanoida, Cyclopoida, Notodel- 
phyoida, Monstrilloida and Caligoida.— Rec- 
ords of the South Australian Museum 8:1-62. 

Tanaka, O. 1966. Neritic Copepoda Calanoida from 
the north-west coast of Kyushu.— Proceedings 
of the Symposium on Crustacea, held at Erna- 
kulam from January 12 to 15, 1965, 1:38-50. 
Marine Biological Association of India, Man- 
dapam Camp. 

Vervoort, W. 1964. Free-living Copepoda from Ifa- 
luk Atoll in the Caroline Islands with notes on 
related species.— Bulletin of the United States 
National Museum 236:1-431. 

(DJB) Department of Biology, Texas 
A&M University, College Station, Texas 
77843-3258; (SO) Hiroshima University, 
Faculty of Applied Biological Science, Si- 
tami, Saijo-cho, Higashi-Hiroshima 724, 
Japan. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 339-361 

A REVISION OF THE GENUS MICROPROTUS 

RICHARDSON WITH DESCRIPTIONS OF 

TWO NEW SPECIES, M. ACUTISPINATUS 

AND M. LOBISPINATUS 

(ASELLOTA, ISOPODA, CRUSTACEA) 

George D. F. Wilson, Oleg G. Kussakin, and Galina S. Vasina 

Russian Abstract.— 

PE3IOME 

PeBH3H5i po,na Microprotus Richardson c onncaHHeM 

AByx hobbix BimoB (Asellota, Isopoda, Crustacea) 

Yhjicoh JJjk.JI.®., KycaKHH OJ., BacHHa T.C. 

B pe3yjibTaTe npo#ejiaHHOH peBH3HH po#a Microprotus, paHee OTHOCHMoro k 
npHMHTHBHOMy cpe^H Asellota ceMencTBy Janiridae, onncaHbi #Ba HOBbix BH#a 
H3 6aTHajiH panona KypnjibCKHx octpobob, bha Storthyngura paradoxa Birstein 
nepeHeceH b stot tkq po#. IIoKa3aHO, hto, HecMOTpa Ha OTcyTCTBHe nuaBaTe- 
jibHbix nepeonoAOB, poA Microprotus AOjmeH 6biTb nepeHeceH b oaho H3 Han- 
6ojiee bwcoko cneuHajiH3HpOBaHHbix ceMeHCTB Asellota— Munnopsidae s.L, zuifl 
npeACTaBHTejien KOTOporo xapaKTepHbi njiaBaTenbHbie 3a,zniHe napbi njieono^OB. 
npHBOA^Tca ^narH03bi ceMeHCTBa Munnopsidae s.L b o6i>eMe, BKjnonaiomeM 
ceMencTBa Eurycopidae, Ilyarachnidae h Munnopsidae s. s., po,a;a Microprotus 

H BCeX nflTH H3BeCTHbIX BHAOB 3TOrO pO^a. 06cy)K£ai0TC5I pOACTBeHHbie CB5I3H 

Microprotus c /royrHMH poflaMH Munnopsidae b nacTHOCTH c po^OM Storthyn- 
gura, k KOTOpOMy oh HanSonee 6jih3ok, xota Storthyngura— THnnnnafl Eury- 
copidae c njiaBaTejibHbiMH nepeono/jaMH, oTcyTCTByiomHMH y Microprotus. 
OScyac/jaeTCfl Bonpoc, flBjiaeTCfl jih OTcyTCTBHe nnaBaTejibHbix nepeono^OB y 
Microprotus njre3HOMOp(J)HbiM npH3HaKOM hjih sto— pe3yjibTaT BTOpHHHoro 
ynpomeHHA. 

Abstract.— The genus Microprotus Richardson, referred earlier to the rela- 
tively plesiomorphic family Janiridae, is revised. Two new species from the 
bathyal zone of the Kurile Islands, M. acutispinatus and M. lobispinatus, and 
Storthyngura paradoxa Birstein are also placed in this genus. Microprotus should 
be included in the Munnopsidae s.L, in spite of its lack of the highly specialized 
posterior natatory pereopods that characterize this family. Diagnoses are given 
for the family Munnopsidae s.L, for the genus Microprotus, and for all five 
species referred to this genus. The relationships between Microprotus and other 
munnopsid genera are discussed, and it is found to be most closely related to 
Storthyngura. Whether the absence of natatory pereopods in Microprotus is a 
plesiomorphic or an apomorphic reversion is discussed; the latter hypothesis 
is favored. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



The status of the janiroidean genus Mi- 
croprotus has been suspect since its incep- 
tion. Richardson (1910) described two in- 
dividuals from the South Bering Sea which 
she put in the family Janiridae, where the 
genus remained until now. Her remarks on 
the genus include the statement (Richard- 
son 1910:116) "This genus has affinities with 
the Munnopsidae" (under Sars' (1899) def- 
inition, this family also included Eurycope 
and Ilyarachna), but later in the text she 
writes "The absence of natatory legs, the 
general form of the body and the style of 
the uropods, however, are characters refer- 
able to the Janiridae." The Munnopsidae 
Sars s.l. (Wilson 1989) is defined by unique 
natatory modifications, including paddle- 
shaped posterior legs with many fringing 
plumose setae. Richardson's (1910) species, 
M. caecus, closely resembles some species 
in the munnopsid genus Storthyngura, ex- 
cept that its posterior legs are not in the least 
paddle-shaped and they have no plumose 
swimming setae. 

Vanhoffen (1914), the author of the genus 
Storthyngura, described a new species, 
Microprotus antarcticus from pleotelson 
fragments. Apparently Vanhoffen did not 
regard this species as a member of Stor- 
thyngura and included it in his currently 
unused and misspelled family Jolellidae. 
Storthyngura, however, has come to include 
a great variety of forms (see reviews by Bir- 
stein 1957, George & Menzies 1968a, b). 
Birstein (1970) included the species para- 
doxa in Storthyngura despite its lack of na- 
tatory posterior pereopods. His comments 
(translated) are revealing: "This inexplica- 
ble trait [lack of swimming limbs] contra- 
dicts not only the diagnosis of the genus but 
even that of the family, although in all the 
other characteristics the new species can be 
regarded as a typical form of the genus Stor- 
thyngura." Birstein (1970) undoubtedly 
overlooked the obscure genus Microprotus 
because all authors, including Wolff (1962), 
had put it in the Janiridae. 

In this paper, we remove the veil of ob- 
scurity from Microprotus by providing new 



descriptions of the its members, and by dis- 
cussing the meaning of its lack of natatory 
limbs, at least from a phylogenetic point of 
view. M. caecus is redescribed, two new 
species are fully described, and diagnoses 
and a key to all species are provided. We 
argue that Microprotus is a derived member 
of the Munnopsidae related to the hetero- 
geneous genus Storthyngura, and that the 
absence of natatory limbs is a reversion to 
a primitive condition, rather than a plesio- 
morphy. 

The Systematic Position of Microprotus 

We present here a formal diagnosis of the 
Munnopsidae Sars, 1899 (sensu Wilson 
1989), to aid the discussion of the system- 
atic position of Microprotus. 

Family Munnopsidae Sars, 1899 s.l. 

Diagnosis (derived from Wilson, 1989).— 
Janiroidea with a distinct natasome: pere- 
onites 5-7 enlarged, muscular, with artic- 
ulations often broadly joined or fused; pos- 
terior ventral nerve cord ganglia fused into 
a single mass in pereonite 5; pleotelson gen- 
erally tapering posteriorly. Pereopods V-VII 
natatory (in most genera) with many long, 
fully plumose setae on margins of broad, 
paddle-like carpi and propodi. Pereopodal 
dactylar claws with trough-like hollow be- 
tween superior and inferior claws enclosing 
distal sensillae. Antennular article 1 broad- 
ened and laterally flattened. 

Discussion. — The above definition of the 
Munnopsidae subsumes the smaller fami- 
lies Eurycopidae, Ilyarachnidae, and Mun- 
nopsidae s.s. The diagnosis includes fea- 
tures that are found in no other family of 
the Janiroidea in this combination. For a 
more detailed discussion, see Wilson 1989). 
The best corroborated phylogenetic trees of 
the Munnopsidae (Wilson 1989) suggest that 
the ancestral munnopsid had the following 
characters in addition to the diagnostic 
characters above: rostrum (a distinct frontal 
projection between the antennulae) present, 
natasomal pereonites and pleotelson freely 



VOLUME 102, NUMBER 2 



341 



articulated, pleopod III rami with many 
plumose setae. 

The diagnosis of the Munnopsidae is pre- 
sented here because Microprotus is a sig- 
nificant exception in its lack of natatory legs 
(Figs. 3D, 7, 10). This genus nevertheless 
has all other diagnostic characters including 
a completely fused natasome, an unusual 
feature for an ambulatory animal (Fig. 1 A- 
B). Given this distribution of characters, two 
possible hypotheses emerge for the classi- 
fication of this genus: (1) the non-natatory 
legs are plesiomorphic within the Munnop- 
sidae, and Microprotus diverged early in the 
evolution of the family before swimming 
was acquired, i.e., it is the sister group of 
the remaining munnopsids; or (2) the non- 
natatory legs are a complete reversion, are 
a distinctive autapomorphy for the genus, 
and other characters must be used to estab- 
lish the affinities of the genus within the 
Munnopsidae. Using the non-natatory pe- 
reopod character alone, one would be forced 
to accept hypothesis 1 (HI) over hypothesis 
2 (H2) because the former is more parsi- 
monious. Several other character com- 
plexes are examined here to establish which 
hypothesis is most likely. The parsimony 
values are counted globally for the Mun- 
nopsidae. 

Cephalon (Fig. IE; Wilson 1989, fig. 38): 
The ancestral munnopsid probably had a 
distinct rostrum, while Microprotus does 
not. HI predicts either that the presence of 
a rostrum in other munnopsids is a rever- 
sion or that Microprotus lost the rostrum 
independently of other munnopsids (two 
evolutionary steps), while H2 predicts that 
only the immediate ancestor of Microprotus 
(and taxa of its clade) lost the rostrum in its 
evolution (one evolutionary step). 

The cephalons of some munnopsids bear 
a synapomorphy called the frontal ridge, a 
supporting bridge between the mandibular 
articulations on both sides of the head. Be- 
cause Microprotus has a frontal ridge, HI 
predicts that the frontal ridge is a synapo- 
morphy of all munnopsids and was subse- 
quently lost in several taxa (e.g., Euryco- 



Table 1 . — A parsimony analysis of two evolutionary 
hypotheses for the non-natatory pereopods V-VII in 
Microprotus. HI = The ancestral munnopsid did not 
have natapods, and Microprotus diverged before na- 
tapods were evolved. H2 = The natapods in Micro- 
protus reverted to an ambulatory state. Values are the 
number of evolutionary steps required in the character 
for the phylogeny of the Munnopsidae (Wilson, 1989) 
given that a hypothesis is true. 



Character 


HI 


H2 


Pereopods V-VII not natatory 


1 


2 


Rostrum absent 


2 


1 


Frontal arch present 


2 


1 


Mandibular notch in cephalon 


2 


1 


Natasome completely fused 


2 


1 


Total parsimony values 


9 


6 



pinae, Munnopsinae) or was developed at 
least twice (two steps). H2, on the other hand, 
only requires that the frontal ridge is a syn- 
apomorphy of the group to which Micro- 
protus belongs (one step). 

In the sister group Acanthaspidiidae and 
in some munnopsid genera, the mandible 
articulates with the head by a rounded pos- 
terolateral border, while other taxa, includ- 
ing Microprotus and Storthyngura, have a 
posterior projection of the mandible that 
articulates with the cephalon in a narrow 
posteriorly-directed slot. HI suggests that 
this feature either evolved twice, or evolved 
once and then was lost in more derived taxa 
(two steps); H2 requires only that it is a 
apomorphy of some group of the munnop- 
sids (one step). 

Natasome (Fig. 1 A-C): The posterior pe- 
reonites and the pleotelson of the ancestral 
munnopsid, although integrated into a unit, 
were probably fully articulated because both 
the Acanthaspidiidae, and several munnop- 
sid genera, such as Amuletta Wilson and 
Thistle, Munnopsurus Richardson, and 
Munnicope Menzies & George are fully ar- 
ticulated both ventrally and dorsally. The 
flexion of the segments, however, is restrict- 
ed and the integration of the natasome is 
obvious in these taxa. Because Microprotus 
has a fully fused natasome, HI requires that 
this was achieved independently from the 



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remaining munnopsids (two steps), while H2 
allows that complete fusion could be a syn- 
apomorphy of a subset of the Munnopsidae. 

Table 1 shows parsimony values for mun- 
nopsid phylogenetic estimates using the 
characters discussed above under HI or H2. 
Although these characters are only a subset 
of the possible features that are useful in 
munnopsid systematics (Wilson 1989), they 
are highly derived features that are unlikely 
to appear more than once in the evolution 
of this family. HI adds nine steps to the 
munnopsid phylogenetic tree, while H2 only 
adds six steps and is therefore more prob- 
able. The ambulatory posterior pereopods 
of Microprotus are best explained as a re- 
version from a swimming form, and must 
be an autapomorphy of this genus alone. 

Furthermore, the natapod to pereopod re- 
version is not simply a reduction of features: 
no natatory landmark is present on pereo- 
pods V-VII of the adult (juveniles are un- 
known at present). There are no natatory 
setae, the carpi and propodi are much longer 
than in any other munnopsid, and these seg- 
ments are tubular rather than flattened. 
Other taxa, such as Syneurycope where the 
pereopodal swimming function may be 
greatly reduced, typically retain the rem- 
nants of these landmarks (e.g., Haugsness & 
Hessler 1979, fig. 31). The detailed similar- 
ity between the anterior and posterior legs 
(Fig. 3B, D) suggests that a genetic transfer 
in pereopodal developmental processes from 
anterior to posterior may underlie this re- 
version (first author's opinion). In the cur- 
rent absence of information on munnopsid 
genetics, further evidence could be obtained 
by examining the pre- and postmarsupial 
development in Microprotus species, once 
juveniles and embryos are collected. 

Microprotus Richardson, 1910 

Microprotus Richardson, 1910:116. 
Storthyngura (pars) Birstein, 1970:352. 

Type species. —Microprotus caecus Rich- 
ardson, 1910. 



Species included. —Microprotus caecus 
Richardson, 1910; M. antarcticus Vanhof- 
fen, 1914; M. paradoxus (Birstein, 1970); 
M. lobispinatus, n. sp.; M. acutispinatus n. 
sp. 

Diagnosis.— Munnopsidae with ambula- 
tory pereopods V-VII: carpus and propodus 
elongate and tubular, not flattened, with op- 
posing margins with rows of unequally-bifid 
setae; all segments lacking plumose swim- 
ming setae. Cephalon with frontal ridge and 
without rostrum. Pereonites 5-7 and pleo- 
telson fully fused, with no free articulations 
either dorsally or ventrally; natasomal pere- 
onites collectively shorter than ambuloso- 
mal pereonites. Body dorsal surface, lateral 
margins of pereonites and pleotelson, coxae 
of pereonites 5-7, and basal articles of an- 
tenna with elongate spines; pleotelson ter- 
minating in pair of elongate spines. Pleo- 
telson with pronounced preanal ridge; ridge 
and anus not covered by opercular pleo- 
pods. Antennular article 1 elongate and sub- 
triangular. Mandible with cuticular projec- 
tion on posterolateral margin articulating 
with cephalon in elongate notch. Uropods 
elongate with tubular protopods and rami. 

Additional description. —Body with elon- 
gate, anteriorly curved spines in following 
pattern: dorsal midline with one spine on 
each pereonite and two on pleotelson; dor- 
solateral spines on pereonites 2-7; lateral 
spines on pereonites 5-7; two pairs of spines 
on lateral margins of pleotelson; one pair of 
spines on posterior tip of pleotelson; one or 
two spines on articles 1-3 of antenna; single 
spines on coxae of pereopods I; paired spines 
on coxae of pereopods II-IV. 

Cephalon without eyes or elongate spines, 
broader than long, with robust laterally pro- 
jecting cheeks at mandibular articulation 
point. Pleotelson broad, in most species 
broader than long; dorsally trilobed, with 
central section separated by troughs from 
dorsally projecting lateral lobes. 

Antennular flagellum elongate, with many 
articles, each article with one or several 
aesthetascs; articles 2, 3 and 5 near same 



VOLUME 102, NUMBER 2 



343 



length, distinctly longer than article 4; fla- 
gellar articles of male very short, wider than 
long, shorter than article 4. Antenna elon- 
gate, length more than two body lengths. 
Mandible normal but body abbreviated 
proximal to dorsal condyle and palp; molar 
process triturative but distally tapering; palp 
functional, longer than mandibular body; 
teeth on incisor process not distinct; lacinia 
mobilis normal with distinct teeth; spine 
row normal with many spines. Maxillipedal 
basis with many coupling hooks; epipod 
elongate, reaching to articulation between 
palp articles 2 and 3, with small rounded 
dorsolateral spine. 

Pereopod I much smaller than posterior 
pereopods, with thin carpus and propodus 
having only simple setae. Pereopods II-VII 
robust, elongate, with strong unequally-bi- 
fid setae on opposing margins of carpus and 
propodus. 

Male pleopod I elongate, much longer than 
wide, with distinct waist midlength, often 
with short thick spines or tubercles on ven- 
tral surface; distal tip with distinct medial 
and lateral lobes; lateral lobe more elongate, 
curving medially posterior to medial lobe. 
Male pleopod II protopod broad, laterally 
rounded, distally curving to short pointed 
tip; protopodal setae on lateral margin short, 
simple, not hemiplumose; endopodal stylet 
shorter than protopod length; exopod very 
short, rounded, with nonprojecting distal 
hook. Female pleopod II broader than long, 
without setae on keel. Pleopod III endopod 
broad and rounded; exopod long, thin, dis- 
tally rounded; exopod and endopod with 
numerous plumose setae. 

Remarks. — The inclusion of M. antarc- 
ticus Vanhoffen is rather dubious because 
the species is known only from fragments. 
Its pleotelson, however, is more similar to 
other species of Microprotus than to Stor- 
thyngura so it must remain in this genus 
until redescribed from new material. We 
(GSV & OGK) have examined the paratype 
of Storthyngura paradox Birstein (1970): this 
species has a strong resemblance to Micro- 



protus and is transferred herein to this ge- 
nus. Two new species, M. lobispinatus and 
M. acutispinatus, extend the known mor- 
phological range of this genus. 

Microprotus has clear affinities with the 
heterogeneous genus Storthyngura, which 
does have well-developed natapods. The two 
genera share the following characters, often 
to small details: cephalon shape and man- 
dibular articulation; dorsal and lateral spi- 
nation; form and orientation of the anten- 
nula; uropod shape; fusion of natasomal 
segments (in some species); shape of venter 
of pleotelson; pleopod shape. Such a long 
list of similar features practically guarantees 
that these two genera form a monophyletic 
taxon within the Munnopsidae. Formal rec- 
ognition of this group as a subfamily, how- 
ever, must wait until the genus Storthyngura 
is revised, and its relationships to Acantho- 
cope are clarified. 

The subgeneric division of Storthyngura 
provided by George & Menzies ( 1 968b) falls 
short of being useful because, although they 
say their analysis was based on 138 char- 
acters, only 3 or 4 characters were given in 
the descriptions of the subgroups; analysis 
of their groupings is therefore extremely dif- 
ficult. Furthermore, the subgroups may have 
dubious meaning because they separate 
species with truncate or forked pleotelson 
tips (as in Microprotus) into different groups, 
even though this may be an important char- 
acter at the generic level. Birstein (1970) 
also found the classification of George & 
Menzies (1968b) to be ineffective. Stor- 
thyngura should be subdivided because the 
type species S. elegans Vanhoffen is flat- 
bodied and broad while S. pulchra Hansen 
(for example) is deep-bodied and has a mus- 
cular natasome. The strongly natatory at- 
tributes of the latter species are more like 
other Munnopsidae, indicating it may have 
diverged early in the evolution of the Stor- 
thyngura group. For the moment, Micro- 
protus is considered to be among a complex 
of genera represented by the polyphyletic or 
paraphyletic genus Storthyngura. 



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Geographic distribution of Micropro- 
tus.— This genus is an Indo-Pacific bathyal 
to upper-abyssal genus with 4 species in the 
northern Pacific: from 40°N to 52°14'N in 
the Japan and Kurile-Kamchatka Trenches, 
and off the South Aleutian Islands. A single 
Antarctic species has been found in the Da- 
vis Sea. This genus inhabits depths in the 
range of 5 50-3400 m. 

Key to the Species of Microprotus 

la. Posterior spines on pleotelson ex- 
tending to tip of uropods; uropodal 
exopod distinctly shorter than en- 

dopod Microprotus antarcticus 

Vanhoffen, 1914 

lb. Posterior spines on pleotelson not 
extending to tip of uropods; uro- 
podal exopod subequal to endo- 
pod 2 

2a. Lateral spines on pleotelson broad, 
nearly as broad as long; male pleo- 
pod II stylet not tapering to thin 
hair-like tip 3 

2b. Lateral spines on pleotelson thin, 
much longer than broad; male 
pleopod II stylet tapering to thin 
hair-like tip 4 

3a. Dorsal and lateral spines on body 
distally thick and typically round- 
ed; pleotelson width (excluding 
spines) distinctly greater than 
length; male pleopod II stylet long, 
distinctly longer than half proto- 

pod length 

Microprotus lobispinatus, n. sp. 

(Figs. 5-7) 

3b. Dorsal and lateral spines on body 
distally thin and pointed; pleotel- 
son width (excluding spines) sub- 
equal to length; male pleopod II 
stylet short, distinctly less than half 

protopod length 

.... Microprotus acutispinatus, n. sp. 

(Figs. 8-10) 

4a. Dorsal and lateral spines on body 
strongly denticulate; posterior 
spines on pleotelson not recurved 



dorsally; pleotelson dorsal surface 
without pair of low bumps anterior 
to uropods; pleotelson width sub- 
equal to length 

Microprotus paradoxus 

(Birstein, 1970) 
4b. Dorsal and lateral spines on body 
finely denticulate; posterior spines 
on pleotelson strongly recurved 
dorsally; pleotelson dorsal surface 
with pair of low bumps anterior to 
uropods; pleotelson width greater 
than length 

Microprotus caecus Richardson, 

1910 (Figs. 1-4) 

The Species of Microprotus 

Microprotus caecus Richardson, 1910 
Figs. 1-4 

Types. — Holotype copulatory male, pleo- 
telson damaged, some legs missing, others 
loose in vial, body length 12.0 mm, USNM 
(United States National Museum no.) 
39521. Paratype copulatory male, dam- 
aged, partly dissected, some limbs missing, 
estimated length 11.2 mm, USNM 39917. 

Type locality.— Albatross station 4781, 
52°14.5'N, 174°13'E, off Cape Sabak, Agat- 
tu Island, Aleutian Archipelago, depth 544 
m, bottom consisting of fine gray sand and 
pebbles. 

Diagnosis. — Cephalon width 1.1 times 
pereonite 1 width. Dorsal and lateral spines 
on body distinctly pointed distally, not 
rounded, finely denticulate. Pleotelson width 
1.3 times length; dorsal surface with pair of 
low bumps anterior to uropods; lateral spines 
narrow, length 2.8 times basal width. Male 
pleopod II stylet short, length 0.45 of pro- 
topod length, with curved hair-like distal 
part. Uropods much longer than posterior 
spines of pleotelson; endopod length sub- 
equal to exopod length. 

Additional description of males.— Body 
(Fig. 1 A-D): Cuticle heavily calcified. Pres- 
ervation color white. All margins and spines 
rugose, with many fine denticles. Body wid- 
est at pereonite 5, body length 2.7 times 




Fig. 1 . Microprotus caecus, holotype male: A-B, Dorsal and lateral views of body; C, Ventral view of 
natasome; D, Ventral oblique view of cephalon and ambulosome, pereopods on right side omitted. Paratype 
male: E, Lateral oblique view of cephalon. Scale bars: 1.0 mm. all. 



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width. Articular margins of anterior per- 
eonites set in shallow transverse grooves. 
Dorsal-most parts of pereonites 1-4 also 
with shallow transverse groove. Pleotelson 
with pair of short broad spines or bumps 
posterior to last median spine; bumps not 
visible in lateral view; lateral spines on pleo- 
telson flattened dorsoventrally, postero- 
lateral spines anterior to uropods triangular 
in cross-section; posterior spines strongly 
curving dorsally. 

Antennula (Fig. 2H, I): Left antennula of 
holotype approximately 1 mm long; length 
0.83 of body length. Surface of articles 1 
and 2 denticulate. Article 1 length 1.9 times 
width; medial margin with dense group of 
denticles adjacent to insertion of article 2; 
dense patch of broom setae proximal to 
group of denticles on medial margin. Article 
2 length 0.39 of article 1 length (including 
distal lobe). Article 3 subequal to article 2. 

Antenna (Fig. 2H): Length 2.8 times body 
length. Articles 2 and 3 with broad flattened 
spines projecting ventrally. Article 3 with 
spine in approximate position of scale. Ar- 
ticles 5 and 6 subequal, length of either 0.22 
of total antennal length. 

Left mandible (Fig. 2A-D): Incisor pro- 
cess with one distinct cusp ventrally, with 
remaining incisive margin sinuous and 
lacking distinct cusps. Lacinia mobilis with 
four cusps, decreasing in length dorsally; 
ventral surface of lacinia mobilis with tuft 
of spinules or cuticular hairs. Spine row with 
1 5 members, distal spines with spinules or 
cuticular hairs on basal ventral surface. Mo- 
lar process tapering distally, distal width 
roughly half proximal width; posterior mar- 
gin of triturative surface with 8 doubly set- 
ulose setae and row of basal denticles, ven- 
tral tooth present. 

Maxillula (Fig. 2E): Outer lobe with 12 
large spine-like toothed setae. Inner lobe 
distal tip broadly rounded, not extending 
beyond medial end of outer lobe setal row. 

Maxilliped (Fig. 2G): Epipod length 0.93 
of basis length; width 0.37 of length; distally 
rounded. Basal endite with 13 coupling 



hooks and approximately 5 apical fan setae 
distally. Palp article 2 width subequal endite 
width. 

Pereopods (Fig. 3A-E): Total length in- 
creasing posteriorly with pereopod I dispro- 
portionately smaller than others: pereopod 
I approximately half body length while pe- 
reopods II- VI increasing from slightly less 
(approximately 5% shorter) to near body 
length. Pereopodal bases: II-IV length sub- 
equal; I length 0.7 1 length of II; V-VII long- 
er than II-IV, increasing in length poste- 
riorly, length ratios with II-IV 1.1, 1.3, 1.5. 
Ischia of all pereopods distinctly shorter than 
corresponding bases: ischia I and VI less 
than half bases length, ischium II length 0.69 
of basis II length. Pereopod I propodus with 
many fine, blunt-tipped, aesthetasc-like se- 
tae; opposing margins of propodus and car- 
pus with only fine setae. Setae of pereopods 
II— VI: single row of unequally-bifid setae on 
opposing margins of propodus and carpus; 
propodus with numerous fine setae both 
dorsally and ventrally. Superior dactylar 
claws II-VI with triangular or blunt projec- 
tion on posterior margin. (Pereopod VII not 
known.) 

Male pleopod I (Fig. 4A-E): Sympod wid- 
est at proximal insertion, tapering to less 
than half proximal width at midlength, and 
expanding to 0.75 of proximal width dis- 
tally. Sympod with two broad irregular rows 
of low spines or bumps on ventral surface. 
Distal tip of in situ sympod resting exactly 
at tip of pleopod II protopod. Lateral lobes 
elongate, pointed, proximally broad, distal 
parts curving medially. Medial lobes short, 
rounded, approximately 0.25 of lateral lobe 
length, with numerous long curved simple 
setae reaching to tip of lateral lobe on distal 
margin. 

Male pleopod II (Fig. 4F, G): Protopod 
broad, laterally rounded, with fringe of sim- 
ple setae laterally and distally; length 1.4 
times width; ventral surface with concavity 
lateral to internal musculature for exopod. 
Endopod inserting 0.65 of protopod length 
from pleopod insertion; exopod short, distal 



VOLUME 102, NUMBER 2 



347 




Fig. 2. Microprotus caecus, paratype male, mouthparts: A-D, Left mandible: A. Mandible, dorsal view; B, 
Incisor process and lacinia mobilis, anterior view; C, Molar process, anteromedial view; D. Distal part of 
mandible showing spine row; E, Right maxillula. ventral view; F, Right maxilla, ventral view; G. Right maxilliped. 
ventral view. Holotype male: H, Cephalon, dorsal view, showing antennula and antenna; I. Antennular basal 
articles, in situ. Scale bars: 0.2 mm. all except for H, 2.0 mm long. 



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Fig. 3. Microprotus caecus, paratype male, pereopods: A, Pereopod I, lateral view; B, Pereopod II, lateral 
view; C, Enlargement of adjacent parts of carpus and propodus of pereopod VI; D, Pereopod VI, medial view; 
E, Pereopod VI, enlargement of dactylar claw, lateral view. Scale bars: 1.0 mm. 



VOLUME 102, NUMBER 2 



349 



hook not protruding, with small distal tuft 
of fine simple setae; exopodal extrinsic mus- 
culature small compared to size of proto- 
pod, extending only halfway to lateral mar- 
gin of protopod. 

Pleopod III: Exopod distally rounded, 
with approximately 17 plumose setae. En- 
dopod distally broad, rounded, very thin, 
fringed with numerous setae. 

Uropod (Fig. 4H): Long, thin, with short 
setae, length 0.69 of pleotelson length. En- 
dopod and exopod subequal, length of either 
0.87 of protopod length. 

Remarks. —Microprotus caecus is known 
only from two adult male specimens; fe- 
males or juveniles have not been collected. 
This species may be distinguished from oth- 
er species of the genus by the following char- 
acters: cephalon only slightly wider than 
pereonite 1; spines distinctly pointed, not 
rounded distally; pleotelson much wider 
than long; lateral pleotelson spines that are 
much longer than their basal width; and 
subequal uropodal rami. Overall, the other 
species are similar to M. caecus, perhaps 
with the exception of M. antarcticus whose 
pleotelson shape differs rather more. 

Richardson (1910: fig. 38) illustrated the 
holotype of Microprotus caecus with distinct 
articulations between the natasomal seg- 
ments. Study of the type material reveals 
that the natasome is fully fused with no free 
articulations. The dorsal surface of the na- 
tasome, however, does have slight ridges 
that are either remnants of the articulations 
or exterior expressions of apodemes. These 
ridges further corroborate the contention 
that pereonites 5-7 are natasomal in origin 
because each ridge extends well into the seg- 
ment anterior to it (Fig. 1A), as is typical 
with most munnopsids. 

Microprotus lobispinatus, new species 
Figs. 5-7 

Types. — Holotype male, 12 mm long, 
(Zoologicheskogo Instituta Akademiya 



NAUK (ZIN) No. 1/8 1 502). Paratype male, 
11 mm, (ZIN No. 2/81503). 

Type locality. —Pacific Ocean, near Itu- 
rup Island, 44°52'N, 149°27'E, depth 910- 
920 m, habitat: muddy sand with stones, 
25 Jul 1984, coll. B. Sirenko. 

Diagnosis. — Dorsal and lateral spines on 
body thick and rounded distally; spines fine- 
ly denticulate. Pleotelson width (excluding 
spines) 1.2 times length; lateral spines broad, 
length subequal to basal width. Male pleo- 
pod II stylet 0.66 of protopod length, with- 
out curved hair-like distal part. Uropods 
much longer than posterior spines of pleo- 
telson; endopod length subequal to exopod 
length. 

Additional description of adult male (Fig. 
5). — Body somewhat robust, front half sub- 
equal in width to natasome. Body length 
approximately 2.2 times body width with- 
out lateral spines; body widest at pereonite 
4. All dorsal spines distinctly flattened, often 
with concave front surface, usually curved 
forwards, sometimes widened in middle 
part. All dorsal spines with finely spinulous, 
rasp-like surface. 

Cephalon broad, approximately three 
times wider than long; frontal margin 
broadly concave. 

Pereonite 1 considerably narrower than 
cephalon, with single medial spine only. 
Pereonite 2 approximately 1 . 5 times wider 
than pereonite 1, slightly wider than pere- 
onite 4 and slightly narrower than pereonite 
3. Dorsal surface of pereonites 2-7 with three 
spines. Medial spines gradually decreasing 
in length from pereonite 2 to 6, medial spine 
on pereonite 7 longer than that on pereonite 
6. Dorsolateral spines nearly as long as me- 
dial spines; on natasomal pereonites, much 
nearer to medial spines than on pereonites 
2-4. Lateral margins of pereonites 1 and 4 
rounded, pereonites 2 and 3 nearly truncate, 
with produced posterolateral angles. Coxal 
plates of pereonites 2-4 long, strongly pro- 
duced, each plate with long spine-like lobes, 
anterior lobe longer than posterior. Coxal 



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Fig. 4. Microprotus caecus, paratype male, pleopods: A-C, Pleopod I, ventral, dorsal, and lateral views 
respectively; D, E, Enlargements of pleopod I distal tip, ventral and dorsal views respectively; F, G, Right 
pleopod II, ventral and enlarged dorsal view respectively. Holotype male: H, Left uropod, in situ ventral view. 
Scale bars: 1.0 mm. 



plates of pereonite 1 with single anterior 
spine-like lobe. 

Segments of natasome fused medially but 
distinct on lateral parts. Anterolateral an- 



gles of pereonites 5-7 produced outwards 
into stout conical spine-like processes. Cox- 
al plates of pereonites 5-7 small, with 
rounded posterolateral angles. 



VOLUME 102, NUMBER 2 



351 



Pleon length not including posterior spines 
0.3 of total body length. Anterodorsal mar- 
gin of first pleonite slightly marked by shal- 
low transverse depression, posterior margin 
somewhat more distinct. Dorsal surface of 
anterior pleonite convex, with median spine 
strongly curved forwards. Pleotelson broad, 
1.3 times wider than long; lateral margins 
each with two long stout conical spines; an- 
terior spine thicker, located nearly at mid- 
length of pleotelson lateral margin. Dorsal 
surface of pleotelson with flattened spoon- 
like median spine curving forwards. 

Antennula (Figs. 5, 6F) when bent back- 
wards slightly exceeding middle of body; 
basal article more than two times longer 
than broad, its outer margin irregularly con- 
vex, inner margin roughly concave; second 
article inserting near midlength of first ar- 
ticle, approximately one third as long as first 
article, slightly increasing in width distally. 
Antennal second article with two curved 
spines, one spine directed outwards, and 
second inwards. Third article also with two 
spines, outer spine longer than inner one. 

Left mandible (Fig. 6C): Incisor process 
with one tooth. Lacinia mobilis with two 
teeth. Spine row with 12 spines, posterior 
spines longer than anterior ones. Molar pro- 
cess forming truncate cone, cut off distally, 
with small tooth on inner side and shallow 
apical excavation in middle. Third article 
of mandibular palp broad, oval, twisted, 
with numerous marginal setae, longer than 
palp article 1 . 

Maxilliped (Fig. 6 A, B): Endite with 10 
coupling hooks. Epipod approximately three 
times longer than broad, distally rounded, 
with triangular lateral projection. 

Pereopods (Fig. 7A, E, F): Pereopod I 
simple, little differentiated; carpus approx- 
imately 2 times longer than propodus; both 
articles with numerous short setae. Carpus 
1.3 times longer than ischium and merus 
together. Propodus of pereopod II slightly 
longer than carpus, ventral margin with 
dense row of bifid spine-like setae, dorsal 
margin with numerous fine setae; merus 




Fig. 5. Microprotus lobispinatus, male holotype, 
body in dorsal view. 



short, bearing setae on both margins. Car- 
pus and propodus of pereopods II-VII ob- 
long, narrow, subequal in length, ventral 
margins with dense row of short spine-like 
bifid setae; on propodus all setae subequally 
long, on carpus small setae alternate with 
larger setae (not shown in Fig. 7E, F); pro- 
podus with one distal plumose seta. Per- 
eopod VI propodus slightly longer and near- 
ly twice as narrow as carpus. 

Male pleopod I (Fig. 7B): Pair complex 
narrowest at midlength, length 2.8 times 
proximal width. Lateral lobe longer than 
medial lobe, curving medially; medial lobe 
with distal tip rounded, lobe-like, bearing 
numerous setae. 

Male pleopod II (Fig. 7C): Somewhat 
broad; protopod suboval, length nearly 2 
times width, inner distal tip triangular, 
pointed; distal part of outer margin and pos- 
terior margin with setae. As compared with 
M. acutispinatus, endopod relatively long. 



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Fig. 6. Microprotus lobispinatus, male holotype, mouthparts and antennula: A, Maxilliped, dorsal view; B, 
Maxillipedal epipod, dorsal view; C, Right mandible, dorsal view; D, Maxillula, ventral view; E, Maxilla, ventral 
view; F, Antennula. Scale bars: 0.2 mm, all except F, 0.3 mm. 



Fig. 7. Microprotus lobispinatus, male holotype, pereopods and uropod: A, Pereopod I; B, Pleopod I; 
C, Pleopod II; D, Uropod; E, Pereopod II; F, Pereopod VII. Scale bars: 0.2 mm— B-D; 0.3 mm— A; 0.5 mm— 
E,F. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 8. Microprotus acutispinatus, male holotype, 
body in dorsal view. 



Stylet extending slightly beyond distal tip of 
protopod, length 0.7 of total protopod 
length. Exopod small, with dense group of 
fine simple setae on posterior curve. 

Uropod (Fig. 7D): Very long and slender, 
extending much beyond posterior spines of 
pleotelson. Endopod slightly shorter than 
exopod; latter ramus nearly as long as pro- 
topod. 

Remarks. —Microprotus lobispinatus is 
represented by only two adult males from 
one sample. This species can be distin- 
guished from all Pacific species by the wide, 
apically flattened and rounded dorsal spines. 
M. lobispinatus also differs distinctly from 
M. acutispinatus and M. paradoxus by its 
relatively wider pleotelson; the species is 
similar in this respect to M. caecus. M. lo- 
bispinatus is easily distinguished from the 
latter species by having thicker and shorter 
lateral spines, especially on the posterior 
pereonites and the pleotelson, by lacking 
dorsal lumps on the pleotelson anterior to 
the uropods, and by uropodal protopods that 
do not extend beyond the posterior spines 
of the pleotelson (as in M. caecus). This new 
species, like all other boreal species, dis- 
tinctly differs from M. antarcticus, a single 
species from the southern hemisphere con- 
ventionally assigned to this genus, by pos- 
terior spines on the pleotelson considerably 
shorter than the uropods, and by uropodal 
rami that are nearly equal in length. 

Etymology.— "Lobispinatus" the adjec- 
tival form of two classical nouns, means 
"provided with lobe-spines." 

Microprotus acutispinatus, new species 
Figs. 8-1 1 

Types. —Holotype, male 1 3 mm (ZIN No. 
1/81500). Paratypes, 2 males, 2 females, all 
fragmented (ZIN No. 2/81501). 

Type locality. —Pacific Ocean, near Itu- 
rup Island, 44°48'N, 149°31'E, depth 1 100- 
1200 m; habitat: sand with pebbles; 25 Jul 
1984, coll. B. Sirenko. 



VOLUME 102, NUMBER 2 



355 



Diagnosis. —Dorsal and lateral spines on 
body thin and pointed distally; spines finely 
denticulate. Pleotelson width (excluding 
spines) subequal to length; lateral spines 
broad, length subequal to basal width. Male 
pleopod II stylet 0.4 protopod length, with- 
out curved hair-like distal part. Uropods 
much longer than posterior spines of pleo- 
telson. 

Additional description of adult males (Figs. 
8, 1 IB, E). — Body relatively slender, slight- 
ly widening posteriorly, so that natasome 
distinctly wider than pereonites 1-4 and ce- 
phalon; length slightly more than three times 
body width across pereonite 5 without lat- 
eral projections. All dorsal spines rather 
slender, narrow-conical, pointed, their sur- 
faces covered with minute spinules. Dorsal 
spines directed forwards and upwards on 
pereonites and backwards and upwards on 
pleon. 

Cephalon broad, width approximately 
three times length; frontal margin broadly 
concave. 

Pereonite 1 distinctly narrower than ce- 
phalon, with single median dorsal spine. 
Pereonites each with three spines; median 
spines subequal in length. Dorsolateral 
spines on natatory pereonites placed much 
nearer to medial spines than on pereonites 
2-4. Lateral margins of pereonites 1 and 4 
rounded, those of pereonites 2 and 3 nearly 
truncate, posterolateral angles produced 
outwards. Coxal plates of pereonites 2-4 
with two produced, long, spine-like lobes 
(damaged on pereopod III in Figs. 8, 1 1 A- 
C); anterior lobe longer than posterior one. 
Segments of natasome lacking dorsal artic- 
ulations; pereonites 5-7 and pleon not clear- 
ly delimited. Anterolateral corners of pere- 
onites 5-7 produced outwards into stout, 
conical spine-like projections. Coxal plates 
of pereonites 5-7 in dorsal view relatively 
short, suboval in form. 

Pleon length excluding posterior spines 
approximately one third total body length. 
First pleonite small, narrow, convex dor- 



sally, with medial dorsal spine curved back- 
wards. Pleotelson (without projections) only 
slightly broader than long, somewhat pro- 
duced posterior part with pair of long spine- 
like pointed projections directed backwards 
and slightly outwards and two pairs of lat- 
eral long stout conical spines: anterior spine 
considerably thicker and slightly longer, lo- 
cated approximately at midlength of lateral 
margin of pleotelson, posterior spine smaller 
and placed near posterolateral angle of pleo- 
telson. Low broad dorsomedial keel with 
recurved spine-like process in middle part. 

Antennula (Fig. 8) when bent backwards, 
reaching midlength of body; basal article 
two times longer than broad, its outer mar- 
gin irregularly convex, inner margin con- 
cave; second article inserting at midlength 
of basal article, approximately one third 
length of basal article, slightly widening dis- 
tally. 

Antennal (Figs. 8, 1 IB) second article with 
two stout spines, one on inner and another 
on outer margin, third article also with two 
subequal spines. 

Left mandible (Fig. 9D): Incisor process 
with one tooth. Lacinia mobilis with three 
teeth. Spine row with eight setae, posterior 
spines longer than anterior ones. Molar pro- 
cess conical, obliquely cut off distally, with 
four setae on distal posterior edge; distal 
triturating surface with conical tooth. Third 
article of mandibular palp longer than first, 
broad, twisted, with setal row. 

Maxilliped (Fig. 9A): Endite with 1 1 cou- 
pling hook on inner margin. Epipod three 
times longer than broad; distally rounded, 
with triangular lateral projection. 

Pereopod I (Fig. IOC) simple, little dif- 
ferentiated; carpus slender and very long, 
length 1.4 times ischium and merus length 
together and 1.6 times propodus length. 
Ventral and dorsal margins of propodus and 
ventral margin of carpus with rows of short 
stout spine-like bifid setae. 

Pereopod II (Fig. 10D): Propodus some- 
what longer, slightly longer than carpus, with 



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Fig. 9. Microprotus acutispinatus, male holotype, mouthparts: A, Maxilliped and epipod, ventral view; B, 
Maxillula, ventral view; C, Maxilla, ventral view; D, Left mandible, dorsal view. Scale bars: 0.2 mm. 



VOLUME 102, NUMBER 2 



357 



setae only along ventral margin. Carpus with 
four bifid setae on ventral margin. Dorsal 
margin of merus serrate distally. 

Pereopods V-VII (Fig. 1 0E): Carpus and 
propodus long, slender, linear, subequal in 
length, without natatory setae, but ventral 
margins with row of short stout spine-like 
bifid setae. Dactylus short. 

Male pleopod I (Fig. 10A): Pair complex 
much narrower in middle part; length ap- 
proximately 2.5 times proximal width. Lat- 
eral lobe much longer than endopod; distal 
tip of medial lobe narrowly rounded, with 
short row of simple setae arranged in fan- 
like order. 

Male pleopod II (Fig. 10B): Protopod very 
broad, length approximately 1.5 times long- 
er than wide, semicircular, distomedial cor- 
ner slightly produced, pointed; posterior 
margin and distal part of outer margin with 
row of thin simple setae. Endopod and ex- 
opod small, reaching well short of distal tip 
of protopod. Stylet relatively short, extend- 
ing only slightly beyond distal margin of 
exopod and not reaching tip of protopod. 
Exopod with dense group of long, fine, sim- 
ple setae on posterolateral margin. 

Uropods (Fig. 8) long and slender, ex- 
tending beyond posterior spines of pleotel- 
son. Exopod slightly shorter than endopod; 
latter nearly as long as protopod. 

Description of female.— -In general, out- 
line (Fig. 11 A, C, D, F) similar to male; 
body length 12 mm. Dorsum somewhat 
wider than in male. Pleotelson bearing two 
tiny low lumps posterior to central dorsal 
median spine and anterior to uropods, sim- 
ilar to those in M. caecus (lumps not visible 
in lateral view, hidden by swelling of dorsal 
surface of pleotelson lateral to depressed 
medial part). Pleopod II (Fig. 10G) very 
broad, 1.3 times broader than long, ventro- 
medial keel without setae. 

Remarks.— Microprotus acutispinatus is 
represented by five specimens: three adult 
males and two females, of which only the 
holotype male is preserved in relatively good 
condition. The other individuals are badly 



damaged, their bodies having broken in half. 
This species is easily distinguished from M. 
caecus and M. lobispinatus by its narrow 
pleotelson, which is nearly as wide as long 
and similar in this respect to M. paradoxus. 
It differs, however, from the latter species 
by the narrowly conical (not flattened) dor- 
sal spines on the pleotelson which are con- 
siderably shorter and stouter than those of 
M. paradoxus. 

Etymology. — "Acutispinatus, " contain- 
ing the adjectival form of "spinus" and 
modified by "acutus," means "provided 
with pointed spines." 

Microprotus paradoxus (Birstein, 1970) 

Storthyngura paradoxa Birstein, 1970, p. 
334, Figs. 19-20. 

Syntypes. —Types were not found by Dr. 
Mezhov, who looked for them at our re- 
quest in the deep-sea isopod collection stud- 
ied by the late Professor Ya. Birstein. The 
collection was first deposited in the De- 
partment of Invertebrate Zoology of Mos- 
cow University, was later handed over to 
the Institute of Oceanology, and finally was 
deposited in the Zoological Museum of 
Moscow University. 

Type locality. — Kurile-Kamchatka 
Trench, Vityaz Station 5601, 46°26'N, 
152°07'E, 2770-2820 m. 

Additional material. —Japan Trench, Vi- 
tyaz Station 6671, 40°15'N, 143°35'E, 2500 
m. Three adult males (14.8, 14.6, and 12.9 
mm long), one male fragment, one damaged 
female with oostegites (13.8 mm long), two 
fragments of females. This material was 
identified by B. Mezhov who kindly showed 
it to one of the authors (GSV). 

Diagnosis.— Dorsal and lateral spines on 
body thin and pointed distally; spines 
coarsely denticulate. Pleotelson width (ex- 
cluding spines) subequal to length; lateral 
spines narrow, length much greater than 
width. Male pleopod II stylet short. 0.44 of 
protopod length, with curved hair-like dis- 
tal part. Uropods much longer than poste- 



VOLUME 102, NUMBER 2 



359 



rior spines of pleotelson; endopod length 
subequal to exopod length. 

Remarks.— The specimens from the Ja- 
pan Trench in general correspond well to 
descriptions and drawings of syntypes given 
by Birstein. This species is clearly distin- 
guished by considerably thinner and longer 
acute spines on the posterior part of the 
body; these spines are not bent dorsally and 
extend far beyond the distal ends of the uro- 
podal protopodites. The dorsal surface of 
the pleotelson lacks lumps anterior to the 
uropods and the pleotelson is relatively nar- 
row. This species differs from M. lobispi- 
natus and M. acutispinatus by considerably 
longer and thinner spines on the pleotelson, 
from M. lobispinatus by thin dorsal spines, 
and from M. acutispinatus by a flattened 
body. 

Microprotus antarcticus Vanhoffen, 1914 

Microprotus antarcticus Vanhoffen, 1914: 
545_546, Fig. 71a-d. 

Type. —Only a 2 mm long abdomen with 
a portion of the posterior pereonal segment 
and a pereopod basis are known. This ma- 
terial (not examined by the authors) is prob- 
ably at the Berlin State Museum, East Ger- 
many, where other Vanhoffen material has 
been found. 

Type locality.— Antarctic Indian Ocean, 
Davis Sea, Gauss station 30.11.1903, 
65°27'S, 80°33'E, 3398 m. 

Diagnosis. —Lateral and posterolateral 
spines on pleotelson robust and elongate, 
posterior spines extending to distal tips of 
uropods. Uropodal exopod distinctly short- 
er than endopod. 

Remarks. —Microprotus antarcticus may 
actually belong in another genus of the Stor- 
thyngura complex, but until more speci- 



mens are discovered, this species must be 
retained in Microprotus. 

With such scarce material at his disposal, 
Vanhoffen (1914) could not describe this 
Antarctic species adequately, and could only 
give comments, comparing the specimen 
with M. caecus, the only species of the genus 
Microprotus known at that time. According 
to Vanhoffen, Pacific (Arctic in Vanhoffen' s 
term) and Antarctic species differ mostly in 
that the latter has a low tubercle on the lon- 
gitudinal thickening of the caudal segment, 
while in the Antarctic species, the middle 
spine between both lateral spines looks 
abrupt. In addition, there are constrictions 
over the two posterolateral spines, and the 
uropods are equal in length to the distal 
spines, with the outer uropodal ramus being 
only one third of the inner ramus length. 
In M. caecus, all spines are considerably 
smaller in comparison with the abdomen, 
so that uropodal rami, which are subequal 
in length, extend beyond the distal spines. 
Judging from the rest of the posterior pe- 
reonal segment, the lateral spines in M. ant- 
arcticus are directed backwards, while in 
M. caecus, they are directed forwards and 
curved, and the dorsal surface of these seg- 
ments bears spines. The operculum and oth- 
er pleopods of the female, which are sche- 
matically drawn by Vanhoffen in his figure 
7 1 d, do not seem useful in his opinion and 
cannot be used in comparison because the 
male was known only for the northern 
species. 

Vanhoffen (1914) put the genus Micro- 
protus in his family Jolellidae (sic), where 
he also assigned the genera Jolella (sic, cor- 
rectly spelled Iolella Richardson), Janthop- 
sis (sic, correctly spelled Ianthopsis Bed- 
dard), Acanthaspidia Stebbing, Jolanthe (sic, 
correctly spelled Iolanthe Beddard), Rhac- 



Fig. 10. Microprotus acutispinatus, male holotype: A, Pleopod I, ventral view; B, Pleopod II, dorsal view; 
C, Pereopod I; D, Pereopod II; E, Pereopod VII; F, Uropod. Female paratype: G, Pleopod II. Scale bars: 0.2 
mm— A, B, F; 0.3 mm— C; 0.5 mm — D, E, G. 



360 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 11. Microprotus acutispinatus, paratypes, body fragments: A, Female, cephalon and pereonites 1-3, 
dorsal view; B, Male, cephalon and pereonites 1-3, lateral view; C, Female, cephalon and pereonites 1-3, lateral 
view; D, Female, pleotelson and pereonites 6-7, lateral view; E, Male, pleotelson and pereonites 5-7, lateral 
view; F, Female, pleotelson and pereonites 6-7, dorsal view. 



VOLUME 102, NUMBER 2 



361 



ura Richardson, and Jaerella Richardson. 
These genera were earlier referred to the 
family Janiridae, owing to the presence of 
a more or less distinct rostrum, elongate 
lateral lobes on the body segments, and two 
or more lateral spines on the abdomen. In 
Vanhoffen's opinion, the anterior body part 
of Microprotus is most similar to that of 
Iolanthe, but in the last case, the abdomen 
lacks distinct distal lobes. At the same time, 
Vanhoffen noted certain similarities be- 
tween Microprotus and the Munnopsidae, 
to which he also assigned the Eurycopidae: 
similar structure of long antennae, which he 
attributed to the deep-sea mode of life. Van- 
hoffen's comments are important, because 
although Microprotus does not belong to his 
family "Jolellidae," his family concept 
clearly indicates that certain of the deep-sea 
"Janiridae," Iolella at least, may need to be 
recognized as belonging to a separate family 
with its name corrected to Iolellidae. 

Acknowledgments 

This paper had its inception when Kus- 
sakin asked Wilson to examine Microprotus 
caecus to clarify its relationships with the 
new species collected during Russian ex- 
peditions that are described in this paper. 
We would like to thank Dr. Thomas E. Bow- 
man, National Museum of Natural History, 
for loaning the type material of Microprotus 
caecus, carefully reading the manuscript, and 
offering suggestions for its improvement. We 
are also grateful to Dr. Boris Mezhov, Zoo- 
logical Museum of the University of Mos- 
cow, for giving us an opportunity to observe 
Storthyngura paradoxa specimens identi- 
fied by him. This research was partially sup- 
ported by National Science Foundation Sys- 
tematic Biology Grants BSR 82-15942 and 
BSR 86-04573 to Wilson. 



Literature Cited 

Birstein, Ya. A. 1957. Certain peculiarities of the 
ultra-abyssal fauna with the example of the ge- 
nus Storthyngura. — Zoologicheskii Zhurnal, 
36(7):96 1-985. 

. 1970. Additions to the fauna of Isopods 

(Crustacea, Isopoda) of the Kurile-Kamchatka 
Trench. Part I. In Fauna of the Kurile-Kam- 
chatka Trench and its environment.— Akade- 
miya Nauk CCCP, Trudy Instituta Okeanologii 
Im. P. P. Schirshova 86:292-340. (In Russian, 
English Translation: Israel Program for Scien- 
tific Translations, Jerusalem, 1972) 

Haugsness, J. A., & R. R. Hessler. 1979. A revision 
of the subfamily Syneurycopinae (Isopoda: 
Asellota: Eurycopidae) with a new genus and 
species (Bellibos buzwilsoni).— Transactions of 
the San Diego Society of Natural History 1 9( 1 0): 
121-151. 

George, R. Y., & R. J. Menzies. 1968a. Species of 
Storthyngura (Isopoda) from the antarctic with 
descriptions of six new species.— Crustaceana 
14(3):27 5-301. 

, & . 1968b. Distribution and probable 

origin of the deep-sea isopod genus Storthyn- 
gura.— Crustaceana 15(2): 17 1-1 87. 

Richardson, H. 1910. Isopods collected in the North- 
west Pacific by the U.S. Bureau of Fisheries 
steamer 'Albatross' in 1906.— Proceedings of the 
United States National Museum 37(1 701): 7 5- 
129. 

Vanhoffen, E. 1914. Die Isopoden der Deutschen 
Siidpolar-Expedition 1 90 1-1 903. — Deutschen 
Siidpolar-Expedition 15 Zoology, 7:447-598. 

Wilson, G. 1989. Deep-sea Lipomerinae and the 
Munnopsidae (Crustacea: Isopoda: Asellota): a 
systematic revision.— Bulletin of the Scripps In- 
stitution of Oceanography 27:i-xiii, 1-138. 

Wolff, T. 1962. The systematics and biology of bathy- 
al and abyssal Isopoda Asellota.— Galathea Re- 
port 6:1-320. 

(GDFW) A-002, Scripps Institution of 
Oceanography, La Jolla, California 92093, 
USA; (OGK & GSV) Institute of Marine 
Biology, Far East Science Center, Academy 
of Sciences of the USSR, Vladivostok 
690022, USSR. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 362-374 

FOUR SPECIES OF SYNOPIIDAE FROM THE 
CARIBBEAN REGION (CRUSTACEA: AMPHIPODA) 

J. L. Barnard and James Darwin Thomas 

Abstract. —Synopia ultramarina is redescribed from the Gulf Stream and the 
Florida Keys and comments are made on the tangled taxonomy in Synopia. 
Tiron bellairsi, originally described from Barbados, is reported from Belize, at 
the opposite side of the Caribbean Basin. Garosyrrhoe bigarra, a Pacific species, 
is also reported from Belize, but no clear subspecific differences are detected 
as yet between Pacific and Atlantic populations. 



Specimens of Synopia ultramarina Dana 
(1853) are described and illustrated from 
the Gulf Stream and Synopia scheeleana 
Bovallius (1886) is also reported. A key to 
seven species of the genus Synopia was giv- 
en by J. L. Barnard (1972); Andres (1984) 
described 2 additional species. Four of the 
nine species remain poorly described: S. an- 
gustifrons Dana (1853, tropical Pacific), S. 
caraibica Bovallius (1886, Venezuela), S. 
gracilis Dana (1853, tropical Atlantic) and 
S. orientalis Kossmann (1880, Red Sea). 
Species of this genus are rarely reported in 
modern times, perhaps because they occur 
mainly in neritic waters or in epipelagic 
waters of the high seas. Collections made in 
neritic and epipelagic waters rarely are ex- 
amined by gammaridean taxonomists, who 
appear to concentrate on benthic samples. 
Two of the species come from the tropical 
Pacific where little neritic collecting has been 
accomplished this century. Individuals of 
Synopia may actually be mostly nighttime 
emergents into neritic waters though they 
appear to occur in daytime in epipelagic 
waters particularly in or near sargasso-like 
flotsam. We have searched for S. caraibica 
in neritic waters in the daytime in Florida, 
Belize and Trinidad, to no avail. Pereopods 
5-7 of that species have ovate article 2 of 
pereopods 5-7 but otherwise the species is 
well enough described by Bovallius to sug- 
gest it is not a species of Tiron, a better 
known genus than Synopia. 



Synopiidae 

Synopia Dana 

Synopia Dana, 1852:315 (Synopia ultra- 
marina Dana, 1853, selected by J. L. Bar- 
nard, 1969b). -J. L. Barnard, 1972:50. 

Diagnosis.— Forehead protuberant (pos- 
sible exception, S. orientalis)', lateral ce- 
phalic lobe not sharp. Eyes and accessory 
eyes present. Antenna 1 of female shorter 
than pereon, of male much longer than pere- 
on. Mandible with huge palp (mainly article 
2), molar large, triturative or swollen, pil- 
lowlike in shape and poorly triturative (S. 
variabilis). Outer plate of maxilla 1 with eight 
or nine spines. Maxilliped inner plate lack- 
ing large smooth spines; outer plate fur- 
nished medially only with plumose setae 
(no inflexible spines). 

Coxae 1-2 small, similar, truncate dis- 
tally; coxae 3-4 pelagont, coxa 3 very large 
and enfolding small coxa 4. Gnathopods 
simple (propodus of gnathopod 1 often 
tumid), defining spines absent; gnathopod 
2 very slender, dactyl tiny. Pereopods 3-4 
slightly to strongly diversified, articles 4-5 
stout; pereopods 5-7 elongate. 

Pleonites 1-6 neither denticulate nor 
toothed. Uropod 3 exceeding uropods 1-2, 
peduncle short. Telson relatively short for 
family. 

Relationship. —Differing from Tiron in the 
shorter peduncle of antenna 1 , thick man- 



VOLUME 102, NUMBER 2 



363 



dibular palp, presence of only 8-9 spines on 
the outer plate of maxilla 1 , the absence of 
large smooth apical spines on the inner plate 
of the maxilliped, the outer plate bearing 
only medial plumose setae, no smooth 
spines; the much enlarged coxa 3; inflated 
propodus of gnathopod 1; tiny dactyl of 
gnathopod 2; diversity and tumidity of pe- 
reopods 3-4; lack of dorsal body teeth; and 
relatively shorter telson. 

Remarks. —We believe that Barnard 
(1965) misrepresented the mandibular mo- 
lar of S. variabilis as being non-triturative; 
he apparently did not turn the molar far 
enough upward to see the triturative sur- 
face. 



Key to the Species of Synopia 

1. Telson entire 2 

- Telson cleft 4 

2. Apex of telson broad and armed with 

3 widely spread spines rotunda 

- Apex of telson narrow and armed 
with 2 or fewer appressed spines . . 3 

3. Telson with 2 notches separated by 
subtruncate protrusion, bearing only 
setules variabilis 

- Telson with single apical notch 
armed with 2 spines triangula 

4. Forehead not protuberant . . orient alis 

- Forehead protuberant 5 

5. Article 2 of pereopods 5-6 rectan- 
gular or with quadrate posteroven- 
tral corners 6 

- Article 2 of pereopods 5-6 ovate or 
with rounded posteroventral cor- 
ners 7 

6. Article 2 of pereopod 5 narrowly 
rectangular, scarcely twice as broad 
as article 3 and with scarcely pro- 
truding posteroventral corner .... 

angustifrons 

- Article 2 of pereopod 5 widely rect- 
angular, nearly twice as broad as ar- 
ticle 3 and with strongly protruding 
posteroventral corner gracilis 



7. Flagellar articles 2, 3, 4 ... n of an- 
tenna 1 with long "hairs"* . . caraibica 

- Flagellar articles 2, 3, 4 ... n of an- 
tenna 1 lacking long "hairs" 8 

8. Telson scarcely longer than broad, 
apicolateral margins smooth, each 
apex with one stout spine . . scheeleana 

- Telson much longer than broad, 
apicolateral margins crenulate, each 
apex with one stout spine and one 
long seta ultramarina 

Synopia ultramarina Dana 
Fig. 1-4, part 

Synopia ultramarina Dana, 1853:995, pi. 
68, fig. 6a-h.-Bovallius, 1886:613, pi. 1, 
figs. 1-21. 

Diagnosis. —Telson much longer than 
broad, deeply cleft, apicolateral margins 
scalloped-serrate, each apex with one long 
seta, one short setule, medial subapex with 
spine in notch. Forehead protuberant, with 
anterodorsal keel. Article 2 of pereopods 5- 
6 broadly rounded posteroventrally, of pe- 
reopod 7 subrectangular and quadrate pos- 
teroventrally. Flagellar articles 1, 2, 3 on 
antenna 1 of both sexes with at least one 
stout spine, short in male, elongate in fe- 
male. 

Description of female "u" 4.53 mm.— 
Head with low anterior keel; rostrum ex- 
tending downward to shallow keel connect- 
ed to epistome, upper lip weakly bilobate 
from lateral view and bilobate from anterior 
view. Eyes purple in alcohol, pigment dif- 
fuse (omitted in drawings), accessory eye of 
2 ommatidia each stained purple on inner 
apices. 

Accessory flagellum 2-articulate. basal ar- 
ticle elongate, armed with stiff spines: pri- 
mary flagellum short. 8-articulate. articles 
1-5 and 7 each with stiffspine, spines longer 
on articles 1-3 than on 5 and 7. Each incisor 



* Structure unknown, possibly setae, aesthetascs. or 
straw-like scales. 



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VOLUME 102, NUMBER 2 



365 



and left lacinia mobilis with 4 teeth, right 
lacinia mobilis bifid and weakly hook- 
shaped, larger rakers about 4 left, 7 right, 
first 2 or 3 when turned to plane view bifid; 
molar heavily triturative, armed basally with 
large brush of apically feathered setae. Inner 
plate of maxilla 1 with 5 large medial setae 
and 2 short apical setae, outer plate with 8 
spines, apex of palp with 5 tooth-spines (one 
illustration showing 4 as aberrancy), medial 
oral face with groups of 3 and 4 apical setae. 
Two principal apical spines on inner plate 
of maxilliped plumose, with two other api- 
cal setae, one apicoventral seta, two medial 
setae and one ventral locking spine; medial 
margin of outer plate with marginal and 
ventral submarginal rows of plumose setae, 
lateral margin with two long setae, one ap- 
icad, the other strongly subapical, lateral 
base raggedly serrate; article 2 of palp evenly 
and densely setose medially; dactyl short, 
nail elongate. 

Many setae on gnathopod 2 apically 
forked; dactyl with shriveled appearance (but 
this appearance normal). Pereopods 3-4 
with three unlocking setae each, pereopods 
5-7 with two unlocking spines each, short 
and simple on pereopod 7, on pereopods 5- 
6 one spine elongate and with apical hooded 
hook. Gills large, unpleated, on coxae 2-7; 
oostegites, thin, strap-shaped, sparsely se- 
tose, on coxae 2-5. 

Epimera naked below, epimeron 1 sharp- 
ly quadrate, epimeron 2 with small tooth 
and epimeron 3 round-quadrate postero- 
ventrally. Each pleopod with two coupling 
hooks and one plumose accessory spine, pe- 
duncles with 1-3 setae only, rami with 12- 
1 4 articles, inner rami with one fewer article 



than outer rami and with forked basomedial 
spines, generally two. 

Urosomite 3 with posterodorsal spine. 
Outer rami of uropods 1-2 with dorsolateral 
comb, similar comb on inner rami ventro- 
lateral. Telson scarcely longer than pedun- 
cle of uropod 3, cleft about two-thirds, cu- 
ticle covered with notch-like lunes. 

Male "q" 5.84 mm. —Antenna 1 about 
4.0 mm long, antenna 2 about 5.9 mm long. 
Eyes slightly larger than in female. Basal 
article of primary flagellum slightly longer 
than in female and callynophore denser; ar- 
ticles of primary flagellum with extra mid- 
ventral male sensory setules, large spines of 
female reduced in size. Dorsal margins of 
articles 4-5 of peduncle of antenna 2 plus 
articles of flagellum bearing dense male sen- 
sory setules. Epimera and pleopods larger 
than in female, epimeron 2 lacking tooth. 
Uropod 3 more setose than in female. 

Intermediate male "s" 4.33 mm. —Eyes 
scarcely enlarged. Antenna 1 elongate but 
basal spines of flagellum elongate as in fe- 
male. Posterior setules on flagellum of an- 
tenna 1 and anterior setules on antenna 2 
poorly developed. Tooth of epimeron 2 in- 
termediate between male and female. Setae 
of uropod 3 moderately developed. 

Illustrations.— Maxilla 1 magnified more 
than maxilla 2; telson magnified more than 
uropod 3. 

Material. — MFP-1, in Gulf Stream, off 
Grand Bahama Island, 27°26'N, 78°57'W, 
219 m, 8 Jul 1986, coll. Jack Morton, at- 
tracted to night light, female "p" 4.35 mm, 
male "q" 5.84 mm (illustrated), female "r" 
4.40 mm, male "s" 4.33 mm, male "t" 4.79 
mm, female "u" 4.53 mm (illustrated) (and 



Fig. 1. Synopia ultramarina, unattributed figures = female "u"; q = male "q." Capital letters as follows refer 
to parts; lower case letters to the left of capital letters refer to specimens noted in legends; lower case letters to 
right of capitals refer to adjectival modifications in list below: B, body; D, dactyl; G, gnathopod; H, head; I, 
inner plate or ramus; J, prebuccal; L, labium; M, mandible; O, outer plate or ramus; P, pereopod; R. uropod; 
S, maxilliped; T, telson; U, labrum; V, palp; W, pleon; X, maxilla; a, anterior; d. dorsal; o, opposite: r. right; 
s, setae removed; t, left; x, apex. 



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Fig. 2. Synopia ultramarina, unattributed figures = female "u"; q = male "q." Letter codes, see Fig. 1. 



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Fig. 3. Synopia ultramarina, unattributed figures = female "u"; q = male "q." Letter codes, see Fig. 1, 



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6 other specimens). LKFR-5, Florida Keys, 
Looe Key Reef, 7 m, sand plain in front of 
fore reef zone, 9 Oct 1983, 2300 h, night 
light, full moon, coll. J. D. Thomas (35 spec- 
imens). 

Remarks. — Said by Dana (1853) to be 
"rich blue, to nearly colorless, with tinge of 
rich blue along the venter or about the ar- 
ticulations"; our specimens and those of the 
first redescriber, Bovallius (1886), however 
were hyaline with yellowish cast when caught 
alive, mouthparts, especially maxillipeds 
with red cast under microscopy seen to be 
represented by tiny red granular inclusions 
below the chitinous exoskeleton. One may 
wonder if Dana might not also have col- 
lected S. scheeleana; he called the "male 
specimen" of S. ultramarina "ultramarine" 
but labeled it on his plate as S. gracilis and 
changed its identification in the published 
text; S. gracilis has since been recognized as 
a distinct species but has never been clari- 
fied adequately; unfortunately Dana's ma- 
terial is presumed to have been lost in the 
great fire of Chicago, 1871; see other notes 
below. 

Despite the color, Bovallius chose this 
species to represent Dana's ultramarina on 
the basis of serrate telson and conformity 
in pereopods 5-7; we agree that this solution 
is correct on morphological grounds. 

Relationship.— Differing from the well 
defined S. scheeleana Bovallius (1886) in 
the longer telson bearing apicolateral ser- 
rations and at least one long apical seta on 
each lobe. 

Synopia caraibica Bovallius (1886) has 
article 2 of pereopod 7 ovate like that of 
pereopods 5-6. Synopia angustifrons Dana 
(1853) and S. gracilis Dana (1853) have ar- 
ticle 2 of pereopods 5-7 rectangular or with 
quadrate postero ventral corners. There is 
the possibility that Dana misinterpreted 
these legs as there is no indication he dis- 
sected them off the animal and pressed them 
flat. Synopia orientalis Kossmann (1880) 
does not have a protuberant forehead. The 
telson of S. variabilis Spandl (1923, see J. 



L. Barnard, 1965), S. triangula Andres 
(1984) and S. rotunda Andres (1984) is un- 
cleft. 

Distribution. —Tropical western Atlantic; 
specific localities include Cuba, Bahamas, 
Florida Keys and Dana's 8-12°S, 11- 
14 1 /4°W, 4-7°S, 21-25°W; neritic and pre- 
sumably epipelagic on the high seas; attract- 
ed to night lights. 

Synopia scheeleana Bovallius 

Synopia scheeleana Bovallius, 1886:16-18, 
pi. 2, figs. 22-29. -Stebbing, 1888:799- 
804, pi. 52.-Chevreux, 1900:54.-Reid, 
1951:233. 

Material. — Gulf Stream, 40 miles off 
Florida coast between Miami and Fort 
Pierce, 10 Dec 1987, R/S Sea Diver, male 
"a" 2.52 mm, specimen "b" 1.80 mm. 

Remarks. —Live color deep blue to s-blue, 
nos. 178 to 179, as matched to ISCC-NBS 
Centroid Color Chart 2106; we described 
this in the laboratory as cobalt blue; spec- 
imens courtesy of Dr. Mary E. Rice, Direc- 
tor, Smithsonian Research Station at Link- 
port. 

Distribution. —Atlantic Ocean, off Ft. 
Pierce, Florida to Barbados; tropical Pacific 
Ocean; assumed to be epipelagic. 

Tiron bellairsi Just 
Fig. 4, part 

Tiron bellairsi Just, 1981:259-263, figs. 1-4. 

Diagnosis. —Accessory eye composed of 
2 and 1 separated ommatidia; accessory fla- 
gellum 2-articulate; labrum densely spi- 
nose; palp of mandible replaced by seta; in- 
ner plate of maxilla 1 long and thin, fully 
setose medially but setae small and widely 
spread, outer plate with 10 spines forming 
elongate multi- sigmoid cluster, palp re- 
duced, about half as long as outer plate; in- 
ner plate of maxilla 2 with row of submar- 
ginal setae; inner plate of maxilliped lacking 
thick simple spines; outer plate of maxil- 



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369 




Fig. 4. Left upper = Synopia ultramarina, male "q." Right, Tiron bellairsi, male "o," note 3 renderings of 
outer plate on maxilliped. Letter codes, see Fig. 1 . 



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liped abnormal, lacking major spines, bear- 
ing plumose and tube-setae; dactyls of pe- 
reopods 3-7 stubby, nails scarcely curved; 
article 2 of pereopods 6-7 without special 
setal row, article 2 of pereopod 7 weakly 
crenulate and setulose posteriorly; pereon- 
ites 5-7 and pleonites 1-3 dorsally crenu- 
late, pleonite 4 with medium dorsal tooth, 
pleonite 5 with large dorsal tooth, pleonite 
6 lacking tooth; outer ramus of uropod 1 as 
long as inner ramus but outer ramus of uro- 
pod 2 slightly shortened; rami of uropod 3 
apically blunt; telson with one apical spine 
and 3 setae on each side, no dorsal or medial 
spines. 

Remarks. —We have redrawn minute de- 
tails of several mouthparts and the telsonic 
apex; we assume these are congruent with 
the Barbados population. The diagnostic 
format above is revised to include new char- 
acters not used by Barnard (1972). 

Material— JDT Bel-53, Belize, Carrie 
Bow Key, 34 m, sand trough near fore reef 
drop-off, in coralgal mud, 1 1 Jun 1980, coll. 
J. D. Thomas, male "o" 2.49 mm (illus- 
trated), Tjuvenile "p" 1.96 mm. 

Distribution.— Barbados, Holetown (type- 
locality), 5-? m ("SCUBA" depths); Belize, 
Carrie Bow Cay, 34 m. 

Garosyrrhoe bigarra (J. L. Barnard), 

new synonymy 

Figs. 5-6 

Syrrhoites bigarra J. L. Barnard, 1962:73- 

75, fig. 1. 
Garosyrrhoe disjuncta J '. L. Barnard, 1969a: 

224-225, fig. 30. 

Taxonomy.— Syrrhoites bigarra and Ga- 
rosyrrhoe disjuncta are the male and female 
of the same species. The male is character- 
ized by the larger head with blunter ros- 
trum, elongate antenna 2 and more aes- 
thetascs and male setae on antennae 1-2. 
Barnard (1 962) overlooked the complex na- 
ture of the dorsal teeth on pereonite 7 to 
pleonite 3. 



The Caribbean specimens are smaller than 
those from the eastern Pacific, where the 
species has been found between about 25°- 
32°N. As yet no specimens from the Pan- 
amanian isthmus region have been found 
and until comparisons between specimens 
from exactly similar latitudes and thermal 
regimes from Pacific and Atlantic waters can 
be compared, the small differences we find 
in the Atlantic specimens cannot be attrib- 
uted to speciation. 

Both sexes of material from Belize have 
fewer serrations on the epimera and the fe- 
male lacks serrations on article 6 of pereo- 
pod 4. Other differences between bigarra 
and disjuncta cited by Barnard (1969a) are 
inconsistencies lacking specific value as 
demonstrated in the present material and 
reexamination of the earlier materials. For 
example, the ventral spine on the inner plate 
of the maxilliped and the dorsally promi- 
nent pleonal teeth were overlooked in var- 
ious specimens; differences among pereo- 
pods are simply variables. Any of the 
differences, albeit minor, between individ- 
uals from different oceans, could be attrib- 
uted to the smaller adult size of Caribbean 
specimens. Small potential distinctions of 
specific value between Atlantic and Pacific 
populations may be present in articular pro- 
portions and armament patterns but con- 
firmation of these requires study of larger 
suites of Pacific material than are now avail- 
able. The following descriptions contain only 
new information and points we judge to be 
necessary to amplify and rectify the former 
depiction of this genus and species; illustra- 
tions are provided for characters which are 
difficult to visualize from descriptions. 

Description of male "x" 3.12 mm. —Pe- 
duncular articles 1-2 of antenna 1 with nu- 
merous dorsal aesthetascs, multiple aesth- 
etascs also present on articles 1-2 of 
accessory flagellum and article 1 of primary 
flagellum; on antenna 2 similar aesthetascs 
on article 4 of peduncle, aesthetascs on ar- 
ticle 5 apically curled, basal articles of fla- 
gellum also with multiple aesthetascs. 



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Fig. 5. Garosyrrhoe bigarra, unattributed figures = male "x"; w = female "w." Letter codes, see Fig. 1 



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Fig. 6. Garosyrrhoe bigarra, male "x." Letter codes, see Fig. 1. 



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373 



Right and left mandibular incisors with 
three large and one small teeth, each lacinia 
mobilis with five teeth, first raker on each 
side broadened, more so on left than on 
right mandible, right and left mandibles each 
with total of two and three rakers. Outer 
plate of maxilla 1 with 1 1 spines (versus 9 
in Pacific material), inner plate with 6 me- 
dial setae, palpar apex with 4 spines, one 
apicolateral seta, 2 mediofacial setae. Inner 
plate of maxilliped with 5 subapical setae, 
2 medial setae, one apicomedial spine, one 
ventral hooked locking spine; outer plate 
with 8 marginal spines and one apical spine- 
seta. 

Articles 4-5 of pereopod 5 more sparsely 
armed than in Pacific male (pereopods 6-7 
in Pacific male unknown). Dactyl of pereo- 
pod 7 lacking comb present on pereopods 
3-6. Serrations on epimera 1,2,3 = 2-4-9 
(versus 4-7-12 on Pacific male). Details of 
one dorsal pleonite plus uropod 3 and telson 
are shown in illustrations. 

Female "w" 3.40 mm (body slightly con- 
tracted).— Serrations on epimera 1,2,3 = 
2-2-9 (versus 5-6-13 in Pacific female). 

Material— JDT BEL-53, Belize, Carrie 
Bow Cay, 1 1 Jun 1980, forereef sand trough, 
29 m, coralgal sand-mud, coll. J. D. Thom- 
as, female "w" 3.40 mm, male "x" 3. 12 mm 
and 4 other specimens. Also found in sim- 
ilar samples: JDT BEL-9B,C,D; JDT BEL- 
14; -19; -53. Associated in amphipod com- 
munity on coralagal sediments at depths of 
27 to 33 m with Netamelita and Hornellia. 

Distribution. —Southern California at 
border with Mexico and in Gulf of Califor- 
nia, 24-44 m; Caribbean Sea, Belize, 27- 
33 m. 

Acknowledgments 

We thank Captain Jack Morton and stu- 
dents of the Marine Field Projects class of 
1986, Florida Institute of Technology, Mel- 
bourne, Florida, for collecting amphipods 
while at sea. The second author was sup- 
ported by NSF grant BSR-85 1 5 186. 



Literature Cited 

Andres, H. G. 1984. Zwei neue Synopiiden (Crus- 
tacea: Amphipoda: Gammaridea) aus dem war- 
men zentralen Nordatlantik. — Mittheilungen aus 
den Hamburg Zoologisch Museum und Institut 
81:109-116. 

Barnard, J. L. 1962. Benthic marine Amphipoda of 
southern California: Families Tironidae to 
Gammaridae— Pacific Naturalist 3:70-1 15. 

. 1965. Marine Amphipoda of atolls in Mi- 
cronesia.— Proceedings of the United States Na- 
tional Museum 117:459-552. 

. 1969a. A biological survey of Bahia de los 

Angeles Gulf of California, Mexico, IV. Benthic 
Amphipoda (Crustacea).— Transactions of the 
San Diego Society of Natural History 15:175- 
228. 

. 1969b. The families and genera of marine 

gammaridean Amphipoda.— United States Na- 
tional Museum Bulletin 271:535 pp. 173 figs. 

. 1972. A review of the family Synopiidae 

(=Tironidae), mainly distributed in the deep sea 
(Crustacea: Amphipoda). — Smithsonian Con- 
tributions to Zoology 124:1-94. 

Bovallius, C. 1886. Amphipoda Synopidea.— Nova 
Acta Regiae Societatis Scientiarum Upsaliensis 
13:1-36. 

Chevreux, E. 1 900. Amphipodes provenant des cam- 
pagnes de /'//z'ra«^//^(1885-1888).-Resultats 
de Campagnes Scientifiques Accomplies par le 
Prince Albert I Monaco 16:iv and 195 pp., 18 
pis. 

Dana, J. D. 1852. On the classification of the Crus- 
tacea Choristopoda or Tetradecapoda. — Amer- 
ican Journal of Science and Arts (2) 14 [Appen- 
dix]:297-316. 

. 1853. Crustacea. Part II. — United States Ex- 
ploring Expedition 14:689-1618, atlas of 94 pis. 
published in 1855. 

Just, J. 1981. Tiron bellairsi sp. n. (Amphipoda, Sy- 
nopiidae) from coral sand in Barbados, with 
notes on behaviour.— Zoologica Scripta 19:259— 
263. 

Kossmann, R. 1880. Malacostraca. Zoologische Er- 
gebnisse einer im Auftrage Koniglichen Aca- 
demie der Wissenschaften zu Berlin ausgefuhr- 
ten Reise in die Kiistengebiete des Rothen 
Meeres. Zweite Halfte, Erste Lieferung. III. Ma- 
lacostraca 67-140, pis. 4-15. Leipzig: Wilhelm 
Engelmann. 

Reid, D. M. 1951. Report on Amphipoda (Gam- 
maridea and Caprellidea) of the Coast of Trop- 
ical West Africa. — Atlantide Report 2: 1 89-29 1 , 
58 figs. 

Spandl, H. 1923. Amphipoden der 'Pola'-Expediti- 
onen in das Rote Meer.— Akademie der Wis- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



senschaftlichen in Wien, Anzeiger, Jahrgang 60: 
17-20, 87-89, 111-112. 
Stebbing. T. R. R. 1888. Report on the Amphipoda 
collected by H.M.S. Challenger during the years 
1873-76. Report on the scientific results of the 
voyage of H.M.S. Challenger during the years 
1873-1876, Zoology 29: xxiv and 1737 pp., 
210 pis. 



(JLB) NHB-163, Division of Crustacea, 
Smithsonian Institution, Washington, D.C. 
20560; (JDT) Newfound Harbor Marine In-, 
stitute, Rt. 3, Box 170, Big Pine Key, Flor- 
ida 33043. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 375-384 

A NEW SPECIES, AMPELISCA BURKEI, 
(CRUSTACEA, AMPHIPODA) FROM FLORIDA 

J. L. Barnard and James Darwin Thomas 

Abstract.— A new species Ampelisca burkei, from Florida, is described. This 
species appears to be very close to A. lobata Holmes, from the eastern Pacific 
Ocean, but differs in the shape and setation of article 4 on pereopod 7, and the 
presence of an antero ventral spine-seta on article 5 of pereopod 7. 



The new species, Ampelisca burkei de- 
scribed herein is a twin of A. lobata Holmes 
from the eastern Pacific. The differences be- 
tween the two species are very small but 
recognition of both is justified by the trend 
for taxonomists to find and validate at the 
specific level small differences between Pa- 
cific and Atlantic amphipods. Small mor- 
phological characters are extremely consis- 
tent over wide geographic areas in the family 
Ampeliscidae. This consistency is not widely 
shared by amphipod groups, and care must 
be taken to document variation in particular 
characters before incorporating them into a 
taxonomic treatment. For example, Bar- 
nard (1980) separated the Pacific Methar- 
pinia oripacifica from the Atlantic M. flor- 
idana by the presence or absence of a single 
spine on the inner plate of the maxillipeds 
(2 spines in Pacific, one in Atlantic popu- 
lations). Dickinson (1982) separated the Pa- 
cific Ampelisca fageri from the Atlantic A. 
schellenbergi on subtle shapes and setation 
patterns of coxa 1, pereopod 7 and uropod 3. 



Ampelisca burkei, new species 
Figs. 1-5 

1 Ampelisca lobata.—]. L. Barnard, 1954b: 2 
(not Holmes, 1908, possible misidenti- 
fied specimens from Colombia and Aru- 
ba). 

Description ofholotype, male, 4.92 mm.— 
Body smooth; urosomite 1 elevated, with 



thick keel, urosomites 2-3 coalesced, with 
strong dorsal saddle. 

Head as long as first three pereonites com- 
bined, slightly longer than tall, with short 
rostrum; head with lobe below attachment 
of antenna 1 , then head sloping down and 
posteriorwards in two steps, no ventral 
tooth. One corneal lens present on each side 
strongly removed from anterior margin, 
second pair on ventral margin far below lat- 
eral lobe. Brown (unusual) pigmentary mass 
present for each lens, similar mass behind 
upper pair. 

Antenna 1 slender, much shorter than an- 
tenna 2, reaching 25 percent along flagellum 
of antenna 2; peduncle short, articles 1-2 
equally long, article 3 short; base of flagel- 
lum forming callynophore; accessory fla- 
gellum absent; main flagellum 5 times as 
long as peduncle, first 2 articles with aes- 
thetascs attached ventrally. Antenna 2 about 
2.4 times as long as antenna 1, about 1.4 
times as long as body, peduncular article 5 
almost as long as 4, anterior margins of ar- 
ticles 3-5 with male setular tufts, flagellum 
3 times as long as peduncle. 

Epistome weakly and obtusely projecting 
anteriorly, labrum incised distally, broader 
than long (tall). Labium normal, with well- 
developed inner lobes. 

Mandible with well-developed strong tri- 
turative molar, incisor toothed; palp 3 -ar- 
ticulate, article 1 short, article 2 longer than 
3, setose, article 3 non-falciform, not dilated 
distally, with 3 D setae, 4 E setae. 

Maxilla 1 : inner lobe with 2 short simple 



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377 



setae, outer lobe with 1 1 spines, three of 
these provided with 2-4 lateral teeth each; 
palp 2-articulate, symmetrical on both sides, 
second article barely dilated distally, pro- 
vided with 4 distal sharp cusps accom- 
panied by 4 spines, 4 subdistal setae and 2 
apicomedial setae. Maxilla 2 with narrow 
lobes, inner lobe shorter than outer, without 
oblique facial row of setae, all medial setae 
marginal. Maxilliped: inner lobe of ordinary 
length, with 2 distal spines intermixed with 
several plumose setae; outer lobe reaching 
apex of palp article 2, bearing row of thick 
spines along medial margin and 2-3 apical 
setae; palp 4-articulate, article 3 not lobed, 
article 4 with main nail about as long as 
remaining part of article. 

Coxae 1-4 much longer than broad, coxa 
1 dilated distally, with convex distal (=ven- 
tral) margin provided with one row of tiny 
setae, lacking posteroventral notch; coxae 
2-4 with increasingly truncate distal mar- 
gins, coxae 2-4 with increasingly sparse set- 
ules, coxa 4 with small posteroproximal lobe 
produced bluntly in proximal part, other- 
wise apposite margins almost parallel. 

Gnathopods 1-2 linear, simple, gnatho- 
pod 1 shorter than gnathopod 2; gnathopod 
1: article 5 scarcely longer than 6, densely 
setose along posterior margins, article 6 ta- 
pering distally with dactyl shorter than ar- 
ticle 6, bearing 5 setules along inferior mar- 
gin and 1 seta on outer margin. Gnathopod 
2: article 5 linear, long, moderately setose; 
article 6 much shorter than 5, tapering dis- 
tally, dactyl like that of gnathopod 1 . 

Pereopod 3 of medium stoutness, with 
articles 4-6 bearing long plumose setae along 
both margins, except posteriorly on article 
6 and anteriorly on article 5; article 4 not 
inflated; dactyl slender, almost straight, 



slightly longer than article 6. Pereopod 4 
like that of pereopod 3, but much longer, 
article 4 setose on both sides to base. 

Pereopod 5: article 2 ovoid, with large 
posterior flangehump, anterior margin with 
one row of plumose setae, article 5 scarcely 
produced distoposteriorly, bearing row of 
spines on apex; article 6 with few long distal 
setae, dactyl short, unguiform, with 2 outer 
cusps. Pereopod 6: article 2 subquadrate, 
with strong posterior lobe and poorly setose 
anterior margin; articles 3-7 like those of 
pereopod 5. Pereopod 7: almost as long as 
pereopod 6 but article 2 large, scarcely ex- 
panding distalwards, lobe reaching middle 
of article 4, bearing sparse plumose setae 
along ventral margin in two disjunct groups; 
article 3 short; article 4 slightly longer than 
3, weakly produced at posterodistal apex 
and bearing one short spine and posterior 
setae; article 5 narrow, with weak postero- 
distal spination and one anterior subdistal 
spine-seta, article 6 weakly tumid, scarcely 
longer than 5, dactyl tumid, tapering rapidly 
and apically pointed, much shorter than ar- 
ticle 6. 

Strongly plaited large gills on coxae 1-6, 
gills generally with 20 + pleats. Pleopods well 
developed, normal, with 2 retinacula and 
one simple accessory retinaculum each. 

Epimera 1-2 weakly convex behind, 
epimeron 1 with posteroventral setule at 
subsharp corner, epimeron 1 with 3 ventral 
setae; epimeron 3 not larger than 2, poste- 
rior margin almost straight, posteroventral 
corner sharply protuberant or obtuse (vari- 
able). 

Uropods 1-2 of ordinary length, uropod 
1 : peduncle slender, outer face of peduncle 
with 5 spines, dorsolateral margin with 3 
spines, apex with cusp, medial corner with 



Fig. 1. Ampelisca burkei, n. sp., unattributed figures, holotype male "h" 4.92 mm; k = male "k" 4.66 mm. 
Capital letters denote main parts in following list; lower case letters to left of capital letters or in body of figure 
indicate modifications as per following list; lower case letters to right of capital letters indicate specimens described 
in captions: A, antenna; C, coxa; D, dactyl; E, epimera; G, gnathopod; H. head; L. lower lip; M, mandible; P. 
pereopod; R, uropod; S, maxilliped; T, telson; U, upper lip; W. pleon; X, maxilla; m. medial. 



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Fig. 2. Ampelisca burkei, n. sp., holotype male "h" 4.92 mm. Letter codes, see Fig. 1. 



VOLUME 102, NUMBER 2 



379 





Fig. 3. Ampelisca burkei, n. sp. Upper, holotype male "h" 4.92 mm. Lower, female "o" 5.66 mm. Letter 
codes, see Fig. 1 . 



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Fig. 4. Ampelisca burkei, n. sp. ? unattributed figures = female "m" 4.40 mm: h = holotype male "h" 4.92 
mm. Arrows on hA2 and mH point to lines denoting undrawn lengths of antennae. Letter codes, see Fig. 1. 





0X2 





oMI 



Fig. 5. Ampelisca burkei, n. sp. Upper, female "m" 4.40 mm. Lower, female "o" 5.66 mm. Letter codes, 
see Fig. 1 . 



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2 spines, medial margin with 5 other spines, 
rami slender, as long as peduncle, curved 
and pointed distally, outer with 2 tiny lat- 
eral spines, medial margin with 4 spines, 
inner ramus lacking lateral spines, with 7 
medial spines. Uropod 2: peduncle slender, 
slightly shorter than outer ramus, with 3 
moderately spread dorsolateral and 3 tightly 
packed dorsomedial spines, with apicolat- 
eral cusp, outer ramus smaller than inner, 
of triangular cross-section, with 4-5 dorsal 
spines, 2 basomedial spines and one tiny 
lateral spine, inner ramus with 1 lateral 
spine, and 7-8 medial spines. Uropod 3 long, 
subfoliaceous, strongly exceeding apex of 
uropod 2: peduncle stout, short, with 2 me- 
dial spines; rami extending subequally, in- 
ner foliaceous, with sharp apices, outer much 
narrower than inner, setose laterally, and 
apicomedially, inner lined with stout me- 
dial spines paired with plumose setae, extra 
setae in gaps between spines. 

Telson reaching one third along rami of 
uropod 3, longer than broad, incised 90 per- 
cent of its length, each lobe expanding in 
middle and then tapering distally, notch 
narrowly beveled apically, each with apical 
spinule, 2 dorsal axial spines and subbasal 
pair plus satellite of penicillate setules (one 
spine occasionally substituted by pair of set- 
ules). 

Male "k. "—Antenna 2 only 0.8 times as 
long as in holotype. 

Female "m" 4.40 mm. — Oostegites on 
coxae 2-5 strap-shaped, narrow, weakly se- 
tose. Antenna 1 slightly exceeding peduncle 
of antenna 2, peduncle short, article 2 as 
long as 1, flagellum with few basal aesthe- 
tascs. Antenna 2 elongate, article 5 of pe- 
duncle slightly shorter than article 4. Gills 
much smaller than in male and with about 
5-6 pleats. Hump on pleonite 4 lower than 
in male. Dorsolateral margin on peduncle 
of uropod 1 with 3 spines, inner apex with 
spine pair, medial margin with 3 other 
spines, outer ramus with 2 lateral and 3 me- 
dial spines, inner ramus with 3 medial 
spines; peduncle of uropod 2 with 2 dor- 



solateral spines, inner apex with triad of 
spines, medial margin with one other spine, 
outer ramus with 3 dorsal and one medial 
spines, inner ramus with 4 medial spines, 
other spine positions noted for male with- 
out spines. Outer ramus of uropod 3 slightly 
shorter than inner, with 5 lateral setal po- 
sitions bearing 1-2 setae each, apex notched 
and with 2 setae; inner ramus with 4 api- 
colateral setules, inner margin with 5 spine- 
notch positions. Telson with only one dorsal 
spine on each lobe besides one apical spine. 

Notes on the redescription of Ampelisca 
lobata by Dickinson, 1982.— Dickinson's 
figure 5 of a female from North Bank Island, 
British Columbia, does not show details of 
pereopod 7 but in the small view presented 
lacks posterior setae on article 6 (a pre- 
sumed oversight) and lacks the anteroven- 
tral seta-spine on article 5 which is present 
in the Floridian material and was shown to 
be absent by Barnard on males (1954a). 
Dickinson does not show the basal aesthe- 
tascs on the flagellum of female antenna 1 . 
Both inner rami of uropods 1-2 of Dick- 
inson's specimen bear one lateral spine but 
none is present on Floridian female mate- 
rial; we assume this may have connection 
with size differences because the British Co- 
lumbian material also has many more pe- 
duncular and inner marginal rami spines 
than our females. 

Illustrations. —Left male uropod 3 drawn 
ventral side up. 

Etymology. —Named for the late William 
W. Burke, III, an original "Son of Bennett" 
(honorary society of biologists at Louisiana 
State University), who first worked with 
Ampelisca in the marshes of Louisiana. 

Holotype. -USNM 195153, male "h" 
4.92 mm, illustrated. 

Type locality. —Florida Keys, Looe Key 
Reef, forereef, 8 m, Apr 1982, full moon 
2200 hours, night diving, attracted to night 
light, coll. J. D. Thomas. 

Material— Type locality, male "i" 4.72 
mm, male "j" 4.84 mm, male "k" 4.66 mm 
and 29 other specimens. Females collected 



VOLUME 102, NUMBER 2 



383 



at same site, LKFR-1C, 18 Apr 1982, from 
algal turf community on Acropora cerxicor- 
nis forereef, 8 m, coll. J. D. Thomas, female 
"1" 4.69 mm, female "m" 4.40 mm (illus- 
trated), female "o" 5.66 mm (illustrated) 
and 10 other specimens. 

Relationship. —Ampelisca burkei is easily 
distinguished from such sympatric species 
as A abdita Mills (1964), A. xadorum Mills 
(1963) and A. declivititis Mills ( 1 967) by the 
short article 3 of pereopod 7. The following 
species of Ampelisca bear similarities to A. 
burkei: A. agassizi Judd (see Dickinson 
1982), A. holmesi Pearse (1908) (see Goeke 
& Gathof 1983), A. panamensis J. L. Bar- 
nard (1954a), A. parapanamensis J. L. Bar- 
nard (1954b), A. verrilli Mills (see Bousfield 
1973), A. lobata Holmes (see Dickinson 
1982), A. hancocki J. L. Barnard (1954a), 
A. millerij. L. Barnard (1954a), A. cucullata 
J. L. Barnard (1954a), and A. romigi = is- 
ocornea J. L. Barnard (1954a). 

Ampelisca burkei differs: from eastern Pa- 
cific A. lobata in the weaker expansion and 
fewer setae on article 4 of pereopod 7, the 
presence of an anteroventral spine-seta on 
article 5 of pereopod 7 and the lack of a 
protrusion apically on article 5 of pereopod 
5 in the female; from A. verrilli in the slender 
pereopod 7, short article 2 of antenna 1 , lack 
of tooth on epimeron 3 and widely expand- 
ed head; from A. cucullata and A. agassizi 
in the slender articles 2-6 of pereopod 7, 
and short article 2 of antenna 1; from A. 
romigi in the smaller postero ventral lobe on 
article 4 of pereopod 7, less beveled article 
2 of pereopod 7, short article 2 of antenna 
1 and narrower posterior flange-lobe on ar- 
ticle 2 of pereopod 5; and from the male A. 
isocornea form by the large spines on female 
uropod 3, much larger process of pleonite 
4 and lack of facial spines on article 6 of 
pereopod 7; from A. hancocki in the shape 
of pereopod 7, numerous spines of uropods 
1-2, longer antenna 1 and shape of coxa 4; 
from A. milleri in the short article 3 and 
general shape of pereopod 7, dense spina- 
tion of uropods 1-2, and shape of coxa 4; 



from A. panamensis and A. parapanamensis 
in the broad head and narrow articles 4-5 
of pereopod 7; and from A. holmesi in the 
short article 2 of antenna 1 , narrower pereo- 
pod 7 and broader head. 

As far as we know, this species differs 
from all other American species of Ampe- 
lisca in the brown (versus red) pigment of 
the eyes; the species therefore looks super- 
ficially like a species of Byblis. 

Distribution.— Florida Keys, Looe Key 
Reef, 0-8 m. 

Acknowledgments 

We thank Linda B. Lutz of Vicksburg, for 
inking our drawings; Janice B. Clark of 
Smithsonian for laboratory assistance; and 
the following referees who helped improve 
this study: Dr. James K. Lowry of the Aus- 
tralian Museum, and Dr. Thomas E. Bow- 
man of Smithsonian Institution; and anon- 
ymous referees. The junior author was 
supported by Grant BSR 8515186 from the 
National Science Foundation. 

Literature Cited 

Barnard. J. L. 1954a. Amphipoda of the family Am- 
peliscidae collected in the Eastern Pacific Ocean 
by the Velero III and Velero IV.— Allan Han- 
cock Pacific Expeditions 18:1-137, pis. 1-38. 

. 1954b. Marine amphipoda of Oregon. — Or- 
egon State Monographs. Studies on Zoology 8: 
103. 

. 1980. Revision of Metharpinia and Micro- 

phoxus (marine Phoxocephalidae Amphipoda 
from the Americas). — Proceedings of the Bio- 
logical Society of Washington 93:104-135. 

Bousfield. E. L. 1973. Shallow-water Gammaridean 
amphipoda of New England. Ithaca & London. 
Cornell University Press, vii-xii. 312 pp.. 13 
figs.. 69 pis. 

Dickinson. J. J. 1982. The systematics and distri- 
butional ecology of the family Ampeliscidae 
(Amphipod: Gammaridea) in the Northeastern 
Pacific region I. The genus Ampelisca. — Publi- 
cations in Biological Oceanography. National 
Museum of Canada 10:1-39. 

Goeke. G. D.. & J. M. Gathof. 1983. Amphipods of 
the family Ampeliscidae (Gammaridea). II. 
Notes on the occurrence of Ampelisca holmesi 
in the northern Gulf of Mexico. — Gulf Research 
Reports 7:289-291. 



384 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Holmes, S. J. 1 908. The amphipoda collected by the 
United States Bureau of Fisheries Steamer, "Al- 
batross," off the west coast of North America, 
in 1903 and 1904, with descriptions of a new 
family and several new genera and species.— 
Proceedings of the United States National Mu- 
seum 35:489-543. 

Mills, E. L. 1963. A new species of Ampelisca (Crus- 
tacea: Amphipoda) from Eastern North Amer- 
ica, with notes on other species of the genus.— 
Canadian Journal of Zoology 41:971-989. 

. 1964. Ampelisca abdita, a new amphipod 

crustacean from Eastern North America.— Ca- 
nadian Journal of Zoology 42:559-575. 

. 1967. The biology of an ampeliscid amphi- 



pod crustacean sibling species pair.— Journal of 
the Fisheries Research Board of Canada 24:305- 
355. 
Pearse, A. S. 1908. Descriptions of four new species 
of amphipodous Crustacea from the Gulf of 
Mexico.— Proceedings of the U.S. National Mu- 
seum 34 (1594):27-32. 

(JLB) NHB-163, Department of Inver- 
tebrate Zoology, Smithsonian Institution, 
Washington, D.C. 20560; (JDT) Newfound 
Harbor Marine Institute, Rt. 3, Box 170, 
Big Pine Key, Florida 33043. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 385-393 

AEGLA DENTICULATA LACUSTRIS, NEW 

SUBSPECIES, FROM LAKE RUPANCO, CHILE 

(CRUSTACEA: DECAPODA: ANOMURA: AEGLIDAE) 

Carlos G. Jara 

Abstract.— A subspecies of Aegla denticulata Nicolet, 1849, is described as 
A. d. lacustris. The new subspecies is restricted to the preandean Lake Rupanco, 
in the River Bueno basin, Osorno Province, Chile, while the nominate sub- 
species is widespread in small, moderately fast running streams in the Central 
Valley of Valdivia, Osorno, and Llanquihue provinces. A. d. lacustris differs 
from A. d. denticulata in having markedly more prominent spinulation along 
the branchial margins of the carapace and upon the chelipeds. 



Aegla denticulata Nicolet, 1849, is one of 
the best known Chilean aeglids. Its mor- 
photype is easily distinguished from that of 
all other species of Aegla Leach (Schmitt 
1942). Extensive collections made by the 
author in central southern Chile have shown 
that A. denticulata ranges between Cautin 
and Chiloe (Jara 1980 and unpublished 
data), living in small to medium-sized rivers 
along the Chilean Central Valley. 

A large population of A. denticulata was 
found occupying the sublittoral of Lake Ru- 
panco (Osorno Province). When I com- 
pared the specimens collected in the lake 
with those collected in rivers, it became ap- 
parent that the lake specimens differed 
markedly in the expression of the carapace 
ornamentation. The spininess of the lacus- 
trine specimens is remarkably more prom- 
inent than that of the riverine animals. 

Since this highly ornamented morpho- 
type was found to be restricted to Lake Ru- 
panco, this variant population was consid- 
ered to represent a geographic isolate of A. 
denticulata, thus deserving a description as 
a new subspecies (as defined by Mayr 1 969). 

Aegla denticulata lacustris, new subspecies 
Fig. la, c, e, g, i, k, m, o 

Holotype. — Instituto de Zoologia, Uni- 
versidad Austral de Chile (IZUA), IZUA 



C-686, adult male, Huillin Cove on north- 
western shore of Lake Rupanco (40°49'S; 
72°28'W), Chile, 25 m depth, baited crab 
trap, 30 Apr 1984, coll. C G. Jara. 

Allotype.— Adult female, 8 Sep 1984, 
IZUA C-686, same locality, collecting de- 
vice and depth as for holotype. 

Paratypes. — IZUA C-687; eight adult 
males (PI to P8), 30 Apr 1984; eight adult 
females (P10 to PI 6), 8 Sep 1984; one ovig- 
erous female (P9), 3 Apr 1984; same local- 
ity, collecting device and depth as for ho- 
lotype. Four juveniles (PI 7 to P20), 7 Sep 
1984, 18 m depth, Scuba hand collection. 

Diagnosis. — External margin of antero- 
lateral lobe of carapace with row of tiny 
spinules; summit of middorsal carina with 
irregular narrow band of minute scales; dis- 
todorsal margin of merus of chelipeds with 
transverse row of small but well defined 
spines; posterior branchial margin of cara- 
pace definitely denticulate; summit of dor- 
solateral carina on second abdominal epi- 
meron with irregular row of scales; 
laterodorsal area of tergum of second ab- 
dominal segment markedly protuberant, its 
apex scaly; dorsum of merus and carpus of 
second to fourth pereiopods clearly dentic- 
ulate; denticles on anterior branchial mar- 
gin conical; palmar crest of chelipeds deeply 
incised. 

Description.— Carapace ovoid, markedly 



386 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



elevated along middorsal line forming well 
denned keel extending from rostrum to pos- 
terior margin of carapace. Dorsal surface of 
carapace and abdomen finely punctate; when 
dry, polished and glossy brilliant. Short stiff 
setae protrude from punctae, some together 
with minute lenslike scales. 

Rostrum prominent, straight, broad- 
based, extended in horizontal plane but 
sometimes slightly recurved at tip. Form 
varying, from strictly triangular to ligulate. 
Rostral margins well defined and slightly 
upturned, fringed with row of closely packed 
minute conical scales pointing towards ros- 
tral apex. Distal third flattened, merging with 
rostral carina into acute conical apex. Ros- 
tral carina narrow, sharp, with row of mi- 
nute scales not reaching rostral apex. Prox- 
imal end of rostral carina marked by two 
small pits just in front of protogastric lobes. 
Epigastric prominences blunt, slightly pro- 
tuberant. Protogastric lobes pronounced, 
their uppermost part with small field of 
lenslike scales. Orbits wide, deep, limited at 
external angle by acute conical orbital spine; 
short row of acute scales just posterior to 
orbital spine on orbital margin. Anterolat- 
eral angle of carapace produced as strong 
acute conical spine extending from broad 
flattened anterolateral lobe. Tip of spine 
reaching at least middle of cornea. Extraor- 
bital sinus a deep V-shaped notch. External 
margin of anterolateral lobe with row of 
small acute spines variable in number and 
size. Free angle of first hepatic lobe pro- 
duced as strong, acute, dorsally recurved 
spine bearing from two to five scales at base. 
Second and third hepatic lobes also well 
marked and bearing spiniform scale at fron- 
tal angle and row of minute scales along free 
margin. Gastric area quite elevated at mid- 
line, then sloping down both sides of cara- 
pace, reaching broad concavity at dorsum 
of hepatic lobes. Uppermost part of mid- 
dorsal carina rugose and scaly throughout. 
Cervical groove distinctly marked but not 
deeply incised. 

Anterior branchial area raised towards 



midline but leveling into horizontal plane 
towards free margin. Frontal angle pro- 
duced as one or two coalescent, strong, 
sharply acute conical spines. Anterior bran- 
chial margin laterally expanded in row of 
from seven to nine single or multiple tipped 
prominent sharply acute denticles that 
scarcely decrease in size from anterior to 
posterior. Row of 9-1 1 tiny conical scales 
mounted atop small tubercles along external 
branchial suture. 

Posterior branchial area with complicat- 
ed topography; inner part raised and leveled 
with cardiac area, then sloping towards lat- 
eral margin, which rises again above mar- 
ginal zone leaving behind wide longitudinal 
trough. Margin markedly expanded in row 
of 9-1 1 clear-cut conical, single or multiple 
tipped, markedly recurved spinelike denti- 
cles. Anteriormost and posteriormost den- 
ticles smallest. At limit between anterior and 
posterior branchial margin, first posterior 
denticle overlaps last anterior denticle. Pos- 
terior margin of carapace thick, more pro- 
tuberant at middle section, which occasion- 
ally forms broad based tubercle. 

Abdominal terga deeply sculptured. Me- 
dian section markedly protuberant espe- 
cially at frontal edge, which forms moder- 
ately sharp transverse carina. On pleural 
area, near laterodorsal pit on second ab- 
dominal segment, dorsal surface raised 
frontally forming rotund broad-based 
prominence bearing at summit from two to 
five tiny scales. Anterolateral angle of sec- 
ond abdominal epimeron produced as strong 
spine tipped with sharp conical scale. It 
stands at frontal end of narrow oblique ca- 
rina extending over anterior two thirds of 
epimeral dorsum. Crest of carina with row 
of closely juxtaposed minute scales decreas- 
ing in size posteriorly. Lateroventral angle 
of epimeron acute; if rounded off, bearing 
two or three acute scales. Lateral angle of 
third and fourth epimera sharply acute. Tel- 
son plate dimerous, subpentagonal. 

Third thoracic sternum broad, its ventral 
surface protuberant, rugose, bearing tiny 



VOLUME 102. NUMBER 2 



3S" 



scales at frontal tip and sometimes one or 
two scales along median line. Fourth ster- 
num with long robust spinelike tubercle 
markedly compressed into sagittal plane, 
tipped with one major and two to three mi- 
nor scales. Fifth sternum slightly swollen 
but not forming tubercle. Frontal margin of 
sixth sternum bearing one or more tiny scales 
in transverse row. 

Chelae subtriangular to ovoid, dorsoven- 
trally depressed. Dorsal surface quite punc- 
tate and covered by numerous small scales 
atop minute tubercles. Scales on proximal 
external surface lenticular and flat, then be- 
coming more acute and prominent towards 
finger tips. Two low. relatively narrow ridges 
on dorsal surface of chelae, one starting from 
upper articulating condylus at carpus-prop- 
odus joint and ending at condylus of prop- 
odus-dactylus joint. Uppermost part of this 
ridge with irregular row of scales larger on 
proximal end. Second ridge along median 
line of chela extending from carpus-propo- 
dus joint to proximal end of cutting edge of 
fixed finger: it is marked by straight row of 
acute scales. Finger cutting edges closely fit- 
ted along distal two thirds but with small 
gap at proximal third. Dorsal edge of dac- 
tylus with sharply prominent conical tuber- 
cle at proximal end. followed distally by row 
of smaller acute tubercles or enlarged scales. 
Palmar crest semicircular in outline, deeply 
incised, formed by from two to three stout 
single or multiple tipped acute spines arising 
from thick bases. Smaller spines or spinules 
attached to basis of body of main spines. 
Frontal margin of propodus above propo- 
dus-dactylus joint with acuminate tubercle: 
spinules are found on both sides of it. Ven- 
tral surface of chela with low but well de- 
fined carina between lower condylus of car- 
pus-propodus joint and tip of fixed finger. 

Carpus strongly armed. Carpal crest with 
two or three sharp, long, robust spines: one 
or two small tubercles behind proximal 
spine. Carpal lobe spiniform. as long as sec- 
ond spine of carpal crest. Row of from five 
to seven spiniform tubercles external to and 



parallel to carpal crest. Second row of small- 
er but acute tubercles external to above row. 
Distodorsal margin of carpus fringed by 
acute scales. Ventral surface with acute con- 
ical spine as long as second spine in carpal 
crest. Distoventral angle below carpus-pro- 
podus condylus with small spiniform tu- 
bercle. 

Dorsal margin of merus with from five to 
eight single or multiple tipped distinct spines 
decreasing in size proximally. Spinules oc- 
casionally present among bases of spines. 
External face roughened by scales, mostly 
on upper half. Distodorsal vertex with large, 
somewhat flattened procurved spine longer 
than the longest in row behind. Margin ex- 
ternal to it with row of sharp conical spi- 
nules. Inner ventral margin with from two 
to five spiniform tubercles in row behind 
single or double tipped spine at distal end. 
External ventral margin scabrous or tuber- 
culate. Distal end with spine as large as that 
on inner ventral margin. One to three small 
acute tubercles variable in size in front of 
latter. Articular lobe at distalmost end of 
same margin spiniform. Inner ventral mar- 
gin of ischium with low broad nodule on 
proximal end. Distal end with conical spine. 
Between both ends from one to four acu- 
minate tubercles variable in size. Distodor- 
sal angle with short but acute spine slightly 
displaced over outer surface. 

Distoventral margin of ischium of pe- 
reiopods fringed with minute scales: one of 
these scales occasionally larger just behind 
ischiomeral joint. Dorsal margin of merus 
with row of small but clear-cut acute den- 
ticles decreasing in size proximally: disto- 
dorsal angle noticeably spiniform: ventral 
margin scabrous bearing minute scales in- 
termingled with short stiff setae. Dorsal 
margin of carpus also with small denticles: 
distodorsal angle spiniform. Dorsal and 
ventral margin of propodus with dense nar- 
row band of scales and setae. Dorsal surface 
of all pereiopodal articles except dactyl mi- 
cronodulose and squamous. 

Color (in life). — Dorsum of cephalotho- 



388 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Aegla denticulata lacustris (holotype), a, c, e, g, i, k, m, o; A. denticulata denticulata Nicolet, 1849, 
male 25.7 mm CL from River Puquitre (IZUA C-686), b, d, f, h, j, 1, n, p. a, b, Dorsal view of carapace; c, d, 
Lateral view of carapace; e, f, Left margin of carapace (ventral); g, h, Second right pereiopod; i, j, Inner view of 
ischium and merus of left cheliped, carpus seen from above; k, 1, Ventral view of right cheliped; m, n, Chela 
and carpus (dorsal); o, p, Lateral view of left second abdominal epimeron. 



rax and abdomen highly variable among in- 
dividuals, ranging from orange-yellow to al- 
most black. Though variation between both 
extremes is continuous, an attempt was 
made to quantify the frequency with which 
different patterns appear in the population. 
The following patterns were defined and 
their frequency recorded (Table 1): 

Pattern A: Dorsum of cephalothorax and 
abdomen uniformly orange-yellow to ter- 
racotta, occasionally scattered with small, 
irregular, dark brownish spots. 

Pattern B: Dorsum of cephalothorax and 
abdomen variegate; extensive reticulate dark 
green or dark brown design on a khaki or 
light brown background. 

Pattern C: Dorsum of cephalothorax and 
abdomen uniformly dark greenish brown or 
dark reddish brown. 



Pattern D: Dorsum of cephalothorax and 
abdomen variegate; small to extensive dark 
green patches on a lighter blue-green or 
brown-green background. 

Pattern E: Dorsum of cephalothorax and 
abdomen variegate; small but numerous 
yellowish or light greenish brown spots on 
a uniform dark brown background; in some 
specimens dorsal carina bluish tinged. 

Pattern F: Dorsum of cephalothorax and 
abdomen variegate; ample irregular blue- 
black or brown-green-black patches on a 
lighter blue-gray or gray- violet background. 

In all specimens, the dorsal surface of the 
pereiopods has the same color pattern as 
the dorsum of the carapace but with darker 
stripes across the carpus and merus. The 
distal half of the chelae is always white or 
yellowish-white, contrasting markedly with 



VOLUME 102. NUMBER 2 



389 



the darker proximal half. The ventral sur- 
face of the body and appendages is milky 
white or slightly yellowish. The articular 
membrane of the appendages is bright pur- 
ple-red. 

In formalin fixed specimens, the dorsum 
of the carapace is wine-red in color, whereas 
in alcohol preserved specimens the color 
changes to light brown or tan. 

Variation and measurements. —Making 
allowances for minor variation in the size 
and number of spines along the upper mar- 
gin of the merus and dorsum of the carpus 
of the chelipeds. adults in the type-series are 
quite similar. Juveniles (PI 7-20) have many 
features of the adult morphotype only 
slightly developed (spines, tubercles, etc.) 
but are still recognizable. The major differ- 
ence between adults and juveniles is that in 
the latter the tegumental scales are replaced 
by short fine setae, most notably on the dor- 
sum of the chelae and on the abdominal 
terga. 

Somatometric data of specimens in the 
type-series are give in Table 2. Measure- 
ments were made with calipers to the near- 
est 0. 1 mm: symbols for the morphometric 
characters in the first column are given by 
Jara & Lopez (1981). 

Distribution. —Known only from the type 
locality. Attempts to collect A. denticulata 
lacustris in Lake Puyehue. 1 2 km to the north 
of Lake Rupanco. failed. In July 1984. dur- 
ing the winter high-water period of Lake 
Rupanco. two small specimens (LC about 
10 mm) were collected in a protected site at 
the bank of River Rahue. 1 50 m from the 
outlet of the lake. These specimens were 
most probably washed out from the lake by 
the effluent current. 

Habitat and natural history. — Lake Ru- 
panco is the southernmost of three major 
preandean piedmont lakes included in the 
River Bueno basin, and is the fifth in size 
among Chilean lakes. It has a maximum 
depth of 274 m and an area of 233.4 km : . 
Limnologically. it is an oligotrophic. cold 
temperate monomictic lake that circulates 
at 9.6°C in midwinter and reaches 19°C at 



Table 1. — Frequency of color patterns in A. dentic- 
ulata lacustris. from Lake Rupanco. Color patterns de- 
fined in text. Sample size: 116 adult specimens col- 
lected at 25 m depth on 30 Apr 1 984. with baited traps. 
Sex unrecorded. 







Frequency 




Pattern 


Absolute 




Relative (%) 


A 


5 




4.31 


B 


11 




9.48 


C 


3 




2.59 


D 


39 




33.62 


E 


29 




25.00 


F 


29 




25.00 



epilimnion in summer. Dissolved oxygen is 
high throughout the year (9.1 to 12.2 mg/ 
liter): conductivity is low (40.6 to 54.0 mS) 
and calcium is scarce (2.1 to 4.8 mg liter) 
(Campos et al. 1985). Lake Rupanco lies at 
118 m above sea level. Its eastern half is 
surrounded by massive mountains and vol- 
canoes (Puntiagudo and Casablanca) of the 
Andes Cordillera. Its affluents are small to 
medium-size rivers of pluvial or pluvio-gla- 
cial regime. Its effluent (River Rahue) dis- 
charges about 100 m 3 sec on the average 
(Campos et al. 1985). Lake Rupanco is of 
glacial origin. The inferred chronology of 
the regional glacial events suggests that Lake 
Rupanco attained its present form about 
1 1.000 years B.P. (Mercer 1972). 

Underwater observations, made by the 
author between and 30 m depth at Huillin 
Cove, showed that A. d. lacustris occupies 
the sublittoral zone from 8 m depth down- 
ward. Its vertical distribution appears closely 
related to that of the soft substrate on which 
crabs alternatively walk and collect particles 
with their maxillipeds. Activity is markedly 
higher at night, while during the day a great 
proportion of the crabs rest buried in the 
substrate. No data are available on the diet 
of A. d. lacustris, but it can be presumed 
that they feed on fine-grained detrital mat- 
ter, though they were seen consuming a dead 
fish lying on the bottom of the lake. Berried 
females appear at the end of the summer 
period (March), and the release of newborn 
crabs from the brooding abdominal cham- 



390 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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VOLUME 102, NUMBER 2 



391 



ber of the females occurs at the beginning 
of the subsequent spring (October). 

At Lake Rupanco, A. d. lacustris coexists 
with Aegla abtao Schmitt and the crayfish 
Samastacus spinifrons (Philippi). A. abtao 
occupies the littoral zone and the upper sub- 
littoral to 1 5 or 20 m depth. However, be- 
low 4 m only large adult males are found. 
The bulk of A. abtao's population lives 
among and under big stones and boulders 
in the littoral. S. spinifrons is found in the 
sublittoral at least down to 80 m depth, only 
on soft substrate. Biological interactions 
among these three species of decapod crus- 
taceans are suspected but the only one ob- 
served so far is predation by large A. abtao 
males upon juveniles of A. d. lacustris. Oth- 
er faunal components of the benthic com- 
munity at Lake Rupanco are freshwater 
mussels of the genus Diplodon and Pisidi- 
um, aquatic snails of the genus Chilina, the 
minute janirid isopod Heterias exul (Bow- 
man et al. 1987), amphipods, acari, oligo- 
chaetes, chironomid larvae and caenid 
ephemeropteran nymphs. 

Etymology.— The trivial name lacustris, 
from Latin, alludes to the fact that this sub- 
species of A. denticulata is apparently re- 
stricted to a lentic environment. 

Comparison.— To compare lacustrine and 
riverine specimens of A. denticulata, four 
crabs (two males and two females) from the 
series from which the neotype of A. dentic- 
ulata was drawn, National Museum of Nat- 
ural History (USNM 169096 = USNM 
80021) (Schmitt 1942) were examined. Ad- 
ditionally, 30 males (LC 18 to 26 mm) col- 
lected in rivers Puquitre and Remehue, 8 
km north of Osorno city, and 50 males (LC 
23 to 33 mm) from Rupanco were examined 
and measured. One male from the Puquitre 
(IZUA C-686) was employed for preparing 
Fig. lb, d, f, h, j, 1 and p. 

A. denticulata lacustris is very similar to 
the nominate subspecies A. denticulata den- 
ticulata Nicolet, 1849, from which it de- 
parts by the morphological features already 
mentioned in the Diagnosis. Additionally, 



Rupanco 




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— i 


n = 50 


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I 




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Valdivia 


V 


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n = 29 




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1.40 1.50 


1.60 


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4> 






Puquitre 




4> 






Valdivia 


\ 


+ 








1.30 


1.40 1.50 


i 
1.60 


CL/ PCL 



Fig. 2. Somatometric comparison of males of A. 
denticulata lacustris (Rupanco) and males of A. den- 
ticulata denticulata (Puquitre and Valdivia). Sample 
labeled Valdivia is a composite made of individuals 
collected all over the Valdivia River Basin (Jara 1 980). 
Vertical line: mean value of somatometric proportion; 
open rectangle: confidence interval for the mean (P = 
0.05). Above: proportion between maximum carapace 
width (CW) and maximum precervical width (PCW). 
Below: proportion between carapace length (CL) and 
precervical length (PCL). 



A. d. lacustris differs from A. d. denticulata 
in the following respects: ridge of dorsal ca- 
rina sharper; dorsal profile of carapace less 
inflated (cf. Fig. lc, d); external margin of 
anterolateral lobe of carapace minutely den- 
ticulate; denticles along ridge of merus of 
chelipeds longer and sharper; tubercles of 
middorsal row of carpus definitely spini- 
form; spine of carpal lobe longer and more 
slender (cf. Fig. li, j); carpal ventral surface 
less hairy; denticles of palmar crest longer 
and slender; ventral angle of second abdom- 
inal epimeron often produced into short 
acute spine (cf. Fig. lo, p). 

Because of the slender and clear-cut ap- 
pearance of the main spines of its carapace, 
together with the cleanness of the carapace 
surface, A. d. lacustris is reminiscent of 
another lacustrine species. A. rostrata Jara, 
but differs from it in that A. rostrata lacks 
the dorsal carina and the median tubercle 
on the fourth thoracic sternum. On the other 
hand, A. d. lacustris lacks the distinctive 
stylet-like rostrum of A. rostrata. 

In Fig. 2 the average and confidence in- 



392 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



tervals (P = 0.05) of two somatometric pro- 
portions for riverine (Puquitre and Valdi- 
via) and lacustrine (Rupanco) samples are 
compared. 

Remarks. — Though at first sight the Lake 
Rupanco morphotype is clearly differentia- 
ble from the riverine morphotype found 
elsewhere in southern Chile, key characters 
that differentiate the two forms appear elu- 
sive. With the exception of the row of tiny 
spinules along the external margin of the 
anterolateral lobe of the carapace, the band 
of tiny scales on the ridge of the dorsal ca- 
rina, and some other minor details, every 
morphological attribute in A. d. lacustris has 
its counterpart in A. d. denticulata. The dif- 
ference is a matter of degree of expression 
rather than of difference of attributes. The 
Rupanco's morphotype appears to be an 
elaborate manifestation of the attributes 
found in riverine specimens. The decision 
to assign the Lake Rupanco population a 
subspecific status is supported by the fact 
that Lake Rupanco's population is, on geo- 
graphical and ecological grounds, an outlier 
as compared to the remaining populations 
of the species. In fact, Lake Rupanco is the 
easternmost locality where A. denticulata has 
been reported in the River Bueno basin, and 
the only one in which the species inhabits 
a lentic environment. The possibility that 
Lake Rupanco's population has already at- 
tained reproductive isolation with respect 
to the riverine populations, and therefore 
qualifies as a different species, remains an 
open question. 

As mentioned elsewhere (Jara 1986), the 
trend towards profuse spinulation in Aegla 
seems to be correlated with living in lentic 
environments. The prominent ornamenta- 
tion seen in A. d. lacustris lends support to 
this hypothesis. However, no explanation is 
available for this overgrown spinulation, 
neither from the ecological nor from the 
genetical point of view. Profuse spinulation 
in A. d. lacustris may well be an adaptive 
response to its immediate biotic or abiotic 
environment. Otherwise, spinulation could 



be a by-product of genetic changes related 
to presently unknown factors. Comparison 
of A. d. lacustris and A. d. denticulata life 
histories could be illuminating in these re- 
spects. 

If the geographical and chronological re- 
lationship between rivers and lakes in 
southern Chile is considered (see comments 
in Jara 1977, 1982, 1986), it appears evi- 
dent that lacustrine populations of aeglids 
descended from riverine stocks. These last 
would have withstood the Quaternary gla- 
cial events in non-glaciated areas, located 
along the western border of the Chilean 
Central Valley and in the Coastal Cordillera 
(lilies 1960). That scenario implies that ae- 
glids could have colonized the lakes rather 
recently (in geological time). Mercer (1972) 
estimates that lakes Llanquihue and Ru- 
panco became free of ice about 1 1 ,000 years 
B.P. Therefore, to colonize the Andean lakes, 
aeglids had to advance from west to east, 
along the rivers that drained the newly 
formed lakes. Most probably, the forerun- 
ners of A. d. lacustris were specimens of A. 
d. denticulata, which could have entered the 
young Lake Rupanco from rivers of the 
Central Valley of Osorno Province. If spi- 
nulation of A. d. lacustris is a derived con- 
dition related to modifications of the former 
riverine invader's genome, one could expect 
to find evidence of these changes by com- 
paring population samples of both subspe- 
cies by suitable techniques (protein electro- 
phoresis, caryological analysis, etc.) (Wiley 
1981). 

Acknowledgments 

The author thanks Rene Navarro and 
Gonzalo Aguilar for their assistance during 
the field work at Lake Rupanco; Dr. Ray- 
mond E. Manning of the National Museum 
of Natural History for the loan of four spec- 
imens of A. denticulata; Mrs. Patricia Araya 
and an anonymous reviewer for correcting 
the MS; Dr. Thomas E. Bowman for making 
available to the author his paper on Heterias 



VOLUME 102, NUMBER 2 



393 



exul; and Direccion de Investigation y De- 
sarrollo, Universidad Austral de Chile, for 
funding field work through grant RS-83-39. 

Literature Cited 

Bowman, T. E., R. Prins, & J. Arenas. 1987. The 
occurrence of the freshwater isopod Heterias 
(Fritzianira) exul in the Lakes Region of Chile, 
with notes on the genus Heterias (Asellota: Jani- 
ridae).-Hydrobiologia 146:275-281. 

Campos, H., J. Arenas, W. Steffen, G. Agiiero, L. Vil- 
lalobos, & G. Gonzalez. 1985. Investigacion 
de la capacidad de cultivo de salmonidos de las 
principales hoyas hidrograficas del pais. Ill An- 
tecedentes limnologicos hoya Lago Rupanco.— 
Corporacion de Fomento de la Production 
(CORFO), Santiago, Chile, 404 pp. 

lilies, H. 1960. Geologie der Gegend von Valdivia 
(Chile).— Neues Jahrbuch fur Geologie und Pa- 
laeontologie 111:30-110. (Translated into 
Spanish by J. Walper, in Publication Ocasional 
de la Universidad Austral de Chile, Valdivia, 
1970.) 

Jara, C. 1977. Aegla rostrata n. sp., (Decapoda, Ae- 
glidae), nuevo crustaceo dulceafuicola del Sur 
de Chile. — Studies on Neotropical Fauna and 
Environment 12:165-176. 

. 1980. Taxonomia y distribution del genero 

Aegla Leach (Crustacea, Decapoda, Anomura) 



en el sistema hidrografico del rio Valdivia 
(Chile).— Universidad Austral de Chile, Facul- 
tad de Ciencias, Thesis, 126 pp. 

. 1982. Aegla bahamondei, new species (Crus- 
tacea: Decapoda: Anomura) from the coastal 
mountain range of Nahuelbuta, Chile.— Journal 
of Crustacean Biology 2(2):232-238. 

. 1986. Aegla spectabilis, a new species of 

freshwater crab from the eastern slope of the 
Nahuelbuta Cordillera, Chile. — Proceedings of 
the Biological Society of Washington 99(1):34— 
41. 

, & M. T. Lopez. 1981. A new species of fresh- 
water crab (Crustacea: Anomura: Aeglidae) from 
insular South Chile. — Proceedings of the Bio- 
logical Society of Washington 94(l):88-93. 

Mayr, E. 1969. Principles of systematic zoology. 
McGraw-Hill Book Co., New York, 428 pp. 

Mercer, J. H. 1 972. Chilean glacial chronology 20,000 
to 11,000 carbon- 14 years ago: some global 
comparisons. — Science 176:1 1 18-1 120. 

Schmitt, W. L. 1942. The species of Aegla, endemic 
South American freshwater crustaceans. — Pro- 
ceedings of the United States National Museum 
91:431-520, pis. 25-28. 

Wiley, E. O. 1981. Phylogenetics. The theory and 
practice of phylogenetic systematics. John Wiley 
& Sons, New York, 439 pp. 

Instituto de Zoologia, Universidad Aus- 
tral de Chile, Casilla 567, Valdivia, Chile. 



PROC. BIOL. SOC. WASH. 
102(2), 1989, pp. 394^00 

FRESHWATER CRABS ASSOCIATED WITH CAVES 

IN SOUTHERN MEXICO AND BELIZE, WITH 

DESCRIPTIONS OF THREE NEW SPECIES 

(CRUSTACEA: DECAPODA) 

Gilberto Rodriguez and Horton H. Hobbs, Jr. 

Abstract. — Records of nine freshwater crabs frequenting subterranean hab- 
itats in Belize and Mexico are reported. Three of them are described as new: 
Potamocarcinus leptomelus, a troglophile from Veracruz; Odontothelphusa 
monodontis, also a troglophilic species, from Tabasco; and Typhlopseudo- 
thelphusa hyba, a troglobite, from Chiapas. The remaining crabs are troglo- 
philes, one of which occurs in Belize, and the others, two of which because of 
inadequate material cannot be fully identified, were found in caves in the states 
of Chiapas, Guerrero, Oaxaca, and Tabasco, Mexico. 



A study of several collections of cave- 
dwelling crabs, kindly donated to the 
Smithsonian Institution by James R. Red- 
dell, has prompted us to record additions 
to his list of Brachyura known to occur in 
subterranean habitats in Mexico, Guate- 
mala, and Belize (Reddell 1981). His list 
includes three troglobites and six others that 
are probable troglophiles. Subsequently, one 
troglobitic form, Typhlopseudothelphusa 
acanthochela Hobbs (1986), was described 
from Blind Crab Cave, 12 km SW Millio- 
nario, Cayo District, Belize, and a troglo- 
philic one, Pseudothelphusa mexicana Al- 
varez-Noguera (1987), from La Jolla [sic, 
Joya?] Cave in the State of Guerrero, Mex- 
ico. Records of the occurrence of eight ad- 
ditional species, two of which cannot be ful- 
ly identified because of inadequate material, 
are reported herein from spelean waters of 
Belize and Mexico. 

All of the "Material" cited herein is de- 
posited in the National Museum of Natural 
History, Smithsonian Institution (USNM). 
Other abbreviations employed are: cl = car- 
apace length, and cb = carapace width. 



Pseudothelphusidae Ortmann, 1893 

Potamocarcinus leptomelus, new species 
Fig. la-h 

Material. —Mexico: Cueva del Tuknel, 
Mahoilca, Porvenir Zongolica, Veracruz, 3 1 
Dec 1986, Steve and Lori Robertson, male 
holotype, cl. 13.3 mm, cb. 2 1 . 1 mm (USNM 
230080). 

Description.— Cervical groove narrow, 
almost straight, very shallow, reaching mar- 
gin of carapace. Postfrontal lobes obsolete, 
their position marked by slight depressions; 
median groove absent except for notch on 
upper margin of front. Surface of carapace 
between postfrontal lobes and front sloping 
gently ventromesially. Front low, of equal 
height throughout, bilobed in dorsal view, 
upper margin defined by series of irregularly 
placed papillae; lower margin V-shaped in 
median part, straight in lateral parts. Dorsal 
margin of orbits sinuous. Lateral orbital an- 
gle forming papillated tooth, followed by 
deep notch; rest of anterolateral margin cov- 
ered by small, irregularly placed, conical pa- 
pillae. Surface of carapace covered by very 



VOLUME 102, NUMBER 2 



395 




Fig. la-o. a-h, Potamocarcinus leptomelus, male holotype: a, Left gonopod, caudal view; b, Same, cephalic 
view; c, Same, lateral view; d, Right cheliped; e, Distal part of chela;/ Opening of efferent channel; g, Left third 
maxilliped; h, Dorsal view of carapace, i-o, Pseudothelphusa mexicana, male, from Resumidor de la Joya: i, 
Left gonopod, mesial view; j, Same, cephalic view; k. Same, subapical view; /, Same, lateral view; m, Same, 
caudal view; n. Opening of efferent channel; o, Left third maxilliped. 



small papillae, invisible to naked eye, par- 
ticularly in branchial region. 

Eyes small, with distinct faceted cornea 
and pigment, of usual shape, not filling or- 
bit. Third maxilliped with conspicuous 
notch on external margin of merus, near 
insertion of palp; external margin of is- 
chium convex; exognath 0.59 times length 
of ischium. As name suggests, all pereio- 



pods conspicuously slender. Chelae only 
slightly dissimilar in size and shape; palm 
of larger one very elongate with straight up- 
per and lower margins; fingers not gaping; 
carpus with strong internal spine preceded 
by four spinules; merus elongate with row 
of spinules on internodorsal and interno- 
ventral ridges and smaller spinules on outer 
ridge. Walking legs very long, length of third 



396 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



pair 1.6 times breadth of carapace, and me- 
rus 6.3 times as long as broad; relations be- 
tween podomeres as follows: merus 1, car- 
pus 0.33, propodus 0.54, dactylus 0.54. 
Dactylus with seven spines in upper row 
and four in two lower ones. 

Gonopod subcylindrical, straight; apical 
part bearing triangular marginal process 
overreaching apex, spiniform mesial pro- 
cess directed distomesially, and bifid ce- 
phalic process with apices pointing mesial- 
ly. Field of spines very narrow, with few 
setae placed on distal surface of apex. 

Remarks.— The gonopod of this species 
resembles that of Potamocarcinus magnus 
(Rathbun 1896:377); the mesial process is 
similar but directed distomesially instead of 
mesially; the bifid process is present in both, 
although distinctly reduced in P. leptome- 
lus; and the marginal process of the latter 
is rudimentary rather than well developed 
and cup-shaped as in P. magnus. 

Potamocarcinus aspoekorum (Pretzmann) 

Pseudothelphusa (Zilchia) aspoekorum 
Pretzmann, 1968a: 12. 

Potamocarcinus (Zilchia) mexicanus Pretz- 
mann, 1968a: 12 [nomen nudum]. 

Potamocarcinus (Zilchia) aspoekorum. — 
Pretzmann, 1971:20; 1972:75, figs. 405- 
407, 417-420. 

Potamocarcinus aspoekorum. —Rodriguez, 
1982:1 15, figs. 69d; 70f, m; 71e, i; 74a-c. 

Material. — Belize: Balam's Cave (Uchen 
Balam), Cayo District, 22-26 Mar 1979, L. 
McNatt, 1 male cl. 18.6 mm, cb. 30.1 mm 
(USNM 230082). 

Remarks. — Troglobitic characters are not 
conspicuously developed in this species. The 
walking legs are moderately slender (third 
right pereiopod of largest male with total 
length 34.0 mm, merus 3.3 times as long as 
broad), but the dorsal surface of the cara- 
pace is not pale, and the eyes are normally 
developed. 



Pseudothelphusa mexicana 

Alvarez-Noguera, 1987 

Fig. li-o 

Pseudothelphusa mexicana Alvarez-No- 
guera, 1987:1. 

Material.— Mexico: Resumidero de la 
Joya, San Gregorio, Guerrero, 23 Feb 1981, 
Steve Robertson, 2 males, cl. 18.6 and 16.5 
mm, cb. 28.7 and 25.8 mm; 1 female, cl. 
18.0 mm, cb. 28.2 mm (USNM 230084). 

Remarks.— Out specimens are from the 
type locality. Troglobitic characters are not 
obvious in this crab, although the walking 
legs are moderately slender (third right pe- 
reiopod of largest male with total length 29.5 
mm, merus width/length = 0.35). The dor- 
sal surface of the carapace, however, is dark 
and the eyes are well developed. 

Odontothelphusa monodontis, new species 
Fig. 2a-f 

Material.— Mexico: Grutas del Cocona, 
Tabasco, 26 Aug 1972, R. W. Mitchell and 
W. H. Russell, 1 male holotype, cl. 17.0 
mm, cb. 25.2 mm (USNM 230078). 

Description. —Cervical groove very shal- 
low and wide, not reaching margin of car- 
apace. Postfrontal lobes wide, flat, clearly 
delimited anteriorly, surface covered by flat 
papillae not visible to naked eye; median 
groove well marked between lobes, but not 
anterior to them. Surface of carapace be- 
tween frontal lobes and front flat. Upper 
border of front well marked by overhanging 
papillae and conspicuous notch in middle; 
lower margin slightly sinuous and lying 
slightly anterior to upper one; front high and 
excavate. Anterior margin of carapace with 
notch behind orbit, about 5 small papillae 
situated between notch and cervical groove, 
and approximately 1 5 papilliform teeth on 
rest of lateral margin. Third maxilliped con- 
spicuously wide; merus 1.43 times as wide 
as long; exognath 0.93 length of ischium of 
endognath. 



VOLUME 102, NUMBER 2 



397 




Fig. 2a-k. a-f, Odontothelphusa monodontis, male holotype: a, Right gonopod, caudal view; b, Same, mesial 
view; c, Same, cephalic view; d, Same, subapical view; e, Left third maxilliped;/ Opening of efferent channel. 
g-k, Typhlopseudothelphusa hyba, male holotype: g. Opening of efferent channel; h. Left third maxilliped; i, Left 
gonopod, caudal view; j, Same, lateral view; k, Same, cephalic view. 



Gonopod slender with apical part bent 
laterally at angle of 45 degrees; apex con- 
sisting of flat quadrangular lobe with long 
spur on proximomesial angle and papilli- 
form tubercle and small finger-like tubercle 
on cephalic surface; field of spines narrow, 
slitlike. 

Remarks. —This crab differs from its con- 
gener in possessing two tubercles on the ce- 
phalic surface of the gonopod and lacking a 
spiniform process on the distal angle of the 
mesial border. Some of the characteristics 
that have been associated with an obligate 
spelean existence are lacking in this species, 



but the dorsal surface of the carapace is pale, 
and the walking legs moderately slender 
(third right pereiopod with total length of 
30.3 mm, merus 2.87 times as long as broad); 
the eyes, however, are well pigmented. 



IRaddus sp. 



Chorrea- 
Syme, 1 

13.8 mm 



Material.— Mexico: Cueva El 
dero, Chiapas, 24 Dec 1973, R 
juvenile male, cl. 9.9 mm, cb. 
(USNM 230090). 

The only specimen available is a distinct 
ly juvenile male; however, the gonopod ex 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



hibits the flat quadrangular form character- 
istic of Raddus, and the features of the 
carapace are those of other members of the 
genus. This cave is located 1 2 km E Tuxtla 
Gutierrez (Reddell 1981:292), near the range 
of Raddus bocourti (A. Milne-Edwards, 
1866). 

Typhlopseudothelphusa hyba, new species 
Fig. 2g-k 

Potamocarcinus {Typhlopseudothelphusa) 
mocinoi. — Hobbs et al. 1977: 145 [in part : 
localities 2 and 3]. 

Material.— Mexico: Cueva de Los Lla- 
nos, 1 5 km ESE San Cristobal de Las Casas, 
Chiapas, 29 Aug 1972, J. Cooke, W. Rus- 
sell, and R. Mitchell, male holotype, cl. 12.4 
mm, cb. 18.1 mm (USNM 150633), 1 fe- 
male paratype, cl. 15.4 mm, cb. 22.8 mm 
(USNM 230081).-same locality, 5 Feb 
1972, D. McKenzie and J. R. Reddell, 1 
female, cl. 1.2 mm, cb. 24.6 mm (USNM 
1436 1 3).— Cueva de los Murcielagos, 1 5 km 
ESE of San Cristobal de Las Casas, Chiapas, 
29 Aug 1972, J. Cooke and R. Mitchell, 1 
male, cl. 12.6 mm, cb. 19.9 mm, 2 females, 
cl. 14.7 and 11.6 mm, cb. 20.9 and 16.8 
mm (USNM 150629). 

Description.— Carapace narrow (cb/cl = 
1 .46), strongly convex in anterior part. Cer- 
vical grooves forming paired, shallow 
depressions in holotype but absent in topo- 
typic female. Postfrontal lobes and median 
groove obsolete, but carapace with slight 
depression immediately posterior to front; 
latter, lacking upper margin, rounded and 
projecting forward, its lower margin bilobed 
in dorsal view. Lateral orbital angle forming 
well developed triangular tooth, followed by 
shallow notch. Margins of orbits and front 
with row of small granules. Lateral margin 
of carapace with larger, irregularly placed 
granules. Eyes reduced, lacking distinct fac- 
eted cornea and pigment. Third maxilliped 
with noticeable impression on external 
margin of merus, near insertion of palp; ex- 
ognath 0.61 times length of ischium. All 



pereiopods extremely slender. Chelipeds 
subequal in size and shape; palm cylindri- 
cal; fingers also very slender, approximately 
twice as long as palm, and armed with mi- 
nute teeth; carpus with small, hooked spine 
on internal margin; merus very elongate, 
overreaching carapace by half its length, and 
bearing rows of granules on its three ridges. 
Third pair of walking legs 1.6 times width 
of carapace; merus 5.87 times as long as 
broad, and relations of its podomeres as fol- 
lows: merus 1, carpus 0.35, propodus 0.66, 
dactyl 0.74; dactylus with six or seven spines 
in five rows. 

Gonopod wide in lateromesial plane, nar- 
row in cephalocaudal plane; distal part 
strongly bent; gonopore and field of spines 
directed cephalad; distal margin rounded 
and armed with prominent conical spines; 
marginal process cup-shaped and directed 
cephalad; and strong, triangular mesial pro- 
cess disposed in same plane as field of spines. 

Remarks. — Typhlopseudothelphusa hyba, 
like its congeners, exhibits a combination 
of advanced characters associated with its 
adaptations to a spelean environment. It 
may be distinguished from them by the tri- 
angular mesial process of the gonopod with 
a single apex. 

Pseudothelphusidae indet. species 

Material.— Mexico: Sumidero de Citla- 
lapa, Zongolica, Veracruz, 23 Mar 1981, 
Steve Robertson and Phillipe Ackamann, 1 
fragmented female, cl. approx. 19.5 mm, cb. 
approx. 33.5 mm (USNM 240112). 

Description. — Cervical groove very shal- 
low and wide, not reaching margin of car- 
apace. Postfrontal lobes rounded, flat, not 
well delimited anteriorly; median groove 
well marked between lobes and narrowly 
linear anterior to them. Surface of carapace 
between frontal lobes and over front flat; 
upper border of latter well marked by over- 
hanging papillae and bearing conspicuous 
median notch; lower margin slightly sin- 
uous and lying slightly anterior to upper. 



VOLUME 102, NUMBER 2 



399 



Front high and excavate. Anterior margin 
of carapace with notch behind orbit, about 
5 small papillae between notch and level of 
cervical groove, and approximately 15 pa- 
pilliform teeth on rest of lateral margin. Eyes 
normally developed, but unpigmented. 

Trichodactylidae 

Avotrichodactylus constrictus 
(Pearse, 1911) 

Trichodactylus constrictus Pearse, 1911:111, 
fig. 4.— Rodriguez & Manrique, 37:183. 

Trichodactylus {Trichodactylus) constric- 
tus. -Coifmann, 1939:111. 

Trichodactylus (Avotrichodactylus) constric- 
tus.— Pretzmann, 1968b:71. 

Trichodactylus (Rodriguezia) constrictus. — 
Bott, 1969:26. — Cottarelli & Argano, 
1977:210. 

Material.— Mexico: 1 km N of Palenque, 
Chiapas, 25 Jul 1973, J. Reddell, N. Ka- 
wakatsu, D. Denson, and S. R. Mitchell, 2 
males, cl. 19.8 and 12.5 mm, cb. 17.9 and 
12.9 mm (USNM 230085). -Cuevade Juan 
Sanchez, NW Acatlan, Oaxaca, 1 Jul 1976, 
T. Byrd, A. G. Grubbs, and M. Cossey, 1 
female, cl. 16.5 mm, cb. 15.8 mm (USNM 
230086). 

Remarks.— The female specimen from 
Cueva de Juan Sanchez is assigned to this 
species with doubt. It has five lateral teeth 
(not including the external orbital angle) on 
the right side and four on the left, whereas 
A. constrictus usually has two or three teeth. 

Avotrichodactylus bidens (Bott, 1969) 

Trichodactylus (Rodriguezia) bidens Bott, 
1969:25, pi. 24: figs. 68, 69. -Cottarelli 
& Argano, 1977:207, figs. 3, 4. 

Material.— Mexico: Cuevita de la Cas- 
cada Azufre, 3 km N Tapijulapa, Tabasco, 
15 Jun 1975, A. G. Grubbs, 1 juvenile male 
(USNM 230083). 

Remarks. — Bott (1969) designated the 
type locality "Arroyo del Solpho" near 



Tapijulapa, which is almost certainly the 
same as, or perhaps an extension of, that 
from which our specimen was taken. This 
juvenile is very small, and the lateral teeth 
on the carapace are not well formed, but 
even at this size, the gonopod is clearly typ- 
ical of A. bidens. 

Acknowledgments 

Thanks are extended to James R. Reddell 
for donating the material cited herein to the 
Smithsonian Institution, thereby making it 
available to us. We are also grateful to Brian 
F. Kensley and Raymond B. Manning for 
their criticisms of the manuscript. 

Literature Cited 

Alvarez-Noguera, Fernando. 1987. Pseudothelphusa 
mexicana, a new freshwater crab from the state 
of Guerrero, Mexico (Brachyura: Pseudothel- 
phusidae). — Proceedings of the Biological So- 
ciety of Washington 100(l):l-3. 

Bott, Richard. 1969. Die Siisswasserkraben Siid- 
Amerikas und ihre Stammesgeschichte. — Ab- 
handlungen der Senckenbergischen Natur- 
forschenden Gesellschaft (Frankfurt am Main) 
518:1-94. 

Coifmann, Isabel. 1939. Potamonidi della Guiana 
Inglesi.— Archivo Zoologico Italiano 27:93-1 16. 

Cottarelli, Vezio, & Roberto Argano. 1977. Tricho- 
dactylus {Rodriguezia) mensabaki n. sp. (Crus- 
tacea, Decapoda, Brachyura) granchio cieco delle 
acque sotterranee del Chiapas (Messico).— 
Quaderni Problemi Attuali di Scienza e di Cul- 
tura, Academia Nazionale dei Lincei 1 7 1(3):207- 
212. 

Hobbs, Horton H., Jr. 1986. A new troglobitic crab 
(Crustacea: Decapoda: Pseudothelphusidae) 
from Belize.— Texas Memorial Museum, Spe- 
leological Monographs 1:1-4. 

, H. H. Hobbs III, & Margaret A. Daniel. 1977. 

A review of the troglobitic decapod crustaceans 
of the Americas. — Smithsonian Contributions 
to Zoology 244:v + 183 pages. 

Milne-Edwards, A. 1866. Description de trois nou- 
velles especes du genre Bosica, Crustaces Brach- 
yures de la tribu des Telphusiens. — Annales de 
la Societe Entomologique de France (4)6:203- 
205. 

Pearse, A. S. 1911. Report on the Crustacea collected 
by the University of Michigan-Walker Expedi- 
tion in the State of Vera Cruz, Mexico.— Thir- 



400 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



teenth Annual Report of the Michigan Academy 
of Science 13:108-113. 

Pretzmann, Gerhard. 1968a. Neue siidamerikanische 
Siisswasserkraben (Vorlaufige Mitteilung).— 
Entomologisches Nachrichtenblatt (Wien) 15:1— 
20. 

. 1968b. Die Familie Trichodactylidae (Milne 

Edwards 1835) Smth [sic] (Vorlaufige Mitteil- 
ung).— Entomologishce Nachrichtenblatt (Wien) 
15:70-76. 

. 1971. Fortschritte in der Klassifizierung der 

Pseudothelphusidae. — Sitzungsberichten der 
Osterreichische Akademie der Wissenschaften, 
Mathematisch-Naturwissenschaftliche Klasse 
(1)1 79(1-4): 14-24. 

. 1972. Die Pseudothelphusidae (Crustacea, 

Brachyura).— Zoologica 42(120), part 1:1-182. 

Rathbun, Mary J. 1896. Descriptions of two new 
species of fresh-water crabs from Costa Rica.— 
Proceedings of the United States National Mu- 
seum 18:(377-389). 

Reddell, James R. 1981. A review of the cavernicole 



fauna of Mexico, Guatemala, and Belize.— Bul- 
letin of the Texas Memorial Museum 27: 1-327. 

Rodriguez, Gilberto. 1982. Les crabes d'eau douce 
d'Amerique. Familie des Pseudothelphusi- 
dae.— Office de la Recherche Scientifique et 
Technique Outre-Mer, Faune Tropicale 22:1— 
223. 

, & Fernando Manrique. 1967. El genero Tri- 

chodactylus en Mexico (Brachyura, Potamoni- 
dae).— Anales del Instituto de Biologia, Uni- 
versidad Nacional Autonoma de Mexico 37:183— 
186. 

(GR) Instituto Venezolano de Investiga- 
ciones Cientificas, Apartado 21827, Cara- 
cas 1 1 A, Venezuela; (HHH) Department 
of Invertebrate Zoology, National Museum 
of Natural History, Smithsonian Institu- 
tion, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 401^04 

A NEW DEEP-SEA CRAB OF THE GENUS 

CHACEON FROM CHILE 

(CRUSTACEA, DECAPODA, GERYONIDAE) 

Luis A. Chirino-Galvez and Raymond B. Manning 

Abstract. — Chaceon chilensis, a large species with five anterolateral teeth on 
the carapace and narrow, compressed dactyli on the walking legs, is described 
from localities off Chile, the Juan Fernandez Islands, and Isla San Felix. It is 
the only Recent East Pacific species of the genus. 



Until recently, carcinologists working with 
deep-sea crabs of the genus Chaceon from 
localities around the world often have iden- 
tified their material with the West Atlantic 
C quinquedens (Smith, 1879), or the East 
Atlantic C affinis (A. Milne Edwards & 
Bouvier, 1894); both of these species for- 
merly had been assigned to the genus Ger- 
yon (see Manning & Holthuis 1989). A 
species of Chaceon from off Chile has been 
identified erroneously in the past with both 
of these species. We describe the Chilean 
species as new herein. 

The holotype and several paratypes are 
in the Museo Nacional de Historia Natural, 
Santiago (MNHN). Other paratypes are in 
the collections of the Museo Instituto de 
Zoologia, Universidad Austral de Chile, 
Valdivia (MIZUA), the National Museum 
of Natural History, Smithsonian Institu- 
tion, Washington (USNM), the Instituto de 
Oceanologia, Universidad Catolica de Val- 
paraiso, Valparaiso (IOUV), the Museo de 
Zoologia, Universidad de Concepcion 
(MZUC), and the Museo de Historia Nat- 
ural, Valparaiso (MHNV). 

We use cl for carapace length, measured 
on the midline, and cb for carapace breadth, 
measured at and including the fifth antero- 
lateral spines, m for meter(s), and mm for 
millimeters. 

Chaceon chilensis, new species 
Figs. 1-2 

Geryon affinis.— Dupre, 1975:34.— Baez & 
Andrade, 1977:215.— Andrade & Baez, 



1980:263, 264. -Chirino-Galvez, 1985: 
R126.-Baez & Ruiz, 1985:103. [Not 
Geryon affinis A. Milne Edwards & Bou- 
vier, 1894.] 
Geryon quinquedens. — Retamal, 1977:249, 
250, fig. 1; 1981:33, fig. 17 r 9. [Not Geryon 
quinquedens Smith, 1879.] 

Previous records. —Chile: Zapallar, 
33°34'S, 71°38'W, 380-450 m (Baez & An- 
drade 1977). — Central Chile, taken in fish- 
ery for Heterocarpus reedi Bahamonde (An- 
drade & Baez 1980). 

Juan Fernandez Islands: Juan Fernandez 
Islands [33°00'S, 80°00'W] (Retamal 
1977, Baez & Ruiz 1985).-Isla Alejandro 
Selkirk [33°45'S, 80°46'W] (Chirino-Galvez 
1985).— East of Isla Robinson Crusoe and 
near Isla Santa Clara [33°42'S, 79°00'W] 
(Dupre 1975). 

Isla San Felix (Baez & Ruiz 1985). 

Material. -Chile: Off Chile, 33°35'S, 
77°42.2'W, 370 m: 3 males (in 3 lots, 
MNHN D-10824, D-10825, D-10875).- 
Isla San Felix, 26°16'S, 80°00'W, 27 May 
1969, in lobster traps: one male (cl 126.8 
mm, cb 145.5 mm, holotype, MNHN 
D-10821).-Off Quintero [32°47'S, 71°32'W], 
400 m: one male (IOUV C-005).- Off Zapa- 
llar [32°33'S, 71°29'W], 400 m: one female 
(IOUV C-003).- Off Zapallar, 360-400 m: 
one male (MHNV).— Islas Juan Fernandez: 
Isla Robinson Crusoe [33°38'S, 78°52'W]: 
three males (IOUV C-001, C-002, C-004).- 
Isla Robinson Crusoe, 270-300 m: one 
specimen (MZUC).— Off Caleta Sanchez, 
Isla Alejandro Selkirk, 150 m, in lobster 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Chaceon chilensis, new species. Male holotype, cl 127 mm, Isla San Felix: a, dorsal view; b, ventral 
view. Male paratype, cl 120.5 mm, off Chile: c, front. Male paratype, cl 1 15 mm, off Caleta Sanchez: d, dorsal 
view; e, anterior part of carapace; f, cheliped. 



traps, 1985, associated with Jasus frontalis 
(H. Milne Edwards) and Paramola rathbuni 
Porter: one male, one female (USNM 
205697), one male (MIZUA C-688). 

All specimens other than the holotype are 
paratypes. 

Description.— A large Chaceon, cl to 152 
mm, cb to 175 mm in adults, with five an- 



terolateral teeth on the carapace and later- 
ally compressed dactyli on the walking legs. 
Carapace broader than long, breadth 1.0- 
1.2 times length, inflated anteriorly, espe- 
cially at protogastric regions. Median pair 
of frontal teeth large, separated by broad, 
V-shaped sinus, overreaching similar lateral 
frontal teeth. Distance between submedian 



VOLUME 102, NUMBER 2 



403 



frontal teeth less than distance between them 
and lateral frontal teeth. Second and fourth 
anterolateral teeth more obtuse and lower 
than first, third, and fifth teeth. Distance 
between first and third teeth subequal to 
distance from third to fifth, distance be- 
tween first and second teeth much less than 
from second to third. Suborbital tooth vis- 
ible in dorsal view, not extending to apex 
of lateral frontal tooth. Carapace with curved 
ridge extending mesially from each fifth lat- 
eral tooth, surface, except for gastric, and 
branchial regions, especially granular. Che- 
liped rough dorsally, with blunt subdistal 
tooth dorsally on merus, carpus lacking out- 
er spine but with well-developed triangular 
inner spine, propodus lacking distal dorsal 
spine. Meri of walking legs lacking distal 
dorsal spine. Fifth leg: merus broad, length 
five times height, about two-thirds cb (0.6 1- 
0.65); propodus broad, length about four 
times height; dactylus about four-fifths as 
long as propodus. Dactyli of walking legs 
compressed, narrow, height at midlength 
much greater than width. 

Size.— Males, cl 107 to 152 mm, cb 124 
to 174.5 mm; females, cl 102 to 124.5 mm, 
cb 112 to 151 mm. Baez & Ruiz (1985) 
recorded males with cl ranging from 105.8 
to 124 mm, cb from 120.8 to 144.3 mm. 
Baez & Andrade (1 977) reported a specimen 
with cl 152.1 mm, cb 174.5 mm, and Re- 
tamal (1977) studied a specimen measuring 
cl 120 mm, cb 140 mm. 

Remarks. — This new species resembles 
C. fenneri (Manning & Holthuis, 1984) in 
general appearance, but differs in numerous 
features. It is a longer-legged species, with 
the merus of the fifth leg greater than 0.6 of 
the carapace breadth (less than 0.6 in C. 
fenneri), and, although the legs are longer, 
the dactylus is shorter. The dorsal ridge on 
the carpus of the walking legs is much 
smoother. The carapace is more inflated 
dorsally and more rounded posterolaterally, 
and is much more granular. The median 
frontal teeth extend further forward. The 
suborbital margin is shaped differently: it is 



.-•## 



hn. 




Fig. 2. Chaceon chilensis, new species. Male para- 
type, cl 1 1 5 mm, off Caleta Sanchez, a, orbit, ventral 
view; b, propodus of fifth leg, posterior view; c, same, 
dorsal view. 

evenly curved in C. chilensis, flattened for 
much of its length in C. fenneri. 

One of the specimens (MNHN D- 10875) 
has but three frontal teeth; it is shown in 
Fig. If 

N. Bahamonde N., in a letter of 8 August 
1973 to John S. Garth, commented that the 
fishermen called this species "jaiba blanca," 
because of its light color in life. 

Retamal (1981) called this species "can- 
grejo de profundidad." There are records in 
the literature from the following depths: 
270-300 m (Dupre 1975, Retamal 1977); 
380-450 m (Baez & Andrade 1977); 200- 
2000 m, in fishery for Heterocarpus reedi 
Bahamonde (Andrade & Baez 1980). 

Etymology.— -The specific name reflects 
the known geographic range of the species, 
from localities off central Chile. 

Distribution. —Known only from locali- 
ties off central Chile, including the Juan Fer- 
nandez Islands and Isla San Felix. 



404 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Acknowledgments 

We thank our colleagues who facilitated 
our study of material of this species in the 
collections under their care. Roy K. Kropp 
photographed the specimens from USNM 
205697. Lilly King Manning prepared all of 
the figures for publication. 

Literature Cited 

Andrade V., H., & P. Baez R. 1980. Crustaceos de- 
capodos asociados a la pesqueria de Heterocar- 
pus reedi Bahamonde 1955 en la zona central 
de Chile. — Boletin de Museo Nacional de His- 
toria Natural (Chile) 37:261-267. 

Baez R., P., & H. Andrade V. 1977. Geryon affinis 
Milne Edwards y Bouvier 1894 frente a las cos- 
tas de Chile (Crustacea, Decapoda, Brachyura, 
Geryonidae).— Boletin de Museo de Historia 
Natural, Valparaiso 10:215-219. 

, & R. Ruiz L. 1985. Crustaceos de las islas 

oceanicas de Chile depositados en el Museo Na- 
cional de Historia Natural de Santiago. Pp. 93- 
108 in P. Arana, ed., Investigaciones Marinas 
en el Archipielago de Juan Fernandez. Univer- 
sidad Catolica de Valparaiso. 

Chirino-Galvez, L. 1985. Stenotermia y biogeografia 
de gerionidos (Crustacea, Decapoda, Brachy- 
ura).— Archivos de Biologia y Medicina Exper- 
imentales 18(2):R126 (abstract). 



Dupre M., E. 1975. Lista de Crustaceos Decapodos 
citados para el Archipielago de Juan Fernan- 
dez. — CIMAR Documentos Tecnicos 8/75: 1—46. 
Centro de Investigaciones del Mar, Universidad 
Catolica de Valparaiso. 

Manning, R. B., & L. B. Holthuis. 1984. Geryon fen- 
neri, a new deep-water crab from Florida (Crus- 
tacea: Decapoda: Geryonidae). — Proceedings of 
the Biological Society of Washington 97(3):666- 
673. 

, & . 1989. Two new genera and nine 

new species of geryonid crabs (Crustacea, De- 
capoda, Geryonidae). — Proceedings of the Bi- 
ological Society of Washington 102(l):50-77. 

Retamal R., M. A. 1977. Sobre Geryon quinquedens 
Smith, 1870, en el Archipielago de Juan Fer- 
nandez.— Boletin de Sociedad de Biologia de 
Conception 51(1):249-251. 

. 1981. Catalogo ilustrado de los crustaceos 

decapodos de Chile.— Gayana (Zoologia) 44: 1 10 
pp. 



(LAC-G) Museo Sociedad Fonck, 4 Norte, 
784 Viiia del Mar, Chile (Present address) 
Department of Geology, Kent State Uni- 
versity, Kent, Ohio 44242; (RBM) Depart- 
ment of Invertebrate Zoology, National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560. 



PROC. BIOL. SOC. WASH. 

102(2), 1989, pp. 405-410 

UPOGEBIA CORALLIFORA, A NEW SPECIES OF 

CORAL-BORING SHRIMP FROM THE WEST INDIES 

(DECAPODA: UPOGEBIIDAE) 

Austin B. Williams and P. J. B. Scott 

Abstract. — Upogebia corallifora, a new species of shrimp from the West In- 
dies, lives in boreholes that it excavates in CaC0 3 substrates, dead coral and 
Pleistocene limestone. The species, here described and illustrated, is similar in 
morphology to U. noronhensis from Arquipelago de Fernando de Noronha off 
northeastern Brazil. The chelae of the new species differ in having a longer 
fixed finger and lacking a row of spines on the lower mesial surface of the palm. 



Recent ecological studies on thalassinid 
shrimps, Upogebia, in the West Indies have 
shown that an undescribed species of this 
genus constructs boreholes in "skeletons of 
dead coral or the dead bases of living corals" 
(Scott, Reiswig & Marcotte 1988), and that 
juveniles, here provisionally included with 
it, bore into limestone (Scott, Moser & Risk 
1988). We describe the new species in this 
paper. 

Upogebia corallifora, new species 
Fig. 1 

Material examined.— Jamaica: USNM 
230075. Holotype 9; Port Royal, Drunken- 
mans Cay, 3-4 m, area of dead Siderastrea 
siderea (Ellis & Solander 1786), P. J. B. Scott, 
Oct 1986. -USNM 230076. Paratypes, 2 6, 
2 9(1 ovig.); Port Royal, Drunkenmans Cay, 
approx. 2-4 m, P. J. B. Scott, 10 Oct 1986. 
One 6 and one 9 from this lot of specimens 
are to be deposited in the National Museum 
of Canada. 

Upogebia corallifora new species ? USNM 
230077. 2 9 Guv.), 2 juvs.; Discovery Bay, 
boring in Pleistocene CaC0 3 of an artificial 
reef, about 5 m depth, P. J. B. Scott, Oct 
1986 (specimens softened in Aerosol OT 
solution after being dried). 

Diagnosis.— Projections to either side of 
rostrum ending in acute spine. Postocular 
spine present. First and second abdominal 



segment lacking ventral spines; sixth seg- 
ment lacking hooked anterolateral spine. 
Telson subrectangular. Carpus of cheliped 
with one small spine on mesiodistal margin. 
Leg 2 with proximal mesioventral spine on 
merus; merus of leg 4 spineless. 

Description. — Rostrum triangular, short, 
slightly downturned; tip exceeding eye- 
stalks; dorsal pair of strong subapical spines 
followed on each side by two spines suc- 
cessively shorter in length, and separated 
from first by wide interval; ornamented dor- 
sal two-thirds of carapace surface anterior 
to cervical groove bearing pilose tufts, 
changing from spiny anteriorly to tubercu- 
late posteriorly, and angling toward sides 
posteriorly; gastric surface posterior to this 
smooth; ornamented anterior part separat- 
ed from and flanked on each side by pos- 
teriorly divergent ridge bearing crest of about 
12 spines, strongest on lateral rostral pro- 
cess and decreasing to obsolescence poste- 
riorly. Cervical groove deep and continu- 
ous, shoulder lateral to it bearing one 
obsolescent tubercle below intersection with 
thalassinidean line; latter continuing to pos- 
terior margin of carapace. Postocular spine 
present. 

Abdomen broadly and smoothly arched 
dorsally; pleura of segment 1 narrowly 
rounded posterolateral^, those of 2-5 
broadly rounded, with dense fine plumose 



406 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Upogebia corallifora, 9 holotype: a, Cephalic region, lateral; b, Anterior carapace, dorsal; c, Right 
cheliped, lateral; d, Same, mesial; e, Leg 2; h, Leg 5; i, Parts of abdominal segment 6, telson and uropods. 
Paratypes: dissociated legs 3-4, f, g. Scales: 1 , a-h; 2, i; each 1 mm. 



setae on posterolateral corner of terga 3-4, 
and in tracts on pleura of segments 3-5; 
segment 6 subrectangular, wider than long, 
lateral margin sinuous, adapted for inter- 
locking with extended uropodal exopod, 
notch anterior to posterolateral lobe contin- 
uous with oblique groove and lunate dorsal 
impression. 

Telson subrectangular, a little wider than 



long, broadest proximally, angle on antero- 
lateral margin interlocking with groove on 
central rib of uropodal endopod; posterior 
margin shallowly biarcuate and densely 
fringed with setae; transverse proximal ridge 
fairly prominent and continuous with low 
lateral ridges at each side; median groove 
obsolescent. 

Eyestalk stout, deepest at about mid- 



VOLUME 102, NUMBER 2 



407 



length, convex ventrally; eye narrower than 
diameter of stalk and directed anterolater- 
al^. 

Antennular peduncle reaching to about 
proximal one-fourth of terminal article of 
antennal peduncle, its proximal two articles 
together slightly longer than terminal arti- 
cle; flagella unequal, lower thinner ramus 
somewhat longer than thicker upper one. 

Antennal peduncle with about one-half 
its length extending beyond tip of rostrum; 
article 2 bearing tiny subdistal ventral spine; 
scale moderate, oval, sometimes with mi- 
nute terminal spine. 

Mouthparts as figured for U. jonesi (Wil- 
liams 1986); maxilliped 3 with epipod. 

Epistomial projection rather broad in lat- 
eral view, bearing small apical spine. 

Chelipeds essentially equal. Ventral mar- 
gin of ischium bearing one tiny spine. Merus 
with single subdistal dorsal spine reaching 
level of postocular spine, row of five small 
to moderate spines on ventral margin. Car- 
pus trigonal, shallow longitudinal groove 
laterally and slender spine at anterior ven- 
trolateral corner; mesiodorsal crest of al- 
most uniform small spines remote behind 
prominent spine on anterior margin and 
partly obscured by setae in proximal part 
of row, all directed distad, and three very 
small spines on anterodorsal margin mesial 
to articulation of carpus; one small spine 
near middle of distomesial margin, and an 
obsolescent spine dorsal to it. Palm oval in 
cross section, length including fixed finger 
about three times maximal height; bearing 
mixed prominent plumose and long setae 
in one sparse dorsal row, plus lateral oblique 
row of setae ending anteriorly in patch of 
long setae near base of fixed finger; mesio- 
dorsal row of small somewhat irregular 
spines ending distally in small acute spine 
above condyle of dactyl; lower mesial sur- 
face spineless but bearing low transversely 
arcuate ridge proximally. Fixed finger 
slightly shorter than dactyl and more slen- 
der, slightly downcurved in middle and ta- 



pering to slender tip, four small teeth on 
proximal prehensile edge. Dactyl much 
stouter than fixed finger, its curved upper 
surface bearing about three small, proximal 
tubercles; tip preceded on prehensile edge 
by more or less straight edge growing more 
elevated proximally and capped by obscure 
small teeth, then a toothless notch basally. 

Leg 2 reaching about to distal one-fourth 
of palm; carpus with obsolescent distodor- 
sal spine and tiny, acute, subdistal ventral 
spine; merus with small subdistal dorsal 
spine and moderate proximal mesioventral 
spine; coxa with strong proximal and small- 
er distal spine mesially. Leg 3 with slender 
ventral spines on proximal half. Leg 4 with 
spineless merus and ischium. Subchelate leg 
5 reaching beyond ischium of cheliped. 

Two arthrobranchs arranged in biserial 
rows of undivided (entire) lamellae on max- 
illiped 3 and legs 1-4. 

Uropod with obsolescent spine on pro- 
topod above base of endopod; nearly straight 
distal margin of endopod exceeded by 
curved margin of exopod, both rami with 
dense fringe of setae and minute scattered 
spines or granules on distal margin; endo- 
pod with low, median longitudinal rib, and 
less prominent lateral rib having shallowly 
concave margin except for rounded, prox- 
imal shoulder; exopod with three ribs, me- 
sial sturdiest one bearing obsolescent spine 
proximally, intermediate rib longer, lateral 
rib least evident of all, slightly curved out- 
ward but shallowly concave proximally. 

Measurements in mm. — Holotype $, 
length anterior carapace 4.6, length cara- 
pace 7.3, length chela including fixed finger 
4.5, greatest height chela 1.2. Paratype 6, 
length anterior carapace 2.4, length cara- 
pace 3.3, chelae missing. 

Known range.— Specimens examined are 
limited to Jamaican localities listed above, 
although one additional specimen of what 
appeared to be U. corallifora was collected 
from dead coral on the reef of Bellairs Re- 
search Institute, Barbados in 1981. The lat- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. a, Mud-lined borehole of juvenile U. corallifora ? exposed in a slabbed section of Pleistocene limestone, 
with part of one individual intact; b, SEM micrograph x 560 of burrow lining composed of organic and inorganic 
constituents. 



VOLUME 102, NUMBER 2 



409 



ter specimen was lost and cannot now be 
verified. It is possible that the species is 
widely distributed, though rare, in other 
parts of the Caribbean. 

Known distribution of U. corallifora is 
patchy. No specimens were found in recent 
reef coral on the north coast of Jamaica, 
where 188 kg of coral rubble and 97 kg of 
living coral were surveyed quantitatively. 
All specimens boring in recent reef rock were 
found in much smaller, qualitative samples 
taken at Drunkenmans Cay on the south 
coast, near Kingston. Bioeroders are gen- 
erally much more abundant in that area than 
on the north coast, including a species of 
boring mussel Lithophaga aristata (Dillwyn 
1817) which does not occur at all in Dis- 
covery Bay on the north coast of the island 
(Scott 1988). Juveniles questionably re- 
ferred to U. corallifora were, however, found 
in Pleistocene limestone placed in Discov- 
ery Bay during the building of an artificial 
reef (Fig. 2). This limestone is much harder 
and denser than natural, recent reef rock. 
These juveniles have more robust chelipeds 
than the adults described above, but resem- 
ble the adults in other respects. We ques- 
tionably refer them to the new species until 
more material is available for comparative 
study. 

Habitat.— The habitat of U. corallifora 
partly overlaps that of the much more abun- 
dant U. operculata Schmitt 1924. Both 
species inhabit dead coral skeletons in Ja- 
maica, but occurrence of the former is much 
restricted. It has not been found with bore- 
holes opening from living corals, which are 
a common habitat for U. operculata (Scott, 
Reiswig & Marcotte 1988); see Warme 
( 1 975) for distinction between boring in hard 
and burrowing in soft substrates. A coral- 
boring species, U. trypeta Sakai (1 970, 1 982), 
has also been reported from Japan. 

Boreholes of U. corallifora are similar in 
appearance to those of U. operculata (Klee- 
mann 1984; Scott, Reiswig & Marcotte 
1988). Pairs of individuals (one 6, one 9) 
live in complex, multi-chambered excava- 



tions which are just wide enough to allow 
the occupants to somersault and reverse di- 
rection. Excavations are lined with a mix- 
ture of mud, coral fragments and organic 
material. Energy dispersive X-ray analysis 
(EDAX) of borehole lining material re- 
vealed the presence of Ca, Si, Mg, Al, CI, K 
& S. Openings to borings are difficult to 
detect on the irregular surface of dead coral; 
each opening lies a few millimeters below 
the surface in a smooth conical pit and is 
much narrower than the remainder of the 
hole to which it gives access. Adult shrimps 
are trapped in the borings by the restrictive 
openings. 

Remarks. — Upogebia corallifora is simi- 
lar to U. noronhensis Fausto-Filho 1969 
(comparison made with paratype 9$, No. 
163, carcinological collection, LABOMAR, 
Universidade Federal do Ceara, Brazil, Fer- 
nando de Noronha, Baia de Sueste, Fausto- 
Filho, 7/8/68), which in turn has some sim- 
ilarity to U. spinigera (Smith 1871) of the 
eastern Pacific, but U. corallifora is appar- 
ently a smaller and less spinose species than 
either of the latter. Upogebia corallifora keys 
out to U. noronhensis in Williams (1986: 
10), but the two can be separated by means 
of the following emendation to the con- 
cluding couplet of his key to Atlantic species 
of the genus. 



- Chelae with no spines proximal to 
fixed finger on ventral keel of palm 



7 



7. Chela with fixed finger about equal 
in length to dactyl; palm spineless 

on lower mesial surface 

U. corallifora, n. sp. 

- Chela with fixed finger about Vi 
length of dactyl; palm with row of 
more or less distinct spines on lower 

mesial surface 

U. noronhensis Fausto-Filho 

Etymology.— The name, a noun form 
from the Latin "corallum," coral, and 
"foro," bore or pierce, alludes to the coral- 
boring capabilities of the species. 



410 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Acknowledgments 

We thank Dr. Jeremy Woodley and staff 
of the Discovery Bay Marine Laboratory, 
University of the West Indies, and Dr. Dun- 
bar Steele, Chairman, Department of Zo- 
ology, University of the West Indies, Mona 
Campus, for facilities and assistance in the 
field. Many thanks also to dive buddy and 
cheerful coral basher Ian Sandeman. Con- 
tribution No. 437, Discovery Bay Marine 
Laboratory, University of the West Indies. 

Literature Cited 

Kleemann, K. 1984. Lebensspuren von Upogebia 
operculata (Crustacea, Decapoda) in karibisch- 
en Steinkorallen (Madreporaria, Anthozoa).— 
Beitrage zur Palaontologie von Osterreich, In- 
stitut fur Palaontologie der Universitat Wien 1 1 : 
35-49. 

Sakai, K. 1970. A new coral burrower, Upogebia try- 
peta sp. nov. (Crustacea, Thalassinidea) col- 
lected from Amami-Oshima, Japan.— Publica- 
tions of the Seto Marine Biological Laboratory 
18(l):49-56. 

. 1982. Revision of Upogebiidae (Decapoda, 

Thalasssinidea) in the Indo-West Pacific re- 
gion.— Researches on Crustacea, The Carcino- 
logical Society of Japan, Special Number 1:1- 
106, plates A-C. 

Scott, P. J. B. 1988. Distribution, habitat and mor- 



phology of the Caribbean coral- and rock-boring 
bivalve, Lithophaga bisulcata (d'Orbigny) (My- 
tilidae: Lithophaginae).— Journal of Molluscan 
Studies 54:83-95. 

, K. A. Moser, & M. J. Risk. 1988. Bioerosion 

of concrete and limestone by marine organisms: 
A 13 year experiment from Jamaica.— Marine 
Pollution Bulletin 19(5):2 19-222. 

, H. M. Reiswig, & B. M. Marcotte. 1988. Ecol- 
ogy, functional morphology, behaviour, and 
feeding in coral- and sponge-boring species of 
Upogebia (Crustacea: Decapoda: Thalassini- 
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495. 

Warme, J. E. 1975. Borings as trace fossils, and the 
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